MICROFICHE REFEREN LlE3RARY

A project sf Volunteers in Asia

Animal Husbandry in the Tropics

by G. Williamson and

W.J.A. Payne

Pub1 ished by:

Longman Group Ltd. Longman House Burnt Mill, Harlow

Essex CM20 2JE

ENGLAND

Available from:
same as above

Reproduced

by

permission. as those
in any I
'

Reproduction of this microfiche document form is jubjoct to the sai:;:ti r&%tions of the original document.

An Introduction to Animal Husbandry in the Tropics

TROPICAL

AGRICULTURE

SERIES

The Tropical Agriculture Series, of which this volume forms part, is published under the editorship of D. Rhind, CMG, OBE, BSc, FLS, FIBiol.
ALREADY PUBLISHED

Tobacco B. C. Akehurst Tropical Pasture and Fodder Plants A. V. Bogdan Coconuts R. Child Yams D. G. Coursey Sorghum H. Doggett Tea T. Eden Rice D. H. Grist Termites W. V. Harris The Oil Palm C. W. S. Hartley Tropical Farming Economics M. R. Haswell Sisal G. IV. Lock Cattle Production in the Tropics Volume I
W. J. A. Pap2

Cotton A. N. Prentice Bananas IV. W. Simmonds Tropical Pulses J. Smartt Agriculture in the Tropics
C. C. Webster and P. N. Wilson

An Introduction to Animal Husbandry in the Tropics G. Williamson and W. J. A. Payne

Cocoa G. A. R.
Wood

An Introduction to

Animal in the Tropics

Third Edition
c.
Wi~hllSOIl

rY

CBE, MRCVS, DVSM MA, Dip. Agric. Sci., PhD, FIBiol.

W.

9.

A.

PaJ4W

Longman

London and New York

Longman Group Limited Longman House, Burnt IMill, Harlow, EssexCM20 2JE, England
Associated Companies throughout the World Published itz the Unlked States of America by Longman Inc., New York

0 *G. Williamson and W. J. A. Payne 1959 This edition 0 Longman Group Limited 1965, 1978 All rights r&served;no part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior written permission of the Publishers,
First published 1959 Second edition 1965 Third edition 1978 Reprinted i 980 Third impression I984 c.

Library of CongressCataloging.in Publication Data Williamson, Grahame. An introduction to animal husbandry in the tropics. (Tropical agriculture series) Includes index. 1. Stock and stock-breeding-Tropics. I. Payne, William John Arthur, joint author. II. Title. SF75.W5 1977 636.00913 76-58000 ISBN O-582-468 13-2 Printed in Hong Kong by Astros Printing Ltd

Preface to third edition

The original concept, when the first edition of this book was published in the late 195Os, to summarize such knowledge of tropical domestic was animals as was available at that time and which could be considered useful for students, livestock extension specialists, progressive livestock owners and administrators concerned with the development of livestock industries and the economy. The dificulties of adequately dealing with such a wide subject in one volume were anticipated and discussed when the first edition was published. These difficulties have now multiplied as the frontiers of fundamental knowledge in animal science have steadily expanded and an ever increasing volume of experimental data in different aspects of tropical animal husbandry has been published. In addition, the total number of animal science students in tropical countries has increased very rapidly, livestock extension agencies have grown and multiplied, there are more technically educated tropical livestock owners and livestock development projects are numerous in tropical countries. In view of all these circumstances it was decided that it was not only essential to expand and upgrade the information content of this third edition but that it was also necessary to rewrite almost all the original, and to add additional chapters. This edition is, therefore, in some respects a different type of book from its predecessors.It is also different in another manner. One ui the authors (G. W.) has now died and the edition is the sole responsibility of the other author (W. J. A. P.). The opportunity is taken by the author to express thanks to all collaborators for the revision of their original contributions or for new contributions, to the General Editor for his advice and to all others who have assisted in the publication of this edition.

Contents

PREFACE TO THIRD

EDITION

Part 1 Basic principles 1 The effect of climate

Trc$ical climates - The effect of climate on livestock: The direct eflect of
climate on the animal: Acclimatization. The indirect eflect of climate on the animal;

2 Maintenance of health by E. R Lindley (including Parasitism by R. P. Lee)

Importance of animal health - Relations between livestock owners and authorities - Diseases:Classification: Transmission: Immunity: Prevention and control - Parasitism - Diseasescaused by worms : Roundworms or nematodes Flukes or trematodes; Tapewormsor cestodes Diseasescaused ; by arthropod parasites.

29

3 Nutrition and feeding

Components of food: Water;
Fats; Minerals; Energy content; Nitrogenous compounds; Carbohydrates; Vitamins; Additives - Evaluation of foods: Digestibility; Pratein content - Feeding standards and requirements Types of feedavailable: Succulents; Roughages; Concentrates; Other, including unconventional, feeds - Storage of feeds - Forage - Types of forage: Natural grazings or range; Cultivated pasture,fodder and browse- Management of forage: Natural grazings; Cultivated pastures and fodder crops Forage conservation: Hay; Silage; Artificially driedforage- Nutritive value of forage: Toxic substances; Intake and digestibility; Effect of climate on nutritive value; Nutritive values.

62

4 Reproduction and breeding

The reproductive cycle: Factors aflecting the oestral cycle;, Managerial
considerations; Fertilization and gestation; Injluence ojhormones on reproduction - The basis of inheritance,: Cell division - The inheritance of characteristics: The inheritance ofsex - Mutation - Changesin the number

1145

of chromosomes - Genetic-en&ronmentaI interactions - Maternal influence- Animal breeding practices- Selection; Inbreeding; Crossbreeding - Breeding livestock for tropical environments: The use of indigenous
breeds; The importation and use grading of indigenous breeds.

of exotic

breeds; Crossbreeding

and/or up

... vlll

Cortenrs

Part II
5 Cattle

Husbandry
196

Numbers and distribution - Origin: Asia; Africa; The Americas; Oceania - Types of tropical cattle - r rsification of breeds; Representative and cattle - Introduction - Breeding: the important tropical breeds Managerial systems: The subsistence most suitable milking breed! dairy farmer; The large-scale producer or peasant producer; The spr -. - Management and feeding: T.. ca!f; The heifer; The milking heifer and cow; The bull - The planning of dairy farms and their equipment: Buildings, Equipment for milking - Beef cattle - Beef production systems: Extensive systems; Irztensive systems - Planning and development of beef enterprises - Transport of trade stock : Movement an foot; Rail transport; Road transport; Air transport - Beef quality - Working ca@le - The working ability of cattle - Choice of animal -. Management - Feeding Diseaseand ill-health - Choice of equipment: Yokes; Implements - Estimation of body and carcase weight - Dentition of cattle as an iodication of age.

6 Buffalo by P. Bhattacharya
Origin and distribution - General characteristics - Four milk breeds of India and Pakistan: Murrah; Nifi; Surti; Jafirabndi - Growth and Reproduction: Growth; Reproduction - Nutrition L Adaptability - Production: Work; Milk; Meat - Disease.

398

7 Sheep
Utility - Origin - Numbers and distribution - Breeds: Africa; Americas; Asia - Reproductive behaviour - Diseases and parasites - Feeding and management; Productivity: Meat; Milk; Wool.

436

8 Goats
by C. Devendra
Importance - Distribution - Breeds - Breeding - Systems of Management - Rearing - Feeding - Potential for increased production.

463

9 Camels
Origin - Numbers and distribution - Breeds - Productivity - Special physiological characteristics of the camel - Reproductive behaviour Feeds and feeding - Management: Working camels; Dentition.

484

10 Llamoids or New World Camelidae

by Saul Fernimdez-Baca
Importance - Origin - Types and distribution - Reproductive behaviour: Puberty; Breeding season; Oestrus and ovulation; Oestrus and mating behaviour; Fertilisation rates and embryonic mortality; Pregnancy diagnosis; Gestation and parturition - Nutritional characteristics - Management: Art[ficiiZl insemination; Dentition - Disease - Productivity: Fibre production; Meat production; Milk production; Work; Factors resulting in low productivity.

499

Contents

ix
519

I1 Game as a source of meat

Reasons for the preservation of wild game - Tropical regions where game could be used for large-scale meat production : Semi-arid regions; Savanna and steppe regions; Humid,forest regions; Montane regions -- Methods of using wild game for meat production: The sustained cropping of game populations; The management of game on ranches and farms; Domestication of wild game - Game Carcases and Meat.
12

Pigs
The worldwide distribution of pigs - origin of domestic pigs - major breeds used in the tropics: British breeds; American breeds; Other heeds from the mid-latitudes: Southeast Asian breeds,. African breeds; Central and South Ati~e/rcan breeds - Reproductive behaviour: Iufertility; Eflect of climate Selection of breeding pigs: Art$cial insemination; Crossbreeding - Disease and parasites - Feeding: Feeds corrtainirlg mainly carbohydrates; Feeds containing mainly protein; Miscellaneous .feeds; Additives; Preparation of feeds; Methods of calculating rations - Management: Adaptive physi&gy; Systems of management; Methods of management.

541

13 Poultry
Fowls - Origin - Breeds - Effect of climate - Feeding: Feed requirements; Specialfeed requirements in a tropical climate; Feeding methods.for the small poultry-keeper - Health and disease - Management: Breeding stock; Eggs for hatching; Incubation; Brooding; Rearing; Layers; Broiler and other meat birds; Housing; Equipment - Ducks- Origin -Breeds and their productivity - Breeding and incubation - Management after hatching: Rearing; Choice of system; Disease; Feeding - Geese - Origin - Breeds and their productivity - Breeding - Management after hatching: Rearing; Adult birds; Feeding - Turkeys - Origin - Breeds - Breeding - Management after hatching: Brooding; Rearing; Feeding; Disease - Guinea fowl - Ostrich Marketing.

595

Part III Animal products 14 Milk and milk products

by (the late) James N. Warner
The dairy situation - Organizing the supply of milk - Milk quality and its conservation - The composition of milk - Preparation of miik for the market - Organizing dairy operations - Milk products: Curd; Cheese; Butter and ghee; Chhena and khoa; Condensed milk; Dried milk; Ice-cream.

654

15 Meat and carcase by-products

by I. Mann
Meat - Avoidance of preventable waste; National livestock development plans - Slaughtering and processing faciJities: Slaughtering facilities Requirements for efficient slaughtering - Freld abattoirs - Preserved meat - Carcase by-products - Hides and skins: Pre-slaughter care; Avoiding damage in the slaughterhouse; Treatment after Paying; Suspension-drying equipment; Treatment after drying; Tools and equipment; By-products Bones - Blood - Meat meal - Commercial by-products pl.ant - Hooves and horns - Casings - Hair - Ruminal contents - Compost-making.

679

Conten Is

16 Wool production by (the late) G. R.

Mode

Genera1 information on tropical wool sheep - Factors influencing wool production - Factors influencing the number of sheep - Breeding sheep for wool production.

APPENDIX APPENDIX
INDEX

1 Nomadism in Africa by P. R. Bake1 2 itiarki;lg livestock for identification

Part I: Basic Prinniples

Chapter 1

The Effect of Climate

Tropical climates
The term tropical is used geographically to designate the area between the Tropics of Cancer and Capricorn. However, climate in this region is not uniform and it is meaningless to talk of a typical tropical climate. It varies with unalterable factors such as latitude, altitude, distribution of land and water, soils and topography and variable factors such as ocean currents, winds, rainfall and vegetation. The interaction of all these factors results in specific micro-climates at specific localities. However, for all their diversity tropical climates exhibit certain common characteristics. Except in the very dry areas daily and seasonal temperature variability is relatively small, being least at the equator. Day length is also fairly constant throughout the area, so that differences in the total hours of sunshine and the total solar radiation depend primarily on the degree of cloud cover. Climate is a combination of elements that include temperature, humidity, rainfall, air movement, radiative conditions, barometric pressure and ionization. Of these, temperature (Figs. 1.1 and 1.2) and rainfall (Fig. 1.3)are the most important. In practice, effective rainfall, that is the amount ultimately available to the vegetation, is a more important index than total rainfall. Many attempts have been made to classify climates, the best-known classifications being those of Koeppen (193 1) and Thornthwaite (1948). Holdridge (1967) has recently proposed a Life Zone Ecology classification combining latitude, altitude, rainfall and mean temperatures that is particularly useful for agriculturists. A simple classification is that proposed by the United States Department of Agriculture (1941). In this system climate in the tropics is classified into the following categories: equatorial or super-humid, humid, sub-humid, semi-arid and arid. As shown below and in Fig. 1.4 and 1.5, it has been found that there is a close relationship between these climatic zones and the major vegetation climax types and soils, and a modified form of this simple classification is used here. 2

Tropical climates

Climate

Vegetation

Soil

Equatorial or super-humid Humid Sub-humid Semi-arid Arid

Rain-forest Forest Grassland Steppe and thornbush Ephemeral and desert

Podzols (grey-brown, red and yellow) and laterites Chernozems and degraded chernozems Chestnut and brown soils Sierozems and desert soils

PACIi-lc

OCEAN

Fig. 1.1 January isotherms of the world in C: slightly simplified.

Fig. 1.2 July isotherms of the world in C; slightly simplified.

The Eflect of Climate

:.

1.

Tropical climates

Latitude Fores
icai Rain Fores

m m m m

Scrub and Thorn

F:wyt

Middle-Latitude

Forests

Mediterranean Scrub Fores1 Br3r~d~k&~oW~d BroadleafConiferous Forest

Steppe (Tropical and Middle latitude) Desert Shrub and Desert Waste

Fig. 1.5 World distribution of the principal plant associations.

Tropical climates

The general extent of these major climatic divisions in the tropics is shown in Fig. 1.4. This map does not, of course, show local variations in climate within the larger climatic zones. This climatic zone is characterized by constant heat, rainfall and humidity. The mean annual temperature varies around 27C (80F) and total annual rainfall is usually within the range of 2,032 to 3,048mm (80 to 120in). Although in any 2- or 3-week period there is almost always an excessof precipitation over evaporation, it is not usual for rainfall to be evenly distributed throughout the year. In some areas, one seasonis slightly wetter, while in others there are two wetter seasons. Equatorial climates are found 5 to 7 latitude north and south of the equator and specifically in the Congo basin and part of the Guinea coast of Africa, in the Indian sub-continent, the Malaysian peninsula, Indonesia, New Guinea, the southern Philippines and the Amazon basin. It is also found in isolated areas further from the equator, such as the west coast of Colombia and the east coast of Madagascar. The typical vegetation of this climatic zone is tropical rain-forest, characterized by a multiplicity of evergreen tree species,some of them hardwood, covered with epiphytes and intertwined with lianes. Climatic stress on domestic livestock is considerable in this region. In general, indigenous domestic breeds of livestock are not numerous, but the climate favours plant growth so that forage could be plentiful and available all the year around. Internal and external parasites are favoured by the climate and animal products rapidly deteriorate when stored. The apparent high fertility of tropical rain-forest soils is illusionary rather than real as they rapidly deteriorate once the forest has been cleared. Although the potential for animal production is high, a great deal of basic and applied researchis neededbefore this region can become capable of supporting a dense and productive animal population.
Equatorial or super-humid climate.

This climatic zone is characterized by high though seasonaltemperatures, humidity and rainfall. Temperature extremes are wider than in the equatorial region. There are usually three seasons: cool-dry, hot-dry and hot-wet. Climates of this type are found adjacent to the equatorial areas,north and south of the equator, in the monsoonal lands. Somewhat similar climates also occur on the windward side of volcanic oceanic tropical islands. The natural vegetation of the zone is rain-forest, but plant growth is not quite so vigorous as in the equatorial zone, deciduous are mixed with evergreen speciesand there are fewer epiphytes and lianes. Climatic stress on domestic livestock is not quite so severe as in the equatorial areas, but forage supplies are more seasonal.
Humid climate. Sub-humid climate.

In general the sub-humid areas are found north and south of the humid forest areasin the northern and southern hemispheres,

The Eflect of Climate

respectively. They are characterized by a more seasonal climate tian in the humid zone. There is usually a relatively short rainy season and a longer dry one, though in some regions there are two rainy seasons, Temperature variations are much wider, with hotter summers due to a high intensity of radiation combined with a longer day. The natural vegetation of theseareas is usually some form of savanna, an open grassland association interspersed with trees. Large areas of savanna are found north and south of the equator in Africa, particularly in the east of the continent. There are also large areas in the Indian peninsula, inland in Southeast Asia, in northern Australia, in Central and South America and on the leeward side of many oceanic tropical islands. There are many types of savanna with various combinations of high, medium and low trees and tall and short grasses.In the wetter areas savanna imperceptibly merges into dry-land forest, such as the miomho of Central Africa, while in the drier areas it merges into open short-grass steppes or desert scrub. This is one of the regions where nomadic livestock husbandry has flourished in the past in Africa and Asia and where European settlers in the Americas and Australia established a ranching industry. It is also a region where, in the past, wild game flourished in vast numbers. Generally, climatic stress on domestic livestock is less intense than in the more humid areas, but forage production is very seasonal and nutritional stress can be a major problem. This is also the region where epizootic animal diseases are rife, though some internal and external parasites of domestic livesto& are easier to control in this region than in the more humid forest areas. This climatic region is characterized by extremely seasonalconditions, with relatively low rainfall and very long dry seasons. Diurnal and seasonal temperature fluctuations are very wide, humidity is low for most of the year and there is a high intensity of solar radiation due to the dry atmosphere and clear skies. Although total rainfall may be within the range of 254 to 508mm (10 to 20 in), it may be very intense when it falls and very irregular in incidence. There are large semi-arid areas in Africa, north and south of the savanna regions, in western Asia and India, and in northern Australia and smaller areas in North, Central and South America. There are also some small lowlying oceanic tropical islands that possess a semi-arid climate. This climatic region is more suited for livestock production than for any other form of agriculture, though productivity is severely curtailed by lack of feed and water, with consequent nutritional stress, as well as by climatic stress. Internal and external parasites can be controlled with relative ease,though control of epizootic disease is more difficult. Often semi-arid areas are more suitable for the management of sheep, goats and camels than they are for cattle, and in the driest areas
Semi-arid climate.

Tropictzl chates

wild game may be the most suitable and economic exploiters of the environment. There are few truly desert areas in the tropics as the great deserts of the world are found in the sub-tropics or even further from the equator. However, small areas of desert are located within the tropics in the southern Sahara, southwest Arabia and the Pacific coast of northern Chile. Tropical desert climates are characterized by temperature extremes of the order of 0 to 52C (32 to 122F) and by the fact tliat there is no seasonal rhythm in rainfall, which is insig,nificant in total amount. Deserts may seasona.lly support very limited numbersof livestock, and in Africa and western Asia nomadic herdsmen may at certain times follow the rain showers across the deserts, feeding their herds and flocks on the ephemeral plants that spring up as soon as it rains. In parts of Arabia and the Sahara a unique and particularly valuable type of desert vegetation occurs irregularly only after a specific sequence of climatic phenomena. It is known in Arabic as gizzu. This simplified account of tropical climates does not take into account one other major zone, the montane region. A very considerable area of land in the tropics is situated at an altitude varying between 305 and 1,524m (1,000 and 5,000ft) and an appreciable area lies above 1,524m (5,000ft). Altitude influences climate in at least four ways. Mean annual temperature decreasesby 1~7C(3F) for every 305 m (1,000 ft) increase in altitude. This decrease is even higher on oceanic islands or where there are very steep mountains. Secondly, the higher the altitude the larger the diurnal though not the seasonalvariation in ambient temperature. Thirdly, rainfall is usually greater at higher altitude and there are more cloudy days. Finally, the higher the altitude the lower the atmospheric pressure, the latter being halved at 5,486m (18,000ft). Three of these influences -namely a decrease in mean annual temperature, an increase in diurnal temperature variation and a higher rainfall -are likely to assist in the improvement of livestock productivity. It is difficult to generalize, but in equatorial and humid tropical rainforest areas the speciescomposition of the forest alters with increasing altitude and gradually merges into other vegetation complexes. In the drier tropical areas the open plains vegetation merges into cool, humid forest as the annual rainfall increases with increasing altitude. Above 3,965m (13,000ft) there is usually an alpine-type vegetation even on the equator, To date little has beendone to exploit these montane areas for livestock production, though they offer opportunity for the development of dairy industries based on temperate-type dairy cattle, temperate-type beefbreeding operations for the production of bulls that could provide semen for use in crossbreeding schemesat lower altitudes, large-scale hill sheep production, and in South America major increases in the production of
Arid climate.

10

The E@cS qf Climate

meat and wool from the domesticated and wild llamoids such as the alpaca, the llama and the vicuiia.

The effect of climate on livestock

Livestock production in all tropical countries is affected by the climate in two ways. First, by a direct influence on the animals utilized, and secondly by indirect effects on the animals environment. The direct effect of climate on the animal Experimental evidence on the direct effect of climate on domestic livestock has been obtained from two sources: direct observations in the field and observations on livestock kept in controlled-temperature laboratories or psychrometric chambers. The disadvantage of direct observations is that it is difficult to set up adequately controlled field experiments, while the disadvantage of using a psychrometric chamber is that only a small number of the larger domestic livestock can be studied at any one time while it is known that there are profound differences between species (Findlay, 1954) breeds or types within a species (Worstell and Brody, 1953)and individuals within a breed (Payne and Hancock, 1957)in their ability to withstand the direct effects of climate. All domestic livestock are homeotherms. That is, they attempt to maintain their body temperatures within a range most suitable for optimal biological activity. The normal range in mammals is 37 to 39OC (98.6 to 102.2F),while in birds it is 40 to 44C (104 to 111*2F)though there are some exceptions. Typical deep-body temperatures of some domestic livestock are shown in Table 1.1.
Table 1.1 Typical deep body temperatures c$ domestic livestock Type of livesrock Deep body temperature -___-__ (C) Horses Asses Cattle Camels Sheep Goats Pigs Fowl (chickens) Ducks Geese Turkeys 37.2-38.2 36,0-38.0 380-39.3 36.0-38.0 38-3-39-9 38.7-40.7 38.9-39.4 41.9 42.1 41.3 41.2 V) (99-O- 100-S) (96.8-l 00.4) (I 00.4-l 02.8) (96.8-I 00.4) (100.9-I 03.8) (101-7-105.3) (102~~103.0) (107.4) [E-~] (106:2)

The effect of climate on livestock

11

In order to maintain their body temperature while subject to a wide range of environmental conditions, domestic livestock must preserve a thermal balance between their heat production or gain from the environment and their heat loss to the environment. This thermal balance can be expressed by the equation:
M-E+F+Cd+Cv+R=O

where M is the metabolic heat production, E the heat loss from skin and respiratory passagesby evaporation, F the heat lost or gained bringing ingested food and/or water to body temperature, Cd heat lost or gained by direct contacts between the body and surrounding surfaces, Cv heat lost or gained by convection due to contact between the air and skin and/or !inings of the respiratory passages, and R heat lost or gained by radiation. Metabolic heat production depends on: 1. Basal heat production for maintaining essential body processessuch as deep body temperature, cardio-respiratory activities and muscle tone ; 2. Digestive heat production that varies with the type of digestive system the animal possesses on the quantity and quality of the food that and it ingests; 3. Muscular heat production that varies according to how much the animal moves about grazing, etc., and 4. Increased metabolism due to productive processessuch as growth, milk production and reproduction. In general, the means by which domestic livestock can vary their heat production are limited in comparison with the methods by which they can dispose of heat. They can reduce productive processesand muscular heat production and to a more limited extent digestive heat production, but they cannot normally reduce basal heat production as minimal body processesmust be maintained. Of the methods of heat loss available to domestic livestock, evaporative loss is potentially the most important under normal circumstances. It depends on the ambient air temperature, the amount of available moisture, the area of evaporating surface, the absolute humidity of the air surrounding the animal and the degree of air movement. The factors in this situation influenced by the animal are some part of the available moisture and the area of the evaporating surface. The amount of available moisture normally depends upon the quantity of sweat and insensible perspiration produced by the animal, unless it is mechanically sprinkled with water. The area of the evaporating surface depends upon the surface area of the animal and the size of the lungs, as considerable evaporation is achieved by the mechanism of panting. In the case of cattle, although all types possesssweat glands, those of the Bos hdicus breeds usually have a higher total volume than those of

12

The Effect of Climate

B. tawus breeds (Yeates, 1965). Even if all cattle possesssweat glands

and some types sweat more freely than others, cattle in general are poor sweaters as compared with some other mammalian species, as are buffaloes, sheep, goats and pigs. Cattle and sheep to some extent make up for their lack of sweating by having relatively high respiration rates. Buffaloes possessfew, if any, sweat glands. Poultry possessno sweat glands, as birds could not possibly evolve sweating as a cooling mechanism and still fly. If they had to evaporate water from their skin the air between the feathers would have to be constantly renewed and this would cause turbulence and drag in flight. They accomplish some evaporative cooling by panting and the extensive air-sac system connected with their lungs may also have an important heat-regulatory function. The heat gained or lost by the animal bringing the food and/or water ingested to body temperature can have a considerable effect on total heat production and loss. Any water consumed in excessof metabolic needs, at a temperature lower than body temperature, and then excreted at body temperature as urine or in the faeces,assists in reducing the heat load on the animal. Lowering the temperature of ingested water is known to have a more marked effect on the heat load than increasing the volume ingested. This effect has practical significance and workers in California, including Kelly et al. (1955),have shown that cooling drinking water can increase the liveweight gain of beef cattle managed at high ambient temperatures. The ability of livestock to lose heat through conduction is very limited. Convection heat loss is of course increased when cool breezes blow on the animal, and increased air movement may also increase evaporative heat loss. Consequently, livestock accommodation in the tropics should always be built in such a way as to encourage maximal air movement on and around the animals. Solar radiation may not only increase the heat load on the animal but also directly affect the skin, causing skin cancers and other photosensitive disorders. This means that the colour and thickness of the skin are of some importance as adaptive mechanisms. A pigmented skin is considered more desirable than an unpigmented skin everywhere in the tropics and the majority of tropical-type breeds possesspigmented skins, while many temperate-type breeds possess unpigmented skins. Individuals of the Hereford breed of cattle often suffer from an eye condition known as epithelioma because they possessunpigmented eyelids, and white-skinned breedsof pigs such as the Large White are particularly susceptible to sunburn. The amount of solar radiation absorbed by the coat of the animal is partly determined by its colour. Approximately half the energy in the solar spectrum is in the visible and half in the invisible infra-red portion. The proportion of the visible portion that will be absorbed by an animal can be approximately estimated by the colour of the coat, as a white

The e@xt of climate 011livestock

13

surface may absorb only 20 per cent while a black surface may absorb 100 per cent of the visible radiation. Energy from the invisible infra-red part of the solar spectrum is completely absorbed whatever the colour of the coat. Colour is not the only factor that affects the influence of solar radiation on the heat load of the animal-length, density and condition of the hair may also have some effect. Smooth-coated animals with short hair appear to be more heat tolerant, so that Bonsma (1949) suggestedthat the degree of felting of samples of cattle hair could be used as a measure of heat tolerance. Yeates (1965) has reviewed the available experimental data and these indicate that a short, light-coloured coat with a smooth and glossy texture is best for minimizing the adverse effects of solar radiation on growth and other productive processes. Posture also has some e&t on minimizing the heat load due to solar radiation, as animals that are standing do not receive as much solar radiation per unit body area as do those that are lying down. There is one effect of length of daylight that influences the ability of some temperate-type cattle to adapt themselves to a tropical climate. It is considered that the length of daylight in the tropics is insufficiently variable to efficiently operate the photoperiodicmechanism that is known partly to control the seasonalhair-growth cycle of temperate--typecattle (Yeates, 1954).Under thesecircumstances some individuals of temperate breeds of cattle are unable to shed their woolly, winter-grown coat and consequently suffer from a heavy heat load when subjected to high ambient temperatures. Individuals of this type never thrive in the tropics and should be culled. 1. Grazing behaviour The effect of climate on cattle is reflected in their grazing behaviour. Data from the experimental work on this subject have been summarized by Payne (1969). The length of daytime grazing of cattle apparently varies according to the degree of climatic stress, the breed and type of cattle utilized and the quantity and quality of the pasture available. When high-grade Bos taurus-type cattle are grazed in a humid tropical climate the length of daylight grazing is radically curtailed and confined almost entirely to early morning and late afternoon periods (Plate 1.l), and the length of the night grazing period fluctuates according to the degree of climatic stress. Even crossbred B. taurus x B. indicus cattle graze for such a short period in the middle of the day that Wilson (1961) reported that yarding such cattle for a 4-hour period at this time, without accessto feed, did not reduce total dry matter intake, and that if the cattle were not yarded they ceasedgrazing and made use of any natural shade available. In many parts of the tropics, indigenous cattle are removed from the grazings and confined during the night. It is usually stated that this is done to protect them from predators, human and animal. However, the majority of observers suggest that whenever possible Bos indicus and

14

The &fleet of Climate

Plate 1.1

Crossbred cattle seeking shade under an oil palm in Brunei.

other indigenous cattle should be allowed to graze at night, particularly when the quantity and quality of the feed available is sub-optimal. If the quantity of feed available is limited total grazing time increases, as it does when the quality is poor, the animal then becoming more selective in grazing. Part of this extra grazing must take place at night if climatic stress is excessiveduring the middle of the day. Joblin (1960), working at Serere in Uganda, where seasonal fluctuations in the quantity and quality of feed are considerable, concluded that the availability of night grazing is critical for B. i~dicus cattle during periods of marginal forage shortage; while under the best conditions the daylight period is sufficiently long for the cattle to obtain their needs, under very dry conditions no amount of grazing time will allow for adequate intake. Restriction of night grazing led, in the Serereenvironment, to a significant decline of 30 per cent in liveweight gain. These experimental findings have since been verified by other workers. No detailed observations have been made of the grazing behaviour of cattle managed under nomadic or semi-nomadic conditions in the semiarid tropics. During the dry season the decreasing water content of the available forage increasesthe animals demand for water at a time when surfacewater resourcesare dwindling, and the animal has to walk further and further to obtain adequate feedand water.Additional walking raises feedand water demand, as increasedmuscular activity requires additional feed and generatesextra heat that has to be dissipated, further depleting the animals water resources. At the same time, the nutrient content of the availa.ble feed decreasesas the dry season advances, the decreasing supplies of free water may become highly mineralized, and ambient

The effect qf climate 011livestock

15

temperatures may rise with a consequent further increase in the animals water requirements. All these factors combine to subject nomadic or partially nomadic livestock in the semi-arid tropics to very considerable physiological stress that may substantially reduce productivity. 2. Intake and utilization (a) Feed intake As would be expectedfrom the effect of temperature on grazing behaviour the available climatic chamber data suggest that high ambient temperatures depress the feed intake of all cattle, but that the feed intake of BOS taurus is depressedat lower ambient temperatures than is that of B. indicus breeds. The effect of very high temperatures is very pronounced, food consumption and rumination practically ceasing in B. taurus-type cattle as ambient temperatures rise above 40C (104F). Increasing humidity at ambient temperatures above 23.9C (75F) also depresses the feed intake of all cattle (Ragsdale et al., 1953) while increasing radiation stress has the same effect on Bos taurus but not on B. indicus-type cattle (Brody et al., 1954). Field data should refIect more accurately than climatic chamber data the overall effect of the climatic environment on the feed intake of livestock. Unfortunately, the available data are limited. Hancock and Payne (1955)and Payne and Hancock (1957)compared the performance of sets of Bos tam-us identical twins fed hay and concentrates, one twin of each set being raised in a tropical and the other in a temperate climate. They showed that hay intake was significantly lower in a tropical climate although overall total digestible nutrient (TDN) intake was approximately the same in both environments. (b) Water intake The direct effect of climate on the water intake of livestock is very complex, as water is required by the animal for at least two different purposes: first as an essential nutrient and component of the body, and secondly to assist the animal lose heat by conductive or evaporative cooling. Although, in general, the water intake of livestock increases with increasing ambient temperature the relationship between water intake and ambient temperature is not simple. For example, in Bos tuurus-type milking cows water intake increaseswith increasing ambient temperature up to 29~4C(85F) but above this temperature it declines. This decline has been attributed to a decline in feed intake and productivity and to a rise in body temperature (Winchester, 1964). Ambient temperature has a differential effect on the water intake of different types of livestock and on different breeds within one type, and it appears that acclimatized animals require less water than unacclimatized when managed at high ambient temperatures. Humidity also affectswater intake, and at ambient temperatures above

16

The Eflect of Climate

23.9C (75F) increasing humidity decreaseswater consumption (Ragsdale et al., 1953)and increasesthe frequency of drinking of cattle. According to Brody et al. (1954) the effect of increased radiation intensity on cattle is to increase water consumption. Presumably this is due to the animal utilizing an increasing quantity of water for evaporative cooling purposes when subject to radiation stress.
(c) l$?ciency of utilization

The experimental evidence available suggeststhat under controlled conditions increasing ambient temperature decreasesthe efficiency of feed utilization, although under field conditions any differences may be insignificant. (d) Loss of nutrients by sweating and drooling Sweating is not of the same importance in all livestock, though it can be important in cattle. In the latter, sweating behaviour differs between breeds as ambient temperature rises, although generally the loss of nutrients and particularly minerals through sweating and drooling is not of practical significance. 3. Growth If climatic stress depressesappetite, reducing feed intake and grazing time, then it is likely to affect productivity as measured by growth and milk production. Although many generalized statements have been made with regard to the adverse effects of a tropical climate on growth there is remarkably little objective field experimental evidenceavailable. In the Fiji-New Zealand twin experiment mentioned in a previous section the growth of the twins was studied from 7: months of age until the end of their first lactation (Hancock and Payne, 1955). The only appreciable differences in growth rates were when air temperatures in Fiji were at their highest: at calving the heifers reared in the temperate climate were only 9.6 per cent heavier than the heifers reared in the tropics, and by the end of the first lactation this difference was substantially reduced. The check in growth rat.ewas reasonably uniform, in so far as all body measurements except belly girth were adversely affected. The belly girth of the twins reared in the tropics was greater and this was attributed to the fact that they had a much greater water intake. Management conditions were good in this experiment, and the results suggestthat an oceanic tropical climate does not appreciably affect the growth rate of temperate-type cattle if management and feeding conditions are good. There is very little experimental evidence on the effect of the different climatic factors on the growth rate of mos iizdicus-type cattle. The birth weights of most B. iudicus calves are low and they often grow rather slowly, but what part if any of this poor growth can be attributed to the direct effect of the climate is unknown.

The eflect of climate on Svestock

17

Information on the effect of climate on the growth of sheep and goats is also scanty. Temperate-type sheep in the Australian tropics exposed to high ambient temperatures often have a low lambing percentage and give birth to small, weak lambs that have a high post-natal mortality. It is also reported from Australia that autumn-born lambs which have been carried through the hot summer are usually smaller at birth than spring-born lambs, but it is very difficult to disentangle the direct from the indirect effects of the climate. The young lamb or kid, like the young calf, is certainly less well adapted to high ambient temperatures than adult animals. The effect of climate on sheep is further discussed in Chapter 7 (Part II). At birth the piglet does not appear to possess a very efficient temperature-regulating mechanism and is incapable of protecting itself against either excessiveheat or cold. During the first 2 days of life the ambient air temperature for piglets should exceed 32.2C (90F) and be gradually lowered as they grow older. It is now normal practice in temperate countries to use infra-red lamps to warm the piglets immediately after birth, so that they do not get chilled. As in the tropics the mean annual temperature varies around 26.7C (80F) and in the daytime temperatures are often of the order of 32.2C (90F) or higher, the problem is not so acute, but it has been found that even in a tropical climate piglet mortality due to overlying may be reduced by the use of an additional heat source during the first few days of life. As the pig ages and grows the optimal air temperature for maximum liveweight gain and efficien.cy of food conversion falls and pigs weighing 3 1.8 to 65.3kg (70 to 144lb) are pro.bably being reared under almost optimal environmental conditions. At higher liveweights normal tropical temperatures are too high for maximum productivity (Heitman and Hughes, 1949). Chicks are more tolerant of high ambient temperatures than are adult birds, but when the air temperature is above 35C (95F) there is a danger of day-old chicks overheating in chick boxes when they are transported from the hatcheries. High ambient temperatures probably reduce the rate of growth of poultry though there are considerable differences in the reactions ofdifferent breeds(Hutchinson, 1954) light breedswithstanding heat better than heavy breeds. 4. Milk production Experimental evidence on the effect of climate on milk, butterfat and solids-not-fat production has been reviewed by Findlay (1954). Most of the available experimental evidence indicates that milk, butterfat and solids-not-fat production are depressed by high ambient temperature, but as in growth studies it is difficult to disentangle the direct and indirect effects of climate. In the Fiji-New Zealand identical twin experiment, where it was possible to assessthe effect of the climatic environment independent of management and feeding (Payne and Hancock, 1957), climate had a marked effect on milk and butterfat but not on solids-not-

18

The Eflect of Climate

fat production. The averagemilk production of the twins in the temperate climate was 44 per cent higher than that of their co-twins in the tropics and their butterfat production was 56 per cent larger. There were, however, profound differences in the reaction of individual twins at the same centre to the climatic environment. Experimental work with dairy cattle in psychrometric chambers has provided more detailed information as to the effect of individual climatic factors on milk and milk solids production. The optimal temperature for milk production in temperate-type cattle breeds appears to be 10C (50F) while the critical temperature after which milk production steeply declines is 21 to 27C (70 to 80F) in Jersey and Holstein, 29 to 32C (85 to 90F) in Brown Swiss and higher in the case of tropical-type cattle. The butterfat content of the milk of temperate cows declines slowly until the ambient temfirature reaches 29C (85F) and then rises.This is presumably due to the fact that above 29C (85F) the decline in milk production is more rapid than the decline in the percentage of butterfat in the milk. High ambient temperatures also affect other constituents of the milk of temperate-type milking cattle. Cobble and Herman (1951) have shown that there is a rise in the chloride content and a fall in the lactose and total nitrogen content of milk when ambient temperatures rise above 27 to 32C (80 to 90F). 5. Reproduction The major climatic factors affecting reproduction are ambient temperature, humidity and the length of daylight. Branton (1970)has reviewed the experimental evidence with regard to the effect of climate on reproduction in cattle and he concluded that high environmental temperatures or sudden violent fluctuations in ambient temperature, such as occur in the sub-tropics, can directly affect the reproductive performance of cattle and that high humidities reinforce the effect of hi.gh temperatures. The known effects of climatic stress on the reproductive behaviour of cattle are depicted in Fig. 1.6. The practical implications of these effects will be discussed in detail in later chapters. Salisbury and Van Demark (1961)have stated that day length appears to be the primary factor affecting the seasonal incidence of fertility, and that where day length varies considerably the highest fertility occurs in the spring with increasing hours of daylight. In the tropics the length of daylight varies very little and this small variation in daylight hours does not appear to have any pronounced effect on the reproductive behaviour of cattle. High environmental temperatures appear to have a marked effect on the reproductive behaviour of sheep (Moule, 1970). In the ewe there is evidence from field observations that both embryonic death and foetal dwarfing occur in a hot environment, and from climate chamber work that continuous exposure to air temperatures that raise rectal tempera-

The eflect of climate on livestock

19

Climatic stress Female cattle Age of puberty Regularity and duration of the oestrus cycle Male cattle Age at puberty

Sexual libido

Incidence of abnormalities of the ova

Interference with the t hermoregulatory function of the scrotum affecting spermatogenesis and semen characteristics

Embryonic mortality

Foetal death rate Gestation length Foetal size

Fig. 1.6 Known effects of climatic stress on the

reproductive behaviour

of cattle.

tures by 1.1 to P7C (2 to 3F) will eventually kill all embryos. Dutt (1960) stated that air temperatures of 32=2C(90F) affect both ova and semenin the female tract of ewes,presumably by raising body temperature. In rams exposed to high air temperatures in the field, degenerative changesin survival characteristics are reported, though there appears to be considerable variation in the testicular temperature of individual Merino rams exposed to high ambient temperatures (Moule, 1970). Male sheep(Moule, 1970)and goats imported into the tropics from the temperate zone appear to be less fecund or even sterile for up to a year after importation and this is presumed to be a photoperiodic effect. High ambient temperatures also appear to affect embryo survival in sows and may have some effect on oestrus (Warnick et al., 1965). Egg production in poultry is highest when air temperatures are within the thermoneutral range (Osbaldiston and Sainsbury, 1963). Constant high ambient temperatures affect the rate of laying of eggs and the total number laid, and there is a diminution in egg weight and shell thickness (Wilson, 1949).Water deprivation enhancesthese inhibiting effects. The fertility and hatchability of eggs is also decreased by high ambient temperatures (Huston and Carmon, 1958). Variation in the intensity of light does not appear to affect egg production significantly.

20

The &fect of Climare

The indirect effect of climate on the animal The major indirect effect of climate on livestock is on the quantity and quality of the feed available for them. Experimental data on this subject have been reviewed by Payne (1969). Other indirect effects are on the incidence of disease and parasites and on the storage and handling of animal products. 1. Feed supply The most important climatic factors that limit plant growth, and hence the quantity of the feed available, are ambient temperature, effective rainfall, length of daylight and the intensity of solar radiation. The quality of feed depends mainly on effective rainfall and on the intensity of solar radiation. The very rl:al differences in climate that exist between the humid and the arid and semi-arid tropical regions thus present two broadly distinct livestock nutritional problems, although there are many exceptions and the distinctions become blurred in the intermediate climatic zones. (a) Equatorial md huinid tropics In general, forage growth is continuous and very rapid though still seasonal, and under high farming conditions very heavy annual yields may be obtained. There are numerous papers detailing yields of humid tropical forage plants and much information has been obtained during the last several decades on their composition and digestibility. Predictions of the productivity of humid tropical forage- in terms of milk production - have been reviewed by Hardison (1966). (i) &fleet on feed and water intake. It is logical to expect that forage grown under conditions of abundant rainfall and high humidity should possessa high water content and most studies show that this is so. Whether this high water content of humid tropical forage inbibits ruminant animals from obtaining a sufficiently high dry matter intake is a controversial question, but the balance of evidence suggests that the high water content of, or the free water on, humid forage may affect the total quantity of feed consumed (Payne, 1969). The experimental evidence (ii) Efltict OIZ tlzenutrimt content oftheforage. is conflicting but, in general, it suggests that forage plants are more nutritious in the wet than in the dry season. Most, but not all, reports conclude that there is a positive correlation between rainfall and the crude protein (CP), silica-free ash and nitrogen-free-extract content of the forage, and an inverse relationship between the rainfall and the crude fibre (CF) content. It is possible that these conflicting reports may be due to an attempt to compare the behaviour of different forage species in what are really different climatic environments, and to the fact that

The qflect of &mute on iivestock

21

although cutting intervals may be the same, the species may be at different stagesof growth at different periods within the same season. Field observations suggest that, after the first flush at the beginning of the rainy season, growth slows down owing to continuous cloudy weather, but that it may increase very rapidly during sunny intervals. It hasgenerally beenassumedthat the nutritive value of humid tropical forage is severely limited by its reported low CP content, but Hardison (1966) concluded that the digestible crude protein (DCP) content of the herbage for which data were available was considerably more variable throughout the year than the TDN content and that the latter was likely to limit milk production for more months of the year than the level of DCP. The CF content of humid tropical forage is not only inversely related to the amount of rainfall but appears to be consistently higher than that of temperate forage at the same stage of growth. This is an additional disadvantage to animals that are already under heat stress and are finding difficulty in eating a sufficient bulk of watery forage. (b) Semi-arid arid urid tropics It is characteristic of semi-arid and arid regions that the total rainfall is low, varying widely in amount from year to year and strongly seasonal and/or erratic in incidence. Thus, it is usual for the rainy season flush of highly nutritious forage to be followed by a long dry period, when growth ceasescompletely and the forage dries up. If a dldught intervenes,the dry seasonmay not be followed by a normal rainy season,but by a few scattered showers and an even longer dry period. Under these circumstances ruminant livestock have to graze for the major part of the year on what is essentially standing hay, and in drought periods they may have to exist on this type of feed for very long periods, at a time when surface-water resources are diminishing. In the truly arid areas forage growth is ephemeral, occurring only after the infrequent rains. Thus, the major problems of the nutrition of livestock in the semi-arid and arid tropics are the intensely seasonalnature of the forage resources and the possibility of low nutrient intake and water deprivation during the dry season. (i) Eflect on feed intake. The dry matter (DM) content of forage in the arid and semi-arid tropics is high throughout most of the year and grazing animals have no difficulty in obtaining an adequate DM intake if ample supplies of forage are available. Thus, the crucial factor in the feeding of livestock in these areas is that of keeping the stocking rate within the carrying capacity of the dry season grazings. (ii) Z$ect of water intake. All domestic livestock require accessto some free water sometimes, but the needs of different types of livestock vary. Cattle require accessto free water at all times, though the water demands

22

The &fect of Climate

of different types of cattle vary and Bos irzdicz~apparently require less free water than B. taurus breeds when managed in the same environment. The water demands of sheep,goats and camels are not as high as those of cattle. When adequate free water is available the water intake of all cattle rises during the dry season but they can be acclimatized to a certain degree of water deprivation. (iii) Water deprivation. This affects water and feed intake, metabolism and productivity. Cattle restrict their DM intake when they are water-deprived, but Bos irzdicus breeds do not reduce their DM intake to the same extent as do B. taurus breeds (Payne, 1965).The higher the ambient temperature the more depressing the effect of the same level of water deprivation on DM intake. If at the sametime cattle are managed in such a way as to restrict their grazing time - being normal managerial practice in many tropical countries (Smith, 1965)-this may decreasefeed intake still further. While the animal is water-deprived, the nutritive content of available feed may decreace,as is normal in the semi-arid tropics with the advancing dry season,and this may lead to a further decreasein voluntary DM intake (Payne, 1965).The cumulative effect of all these factors may mean that the nutritive intake of cattle in this environment is very inadequate As rumen fluid provides the most suitable source of water to offset water lossesduring the initial stagesof dehydration it could be expected that water deprivalion would affect rumen function. It is thought that this happens, and there is experimental evidence that water deprivation in cattle increasesthe digestibility of the feed ingested and in particular the digestibility of the crude fibre component of the feed. Of course, this improvement in digestibility need not be the direct result of water deprivation but could be due to an indirect effect, such as reduced feed intake. Subjective field observations suggestthat some cattle thrive better than could be expected in the very severe nutritional stress environment experienced towards the end of the dry seasonin semi-arid tropical areas. One obvious reason for the better-than-expected performance is an animals undoubted ability to select those parts of the forage plant that are usually of the highest nutritive value. However, towards the end of the dry season,there is often no leaf material left on the grazing or browse speciesand the animal is forced to ingest feed of extremely low nutritive value. It is therefore likely that there are other factors that assist the survival of cattle under extreme nutritional stressconditions. Livingston et al. (1962) investigated the metabolism of Bos taurus and B. indicus cattle under simulated nutritional stress conditions and found that they were able to reduce nitrogen output and particularly urea excretion in their urine to a low level. Payne (1965) has since reported that when the CP intake of B. Taurusand B. indicus cattle was so low that they were in negative nitrogen balance, severe water deprivation improved

The efict of ciimatc 011livestock

23

their nitrogen balance. Roger-son(1963) investigated the effect of declining nutrient intake and water deprivation on energy metabolism in B. tazms and B. irzdicus and concluded that cattle on a low-protein roughage diet might also be better able to maintain themselvesin positive energy balance if they were subjected to some form of water restriction. Water-deprived sheep and camels also appear to behave in a similar manner and it has been suggested that when nitrogen intake is low, nitrogen output is reduced and nitrogen is recycled in the body (SchmidtNielsen et al., 1958; Houpt, 1959),possibly via the salivary glands and/or other channels. Severe water deprivation reduces nitrogen output still further, thus enhancing the value of any recycling mechanism. In many semi-arid areas the mineral content of the only free water available progressively rises as the dry seasonadvances,and may become so concentrated as to be unsuitable for drinking purposes. This is yet another hazard for cattle living in these areas. Tolerance to highly mineralized water can be induced (Mulhearn, 1957), is a function of adaptation, and presumably happens in practice during the dry season in the semi-arid areas as the free water supplies contract. The short-term effect of water deprivation on the liveweight of cattle can be very dramatic (Payne, 1965),but this may be mainly due to loss of body water. The long-term effects are not so well documented, but as water deprivation reducesfeed intake it would be expected that it would also decreaseliveweight gain. The experimental evidence available suggests that this is so. Payne (1965) reported that over a 2-year experimental period intermittently water-deprived Bos irzdicus-type identical twins weighed 14.9 per cent less than their normally watered co-twins, but that the difference in liveweight gain between the two groups occurred during the first 6 months of the experinment.In this experiment the effect of seasonal changes in the quantity and quality of the forage available was far more marked than the effect of water deprivation on liveweight gain. (iv) Eflect on the nutrient content of the forage. Cattle in the arid and semi-arid regions have to exist for long periods on forage that is essentially mature standing hay of low nutrient value. The low protein content, often only averaging between 2 and 4 per cent CP, and the even lower protein digestibility of this forage, is one of the major reasons for the poor performance of cattle in these regions. The reasons for this are that many of the indigenous forage plants are inherently not particularly nutritious, that the forage is usually mature when it is consumed, and that as the forage plants dry out leaf fall increases, leaving feed material with a high stem : leaf ratio. The crude fibre content of the standing hay is high as the forage plants are mature and there is evidencethat environmental conditions in the arid and semi-arid tropics favour the early onset of lignification. Livestock are therefore forced to digest highly lignified fibrous feeds,

24

The Eflect of Climate

and as the digestion of fibre increases the heat output the heat load on the animal is increased at a time when it is already under considerable heat stress. 2. Parasites and disease High ambient temperatures and humidities provide a favourable breeding environment for internal- and external parasites, fungi and disease vectors. Internal parasites are not so important in the semi-arid tropics, but external parasites usually remain very important, though their importance diminishes in the very arid tropics. In so far as the type of vegetation in a region influences the incidence of insect vectors of disease, so climate has quite dramatic indirect effects on animal production. In those regions of tropical Africa where the rainfall is sufficiently high to support a dense growth of bush, and other factors are favourable, the high incidence of tsetsefly (Glossina spp.) makes some forms of livestock production difficult, if not impossible. Similarly, climatic conditions that favour Stomoxys spp. make it impossible in a country such as Mauritius to graze livestock outdoors at certain times of the year and compel livestock owners to build relatively expensive housing to protect their animals from the swarms of flies. In other countries, such as the Sudan, the seasonal incidence of biting flies greatly influences managerial methods. 3. Storage and handling of animal products Any tropical climate, humid or arid, favours the rapid deterioration of stored animal products, thus increasing processing and handling costs. This indirectly affects animal production as increased processing, handling and storage charges, such as the provision of additional refrigerator capacity, may make increased production uneconomic in certain marginal areas that are otherwise suitable for the development of a livestock industry.

Acclimatization
Acclimatization is the name given to the complex of processesby which an animal adapts itself to the environment in which it has to live. If an animal is introduced into a new environment and the stresseson it are too great it will fail to acclimatize and will deteriorate. This often happens when temperate types of domestic livestock have been introduced into a tropical environment. Of course, climatic stress is only one of many stressesthat the temperate type of animal has to withstand in a tropical environment, other major factors being nutritional and disease stress. Acclimatization to heat stress may be temporary or permanent and depends either upon the animal increasing its heat loss, reducing its heat

Acclimatization

25

production, or increasing the tolerance of its tissues to more fluctuating and higher body temperatures. Temperate-type domestic livestock are able to acclimatize more easily to considerable intermittent heat stress than to more moderate continuous heat stress. In the arid sub-tropical areas of the Americas and Australia, temperate-type beef breeds such as the Hereford are exposed to very considerable heat stress during the daytime in the summer months, yet this breed thrives in these areas presumably because heat stress is intermittent in effect - falling off at night during the summer months and non-existent during the day or night during the winter months. The same breeds do not thrive in the humid tropical areas of the Americas and Australia where heat stress during the daytime in the summer months is far lets, hrrt where there is some heat stress all through the year. Permanent acclimatization to climatic stress may be due to changes in the behaviour of the animals or to changes in physiological reactions that may or may not be inherited. Natural or artificial seiection for morphological characteristics that assist the animal to acclimatize may also take place. Changes in the behaviour of domestic livestock are important in assisting acclimatization, and it should be the aim of good management of livestock in the tropics to facilitate these changes. Livestock become more sluggish in their movements in a tropical environment, thus reducing muscular heat production. Well-known examples of this are as follows: bulls are more tractable in the tropics than in the temperate zone, other factors being equal; poultry become more sluggish in their movements and when standing hold their wings slightly separated and if lying down adopt an extended position; temperate-type cattle in the tropics graze more at night and seek shade more often during the day than they do in the temperate zone; and all domestic livestock that are not specifically adapted to arid conditions drink and use more water in the tropics than they do in the temperate zone. Physiological adaptation may be achieved by changes in hormonal activity. There is some evidence that when temperate-type stock are introduced into the tropics there is a reduction in their thyro-adrenal activity, with a consequent reduction in basal energy production and perhaps some effect on reproductive performance. Yousef and Johnson (.1965)stated that part of this depression can be directly attributed to high ambient temperature. Some knowledge of the factors concerned with acclimatization is very important if deliberate attempts are to be made to breed productive animals that are acclimatized to a tropical environment. Animals may be selected for certain morphological characteristics that possibly assist acclimatization, such as a large skin area in relation to body size and short, light-coloured hair and a pigmented skin. When the importation of a breed into a new environment is con-

26

The Effect of Climate

IScorching
I Large zebu cattle f-

I I

Humid

I Dwarf z&u cattle

(VF (60F) --,,, (40F) -

@t:::;~~Rah 2;) I 30 I 40 1 50 II 60 I 70 I x0 I IO

Relative humidity (%)

Fig. I.7

Typical climographs for cattle in Europe and Asia (modified from Wright. 1954).

templated a method of evaluating the possibility of the breed acclimatizing easily has beendescribed by Wright (1954).This consists of constructing climographs made by plotting the mean monthly air temperatures against the mean monthly relative humidities, using climate data collected in both environments. If, when the resulting points are joined, the position, shape and area of the two climographs are similar, then there is a reasonablepossibility that the breed will readily acclimatize to its new climatic environment (Fig. 1.7).

References
Bonsma, J. C. (1949) Breeding cattle for increased adaptability to tropical and subtropical environments. J. agric. Sci. (Cam/x), 39, 204-21. Branton, C. (1970) Fertility, Chapter 7 in Payne, W. J. A. (ed.), Cattle Productiorz irl the Tropics, Vol. 1. Longman : London. Brody, S., Ragsdale, A. C., Thompson, H. J. and Worstell, D. M. (1954) Environmental Physiology and Shelter Engineering with Special Reference to Domestic f.nimals. XXV. The effect of wind on milk production, feed and water consumption and body weight in dairy cattle. Res. Bull. MO. agric. Esp. Stn, No. 545. Cobble, J. W. and Herman, H. A. (1951) The influence of environmental temperatures on the composition of milk of the dairy cow. Res. Bull. MO. agric. E-up. St:z, No. 485. Dutt, R. H. (1960) Temperature and light as factors in reproduction amongst farm animals. Proc. 4th Biennial Symp. Anim. Reprod., J. Dairy Sci.. 43 (Suppl.), 123-44. Findlay, J. D. (1954) The climatic physiology of farm animals. Met. Mortogr.. 2, 19-29. Hancock, J. and Payne, W. (1955) The direct effect of tropical climate on the performance of European-type cattle. I. Growth. Emp. J. esp. Agric., 23, 55-74. Hardison, W. A. (1966) Chemical Composition, Nutrient Content and Potential Milk-

References

27

producing Capacity of Fresh Tropical Herbage. Dairy Train. Res. Inst. Univ. Philipp., Res. BuK, No. I. DTRI: Los Baiios. Philippines. Heitman, H. Jr. and Hughes, E. H. (1949) The effects of air temperature and relative humidity on the physiological wellbeing of swine. J. Anim. Sci.. 8, I7 l-8 1. Holdridge, L. R. (1967) Life Zone Ecology (rev. edn). Tropical Science Centre: San Jo&. Costa Rica. Houpt, T. R. (1959) Utilisation of blood urea in ruminants. Am. J. Physiol., 197, 115.-20. Huston, T. M. and Carmon, J. L. (1958) Influence of high environmental temperature on fertility and hatchability of eggs of domestic fowl. Physiol. Zool.. 31, 232-5. Hutchinson, J. C. D. (1954) Heat Regulation in Birds. in Hammond, J. (ed.). Progress in the Physiology of Farm Animals. Vol. 1. Butterworths: London. Joblin, A. D. H. (1960) The influence of night grazing on the growth rates of Zebu cattle in East Africa. J. Brit. Grassland Sot.. 15, 212-l 5. Kelly, C. F., Bond, T. E. and Ittner, N. R. (1955) Water cooling for livestock in hot climates. Agric. Engrlg (St Joseph, Mich.). 34, 173-80. Kiieppen, W. (1931) Grudriss der Klimakunde. Borntraeger: Berlin. Livingston, H. G., Payne, W. J. A. and Friend, M. T. (1962) Urea excretion in ruminants. Nuture (Land.), 194, 1057-8. Moule, G. R. (1970) Australian research into reproduction in the ram. Anim. Breed. Ahstr.. 38, 185-202. Mulhearn, C. J. (1957) Assessing the suitability of water for livestock. J. Dep. Agric. S. Austrul.. 61. 49-58. Osbaldiston, G. W. and Sainsbury, D. W. B. (1963) Control of the Environment in a Poultry House. I. Principles and practice. II. Broiler house experiments. III. Practical aspects of controlled environment poultry housing. Vpf. Rec.. 75, 159-70, 193-202, 223-9. Payne, W. J. A. and Hancock, J. (1957) The direct effect of tropical climate on the performance of European-type cattle. II. Production. Emp. J. exp. Agric., 25, 321-38. Payne, W. J. A. (1965) Specific problems of semi-arid environments. Qzrulirus PI. Muter. veg.. 12, 268-94. Payne, W. J. A. (1969) Problems of the Nutrition of Ruminants in the Tropics. in Cuthbertson. Sir D. P. (ed.). Nutrition ef AnimuIs of Agricultural Importance. Part 2. Int. Eltcyc. Food Nutrit., Vol. 17. Pergamon: Oxford. Ragsdale, A. C., Thompson, H. J., Worstell, D. M. and Brody, S. (1953) Environmental Physiology and Shelter Engineering with Special Reference to Domestic Animals. XXI. The effect of humidity on milk production and composition, feed and water composition and body weight of cattle. Res. Bull. MO. agric. E-up. Stn. No. 521. Rogerson, A. (1963) Energy utilisation of poor quality roughage by water-deprived steers. Nature (Lond.). 197, 1222. Salisbury, G. W. and van Demark, N. E. (1961) Physiclogy of Reproduction and AI of Cattle. Freeman: San Francisco. Schmidt-Nielsen, B., Osaki, H., Murdaugh, H. V., Jr. and ODell, R. (1958) Renal regulation of urea excretion in sheep. Amer. J. Physiol.. 194, 22 I-8. Smith, C. A. (1965) The grazing habits of African-owned. East African Zebu cattle in Zambia (Northern Rhodesia). J. agric. Sci. (Camh.). 64, 295-8. Thornthwaite, C. W. (1948) An approach towards a rational classification of climate. Geog. Rev., 38, 55-94. USDA (1941) Climate and Man. USDA Yearbook ofAgriculture. United States Department of Agriculture: Washington. Warnick, A. C., Wallace, H. D., Palmer, A. Z., Soza, E., Duerre, D. J. and Caldwell, V. E. (1965) Effect of temperature 0,~ early embryo survival in gilts. J. Anim. Sci.. 24, 89-92. Wilson, P. N. (1961) Observations on the grazing behaviour of crossbred Zebu Holstein cattle managed on Pangola pasture in Trinidad. Turrialhu. 11, 57-71. Wilson, W. 0. (I 949) High environmental temperatures as affecting the reaction of laying hens to iodized casein. Poult. Sci., 28, 581-92.

28

The E$&ectof Climate

Winchester, C. F. (1964) Symposium on growth. Environment and growth. J. Anim. Sci.. 23, 254-64. WorstelI, D. M. and Brody, S. (I 953) Environmental Physiology and Shelter Engineering Reference to Domestic Animals. XX. Comparative physiological reactions with Special of European Indian cattle to changing temperatures. Res. Bull. MO. agric. Esp. and Srn, No. 515. Wright, N. C. (1954) The Ecology of Domesticated Animals. in Hammond. J. (ed.). Progress in the physiology of Furm Atlimnls. Vol. 1. Butterworths: London. Yeates, N. T. M. (1954) Environmental control of coat changes in cattle. Nature (Land.). 174, 609-10. Yeates, N. T. M. (1965) M odern Aspects of Animal Production. Butterworths: London. You&, M. K. and Johnson, H. D. (1965) Feed and temperature effects on thyroid activity in cattle. J. Dairy Sci.. 48. 8 13.

Further Reading
Bligh, J. Temperatlrre Regulatiorl in Mammals ad Other Vertebrates. North-Holland:

Amsterdam. 1973.
Brody, S. Bioenergetics and Growth. Reinhold: New York. 1945. Moloiy, C. M. 0. (ed.) Comparative Physiology of Dew-r Animals. Academic Press:

London. 1972.
Mount, L. E. The Climatic Physiology of the Pig. Edward Arnold: London. 1968. Schmidt-Nielsen, K. S. Desert Animals: Physiological Problems of Heat arzd Water. Oxford

Univ. Press: London. 1964.

Sturkie, P. D. Avim Physiology (2nd edn.). Comstock: Ithaca, New York. 1965.

Chapter 2

Maintenance of Health
by E. P. Lindley
cfo FAO, Rome, Itdy

(including a sub-chapter on Parasitism by R. P. Lee)

Importance of animal health

Profitable animal production demands efficient husbandry of healthy animals; as disease remains a profit-limiting factor in most tropical territories. Even in countries where there is intensive veterinary control it may cause losses of between 15 and 20 per cent of total production. Details of some of the more important diseasesprevalent in the tropics are given in Table 2.1. Somemajor epizootics are now being brought under control. Since the first edition of this book was published in 1959, rinderpest has been eliminated from many countries in Africa; certainly with the efficient vaccinesnow available this diseaseshould present no problem to animalproduction projects. Contagious bovine pleuro-pneumonia has been almost eradicated from Australia, but it remains widespread and prevalent in Africa. There is, however, ample evidence to suggest that with the improved stable and standardized vaccines now available it, too, should not jeopardize cattle-raising schemes.There still remains, however, d long list of diseasesranging from the acute enzootics such as foot and mouth diseaseto chronic parasitic infestations that present serious hazards to profitable livestock production. Of course, low production may aiso be the result of other factors than disease,such as the climate, a low plane of nutrition and poor management. Good husbandry will minimize lossesfrom thesecauses.Additional feeding and continuous medication can also sometimes reduce the effects of disease and parasitism, but these are usually expensive methods of maintaining production.

Relations between livestock owners and authorities

In developing. countries several levels of husbandry often exist side by side, varying from primitive methods based on a subsistenceeconomy to those found on state farms and commercial undertakings that may be sophisticated and efficient. These diverse patterns of husbandry are
29

Tabie 2.1

Basic information concerning some infectious diseases of domestic livestock Iwhhw period Direct contact or through eating material contaminated with excretions Water and food contaminated with blood and excretions or by wound infection Infective ticks 6 to 9 days

predent

in the tropics

Disease

First symptonts

African swine fever

Sudden death, high fever, blotched skin. diarrhoea Sudden death or very high fever

Pigs

Sanitary segregation and destruction of affected animals Annual vaccination

Anthrax

1 to 3 days or ionger

Avian spirochaetosis

2 to 7 days

Intense thirst, profuse diarrhoea Gas gangrene, sudden death

Blackquarter

Contagious bovine abortion Contagious bovine pleuro-pneumonia

Water and food contaminated with blood and excretions or by wound infection Food, water, etc., contaminated by discharge and aborted foetus Direct contact

2 to 5 days

Mostly cattle. Buffaloes, sheep, goats, pigs and horses less frequently Poultry, ducks and turkeys. guineafowl and some wild birds Cattle and sheep

Tick elimination. Vaccination Annual vaccination

7 days to several months 2 weeks to 4 months

Abortions, fullterm stillbirths, retained afterbirths Frequent painful subdued cough. Prolonged unthriftiness

Cattle and buffaloes Cattle

Segregation and other sanitary measures. Vaccination before breeding Annual vaccination, slaughter of affected animals

East Coast fever

Infective ticks

1 to 4 weeks

High fever, unthriftiness, weakness

Foot and mouth disease

Fowl cholera

Fowl plague

Direct contact or with material contaminated with discharge from lesion Direct contact or by eating or inhaling material contaminated with excretions Direct contact or by eating or inhaling material contamined with excretions Direct contact

3 to 8 days

2 to 5 days

Salivation, vesicles on tongue and feet, lameness, fever Sudden death or discharge from eyes and nose, inability to stand Sudden death or discharge from eyes and nose, inability to stand Rough, brown wartlike sores about the head, sore mouth and eyes Diarrhoea, dejection

Cattle. Sometimes buffaloes in East Africa. Cattle indigenous to enzootic areas are immune All domestic animals

Tick control and elimination

Segregation and other sanitary measures. Vaccination Sanitary segregation

3 to 5 days

Fowl pox

3 to I5 days

Poultry, ducks and geese. Occasionally turkeys, guineafowl and some wild birds Poultry, ducks and geese. Occasionally turkeys, guineafowl and some wild birds Poultry, turkeys. Sometimes guineafowl, ducks and geese Poultry, ducks and geese. Occasionally turkeys, guineafowl and some wild birds

Sanitary segregation. Vaccination

Vaccination

Fowl typhoid

Ingestion of contaminated material

4 to 5 days

Sanitary segregation. Vaccination

w Y

Table 2.1 - continued

z Trartsmissiorz Imxhatiorl period 9 to 28 days First symptoms Preventive meusures Sheep, goats and cattle Poultry. Occasionally turkeys, guineafowl and some wild birds All domestic animals Goats Cattle, buffaloes. Sometimes sheep and goats Sheep
_,.

Disease

Heartwater Newcastle disease

Infective ticks Direct contact or by eating or inhaling material contaminated with excretions Infective ticks Direct contact Direct contact or with material contaminated with discharge from lesions Inhalation and through broken skin of material contaminated with discharge Tsetse and other infective flies

3 to

5 days

Piroplasmosis (babesiosis) Pleuro-pneumor i:i of goats Rinderpest

1 to 4 weeks

2 to 3 to

6 days 15 days

Fever, nervous signs, convulsions Sudden death. Stringy discharge from beak, shaking of head and stretching of neck, prostration, diarrhoea Fever, red urine, progressive weakness Fever, nasal discharge, lethargy High fever, bloodstained diarrhoea, mouth lesions High fever, discharge from eyes and nose, skin lesions Intermittent fever, unthriftiness

Tick control and elimination Vaccination

Tick control and elimination Sanitary segregation. Vaccination Vaccination

Sheep pox

2 to

7 days

Vaccination

Trypanosomiasis (surra, nagana, etc.)

Few days to some weeks

All domestic animals

Chemoprophyiaxis and fly eradication

Relations between Zivestock owners and authorities

33

naturally associated with different attitudes to animal health and sickness and are influenced not only by economic but also by social and educational factors. Treatment and nursing of animals is a matter of cost, as well as of experience and training, so that any policy of disease control must be related to the economic level of the animal industry. A good example is bovine contagious abortion or brucellosis, control measuresfor which depend not only on the incidence of the diseaseand the degree of infection in the herd and surrounding local herds but also upon the monetary value of individual animals and social factors. On a government ranch or farm drastic culling or segregation of infected animals can be undertaken ; for most commercial undertakings a vaccination scheme may be suitable; but in herds kept at subsistence level even the necessarycontinued vaccination coverage may not be possible so that the villager has to tolerate the diseasein his herd. On economic grounds it is always advisable to consider the timely and prudent culling of chronically sick animals rather than embark on a long period of cure. Whatever action is taken will depend, among other things, upon the nature of the diseaseand the attendant risk of extension to other stock. The fundamental approach should be the maintenance of health and the avoidance of disease,and owners should do everything in their power, using all available sources of information and expertise, to keep their animals healthy. The stockman cannot expect to be an expert in veterinary sciencenor is it necessary that he should be. However, all governments expect him to be able to recognize the symptoms of certain diseasesthat are notifiable to the authorities. In most countries diseases such as rabies, glanders, rinderpest, fowl plague, anthrax, contagious bovine pleuropneumonia and swine fever are notifiable. The FAO/WHO/OIE Animal Health Yearbook contains details of the distribution of these diseases and the local government veterinary or agricultural officer has ready information and probably a copy of the relevant laws. Government services are available to help owners maintain animal health becauseit is in the national interest to preserve the livestock wealth of the country. State animal health servicesare also organized to combat certain contagious diseases including those which are notifiable. This they achieve by applying regulations to control the internal movement and importation of animals, the cleansing of vehicles, isolation of premises, compulsory vaccination, etc. Usually there is also a government animal health and production advisory or extension service. Stockowners are strongly advised to acquaint themselveswith the local representatives of these bodies and to use their servicesand advice as far as is practicable. For the treatment of individual animals, with which state services do not usually concern themselves, and for advice on general animal health matters, owners may consult a private veterinarian. Although there are as yet few private veterinarians outside the cities in most developing countries, there may be government veterinary clinics. As it is always

34

Maintenance of Health

through the owners initiative that remedial action must be taken when an animal is sick he should not delay in seeking technical help. An animal which is well fed and watered and in good condition will resist disease better than one that is undernourished or suffering from a mineral deficiency, climatic stress or parasitic infestation. Thus, in discussing the determinants of disease the underlying predisposing causes, especially those associated with the environment, cannot be ignored. Good management can do much to remove or reduce the effects of adverse environmental factors.

Diseases
Classification Animal diseasesmay be conveniently classified according to their causative agent as follows: 1. 2. 3. 4. 5. 6. 7. microbiological (bacteria, viruses, protozoa and rickettsia); parasitic (external and internal); metabolic disorders (including nutritional deficiencies); fungal; poisoning (plants, snakes, and chemicals); neoplasms; physical injuries.

Diseases may also be referred to by the particular disability they cause or the part of the body principally affected, without reference to the causative agent, for instance pleuro-pneumonia and enteritis. On the other hand some micro-organisms are so specific in their effect that this may be included in the name, as for example infectious bovine rhinotracheitis and chronic respiratory disease. However, many infectious agents may affect more than one organ or have a generalized effect on the whole animal. A disease is said to be contagious if transmitted by contact between sick and healthy animals, and infectious if spread from the sick to the healthy without direct contact. In the classification above, the infectious and contagious diseasesare included in the first two groups. The bacteria are a large group of unicellular micro-organisms of diverse shapesO-5to 5 pm in size and hence visible under the light microscope. Given favourable conditions they can multiply very rapidly, producing populations of millions within a few hours. Some of them may form resistant spores capable of survival under adverse conditions. Disease is causedwhen certain pathogenic bacteria invade the tissues,multiply and produce toxins. Examples are Brucellu abortus, the agent of contagious bovine abortion, and Clostridium tetani, the cause of tetanus in man and animals. Most of the pathogenic bacteria are susceptible to one or other of the antibiotics.

Diseases

35

Viruses form another large and very important group of diseasedeterminants. Smaller than bacteria and generally comprising a nucleic acid core in a protein shell, they cannot be seen using ordinary techniques with the light microscopeand are designated filter-passing to differentiate them from bacteria. Viruses require living cells in which to multiply. Their morphology and mode of action have been much studied in recent years using electron microscopy. They are not susceptible to most available antibiotics and there is extensive rese.archto find more efficient antiviral agents. Pathogenic viruses, such as those of rinderpest and foot and mouth disease, inv;r host cells, destroying them, and in this way, as de many millioiis crfcells may be involved, they produce cell dysfunction, tissue damage and hence disease. Arbor viruses that cause diseases,for example blue tongue and equine encephalomyelitis, are spread by arthropods such as mosquitoes-a complicating factor in their control. Protozoa are unicellular micro-organisms belonging to the animal kingdom. They causesomeof the most important, widespread and serious diseasesin tropical countries. Some, for example the coccidia, are transmitted directly, but others, like the protozoa1 blood parasites such as trypanosomes, babesia and anaplasms, are spread by arthropod vectors such as ticks and insects,and the control of these vectors presents additional problems. A further complication with protozoa1 infections is that in some casesanimals support a certain level of infection as long as they are under good conditions and well nourished, but as soon as they are stressed,they becomeclinically ill, for example coccidiosis in adult birds and trypanosomiasis in the West African humpless breeds of cattle. There are other families of unicellular micro-organisms which are important pathogens and which cannot strictly be included in the aforementioned groups. Examples are the rickettsias which may cause Q-fever and heartwater and are spread directly by mites and ticks, and also the mycoplasmas which can cause contagious pleuro-pneumonia in goats and cattle as well as contagious agalactia in goats and sheep. External and internal parasitic diseasesare so important to the stockowner and their control is so dependent upon his competence and efficiency that they are discussed later in a separate section. Metabolic diseasesarise from dysfunction within the animals body. Examples in tropical areas are bloat and hypomagnesemia. Milk fever and acetonaemia are not so common where milk production is lessintensive. Nutritional and deficiency diseases,leading to abnormalities, arise from mineral imbalance or a faulty diet, and in particular may occur in rapidly growing animals. Examples are piglet anaemia and curled toe paralysis of poultry. At the end of the dry season in the tropics grazing animals are subject to a low plane of nutrition and/or nutritional imbalance and all species-but especially cattle- are particularly liable to secondary infection from parasitic and protozoa1 disease at this time. Most fungi are saprophytic and beneficial, as for example the yeasts in fermenting processes, but some are pathogenic and cause external

36

Maintenance of Health

mycotic diseasessuch as ringworm or internal mycoses such as aspergillosis in poultry or epizootic lymphangitis in horses. In general, mycotic diseasesare very contagious and less host-specific than most pathogens; though some are found as saprophytes in and on the animals body and only produce diseaseas a result of an increase in the susceptibility of the animal brought about by stress,for example streptothricosis caused by Dematophyllus congolensis. Animals do not normally eat poisonous plauts, but starving cattle or sheep or any grazing animal recently introduced to a new area may do so; and owners should endeavour to remedy underlying mismanagement as it is usually impossible to remove all the offending plants. A stockman who can recognize the poisonous plants of his locality and the symptoms of their poisoning is very valuable indeed, as he is able to make an early differential diagnosis and take necessary remedial action before losses become serious. Although the bites of most poisonous snakes are not usually fatal for cattle they often causelarge oedematousswellings on forelimbs or brisket. If an animal is bitten in the region of the head, the subsequent swelling may interfere with breathing and lead to death. Prompt use of anti-snake venom is justified if the life of the animal is endangered, while pasture improvement naturally reduces the snake population. Most domestic animals are slaughtered at a relatively young age and tumours or neoplasms are seldom a cause of loss of production. If the converse is the case the owner should seek expert advice. If accidents due to staff thoughtlessness or negligence account for regular losses-Monday-morning diseases- the management should make a thorough investigation to ascertain the underlying causesof these mishaps. Transmission Healthy intact skin forms a natural obstacle to invading pathogens. These gain access to the body when cuts, scratches, continued heavy rain, chemicals or malnutrition reduce the effectivenessof the skin as a defensive barrier. Biting flies and ticks also breach normal skin, transmitting blood parasites, streptothricosis, etc. Airborne micro-organisms enter with inspired air. Many are arrested on the mucosal surfaces of the nasal passages,larynx and trachea, but some, depending on the droplet size, may reach the inner membranes of the lung. Even at this stage infection is not inevitable for it depends on the dose, the susceptibility of the animal and the virulence of the organism. The digestive tract is a continuous tube from mouth to anus and its contents are not, technically speaking, inside the tissues of the body. It provides a seriesof very special environments each endowed with its own specialized flora and/or sometimesfauna. In healthy animals these micro-

Diseases

37

organisms are commensals, often necessary,as in the ruminant animal, for proper functioning of the digestive system. Under certain circumstances thesecommensals may themselvesset up disease,but usually the disease-causingorganisms in the digestive tract are other specific pathogenic micro-organisms, perhaps closely related, introduced with the feed and water consumed by the animal. Pathogenscapable of causing generalized disease,such as vibriosis and brucellosis, may gain accessvia the genital tract during copulation. A similar transmission risk exists if artificial insemination is not carried out with strict sterile precautions. Recent studies on nocturnal moths which suck lachrymal secretions in cattle have revealed another mode of diseasetransmission, although so far only infectious bovine kerato-conjunctivitis has been associated with these vectors. For a clear understanding of the objectives and limitation of sanitary control measuressuch as isolation of sick animals, quarantine and disinfection one must appreciate the ways in which diseasesare transmitted. In cases of direct transmission, physical separation of infected from healthy animals will prevent further spread, although most bacteria and viruses can also be transmitted by other means, for instance by soil. manure, sacks, food, boots of workers, water, etc., or by vectors like biting insects or ticks. Quite apart from the innate resistance of stock there is the important factor of infective dose. Very few maladies can be initiated by one single micro-organism. Usually a certain minimal number of microbes, sometimes in repeated doses, is necessary. Thus, in spite of the seemingly alarming possibilities of extension, if practical sanitary precautions are formulated and consistently carried out, then the spread of infection can be arrested. Included in Table 2.1 is the host range of some infections. In recent years it has become more and more evident that many wild animals and birds act as carriers of some of the infective agents which cause disease in domestic stock. In these cases a more rigid control of the environment is necessary to limit transmission, For this reason intensive methods of production based on control of the whole environment have been very successful. Modern pig and poultry units are not only completely isolated but their managers regulate the temperature, humidity and flow of air in the housing, in addition to the food and water supplies. Inflammation, characterized by congestion, pain, swelling and the affected parts feeling hot to the touch, is the body response to injury or irritants -chemical, physical or biological. The physical result of the structural changes produced in the tissues by an inflammation or neoplasm is called a lesion, Examples of gross lesions include abscesses, ulcers, blisters, swollen lymph glands and cornea1opacities. Lesions may not always be visible to the naked eye, even on post-mortem examina-

38

Maintenance of Health

tion, for some diseasesare the result of cellular damage or dysfunction due to biological changewithin cells. The study of the basic biochemical determinants of diseaseis not yet far advanced for it involves very complex chemistry and intricate experimentation. Once organisms have invaded the body they multiply and produce specific toxins, with the rate of multiplication depending upon the virulence of the microbe and effectivenessof the defence mechanisms of the host. The incubation period is the interval between the introduction of the infective agent and the first clinical signsin the host. Although the incubation period varies within certain limits it is specific for each disease (Table 2.1). During the incubation period the organism proliferates in the tissues, but eventually some abnormality becomes apparent such as liquid faeces, lachrymation, discharges, pus, etc., and animals present signs of discomfort such as coughing, straining or merely looking dejected. Immunity The defence mechanisms of the body and their reactions to invading micro-organisms make up one aspect of the study of immunology. Briefly, pathogenic bacteria, viruses, etc., and their toxic products consist of antigens which stimulate the immune system of the host which in turn produces antibodies capable of reacting with, or neutralizing, the antigens. The battle is fought on many fronts, but the specific humoral antibodies of the blood and the white blood cells are very important participants. The speedof their production and their specific effectiveness decide the fate of the animal, which may succumb in hours, may be ailing for only a few days or not at all, or may continue to suffer from chronic diseasefor years. An animals condition affects its responseto infection. If it is weakened by inadequate feeding, bad accommodation, overwork or concurrent diseaseits resistanceis low. On the other hand the robust vitality associated with proper management, good feeding and absence of stress minimizes the effects of the infection, for the bodys natural defences a&quickly brought into action, antibodies are elaborated and immunity acquired. It is this ability of the body to react to the inoculation of foreign antigens which is the basis of the protection offered by vaccines and certain other biological products. These biologicals take three forms : 1. Live vaccines - the infective agent is so attenuated that after inoculation as vaccine it invades the tissuesonly to an extent sufficient for the stimulation of an active immunity - that is, to produce antibodies but not suficient to set up the natural disease. 2. Inactivated vaccines and toxoids- the dead infective agent or its modified toxins are also used to evoke an active immunity.

Diseases

39

3. The specific antibodies themselves either as serum or in a more purified

form- these protective sera are obtained from donor animals, usually horses, which have been hyperimmunized using concentrated doses of the specific antigens made from the disease agents or their toxic products, and which have been altered to make them safe for inoculation. Using the first two products, animals can be vaccinated prophylactically at the convenience of the owner while they are in good health and not exposed to the disease. Following active immunization, 2 to 4 weeks elapse before satisfactory antibody blood levels develop, but such immunity, whether from vaccination or infection, is long-lasting being effective for months, even years and sometimes for life. Vaccination of animals already infected is generally uselessand may lead to exacerbation of the existing malady. Specific immune serum and gammaglobulins, that is the purified antibody fraction of the immune serum, may be injected for passive immunization. Such passiveimmunity is only temporary, lasting days rather than weeks. It is conferred immediately on inoculation and can thus be used to aid the immune response of a sick animal. These products are especially useful in young animals in which the immune system has not fully developed, for example calves with coli-enteritis (calf scours). Although in the pig some immunoglobulin may pass the placenta to the foetus, new-born stock usually receive the maternal antibodies in the colostrum. These antibodies are absorbed for about 36 hours after birth, and the young are thus enabled to survive the first few weeks of life before their own immune system has developed sufficiently to combat infection. It is for this reason that the newly born should be fed the colostrum. Biological products designed to give protection against nearly all the bacterial and viral diseasesare generally available, but vary in effectivenessowing to different inherent properties of the organisms and methods of preparation. Thus, it is possible to employ either a live or an inactivated vaccine depending on the circumstances, as for example against Newcastle disease or brucellosis. Commercial houses compete to produce more effective, better standardized, safer and cheaper vaccines, and the wide choice and differing claims made for the products may present a bewildering complex to the stockowner. For technical as well as economic reasons he should consult a veterinary adviser when planning his programme of prophylactic vaccinations. The instructions issued by the manufacturers regarding the storage and use of their vaccines should be followed carefully if the best results are to be obtained. As yet, satisfactory vaccineshave not been produced far the protozoa1 diseases,although procedures analogous to vaccination - such as the deliberate infection of calves-are successfulwith, for example, babesiosis. Generally, chemical medication has been satisfactory as in the cases of

40

Maintenance of Health

the phenanthridium compounds for the treatment of trypanosomiasis and the nitro drugs for coccidiosis in poultry. Antibiotics have been shown to be useful in treating anaplasmosis. Prophylactic vaccination should be considered as an additional safeguard rather than as an excuse for neglecting ordinary sanitary precautions. Vaccination cover is rarely 100 per cent and young animals may be susceptible so that strict measuresshould always be taken to reduce the chance, the intensity and the spread of infection. These measuresare basedon avoiding contact with infected animals, their discharges, faeces and objects in their immediate neighbourhood which may be contaminated. Indigenous livestock which have been already exposed to certain diseasespossessantibodies, therefore their immune system responds more quickly than that of imported animals with no such disease experience. An infection unimportant to the former may be fatal to the latter on first exposure. Thus newly imported animals are often exposed to considerable risk in their new environment, requiring good management, special vaccinations and treatments to acclimatize. This is particularly true in the humid tropics where even to move stock only a few hundred miles from their place of origin may entail risk of loss from disease. Prevention and control The ideal, for maintenance of health, is to kee!: .:-Lnals in small herds or flocks, segregatedon non-contaminated grouk:d in clean accommodation, watered and fed apart from other stock and kept free of external .and internal parasites. However, such counsel of perfection can seldom be entirely followed. Even though newly acquired animals may look fit and come from reputed disease-free areas, it is a mistake to introduce them without adequate quarantine precautions. Animals returning from markets or agricultural shows which have been exposed to the risk of infection must also be quarantined before being allowed back in the herd. Although quarantining is often inconvenient to management and staff, the temptation to dispense with it must be totally resisted. The duration of time spent in quarantine should be the maximum incubation period of the diseaseinvolved. Routine day-to-day management should be organized to reduce the risk of introducing disease and to control any outbreak which occurs with the least possible inconvenience and expenditure. On account of increasing labour and construction costs livestock units tend to become more intensive and the herds and flocks larger and more crowded, but unlessstandards of hygiene are correspondingly high losses from disease may jeopardize such enterprises. Sanitary-control measuresmust be taken into consideration and incorporated into these 4ecting to do so is to invite trouble at a projects from the beginning; n,, later date.

Diseases

41

Disease is a departure from the normal and time spent in observing healthy stock is never wasted. The first essential is to recognize all aspects of the sound animal- healthy eyes, skin, membranes, excretions, discharges, conformation, etc. Normal animals follow regular patterns of behaviour in their eating, resting, defaecating, etc., and they show definite response to outside stimuli and their sounds, odours and movementsor lack of them - are significant. One of the duties of the stockowner is to be able to detect abnormality in his animals as a first sign of disease and to keep in mind the possibility of its infectious nature. It is of the utmost importance that infectious diseasesshould be detected early and, if in doubt, he should seek the help of his veterinarian. First signs of disease may be obscure and often it is only when there is a radical change in behaviour or a drop in production that suspicions are aroused. Fortunately, most bouts of indisposition are temporary and after a few days the animal is back to normal, but any deviation from the usual state of health must be taken as a warning. A disease which may be catastrophic in its final effects cannot, in the early stages, be differentiated from a minor or temporary malady; therefore it is prudent to take immediate precautions, keeping all sick animals under observation and, wherever possible, isolating them. A consequenceof some infections is that animals show a rise in body temperature. This is usually accompanied by other disturbances such as inappetence and shivering which indicate the presence of fever. As the febrile reaction may occur in the sick animal before other more specific signs are evident it can be used to differentiate some infected animals, which can then be isolated. On account of diurnal fluctuations in body temperatures measurementsof body temperature should be taken at the same time each day, preferably early in the morning. Laboratory examination of material from the sick or dead animal may be necessary in order to make a diagnosis. Samples required by the diagnostic laboratory differ according to the examination requested and the disease.Today specialized transport media and other refined techniques are available to facilitate veterinary investigations in rem,oteareas. It is, however, essential that the correct material reaches the laboratory in a condition fit for examination. This point cannot be overemphasized and for this reason it is better that samples be collected and dispatched by someone trained for the work, but if this is impossible, technical guidance should be obtained from the laboratory concerned. Theactual mechanicsof the application of sanitary control must depend upon local circumstances and the disease suspected. The quicker the control measuresare implemented the more successfulthey are likely to be. Micro-organisms do not sleep from 22.00 to 06.00 hours. The sick animal should be placed in an isolation paddock or stable kept exclusively for this purpose. In-contacts must be segregatedfrom the herd and other herds and kent under close surveillance. If the diseaseis notifiable

42

Maintenance of Health

or if doubt exists, the authorities should be informed. Movement of stock, staff and equipment should be organized to minimize possible spread of infection. Necessary vaccination and treatments of the various groups of stock should be initiated, and as these measures may have to be continued for several weeks, division and segregation of the herd may perhaps require the employment of extra labour for feeding and herding until all risk of spread of infection has passed. Under no circumstances should there be any attempt at concealment. This invariably leads to greater loss and if the disease does eventtyally prove to be notifiable this will make the owner culpable. Disinfection involves the removal or destruction of infection and successdemands a clear understanding of the methods and materials available and their conscientious application. Heat is a most efficient disinfecting agent and burning is an excellent way of destroying infection in bedding, litter, brushes, sacks, etc., although the blowlamp and flame gun, albeit very efficient, are particularly dangerous for use on wooden structures in dry climates becauseof the risk of fire. Some microorganisms are destroyed by currents of hot dry air, but it is better to cleansethoroughly yards, fittings and materials and then expose them to strong sunlight. Small items such as scissorsand clippers can be sterilized by boiling in water for 30 minutes. Many chemical disinfectants are manufactured, but often one has to usewhat is available. In all casesthe manufacturers instructions should be followed and the product used at the correct concentration. It is also essential that the disinfectant be left in contact with the surface to be treated for a sufficient time, the longer the better. To obtain the best results, all traces of dung, litter, etc., should be removed from posts, floors and walls before applying the disinfectant. However, if anthrax is suspected,becauseof the danger to personnel, surfacesshould be treated at once with strong disinfectant and the cleaning carried out later. Certain products are recommended for particular purposes. For example, formaldehyde for fumigation, 10 per cent ammonium hydroxide for coccidiosis, 70 per cent ethyl alcohol for skin disinfection, caustic and washing soda for foot and mouth and other virus diseases,chlorine and iodine for water sterilization, quaternary ammonium compounds becauseof their low toxicity-for wound and skin disinfection, and lysol, creosote or limewash for buildings and other large surfaces. Infected carcasesshould be buried or burned. Burying involves digging a deep pit and covering the carcasewith quicklime under at least 2.0 m (6.6 ft) of earth. Burning can be carried out in a Bostock pit or over a crossed slit trench, each trench being 2.0 m (6.6 ft) long by 05 m (1.6 ft) wide by O-5m (1.6 ft) deep, the soil being piled in the angles formed by the cross and the iron bars lying across the heaps supporting wood, carcaseand more wood. Several bales of straw, about a ton of wood and 2.0 litres (0.4 gal) of kerosene are required to burn completely a 250 to 300 kg (55 1 to 661 lb) carcase.

Parasitism

43

In conclusion, profitable animal production demands good husbandry with a positive endeavour to maintain health. To achieve this the stockman should useall available sources of technical assistance.He should be vigilant for early signs of diseaseand inculcate similar awareness in his staff. Efficient organization of the day-to-day running of the farm with an enlightened approach to preventive medicine and an intelligent application of sanitary-control measureswill do much to prevent diseaseand to limit losses.

Parasitism
It- P. Lee
Department of Clinical Veterinary Sciences, Veterinary College of Ireland, Ballsbridge, Dublin, Ireland

Diseasescausedby metazoan and protozoan parasites constitute a major obstacle to the development of profitable livestock industries in the tropics. In some of the drier pastoral areas that are at present generally more suitable for livestock, generations of cattle and sheep owners learned that constant movement of stock to fresh pastures was attended with better health, provided the risk of exposure to other herds and flocks suffering infectious diseasewas avoided. Animals kept under such systems escaped the dangers of parasitism associated with their remaining indefinitely on pastures which they themselveshad contaminated. Now that better utilization of land, involving set stocking, has become necessaryparasitic diseases haveassumedgreater importance and they will become even more important as carrying capacities improve and stocking rates increase. The tropical environment is for various reasons eminently suitable for the development of parasitic diseases.In the first place, many of the parasites encountered in temperate regions are also found in the tropics, together with the numerous speciesthat are peculiar to the warmer parts of the world. Practically all parasites require to undergo a period of development outside the host before becoming infective for another host, and during this time in the outside world the rate of development and their chances of survival are influenced by climatic and other environmental factors. The rate of development of the extra-host stages tends to rise with increasing temperature, moisture is required for survival and extreme desiccation is usually lethal. In the wetter parts of the tropics extra-host stages are seldom exposed for long to the destructive effects of desiccation and temperatures are commonly optimal throughout the year. Under these conditions the survival rate of parasitic forms outside the host is high; they develop rapidly to the infective stage and large populations are established. Even where prolonged dry seasonsalternate

44

Maintenance of Health

with very short wet seasonsthe extra-host stages of many endoparasites can take full advantage of the warm, wet conditions associated with the latter, while the adult stageswhich subsequently develop within the host are well protected and are safely carried over the ensuing dry season. A second factor contributing to the importance of parasitism is the prevalence of malnutrition in the tropics. Animals with a high nutrient intake would, in many instances, have little difficulty in compensating for the harmful effects of parasitism such as blood loss, leakage of albumen or impaired digestive efficiency. On the other hand the combined effects of evenlight parasitic infestations and malnutrition may harm the animal, and a high incidence of parasitism may seriously reduce the production efficiency and even threaten the survival of individuals during seasonal periods of nutritional stress. Some traditional husbandry systemscommonly practised in the tropics are also conducive to parasitism. For instance, in many countries there is a tendency for the smaller ruminants, pigs and poultry to wander at large round villages where they scavengefor their food, mainly on fouled land. Under these conditions the infective stages of worms, intestinal protozoa and certain arthropods accumulate on the ground and stock are exposed to high infection pressures. Also, during drought periods, all types of livestock, unconfined as they usually are, concentrate in the vicinity of river beds and standing water where green herbage persists. Increased contamination occurs in such places and the moist environment favours the development and survival of infective material. Stock that concentrate in these areas are also exposed to specialized parasites which utilize water-snails and other aquatic invertebrates as intermediate hosts for completion of their life cycles. Furthermore, in many parts of the tropics cattle and sheep are herded or otherwise confined by night on permanent or semi-permanent sites. The surroundings become covered with a mixture of earth, organic matter and infected faecesthat constitutes an ideal medium for the development of the infective stagesof a wide variety of internal parasites. These are readily transferred to the adjacent herbage.

Diseases caused by worms

The common worm parasites are broadly classified into three groups: roundworms or nematodes, flukes or trematodes, and tapeworms or cestodes.Worms are ubiquitous in the tropics and one rarely examines an animal which does not harbour at least a few species.While worms occur in most parts of the body, including notably the respiratory, urinary, circulatory and the central nervous systems and the body cavities, the majority of speciesare found in their adult stages in the gastro-intestinal tract. All types of domestic animals have their own particular fauna of parasitic worms, and while there is a marked tendency to host specifi-

Diseases caused by worms

45

city several economically important species are capable of infecting a variety of domestic animals. Some worms which are essentially parasites of domestic animals are also capable of infecting man; others occur in man in the adult stages but use domestic animals as intermediate hosts. Therefore, worm parasites of domestic animals have public health as well as economic significance. Roundworms or nematodes Roundworms of the gastro-intestinal tract exceed in number and in economic importance those found in other organs. Those that generally causemajor troub!e to livPrtn& in the t.rnnirr: mqv be considered in two .. . ~..v-~*a uF--U ___J broad groups, namely the strongyle worms and the ascarids or large roundworms. The strongyle worms include such cornrnon species as Haemonchus contorlus, a bright red worm with a twisted appearance which is from 10 to 30 mm (0.4 to 1.2 in) long, readily visible and occurs in the abomasum of ruminants; the voracious blood-sucking hookworms, Bunostomum phlebotomum, B. trigonocephalum and Gaigeria pachyscelis, which are found in the small intestine of ruminants and are easily seen by the naked eye; Oesophagostomum spp.,distinctly white worms of about the samesize as H. contortus occurring in the large intestine of ruminants and pigs where they produce nodules in the bowel wall; the somewhat bigger red worms such as Strongylus spp. of the large intestine of members of the Equidae, and various genera of less easily seen slender, threadlike worms, too numerous to name in an introductory work, which occur in the stomach and intestine of ruminants, pigs and poultry. The ascarids include Ascaris suum, Neoascaris vitulorum, Parascaris equorum and Ascaridia spp., of pigs, calves, horses and poultry, respectively. The sexually mature stagesall occur in the small intestine of their host and are easily recognized by their large size, as they measure from about 6 cm (2.4 in) long in poultry to between 15 and 50 cm (5-9 and 19.7in) in the other hosts. The life cycles of all these worms have certain features in common. The sexually mature females,living in the digestive system of their hosts, lay enormous numbers of eggs whiGh reach the exterior in the faeces where they develop further. In the case of the strongyle worms development generally takes place in the faecal mass to the infective stage, which is a free larva capable of considerable activity. This assists it in moving from the faeces to herbage where it is more likely to be ingested. Oxygen, warmth and moisture are necessaryfor its development, but desiccation is soon lethal to both the pre-infective and infective stages so that the faecal mass, which retains moisture longer than the surrounding herbage, tends to serve as a reservoir of infection during short periods of drought. The

d6

Maintenance of Health

eggsof ascarid worms develop in other media such as soil as well as in faeces,but the infective larvae remain within the eggshell which affords them considerable protection. In moist conditions the infective eggs remain viable for months or even years and are capable of resisting quite powerful chemical disinfectants, but they are susceptible to desiccation and drying is probably one of the most effective, practical means of killing these eggs. Ingestion of infective larvae or infective eggs by the host is the most common mechanism of infection, though hookworms and a few other speciesare capable of penetrating the intact skin. In the case of hookworms and Neoascaris vitulorum (the large roundworm of the calf) the foetus or the neonate may become infected with immature, migratory stageswhich pass from the tissue of the dam across the placenta or into the colostrum. This is an example of the important link that exists in the transmission of worm populations from one generation of a host to the next. Another important example is the post-parturient rise in egg output of lactating animals which is briefly discussed in a later paragraph. After entering the host some species of both strongyle and ascarid worms migrate extensively in the body before returning to the gastrointestinal tract where they grow to maturity, copulate and begin to lay eggs; others do not migrate further than the mucous membrane lining the stomach and intestines. Nematode parasites of the digestive tract produce their harmful effects in a variety of ways. Those that migrate extensively, notably Ascaris suum and Parascaris equorum, causetraumatic and inflammatory damage to the liver and lungs. Haemonchus contortus, hookworms and other blood-letting speciesproduce anaemia which in suddenly acquired, heavy infestation may result in a fairly early death of the host. In lighter infestations the haemopoietic reservesof the body may be unable to compensate indefinitely for the more gradual loss of blood, especially where an inadequate intake of nutrients does not allow the blood-forming tissuesto expresstheir full potential, and a progressive anaemia develops. In other instances inflammatory reactions of the gastro-intestinal mucous membranes result in impaired digestive efficiency of the stomach and small intestine and in leakage of albumen into the lumen of the gut. Malnourished animals show clinical evidence of these harmful effects earlier and more noticeably than well-fed individuals. In view of the variety of pathogenic effects it is not surprising that symptoms of parasitism vary considerably, depending on the type of worms predominating. In general, however, most casesshow a progressive loss of condition, with or without diarrhoea, and productive efficiency is impaired. As the diseaseprogresses weight is lost, the body is depleted of fat, and bony structures become prominent, the tissuesmay become oedematous and swellings may then appear on the dependent parts of the body. In ruminants these swellings are particularly apparent under the jaw and along the brisket. Finally, the animal becomes

Diseases caused by worms

47

emaciated and so weak that it is unable to rise. In sub-clinical cases a vague unthriftiness is often all that is observed.The prevalence as well as the severity of worm diseaseis much higher in young than in older stock. In ascarid infestations trouble is more likely to be encountered before weaning, while the post-weaning period is generally a more critical time where strongyle worms are involved. Diseases caused by the nematode parasites result from exposure of susceptible animals to the intake of infective eggsor larvae at an excessive rate, so that harmful effects are produced before the host succeeds in mounting an effective resistanceto its parasites. Control measuresshould therefore be directed towards limiting the number of infective eggs and larvae on the ground to a level likely to result in susceptible animals developing an effective acquired resistancethrough continuous exposure to low infection pressures. At the same time the animals should be adequately fed so that they are given every opportunity to mount an immune response and to tolerate the harmful effects of any worms they happen to acquire. Another, more ambitious, approach is to attempt to maintain herds and flocks completely free of worms. This latter approach is now almost technically feasible in specially favourable circumstances where modem anthelmintics can be used intensively in conjunction with appropriate husbandry techniques; but it is seldom economically feasible. The first approach has more general application and a combination of the following measures may be used where this approach is favoured. Herds and flocks should be maintained on an adequate plane of nutrition throughout the year, special attention being given to breeding stock towards the end of the gestation period and during lactation and to their progeny at and immediateiy after weaning. Where it is feasible to do so, overcrowding of stock, particularly young stock, should be avoided because dense stocking increases the contamination rate and consequently the infection pressure to which animals are exposed. Heavy stocking could be practised with less risk during prolonged periods of drought where extreme desiccation can be relied upon to destroy the pre-infective and infective stages on the ground, but it is unlikely that adequate feed would be available at such times. However, in these circumstancesexcessivelocalized contamination may occur in permanently damp patches such as in the vicinity of standing water. If possible, stock should be denied accessto these places and the surrounds of watering points should be kept as dry as possible. In general, sheep and goats are relatively resistant to the nematodes of cattle. The reverse is also true. Horses are even more resistant to the nematodes of ruminants. Therefore, if a given area is grazed by mixed stock, such as cattle and sheep,the contamination rate will be less than if it were grazed by only one of these speciesand the risk of disease will tend to be reduced. This is essentially an avoidance of overstocking with susceptible stock. Adult animals of a given species are much more resistant to their

48

Maintenance of Health

nematodes than are younger animals, though lactating females may pass greatly increased numbers of eggsand are then a most important source of infection for their progeny. Therefore, non-lactating breeding stock and other adult animals should be used to graze pastures known to be highly infective for young stock. Rotation of pastures was frequently recommended in the past as a method of worm control. Theoretically, animals can be reared free of strongyle worms if they are moved to worm-free ground within the minimum period liable to elapse between passageof eggs in the faeces and the development of infective larvae. Unfortunately, at the high temperatures encountered in the tropics this period may be as short as 3 days so it would be necessary to move stock at short intervals. It is often difficult to apply this principle because vacated pastures would have to be rested for very long periods, especially during the wet season, before larval mortality rendered the grazed areas as safe as they were originally. During these extended periods the new growth of grass must be used. Thus, while pasture rotation may well result in better nutrition by improved management of the sward it is unlikely to result in a decreased intake of infective material. Anthelmintics may be used prophylactically to prevent potentially dangerous numbers of worm eggsreaching the ground, thereby reducing the infection pressure to which susceptible hosts are subsequently exposed. Most modern anthelmintics have a broad spectrum of activity. That is to say, they are highly effective against the adult stagesof a wide variety of nematodes found in the gastro-intestinal tract of domestic animals and they are fairly effectiveagainst the youngest, immature stagesof many -though not all - of theseparasites.Thus, a single treatment may remove practically all the adult nematodes and a high proportion of their immature stages from the gastro-intestinal tract, and some weeks may elapse,even where animals are kept on infective ground, before burdens of adult worms return to pre-treatment levels. These properties of .nodern anthelmintics have improved the effectiveness of prophylactic anthelmintic medication, but a knowledge of the epidemiology of the worm diseasesprevalent in an area is required for full benefits to be derived from their use. Such knowledge indicates when treatments should be given; it shows which categories of stock should be treated at a particular time and it enables appropriate husbandry techniques to be applied in order to supplement the effect of anthelmintic treatments on the worm populations. It is difficult to generalize on such matters becausethe epidemiology of worm disease varies from area to area depending on differences in climate, husbandry practices and other considerations, and systems of prophylactic medication for particular localities, and indeed for individual farms, can be devised only on expert veterinary advice based on local knowlege. Nevertheless, a few broad recommendations can be made.

Diseases caused by worms

49

In areas where the dry season is prolonged and severe, pastures are often free of infection long before the beginning of the rains. In such areas worm infection is carried over from one wet seasonto the next by infected carrier animals, therefore anthelmintic treatment of all potential carriers some weeks before the onset of the rains may delay the build-up of infection on the ground to dangerous levels after the return of wet weather provides suitable conditions for the development and survival of parasites outside the host. Unfortunately, such treatments cannot be relied upon to clear animals of their entire worm burdens hence some contamination of the pasture will occur after the return of wet weather; young susceptible stock will become lightly infected and they in turn will greatly increase the rate of contamination. It is therefore necessary to treat stock at appropriate intervals during the wet season, but it is impossible to make any generalizations as to how often such treatments should be given without a knowledge of the epidemiology of worm diseasesin the area. The beneficial effectsof such anthelmintic treatments are more lasting where treated groups can be moved immediately after medication to worm-freeground or to the cleanestpasture available. A final anthelmintic treatment administered after the end of the rains when pastures are judged to be free of infection enablestreated stock to be carried through the dry season with minimal worm burdens. In areas where the wet seasonis prolonged, prophylactic anthelmintic medication may require so many treatments that the cost becomes prohibitive. Under these conditions very low stocking rates may be the only means of avoiding serious outbreaks of disease. Finally, in considering the prophylactic use of anthelmintics, the special role which the dam plays after parturition in contaminating pasture should be appreciated. The number of worm eggs passed by adult females increasesgreatly at about the time of parturition to reach a peak some weeks later, and there is now evidence to show that this increase in the egg output of lactating animals is brought about by changes in the hormonal rhythm of the host. In many epidemiological situations the rise in worm-egg output occurs at a time when climatic conditions are becoming favourable for the development of the extra-host stages of the worms and, being linked to the post-parturient period, it coincides with the presence of relatively large numbers of highly susceptible young stock. Under these circumstances, anthelmintic treatment of breeding stock shortly before parturition, or immediately after, affords protection to their progeny by eliminating or at least reducing this rise in the output of potentially infective material. The above remarks on prophylactic anthelmintic medication refer principally to the controi ofstrongyle worms. The approach to the control of ascarid worms is rather different in that young animals tend to be the chief source of eggs,although their dams also play some part as carriers. Ascarid eggs are remarkably resistant and the large numbers passed by

50

Maintenance of Health

young stock survive long enough to infect later batches of young animals, such infection being acquired, in the main, within the first few weeks of life. The principle to be followed, then, in using anthelmintics prophylactically for the controT of ascarid worms is to treat young stock when their ascarid worms are about to begin egg-laying. Accordingly, chickens should be treated with a suitable anthelmintic when they are about 6 weeks of age while young pigs and foals should be treated when aged 2 and 3 months respectively. In all cases treatment should be repeated once or twice at monthly intervals. The aim here is to reduce the amount of infective material to which later generations of young stock are exposed,and if this system is adopted it must become a routine management procedure if satisfactory results are to be obtained. Treatment of sows and mares shortly before parturition is advisable because of the role they may play as carriers. For reasons which cannot be discussed in this brief review the epidemiology of Neoascaris vitulorum infestation of calves is rather different, but it may be noted that treatment of calves as a routine at 1 and 2 months of age is generally adequate in herds where this worm is troublesome. Flukes or trematodes In the tropics, fluke infections are found in all speciesof domestic animals, including poultry. The life cycles of flukes which infect domestic animals involve the useof one or two intermediate hosts. One of the intermediate hosts is invariably a snail, but where the life cycle involves the use of two intermediate hosts the first is always a snail in which development of cercariae, representing the penultimate stage in the development of the parasite outside the final host, is completed. In such cases, the cercariae encyst as metacercariae in or on the bodies of a wide variety of second intermediate hosts such as other snails, arthropods, crustaceans, frogs and fish. The metacercaria is the infective stage for the final host. The type of second intermediate host, where utilized, is related to the food chain of the final host. For example, the first imermediate host of Dicrocoelium dendriticum, a trematode parasite of the bile ducts of ruminants, is a land snail while the second intermediate host is an ant which, by the nature of its habits, is likely to be ingested by sheep and cattle grazing on dry terrain. Snails, crustaceans, arthropods and frogs frequently serveas second intermediate hosts for the trematodes of birds. In many parts of the tropics, particularly in Africa, the Indian sub continent, Southeast Asia and many of the Pacific islands, Fasciofa gigantica is economically the most important trematode of domestic animals, affecting cattle and sheep. However, the common liver fluke F. hepatica, which is so widely distributed in temperate and sub-tropical regions, also occurs in some parts of the tropics, where it tends to be confined to cool, higher aititude areas. This latter parasite generally,

Diseases caused bv worms

51

though not invariably, usesa mud snail as an intermediate host so that infection with this species is usually associated with herds and flocks grazing wet, marshy land. On the other hand, F. gigantica generally uses a water-snail as its intermediate host. Therefore, infection with this species is associated with stock drinking from snail-infected watering places as well as with grazing wet land which may be seasonally inundated. The ecology and therefore the control of another group of trematodes known collectively as rumen flukes is similar to that of F. gigantica. The latter trematode may be encountered in a variety of domestic and feral animals, but cattle and sheep are the principal victims. Eggs, laid by adult parasites in the bile ducts, are passed in the faeces and provided they reach water motile organisms hatch in 10 days or more. These organisms seek a suitable snail which they penetrate and after a process of asexual multiplication has been completed within the tissues of the snail numerous cercariae emerge some 6 weeks later. The cercariae are very active and swim about in search of a suitable medium such as herbage on which to encyst as metacercariae and as such are infective for the final host. Metacercariae are fairly resistant although they are unable to withstand prolonged desiccation and on very dry herbage die in a matter of weeks. Sheep and cattle become infected by ingesting contaminated herbage or by drinking water containing metacercariae. The severity of the diseasecaused by this parasite is largely associated with the number of metacercariae ingested. Where massive infections are suddenly acquired the host may succumb to an acute, rapidly fatal hepatitis characterized by haemorrhagic tracts in the liver caused by the migration of numerous immature flukes throughout the tissues of the organ. In lighter infestations a chronic condition develops as the flukes mature in the bile ducts some 16 weeks after infection, giving rise to a progressive loss of blood. Affected animals develop anaemia, become unthrifty, lose condition and in the terminal stages of the disease they are emaciated and watery swellings may appear on dependent parts of the body. Control measures may be directed towards preventing access of animals to snail habitats where metacercariae have encysted, towards reducing snail populations or by maintaining the fluke burdens of the final host at the lowest possible level by regular anthelmintic medication. Removal of stock from snail habitats cannot be relied upon to eliminate infection of the snails becausewild speciesserve as reservoirs of infection. However, if possible stock should be kept away from the margins of slowrunning water and from wet places,pools and reservoirs, especially during the dry seasonwhen recessionof water levels frees previously submerged land for grazing. Theoretically, once such pastures are well dried out they should tend to become safer for grazing stock through the destruction of metacercariae by desiccation. The use of watering troughs sited on dry ground and fed with water from reservoirs or from deep wells should be considered, but the inlets

52

Maintenance of Health

should be screenedto reduce the risk of infected snails being introduced in the water and the troughs should be inspected regularly to ensure that they do not become colonized with snails. Tapeworms or cestodes A wide variety of tapeworms occur in all the domestic animals, although in general they are of little pathogenic importance in their adult stages in these hosts. The tapeworm problems of greatest economic importance in the tropics are Cysticercus hovis and C. cellufosae infection of cattle and pigs, respectively.They are the intermediate stagesof the adult tapeworms Taenia saginata and T. s&urn which parasitize the small intestine of man. Cattle and pigs become infected by ingesting eggs released by mature tapeworm segmentspassedin the faecesof man and they develop eventually into milky-white cysts about 1Qcm (0.4in) long situated in the musculature of these animals. Man acquires the adult tapeworm by eating raw or improperly cooked beef and pork. Eoth theseinfestations are prevalent in the tropics and causeeconomic loss through condemnation or special treatment of affected carcases.In the case of Cysticercus bovis, in particular, its high incidence in certain African countries complicates the development of an export trade in beef. As far as the stock-keeper is concerned, control is largely a matter of preventing humans defecating on pasture, avoiding the use of human manure and sewageon the land and encouraging employees and others living on the property to refrain from eating improperly cooked beef and pork. Young stock are particularly susceptible to infection and as ripe segmentsof Cysticercus bovis are capable of passing through the anus of man of their own volition special care should be taken to see that calf attendants are kept free of infection. Adult tapeworms are more pathogenic to poultry than to other domesticated animals, and birds are susceptible to many specieswhich may be minute and barely visible to the naked eye or up to 25 cm (9.8 in) long. They utilize various intermediate hosts, including ants, beetles, house flies, slugs and earthworms for completion of their life cycles; consequently, tapeworms are more common in birds on free range and are seldom a problem under semi-intensive or intensive systems of management. Under the former conditions regular treatment with anthelmintics is frequently the only practicable control measure.

Diseases caused by arthropod parasites

Arthropod parasitesassumeeconomic significance in two principal ways. Firstly, they impair productivity or cause disease as pests or as direct

Diseases caused by arthropod parasites

53

pathogens. For instance, biting flies such as tabanids may worry stock so much that the animals have to be housed by day and allowed out to graze only after dark, The larvae of the sheep nostril fly, Oestrus ovis, which hatches from eggslaid on the muzzle, causescatarrhal inflammation of the nasal passages sheep and goats. The mange mite, Dmode?c of bovis, and other speciesproduce lesions in skin follicles that render vast quantities of hides and skins uselessfor leather manufacture. Secondly, arthropod parasites assume even greater importance for the part that they play in the transmission of infectious diseases.For example, midges belonging to the genus Culicoides transmit the viral diseasesblue-tongue (Febris catarrhalis ovium) of sheep and African horse sickness (Pestis equorum), as well as acting as intermediate hosts for certain nematodes. Mosquitoes transmit Rift Vallejr fever (Hepatitis enzootica) of sheep and other animals, African horse sicknessand other important viral infections of animals and man. Horse flies, belonging to the family Tabanidae, are restless feeders and may bite a number of animals in a short space of time. Thus, they are efficient mechanical transmitters of a variety of viral, bacterial and protozoan diseases such as equine infectious anaemia (Anaemic icfectiosa equorum), anthrax and trypanosomiases. The bloodsucking stable fly, Stomoxys calcitrans, also transmits diseasemechanically and the common house fly, Musca domestica, transmits a variety of bacterial and several viral infections as a result of its feeding on faeces, and on the food of man and animals. Among the arthropods the tsetse fly and ticks are the most important transmitters of diseasesof domestic animals. The tsetse fly is important because of its role in the transmission of trypanosomiasis, or nagana,of cattle and other animals in Africa. It is estimated that some 44 million square miles of tropical Africa are infested with tsetse flies and vast tracts (Fig. 2.1) are literally closed to successfulanimal husbandry becauseof trypanosomiasis. There are over twenty different speciesof tsetse fly and various groups have particular biological requirements which may be exploited for their eradication or control. For example, riverine species such as Glossina paZpalis and G. tachinoides require the cool, moist environment associated with tree-lined river banks. In certain areas clearing and spraying the heavier vegetation which is often confined to the banks of streams may result in eradication or an effective reduction in fly numbers, provided sufficiently large areas are treated. On the other hand Glossina morsitans, a savanna species,inhabits dry, open parkland through which it ranges widely, and bush-clearing over such large areas would be generally impracticable. This species prefers to feed on big game and in order to survive it must find a suitable host within a few days of emerging from the pupal case. This suggests that the systematic destruction of big gamemight control G. morsitans populations and for some years this action was recommended. This type of control is now considered by many to be unnecessarily wasteful as it has

54

Maintenance of Health

Fig. 2.1

Approximate distribution

of tsetse flies in Africa.

not yet been shown that the fly is incapable of adapting itself to other hosts, such as small game which are practically impossible to eliminate. In the caseof somespeciesthe breeding places are confined to relatively restricted areas,and as the fly populations rise in these locations a point is reached where dispersion over much larger tracts occurs. Also, during very dry periods savannatsetsemay be forced to retreat to thickets from which they range in search of hosts. Control may be attempted under these circumstances by the wholesale destruction or treatment with insecticides of restricted breeding and resting places and of dry season refuges. While eradication of the tsetse fly remains the fundamental approach to elimination of trypanosomiasis it is usually an expensive undertaking and requires highly skilled personnel for its direction. Consequently, much attention has been paid to chemotherapy of trypanosome infec-

Diseases caused by arthropod parasites

55

tions, and for many years a number of drugs have been in common use for treatment of established cases. Reinfection presents a problem, but drugs are also available which, in addition to curing existing infections, give protection for up to 6 months against further infection Such long periods of protection are afforded only under conditions of low fly density; where the challenge is high the period of protection may be drastically reduced. However, the use of thesedrugs enablescattle to pass through and to use infested bush, but unfortunately their use has also contributed to the development of strains of trypanosomes resistant to a wide variety of these drugs and it is now known that some of these strains can be transmitted by the tsetse. Thus, there is a pressing need to develop new chemotherapeutic agents, unrelated chemically and in mode of action to the existing drugs. Ticks occur in all parts of the world, and on a global scale they are of even greater importance than the tsetse fly in the transmission of disease. They transmit many of the more serious viral and protozoan diseasesfound in the tropics and sub-tropics, and their significance in this respect is such that tick control is one of the first requirements for an efficient animal industry in most warm countries. It is thus essential for the livestock owner to be acquainted with the principles underlying tick control. It is so important that it is considered necessary to devote the remainder of this chapter to the subject. The engorgedfemale tick lays her eggson the ground in sheltered places, under stones and in crevices, in a single cluster comprising thousands of eggs. She then dies. The eggs hatch in a matter of weeks or months, depending on the species and prevailing temperatures, and the larvae or seed ticks which emerge climb blades of grass and shrubs to reach a favourable position for attaching themselves to a passing host. Certain species are one-host ticks. That is to say, the ticks spend all their time continuously on the same individual during their larval, nymphal and adult stages,and the females drop to the ground to lay eggsafter they have finished their blood meal. Boophilus decolorutus, the blue tick, which transmits babesiosisor red-water fever and anaplasmosis or gall sicknessof cattle and other diseases domestic animals is a typical of one-host tick. The parasitic stages from larvae to adults require about 3 consecutive weeks on the host. Therefore, spraying or dipping at intervals of 2 to 3 weeks constitutes effective control. With two-host ticks the larvae and nymphs require to spend about 2 weeks or longer on the host. The engorged nymph falls to the ground, mouhs and the adult seeksa fresh host on which it remains for 5 or 6 days or more. Since the immature stagesare frequently spent on small ground game, action is directed towards the adults and dipping at intervals of 7 days or less is required for the control of these ticks. Finally, in the case of three-host ticks each stage drops to the ground after a feeding period which may last for as short a time as 3 days.

56

Maintenance of Health

Plate 2.1 A spray race used in the Ankole District tion. Uganda).

of Uganda (Department

of Informa-

Treatment may be required at intervals of a few days for complete control of three-host ticks. Various measuressuch as grass-burning, cultivation of land and starvation have been recommended for the control of tick populations, but their destruction on the host by the application of chemical substances is still the most practical and effective method. Application is by dipping or spraying (Plate 2.1). The essential features of a good cattle-dipping tank are shown in Fig. 2.2. The tank should be sited on well-drained lanil with adequate space for mustering and holding cattle. A permanent supply of clean water, preferably piped to the tank, should be available. The collecting pen should be easy to clean and the entrance race, at least, should be constructed of concrete to reduce the amount of mud and dung carried into the tank on the feet.Indeed, the construction of a foot-bath at the entrance is advisable since exclusion of organic matter prolongs the effective life of the dip wash, especially where the active principle is one of the synthetic insecticides. This is an important consideration becauseof the high cost of filling the tank. The actual swim-bath, excluding the exit ramp, must be at least 4*6m (15 ft) long so that animals are exposed to the dip wash for an adequate

Diseases caused by arthropod parasites u L

57

u --

r . !,

b. -

58

Maintenance of Health

time. The officially recommended South African dips are 7.3 m (24 ft) long at this point. Similarly, the depth must be sufficient to ensure that the cattle submerge completely following the plunge, and it is usual to have an attendant with a forked stick to push down individuals that have learned how to keep their heads dry. Considerable splashing occurs as the cattle hit the surface and the extension of the sides of the tank above ground level as splash walls effects a great saving in dip wash. Another feature essential for economy is the provision of a concrete draining race to hold dripping animals as they leave the tank. It should be sufficiently long to accommodate animals while the surplus dip drains off without interrupting the steady passageof cattle through the tank. For satisfactory results it is necessary to maintain the dip wash at a constant strength, Precautions must therefore be taken to prevent stormwater from entering the tank from the drainage and entrance races. Similarly, the tank itself should be roofed. Apart from diverting rainwater the roof reducesevaporation which in the tropics can very rapidly raise the concentration of the dip wash. This precaution was formerly very important when arsenical dips were used almost exclusively, because toxic levels of arsenic can easily be reached. Inexperienced stock may require a little persuasion to take the plunge, so once they start to go through they should be kept moving in a steady stream. Not infrequently individuals experience difficulty in the tank and attendants should be prepared with ropes and halters to deal with emergencies. For many years arsenic was the most common active principle of dipping solutions and it had the advantage of being cheap and effective. It is, of course, toxic to animals and to man and its use has been prohibited in the major meat-exporting countries although it is still used in certain African countries. On account of its toxic properties strict precautions must always be taken to see that the wash is maintained at the correct concentration, and when the tank is emptied the old wash must be disposed of carefully to prevent pollution of food and water. A further disadvantage is that strains of ticks resistant to arsenic have now developed after many years of its use. Arsenic has been largely replaced by chlorinated hydrocarbons, such as benzene hexachloride and toxaphene and by organo-phosphorus compounds. They are very effective and can have some residual effect, but this varies depending on the chemical employed as well as the spcies ~and stage of tick against which they are used. All are toxic to man and animals, therefore recommended dipping strengths should no: be exceeded and the precautions issued by manufacturers in connection with dipping and spraying procedures and the general handling of containers and solutions must be heeded. Unfortunately, in some countries, ticks -especially those b&c& I:> the genus Boophilus - have developed resistance to the non-arsenical compounds and it is likely that in time all species of ticks exposed to

Entrance race with at least one foot-bath: preferably two baths should be Drovided (IO ft) long and 20 cm

Entrance frame with-

2x5 16 nozzles, in
8~ 14in nozzles and pressure gauge Rail joining sleeves n

each at least 3.2m

Side wall cut away to show spray pipe system and internal crush race Galvanized metal chute, Priming valve and funnel Draining race fence

6.4 cm (2.5 in) bore suet ion pipe bore aelrvery pipe Perforated metal filter basket standing in return wash channel Suction pit Y Foot valve Pump foundation block Horizontal spray pipes with 4 xl,4 in nozzles each (board not shown)

10 cm (4 in) dia. bolster

Draining race

Connect ion to soakaway for rain- water

Fig. 2.3 The essential features of a good cattle spray race. (Coopers Improved Spray Race. Patentedcourtesy of The Wellcome Foundation Ltd, London and Berkhamsted, England.) Note: Imperial measurements only given for the spray nozzles.

60

Maintenance of Health

regular treatment will become resistant. The stockowner should be.conscious of this possibility, and he should see that formulations are not used below the recommended strength and should seek expert advice at once if he has any reason whatsoever to believe that his dipping or spraying is ineffective. By reason of the high capital cost of dipping tanks - which, if they are to be effective, must be $Jf considerable capacity and are therefore expensive to fill with the newer non-arsenical dips-spray races have tended to replace dipping tanks in many countries in recent years. These consist of an enclosed race in which cattle are exposed t.o a dense spray delivered at high pressure from a system of appropriately arranged jets (Fig. 2.3; Plate 2.1). The discharged fluid drains to a sump from which it is circulated by a pump operated by a small stationary engine or a tractor power take-off. In addition to being cheaper to install than a dipping tank it usesonly a small quantity of wash which can be freshly made up each day. Thus, the risk of using under-strength solutions is avoided and the operator can switch to new formulations without the expenseof having to refill a large-capacity tank. Spraying is quicker than dipping and causesless disturbance to cattle. Other methods of treatment inciude hand spraying and hand dressing. fland spraying can give satisfactory results when carried out by an experienced and conscientious operator, but it is impracticable where there are large numbers of animals. In hand dressing, dip washes or even greasy substances,such as used engine oil, are applied to the predilection sites of ticks and it can be a useful last resort in the control of outbreaks of disease,such as East Coast fever (Theileria purva infection), in backward areas where facilities of dipping or spraying do not exist and, indeed, where supplies of dip materials may not be readily obtainable. Finally, it should be appreciated by the livestock owner that the control of ticks is a complex undertaking, and in planning a programme much thought must be given to such matters as the frequency of dipping or spraying, whether such treatments should be seasonal or continue throughout the year, the best formulation to use and the most suitable rate of application. Decisions depend on such considerations as the speciesand ecology of the ticks in the area and whether the purpose of the programme is merely to reduce tick infestations on the host or to control tick-borne disease. Expert advice, which is generally available locally from government departments, should be sought.

Further Reading

61

Further Reading
British Veterinary Association. Hurldbook C$Animal Diseases in the Tropics (rev. edn). Brit. Vet. Ass.: London, 1970. Dunn, A. M. Veterinary Helminthology. Heinemann : London, 1969. FAO/WHO/OIE. Animal Health Yearbook. FAO: Rome, 1975. Hall, H. T. B. Diseases and Parasites of Livestock in the Tropics. Longman: London. 1977. Parker, W. H. Health and Diseases in Farm Animals. Pergamon: Oxford, 1970. Soulsby, E. J. C. Helmirrths, Arthropods and Protozoa oj Domesticated Animals. Baillikre, Tindall and Cassell: London, 1968.

Chapter 3

Nutrition and Feeding

[with a sub-chapter on Forage]

One of the ultimate objectives of any livestock industry is the conversion into animal products of feedswhich are either inedible by man or surplus to his immediate requirements. In a world where the ever-increasing human population exercises continual and probably mounting pressure on world feed resources it is inevitable that conventional animal feeds should become increasingly more expensive and that the range of basic feeds available for animal feeding should decreaseas more by-products are processed so that they can be directly consumed by man. These handicaps for the animal feeders are reinforced by the fact that the production of animal products inevitably includes a double conversion of basic food constituents. First, soil nutrients are converted into plant products. The latter are then fed to animals for conversion into animal products. One example of this double conversion is that the average efficiency of conversion of fertilizer nitrogen into plant protein is approximately 50 per cent, while the average efficiencies of conversion of plant into animal protein in the temperate zone are approximately 40, 25 to 18, 24, 19 and 11 per cent for milk, eggs,poultry meat, pork and beef, respectively (Table 3.1). For a variety of reasons conversion in the tropics is even less efficient. For example Cuthbertson (1969) has stated that the average efficiency of conversion of plant to animal protein during milk production in the tropics is only 25 per cent. While it may be argued that research during the years ahead will undoubtedly discover new methods of increasing the efiiciency of nitrogen conversion it has to be accepted that two steps in the conversion of nitrogen fertilizer into edible protein can never be as efficient as one. This means that it will always be possible to feed more humans per unit of cultivable land by growing crops for direct human consumption than by growing crops for conversion by domestic livestock into edible animal products. Wilcke (1966), for example, has calculated that 0.4 ha (1.0 acre) of productive land will provide for the protein requirements of one man for only 77 days if the crop is beef, but for 2,224 days if the crop is soybean. What then are the justifications for feeding any livestock? First, most humans demand a mixed diet as man is an omnivorous species and the 62

Nutrition and Feeding Table 3.1 Approximate eficiency of some domestic livestock in converting the energy and protebl qf animal feeds into food for man EfJiciency of conversion (%) ---.Energy Protein

63

Type of animal

Cattle

Milk production during lactation Beef production during fattening Pigs Pork production Poultry Broiler production Egg production

27 11 27 7 13-17

40 11 19 24 18-25

Note: The data have been calculated for temperate-zone countries and not specifically for the tropics. Source: Cuthbertson (1969).

majority are willing to pay higher prices for foods of animal origin than they are for foods of plant origin. Nutritionally, it may be possible for man to exist solely on plant foods, but less total bulk is required of a good mixed diet than of one containing only foods of plant origin. In addition, foods of animal origin are very palatable and usually possessa high protein, fat and mineral content. Secondly, there are many plant foods, particularly forage, that cannot be properly digested by man and that at present can only be processedinto foods suitable for man by the feeding of them to ruminant livestock. Thirdly, in the more ecologically stable agricultural systems plants and animals are complementary and the utilization of both as sources of food may increase total food production per unit area of available land. The latter justification may become more important as the necessity increases for the establishment of ecologically stable agricultural systems. In the past all animal feeds were derived from three sources. These were as follows: crops other than forage, the by-products of crop and animal processing, and natural or planted forage. Today there is a fourth possible source that will probably increasein importance: synthesis from non-biological materials, for example urea from nitrogen or protein concentrate from oil or natural gas. Most feeds for livestock may be conveniently classified into two major types, roughage and concentrates. Roughages are characterized by the relatively large amounts of crude fibre that their dry matter contains. As a group they can be further sub-divided and classified into dry roughages and succulents, the latter containing large quantities of water. A major source of roughage is forage, whose production and utilization are

64

Nutrition and Feeding

discussed in some detail in the second part of this chapter. In general only ruminants such as buffaloes, cattle, sheep and goats, camels and llamoids can properly utilize large quantities of roughage feeds. This is because ruminant livestock differ from non-ruminant livestock in possessingcomplex digestive tracts capable of digesting forage. These differences between ruminant and non-ruminant livestock are fully discussedin the appropriate chapters in Part II. In general, concentrate feeds contain less crude fibre than roughages and relatively large but varying quantities of carbohydrates, crude protein and fat, together with relatively little water. They can be utilized by non-ruminant livestock such as pigs and poultry as well as by ruminants. In addition there are a limited number of feedsthat are difficult to classify into either category. A knowledge of the chemical components of different feeds and of the basic principles of feeding are essential for successfullivestock production. The fundamental principles of nutrition are universally applicable, but all aspects of their utilization for all species of livestock in tropical climates are not as yet fully understood. Under these circumstances discussion will include the components of animal feedsand their evaluation and the feeding standards and requirements of different types of iivesrock utilized in the tropics -as far as these requirements are at present known. One of the major differences is that for some par1 of the year in the temperate zones a part of the energy content of feed has to be expended to maintain the animals body temperature, whereas for the major part of the year in all except the montane tropics, the animals problem is to dispose of surplus body heat. The effect of this basic difference-and other differences that may be induced by climate on the feed requirements of livestock -has not yet been properly evaluated.

Components of food
The food that the animal eats is essentially composed of the same elements that form its bod:y and products. The total amount and the relative proportions of the elements in each food vary greatly, but they are integrated to form compounds or groups of similar substancesupon which a classification of food ingredients can be based. Thus, all foods contain water and their dry matter consists of inorganic material or minerals and organic matter. The latter includes three major groups of substances: nitrogenous compounds, carbohydrates and fats or oils, together with a quantitatively small but qualitatively important group of organic accessory foods known as vitamins. A simple classification of these food constituents gives no indication of their relative importance, either quantitatively or qualitatively. Each must be considered with reference to their function within the animals body and in relation to each other. Therefore, each will be discussed separately.

Components of food

65

Water Water is more vital for the maintenance of the animals life than any other food component. It is the main constituent of all body fluids, being essential for the transport of nutrients to body tissues and the excretion of waste products through the urine and faeces. It is also vital for the proper functioning of most enzymic reactions as these take place in solution and involve hydrolysis. In addition body water plays an all important role in the animals thermoregulatory mechanism. Evaporation of water from lung and skin surfaces helps the animal to dispose of unwanted heat and the high specific heat of the body water assiststhe animal to accommodate itself to large changes in heat production with little change in body temperature. The water content of the animals body varies with age. New-born animals consist of 75 to 80 per cent water, whereas mature, fat animals may contain only 50 per cent or less.The average animal probably consists of between 55 and 65 per cent water. Water requirement increases with growth, with an increase in productive processes such as lactation and egg-laying and with increased physical exercise as when animals are worked and when the animal is subject to heat stress. Requirements apparently vary between different species, between breeds or varieties within species and between individuals within breeds. In fact, water requirement appears to be a very individual and specific characteristic. Average water requirements of some livestock in the semi-arid tropics are shown in Table 3.2.
Table 3.2 Average water requirements of some types qf livestock in the semi-arid tropics during the dry season Type of livestock Average daily water requirement Frequency of drinking

Camel Cattle Sheep Goats Source: Modifications

60-80 30-40 4-5 4-5 of data from Baudelaire (1972).

Every 4 or Every 1 to Every 1 or Preferably

5 days or longer 3 days 2 days once a day

Animals acquire water in three ways. Mainly by drinking free water, but also by utilizing the water that forms part of the food that they consume, and occasionally by manufacturing small quantities of metabolic water obtained by the oxidation of fat within their own body. Some wild animals, particularly those whose habitat is the desert or desert fringe, appear to be able to survive without accessto free water.

66

Nutrition and Feeding

They presumably manageon dew, on the small amount of water available in the dry vegetation that they consume and perhaps to a very limited extent on metabolic water. All domestic animals require ultimate access to free water, some daily and some at less frequent intervals. The camel is the outstanding example of a domestic animal which can withstand infrequent watering intervals. An interesting example on how domestic livestock can thrive on the water available in the vegetation, without accessto free water, occurs in the desert of northwest Sudan. After a specific set of climatic conditions that do not necessarily occur annually, the nomads are able to graze their camelsand sheep for quite long periods across considerable tracts of waterless country. This is because the ephemeral vegetation that occurs, known in the Sudan as the gizzu, contains sufficient water to satisfy the normal needs of the livestock. As stated above, different species of domestic animals differ in their water requirements and these differences are reflected in their respective abilities to withstand dehydration and in their demand for free water. The camel, as stated above, has an exceptional tolerance of heat and water deprivation (seeCh. 9, Pt II), can withstand the loss of up to 27 per cent of its body weight and is able to drink exceptional quantities of water at any one time (Schmidt-Nielsen, 1964). Some breeds of sheep also possess exceptionai tolerance of dehydration (Schmidt-Nielsen, 1964) an and it is likely that many breeds of goats are also very tolerant of partial dehydration. Cattle are not generally as tolerant as camels and sheep,but their requirements for free water vary with type, Bos indicus breeds requiring less water than most B. taurus breeds when managed in the same environment (French, 1956).However, the Ndama, an indigenous B. taurus breed from West Africa, is likely to be as tolerant of dehydration as any B. indicus breed. Payne (1965) reported that acclimatized East African B. indicus-type cattle can lose up to 17 per cent of their body weight between drinking, without apparent harm. Buffaloes and pigs are relatively intolerant of water deprivation. In the humid tropics the grazing animals demand for water may be largely satisfied by the high water content of the forage and by the free water remaining on it after rainfall. Indeed, the animals intake of highwater-content forage may make for difliculties in obtaining a sufficiently high dry matter intake. Nevertheless, daily accessto free water is always desirable. All yarded or housed animals in the wet tropics must have accessto free water, preferably throughout the 24 hours. In the dry tropics the direct and indirect effects of low rainfall are additive in their effect on grazing animals during the dry season. The decreasing water content of the available forage increases the animals demand for water at a time when surface water resourcesare diminishing and the animal has to walk further to obtain both feed and water. Additional walking raisesfood and water demand, as increased muscular activity requires additional feed and generates additional heat that has to be dissipated, further depleting the animals water resources. At the

Components

of food

67

same time ambient day temperatures rise during the dry season, with a consequent further increase in the animals water requirements. In practice this means that often in the arid or semi-arid tropics livestock are watered only every secondor third day. If adequate feed is available this reducesproductivity, but experimental work in East Africa (Payne, 1965) has shown that if inadequate feed is available water deprivation, if not too prolonged, can be advantageous to the animal, as it assists in the conservation of nitrogen and possibly of other food constituents in its body. Water deprivation also conserveswater if it is in short supply. In the East African experiments referred to above, water deprivation for 48 and 72 hours reduced total water intake by 10 and 26 per cent, respectively. Withholding water from sheep also appears to have the same sparing effect on total water consumption. Animals living in the dry tropics are also subject to another hazard. In many regions the mineral content of the only free water available progressively rises as the dry seasonadvances and may become so concentrated as to be unsuitable for drinking. Camels, cattle, sheepand goats can acclim : ize themselvesto such conditions and becomecomparatively tolerant of highly mineralized water. For example, Denton et ~1. (1961) stated that livestock adapt to highly mineralized water and that in certain regions in Australia cattle and sheepare known to thrive on water containing 1.5 to 1.9 per cent total solids, respectively. It would be expected that the consumption of highly mineralized water might have a considerable effect on the mineral metabolism of animals. This appears to be the case. For example, French (1956) reported that the consumption of alkaline water (O-37per cent sodium carbonate) in the dry regions of East Africa increased sodium and chlorine retention but reduced the retention of calcium, potassium, magnesium and phosphorus. When water is freely available, housed or yarded animals usually drink at frequent, short intervals, imbibing a small quantity at any one time. Grazing animals tend to drink in the cool of the morning or in the evening if they have a free choice, but usually they are watered only every 24 hours. Often this single watering takes place during the heat of the middle of the day as the animals are grazed to the watering place, watered and then grazed away from the watering place. The temperature of drinking water is of some importance, cool water being preferable to warm, as drinking cool water assists the thermoregulatory mechanisms of the animal to dissipate body heat (see Ch. 1, Pt I). As the demand of the individual animal for water is normally very variable only average estimates of water requirements in a specific climatic environment can be given (Table 3.2). French (1956) reported on the water consumption of well-managed and fed East African Zebu cattle and grade European-type cattle in Kenya (Table 3.3) and his data

68

Nutrition and Feeding The water consumption of tropical- and temperate-type cattle in Kenya Average water consumption kg(lh) per dart 1: years old Tropical* Temperate+ 18.3 (40.3) 16.2 (35.7) 3; years old Tropical* 19.3 (42.5) 16.9 (37.2) - -Temperate? 27-O (59.5) 23.5 (5 1.8)

Table 3.3

Frequency qf watering

Once daily Every second day

7.8 (17.2) 6.7 (14.8)

* Indigenous cattle. 1-Three-quarter grade Ayrshires. Source: French (1956).

should be compared with those giver! in Table 3.2. Also in East Africa, Payne (1965) found that the range of water intake of growing, indigenous Zebu cattle managed in a semi-arid environment for 2 years was 3 to 43 kg (6 to 94 lb) per day or O-2to 4-O kg (0.4 to 9 lb) per 45 kg (100 lb) of liveweight. These results are comparable with those published by other workers. Nitrogenous compounds The nitrogenous compounds in a feed include the proteins and the nonprotein nitrogenous compounds. The latter include amino acids, amines, amides, nitrates and alkaloids, etc. Nitrogenous substancesin a feed are usually expressed in terms of crude protein (CP) content. Proteins Proteins are complex organic compounds of high molecular weight with many and diverse functions within the animals body. They are found in all living cells where they participate in all phases of cell activity. They also serve as structural elements in all soft body tissues. All known enzymes, many hormones, the oxygen-carrying pigment of the blood, collagen, antibodies and the chemical units of hereditary transmission are all proteins. They consist of chains of amino acids. Most of these are characterized by having a carboxyl group (-COOH) and an amino group ( - NHZ) attached to the samecarbon atom (Fig. 3.1). Also attached to the carbon atom at the position (R) in Fig. 3.1 is the remainder of the amino-acid molecule which will of course vary in composition. Approximately 100 amino acids have been isolated from biological materials, but only 25 of theseare generally considered to be components of proteins. There is, therefore, a great variety of proteins, the characteristics of each depending upon the number and the type of amino acids composing their molecule. Plants and many micro-organisms are able to synthesize proteins from

Components offood
Amino acids are characterized by the possesion of a basic nitrogenous group (-NH,?) and an acidic carboxyl group (-COOH) as shown below.

69

NH2 (R)--&I

I COOH
The nature of the (R) group varies with the amino acid. It may be an hydrogen atom as in glycine;

NH2
I

H-C

COOH

H &cine

or a more complex group as in isoleucine: w \

CH2
\ H ---Cc~/

NH2
I

H imleucine

CH, Fig. 3.1 The amino-acid molecule.

I COOH

simple nitrogenous compounds such as nitrates. Non-ruminant animals must have a dietary supply of amino acids, but they are able to synthesize some that they require from others. Those that animals cannot synthesize are known as essential amino acids. At present it is recognized that growing non-ruminant animals require to ingest ten essential amino acids. These are : arginine histidine isoleucine leucine 1 ysine methionine phenylalanine threonine tryptophan valine

Birds, in addition, require to ingest glycine. Arginine and histidine may not be essential in the adult non-ruminant animal. Ruminant animals possessbiological systems that can synthesize all the amino acids that they require within their digestive tract. The bacteria and protozoa in the tract synthesize amino acids (Muding the essential ones) from dietary protein and non-protein nitrogen. The microbial protein so produced is later digested by the animal at a lower point in the digestive tract with the production of all the amino acids required by the animal for protein synthesis.

70

Nutrition and Feeding

Naturally, those proteins in the ingested food whose amino-acid composition most closely resemble that of the amino-acid composition of the required body protein possessa superior feeding value or a higher protein quality. Food proteins derived from the animals body, such as the protein in blood meal, can be broken down and synthesized into required body proteins with lesswaste than the protein of a grain such as sorghum. Blood-meal protein is therefore said to possessa higher biological value than sorghum grain protein. The biological value of a protein can therefore be used as an assessmentof the relative values of different proteins for non-ruminant animals (Table 3.4). Protein feeds of animal origin usually possessa higher biological value than those of plant origin.
Table 3.4 The biological value of the protein in selected feeds used in pig feeding Feed Biological value of the protein 95-97 74-89 63-76 63 49-6 1

Milk Fish meal Soybean meal Cottonseed meal Maize

Source: Armstrong and Mitchell (1955).

Deficiency of an essential amino acid in the diet of non-ruminants may lead to a failure of growth and eventual death. Different nonruminants have different requirements for the essential amino acids. In birds glycine cannot be synthesized sufficiently rapidly for adequate feathering since the protein keratin, required for feathering, contains a high proportion of glycine. Lysine, methionine, threonine and tryptophan, in that order, are the most likely amino acids to limit growth and reproduction in most non-ruminants. Their deficiency can be made good by adding them individually to the feed. Thus plant protein diets of low biologictil value can be supplemented with individual amino acids and their biological value improved to equal that of protein of animal origin. This supplementation technique is now widely used in the manufacture of feeds for pigs and poultry. While the amino-acid content of the protein in the ration is important for non-ruminants and for the pre-rumination period of young ruminants it is of less importance in older ruminants. What is of major importance is the total nitrogenous content of the ration. Thus the protein requirements of adult ruminants are best expressedin terms of total digestible crude protein. In feeding practice the quality of the proteins fed to ruminants should not be entirely ignored. It is usually economic to feed a variety of protein concentrates as the
Protein requirements.

Components offood

71

pooling of proteins containing different types and quantities of the various amino acids improves the ability of the microbial population in the gut of the ruminant to synthesize rapidly all the amino acids required. In the case of non-ruminant animals requirement can be expressed in terms of digestible crude protein with the stipulation that the nitrogenous content of the ration should contain x per cent lysine, y per cent methionine, etc. The difficulty of this methodology is that the need for any particular amino acid expressedas a percentage of the total nitrogen ingested may change with the rate and stageof growth of the animal and with the breed. Calculation of minimal protein requirements involves: 1. assessment nitrogen retention within the body for growth and other of productive purposes and for maintenance (NR); 2. assessment of endogenous urinary nitrogen and metabolic faecal nitrogen losses (NE). The sum of these, i.e. (NRf NE), is multiplied by 6.25 to provide the minimal protein requirement, i.e. 625 (NR 6 NE). If this figure is divided by the average biological value expressed as a decimal an estimate is obtained of the true digestible crude protein requirement. As almost all living cells are composed of protein and the cells are continuously undergoing degeneration and distintegration a continuous supply of new protein is required by the body even when it is at rest. Thus every animal must receive a continuous though limited supply of protein if it is to maintain its health and not lose weight and condition. In addition any increasein productivity, with the exception of an increase in work, greatly increases the demand for dietary protein. Excess protein in the diet will be deaminated in the digestive tract, the excessnitrogen being excreted in the urine and the faeces.The energy component of the protein can then be used by the animal in the normal metabolic manner. This is of course a wasteful process as protein concentrate feeds are usually more expensive to purchase than energy concentrate feeds.In addition the ingestion of excessivequantities of protein may disturb to some extent the normal working of the digestive system. It is, however, usually desirable to feed rather more protein than the minimal requirements as protein usually improves the palatability of feeds and may thus increase total feed intake. In addition protein is associated in many feeds with desirable minerals and/or vitamins. The specific protein requirements of the different types of livestock are further discussed in the various chapters in Part II concerned with each type of livestock. Non-protein nitrogenous compounds A considerable variety of nitrogenous compounds which cannot be classified as proteins occur in both plants and animals. These include amino acids, nitrogenous lipids, amines, amides, purines, pyrimidines,

72

Nutrition and Feeding

nitrates, alkaloids and some compounds in the vitamin B complex. In this general text only the briefest mention can be made of some of these
Amines.

These are often produced from the amino acids-which comprise the major part of the non-protein nitrogen fraction- by microorganisms. Many thus occur as decomposition end-products and possess toxic properties. A major am le i., urea, the main end-product of nitrogen c metabolism in mammals.
Amides.

Large amounts of nitrates are often found in forage that has been heavily fertilized. Nitrates may be reduced in the rumen to nitrites which possesstoxic properties.
Nitrates. AZkaZoids. These occur in specific plants and are of practical interest

as many of them possesstoxic properties. Well-known examples of toxic or semi-toxic alkaloids in tropical plants are the occurrence of ricinine in Ricinus communis (castor) and mimosine in Lucaena Zeucocephaia. Other plants contain valuable drugs; for example morphine. Carbohydrates The carbohydrates of feedsare generally classified according to the complexity of their structure. There are three major groups. These are simple five- or six-carbon sugars. Ribose, for example, has the general formula CsHlo05, while glucose and fructose have the formula C6Hl 206. They are readily digested and utilized by animals.
1. Monosaccharides. 2. Disaccharides.

These have the general formula Cl 2H22011; examples are sucroseand lactose. They are also normally readily digested and utilized by animals. These are complexes made up from the simple sugars and they usually possessa high molecular weight. They include the following:
3. Polysaccharides.

(a) Starches, the major reserve carbohydrate in plants, and dextrins which are intermediate products in the breakdown or synthesis of starch. Non-ruminant animals can digest and absorb starches. (b) Pectins, cellulose, lignin, hemicelluloses, pentosans and polyuronides. Of these cellulose and lignin are the most important and abundant. forming part of the structure of most plant cells. Non-ruminant livestock, such as pigs and poultry, cannot efficiently digest most of these compounds. Cellulose is, however, broken down by bacteria

Components oj.food

73

and protozoa in the ruminant digestive tract. The products of cellulose breakdown in the ruminant digestive tract are not simple sugars but the volatile fatty acids (VFA), acetic, propionic and butyric, together with methane and some microbiological starch. The VFA are used by the animal for maintenance and for the synthesis of body fat and milk constituents. Acetic acid is used for the synthesis of milk fat, therefore the percentage of fat in milk depends to some extent on the quantity of acetic acid that is produced in the digestive tract and this depends in turn upon the composition of the diet. For example, dairy cows fed on diets low in roughage tend to produce milk with a low butterfat content- Lignin is not appreciably broken down by the1microbia of the digestive tract. Thus the process of lignification, associatedwith the ageing of the plant, may significantly decrease its nutriti,ve value. When feeds are analysed the residue of lignified material, insoluble in dilute acid and dilute alkali, is known as the crude jibre. In a tropical climatic environment the process of lignification in plants appears to commence at an earlier age than it normally would in a temperate climate. Carbohydrates form the largest constituent of plants and are therefore the major part of the food of domestic livestock. However, very little carbohydrate is found in the animals body. The reason for this is that after digestion and absorption carbohydrates are either oxidized directly for the production of energy or they are transferred and stored in the form of fat. Cereals, tubers and roots are the feeds that are richest in sugars and starches, whereas forage, particularly straws, contains less sugar and starch and very large quantities of fibre. Although crude fibre cannot normally be easily digested, even by ruminant livestock, a certain amount is required in the diet of all animals after they have been weaned. Ruminants, of course, require more than non-ruminants. However, even the pig requires some fibre in its diet. In moderate quantities fibre ensures the proper working of the digestive system and gives a feeling of repletion which is one essential of proper feeding. However, as it is the cheapest feed and as it is usually in such plentiful supply the tendency is to always feed too much. When this occurs the digestion of all food constituents is depressed and the sheer bulk involved may depress total intake below nutritional needs. This often happens when cattle graze old, dead forage at the end of the dry season or when pigs are fed only rice bran. There are major differences between ruminant species in their ability to utilize roughages. For example, buffaloes are known to thrive on forage that will not support cattle, while camels, llamoids, tropical breeds of sheep and goats all appear to be able to utilize roughages rather more efficiently than cattle. The reasons for this are not as yet well understood.

74

Nutrition and Feeding

If a sufficiency of carbohydrate food is not available the animal can use excessdietary protein, or if neither is available it can utilize its own fat reserves as a source of energy. Fats When fats are estimated in food not only the true fats or glycerides but also resins, organic acids, essential oils, sterols and plant pigments are extracted and estimated. In general the glycerides possessa higher feeding value than the non-glycerides, but the latter do include essential nutrients such as vitamins A, D, E and K. In non-ruminants absorption of glycerides takes place in the upper section of the small intestine. If the gut is full of dietary fat some may be absorbed in particulate form. Normally, however, triglycerides are transformed into monoglycerides and fatty acids through the agency of the enzyme lipase. Bile salts then mix with the monoglycerides and fatty acids to form completely dispersed micelles. These are absorbed by the mucosal cells of the digestive tract, the bile salts being returned to the lumen of the tract and the monoglycerides and fatty acids being resynthesized into triglycerides. In ruminants lysolecithin may take the place of the bile salts as no monoglycerides reach the small intestine and bile salts do not form micelles with free fatty acids. The resynthesized triglycerides are utilized directly or they are stored in the fat deposits of the body. As triglycerides are a combination of different fatty acids and glycerol, different speciesof animals may possess different and specific types of body fat. Fat is deposited everywhere in the body, either as a protective or as a supporting material. A considerable part of the body fat forms a subcutaneous layer which is usually more conspicuous in temperate-type livestock. Ledger (1959) has shown, for example, that in East Africa Bos taurus and B. indicus cattle exhibit different patterns of fat deposition. At any particular degree of fatness B. indicus cattle deposit more fat intramuscularly than subcutaneously compared with B. taurus cattle. This is presumably because subcutaneous fat deposits form an insulating layer that creates a barrier to heat flow from the deep body tissues to the skin, thus increasing heat stress. It has also been reported from South America that Criollo-type cattle, which have been acclimatized to a tropical climate for several hundred years, deposit their subcutaneous fat in blobs and not as a continuous layer. This also would facilitate heat transfer from the deep body tissues to the outer skin of the cattle. Other types of tropical livestock also deposit surplus fat at specific body sites-camels and Zebu cattle in their humps and fat-tailed and fat-rumped sheep in their tails or rumps. In the past it has been suggested that these fat deposits were useful in times of water shortage as the animal could oxidize the fat in order to obtain metabolic water. However,a simple calculation will demonstrate that the quantity of water thus

Components offood

75

obtained would be very small. It is more likely that the deposition of fat reserves at specific sites, rather than subcutaneously, is a device that assists the animal to rid itself more easily of surplus heat. The deposition of fat usually increaseswith age, and in quick-maturing animals the final deposition takes place between and around the muscle fibres. This process is known as marblirig and has considerable commercial importance as marbling is associated with tenderness in meat. With present methods of husbandry, marbling is not as common in tropical as in temperate breeds of cattle. As stated above, different species of livestock possessdifferent and specific types of fat, and both the quantity and the type of fat are influenced by feeding. The quantity of fat that may be stored by the animal depends upon the total quantity of feed that it consumes rather than on how much fat there is in the food. Herbivora normally eat little fat, unless they are fed on oil-cake, as forage usually contains only quite small quantities. Young animals can consume more fat than older animals and the milk consumed by all young often contains considerable quantities of fat. Some breeds of buffaloes, for example, secrete milk that contains as much as 14 per cent fat. Young pigs can consume milk sub stitutes containing as much as 25 per cent lard. The quality of fat as expressedin terms of hardness can be very easily influenced by feeding. For example, pigs fed large quantities of rice bran exhibit very soft fat, while those fed large quantities of cassavapossessvery hard fat. The fat of different speciesmay differ considerably in colour. Species and breeds of animals that convert the food pigments as soon as they are assimilated normally possess white fat, while those that do not rapidly convert the pigments may accumulate them in their fat deposits. For example, buffalo fat is white while the butterfat and body fat of certain breeds of cattle, such as the Jersey, are yellow. Some fat must be directly consumed by the animal as a deficiency adversely affects carbohydrate metabolism and increasesthe demand for certain I3 complex vitamins. Fats are also the source of the fat-soluble A, D and E vitamins. In omnivorous animals fat also retards the emptying of the stomach and delays the onset of the feeling of hunger and restlessnessassociated with an empty stomach. Minerals The inclusion in the diet of a number of mineral elements that possess important metabolic roles is essential. If one or more of these elements are deficient in the diet, animals wiil ultimately exhibit clinical symptoms of deficiency. These essential mineral elements may be classified into two groups (Table 3.5). Th ere are firstly the macro-elements: calcium, phosphorus, potassium, sodium, chlorine, sulphur and magnesium which are required by animals in relatively la,rgequantities. Secondly, there are the trace or

76

Nutrition and Feeding Essential mineral elements and their approximate concentrations in animal

Table 3.5 bodies

Essential -_______ Macro-elements Calcium Phosphorus Potassium Sodium Chlorine Sulphur Magnesium Source: McDonald,

(%) 1.50 l*OO 0.20 0.16 011 0.15 0.04

~.-~ Micro-elements Iron Zinc Copper Manganese Iodine Cobalt Molybdenum Selenium

hm) 20-80 10-50 l-5 0.2-0.5

Prohahly essential -..__.-___Micro-elements (PP4 Fluorine Bromine Barium Strontium No No No No data data data data

0.3-0.6 0~0241
l-4 No data

Edwards and Greenhalgh (1973).

micro-elements which are required by animals only in very small quantities. At the present time the elements iron, copper, cobalt, iodine, manganese, zinc, molybdenum, selenium and fluorine are included in this group. In addition the elements bromine, barium and strontium may also be essential, but the evidencefor their inclusion is not yet conclusive. The total amount of minerals in the animals body is a very small
Table 3.6 Element Macro-element deficiency symptoms and common sources of the e!ement Type of animal Deficiency symptoms Source of element and/or cure for deficiency

Calcium

Young animals Mature animals Hens

Milking animal Phosphorus Young animals Mature animals All animals

Rickets Milk; green plants; fish Osteomalacia and meat and bone Soft beak and bones; by-products; ground retarded growth; limestone; steamed thin egg shells; bone flour; dicalcium reduced egg phosphate; rock production calcium phosphate (the latter must be free of fluorine) Milk fever Intravenous injection of calcium gluconate Rickets; stunted growth Osteomalacia; low milk yield Pica (depraved appetite); aphosphorosis; stiff joints; muscular weakness; low fertility; low milk yield Milk; cereal grains; fish and meat and bone by-products; dicalcium phosphate; rock calcium phosphate

Components of food Table 3.6 - continued Element Type of animal Deficiency symptoms

77

Source of element and/or cure for deficiency All green plants

Potassium

All animals Calves fed on synthetic milk low in potassium Chicks fed deficient diets All animals Hens

Unlikely to occur in practice Severe paralysis Retarded growth; tetany --Retardation of growth Reduced growth and egg production Decline in appetite; reduction in growth Feather picking; cannibalism

___-I Sodium

-__ - ___ ----- -_ Fish ant meat and bone by-pruJucts; common salt

-_~ _-.-.__-~Chlorine

_-_ All animals Hens -.

___ As for sodium

Sulphur

All animals

Magnesium

Calves; milk fed for XI-70 days Mature cattle and sheep

-____-__-Protein feeds; sodium Limits synthesis of su!phate; elemental the amino acids cysteine, cystine sulphur and methionine -____Wheat bran; cottonseed Tetany; death cake; linseed cake Injection of magnesium Grass staggers or sulphate; prophylactic; hypomagnesaemic magnesium oxide at tetany;* when rate of 5Og per head blood serum Mg per day; magnesium 0.5 mg per IOOml fertilizer on the death may result pastures

*The exact cause is unknown; it may be that dietary magnesium is poorly absorbed.

proportion of the total body weight and the major part are found in the skeletal tissue. Nevertheless,small quantities of minerals are found in all parts of the body. A short account of the metabolic role of each of these essential rninerals is given below and in Tables 3.6 and 3.7 the various mineral deficiency symptoms encountered, some feed sources of the minerals and cures for specific deficiencies are listed. The role of minerals in the nutrition of animals is complicated becauseexcessof some of them may also cause toxicities (Table 3.8) and there are, in addition, a number of rather complex interactions between different essential mineral elements.

78

Nutrition and Feeding Micro-element deficiency symptoms and common sources of the element Type of animal Deficiency symptoms Source of element andfor cure for deficiency Iron dextran injections for pigs Iron is well distributed in green leafy materials -~-~~___~ Plant seeds; copper salts

Table 3.7 Element

Iron

Sue;:ling pigs

Anaemia

Copper

Cattle and sheep

Cattle Lambs

.-_I___Cobalt

__--Cattle and sheep

-Iodine

._--. All animals

_______-____ Anaemia ; poor growth; bone disorders; scouring; depigmentation of hair and wool ; gastro-intestinal disorders Teart, with excess molybdenum and sulphate Lesions in brain and spinal column; muscular incoordination -Cobalt salts; cobalt Emaciation and bullet containing 90% listlessness occurs cobaltic oxide; vitamin when cobalt content of forage BIZ less than 0.08 ppm; pining (vitamin I3i 2 deficiency) _~_~~--___Fis hmeal ; seaweed ; Endemic goitre or iodized salt big neck; reproductive failure Poor growth; leg deformities; poor fertility; frequent abortion Lameness Perosis or slipped tendon Reduced hatchability; reduced shell thickness in eggs; head retraction Only very small quantities of manganese are required Rice and wheat offals; manganese salts

Manganese

Cattle grazing sand and peat soils

Pigs Chicks Hens

--

Components
Table 3.7 - continued

offood

79

Element

Type of animal

Deficiency symptoms

Source of element and/or cure for defciency Widely distributed; yeast; bran and germ of cereal grains Zinc at the rate of 40 to 1OOppm in the diet; as zinc carbonate or suphate

Zinc

Grazing animals Pigs; intensively housed and fed on a dry diet

Unlikely to occur Para-keratosis; sub-normal growth; low efficiency of feed conversion; skin lesions Poor growth; poor feathering and calcification; skin lesions None under practical farming conditions Poor liveweight gain Poor growth Liver necrosis due to vitamin E deficiency Muscular dystrophy due to vitamin E deficiency -i;)ental caries 5s

Chicks

Molybdenum

All animals Lambs on diets low in molybdenum Chicks on purified soybean diets ---___Pigs Calves Lambs .___

Moiybdenum salts

Selenium

----___-Vitamin E or sodium selenite Vitamin E or sodium selenite Fluorides in very small quantities

Fluorine

All animals

Table 3.8

Toxic mineral elements Symptoms Limits of concentration of element or compound

Element or compound

Potassium

High intake of potassium may interfere with the absorption and metabolism of magnesium Too high a level of intake causes excessive thirst, muscular weakness and oedema; salt poisoning quite common in pigs and poultry, particularly when water is limited When water is limited; 4% hens, 2% chicks and 1% turkey poults

Sodium chloride

80

Nutrition and Feeding

Table 3.8 - continued Element or compound Symptoms Limits of concentration of element or compound

Iron ___Copper

Excessive intake causes digestive disturbances ----. ___-Continuous ingestion of excess copper leads to an accumulation in the body tissues; sheep are particularly susceptible; copper poisoning occurs naturally in parts of Australia Unlikely to occur under practical farming conditions ~-. Toxic in very large doses; toxicity unlikely to occur under normal farming conditions ----.Depression of feed intake and induction of a copper deficiency; unlikely to occur under normal farming conditions Induces copper deficiency under certain conditions

---

-~

.- -..-- -. --- -.

Cobalt

40-50mg daily of Co per 45 kg ( 100 1b) body weight - _-.- --_ - - -... .- ._- -

Manganese

Zinc

Molybdenum Selenium

Causes toxicity in horses, cattle 1O-30 ppm and sheep; known as alkali disease or blind staggers in the United States; dullness; stiffness of joints; lameness; loss of hair; some plants accumulate selenium; Astragalus hisulcatus may contain up to 4,OOOppm on a dry-matter basis; the toxic effect is reduced when high-protein feeds are given to the livestock > 20ppm in diet Causes fluorosis; teeth become pitted and worn; intake affected; fluorine is an accumulative poison; the major sources are water, rock phosphates used as a fertilizer, and industrial plant in some regions

Fluorine

As it is impossible to consider this subject in depth in an introduction and as the subject is a dynamic one with new information constantly being made available, the reader who is particularly interested should consult specialized texts and the current literature.

Components of food

Macro-elements This is the most common mineral element in the animal body. It is a constituent of bone and teeth and most living cells and tissue fluids. It also performs a role in the coagulation of the blood, the normal action of skeletal and heart muscle and in the regulation of the excitability of the nervous system. The deficiency symptoms described in Table 3.6 may also be caused by a deficiency of phosphorus and/or vitamin D or by an abnormal calcium : phosphorus ratio. Normal ratios are 1 : 1 to 2 : 1 in mammals and are rather wider in birds. Calcium metabolism is under the control of a hormone secreted by the parathyroid gland.
Calcium,

It is a constituent of bone, phosphoproteins, nucleic acids and phospholipids and has a role in calcium metabolism. Phosphorus deficiency (Table 3.6) is usually more common in cattle than in sheep. Pica (Table 3.6) is not a conclusive symptom of phosphorus deficiency as it may be caused by other factors. Although cereal grains are a good source of phosphorus, much of what they contain may not be available for non-ruminants if it occurs in the form of phytates. These are more digestible by ruminant than by nonruminant livestock.
Phosphorus.

This element has a role, together with sodium, chlorine and bicarbonate ions, in the osmotic regulation of the body fluids, in which it functions primarily as the cation of cells. It also plays a part in nerve and muscle excitability and in carbohydrate metabolism.
Potassium. Sodium.

Like potassium this element is concerned with the osmotic regulation of body fluids, being the main cation of blood plasma and other extra-cellular fluids.
Chlorine.

The element is associated with potassium and sodium in osmotic regulation and has an important role in gastric secretion in the true stomach of animals. Proteins containing the amino acids cystine, cysteine and methionine, the vitamins biotin and thiamine and the hormone insulin all contain sulphur. Wool contains up to 4 per cent sulphur. Small quantities of sulphates also occur in the blood. Deficiency of sulphur normally denotes a protein deficiency. When a non-protein nitrogenous compound such as urea is fed to ruminants additional sulphur may assist microbial synthesis of sulphur-containing proteins.
Sulphur. Magnesium.

This element is closely associated with calcium and phosphorus in the skeletal structure, it is an activator of phosphates and it

82

Nutrition and Feeding

is also concerned :vitlr calcium metabolism. As adult animals possessonly a very small available reserveof magnesium in their bodies they are very dependent upon a regular supply of this mineral. Micro-elements Iron. Most iron in the body occurs in the haemoglobin of the red blood cells, but it is also found in a blood serum protein known as siderophilin. This protein is believed to have a role in the transport of blood from one part of the body to the other. Iron is also found in the protein ferritin which is present in the spleen, liver, kidney and bone marrow. It is also a component of many enzymes and some flavoproteins. The daily requirement of iron by the normal healthy animal is small. Anaemia due to iron deficiency could occur after prolonged haemorrhage, but normally only occurs in baby piglets. This is because sows milk is particularly deficient in iron. The absorption of iron appears to be to some extent independent of the dietary source. Copper. This element is essential for the production of red blood corpuscles and for maintaining their activity. It is a component of many enzyme systems and is necessary for the normal pigmentation of hair, fur and wool. Storage takes place mainly in the liver, but copper is probably present in all body cells. As will be seen from Table 3.6 there are a variety of deficiency symptoms. In Australia a copper deficiency in lambs is known as enzootic ataxia and is associated with the grazing of pasture with a low copper content (2 to 4 ppm Cu in the dry matter). Similar clinical symptoms occur in lambs in the United Kingdom suffering from a condition known as swayback although the copper content of the grazings is normal (7 to 15 ppm Cu in the dry matter). A clinical condition known as teart, characterized by unthriftiness and scouring, occurs in cattle in the United Kingdom. A similar condition is known as peat scours in New Zealand. The feeding of copper sulphate to animals controls this condition although the copper content of the pasture may be normal. However, thesepastures do possess high molyba denum content (20 to 100 ppm compared with O-5to 3.0 ppm in the dry matter of normal pastures).It is believed that molybdenum affects copper retention by the animal, but limits it only in the presence of sulphate. This element is required by microbia in the rumen for the synthesis of vitamin B 12 and it is an activating ion in certain enzymic reactions. Sheep are more liable to exhibit cobalt deficiency than other types of livestock.
Cobalt.

This is a constituent of the hormone thyroxine that controls the metabolic rate of animals. Although a deficiency of iodine in the diet causes goitre, it is not the sole cause. Some feeds contain goitrogenic compounds, particularly Brassica spp., soybeans, peas, groundnuts and
Iodine.

Components

of food

83

linseed. These goitrogenic compounds appear to block the absorption of iodine in what would otherwise be an adequate diet.
Manganese.

This is important as an activator of enzymic reactions concerned with carbohydrate, fat and protein metabolism. It is found in traces in most tissues. The highest concentrations are in the bones, liver, kidney, pancreas and pituitary gland. Manganese deficiency is not the only factor concerned in the clinical condit& known as perosis in chicks (Table 3.7).The condition is aggravated by high intakes of calcium and phosphorus.
Zinc.

This element appears to be present in all tissues and is concerned in some enzymic reactions. The clinical condition in pigs known as parakeratosis (Table 3.7) is aggravated by increased calcium levels in the diet and decreased by reduced calcium and improved phosphorus levels.
Molybdenum.

This is a constituent of the enzyme xanthine oxidase that has an important role in purine metabolism, of nitrate reductase and of a bacterial hydrogenase. The exact relationship between selenium and vitamin E (Table 3.7) has not yet beendetermined. Selenium can replace sulphur in the amino acids methionine and cystine found in seleniferous plants.
Selenium.

This is distributed throughout the body but is concentrated in the bones and teeth. The role of this element in the metabolism is not yet fully understood.
Fluorine. Bromine.

.This element may possesssome role in the growth of chicks.

Conclusive evidence that these are essential elements is not yet available.
Barium and strontium.

In tropical environments mineral deficiencies may be enhanced by the animal losing considerable quantities of the elements during sweating or by dribbling saliva. For example, Bonsma (1940) stated that unacclimatized bulls may dribble 13 to 18 litres (3 to 4 gal) of saliva a day and lose 51 to 71 g (2 to 25 oz) of minerals. However, it is not considered that this is a problem in acclimatized livestock. Cereals, and to a lesserextent cereal by-products, normally possessa low proportion of calcium and a relatively high proportion of phosphorus. Most forages possessmore calcium than phosphorus. Legume forages normally possessa relatively high calcium content and browse is usually relatively rich in all mineral elements. Feeds of animal origin are usually well supplied with minerals. Our knowledge of mineral deficiencies in tropical regions is very

84

Nutrition and Feeding

limited at the present time. In Guyana very large areas are known to be deficient in minerals as are extensive areas in Brazil and Central America. There is at present a.major project, with headquarters at the University of Florida, that is concerned with ascertaining the extent of mineral deficiencies in tropical America. In Africa it is known that very large areas of range, particularly in East, Central and South Africa, are deficient in phosphorus and that this deficiency reduces the fertility of cattle in these regions. In Kenya and Zaire cobalt-deficient areas have been identified (Suter, 1962).It is likely that the soils in almost all humid tropical regions are deficient to a greater or lesserdegree in one or more mineral elements.Obviously a very great effort will be required to map mineral-deficient areas in the tropics. In the interim period the livestock owner can only insure himself against mineral deficiencies by the feeding of mineral supplements. Nomadic peoplesin the tropics have known for a very long time that it was necessaryfor them to insure against mineral deficiencies, although their livestock range over wide areas thus reducing the risk of mineral deficiency in any one locality. For example, the Baggara people in the western Sudan know which of their ranges are salty and they herd their animals so that they spend as long a time as is possible on the salty range. Nomadic herders also usually carry salt with them for their livestock and/or visit areas where there are edible earths. Certainly eartheating or geoplzagia is a very common habit of both domestic and wild animals in the tropics. It is interesting that chemical analyses of edible earths from both East and West Africa, although demonstrating the variety of their mineral constituents, have failed to establish direct links between known mineral deficiencies and geophagla (French, 1945). In the humid forest regions of South and Southeast Asia salt has been used to attract wild cattle of the Bos (Bibos)-type to breed with domestic cattle and it has been suggested that the domestication of B. (Bibos)type cattle may have been originally achieved by man attracting the wild cattle by the provision of salt or highly mineralized earths (Simoons and Simoons, 1968). We suggest that in the tropics all grazing livestock should be fed a simple mineral supplement and that all yarded and/or housed livestock should be fed complete mineral supplements. A very suitable and simple mineral supplement for ruminant livestock is as follows: Common salt Ground chalk, limestone or shell Steam bone flour 40 parts 40 parts 20 parts

This mixture should be fed ad lib. to grazing animals. If concentrates are fed the mixture may be added to them at the rate of 2 to 3 per cent of the total ration. If there is a known mineral deficiency in the regionapart from calcium, phosphate, sodium and chlorine- then a salt of the

Components of food

85

specific mineral that is known to be deficient should be added to the above mixture at the recomme:lded rate. We do not advocate the indiscriminate feeding of complete mineral mixtures to grazing animals unless the latter are exhibiting some symptoms of minerai deficiency. However, it should be realized that in this context the deficiency symptoms may be sub-clinical in their manifestation, i.e. poor growth and low fertility. Range livestock may have to be supplemented in the dry season with other nutrients, and where urea-molasses-mineral blocks or liquid feeds are used the animals will be able to obtain their mineral requirements from the block or liquid feed. Yarded or housed livestock must of course be considered to be in a different category. They should all receive rations containing complete mineral supplements. In most tropical countries the livestock-feeding industry has now developed to such an extent that compl;te mineral mixtures are readily available. If this is not the case then the livestock owner requiring information on the composition of complete mineral mixtures should consult his local extension adviser. Vitamins Vitamins are essential food substances that are required in very small quantities by animals. Those known to be of any importance in animal nutrition are listed in Table 3.9. For convenience they are classified in
Table 3.9 Vitamins of importance i,t animal rtutritiorr

Fat soluble A retinol: precursor: carotenoids, the most important being /?-carotene D there are ten to twelve different forms: only ergocalciferol (Dz) and choiecalciferol (DJ) are of importance E collective name for a group of closely related compounds known as tocopherols: a-tocopherol is the most important K collective name for a number of compounds: phyloquinone (Kr) is the most important Water soluble B complex B r thiamine Bz riboflavine nicotinamide: formerly known as nicotinic acid or niacin Bg pyridoxine: exists in three forms that are interconvertible pantothenic acid biotin choline folic acid Bra cobaltamin: several forms of the vitamin are known C l-ascorbic acid

86

Nutrition and Feeding

two groups : fat soluble and water soluble. Details of the clinical symptoms that deficiencies of these vitamins may cause, together with some of their common sources, are listed in Table 3.10. Brief accounts of the roles that these essential foodstuffs occupy in the metabolism of animals are summarized below.
Table 3.10 Vitamin Vitamin deficiency symptoms and common sources of the vitamin Type of animal DeJiciency symptoms Source of vitamin and/or cure for deficiency Green, leafy materials; fish-liver oils; synthetic vitamin A

Cattle

Pigs Poultry

All animals D Young animals Older animals

Rough coat and scaly skin; excessive watering of eyes culminating in xerophthalmia; low fertility; abortion Poor growth; low fertility Retarded growth; staggering gait; low egg production and hatchability Secondary bacterial infections Rickets Osteomalacia

Synthesis in the skin of outdoor-managed animals; sun-dried feeds such as hay; fish-liver oils Green leafy materials; cereal grains; synthetic a-tocopherol; selenium salts Green leafy materials; fish meal Cereal grains; generally wide distribution in feeds

Young cattle and lambs All animals Chicks Chicks Chicks

Muscular dystrophy Reproductive failure Encephalomalacia; exudative diathesis Delayed clotting time of the blood Polyneuritis

Bi

B2

Pigs Chicks Hens

Loss of appetite; severe Green leafy material ; milk; dried whey; diarrhoea dried skim-milk Curled toe paralysis Decreased hatchability of e!xs Poor growth; enteritis; dermatitis Black tongue Any source of tryptophan as nicotinamide can be synthesized from it

Nicotinamide

Pigs Poultry

-__---

Components of food Table 3.10-continued vitamin Type of animal DeficBency symptoms

87

Source of vitamin and/or cure for de$ciency Widely distributed in feeds so that deficiencies are unlikely to occur in practice Widely distributed in feeds ; deficiencies unlikely to occur in farm practice Widely distributed in feeds; also usually synthesized in the alimentary tract Widely distributed in feeds and can be replaced with methionine and betaine in the diet Widely distributed in feeds; also synthesized by intestinal microbia -.--__ Foods of animal origin; synthesized by intestinal microbia when cobalt is present; also synthesized in poultry litter Farm animals do not require any dietary source of this vitamin as they can synthesize it

B6

Pigs Chicks Hens

-__ Pantothenic acid

-__-.-.--.Pigs

Poor growth rate; anaemia; convulsions Poor growth; convulsions Poor egg production; reduced hatchability Poor growth; dermatitis; characteristic goosestepping gait Poor growth; dermatitis No deficiency symptoms seen under practical farming conditions

-.- __-Biotin

Chicks ---

-_~-.Choline

Chicks

-___ ~Perosis or slipped tendon (see Table 3.7); can be prevented by feeding choline Anaemia; poor growth

Folic acid

Chicks

B 12

All animals

Poor growth

All animals

None

Fat-soluble vitamins
Vitamin A. The animal has to synthesizevitamin A from the precursors

found in plants. These are known as carotenoids, the most widely distributed one being /?-carotene.Vitamin A is a constituent of the pigment of the red cells of the retina of the eye. It is concerned with the main-

88

Nutrition and Feeding

tenance of the mucous membranes of the respiratory tract, intestinal tract, urethra, kidneys and eyesand also plays a role in bone formation. Species and breeds within species differ in their ability to convert carotenoids into vitamin A. Sheepand buffaloes are efficient converters while some breeds of cattle such as the Jersey are relatively inefficient converters, their butterfat and body fat containing large quantities of the unconverted yellow carotenoids. Vitamin A is stored in the liver so that deficiency symptoms may not appear for quite a long period after animals are fed on a carotenoiddeficient diet. In the monsoonal and dry tropics grazing ruminant livestock may suffer from vitamin A deficiency towards the end of prolonged dry seasons.It is unlikely that grazing animals in the wet tropics suffer from a deficiency of the vitamin. All yarded or housed livestock should receive adequate green feed or synthetic vitamin A supplcaents. Poultry, in particular, are likely to suffer from a deficiency of the vitamin.
Vitamin D. This vitamin facilitates the deposition of calcium and phos-

phorus in the bonesand improves the absorption of these elements from the intestinal tract. Grazing animals in the tropics are unlikely to suffer from vitamin D deficiency as sunlight helps to synthesize it in the skin. Housed animals without accessto sunlight should receive supplements.
Vitamh E. The exact biological functions of vitamin E are not yet com-

pletely understood, but it appears to be essential for the proper functioning of a number of biological systems including the synthesis of vitamin C. Deficiency of the vitamin can cause a muscular dystrophy in calves and lambs; this can be cured by the administration of either vitamin E or a selenium salt. As the vitamin is widely distributed, particularly in green leafy material, deficiencies are unlikely to occur in grazing animals in the humid tropics. Housed animals should receiveadequate green feed or synthetic vitamin E supplements. The complete role of this vitamin has not yet been established although it is known to be necessary for the formation of prothrombin, an essential component of the blood-clotting mechanism. Symptoms of vitamin IS deficiency have not been reported in ruminants or pigs under practical farming conditions. A diseaseof cattle associated with the feeding of spoiled sweet clover (Melilotus alba), containing a compound known as dicoumarol, can be cured by the administration of vitamin K. Poultry feed should contain up to 2.5 per cent dried green forage material to ensure that the birds do not develop deficiency symptoms.
Vitamin K.

Components

offood

89

Water-soluble vitamins
Vitamin B1. A derivative of the vitamin - thiamine pyrophosphate - is a

coenzyme involved in the oxidative decarboxylation of pyruvic acid. Ruminants can synthesize the vitamin, and becauseof the fact that cereal grains are a rich source of the vitamin pigs and poultry are unlikely to suffer deficiencies. In some areas of the humid tropics the feeding of raw fish to livestock could induce a thiamine deficiency as some fish contain an enzyme known as thiaminase that destroys thiamine in the remainder of the feed.
Vitamin Bz. Riboflavin is an important constituent of the flavoproteins.

These are concerned in carbohydrate metabolism. In practice, deficiencies may occur in the diets of pigs and poultry fed mainly on cereals. As poultry excreta is often richer in riboflavin than their diet, the floor brooding of chicks is advantageous when high cereal content diets are fed. The precursor for this vitamin is the amino acid tryptophan. The vitamin functions as part of two important coenzymes that are involved in hydrogen transfer in living cells. In practice, deficiencies are only likely to occur where pigs and poultry are fed very high maize content diets.
Nicotinamide. Vitamin Be. Pyridoxal phosphate, a derivative of the vitamin, serves

as a coenzyme in a number of metabolic reactions. Deficiencies do not normally occur under practical farming conditions.
Pantothenic acid.

This vitamin is a constituent of coenzyme A. Because of the wide distribution of the vitamin in feeds, deficiencies are rarely reported in farming practice.
Biotin. All the metabolic functions of this vitamin have not yet been ascertained, but it is known to possessan important role in fat synthesis. Deficiencies aie rare in farming practice.

This vitamin hasa role in several metabolic reactions, a derivative of it being important in the transmission of nervous impulses. AS it can be replaced in metabolic functions by the amino acids methionine and betaine and is widely distributed in feeds, deficiencies are rare in farming practice.
Choline.

The vitamin plays a role in various enzyme systems. In practice only chicks are likely to suffer from deficiencies although the prolonged oral administration of sulpha drugs may induce deficiency symptoms in other livestock due to the depression by the drug of bacterial synthesis of the vitamin.
Folic acid.

90

Nutrition

and Feeding

Vitamin B12. Compounds formed from this vitamin are of importance

in many metabolic reactions, particularly in propionic acid metabolism in the ruminant. As long as adequate cobalt is available and its utilization is not blocked, vitamin B 12 can be synthesized by ruminants and non-ruminants. Synthesis of vitamin B 12can also apparently take place in deep litter in poultry houses. All domestic livestock can synthesize this vitamin. It has an important role in the oxidation-reduction mechanisms of living cells.
Vitamin C.

It will be realized from these brief notes that, except under special circumstances, vitamin deficiencies are rare in free-grazing animals. However, the situation with housed animals is different. Non-ruminant livestock and particularly poultry are liable to suffer from vitamin deficiencies if their rations are not supplemented. Commercial poultry rations are usually fortified with adequate vitamins, and farmers who mix their own rations can normally purchase vitamin premixes in most tropical countries. The small pig and poultry farmer with no accessto commercial mixed rations or vitamin premixes should always provide some green feed for his livestock. Poultry are less likely to suffer from vitamin deficiencies if they are managed on deep litter. Additives Compounde rs of animal feeds normally add vitamin-mineral supplements to their rations. In recent years they have also been adding specific amino acids, such as lysine, to rations that are considered to be deficient in the amino acids. These constituents can all be considered essential feed additives. In addition, however, compounders have been increasingly adding, in those countries where it is legal, other substancessuch as antibiotics, hormones, arsenicals, tranquillizers, detergents and - in the case of pig rations-additional copper sulphate. These are chemical substances produced by microorganisms which in minute quantities inhibit the growth of other microorganisms or even destroy them. Antibiotics can be classified into two groups: broad-spectrum antibiotics that inhibit the growth of a wide range of micro-organisms and narrow-spectrum antibiotics that only inhibit the growth of one or a small number of other micro-organisms. It has been shown that low-level intakes of antibiotics can improve the productivity of pigs, poultry and young calves. In pigs the average response to feeding antibiotics has been a 10 to 15 per cent increase in growth rate and a 3 to 5 per cent improvement in the efficiency of feed utilization. The response is greatest where standards of hygiene and management are poor, in young rather than older animals and when all-vegetable protein diets are fed. The optimum
Antibiotics.

Components

offood

level of incmsion of antibiotics is considered to be within the range of 5 to 15 gm per tonne of the rations and it is generally recommended that feeding should continue throughout the life of the fattening pig since abrupt withdrawal may reduce liveweight gain and nullify the initial advantages. With poultry the degree of response to antibiotics varies with the standards of health and management and the age of the birds, Turkey poults respond better than chicks with increases of up to 15 per cent in growth rate being recorded. In young calves the inclusion of antibiotics in the diet reduces the incidence and severity of scours and improves growth rates. There is also some evidence that the inclusion of antibiotics in the rations of mature ruminants, fed rations that are composed mainly of concentrates, can be of value. It is probable that the improvements in productivity achieved by the use of antibiotics are mainly due to the suppression of sub-clinical infections, although there may also be other effects. The disadvantage of using antibiotics in feedsis that resistant strains of micro-organisms may ultimately develop. However, the prolonged useof antibiotics in livestock feeds at the normal recommended levels is unlikely to become a health hazard to the consumers of animal products. In the United Kingdom and in some other countries recommended inclusion rates of feed antibiotics such as zinc bacitracin have been promulgated by the authorities. Readers interested in maximum and recommended inclusion rates for pig, poultry and calf rations should consult Lucas (1972). Hormones. Synthetic oestrogenic hormones, such as stilboestrol and hexoestrol, possess growth-promoting properties, while thyroxine can stimulate growth and milk and wool production. Hormones may be administered to farm animals either orally or by subcutaneous implantation. With implantation, pellets are placed at the base of the ear of ruminant livestock, and in the neck in the chemical caponization of cockerels. At slaughter of the livestock the tissues from these sites can be discarded. The action of hormones in caponized cockerels encourages fat to accumulate in the body tissues.In ruminants the action of the hormones is quite different. The carcasesof treated animals contain more muscle, bone and water and less fat than the carcasesof untreated animals. The normal amounts of hormone implanted or fed orally to cattle and sheep are as follows: 45 to 75 mg of hexoestrol implanted or 10 mg in feed per day for beef cattle, and 10 to 15 mg implanted or 2 to 4 mg in feed per day for sheep. The best results are obtained during the final stages of growth when the animals are maintained on a high plane of nutrition. Average increasesin liveweight gain can be of the order of 25 per cent.

92

Nutrition

and Feeding

The use of hormones is a very controversial subject at the present time and their usefor implantation purposes or in feedsis illegal in many countries. The most serious criticism is that there is a human health hazard from the possible carcinogenic properties of residues of the hormones in carcases.Excretion from hormone-treated animals could also contaminate pastures and endanger the reproductive cycle of breeding animals grazed at a later date on the pastures. Somecompounds containing arsenic, such as arsanilic acid, sodium arsanilate, arsenic acid and arsenobenzene, appear to possess growth-promoting properties. This is probably due to their effect on the intestinal microbia. As arsenic is an accumulative poison very considerable precautions must be taken if these compounds are to be used in feeds,although there is no evidence that the feeding of these compounds produces carcases with unacceptably high tissue concentrations of arsenic.
Arsenicals.

There is evidencethat the feeding of some of thesecompounds, which are normally used to reduce hypertension and nervousness, may improve liveweight gain.
Tranquillizers. Detergents.

Evidencefor the useof some of thesecompounds as growthpromoters is contradictory.

Copper sdphate. The use of additional copper sulphate in the diet of

pigs is discussed in Chapter 12 (Part II). There are also a number of other feed additives, such as nitrovin -a guanidine derivative - and quinoxaline compounds, that appear to improve the growth rate of some classesof livestock. Coccidiostats used in poultry rations and the drugs used in the treatment of histomoniasis in turk.eys also act as growth stimulants. Apart from the use of mineral-vitamin premixes, the fortification of amino-acid-deficient rations with individual amino acids and the use of feed antibiotics, the feeder in the tropics should be very cautious in his use of the very large number of additives that are available. This is because virtually all investigational work on additives has been conducted in the temperate zone and little is known of the effects of many of them on animals managed in a tropical environment. Readers with a special interest in the subject should consult Lucas (1972).

Evaluation of foods
Although it is possible with modern equipment and methods to determine the individual components of foods, most of the information that

Evaluation of foods

93

we possess their composition is based on what is known as proximate on

analysis.

In this system of analysis the food is divided into six fractions. These are as follows: 1. Moisture: including any volatile acids and basesthat may be present; 2. Ash: this fraction includes essential and inessential mineral elements; 3. Crude protein: this fraction includes protein, amino acids, amines, nitrogenous glycosides, glycolipids, B-vitamins and nitrates; 4. Ether extract: this fraction includes fats, oils, waxes, organic acids, pigments, sterols and the fat-soluble vitamins; 5. Crude $fibve:this fraction includes insoluble cellulose, hemicellulose and lignin ; 6. Nitrogen-free extractives: theseinclude soluble cellulose, hemicellulose and lignin, sugars, fructosans, starch, pectins, organic acids, resins, tannins, pigments and water-soluble vitamins. The moisture content is determined by drying food to a constant weight at 100C(212F). The ash content by ignition of food at 500C (1,060F) until all the carbon has been removed. The CP content is calculated from the nitrogen content of the food, determined by some modification of the Kjeldahl sulphuric acid digestion technique. Where N is the total nitrogen content, N x 6*25is assumed to be the CP content of the food. The ether extract content is determined by subjecting the food to continuous extraction with petroleum ether for a defined period. The residue after the evaporation of the ether is the ether extract content. The crude GL;: {CF) content is determined by subjecting the residual food from the ether extraction to successivetreatments with boiling acid and alkali ofdefined concentration. The organic residue is the CF content. The nitrogen-free extractives content is determined by subtraction of the sum of the percentagesof moisture, ash, CF, ether extract and CF from 100. Digestibility Although the potential value of a food can be approximately determined by proximate analysis, the actual value of the food to the animal can be determined only if the digestibility is known. Digested food is that portion that is not excreted and which is assumed to be absorbed by the animal. Digestibility can be measured by the use of in vivo or itz vitro methods. In in vivo methods the food under investigation is fed to an animal and the total input and output of the food constituent are measured. Thus if I representsinput and 0 output of an ingredient the digestibility can be calculated using the following formula: I-O Percentage digestibility= -I- x 100

94

Nutrition

and Feeding

Special methods using indicators have to be used for determining digestibility in grazing animals. Suitable indicators are lignin and chromic oxide. If indicators are used then when If represents the percentage of indicator in the faecesand Ir the percentage of indicator in the ration then If-Ir Percentage digestibility= --x 100 The digestibility of the same feed varies according to the species consuming it (Devendra, 197l), as will be seen from Table 3.11. Thus digestibility coefficients for a feed are probably specific for only one type of animal.
Table 3.11 The range of digestihilities of the dry-matter (DM) content of guinea grass (Panicum maximum) in cattle, water hufluloes, sheep and gouts
Type of livestock Range of digestibility of the DM of guinea grass Site of trials

Cattle Water buffaloes Sheep Goats

51-60 58-64 50-59 57

Uganda, India and the Philippines Philippines Australia and Puerto Rico Malaysia

Source: Devendra (197 1).

As any type of in vivo digestibility determination is costly and timeconsuming, efforts have been made to reproduce in the laboratory the reactions that take place in the digestive tract of animals. As a consequence,in vitro methods have beendevised that are relatively cheap and rapid to perform. The digestibility coefficients determined by these methods are usually lower than those determined on the same foods by in vivo methods so that correction factors must be used. Further correction factors are also probably required in order to equate the data for different species.Nevertheless, these new methods are invaluable for the rapid determination of the digestibility of a large number of food samples. Interested readers should consult standard textbooks and publications for details of the methods employed in both in vivo and in vitro digestibility determinations. Energy content The ability of the food to supply energy is of major importance in the evaluation of its nutritive value. The total energy content of a food is known as the gross energy. There are, however, many losses of energy within the body due to defaecation, urination, the production of methane in the digestive tract of ruminants and the heat increment or specific dynamic action of the food. These are shown diagrammatically in Fig.

Evaluation

of foods

95

3.2.That fraction of the gross energy that remains available to the animal for maintenance and productive purposes is known as the net energy. The first demand on net energy is for maintenance of the body processes, that remaining may be used for productive purposes. Total heat production - that is the heat released by specific dynamic action of the food, together with the heat resulting from the maintenance of body processes and production -can be a source of major difficulty for the animal in the tropics (see Ch. 1, Pt I).
Gross energy

rr Faecal energy I Methane energy

Digestible energy I

I Urinary energy

I Metabolizable energy

h Heat irkrement I 1 1 I I I Maintenance energy I (BMR activity; 1 body temperature) I I 1 Heat production I Total
L --a--L --------------

Net energy

Productibn energy I I I
I ---J

Fig. 3.2 Diagrammatic representation (McDonald et al., 1973).

of the utilization

of energy in the animal

The two major systems used in practice for the evaluation of food energy in ruminant rations are the starch equivalent (SE) system that is used in Europe and many British Commonwealth countries, and the total digestible nutrients (TDN) system that is used in the Americas and in some countries in Africa and Asia. The SE system is based on net energy. The value of the food is not expressed in absolute terms but as the fat-producing ability of a food relative to the fat-producing ability of a unit of starch. The SE of a food is therefore: Weight of fat stored per unit weight of food ingested x loo Weight of fat stored per<nit weight of starch ingested The calculation can be expressed on a net energy basis by assuming that 1 g of fat contains 9.5 kcal of energy. TDN, however, is calculated by adding together the quantities in 100 units of food of the digestible

96

Nutrition

and Feeding

crude protein, crude fibre, nitrogen-free extractives and the digestible ether extract multiplied by 225. In Table 3.12 calculations are shown of the respective SE and TDN values of barley meal.
Table 3.12
Calculations of the starch equivalent (SE) and total digestible nutrients (TDN) s/-barley meal from data on the digestible nutrients Constituent Digestible nutrient in dry matter SE factor SE TDN

Digestible crude protein True protein Ether extract Crude fibre Nitrogen-free extractives Totals Source: McDonald

12.6 11.3 i-6 0.2 65.8

0.94 2.12 I*00 140

TO6 3.4 o-2 65.8 80.0

12.6 3.6 0.2 65-8 82.2

et al. (1973).

It is generally agreed that neither of these major systems achieves an accurate prediction of the effect of food intake on the production of animals. Blaxter (1962) has therefore proposed a new system. This is a more accurate system but greater accuracy is achieved at the expense of simplicity. Evaluation of foods for pigs is considered in Chapter 12, Part II. Energy values if required are usually stated in terms of TDN, particularly in the Americas. The energy value of poultry foods is now normally expressedin terms of metabolizable energy as this component is relatively easily measured. Protein content As stated previously the CP content of a feed measures both the true protein content and the non-protein nitrogen content. In practice this is not very important as in ruminants the non-protein nitrogen fraction can be synthesized into protein by the microbia of the digestive tract and the diet of non-ruminants does not usually contain substantial quantities of non-protein nitrogen. True protein (TP) can be determined chemically. The apparent digestible crude protein (DCP) content can also be determined using the digestibility techniques discussed in a previous section. As the DCP content of a feed is not an entirely satisfactory assessment protein value for an animal, becausethe efficiency with of which a protein is used differs according to its source, several other methods of evaluating protein have been devised. The major ones are:
Non-ruminants.

Evaluation

offoods

97

the protein eficiency ratio (PER), the gross protein value (GPV), the protein replacement value (PRV) and the biological value (BV). The BV is probably the method that is mainly used. It may be calculated from the formula N-(FN-MFN)-(UN-EUN)X N-(FN-MFN) 1oo

where N is nitrogen intake, FN is faecal nitrogen, MFN is metabolic faecal nitrogen, UN is urinary nitrogen and EUN is endogenous urinary nitrogen. Some BVs of the proteins of common foods are shown in Table 3.4. The BV of a food protein depends upon the number and the types of amino acids present in the protein. The closer the amino-acid composition of the food protein approaches to the amino-acid composition of the body protein the higher will be its biological value. It will readily be seen that the BV of a mixture of foods will not be a mean of the individual B;Vs.In general, animal proteins will possesshigher BVs than plant proteins. Amino-acid estimations can be made by chemical (lysine) or biological methods (methionine and cystine) or by microbiological assay. Relevant textbooks should be consulted as to the methods employed. In practice a CP estimation is normally used for the evaluation of the protein content of feeds in pig and poultry rations, together with an assessment the ability of the proteins to supplement the known aminoof acid deficiencies of the cereal part of the rations. Proteins in ruminant rations are normally evaluated in terms of CP or DCP. The concept of protein equivalent (PE) was introduced to allow for the value of the non-protein nitrogen fraction in the CP. The PE was calculated as follows: PE Percentage digestible TP + percentage digestible CP = 2 or
Ruminants.

PE Percentage DTP + percentage DCP = 2 where DTP is the digestible true protein. There does not appear to be any particular justification for the use of PE rather than DCP in the evaluation of the protein content of ruminant feeds. Any assessmentof protein quality in ruminant feedsis dimcult as food proteins are utilized by microbia and transformed into microbial protein. This has a high biological value of approximately 80. Usually the ruminant cannot take full advantage of this situation as some food

98

Nutrition

and Feeding

protein is transformed into ammonia. Although some of this ammonia is used by the microbia most of it will be absorbed into the bloodstream and converted into urea, only a part of which will be returned to the digestive tract via the saliva and other metabolic pathways. The remainder is excreted in the urine. The biological value of food proteins in ruminants therefore depends to some extent on how much ammonia is found in the rumen and how this ammonia is then utilized. The latter will depend upon the availabi!ity of an adequate source of energy for the microbia in the digestive tract.

Feeding standards and requirements

Feeding standards for the different classes of tropical livestock, where they are known, are detailed in the chapters concerned with the various types of livestock in Part II of this text. Information on temperate-zone feeding standards, that have at present to be generally used in the tropics, is available in Morrison (1957) Evans (1960) Agricultural Research Council (1963, 1965, 1967) National Academy of Sciences-National ResearchCouncil (1966, 1968a, 1968b, 1970, 1971) and McDonald et al, (1973). A major difference in the nutrition of the animal in the tropics is that heat produced within the body generally has to be dissipated, whereas in the temperate zone it can often be conserved in order to maintain normal body temperatures. Thus it might be expected that energy requirements for maintenance would be lower in the tropics. Kehar (1954) suggested that the energy requirements of cattle in India could be as much as 40 per cent less than generally accepted requirements in the temperate zone. A difference of this magnitude is not apparent in practice, but other workers at the Indian Dairy Research Institute have suggested that animals need less energy during the hottest periods of the year. It has also been suggestedthat the protein requirements of animals may be lower in hot climates, but it is likely that mineral and vitamin requirements will be the same at similar levels of production. Payne (1969) has discussed these and other problems in some detail in a review of the nutrition of animals in the tropics.

Types of feed available

There are obviously many methods that could be used for the classification of animal feeds.One method used in practice is to classify them into groups according to the predominant nutrient or nutrients which each contains. A selection can then be made from feeds within any one group for availability and cost, and the most suitable feed from any one group can be used in a mixed ration. In practice, any food from one group

Types offeed available

99

can be substituted in a mixed ration for any other food within the same group without substantially altering the nutrient balance of the mixed feed. It would, however, be a mistake to assume that all feeds within a group possess exactly the samefeeding value or that the samefeed grown in different parts of the tropics will have exactly the same composition. One suitable classification by feed group is as follows: 1. Succulents.These are feedswhose principal constituent is water. They may be sub-divided into two major sub-groups: green forages and root crops. 2. Roughages. These feeds are characterized by their high CF content. Some feeds that can be classified as succulents when they are young would be classified as roughages when they are mature. 3. Concentrates. The main characteristic of all concentrate feeds is that they contain relatively large quantities of a major food constituent. They may be sub-divided into concentrates of plant or animal origin, and concentrates of plant origin may be divided into a further two groups - those that are energy-rich and those with a high CP content. 4. OrCtmfeeds.Some feedscannot be easily classified in any of the three major groups. This group includes new types of feed that are manufactured from inorganic or organic materials. In compounding any specific ration the relative quantities of feed used from these different groups will vary according to the species or breed within the speciesof animal which is being fed, and in accordance with the animals production. For instance, ruminant animals can utilize food from all groups, but they utilize succulents and roughages more efficiently than other livestock and it may sometimes be wasteful to feed concentrates to them. Non-ruminants require mainly concentrate feeds, as do highly productive milking cows. Some data on the nutrient content of representative feeds from each of the groups are shown in Table 3.13. Succulents This group includes practically all growing or fresh vegetation of which forages form a major part. The water content of succulents is always high-usually between 75 and 95 per cent. These are of such major importance in the feeding of livestock that they are considered at length in the second section of this chapter. During their growth stage most forages constitute a complete food for ruminant livestock, as long as they are not grown on soils that are deficient in an essential nutrient. Within the forage group, legumes usually possesshigher CP, mineral and vitamin contents than grasses. Browse legumes normally possess particularly high CP and mineral contents, though their digestibility may be somewhat less than those of forage legumes.
Forages.

Table 3.13
Group

The proximate analyses of some representativefeedstufi Feedstuff Dry matter (%I Crude fibre (%I Ether extract (%I On DM basis N-free extractive CP (%I (%I Phosphorus (%I $

Ash (%I

TDN (%)

Calcium (%)

Succulents

Panicum maximum (guinea); IS to 28 days growth

Centrosema pubwcens LZ$fZ) esculenta

19.4 24.2 37-l

14.3 8.3 3.0

30.9 31.0 4.3

3.4 3.9 0.9

39.2 357 88.3

12.2 21-l 3.5

10.2 14.9 31.6

O-86 I.30 026

O-36 o-3 I O-16

(cassava) roots Roughages

Cynodon dactylon

(Bermuda) hay
Oryza sativa

87.4 89-O

10.2 17.7

32.0 35.7

5.3 2.4

39.2 38-7

13.2 54

43.2 37.7

o-75 O-29

O-23 O-36

(rice) straw

Concentrates
Plant origin

Energy rich

Oryza sativa Zea mays

(paddy)

89.4 87.2 90.2 91.1 93.7 91.0 907 93.7 93.9 73.4 23.7

7.9 3.3 5.2 6.6 II.6 6.2 20.5 28.1 8.0 8.6 1-O

9.5 3.0 14.6 13.3 6.2 5.9 02 2.2 0.6 0.0 3.6

3.8 3.7 22.6 2.4 9.0 4.9 6.8 10.6 l-1 o-0 t-6

68.5 79.5 36.5 47.4 23.6 30.0 0.3 3-l 51-I 61.7 t 1.8

102
IO-5

51.9 66.5 53.9 68.6 71.3 77.9 72.4 65.3 79.8 53.7 16.1

0.41 0.20

o-50

(maize) cracked grain

Gossypium spp.

0.22

Protein rich

Animal origin

(cotton) seed Coconut meal (solvent extracted) Sesame meal Soybean meal (expeller extracted) White fish meal Meat and bone meal Dried skim-milk Cane molasses Wet brewers grain

20.8 21)4 43.3 44.0 62.9 49.7 33.1 3.0 5.7

0.21 2.02 0.27 6.76 IO.67 1.28 0.66 0.07

0.64 1.61 0.63 3.69 5.27 1.04 0.08 0.12

Other

Sources: Composite data from Morrison

(1957) and McDowell et al. (1974).

102

Nutrition

and Feeding

Roots.

These are grown in large quantities in the monsoonal and wet tropics. The major root crops in the tropics are cassava (Munihot es&en@, yams (Dioscoreu spp.), sweet potato (Ipomoeu bututus), taro (Colocusiu esculentu) and arrowroots (Muranta urundinaceu and Cunna edulis). Breadfruit (Artocarpus ultilis), although not a root crop, has approximately the same feed value as the typical root crops. The energy content of the dry matter of root crops is high, the water content varies, but the CP, mineral and vitamin contents are normally low. Silagesmade from succulents will vary in nutritive value according to what feed was originally ensiled. The feeding value will always be lower than the fresh material due to wastage during the ensiling process. Roughages Jt is probable that a majority of ruminant livestock in the tropics subs&t on roughage for a major part of the year. This includes hay, standing hay and artificially dried forage. Good hay made at the correct stage of growth can be an excellent maintenance roughage feed. UnforYunately, little good hay is made in the tropics. Most of the hay that animals consume is standing hay or forage that has matured in situ. Under, favourable circumstances, such as occur in the Sahelclimatic zone of Africa and in some regions of the savanna, standing hay can be a medium-quality roughage. Normally. however, it is fully matured forage with a very low feeding value, containing a high content of indigestible cellulose and lignin. Artificially dried forage may bc classified as a roughage or as a concentrate feed, depending on the nutrient content of the original forage and the stage at which it was dried.
Forage roughage. Straws and huulm. The straws of the cereal crops grown in the tropics such as rice (Oryzu sutivu), maize (2%~muys), sorghum (Sorghum vulgure) and millet (Pennisetum typhoideum) are almost universally used for feed-

ing purposes, as are the haulms of legume crops such as groundnut (Aruchis hypogueu) and the dried stalk materials of such crops as sesame (Sesumumindicum) and cotton (Gossypium spp.) These roughagesgenerally possessa low feeding value, but livestock grazing them also have access to the weeds in the stubbles and these improve the overall feed value of straws and haulms. Legukmestraw or haulm possesses higher feeda ing value than other straws. There are other roughage feeds available at specific locations in the tropics such as the waste materials from oil palm processing plants, sisal waste, pineapple waste, bagasse,&coa pods, coffee hulls, etc. The feeding value of these materials varies, but it is usually low.

Types of feed available

103

Concentrates There are of course a wide variety of concentrate feeds produced in the tropics, but those that are normally economic to feed are usually the byproducts of crop or animal production. Concentrates of plant origin These include both energy-rich and protein-rich feeds. The major group are the cereals -rice, maize, sorghum and millet. Their SE and TDN are high, CP content medium and CF content low. Their mineral content varies but is often unbalanced. As they are normally the staple human food in the countries in which they are grown it is unusual for them to be used as animal feedsexcept in restricted quantities in specific regions. If a surplus of cereals is available it is usually most economic to feed it to nonruminants and in particular to poultry. Other energy-rich concentrates are sugar and dried roots such as dried cassava, a major product in some parts of Southeast Asia where it is produced for export to Europe. Sugar is not normally fed to livestock, but when sugar prices are depressed it is an excellent feed to incorporate in the ration of young pigs. Oil seeds,from which many of the protein-rich cakes are manufactured, are high energy content feeds, principally on account of their high fat content. They are not usually used for feeding purposes, the excep tion being cottonseed.
Energy-rich concentrates of plant origin.

Grain legumes, like cereals, are normally used for human consumption, but where there is a surplus they can be fed to animals. The quality of their protein is relatively high. Oil cakes and meals are the most common protein-rich concentrates. They include groundnut, soybean, sesame,palm kernel, cotton, rubber seedand coconut. Usually they have a relatively high SE or TDN content and a CP content that varies from 15 to 45 per cent. Their feeding value depends partly on the amount and quality of the protein that they contain, partly on the amount of fibre and other inedible material retained in the cake after processing of the original plant material and partly on whether they contain any toxic materials after processing. Their oil content varies according to the method of processing. Those cakes processedby primitive methods wiIZcontain quite a high oil content-sometimes as much as 10 per cent. Their mineral content varies considerably, but they are often rich in phosphorus. Their vitamin content will be minimal on account of the processing. Some oil seeds, such as cotton and rubber, contain compounds that are toxic to non-ruminants. Cotton seedscontain an aromatic aldehyde known as gossypol, toxic to nonruminants at low levels, but inactivated by special processing. Rubber
Protein-rich concentrates of plant origin.

104

Nutrition

and Feeding

seed contains a cyanogenetic glucoside known as linamarin and an enzyme known as limase which hydrolyses the glucoside, The raw cake can therefore be toxic and should be boiled or processed in some other way before feeding. Concentrates of animal origin These include the following: the by-products of meat animal processing such as meat meal, meat and bone meal and blood meal, the by-products of fish processing such as fishmeal and shrimp meal, and the by-products of milk processing such as skim milk powder, whey and buttermilk. They are characterized by the relatively large quantities of high-quality protein that they contain and by a high mineral content. The vitamin content of some of these feeds may also be considerable. In general they are not freely available in the tropics as most animal and fish offals are consumed by the human population and there is insufficient milk available for processing. In some tropical countries there is a waste of these by-products, either becausesuitable processing plants are non-existent, existing processing plants are inefficient or becauseof local ignorance of the value of the by-products. Other, including unconventional, feeds These include by-product feeds such as sugar and citrus molasses, brewery and distillery by-products like brewers grains and fermented feeds.The latter are of some importance in certain regions of Southeast Asia. There is at present in industrialized countries a very considerable interest in new sources of feed. Many of these new feeds could be of value in the tropics. The possibility now exists for plant breeders to breed plants that are more nutritious. An example is the breeding of high-lysine varieties of maize. Obviously this idea could be of the greatest interest in the tropics where new highly nutritious varieties of all types of crops are urgently required.. The use of waste faecal material for the culturing of chlorella is at present practised only in some countries in Asia, such as Taiwan. The chlorella can then be used as a protein concentrate for pig feeding (see Ch. 12, Pt II). Possibilities for the processing and utilization of leaf protein or grass juice containing leaf protein are particularly promising in the humid tropics. The leaf protein concentrate with a CP of 60 to 70 per cent could be used for non-ruminant feeding while the discarded material could be used as a roughage for the feeding of ruminants. Readers with a particular interest in this unconventional feed should consult Pirie (1969). The production of single-cell protein concentrates using specific yeasts

Forage

105

from waste carbohydrate materials such as the contaminated flour from cereal processing mills or from paraffins or gas oil is slowly developing. The process that useswaste flour as a substrate produces a high-quality protein suitable for human consumption while that which usesparaffins or gas oil produces a 63 to 70 per cent CP concentrate suitable for feeding to pigs, poultry and calves. In tropical countries where there are by-product carbohydrate wastes or plentiful supplies of natural gas these new processescould be of value. In the Philippines an edible yeast has been successfully cultured on coconut water, a product that is normally wasted at copra-processing plants. A protein of bacterial origin is already on the market in the United Kingdom. New ideas are also now current as to how waste wood materials could be hydrolysed in order to break down the cellulose and lignin into feed products that could be digested by animals, and promising new and apparently economic methods are emerging. The possibilities for the utilization of these new methods in the tropics, both for the processing of waste wood in the wet tropics and straws in all regions of the tropics, are obviously immense. Readers who are particularly interested in this subject should consult Bender et al. (1970) and Heany and Bender (1970). A protein known as pekiloprotein obtained by the growth of a fungus on wood pulp substrate is at the pilot stage of production and may also offer considerable possibilities in the tropics.

Storage of feeds
The storage of all feeds is a very considerable problem in the tropics. Some information on the storage of forage as hay or silage is given in the next section of this chapter. All feeds should be stored in dry, vermin-proof accommodation, in which air can circulate freely and where ambient temperatures can be kept as low as possible. Concentrate feed stores should be regularly treated with an insecticide. The normal method of treatment is fumigation. Protein concentrates are particularly liable to deterioration. The fats in them may undergo oxidative changesand become rancid, making the feed unpalatable. Fungal infection of some concentrate feeds can lead to the feed becoming toxic. This is particularly likely to occur in hot, humid climates.

Forage
It is estimated that 20 per cent of the worlds surface is covered with planted pasture or fodder, range plants or some other form of forage.

106

Nutrition

and Feeding

Figure 1.5 provides some information on the world distribution of the principal plant associations and it can be seen that there is a considerable area of rangeland in the tropics, principally savanna, but also some steppe and desert scrub. In addition there are very large areas of grasslands occurring within other associations, such as the rain-forest or the undifferentiated highland regions. At present about 10 per cent of mans global supply of food is produced from pasture or range while about 90 per cent comes from cropped land, though it is unlikely that man in the tropics obtains 10 per cent of his food from tropical pasture and range land. Quantitatively, therefore, the contribution of pasture and range to mans food supply is not large, though qualitatively it is important as animal products such as milk and meat have a high dietary value. The contribution of pasture and range to the feeding of mans domestic, ruminant livestock in the tropics is very great, as an extremely high proportion of them subsist on natural grazings. In the tropics, as elsewhere,man utilizes for his ruminant livestock land that cannot be used for any other purpose at the present time; broken hill country, swamps, semi-deserts and heavily leached soils of low fertility. There are some regions, however, such as northern South America and parts of Africa where man at present utilizes areas of good fertile soil as ruminant livestock range. As total population increases and demand for food rises it is inevitable that most of the fertile land now utilized as range will be cropped. In addition, by the drainage of swamps, the irrigation of semi-arid areas and the improvement of less fertile soils by the increasing use of fertilizers, even large areas now considered only economic for use as rough grazings will also be cultivated. Thus, it may be forecast with someconfidence that the total area of range available for ruminant livestock in the tropics will diminish in the future. This situation emphasizes the growing importance of making provision for alternative sources of feed for ruminant livestock. One major alternative is forage, considered as a crop. Forage may be produced from perennial or permanent pastures and/or fodder crops and trees, from short-term pastures or leys, from annual or catch-crop pastures and fodders, as a by-product of other crops such as cereals and pulses or in association with other crops such as fruit, nuts, oil palm and rubber. Forage, and particularly pasture, could also be important in the cropped area as it could help to restore the fertility of soils that have been worn out by continuous cropping or used to maintain or even improve fertility by alternating it with crops in a rotation. The system of alternating forage with other crops is known as alternate husbandry and this could be a viable system in some tropical countries, particularly in the montane areas.The association of forage with other crops is known as integrated husbandry and is particularly relevant to conditions in the humid tropics where, in the forrn of pasture, it can be associated with tree crops. The reasonswhy forage crops, and particularly mixed forage crops such

Types offorage

107

as pasture, are so useful are complex. The root system of grasses tends to bind the soil together, thus reducing the risk of erosion. At the same time it improves the physical condition or tilth of the soil, adds humus and thus improves both fertility and water-retention properties. In addition, legumes in a pasture mixture will add to the total availability of nitrogenous plant foods in the soil, as bacteria in the nodules on their roots possess facility to fix nitrogen obtained from the air and make the it available to other plants. Finally, the grazing animal returns organic matter to the soil in the form of faecesand its urine appears to possess some special plant growth-promoting properties. Thus the soil, plant and animal are all components of a biological system that, when manipulated intelligently, can be used to maintain or even raise the fertility of most soils.

Types of forage
Natural grazings or range Davies (1960) has suggestedthat kmajorchanges in the type and composition of the vegetation occur approximately at latitudes 30N and S of the equator and that the area between could be conceived as the biological tropics. One-half of the worlds natural grazings are found within this area. Climate, and in particular the total amount and the seasonaldistribution of rainfall, is probably the most important factor affecting the use of land for natural grazings. Ranges occur most frequently in regions receiving from 250 to 2,000mm (10 to 79 in) total rainfall per annum and less frequently in regions of lower and higher rainfall. Topography, soil and the occurrence of natural fires are also factors, but the other most important single factor is undoubtedly the influence of man. Very large areas of range have beenderived from forest as the direct result of mans activities. The major types of natural grazings in the tropics are those associated with rain-forest, dry woodlands, savanna, tropical steppe, semi-arid regions, montane areas,seasonally flooded land and permanent swamps. It is impossible in a brief account to list and describe all the forage species found in these areas, and only a selectedfew will be mentioned. Readers who have a special interest in specific forage plants or plant associations should consult such general texts as Whyte et al. (1953, 1959)and Whyte (1974),or specifically for Australia (Moore, 1970), Africa (Rattray, 1960), the Americas (Beard, 1944; Blydenstein, 1967) and Asia (Whyte, 1964, 1968; Dabadghao and Shankamarayan, 1972). Rain-forest grazings These include evergreen equatorial, evergreen and semi-evergreen monsoonal forests and monsoonal deciduous forests. Annual rainfall in these

108

Nutrition

and Feeding

forests is usually above 1,520mm (60 in), but in some monsoonal regions moist deciduous forests occur in areas where the annual rainfall may be as low as 1,020mm (40 in). In virgin forest grasses are either absent or extremely sparse. However, if the forest is clean cleared or slash-bum agriculturists return too frequently to the same area, forest trees are replaced by a mixture of forage plants and shrubs. If these are then burnt annually neither necessarily regenerate and the forest is replaced by an ecologically unstable range association. On account of this processmajor forests have largely disappeared in the monsoonal regions; in particular the monsoonal deciduous forests are disappearing at an ever-accelerating rate in the equatorial regions of Southeast Asia, Africa and tht: Americas. Some of the dominant grasses found in grazings derived from these forests are: Imperata cylindrica, Themeda trianda (red oat grass), Dichanthium spp., Schima spp. and Ischaemum spp. in South Asia; Imperata cylindrica and Ischaemum spp. in Southeast Asia; Andropogon spp., Digitaria spp., Hyparrhenia spp., Pennisetum purpureum (elephant grass), Ctenium newtonii and lmperata cylindrica in Africa; Axonopus spp., Bouteloua spp., Paspalum spp., Hyparrhenia spp., imperata spp. and Panicum maximum (guinea grass) in Central and South America; and Axonopus compressus (carpet grass) and guinea grass in the tropical oceanic islands. Although there are some leguminous tree species in the rain-forests, herbaceous legumesappear to establish themselves rather slowly in the grass-bush associations that replace the forest on account of the activities of man. Nevertheless, conditions can be favourable for legumes as moisture may be more or less continuously available, so that ultimately a considerable number of legumes become established, although they may not make any major contribution to the total quantity of feed available. Commonly found are Alisicarpus spp.. Calapogonium spp., Centrosema spp., Crotalaria spp., Desmodium spp., Indigofera spp., Pueraria spp. and Stylosanthes spp. In those areaswhere tree crop plantations have replaced the rain-forest the spread of such legumesas Calapogonium muconoides (calapo), Centrosema pubescens (centro), Pueraria phaseoloides (puero) and Stylosanthes guianensis (stylo) has been encouraged by their use as ground cover beneath and between tree crops. Dry woodland grazings These occur in regions where the average annual rainfall is within the range 640 to 1,400mm (25 to 55 in) and where there is a dry season of from 4 to 7 months duration. The largest area of dry woodland in the tropics is found in Central and East Africa, where it is known as miombo. Somewhat similar associations are found in West Africa, Southeast Asia, Australia and Central and South America. In African miombo where the rainfall is within the range 890 to 1,400mm (35 to 55 in) a low tree-high grass, Combretum-Hyparrhenia association is dominant, whereas where

Types of forage

109

the rainfall is between 640 and 890 mm (25 and 35 in) trees of the genera Brachystegia, Julbernardia and Isoberlinia dominate, while the ground cover consists of a mixture of tall tussock and fine stoloniferous grass species. The grass cover may include Hyparrhenia spp., Aristida spp., Eragrostis spp. and Cympobogon spp. Dry open woodlands of this type are relatively easily fired during the dry season with the consequencethat large areas have been converted to savanna. Where these woodlands persist the relatively sparse ground cover is usually grazed. Savanna grazings Savanna is a collective tern? embracing many different types of open grasslands interspersed with trees. It may have developed as the result of climate or have been derived from rain-forest or dry woodlands. Continuous firing or periodic flooding of forest or dry woodlands may preclude the growth of the majority of trees so that they are replaced by a savanna association. It is often difficult to ascertain whether a specific savanna has developed as the result of natural causes or whether it is a result of the activities of man. In addition this confused situation is compounded by other grassland associations, such as those in the semiarid thornbush areas or those in floodplains such as the llanos of Venezuela and Colombia, being often referred to as savanna. At present, a major proportion of the range in the tropics may be termed savanna as associations of this type exist in areas where the annual rainfall varies from 508 to 1,520mm (20 to 6Oin), that is from the boundaries of the semi-arid regions to the rain-forest. Consequently there are many types of savanna association, such as high tree/high grass, high tree/low grass, low tree/high grass and low tree/low grass. Those savannas where the trees are large and relatively well spaced are known as orchard savanna, while those where the trees are usually smaller and generally grouped in clusters are known as parkland savanna. The grasses and legumes found in savanna are usually similar to those in the association from which the savanna has been derived or to adjacent associations. In general, ground legumes are scarce and the introduction of Stylosanthes humilis (Townsville stylo) has been very successful in improving the carrying capacity of many savannas. Steppe and semi-arid thombush grazings Associations of this type are found in regions that generally receive 380 to 74Omm (15 to 30 in) or less rainfall per annum. They are found both within the tropics and more extensively immediately north and south of the tropics. Generally, tropical steppe is found at medium altitudes and semi-arid thornbush at lower altitudes. Thus, in East Africa tropical steppe associations occur at altitudes varying from 1,524 to 2,134m

110

Nutrition

and Feeding

(5,000 to 7,OOOft) while semi-arid thornbush usually occurs at altitudes below 1,219m (4,000ft). These are often rather unstable associations. Perennial grass species are rapidly replaced by annuals if tropical steppe is overgrazed, while there is a tendency for bush species to spread in the thombush areas unless they are restrained by annual burning. Perennial and annual grassesare medium to short in height, providing an open and sparse cover. Some of the dominant grasses are Aristida spp. Cenchrus spp., Eragrostis spp. and Themeda spp. Herbaceous legumes are scarce or absent. There are, however, large numbers of bush legume species in the semi-arid thombush areas, particularly Acacia spp. and Cassia spp. in Africa and Prosopis spp. in the Americas. As a consequence browse provides a considerable proportion of the forage available for ruminant livestock, particularly in the dry season. Where the mean annual rainfall is below 250 mm (10 in) the semi-arid thornbush associationsmerge into desert scrub and true desert. Perennial speciesdisappear on the desert margins and are replaced by ephemerals that grow for a short period after rain. Montane grazings Evergreen and/or semi-evergreentropical montane forests are found at altitudes ranging from 1,070 to 3,050m (3,500 to lO,OOOft),depending upon the latitude and the climatic environment. Annual rainfall in these forests varies from 1,020to 5,000mm (40 to 200 in) and there may or may not be a dry season. In equatorial regions lowland tropical rain-forest gradually mergesinto humid montane forest, but in the drier tropics the montane forest may resemblea girdle encircling the higher altitude areas, with different ecological associations above and below. In these montane regions there are often tracts of open grasslands completely surrounded by forest. Grasslands of this type have probably beenderived as the result of timber cutting and fire, but some may be the result of inadequate drainage. In addition there may be high-altitude natural grazing above the forest and below the snow line, usually at more than 3,050m (lO,ol>oft) altitude. These grasslandsare characterized by short grasses,few legumes and numerous low-growing herbs. Grazings associated with montane forests are common in South and Southeast Asia. In Sri Lanka, where they are known as wet patanas, they are usually dominated by the grass Chrysopogon zeylanicus. In the montane areas of East Africa Pennisetum clandestinum (Kikuyu grass) and Trifolium semipilosum (Kenya wild white clover) are common in the wetter and more fertile areas, while in the drier areas the grasses Pennisetum schimperi and Eleusine jaegeri are common. Loudetia simplex is a common grass in the montane areas of Rhodesia. High-altitude natural grazings are of major importance in South America where they are known as parunas.

Types of forage

111

Many temperate grass and legume species will of course thrive and naturalize in the montane regions. Seasonally flooded and permanent swamp grazings Where there is seasonal alternation of flooding with dry conditions, growth of trees may be restricted to slightly higher land on the banks r#fthe rivers and streams,where they form gallery forests, or to surrounding higher land. Natural climax grasslands result. In these regions there is normally abundant forage available as the flood waters recede, but it quickly dries out and becomes unpalatable. It is then often burnt to encourage regrowth. This alternation of flooding, drought and burning produces associations in which the speciesthat survive are those that are resistant to flood, drought and fire. There are very extensive areas of this type of range in South America, particularly in Colombia and Venezuela where they are known as llanos. Grasses of major importance in the llanos are Paspalum spp., Trachypogon spp. and Leersia hexandru. Smaller areas are also found in other regions of the tropical Americas. In Africa such ranges are found primarily in the Niger and Nile valleys, and around Lake Victora in Uganda and Lake Chad in West Africa where Echinoclou spp. are of importance, but smaller areas of the samegeneral type of range are common in East Africa where they are known as vleis or mbugni. Tviany large areas are also found in Southeast Asia, particularly in the islands of Borneo and Sumatra and west of the dividing mountain range in east,c,n sub--tropical and tropical Australia. Permanent swamps may be grazed on their margins for a part of the year. They are of importance if they can be used during the dry season in association with upland areas of range that are available during the wet season. Cultivated pasture, fodder and browse Three major types of cultivated pasture can be differentiated : permanent or perennial pastures, short-term pastures or leys and temporary or annual pastures. Permanent or perennial pastures These consist of associations of perennial grasses, with or without legumesand herbs, that are grazed year after year. They may be reseeded at intervals or renovated using a number of different techniques. They are characterized by a high productivity per unit area of land and they can possessa high annual stocking capacity. They are particularly suitably for dairying or for the fattening of livestock. At present there is only a limited area of permanent pasture in the tropics, concentrated mainly in Australia, the Americas, th,eoceanic islands and certain regions of the African and Asian humid tropics. There are, however, very considerable possibilities for the rapid development of perennial pastures in most wet

112

Nutrition

and Feeding

tropical regions and in some monsoonal tropical regions, particularly in association with tree crops. Perennial pastures are not generally suited to the semi-arid and arid areas unless irrigation is available. When irrigation is available a single species forage crop may be grown. One example is the use of Medicago spy. (lucerne or alfalfa) grown under irrigation in the very dry but tropical region of southwest Arabia. Some of the most suitable grassesto use in perennial pastures in the wet tropics are: Brachiaria spp. including B. mutica (para) and B. brizanthu (signal), Panicum spp. including guinea, Digitaria spp. including D. decumbens (pangola), Ischaemum spp. and Echinochloa polystachya (Aleman). Elephant grass may also be used in perennial pastures, although it has to be managed very carefully. Melinis minutaejora (molasses)and Hyparrhenia rufu (jarhgua) are also sometimes planted, especially in the American tropics. Cynodon plectostachyus (African star grass) is increasingly used in the relatively drier regions of the wet tropics. Paspalum spp. are also of utility in specific regions. The major legumes used are centro, puero, calapo, stylo, Desmodium spp. such as D. intortum, D. uncinatum and Macroptilium atropurpureum (siratro). Cjitoria ternatea has also shown promise in some regions. Indigofera spp. are widely distributed throughout the tropics and agronomically some-speciespossess excellent characteristics. Unfortunately they contain toxir ;lhctanccs. and the plant breeders are now attempting to produce non-toxir strains. Mimosa pudica is also widely distributed throughout the wet tf epics. In some countries it is considered a valuable component 01 p,itsrure and in others a weed.Aeschynomene americana is useful in the wetter sub-tropics. Short-term pastures or leys These consist of associations of perennial grasses and/or legumes and other forage plants that are grown in rotation with cultivated crops. They may be grazed for from 2 to 5 or more years before they are ploughed and replaced by a crop. The advantage of this system is that the grass ley improves both the texture and the fertility of the soil, particularly if it consists partly of legume species.When the area is ploughed the crops that follow benefit and are more productive. After a few years cropping, the ley is established once again. Grass leys can be characterized by a very high level of productivity per unit area of land and are normally associatedwith high-level farming practices. They can be successful in those regions of the tropics only where it is practical to plant and plough-up pastures easily and where advantage can be taken of the inclusion of pasture in the rotation-that is in those climatic environments that are neither too wet nor tco dry. They are usually not suitable for use in the wet or the semi-arid tropics. For example, Meiklejohn (1962) reported that grass leys ploughed up in Ghana Cd not, as would be expected, improve the yield of the following crops and he attributed this situation to the fact that there was a lack of nitrite-oxidizing bacteria in the soil. This managerial system is likely to

Types of forage

113

be most successful in montane and some monsoonal tropica! environments. The speciesincluded in leys must establish easily and quickly and the grasses used must not be stoloniferous, or they wil! be difficult to eradicate during the cropping period of the rotation. As grass leys are particularly suited to the montane tropics, temperate-type grass and legume speciescan often be used. Temporary or annual pastures These are usually single-species grass, legume or other plants grown specifically as forage within a crop rotation. They are usually characterized by a high yield per unit area of land and a high cost per unit weight of forage compared with that produced from perennial pastures. In the wet tropics the major fodder used is one or the other strain of elephant grass, but Saccharum spp., Sorghum spp., Punicum spp. and Tripsacum hum (Guatemala grass)are also used in specific areas. In the drier tropics the main fodder grassesare Sorghum spp. Legumes are not usually grown as annual crops for forage purposes, but certain annual pulses such as Stizolohium spp. (Mauritius or velvet bean) may be used for this purpose. Cultivated browse Wild trees and bushes, particularly the leguminous species,are a very important source of forage for livestock. They are of major importance in the dry tropics as many of them remain green long into the dry season after the ground vegetation has dried out and their leaves generally possess high nutrient content. However, the number of trees and bushes a specifically planted for forage production purposes is limited. As desirable browse species may be left intact when other bush is cleared it is often d&cult to distinguish between cultivated and uncultivated browse. In the drier regions of Australia the salt bushes Atriplex spp. and Kmhiu uphylla are important wild fodder species while Leucaena Zeucocephdu is also an important forage, both cultivated and wild, in the oceanic Pacific islands. Roseveare(1948) listed 385 browse species that are utilized in the Americas. Some of these,such as Gliricidiu sepium, are semi-cultivated as they are us,edfor live fencing. In the drier areas of Africa a high percentage of the trees and shrubs are browsed to some extent by livestock and/or game; in East Africa the percentage is said to be as high as 75. Many trees are also lopped during the dry season (Plate 3.1),so that the leaf material on the brarrclhescan be eaten by small stock such as sheep and goats. 2,ezii; sgi;. are particularly usefur itidrthis purpose and one species,A. ulbida, even if it is not now considered to be a cultivated plant, is believed to have been planted in the past. It is particularly useful as it remains green throughout the dry season.In the Sahel and savanna regions of Africa the tree from which gum arabic is produced, A. senegul, is also useful as a browse tree. It might be possible

114

iduutrition and Feeding

Plate 3.1 Stripping trees for fodder during the dry season in the African savanna (Dr N. C. Wright).

to integrate gum arabic with livestock production in these regions by planting A. Senegalas a field hedgeor in association with pasture. Browse from trees and bushesis also widely used for livestock feeding in South and Southeast Asia; the Commonwealth Agricultural Bureau (1947) listed ninety-one speciesthat are used for browse in India and Sri Lanka. Sesbuniu grundijloru is certainly widely cultivated for use as a browse tree, particularly in Indonesia. Leucuena leucocephala is also used everywhere that it grows wild and it is increasingly cultivated in Southeast Asia.

Management offorage

115

Management of forage
The principal objective in the management of both natural and planted forage should be to maintain, as far as is possible, maximum productivity on a year-row4 hzf k of the more desirable specieswith the intention of obtaining maximlum production from the livestock that consume them. The principal problem is an imbalance between forage availability and livestock requirements that is worse on natural than on planted grazings and in dry compared with wet climatic environments. In order to attempt to attain the objective and mitigate the problem the livestock owner must manipulate the complex ecological factors that are involved (Fig. 3.3) to

Rr----7
Run-off: streams, rivers, dams Land Water Soil moisture: ground water I Soil nutrients I Plant nutrients t Faeces and urine Rain-fed and/or irrigation Solar energy 1 Forage t Management of animals

Fertilizers

Fig. 3.3 Diagrammatic representation of the major ecological factors involved in forage production.

best advantage. Methods by which the ecological complex can be most economically manipulated will obviously vary from region to region. It is our intention, therefore, to consider some of the more important factors involved in the production oic Ir.age under extensive and intensive managerial conditions. Natural grazings Natural grazings often provit,30the only major source of forage available (Plate 3.2). They are often, but not always, the most economical source

116

Nutrition

and Feeding

Plate 3.2 Unit).

Nomadic cattle herded on natural grazings in Africa (Shell Photographic

of forage. The problems of managing extensive natural grazings vary widely from the wet to the semi-arid and arid tropics. In an introduction, however, it is only possible to provide a general outline of managerial requirements. Rain-forest grazings Associated grazings, with few exceptions such as the natural stands of elephant grass in Africa, are usually of low productivity. It is often economic to replace them by planted pastures and fodder crops or with integrated tree-crop pastures.The productivity of the Imperutu spp. grasslands that are so common in Southeast Asia may be improved by the introduction of a legume. The legume generally recommended is stylo. Once the introduced legume has improved the fertility of the soil, improved grassescan be introduced. Dry woodland grazings These grazings are only moderately productive. When the available species are young they are readily eaten by livestock, but they soon

Management offorage

117

become rank, unpalatable and of low nutrient quality. The density of the grazings can be improved by thinning of the trees and coarse grassesand bushes can be controlled by the judicious use of fire In general, the length and severity of the dry season precludes the use of cultivated pasture in these regions. Savanna, steppe and semi-arid thornbush grazings These are often unstable associations under grazing conditions. Many of them are dominated by coarse grassesthat are palatable and nutritious only when they are very young. If ungrazed they dry out to form lowquality standing hay. There is a general tendency for the grazings to be invaded by shrubs and this is encouraged by the exclusion of fire and by overgrazing. Thus the judicious useof fire is an important managerial tool. In Africa, bush invasion may not only reduce the total availability of forage but may also encourage the spread of the tsetse fly. As total annual rainfall decreasesand the dry seasonlengthens the ground cover may become very sparse, perhaps forming only 5 per cent of the total area, and perennials are replaced by annuals. Supplies of free water may also become very limited, particularly in the dry season,so that livestock may have to travel long distances between watering points. In some of the savannaareas that are derived from forest and in other areas where rainfall is regular, planted pasture and forage crops may be economic. This is not usually the casein the tropical steppe or semi-arid thornbush areas. The potential and actual carrying capacities are therefore very varied -quite high in the most favoured savanna regions and very low in the semi-arid thornbush regions. Montane grazings The productivity of these ranges may be relatively high, but of course it varies with the total and seasonal incidence of rainfall, the fertility of the soil and the species available. High rainfall often causes leaching and increasesthe acidity of the soils. This is the situation in Sri Lanka where montane pastures require regular applications of lime and other fertilizers. Even if productivity is quite high on the natural range it is often economic to plant pasturesand fodder crops in these areasand to practise ley farming. Seasonally flooded and permanent swamp grazings Where the grazing of these during the dry season can be combined with the grazing of upland range during the wet season a relatively stable managerial system can be established. There are often many nutritious and palatable perennial forage plants in these ecological associations and if they are properly managed the stocking rate can be quite high. However, uncontrolled grazing will eliminate the superior grasses in seasonally flooded areas. This is essentially what has happened in the llunos of Colombia and Venezuela.

118

hbtrition

and Feeding

Livestock management on natural grazings On unimproved ranges indigenous livestock are likely to thrive better than exotic. In the wet tropics Bs: (Bibos)-type cattle such as the IBali may be particularly valuable. In dry and semiarid areas Zebu or crossbred Zebu cattle are likely to thrive best. In the tsetsefly infested areas of Africa., trypanosomiasis-tolerant cattle such as the Ndama and the Dwarf Shorthorn must be used.In high grass regions cattle will thrive better than sheep,whereas on short grass ranges sheep will usually thrive better than cattle. In the arid areas goats, sheep and camels will thrive better than cattle. In seasonally flooded and swamp areas water buffalo probably thrive better than cattle. In the montane areas temperate-type livestock can be utilized, while on the highest altitude ranges the llamoids of South America possess special advantages. It is likely that on most range areas it is a desirable managerial practice to herd two or three different species at the same time. For example, in unstable ecotypes where bush easily replacesgrassesa mixed population of cattle and goats may be more satisfactory than grazing either goats or cattle alone. The value of goats under these circumstances has in the past beendemonstrated at Mpwapa in Tanzania (Staples et al., 1942).In very dry areasand particularly in those where free water is very scarcewild game may be more productive than domestic livestock (see Ch. 11, Pt II).
Choice oJ livestock.

Control of livestock by fencing or herding is as important on range as on planted pasture. Enclosure by fencing is usually more desirable than herding if it is economically viable, but herding is still the main method of controlling livestock on tropical ranges (Plate 3.2).The essential factor is that the livestock should be controlled so that they never graze out all the desirable species, i.e. overgraze. In general, short grasseswill withstand some degree of overgrazing better than the tall grasses.
Fencing and/or herding. Shade. This is desirable at certain times of the year for almost all types

of livestock. Under range conditions shade may or may not be available, depending upon the ecotype that is grazed. The control of water resources is an essential managerial tool on range. All man-made watering points should be adequately fenced and be subject to overall managerial control. The water requirements of range livestock have been given in a previous section.
Water.

In practice, stocking rates are often too low on range in the wet tropics and invariably too high on range in the dry tropics. As a consequence vast quantities of dry matter are burnt annually and therefore wasted in the wetter areas, while the vegetative cover of the
Stocking rates.

Management of forage

overstocked dry ranges steadily deteriorates both in quantity and in quality. It is now considered by some authorities that chronic overgrazing, resulting in radical changesin the vegetative cover CC range, may the so modify the local climate as to causea decreas:;: annual rainfall, thus in hastening further deterioration of the vegeta.tivecover of the range. What is certain is that badly degraded range recovers very slowly, even after overgrazing ceases. In the wet tropics stocking rates can be based on the year-round carrying capacity, but in the dry tropics, particularly within enclosed areas such as ranches, they must be based on the dry season carrying capacity. It would be expected that, other conditions being equal, the stocking rates could be somewhat higher under nomadic and transhumant managerial systemswhere the livestock essentially rotate over vast areas of range. In assessingthe livestock-carrying capacity of a grazing it is usual to express it in terms of the number of livestock units (LSUs) that can be carried by a unit area of the grazing for one year. The LSU denotes a standard liveweight of animal and this standard may vary from country to country, but it is usually within the limits of 300 to 500 kg (660 to 1,100lb). In order to be able to convert all speciesof livestock to the same common unit FAO (1974) have used the conversion factors detailed in Table 3.14. These conversion units are recommended for general use.
Table 3.14
Livestock unit (LSU) conversion factors Species Factor

Camel Buffaloes, horses, mules Cattle, asses Pigs Sheep, goats Note: The number of the species in-

volved are multiplied by the above factor to obtain the conversion factor, i.e. 100 sheep equal 100 x O-1 LSU or 10 LSU. Source: FAO (1974).

Within any species or type of livestock similar factors can be used to convert the average liveweight of all classes of stock to the common LSU. An example of such a conversion is given in Table 3.15. To obtain maximum productivity on range some form of supplementation of the feed supply during the dry season is usually essential.The form that this supplementation should take will depend on
Supplementation.

120

Nutrition

and Feeding

Table 3.15

AH example of the conversion of the average liveweights of diflerent classes of the same species to a common livestock unit (LSU) Baggara cattle in southerrr Ddrfur, Sudan Class of cattle
----___

LSU equivalent Males Females

Calves Weaners and yearlings 2 -2: years old 3&3$ years old 34-4 years old
Sourer: Hunting Technical Services (1974).

local ecological and economic factors. Supplementation is dealt with in some detail in the specific chapters concerned with the management of different types of livestock, in Part II of this text. Other aspects of the management of natural grazings Although the speciesthat are present are those that have of necessity to be utilized, overall management can influence the type of speciesgrazed by encouraging the desirable at the expense of the undesirable. Overgrazing must be avoided, as must the encroachment of undesirable bush species.
Forage species utilized.

In all except exceptional casesthe use of fertilizers on range forage is likely to be uneconomic. Phosphatic fertilizers may, however, be used under certain circumstances to encourage the growth of oversown legumes. Such managerial practices are most likely to be economic on wet tropical ranges.
Use offertilizers. Control of bush and weeds. Mechanical or c.hemical methods of control

are unlikely to be economic except in favoured areas. These are more likely to be in the wet tropics or in some derived savanna regions. In the drier tropics the only economic method 2 bush cnntrr,l is probably burning and, under special circumstances, biological control. Burning is a very controversial practice. It is condemned by most foresters, although controlled burning is at present used to clear undergrowth in the coniferous forests of the southern United States, and by the soil conservationists. Nevertheless, it is often the only economic method of controlling bush and weeds on range land. The major objection to bufiiing is that it wastes organic matter in the soil, particularly if the fire is very hot, and that this encourages erosion. It also kills young treesthat may be required for shade, destroys fences and encourages the proliferation of fire-resistant bush species that may

Management

offorage

121

not be suitable for browse. These objections may be overcome if the burning is conducted with intelligence and foresight. In the wet tropics the burn should always be made before the period of major growth. This ensuresthat the burnt area will quickly be covered with vegetation, thus minimizvng the risk of extensive erosion. Precautions should always be taken to see that proper fire-breaks are cut or burnt and fires should not be started when there is ,a high wind. In order to get a good burn, particularly in the dry tropics, the livestock should be removed from the area to be burnt many months before the burn, so that there will be a dense mat of vegetation that will carry the burn. Controlled and uncontrolled burning is practiscd in all tropical regions and the growth of some very desirable species can be encouraged by this practice. For example, in East Africa the dominance of the desirable red oat grass is strongly favoured by burning and in the Pacific islands grasses become dominant in Psidium guayana (guava)-infested hill pastures if they are closed and burnt for 2 or 3 successiveyears. If burning is to be successful it must be accompanied by an intelligent stocking policy. It is of little use to destroy one weed only to replace it by others. The most spectacular instance of the biological control of bush has been the control of Opuntia spp. (prickly pear) in Australia. In 1920 this plant covered some 24 million ha (60 million acres) of fertile soils. In 192627 the large-scale distribution of the insect Cactohlustis cactorwr. whose caterpillars live on the prickly pear, began and the insect established itself so rapidly that by the end of 1931the bulk of the-prickly pear had been destroyed. Mopping-up operations included the introduction of other insect pests and conventional control methods. The prickly pear was completely destroyed in 7 years. It is unlikely that this spectacular demonstration of biological control can easily be repeated elsewhere, but biological control methods might be used to reduce and weaken invading bush species.The whole subject must of course be approached with the greatest caution. Cultivated pastures and fodder crops Although it has been widely demonstrated that grazed forage is the cheapest feed for ruminant livestock and that natural grasslands are necessarily relatively low yielding while cultivated forage can yield more nutrients per unit area than any other crop, the concept of treating forage as a crop is, in general, still a novel idea in the tropics. This is understandable for the following reasons: a Cultivated forage is generally only practicable and economic in the equatorial, monsoonal and medium-altitude tropics. Without irrigation, the cultivation of forage in the drier tropics is difficult and yields do not usually justify necessaryinputs. Cultivation requires increased expenditure and the majority of

122

Nutrition

and Feeding

peasant farmers in the tropics cannot afford to invest in crops that do not provide a direct cash or s&sistence return. e In many regions where forage could be cultivated existing population pressure is intense and is increasing. The average size of holding is very small and often all available land must be used for the production of crops for direct human consumption. @ The cultivation and subsequent utilization of forage require a higher degree of managerial sophistication than the average peasant farmer at present possesses. @ Even if farmers wished to cultivate forage, in most tropical regions there is a lack of knowledge of the most suitable species to use and of satisfactory methods of establishment and management. e It is often pointless to improve forage productivity unless superior types of livestock can be made available to utilize the improved forage. They are not usually available and in addition the lack of organization of the livestock industry often mitigates against the economic marketing of any additional output of animal products. Establishment As the establishment of cultivated pastures is necessarily expensive it is important that it should be carried out as economically and efficiently as possible. Some of the major factors involved are as follows:
Choice of species. There are virtually no forage speciesthat grow equally

well in all tropical environments and even subtle differences in the environment can exert major influences on the growth of the forage species.Generally, pure grass stands can be made more productive than mixtures, but they are often less economic as they require more fertilizer and normally exhibit a more seasonal productivity. Choice is of course limited by the availability of seedand/or vegetative planting material. The introduction and testing of a large number of forage species are only now beginning in many tropical countries and the most suitable speciesmay not be everywhere available. If seed or vegetative planting material is freely available then it is important that the forage selectedshould be high yielding in the specific environment in which it is to be used. Palatability and nutritive value are not so important, as these factors depend primarily on the way in which the forage is managed. Species that produce viable seed are obviously to be preferred to speciesthat have to be propagated vegetatively. The way in which the forage is ultimately to be managed is also of some importance as particular species may be more suitable for cutting than for grazing or vice versa. Topography must also be taken into consideration, as on flat areas mechanization of harvesting may be easy and drainage difficult, while on steep slopes drainage may be easy and mechanization difficult. For grazing species,persistency is important, but if the speciesis used in a ley it must not be difficult to eradicate. In

Management @forage

123

the wet and dry or the semi-arid tropics species must be able to persist through the long dry seasons. A list of some of the grassspeciesthat may be used in planted pastures in different environments is provided in Table 3.16. Legumes should also be included in planted pastures as at the same stage of growth they normally possessa higher nutrient content than grasses,often possessa different growth cycle, add variety to the diet of grazing animals and. most important, they fix atmospheric nitrogen that eventually becomes available to other plants in the pasture association (Plates 3.3 an& 3.4). Some doubts have been expressed in the past as to the value and efficiency of tropical legumesin symbiotic nitrogen fixation. It is now believed, however, that the comparatively poor results obtained in the past have been due in part to the use of unsuitable speciesand rhizobium
Table 3.16 Some grass species thar may he used-for planted pastures in diflerent tropical
enr~ironmi rlts Environment Common name Species Pennisetum purpureum Notes

Humid tropics (this includes the monsoona .I areas of the tropics)

Elephan t or Napier Para

Very erect plant with a deep root system Also suitable for seasonally flooded land

Brachiaria

mutica

Signal In general, those grasses listed first wil I withstand higher rainfall Pangola conditions Guinea Jaragua Molasses African star or giant star

Brachiaria Brachiaria

hrizantha decumhens

Forms a dense sward Stoloniferous Many different varieties A rather coarse fodder Very suitable for the drier areas of the humid tropics A good hay grass Many different varieties

Digitaria decumhens Ischaemum aristatum Panicum maximum Hyparrhenia rufa Melinis minutiflora Cynodon plectostachyus

Dichanthium

caricosum

Digitaria pentzii Bermuda or Coastal Bermuda. etc.

-

Cynodon dactylon

Setaria splendida

Seasonally flooded regions

Aleman

Echinocloa polystachya Echinocloa pyramidalis Leersia hexandra Panicum repens

124

Nutrilian

ana Feeding

Table 3.16 - continued

Environment Common name Species

Notes A very great diversity of types Some varieties more stoloniferous than others A good hay grass Very drought resistant Tall and erect with strong rhizomes Mat-forming species Very drought reslstant A great diversity of forms

Dry tropics

Buffel Rhodes

Cenchrus ciliaris Chloris gayana

Makarikari Columbus Sabi - ~--~---~~ _- -. Montane tropics Nandi setaria Kik uyu Dallis Bahia Many temperatezone species

Andropogon gayanus Panicum coloratum Sorghum alnum Urochloa mosamhicensis Erugrostis curvula Maria sphacelata

Phaiaris tuherosa Pennisetum clandesrinum Paspalum dilatar rnn Paspalum nolatum

Norm: No attempt ilas been made to detail all the available and suitable species but only those that are well known and continuously used.

Plate 3.3

Jamaican Red Poll cattle grazing good enclosed pastures.

Management qf.forage

125

(guinea) Plate 3.4 Close-up of a productive grass/legume pasture: Parzicum m~~.~ini~ui~ and Ccntroserntl p~hc.sccns(centro).

(inoculant) strains. doubtful fertilizer practices and unsatisfactory management. It has been.conclusively demonstrated that suitable tropical legumes can fix considerable quantities of readily available nitrogen. Bryan (1962) stated that in grass-legume mixtures in tropical Australia. St~~losmtlws hr?jrr*i.Dcwwditm m%~tm and I~ldigc$w spimtu yielded 114. 180 irnd 264kg per ha (101, 161 and 235 lb per acre) of nitrogen per annum respectively. while Moore (1962) stated that in the humid. tropical region of Nigeria centro in a 2-year-old centro-African star grass mixture yielded 279 kg per ha (250 lb per acre) of nitrogen and that the nitrogen content of the African star grass was raised from 1.8 per cent in a pure stand to 2.4 per cent in the grass-legume mixture. Henzell et (11. (1966) have shown that in Australia a 4-year-old stand of Lwcu~r~cr l~~zrc~~~ph&yielded 578kg per ha (515 lb per acre) of nitrogen per annum while Townsville stylo yielded 109kg per ha (96 lb per acre). A

126

Nutrition

and Feeding

Table 3.17 Some legume species that may be usedfor planted pastures in d@erent tropical
environments Environment Common name Species Calopogonium muconoides hotes

I-Iumid tropics In general, those legumes listed first will withstand higher rainfall conditions and higher ambient temperatures

Calopo Centro

A trailing species; not very palatable A trailing species; palatable when young Wide distribution in natural pastures A sprawling plant A sprawling plant A trailing species; not very palatable Four major varieties in Australia A small tree; many different varieties Three major varieties in Australia Can also be used in montane areas A trailing species An annual species An annual species An annual species Does well on heavy soils; an annual A profuse seeder; annual An annual Normally used as a food crop Grows well in arid areas with irrigation and some light shade Grows well when adequate moisture is available

Centrosema pubescens

Desmodium heterophyllum

Silver-leaf desmodium Green-leaf desmodium Puero Sty10 Siratro

Desmodium uncinatum Desmodium intortum Pueraria phaseoloides Stylosanthes guyanensis Macroptilium atropurpureum Lucaena leucocephalg GIycine wightii Lotononis bainesii Dolichos axillaris Desmodium uniflora Phaseolus aureus Vigna sinensis

Seasonally flooded regions Dry tropics

Phasey bean Townsville sty10 Pigeon pea Lucerne or alfalfa Berseem

Phaseolus lathyroides

Stylosanthes humilis Stylosanthes hamida Dolichos lablab Cajanus cajan Medicago sativa

Trt$olium alexandrinunt Trifolium repens

Montane tropics

White clover Kenya wild white clover

Trifolium

semipilosum

---

Note: No attempt has been made to detail all the available and suitable species but only

those that are well known and commonly used.

Management ojforage

127

list of legumes that may be used in different environments is provided in Table 3.17. Development should proceed in accordance with a plan and the best land should be improved first, as the return on a given investment is likely to be higher on good rather than poor land.
Choice of land.

Cultivated pastures must be treated as a crop and the land prepared accordingly. For seededpastures the soil should be weed free and consolidated, but the tilth should not be too fine as this may cause packing of the soil once it rains. Where speciesare to be planted vegetatively the tilth can be rougher. There are areas of rain-forest, for example in the Amazon basin in South America, where the soils are so poor that after felling the forest and 1 or 2 years cropping the only possible economic crop is forage. Under these conditions grass and legume seedsshould be oversown in the first or second crop or vegetative material planted by hand between the tree stumps. Often the forest should never be felled in such regions.
Land preparation.

It is very important to obtain a good plant cover as quickly as possible. This will occur only if the soil moisture level remains high for several weeks after planting. For rain-fed areas planting should normally take place at the beginning of the rainy season.Where irrigation is available there may be advantage in planting in the middle of the dry seasonso that a very clean seedbedcan be obtained. In the wet tropics it can be advantageousto sow beneath a nurse crop. If the latter is maize the row spacing should be 1.8 to 2.1m (6 to 7 ft). There are normally no advantagesin planting beneath a nurse crop in the drier areas.
Time of planting.

When using vegetative material the nursery should be sited close to the planting area, and provision must be made for a sufficient quantity of planting material. It has been calcu!ated, for example, that O-5ha (1.2 acres) of guinea grass spaced at 50 x 50 cm (20 x 20 in) provides sufficient planting material for 12 to 15ha (30 to 37 acre) of land if each parent plant is split into four tufts. There are three major methods of vegetative planting:
Method of planting.

1. Using tufted grassessuch as guinea the crown of the parent plant can be split. These splits can be dibbled into the ground and consolidated by trampling. It is very difficult to mechanize this operation successfully. 2. In the case of tall, upright fodder grasses such as elephant and Guatemala, short lengths of cane, each with at least two nodes, can be pushed into the ground or the operation can be partially

128

Nutrition

and Feeding

mechanized by laying lengths of cane in furrows and covering them with soil in the same way as sugar cane is planted. 3. With stoloniferous or rhizomatous speciessuch as African star grass, para or pangola, cuttings can be distributed in plough furrows or spread on and disced into the soil. This is a very suitable method to employ where irrigation is available and the operation can be completely mechanized. If seed can be used and it is normally preferable to use seed, particularly in the drier areas, it may be either broadcast by hand or sown through a fertilizer distributor or a special seeddrill. Viability tests should be made with the seedbefore sowing. Seeding rates must be higher when the seed is broadcast than when it is drilled. Drilling should be shallow. Very hard legumeseedsmay require scarification or some other special treatment before sowing and they should always be treated with the correct inoculant. Management during the early stagesof growth This is essential in the early stages of establishment and is assisted by good land preparation prior to planting. Weedsmay be controlled by chemical methods: either pre-emergence sprays such as paraquat before the grass is planted or post-emergence sprays. The latter cannot be normally used where legumes have been planted. Chemical weed control is usually expensive. In general, an early mowing to prevent the first weeds flowering, followed by systematic hand or spot-chemical weeding, is likely to be the most economic form of weed control.
Weed control. Grazing control. Early grazing is almost always desirable where stoloniferous or rhizomatous speciesare used. Where tufted speciesare used in drier areas, too early grazing may result in the animals pulling out the young plants. In theequatorial and wetter monsoonal areas tufted speciescan be grazed quite soon after planting a!!:,qrazing encourages them to produce more leaf.

Farmers should understand something of the fertilizer requirements of their pastures and fodder crops and if a service is available soil samples from the fields should be taken and analysed. Potash, phosphate and lime, if considered necessary,should be applied before planting and nitrogen soon after planting. Phosphatic fertilization is often essentialfor the proper growth of legumesand in some regions additional sulphur may be required. For example, sulphur deficiencies have been reported in pastures in Uganda and in the Trans-Nzoia area of Kenya. Lime is not as essential for legume growth in the tropics as it is in the temperate zone. Heavy nitrogen applications may be economic on fodder crops but are unlikely to be economic on most planted pastures. As
Use offertilizers.

Management of forage

129

suggested in a previous paragraph the use of legumes in a pasture mixture should reduce the need for nitrogenous fertilizers. There is now some evidencethat certain tropical grassspecies,such as guinea, can also fix atmospheric nitrogen under certain specific conditions, so that the choice of grass speciesto be utilized may become of major importance. Management after full establishment The aim of the farmer, once he has establishedcultivated pastures, should be to obtain the maximum quantity and quality of feed from them, to reduce their seasonality of production, to maintain them in a productive state for as long a time as is possible and to ensure the efficient utilization of the forage obtained from them. Dry matter yield of forage per unit areadependsupon the speciesutilized, the amount of solar radiation, the availability of soil moisture and plant nutrients and the method of management.The choice of specieshas already been discussedin a previous paragraph. The amount of solar radiation dependsupon the total number of hours of sunshineand the cloud cover, etc. Total radiation per annum is almost always higher in the tropics than in the mid-latitude regions and it is usually much higher in the dry than in the :i;et tropics. Availability of soil moisture depends upon the total annual rainfall, its seasonality and repeatability and the type of soil. Soils with a high organic matter content normally hold water better than soils with a low organic matter content. In the drier tropics soil moisture content will be a limiting factor in pasture growth for a greater or lesser period during the year, but in the humid tropics it may limit growth for only relatively short periods. If irrigation is economically feasible,then availability of soil moisture ceasesto be a limiting factor. The availability of plant nutrients other than water depends on the inherent fertility of the soil and the rate at which nutrients in the soil becomeavailable to the plant. The latter dependson many factors, including the rate at which nutrients are leachedout of the soil and the quantities returned to the soil by the grazing animals. In general, tropical soils are not inherently very fertile, but there are exceptions. Grazed forages certainly require less fertilizer than continuously cut forages. The amount of nutrients removed by grazing animals depends upon their productivity. Approximately 80 per cent of the nitrogen, phosphate and potash consumed by animals are excreted, but much of the excreted nitrogen in the urine and faeces is lost in a tropical environment (Smith, 1965). Even if considerable quantities of nutrients are returned to the soil by grazing animals there is a problem of the poor distribution of excreta on the pasture. This can be improved by an occasional harrowing. Very large quantities of nutrients may be removed when pastures or
Production of the maximum quantity and quality of&wage.

130

Nutrition

and Feeding

fodders are continuously cut. Vicente-Chandler el al. (1964) have produced data (Table 3.18)from Puerto Rico that dramatically demonstrate this effect.
Table 3.18
The &feet of cuttingforage on the removal of nutrients from the soil

Yield of DM per annum (kg/ha (/h/acre))

Quantity of nutrients removed per annum (kg/ha (lb/acre)) Nitrogen Phosphate Potassium Calcium

-. Magnesium

_.

2r,423 (2 1,800)
Source: Vicente-Chandler

314 (281) et al. (1964)

52 (46)

408 (363)

119 (105)

79 (70)

The principal nutrient required by tropic31 grasses is nitrogen. The extent to which it is economic to utilize nitrogenous fertilizers depends upon many factors, not the least being the cost of the nitrogenous fertilizer. Very detailed experiments have been conducted by VicenteChandler et al (1964) on the response of various tropical grasses to nitrogen application in Puerto Rico. They found that at the time that the experiments were conducted molassesgrass gave an economic response when nitrogen was used at the rate of 224 kg per ha (200 lb per acre), while guinea, para and pangola gave economic responses2t.g rat& :-ii the 448 kg per ha (400 lb per acre) and elephant grass at the r-ate oi: 8% kg per ha (800 lb per acre). They also found that the greatest respot:yioccurred when grasses were cut at infrequent intervals, that there tias 1:~ advantage gained in splitting the nitrogen application into more than six parts to be applied in any one year and that the responsewas greatest in the flush growth period, thus increasing the seasonality of prcductr\oa. Irrigation in the drier areas improved the response of the graE3 to nitrogen application (Table 3.19). However, the application of nitrogen to grass-legume rillixturcs sup presseslegume growth. Grasses respond more rapidly than legumes to
Table 3.19
fertilized The eflect ofirrigation with nitrogen on the DM yield of guinea grass (Panicum maximum)

Total N per annum (kg/ha (B/acre))

DM yield of guinea grass per annum (kg/ha (lb/acre))

-

Without irrigation

With irrigation

225 (200) 447 (400)

Source: Vicente-Chandler

0 (0)

10,883 (9,710) 13,830 (12,346) 14,223 (12,690)

et ai. (1964).

18,168 (16,210) 31,664 (28,250) 42,290 (37,730)

Management of forage

131

the nitrogen and the legumes in the mixture suffer from the effect of shading and increasedcompetition for root space,soil moisture and other nutrients. It is possible that creeping legumes such as centro are not so badly affected as those with more upright growth habits. Under certain circumstances the application of phosphatic fertilizer is essential for the proper growth of legumes.Such action can also raise the phosphorus content of grasses with consequentadvantage to the grazing animals in phosphate-deficient areas. Vicente-Chandler et al. (1964) recommended the annual application of 74 kg per ha (65 lb per acre) on grassesthat are continuously cut and heavily fertilized with nitrogen in the wet tropics. On grazed pastures animal urine returns potash to the soil, but where grass is heavily fertiiized with nitrogen and continuously cut VicenteChandler et al. (1964) recommended applications o.potash. The response of tropical grass and legume species to applications of lime has been negligible, although lime?may be essential under specific circumstances. Whenever heavy fertilizer applications are used trace elements may also be required. The method of managementdependsto a very large extent on the way the forage is to be utilized. Species that are to be continuously grazed should be spacedcloser than speciesthat are to be cut, and tufted species require a closer spacing than rhizomatous or stoloniferous species.The manner in which the pasture is cut or grazed determines to a considerable extent the amount of leaf material present at any one time. Most tropical pasturesshould be managed so that their averageheight is somewhat above that of similar pastures in the temperate zone. They can be controlled, even when grazed, by the judicious use of the mower. This Lan prevent flowering, encourage tillering, remove coarse, unpalatable growth and help in the control of weeds. There is not only a problem of seasonality in the growth and total dry matter production but also of variation in nutritive value owing to the effect of the seasonon maturation of the species.Somemanagerial practices that may reduceseasonality in the quantity and quality of forage are as follows:
Reduction in the seasonality of production.

1. The well-planned application of fertilizers. These should not be applied during the flush seasonbut before the major growing season begins and perhaps again towards its end. 2. The conservation of excessforage during the flush season and the useof the conserved material at other times. This cannot be achieved without some loss of potential nutrients. This loss has been estimated to averageabout 25 per cent of the total conserved nutrients. Details of methods of conservation are given in a later section. 3. The implementation of seasonal livestock-breeding programmes so

132

Nirtrition

and Feeding

that the total animal demand for feed achieves a peak at approximately the same time as forage growth is maximal (Fig. 3.4). 4. The use of grass-legume mixtures rather than pure stands of grass. The peak growth of a grass-legume mixture is likely to be somewhat less than that of a pure stand of grass. Generally legumes grow and mature more slowly than grassesso thp.t ra)t only is the overall nutritive value of the grass-legume mixture superior to all except the youngest pure grassstands, but the nutritive value remains higher for a much longer period. Legumes also tend to be deeper rooted than

C&kg

1
BreLding

Weaning

I v

Wet season 4-

Dry season

Fig. 3.4 Schematic illustration of a managerial system that relates maximum herd demand for forage to the period of maximum forage production (----~---- total forage production per unit area; - - - - - - herd demand for forage).

grassesand continue to grow in the early stages of a dry season long after their companion grasseshave ceasedgrowth. 5. The use of pure stands of legume that could be grazed alternatively with grasses.Under certain environmental conditions this might be an alternative to normal mixed grazings. 6. The use of legume browse in drier areas. Many legume bushes and trees in the dry areas will produce high-quality browse during the dry season. The possibility of the integration of browse production into forage production systems in the dry tropics has not yet been fully investigated.

Management of forage

133

7. The intensification of weed control during the period of maximum growth. At this time frequent cutting for weed control could have a minimal effect on the growth of the desirable forage species. 8. The use of irrigation.
Maintenance of production.

This depends primarily on choice of suitable speciesand their subsequentmanagement with particular reference to the management of the animals that graze them.
ESJcient utilization of forage by livestock. This depends upon the
pro-

portion of the total forage grown that is consumed and the efficiency with which the consumed forage is converted into milk, meat or energy for work purposes in the animals body. Major factors that alIect the efficiency of utilization are as follows:
1. Type of livestock used. This factor has been discussed in a previous

section with referenceto the most suitable type of livestock for use on range grazings. Obviously it will depend upon the climatic environment, the type of agriculture practised, the relative demand for different animal products and the speciesand breedsavailable. In general, as nutrition should be generally superior on planted pasture to that on natural grazings, it should be possible to use more productive types of livestock on planted pasture. 2. Palutability of the foruge. Palatability and quality as expressed by digestibility and nutritional value depend upon the speciesused, the climatic environment and the method and age of utilization. Young forage is always more palatable than older forage, with a higher crude protein and mineral content and a lower crude fibre content. Young forage is not only more palatable, but because its fibre content is relatively low, consumption of large q-uantitiesproduces lessof a heat load on the animal, with the consequencethat the animal can increase its intake. Thus livestock eat more feed when it is highly palatable even in a tropical climatic environment. However, the availability of forage may determine the level of intake as the total quantity available per animal decreasesas the stocking rate increases. At any one grazing it is essential that livestock should eat a high proportion of the forage available so that there will be a minimum quantity of mature forage in the regrowth, as increasing quantities of mature forage reduce the quality of the forage available for the next grazing. In order to ensure that livestock do eat a high proportion of the available forage, the stocking rate must be high and the livestock must graze closely and evenly. However, under thesemanagerial conditions individual animal intake may decline. Therefore if grazing intensity is increased,production per unit area will increase for a time because of the increase in the stocking rate (Fig. 3.5) but the increase in total output is ulti-

134

Nutrition

and Feeding

m Increasing stocking rate per unit area Fig. 3.5 Diagrammatic illustration of the relationship between stocking rate, individual LWG per head; -----animal productivity and total productivity per unit area (--LWG per unit area).

mately achieved at the expense of a decrease in output of the individual animal. This inherent contradiction in practice is a major problem in assuring the most effective utilization of forage in any climatic environment. 3. Methods of management of the livestock. These are discussed in detail in the chapters concerned with specific types of livestock in Part II.

Forage conservation
The unavoidable seasonalnature of range and permanent pasture production means that at one season of the year there is a surplus of feed while at another there is a deficiency. Unless supplementary feed can be fed, or some animals can be removed from the grazings, the stocking rate must be geared to the season of low pasture productivity, and in the seasonof high pasture productivity a great deal of feed is wasted as there is an insufficient number of animals to consume all the forage available. It is recognized that there must be a wide fluctuation in the yield of forage from seasonto seasonin the dry tropics, but it might be thought that in the wet tropics, where growth is continuous, there would be little seasonalfluctuation. Unfortunately, this is not necessarilythe case,as will

Forage conservation

135

Tropics 5% Temperate Zone

27% 18% 15,400 (13,700) 9.100 (25,900)

kg (lb) dry matter per ha (acre) per annum Fig. 3.6 A comparison of seasonal forage production at localities in the tropical and temperate climatic zones (kg(lb) dry matter per ha (acre) per annum).

be seen from Fig. 3.6. At Sigatoka in Fiji, where the average annual rainfall is 1,780 to 2,030 mm (70 to 80 in) and is quite well distributed throughout the year, the seasonalvariation in growth of pasture is considerable and is as large as it is at Palmerston North in New Zealand, where forage conservation is a normal practice. This seasonalfluctuation in the production of forage is a major factor limiting animal production on range and permanent pasture. All over the world millions of tons of dried-out and unused potential animal feed are burnt annually after the flush season.It is obvious that if part of this surplus growth of forage could be preserved for the season when there is insufficient feed, then the annual carrying capacity of range and permanent pasture could be effectively improved. The preservation of surplus forage is a normal feature of temperatezone management of range and permanent pasture, but it is hardly practised in the tropics. Forage may be preserved as hay, silage or as an artificially dried product everywhere in the tropics, or as standing hay in the arid tropics. However, a word of warning is necessary.Forage conservation is an intensive husbandry practice that yields less return on investment than some other improvement practices. It can be argued that only feed that is truly surplus should be conserved. If the grazing area of animals has to be restricted in order to save forage for conservation then the depression in their growth rate during the period of restricted grazing may not be compensatedfor by an increased growth rate during the period when the conserved material is fed, as there is an inevitable loss of nutrients during the period of storage.This effect has been demonstrated by Smith (1967) in Queensland and some details of his investigations are shown in Table 3.20. Another factor that must also be considered under extensive grazing conditions is that losses in liveweight of meat-producing animals during the dry seasonwill to some extent be made up by compensatory liveweight gain during the wet season.This effect appears to

136

Nutrition

and Feeding olforage on liveuteight production of cattle

Table 3.20

IZflecl ofconservution

kg/ha (lb/acre) liveweight produced per annum

Stocking rate No conservation Conservation and hay feeding

0,s ha (1.24 acre) per steer 590 (530) 580 (520)

0.3 ha (O-74 acre) per steer 760 (680) 670 (600)

Nores: 1. Reduced growth occurred before feeding-back of the conserved material took place. 2. The pasture was predominant!y a Sorghum a/mum -~luccrne mixture and the hay that was conserved was predominantly lucerne with a 60 per cent digestibility. Sowce: Smith, C. A. (1967).

be due to the fact that animals, and particularly older animals that have been on a low plane of nutrition, eat far more forage when they suddenly find themselveson a high plane of nutrition than animals that have always been raised on a high plane. This phenomenon is further discussed in Chapter 5 (Part II). Under extensivegrazing conditions it could be expedient to conserve some forage for drought or other emergency feeding of breeding stock, but generally forage conservation is a practice that should be used only where the production of animal product per unit area is high and where increased production will justify the necessaryincreased expenditure, as in dairy farming or in baby beef or fat lamb production. It will usually be more economic under extensivegrazing conditions to attempt to solve the problem of seasonalforage production in other ways : perhaps by using somespecieswhose growth cycle is out of step with the majority of the speciesused or specieswhich grow for a longer period into the dry season; perhaps by utilizing high fertilizer inputs or by fodder cropping on a limited but improved area of the extensive grazing. Hay Hay is generally considered a roughage feed, but forage used for hay should be cut before the plants flower in order to obtain the most nutritious and palatable product. If, as so often happens, the plants are allowed to Mly mature and wilt, the hay may still be useful for the feeding of beef cattle if it is properly supplemented, for instance, with urea. In the dry tropics it is preferable under these conditions to leave it standing in the field and not to cut and store it. Hay of medium quality may also be useful for feeding as a supplement in the middle of the wet seasonwhen the dry matter intake of grazing animals may be low, owing to the very high moisture content of forage. Hay can be made in severalways and the choice of method will depend upon local circumstances.The usual method is to cut it in the field, let it

Forage conservation

137

dry in the row and to stack it as soon as it is dry. In the semi-arid tropics it may be possible to make hay very rapidly, but in the wet tropics hay can easily be spoiled in the field as it lies in the row. Even in the semi-arid tropics hay may mould in the field as it usually has to be made at the end of the growing seasonwhile the humidity is still high. If there is sufficient labour available the forage can be cut and stacked on tripods in the field. This allows it to dry out without spoiling in a wet climate, and hay produced by this method is usually of high quality. Certain new methods,such as crushing of the stalks while cutting or fluffing in the row, may assist rapid drying out in the field and might be useful practices in the tropics. In any event, the cut forage should not be allowed to become sunbleachedbut shou!d be gathered as soon as it is dry and while it still retains an appreciable amount of colour. Nor should it be exposed unnecessarily to the leaching effect of tropical showers. When it is stacked, or even before it is carted, it may be quite seriously attacked by termites unless it is raised above ground surface. Despite the fact that it is possible to make satisfactory hay at certain seasonsin some tropical countries and there are tropical grassessuch as Dichanthium curicosum (Antigua hay grass)that are very suitable for haymaking, very little hay is at present made outside the American tropics. There are many reasons for this situation: the generally small size of farms, the lack of suitable machinery, etc., and perhaps most important the fact that even if good hay is harvested it is almost certain to mould during storage in the wet tropics. Recently, new types of hay-collection machinery have been introduced in temperate countries. These roll the hay into large bundles or gather it into small stacks that can be mechanically handled. Such machinery would be economic on only the largest farms in the tropics. Silage Almost any forage can be ensiled, this being a simple method of preservation. There is always somewastage, but good management will reduce it to a minimum. Forage is ensiled by compacting it in some form of container from which air is excluded. Plant respiration goes on for some time after the silo is filled and the bacteria, yeasts,moulds and enzymes present initiate a fermentation process. The temperature inside the silo rises and acids are releasedby the fermentation. As the acidity of the silage rises, further fermentation is prevented and the great bulk of the silage is preserved. The quality of silage depends on the following factors: the stage of maturity and chemical composition of the crop ensiled; the ratio of soluble carbohydrates to the mineral base content of the crop.; the percentage of moisture in the crop; the rapidity and comp!eteness with which air is excluded during the filling of the silo; and the extent to which the temperature of the forage rises.

138

Nutrition

and Feeding

Forage of almost any quality may be ens-led, but it should be remembered that to ensile low-quality roughage is hardly likely to be economic, and that the nutrient quality of the silage will always be somewh,at less than the nutrient q-uality of the forage that has been ensiled, as even with the best management there will be some loss of nutrients, Unripe sorghum and maize make excellent silage, but many tropical grassespossess relatively low sugar content and they should be ensiled a using molassesat the rate of 3.5 to 4 per cent of the green weight of the forage. There are many other types of additives that may be used. These include : 1. Those that are a substitute for fermentation such as the mixture of suiphuric and hydrochloric acids used in the well-known AIV method and formic acid; 2. those that enhance fermentation such as enzymes, bacterial cultures and antioxidants; and 3. those that add nutrients such as the molasses referred to above. As it is unlikely that the majority of farmers in the tropics will have access to acids or special fermentation products the most practical method that they can normally utilize is the addition of nutrients. A silo should not be filled while it is raining. Succulent fodder should be wilted in the field before fi!ling into the silo, but water may have to be added if the forage is too dry. It is essential that the forage should be well compacted and covered so that air is excluded as much as possible. This means that with the exception of very young, short grass, forage must be chopped in order to ensure adequate compaction and exclusion of air. The container may be a structure built above the ground such as the tower silos so common in America and parts of Europe or a silage clamp or bunker or it can be a trench or pit in the ground. Some modem tower silos are glass lined and are both efficient and expensive. Sheet plastic is now widely used to cover silage made in bunkers, trenches and pits. Silage can also be stored very successfully in large, evacuated plastic bags and this would appear to be a method that might be used by small farmers in the tropics as the only equipment that is needed to create a vacuum in the silage bag is a small pump. There are difficulties, however, as it has been found that termites attack and puncture the bags. It is not necessaryto have any container at all as the silage can be stacked, but the wastage in a stack silo is very considerable. On large mechanized farms forage is usually cut with a forage harvester that loads the chopped fodder on to a trailer. On smaller farms the forage may be cut with a mower and transported to the silo on a buck-rake mounted on a small tractor. At the silo the forage can be chopped using a portable chaffing machine. On the larger farms it is becoming custom.ary to ensile forage in above-ground, open-ended pits or bunkers. With this type of silo the tractors can run over the silage to

Forage conservation

139

help consolidation and the silage can be mechanically loaded when it is required for feeding. Sifage bunkers or trenches can also be constructed so that animals can feed themselves.At a self-feedingsilo the larger types of cattle can eat up to a height of 2.1 m (7 ft). A barrier such as a wooden fence is needed to prevent burrowing. A feeding width of 17 to 22 cm (7 to 9 in) for each adult animal is required unless feeding time is restricted, when a width of up to 76 cm (30 in) is needed. The making of siiage is equally important but more difficult for the small peasantfarmer possessing few head of stock. A silage pit or trench a is the most easily constructed and most economic form of container, but it can be used only in the dry tropics or at a very well-drained site in the wet tropics. In the wet tropics experience has shown that the silage is usually spoilt by water seepage.The pit or trench must be of such a size that to obtain sufficient feed for eachdays requirements the exposed surface must be removed only to a depth of a few inches, otherwise much silage becomesmouldy and is wasted. The most difficult operation for the smafl peasantfarmer is the chaffing of the forage during the filling of the silo and this operation is usually essential if efficient compaction is to be ensured, particularly in the drier tropics. This difficulty might be overcome by the cooperative use of small power chaffers. Supplementary fodder crops such as maize, sorghum or millet are particularly suitable for silage making as are many tall tropical fodder grasses such as elephant, guinea, jaragua and Sudan *grass (Sorghum
sudanense).

Despite the fact that good silage can be matie in both the wet and the dry tropics and a major problem of all livestock farmers is the seasonality of the forage supply, little progress has been made in the tropics during the last decade in the development of the concept of conserving surplus forage as silage. Artificially dried forage Artificially dried forage is produced by placing forage on some form of drying tray and passing through it a blast of hot air until it is sufficiently dry to store. It may then be baled or milled and the fine milled material may be pelleted before storage. Large forage-drying plants operate on a continuous flow, rather than on a batch system. Machines that harvest, dry and pellet forage in one operation have recently been developed in the United States. Capital requirements are obviously high, but very considerable improvements have been made in forage-drying machinery during the last decade, and where high-quality forage can be cheaply produced and energy is not too expensivethe production of artificially dried forage in the tropics for specialist markets could now be economic. The continuous growth of the livestock feed industry and the development of industrialized pig and poultry industries in some tropical countries has

140

Nutrition

and Feeding

created a continuously expanding demand for dried leaf meal. In addition, if prices can be made competitive, there is the possibility of develop ing an export market for dried leaf meal in some humid tropical countries. There are at present some organizations evaluating the economics of producing artificially dried forage in the wet tropics, including a private enterprise group in west Malaysia. Severa!tropical forage speciescould be dried to produce leaf meal comparable in quality with lucerne meal, one of these being Leucaena 1eucocephaZa.It has also been suggested in the Caribbean region that milled, dried young elephant grass with a CP content of 10 to 12 per cent could be used to absorb a mixture of molasses,urea and minerals ?o produce an economic dairy concentrate feed. A feed of this type could either be produced at existing sugar mills or in association with sugar mills.

Nutritive value of forage

The nutritive value of a forage depends upon its composition, digestibility and intake, or the total amount that any animal will ingest within a specific period. Total energy intake of the animal ingesting forage is undoubtedly the most important factor determining total output of animal product, as long as protein and mineral requirements are met and the forage contains no toxic substances. The CP content of forage, though important and a useful indicator of quality, is not of such major significance in ruminant nutrition as was once alleged, as ruminants can utilize non-protein nitrogen as well as protein nitrogen. It is generally accepted that the minimum percentage values of total nitrogen in forage ingested by cattle should be 1.6, 1=6 and 1.9 for maintenance, beef and dairy production, respectively. If the mineral content of the forage is low, minerals can be provided in the form of a supplement. Indeed, this is the normal practice. It should also be remembered that the mineral content of legumes, herbs and browse trees is uswily somewhat higher than that of grasses grown in the same environment. An exception is Townsville stylo. This legume generally possesses low phosphorus content. a Toxic substances Severalwell-known tropical forage plants contain toxic substances.Some of these are: Leucaena leucocephala, a small legume tree that contains mimosine, a substance inhibiting cell division; the legume Indigqfera spicata that produces a dangerous hepatoxin known as indospicine; and speciesof the grassgenus Setaria that possesshigh oxalate contents. As a consequenceintroduced forage plants should always be screenedby the

Nutritive

value oj,firage

141

appropriate authority for toxic substancesbefore they are released for general farm use. Intake and digestibibty The digestibility of forage can be expressed in terms of dry matter, organic matter or energy, or of a single component such as CP. The determination of the digestibility of feedshas already been discussedin a previous section. The digestibility of forage dependsabove all on its stage of maturity, as will be seenfrom Table 3.21.
Table 3.21
The eflect of variety and stage of maturity on the digestibility DM intake of guinea (Panicum maximum) forage in Australia No. qf days crf regrowth Digestibility (%I Green panic of DM Voluntary DM intake k/k W ) Green panic and voluntary

Coloniao 62.2 53.4 50-2

CoIoniao 69.2 47.6 42.3

28 63 91

62-O 47.4 47-7

69.2 47.6 42-3

!iotes: 1. Here, DM intake is expressed in terms of the metabolic size of the animal, i.e.

g/kg body weight. 2. Green panic and coloniao are two varieties of guinea grass (Panicum maximum). Source: Minson, D. J. (1971).

The voluntary intake of forage by any class of ruminant livestock under specific environmental conditions is greatly influenced by the type of forage ingested and by its stage of maturity. As will be seen from the data in Table 3.21,voluntary intake varies considerably even between varieties of the same forage plant and declines rapidly as the plant ages. In fact the variation in voluntary intake due to varietal differences is much larger than the data shown in Table 3.21 suggest,as the intake of six varieties of guinea grass by individual animals varied from 35.7 to (Minson, 1971). 80.8 g per kg W oD75 Proximate analysis and the determination of the SE and TDN values of tropical forage are said to underestimate their feeding value and it is probable that the approximate nutritive value and digestibility of forage can most easily be determined in the laboratory by measuring the total nitrogen content as an index of quality and the in vitro digestibility. Voluntary intake and digestibility can be determined using live animals either in an animal house or out on grazing. The advantage of using the grazing animal is that measurements are taken on the forage that the animal actually selects and eats. Some details as to how digestibility is determined under such circumstances have been given in a previous section. A further advantage of determinations made on the grazing

142

Nutrition

and Feeding

animal is that pasture measurements,such as the amount and quality of the forage available and the persistenceand botanical composition of the pasture, can also be measured. Those requiring further information on the various techniques mentioned above should consult specialized papers and textbooks concerned with this subject. Effect of climate on nutritive value The most important climatic factors that limit forage growth, and hence the quantity of feed available, are temperature, effective precipitation, length of daylight and the intensity of solar radiation. The quality of feed depends mainly on effective precipitation and on the intensity of solar radiation. The very real differences in climate that exist between the humid and the semi-arid tropical areasthus present two broadly distinct nutritional problems. These have already been discussed in Chapter 1, Part I. Nutritive values It is not possible in an introductory text to present details of the nutritive value of all the different tropical forage species.Fortunately a considerable volume of data has now been collected, collated and analysed,and readerswho are interested are advised to consult publications by Morrison (1957) Crampton and Harris (1969),McDowell et al. (1974) and Gohl (1975).

References
Agricultural Research Council (1963) The Nutrient Requirements of Farm Livestock. (1) Poultry. Summary of Recommendations. ARC: London. Agricultural Research Council (1965) The Nutrient Requirements of Farm Livestock. (2) Ruminants. Technical Reviews and Summaries. ARC: London. Agricultural Research Council (1967) The Nutrient Requirements of Farm Livestock. (3) Pigs. Technical Reviews and Summaries. ARC: London. Armstrong, D. G. and Mitchell, H. H. (1955) Protein nutrition and the utihsation of dietary protein at different levels of intake by growing swine. J. Anim. Sci., 14,49-53. Baudelaire, J. P. (1972) Water for livestock in semi-arid zones. World Anim. Rev. (3), 1-9. Beard, J. S. (1944) Climax vegetation in tropical America. Ecology, 25, 127-58. Bender, F., Heany, D. P. and Bowden, A. (1970) Potential of steamed wood as a feed for ruminants. Forest Prod. J., 20, 36-41. Blaxter, K. L. (1962) The Energy Metabolism of Ruminants. Hutchinson : London. Blydenstein, J. (1967) Tropical savanna vegetation of the llunos of Colombia. Ecology.
48, l-15.

Bonsma, J. C. (1940) The influence of climatological factors on cattle: observations on cattle in the tropical regions. Fmg. S. AI;.. 15, 373-85.

References

143

Bryan, W. W. (1962) The role of tAe legume in legume/grass pastures. Bull. Comw. But-. Past. Fld. Crops, 46, 14760. Commonwealth Agricultural Bureaux (1947) The use and misuse of trees and shrubs as fodder. COWJ. Agric. Bur. Joint Pub., No. 10. CAB: Farnham Royal, U.K. Crampton, E. W. and Harris, L. EL(1969) United States-Cunadian Tables of Feed Composition. NAS-NRC, Pub. 1084. NRC: Washington. Cuthbertsom, Sir D. P. (1969) The Science of Nutrition of Farm Livestock, in Cuthbertson, Sir D. P. (ed.), Nutrition of Animals of Agricultural Importance. Part I. Int. cncylp. Food Nutrit., Vol. 17. Pergamon: Oxford. Dahadghao, P. M. and Shankarnarayan, K. A. (1972) The Grass Cover of India. Indian Count. Agric. Res. Sci. Mono. ICAR: New Delhi. Davies, W. (1960) Temperate and tropical grasslands. Proc. 8th Int. Grussld. Congr.. UK. Denton, D. A., Goding. J. R., Sahine, R. and Wright, R. D. (1961) Salinity problems in the arid zones. Proc. Teheran Symp. Arid Zone Res., 14. UNESCO: Paris. Devendra, C. (1971) The comparative efficiency of feed utilisation of ruminants in the tropics. Trop. Sci., 13, 123-32. Evans, R. E. (1960) Rations for livestock. Min. Agric. Fish. Food Bull., No. 48. HMSO: London. FAO ( 1956) Prodtiction Yearbook. FAO : Rome. FAO (1974) Production-Yearbook, Vol. 27, p. 447. FAO: Rome. French, M. H. (1945) Geophagia in animals. E. Afric. med. J., 22, 103-10. French, M. H. (1956) The importance of water in the management of cattle. E. Afric. ugric. J.. 21, 171-81. Gohll, B. (1975) Tropical Feeds. FAO : Rome. Heany, D. P. and Bender, F. (1970) The feeding value of steamed aspen for sheep. Forest
Prod. J., 20, 98-102.

Denzell, E. F., Fergus, I. F. and Martin, A. E. (1966) Accumulation of soil nitrogen and carbon under a Desmodium uncirtatum pasture. Aust. J. exp. Agric. Anim. Hush.. 6, 157-60. Hunting Technical Services (1974) Southern But-fur Lund-use Planning Survey. Annex 3. Animal Resources and Range Ecology. Hunting Technical Services: Boreham Wood, UK. Kehar, N. D. (1954) Indian Coun. Agric. Res. Silver Jubilee Souvenir. Govt. India Press: New Delhi. Ledger, H. P. (1959) A possible explanation for part of the difference in heat tolerance exhibited by Bos tuurus and Bos indicus beef cattle. Nature (Land.), 184, 1405. Lucas, I. A. M. (1972) The use of antibiotics as feed additives for farm animals. Proc.
Nutrit. Sot.. 31, 1-8.

McDonald, P., Edwards, R. A. and Grcenhalgh, J. F. D. (1973) Animal Nutrition (2nd edn). Longman : London. McDowell, W. R., Conrad, J. H., Thomas, J. E. and Harris, L. E. (1974) Latin American Tables of Feed Composition. Univ. Florida: Gainesville, Fla. Meiklejohn, J. (1962) Microbiology of the nitrogen cycle in some Ghana soils. Emp. J.
exp. Agric., 30, 115-26.

Minson, D. J. (1971) The digestibility and voluntary intake of six varieties of Punicum.
Aust. J. ugric. Anim. Hush., 11, 18-25.

Moore, A. W. (1962) The influence of a legume on soil fertility under a grazed tropical pasture. Emp. J. exp. Agric., 30, 239-48. Moore, R. M. (ed.) (1970) Australian Grasslands. Aust. Nat. Univ. Press: Canberra. Morrison, F. B. (1957) Feeds and Feeding (22nd edn). Morrison Publishing: Ithaca, NY. Nditional Academy of Sciences-National Research Council (1966) Nutrient Requirements of13airy Cattle (3rd rev. edn). NAC-NRC, Pub. No. 1349. NAC-NRC: Washington. National Academy of Sciences (1968a) Nutrient Requirements of Sheep (4th rev. edn). NAS, Pub. No. 1693. NAS: Washington. National Academy of Sciences (1968b) Nutrient Requirements of Swine (6th rev. edn). NAS, Pub. No. 1599. NAS: Washington.

14

Nutrition

and Feeding

National Academy of Sciences (1970) Nutrierzt Requirements qf Beef Cattle (4th rev. edn). NAS:Washington. National Academy of Sciences (1971) Nutrient Requirements of Poultry (6th rev. edn). r AS: Washingion. , Payne, W. J. A. (1965) Specific problems of semi-arid environments. Qzcalitas PI. Muter. veg.. 12, 268-94. Payne, W. J. A. (1969) Problems of the Nutrition of Ruminants in the Tropics, in Cuthbertson, Sir D. P. (ed.), Nzctrition of Animals of Agricz~lturul Zmportunce. Part 2. Znt. Encylp Food Nutrit., Voi. 17. Pergamon: Oxford. Pirie, N. W. (1969) The production and use of leaf protein. Proc. Nutrit. Sot., 28,85-9 1. Rattray, J. M. (1960) The Grass Cover of A,fricu. FAO Agric. Studies, No. 49. FAO: Rome. Roseveare, G. M. (1948) The Grasslands of Latin America. Comw. Bzo-. Past. Field Crops Bull., No. 36. Comw. Agric. Bur.: Farnham Royal, UK. Schmidt-Nielsen, K. (1964) Desert Animuls. Physiological Prohlems of Heat and Water. Ciarendon Press : Oxford. Simoons, F. J. and Simoons, E. S. (1968) A Ceremonial 0.x qf Zndiu. Univ. Wisconsin Press : Madison. Smith, C. A. (1965) Studies on the Hypurrheniu Veld. VI. The fertiliser value of cattle excreta. J. ugric. Sri. (Cumh.), 64, 403-6. Smith, C. A. (1967) Brigalow country. Rep. Div. Trop. Past.. CSIRO 1966j67, pp. 25-34. CSIRO: Australia. Staples, R. R., Homby, H. E. and Homby, R. M. (1942) A study of the comparative effects of goats and cattle on a mixed grass-bush pasture. E. Afi. ugric. J.. 8. 62-70. Suter, H. E. (1962) Carence en cobalt dans un tlevage des bovides au Katanga. Re:. Elevuge Med. Vtt. Pays Trop., 15, 31-41. Abstracted: Nutrit. Abst. Rev., 33, 1818. Vicente-Chandler, J., Cam-Costas, R., Pearson, R. W., Abruiia, F., Figarella, J. and Silva, S. (1964) The intensive management of tropical forages in Puerto Rico. Unir. Puerto Rico ugric. Exp. Stn. Bull., NO. 187. Whyte, R. 0. (1964) The Grassland and Fodder Resources of India (2nd edn). Indian Count. Agric. Res.. Mono. No. 22. ICTA; New Delhi. Whyte, R. 0. (1968) Grasslands of the Monsoon. Faber and Faber: London. Wbyte, R. 0. (1974) Tropical Grazing La&s. Dr W. Junk: The Hague, Whyte, R. O., Moir, T. R. G. and Cooper, J. P. (1959) Grasses in Agriculrnre. FAO Agric. Studies. No. 42. FAO: Rome. Whyte, R. O., Nilsson-Leissner, G. and Trumble, H. C. (1953) Legzzmes in Agricultzrre. FAO Agric. Studies No. 21. FAO: Rome. Wilcke, H. L. (1966) Feedsty&, 38 (5).

Further Reading
Davies, J. G. Pasture improvement in the tropics. Proc. Ninth Znt. Grassld. Congr.,
Brazil (1965).

Davies, W. and Skidmore, C. L. (eds.). Tropical Pastures. Faber and Faber: London, 1966. Spedding, C. R. W. Grassland Ecology. Oxford Univ. Press: London, 1971. Whyte, R. 0. The myth of tropical grasslands. Trop. Agric. (?rin.), 39, l-l 1 (1962).

Chapter 4

eproduction and Breeding*

Although someof the very first animal-breeding practices must have been initiated when Neolithic man domesticated animals. the major part of modern animal-breeding practice has developed in the temperate zone during the last two centuries. In some temperate-zone countries. particularly in northwest Europe, the overall environment for livestock had been improved to such an extent by the beginning of the nineteenth century that attention could be directed towards the breeding of improved types of livestock that could take economic advantage of this situation. In addition, in northwest Europe the most adverse effects of climate on livestock could be avoided by the provision of adequate housing. As a consequence,during the nineteenth century spectacular advances were made in livestock breeding, particularly in the United Kingdom. However, theseadvances, although basedon acute observation and limited records, were still very much the result of trial and error, as nothing was known of modern genetic theory. The scienceof genetics, based on Mendelian theories of inheritance, began to gain general recognition at the beginning of the twentieth century, and during the lhst 50 years the application of genetic principles, combined with new knowledge gained on the physiology of reproduction, has revolutionized animal-breeding practices. These practices are slowly being applied in tropical countries. In the latter, few of the .livestock-owning peoples have in the past developed rules or customs concerned with the better breeding of their animals. The majority have allowed their livestock to breed more or less hap hazardly and the results have been accepted as a matter of fate and therefore inevitable. Nevertheless.a number of factors have contributed to the evolution of a multiplicity of different breeds of tropical livestock. In the first place the original domestication of most species probably occurred in the tropics and/or sub-tropics. For example, there are three major types of domestic cattle in the tropics, i.e. Bos taurus, B. irzdicus and B. (Bihos), together with their crossbreds, while there is only one
* Dr R. S. Temple, Director: Animal Science, International Livestock Centrc for Africa, Addis Ababa, Ethiopia, assisted in the writing of this chapter.

145

146

Reproduction

and Breeding

major type of cattle in the temperate zone, i.e. B. taurus. Secondly, the large-scalemovement of some livestock-owning peopleswithin the EuroAsian-African land mass, followed by periods of isolation due to intermittent warfare and/or the natural hazards of communication of small groups of people in large land areas, meant that livestock were moved into isolated areas where they naturally developed over a long period characteristics that fitted them to their new environment. Some peoples probably also bred their livestock for specific features, and originally this action may have had magico-religious connotations. For example, the B. taurus breeds of Hamitic Longhorn cattle that were first taken by migratory people from western Asia to the tsetse-fly infested forest regions of West Africa, perhaps some 5,000 years ago, have developed a tolerance of trypanosomiasis that is certainly not possessedby any other major cattle breed originating from western Asia. Not only have both the Ndama and Dwarf Shorthorn breeds developed this tolerance, but the tolerance of the Ndama cattle, which were the first to arrive in West Africa, is more developed than that of the Dwarf Shorthorn cattle. On the other hand, there must have been deliberate selection by man for very large horns within cattle of the Ankole breed in East Africa. Efforts to improve the productivity of livestock in the tropics have moved in a cyclic manner ever since the sixteenth century. At that time, when Europeans first discovered and then colonized the Americas, they imported their unimproved domestic livestock with them. These livestock thrived in the temperate but not in the tropical regions of the Americas. However, some of the breeds in the tropical regions did gradually acclimatize. One example is the Criollo cattle of tropical Central and South America. However, the parallel importation of European livestock into Africa and Asia was almost totally unsuccessful except in some favoured tropical islands such as Mauritius. Later, with the advent of a large-scale slave trade between Africa and the Americas some African livestock such as Ndama cattle and Dwarf Forest sheepand goats were imported into the Americas. These livestock were crossed with imported temperate-type livestock and form the basis for some modem American breedssuch as the Nelthropp cattle and the Black Bellied Barbados sheep of the West Indies and several sheepbreeds in northeast Brazil. In the eighteenth and nineteenth centuries there were also considerable importations of tropical livestock from Asia into Europe, an important example being the Siamese and Chinese pigs which have contributed characteristics to modem British pig breeds (seeCh. 12, Pt II). During the latter part of the nineteenth century, as it began to be realized that many of the original importations of temperate-type livestock into the tropics were not flourishing there was a considerable importation of Asian cattle into tropical and sub-tropical America. Improvements in the breeding of European and North American livestock, with the realization that the productivity of the new breeds far

The reproductive cycle

147

surpassedthe productivity of any indigenous tropical breeds, led at the end of the nineteenth and at the beginning of the twentieth century to a new cycle of importation of temperate-type livestock into the tropics. The failure of these importations was immediately apparent as by this time governments were monitoring importations and beginning to keep records. It was ultimately realized that until such time as epizootic diseasewas controlled and nutrition and management improved the possibilities for the successfulintroduction of highly productive temperatetype livestock into the tropics were poor. A major effort was then made to identify tropical breeds and to initiate selection within them. The result of these efforts has been disappointing and another major effort is now being made to introduce productive, temperate-type livestock into the tropics. The possibilities for successare now much greater than they were in the past, as we possess more knowledge of the characteristics of tropical and temperate-type livestock and we are better able to control and improve the tropical environment. In order to promote a rapid improvement in the productivity of tropical livestock we must first understand the basic facts of reproduction and the inheritance of characteristics and be able to apply suitable animal-breeding practices that have been developed in the temperate zone. The development of the livestock industry in the tropics depends upon improvements on a broad front, but it is also important that development priorities should be established. In areas where epizootic disease is still prevalent it is obvious that disease control must be accorded some priority. Elsewhere-and in those regions where the major epizootics have already been brought under control - improved feeding and management should be accorded priority. It is axiomatic that it is uselessto improve the genetic merit of livestock if environmental factors remain unimproved, and that the upgrading of livestock must be accompanied by the upgrading of management and by improvements in the overall environment. On the other hand, improvements in management and in the environment should not be allowed to outpace improvements in the genetic merit of the animals managed within that environment. It is just as wasteful to provide an environment that cannot be fully exploited on account of the low genetic merit of the livestock used, as it is to provide animals of high genetic merit for a poor environment.

The reproductive cycle

The reproductive cycle begins when the ovum or egg of the female is fertilized by the spermatozoa or sperm of the male and it ends at parturition. Male reproductive cells or sperm differ markedly from other body cells

148

Reproduction

and Breeding

Seminal vesicles

Rectum Cowoers glands

Vas deferens \1

Prostate Urethra Epididymis Test is

Fig. 4.1

Diagrammatic representation of the male reproductive organs of cattle.

in that they possessa tail-like structure which enables them to move freely. Sperm are produced in the testes, which are encased in and protected by a scrotumto form the testicles (Fig. 4.1). The scrotum possesses a muscular attachment which enables a male to lower or raise his testicles. During hot weather the testicles are lowered away from the body, in an effort to ensure that their temperature remains somewhat below that of the remainder of the body, a condition necessaryfor good viability of the sperm as these are adversely affected by high temperature. In the wet tropics it is not unknown for the testicles of an exotic bull to hang so low that they almost touch the ground. During cold weather the scrotum is contracted so that the testicles receive warmth from the animals body. Sperm are produced in vast numbers in the epithelium of the convoluted, seminiferous tubules of the testis and are stored in a retaining reservoir, known as the ductus epididimis (Fig. 4.1). The sperm mature during this period of storage, which may be for as long as 1 or 2 months. Thereafter they degenerateand lose their viability. The male sex hormone is also produced in the testes. This activates the sex instinct and sexual activities of the male. There are also various other glands accessoryto the testeswhich produce the fluid medium in which sperm are conveyed from the male into the genital organs of the female during the act of copulation at mating. The mix of the fluid medium and sperm % known as semen. The volume of semen produced at any one time varies considerably as between different species of livestock (Table 4.1) as does the average sperm content of the semen. The period during which sperm retain their viability within the female genital tract varies from speciesto species.In the cow the average duration is

The reproductive cycle

149

Table 4.1
Species

Data on the semen of some domestic livestock species Volume of male ejaculate (ml) _~_ Average Range Average sperm content
of semen

(million/ml)

Buffalo Cattle Goat Sheep Pig Fowl

2.0-I 3.0 2-O--10.0 0.1-3.5 0.2-3-O 5o*cL4000 01-1.0

3.0 5.0 0.7 1-O 250.0 0.2

900 1,ooo 2,700 2,500 125 4,000

30 hours, in the ewe 22 to 24 hours and in the sow up to 35 hours. The spermatozoa of some birds retain their viability for as long as 2 to 3 weeks. In the majority of speciesmating can be accomplished only when the female is at a specific stage in her sexual or oestral cycle. That is, when she is in heat. Camels and llamoids are an exception, as in their case copulation mduces ovulation (seeChs. 9 and 10, Pt II). After the semen has been deposited in the femalesgenital tract (Fig. 4.2) depending on where it has been deposited, the sperm move through the cervix and the uterus into one or other of the Fallopian tubes, where fertilization may take place if an egg is present. The movement of the sperm, from where they are deposited in the female genital tract, to the Fallopian tube usually takes from 2 to 15 minutes and is accomplished partly by their own movement and partly by the aid of other factors, such as rhythmic contractions of the vagina, cervix and uterus. The main site in the female genital tract where sperm are deposited during mating varies from speciesto species.In cows and
Rect urn Vulva Clitoris Vagina Cervix Bladder
,

1Jdder

Fig. 4.2

Diagrammatic representation of the female reproductive organs of cattle.

150

Reproduction and Breeding

ewes it is usually in the lower part of the vagina, while in sows it is in the cervical canal. Actual penetration of a sperm into an egg is probably the result of the sperms own powers of movement and the exudation by it of a chemical which helps break down the cell wall of the egg. Several sperm probably play some part in penetrating the outer layer of the egg, but only one actually fertilizes it. Eggs are produced in the ovaries of the female (Fig. 4.2). The process corresponds to that which takes place in the testes of the male when sperm are produced. Thus the ovary is analogous in its role in the reproductive cycle to the testis. It also produces hormones that activate and regulate the sexual activities of the female. Females normally possess two ovaries, eggs being produced alternately from one or the other. Ovaries are usually only a fraction of the size of the male testes. For example, a cows ovary is about 2.5 cm (1-Oin) in diameter, though the eggs that it produces are of microscopic size. Eggs are produced in the ovarys outer layer which is composed of germinal epithelium. Within this tissue specialized groups of cells, known as primary follicles, are the potential source of the eggs.At sexual maturity, and at specific intervals thereafter, one or more of the primary follicles rapidly mature within a thin, fibrous, capsulated structure known as a Gractfia~z jblricle. In cows these follicles are of a sufficient size to be visible to the naked eye. With increasing maturity the wall of the follicle becomesthinner and thinner and finally ruptures at the surface of the ovary, releasing the egg into the fibrillated, wide-mouthed end of the Fallopian tube. This process is known as ovulation. In the Fallopian tube the egg is held for several hours in a coagulated fluid which is formed from the viscous fluid that is released from the follicle at the same time as the egg. After a period that varies from species to species but is usually a few hours, the coagulated fluid reliquefies, thus releasing the eggand allowing it to move along the Fallopian tube towards the uterus. It is at this stage that fertilization by a sperm can take place. After ovulation the cavity in the empty follicle on the surface of the mammalian ovary is filled by a yellow cell, variously known as the yellow body or the corpus Zuteum. The function of this structure is to secrete hormones that seek to ensure the continuation of gestation and the suspensionof further ovulation until after parturition. Should the egg fail to be fertilized it loses its viability within a few hours, the yellow body disappears, the balance of hormones within the females body changes and the sexual or oestral cycle recommences. Factors affecting the oestral cycle Although the oestral cycle of the female is mainly governed by hormones that are secreted internally, there are other factors that exert a considerable influence on it, either directly or indirectly. The extent of their influence varies both between and within speciesand breeds.

The reproductive cycle

151

Apart from the abnormality of disease the most important factors affecting the oestral cycle appear to be the plane of nutrition, the length of daylight and ambient temperature. The inadequately fed female animal grows slowly and her sexual maturity, and hence the onset of her oestral cycle, is delayed. Very large numbers of tropical cattle have to subsist on a low level of nutrient intake for long periods during the year. As a consequence the first effective heat periods of heifers are often delayed until they are 2 years or older. On the other hand, in some species such as sheep, an abundanceof feed during or just before the normal breeding seasonoften increases fertility, as more than one egg is liberated at ovulation with resultant multiple births. Certain individual forage plants can also affect the sexual cycle of the livestock that consume them. For example, there are grassesand legumes that at certain stages of growth contain sub stancessimilar in chemical composition to the sex hormones produced by animals. The consumption of these plants may have either a beneficial or an adverse effect, according to the time of the year and the amount consumed.In Australia, Merino ewesconsuming considerable quantities of a subterranean clover (Trifolium subterraneum var. Dwalganup) exhibit infertility and difficulties in gestation and parturition. Pigs fed large quantities of the legume Leucaena leucocephala lose hair and often become infertile.
Nutrition. Length of daylight. Changesin the length of daylight may affect the sexual

cycle of all domestic livestock, but the effect is only of major importance for sheep,goats and poultry. In the temperate zonesthe breeding seasonof sheep is associated with a shortening day. As a consequenceewes transferred from the northern hemispheretemperate zone to the southern change the months in which they breed. In the tropics, where the length of daylight varies little throughout the year, most breeds of sheep reproduce at any season, although there may still be some seasonal variation in the intensity of breeding. Rams transferred from the temperate to the tropical regions of Australia are usually infertile for the first year after the transfer. Male goats transferred from the temperate region of Australia to Fiji have been reported to behave in a similar manner. The effect of light periodicity on poultry has been used commercially to increase egg production. This can be raised appreciably by slowly increasing the length of exposure of the birds to natural or artificial light. There is considerable experimental evidence that high ambient temperatures directly affect the sexual cycle of female livestock (seeCh. 1, Pt I), the libido of males and viability of the sperm of males.
Ambient temperature.

152

Reproduction

and Breeding

Table 4.2 Average length ojoestral cycle, duration qf heat and time of ovulation of some domestic livestock species
Species Oestral cycle (days) Average Range Duration of heat (hours)
Average

Time of ovulation (hours) Average Range

Range

Buffalo Cattle Goats Sheep Pigs

21 21 19 17 21

very 36 variable 16-26 18* 18-21 14-21 16-30 28t 26 48

12-I 20 14-22 24-72 24-48 40-65

very 5-24 after end variable of oestrus 11 2-22 after end of ocstrus 12.-36 after onset of oestrus 24 before end of oestrus 36 1847 after onset of oestrus

*Bos indicus cattle often exhibit shorter heat periods as do B. taut-us milking cattle managed in the tropics. +Considerably shorter in some tropical breeds.

Managerial considerations Since, with the exception of camels and llamoids, ovulation occurs at a definite time measurablein hours after the onset of heat and in all mammalian speciesthe egg and the sperm are relatively short-lived, it is very important that mating should be planned so as to coincide with ovulation. As far as is practicable mating should take place about the same time-or somewhat before- ovulation occurs so that the sperm will have the maximum chance of meeting with an egg that is passing down the Fallopian tube. Thus ir. order to properly plan matings it is important to know the length of the heat period, the duration of heat and the time of ovulation. These vary from speciesto species.Some details of average data for various speciesare shown in Table 4.2. The first occurrence of heat is determined mainly by environmental factors, but within the same environment there is considerable individual animal and breed variation. The occurrence of heat is evident in most species by some form of restlessness the part of the femaie and by her seeking out a male. In on some species, particularly the pig, there is a slight relaxation of the external genitalia. There may also be characteristic exudations from the external genitalia. However, even within species there is a very wide variation in the manner in which females exhibit the onset of heat. In some females heat is easy to detect and in others it is very difficult. For example, it is well known that heat detection can be very difficult in some breeds of Bos indicus cattle while it is particularly easy in the Bali breed of B. (Bibos) cattle. What is certain is that the easiest manner of finding out if a female is in heat is to see if she will accept a male.

The reproductive cycle

153

Fertihtion and gestation When a female accepts a male and he copulates and ejaculates, millions of sperm are deposited in her vagina. Only a portion of these, however, reach the extremity of the l\terus and only then if the genital passages are relaxed, are free from inflammation and the secretions in the tract are chemically and physically appropriate. Eventually only one sperm will fuse with the egg. Which one of the many will accomplish this feat is probably a matter of pure chance. That is not to say that only one sperm is necessary for fertilization to take place. On the contrary, it has been well established that there must be a certain concentration of sperm at the site of fertilization if one of them is to be effective. The process is somewhat different in the case of birds. As no yellow body replaces the ruptured follicle in the ovary and therefore no hormone is excreted to inhibit further ovulation, an egg can be released each day. In the case of the hen, the cocks sperm accumulates towards the end of her Fallopian tube and each egg is fertilized as it moves along the tube. 1 The fusion of one sperm from the male and the single egg of the female constitutes fertilization and is the only physical link between one generation and another, that is to say that the genetical make-up of the male and the female are combined at the moment of fusion to produce a fertilized egg,called a zygote.Shortly after fertilization this zygote begins to divide. The first division is generally completed within 24 hours. As the result of repeated cell divisions and the formation of special structures the embryo of the newly created individual begins to take shape. There are various stagesof embryonic development, the length of each depending on the total length of the embryonic period or the length of the gestation period of the species.In cows, for example, the zygote takes about 4 days to reach the eight- to sixteen-cell stage. The next phase in the development of the embryo is further cell division with the formation of a group of specialized cells which become the embryonic membranes, as shown in Fig. 4.3. Three membranesenvelop the embryo: the outer chorion which directly covers the allantois; the outer layer of the allantois which is joined with the chorion to form the allanto-chorion; and the inner amnion. The amnion, coupled with the inner layer of the allantois, forms the aIlanto-amnion. The allantois and the amnion are filled with fluid and surround the embryo. At an early stage in the development of the embryo, the allanto-chorion makes contact with the developed uterine mucosa. At this stage implantation into the uterus begins. Implantation is a gradual process. In the cow it begins about one month after fertilization and is completed during the third month. In specieswith shorter gestation periods, implantation usually takes a shorter period of time. For example, in sows implantation takes place during the second and third weeks of pregnancy. Implantation takes placeso that there can be an exchangeof substances between the maternal and embryonic tissues.This exchange takes place

154

Reproduction

and Breeding

~Umbiiical stalk Amniotic cavity,

Allanto-chorion Allanto-amnion Umbilical cord

Cord - coelom Fig. 4.3 Schematic illustration of the foetus and foetal membranes in the cow about 27 days after fertilization of the egg (Johansson and Rendel, 1968).

through a composite structure known as the placenta. It consists of the allanto-chorion, which is the embryonic part, and the mucous membrane of the uterus, which is the maternal part. Different speciespossess different types of placenta. The placenta of the pig is diffuse, the allanto-chorion making direct contact with the uterine mucosa, while in ruminant species the exchange of materials between the mother and the foetus occurs through button-type structures known as cotyledons. The development of the foetus appearsto be very slow during the early stagesof gestation, but the pace gradually quickens. In Bos taurus cattle, for example, the average 45day-old foetus weighs approximately 85g (3 oz), while at 90 days it will weigh approximately 1.0kg (2.2 lb) and at 200 days approximately 10kg (22 lb). There is considerable variation in the length of the gestation period as between speciesand some variation between breeds and types within species(Table 4.3). At parturition the placenta is normally evacuatedfrom the uterus after the young animal is born. Influence of hormoneson reproduction Hormones have a role at all stagesin the reproductive cycle, for example the development of the egg in the ovary, the rupture of the follicle and the conditioning of the uterus to receivethe fertilized egg are all initia.ted and controlled by hormones. Hormones are chemical substances that are produced in ductless glands known as endocrine glands. These secrete the hormone directly into the bloodstream which transports it to the various body tissues on which it acts. Hormones are specific in that they affect particular tissues

The reproductive cycle

155

Table 4.3
Species

Gestation periods of some domestic livestock species Type Breed Gestation period (days) A verage Range

Buffaloes Camels Cattle

River Swamp Dromedary Temperate

Murrah Egyptian

308 317 332

-

280 Angus Brown Swiss Charolais Friesian Hereford Jersey

-

302-313 313-319 301-343 360-390 273-289 273-282

-

289 289 279

Goats Llamoids Sheep Pigs

Tropicai Temperate Tropical Alpaca Llama Temperate Tropical Temperate

283-286 278-280 284-288 154 146 342 345 330

-

Saanen H uacaya Surti Dorset Horn Merino

145-148
-

114

140-148 148-152 140-160 110-117

that will react only to that hormone. The testesof the male and the ovaries of the female serve a dual role as ductlessglands. They not only produce, respectively,male and femalesex hormones but also produce the sex cells or gametes that transmit characteristics from one generation to the next. The orchestration or integration of hormone production within the reproductive cycle is conducted from a site in the brain of the animal known as the hypothalamus (Fig. 4.4). This controls the pituitary gland, which produces the gonadotrophic hormones, including the folliclestimulating hormone (FSH), the luteinizing hormone (LH) and the luteotrophic hormone (LTH) which stimulates the yellow body in the follicle to produce the hormone progesterone. The pituitary also produces several other hormones which regulate other specific bodily functions not directly concerned with the reproductive cycle such as the growth hormone (GH) and the thyro-trophic hormone (TSH), etc. The major hormones concerned in the reproductive cycle are as follows:
Follicle-stimulating

The main function of FSH in the female is to stimulate follicular growth in the ovaries and to control the maturation of the egg.In the male, FSH initiates the growth of the testes and induces the production of sperm.
hormone (FSH).

1%

Reproduction

and Breeding

,Nerve

irIses,

I Pituitary 1 gland podeerior I,,,/ ppior

/m fl py%q pijiq mi

1 Androgen Male sex hormone

Lb Oestrogen Progesterone Female sex hormones

Fig. 4.4 Simplified schematic representation of the endocrine glands concerned with reproduction.

In the female LH controls the rupturing of the follicle, thus initiating ovulation. It is also responsible for the production of the yellow body in the ruptured follicle. In the male LH stimulates the production in the testes of androgen, the male sex hormone. This hormone controls the production, by the yellow body in the ovarian follicle, of progesterone.
Luteo-trophic hormone (LTH). Female sex hormone (oestrogen). The ovarian follicles produce this

Luteinizing hormone (LH).

hormone while they are developing under the influence of FSH. It is responsible for bringing the female to an excitable stage during heat, so that she will accept the male. It also has an effect on the vagina, causing an increase in the secretion of mucus during heat. As stated above, this hcrmone is produced by the yellow body in the ovarian follicle. It prepares the uterus for the implantation of the foetus and generally ensuresthe continuation of the normal course of pregnancy. Progesterone, together with oestrogen, also has an effect on the mammary glands of the female, preparing them for t$ end of pregnancy when they will be required to secrete milk.
Progesterone.

The basis of inheritawe

157

As stated above, this is under the control of LH and is secreted by the testes. It is responsible for stimulating the male in order to increasehis libido and for the development of secondary sexual characteristics in the growing male. The role of hormones in the reproductive cyc!e is obviously extremely important and in certain casestreatment of reproductive disorders can be affected by the therapeutic use of hormones. It must be emphasized, however, that such treatment is highly specialized and should be conducted only under professional guidance. Synchronization of the oestrus cycle using hormones has been experimentally performed in recent years in cattle, pigs, sheep and goats. Quantitative relationships between the hormones and fertility are still being intensively investigated and when more is known of their interrelationships greater successin controlling oestrus can be expected. At present oestrus can be conveniently synchronized in the sheep and goat and the breeding seasonslightly advanced by the use of progestogenimpregnated intravaginal sponges.Some degree of control of ovulation in yarded cattle is presently possible by the feeding of hormonal compounds which suppress heat and ovulation during the period of their administration. While ovulation is being prevented, the corpora lutea of the ovaries regress as the follicles continue to develop. At termination of the treatment the majority of the cattle come into heat at approximately the sametime. Although there is a relatively precise method of controlling ovulation in the sow using a product known as otethalli bure-a gonadotrophin-suppressing compound-it is not used extensively and in some countries its use has been prohibited as it has been found that if accidentally included in the feed of pregnant sows it causesthe birth of deformed piglets. The genetic impact and advantage of having a convenient practical method of synchronizing oestrus in farm animals would be through a greater useof artificial insemination (AI) so that a smaller number of sires would be required, thereby considerably increasing the possible selection differential. The best sires could be used on a larger percentage of the total population. Such a practice would, of course, have to be closely supervised by animal breedersso as not to cause any major increase in the level of inbreeding in a breed. Another practical advantage of oestrus synchronization would be the possibility of closer managerial control of the female breeding herd during the breeding season and in producing large groups of marketable animals of about the same age and of similar genetic make-up.
Male sex hormone (androgen).

The basis of inheritance

All tissues of the body consist of cells, differing in size and shape in accordance with their bodily function. They contain a substance known

1%

Reproduction

and Breeding

as protoplasm and among other characteristics they can metabolize nutrients, grow and multiply. A typical cell (Fig. 4.5) is surrounded by a membrane in which the cytoplasm, nucleus and centrosome can be distinguished. In the nucleus of the resting cell one or more nucleoli are present together with a lar= number of smafi particles known as EjL clzromatir~g~am&s that at specific stagesof cell division appear as threadlike structures or chromosom,os. These chromosomes occur in pairs. The total number of pairs and the size and shape of individual pairs vary from speciesto species(Table 4.4). There can also be differences between chromosome pairs within the samespecies,particularly those that determine the sex of the individual. Chromosomes possess the ability to multiply by division in step with the multiplication of cells while still

Mitochondria Cell membrane Cytoplasm Fig. 4.5 Schematic representation of an animal cell.

retaining their individual characteristics. They are of paramount importance because they carry the coded messagesor genes that transmit inherited characteristics from one generation to the next. The genetically active substance in the genesis deoxyribonucleic acid or DNA, which is considered to consist of two chains of nucleolides twisted together in a spiral formation, the sequenceof nucleolides in the formation determining the genetic information that is carried in the gene. As the possible total combinations of nucleolides is very large the quantity of genetic information that can be stored in this manner is extraordinary. When a cell divides, the spiral formation of the DNA molecule splits lengthways, one chain of nucleolides going to one cell and the other chain to the other cell. Tnthe two new cells the single chains of the split DNA molecule eachact as templates,an exact copy of the single chain being synthesized in order to form a new DNA molecule exactly similar to the one from which the split chains originated. In this way each of the new cells acquires its full complement of chromosomes and genes.

The basis of inheritance

159

Table 4.4

The diploid number qf chromosomes in some domestic livestock species Species

Type River Swamp

Diploid (2n) number of Chlm?lrISOme.~

Buffaloes Camels Cattle Goats Llamas Sheep Pigs Fowls Ducks Turkeys

50 48 74 60 60 74 54 40 78 80 82

Cell division There are two sharply differentiated types of cell in the animals body. Somatic cells that make up the body of the individual animal and are concerned with all body structures and most body functions, and germ cells that are concerned with the reproduction of a new generation of individual animals. When somatic cells multiply, the daughter cells are exact replicas of the mother cell, whereaswhen germ cells are produced the number ofchromosomes is reduced to half the number for the species. Somatic cells are diploid and ;?ossess chromosomes whilst germ cells 2n are haploid and possess chromosomes. n
Somatic cell division or mitosis.

In mitotic division each daughter cell receives the same chromosome complement as was possessedby the mother cell by the mechanismdescribed in a previous paragraph. Mitosis is a continuing processand it is the meansby which the individual animal grows and replaces worn-out cells with new, healthy cells. A schematic illustration of mitosis is shown in Fig. 4.6.
Germ cell division, reduction division or meiosis. The process of meiosis

occurs in the same way in both the male and the female of the species. The male and female germ cells are produced in the manner described in the section concerned with the reproductive cycle. It was stated in a previous paragraph that there are differencesbetween chromosome pairs within the same species,in particular those that determine the sex of the individual. These chromosomesare known as the sex chromosomes. They also carry many other genesfor other characteristics and these are called sex-linked characteristics. The other chromosomes in the cell are known as autosomes. Although the mechanics of meiosis are the same in the male and the female each process will be described separately.

160

Reproduction

and Breeding

Cent romere

Interphase

Early Prophase

Mid Prophase

Late Prophase

Metaphase

Early Anaphase

Anaphase

Early Telophase

Late Telophase

Daughter Cells

Fig. 4.5 Schematic illustration of mitosis (Gardner, 1960).

In the male the germ cells are found on the inner wall of the seminiferous tubules of the testicles.At sexualmaturity thesecells are stimulated by hormonal action to produce functional sperm. The process of meiosis begins with the pM~~ry spermatocyte growing and enlarging and finally dividing into two secondaryspermatocytes. The division follows a process known as synapsis in which the pairs of like chromosomes known as homologues- one member of each pair having been derived from each parent -fuse together. During this process filaments of chromatin twist around each other, thus providing an opportunity for the exchange of genesfrom one homologue to the other. This is one of the mechanisms by which variation occurs, but there is of course no complete interchange of genesbetween one homologue and another. After synapsis there is a separation of the fusedpair of chromosomes,one homologue of eachpair moving to one or other side of the cell (Fig. 4.7). After the polarization of the chromosomes is completed a new cell wall is formed with the production of two secondary spermatocytes, each containing one-half

The basis of inheritance

161

the total number of chromosomes normal for the species.The haploid cells containing half the number of chromosomes are not of course identical with the parent cell or with the normal body somatic cells. The two secondary spermatocytesthen divide by the processof ordinary mitosis to produce four spematids. After a pericd of maturation the four spermatids develop into functional sperm with the characteristic tail, body and head. As the cell division proceeds,the mature sperm migrate from the wall of the tubule to the lumen or centre of the tubule through which they exit from the testicle in the manner described in the section concerned with the reproductive cycle. The processof meiosis in the female is similar to that in the male with the exception that the primary germ cell or ovacyte forms one secondary ~WKJWand one non-functional polar body (Fig. 4.7). Each contains onehalf the normal number of chromosomes. The polar body differs from the secondary ovacyte in that it has no yoke. The secondary ovacyte and
Spermatogenesis Spermatogonium

0 0
I$ $

Oiigenesis 0 /I vu t 3. ; 04% 0 &I t

Oiigonium

t 9; we Primary Cl 0 spermatocyte ill1

Primary oijcyte

Fig. 4.7 Schematic illustration of meiosis (Gardner, 1960).

162

Reproduct ion and Breeding

the polar body then further divide by the process of mitosis: the ovacyte producing an ovum or eggand an additional polar body and the first polar body producing two polar bodies. The ovurn thus produced is capable of being fertilized by sperm. The polar bodies appear to be non-functional and are absorbed. The function of meiosis is to retain constancy in the chromosome number of the speciesand to ensure that both the male and female contribute genesto their offspring that are randomly representative of their characteristics. The haploid (n) egg contains half the number of chromosomesof the female parent when it fuseswith the haploid (IZ)sperm that contains half the number of chromosomesof the male, forming a diploid (2n) zygote or fertilized egg that contains the number of chromosomes characteristic of the species.

The inheritance of characteristics

As the distribution of chromosomesand consequently of genesin meiosis is at random, this permits new combinations of paternal and maternal characteristics in the offspring. A knowledge of this process and the ability to select individuals through various selection methods provides an opportunity for the selection of animals possessingnew combinations of genes that provide desirable characteristics required by man in his domestic livestock. It is also obvious that new combinations of genes may alternatively produce individuals with undesirable characteristics. These animals can be culled before they are allowed to breed and reproduce. The process of meiosis described above does not necessarily proceed smoothly on all occasions.There can be accidents during the processso that the normal number of chromosomes for the speciesis not always attained. These accidents may be due to non-disjunction, separation of parts of the chromosomes, abnormal pairing or non-pairing of homologues,etc. In addition, as explained in a previous paragraph, the:arrangement of genes on the chromosomes can vary due to crossing over of genesduring the processof synapsis.When variations due to these causes occur they will, of course, produce considerable and in some ways abnormal variations in the characteristics of the offspring. These phenomena are discussedin some detail in later sections. Phenotypic differences-or the difference in the appearance of individuals both within and between species- have been studied and theorized on by naturalists for a long time. In 1900, three botanists working independently- De Vries in the Netherlands, Correns in Germany and von Tschermak in Austria - proposed theories to explain the mechanism of the inheritance of individual characteristics. The work of these three botanists independently substantiated the work of an Augustinian monk, Gregor Mendel, who in 1866 had published papers that reported on data from 8 years of crossbreeding experiments using

The inheritance

of characteristics

163

common garden peas. Mendels work was known in both Europe and America, but the significance of his findings was not appreciated until De Vries, Correns and von Tschermak published their -papers. In his papers Mendel proposed two basic laws on the inheritance of characteristics. These were: (1) the law of segregation -that the characteristics of an individual are determined by pairs of genes, but that their germ cells possess only one genefrom each pair; and (2) the law qf independent assortment-that the genescombine at random with each other, both at the formation of germ cells and at fertilization. Since Mendels original publication there have been many modifications of these laws, but they are still fundamental for the whole science of genetics. The resulting overall interpretation of the laws of inheritance of characteristics is now known as the Mendelian theory of inheritance. The practical significance of the first two laws of Mendelian theory are of extreme importance when man is selecting for specific traits in plants and animals. They will now be considered with reference to examples from domestic livestock. As stated in a previous paragraph the chromosomes occur in pairs and each homologue contains identical genesthat occur in the same order in eachhomologue. Theseidentical genesare known as alleles. Although the opposite genesare identical in that they affect the same phenotypic characteristic or development processof a character, they do not necessarily influence it in the sameway. If both alleles have the same influence on a characteristic the individual possessingthem is said to be homozygousfor that characteristic, but if they differ in their influence the individual is said to be heterozygous for the characteristic. If the effect of one allele is stronger than that of the other to the extent that it masks the effect of the other, the masking allele is said to be dominant, while the allele that has been masked is said to be recessive. For example, in cattle the polled or absence-of-hornstrait is dominant over the horned trait. Thus when homozygous horned cattle are mated with homozygous polled cattle the offspring possess genefor the presenceof horns and another one for the absenceof horns, i.e. they are heterozygous for the characteristic. Since the polled gene is dominant all the offspring are polled (Fig. 4.8). For graphic purposes, in order to expressthe actions of such genes,a capital letter is used to denote the dominant allele and a small letter to indicate the recessive allele. Thus, in the example given above the homozygous horned individual would be designated as hh, the homozygous polled individual as HH and the heterozygous individuals resulting from the mating as Hh. The situation could then be expressed in the following manner.
HH polled
X

Ml horned

I Hh polled

164

Reproduction

and Breeding

Reproductive cells

HH Polled

hh

Homed

FI generation Reproductive cells

F2 generation HH Polled Fig. 4.8 Illustration Hh Polled hH hh Horned

of the inheritance of the polled trait in cattle.

The phenotypes of the HH and Hh individuals will be the same, but their genotypes-or their potential for transmitting characteristics to their offspring - will be different. In some traits dominance is not complete. If this is the case and dominance is only partial then the offspring may exhibit phenotypic characteristics that are a blend of those of the two parents. A well-known example occurs in the Andalusian breed of poultry in which there are black and white varieties. If a black homozygous individual is mated to a white homozygous individual the offspring possess blue feathers. When an individual is homozygous for a recessivecharacteristic, such as is the case in the white variety of Andalusian poultry, the phenotype is the same as the genotype since the recessivetrait has neither been masked nor modified. An individual that is homozygous for one pair of genes has only one type of that gene to transmit and therefore breeds true. However, if an individual is heterozygous, it will have two types of the same gene to transmit. As only one of each pair of genes is transmitted to each germ cell at meiosis, any resulting fertilized egg could contain either of the two

The inheritance of characteristics

165

diflerent types of the same gene. If the example of the transmission of horn or absenceof horn characteristics in cattle is reconsidered it will be seen that when the polled herozygotes (Hh) are intermated then, as both parents produce equal numbers of germ cells carrying either allele, the chancesof any sperm cell from the male fusing with any egg of the same type or of a different type are equal and so four combinations of the genes are possible. These combinations are HH, Hh, hH and hh (Fig. 4.8). Thus the result of a random fertilization is on average one homozygous dominant (HH) polled animal, two heterozygous (Hh) polled animals and one homozygous recessive(hh) horned animal. The phenotype grouping is therefore three polled to one horned animal, but since fertilization is at random it must not be expected in practice that among every four progeny of such matings the result will be three polled and one horned animals. This will occur only when a very large number of matings are evaluated. The chance of a recessivetrait being exhibited under these circumstances is obviously one in four, and the chance of obtaining a true breeding individual from among those which phenotypically exhibit the dominant trait is one in three. The practical importance of this type of information is considerable when the animal breeder is selecting for specific characteristics. An example of what happenswhen dominance is lacking in alleles can be demonstrated by referenceto the inheritance of coat colour in Shorthorn cattle. If a red-coated Shorthorn is represented by RR and a whitecoated Shorthorn by rr, then when red and white Shorthorns are mated the offspring are of a Rr type and their coat colour is roan rather than either red or white. This may be represenicd as follows:
RR red coat
X

rr white coat

1 Rr roan coat

If the roan offspring are inter-mated then segregation takes place as in the previous example. This may be represented as follows:
RIroan coat RR red coat Rr roan coat
X

Rr roan coat rR roan coat rr white coat

Thus in this case the phenotypic and genotypic ratios are the sameone homozygous (RR) red animal, one homozygous (rr) white animal and two heterozygous (Rr) roan animals and the heterozygotes are distinguished in appea.rancefrom either of the homozygotes. This was not the case in the example discussed previously of the inheritance of the

166

Reproduction

and Breeding

polled trait. The genotypes HH and Hh appeared to be phenotypically the sameand it would only be possible to distinguish them genotypically by performing a further breeding test, i.e. by mating them with animals of a known genotype and observing the progeny. For example, if the polled HH animals were mated with horned animals then all the progeny would be polled, whereas if the polled Hh animals were mated with horned animals half of the progeny would be polled and half horned. The examples given above demonstrate the operation of the first Mendelian law on the segregationof inherited characteristics. The second Mendelian law, that of independent assortment, can be demonstrated in an example where the inheritance of two pairs of genes are considered at the same time. For example we can consider two dominant traits, polled&s and snorter dwarfism in Hereford cattle. If a homozygous
HHDD polled : non-dwarf
X

hhdd horned : dwarf

I HhDd FI generation polled : non-dwarf --------------------___________c________----------HhDd polled : non-dwarf F? generation HHDD polled : nondwarf HHDd polled : nondwarf HhDD polled : nondwarf HhDd polled : nondwarf HHDd polled : nondwarf HIIdd polled : dwarf HhDd polled : nondwarf Hhdd polled : dwarf HhDD polled : nondwarf HhDd polled : nondwarf hhDD horned : nondwarf hhDd horned : nondwarf HhDd polled : nondwarf Hhdd polled : dwarf hhDd horned : nondwarf hhdd horned : dwarf HhDd polled : non-dwarf

Ratio of phenotypes in the F2 generation (polled : non-dwarf) 9: (polled : dwarf) 3: (horned: non-dwarf) 3: (horned : dwarf) I Fig. 4.9 Schematic representation of the inheritance of the polled and snorter dwarf traits in Hereford cattle. Note: H represents the polled, h the horned, D the non-dwarf and d the dwarf traits.

The inheritance of characteristics

167

polled, non-dwarf individual with the genotype HHDD is mated to a homozygous horned, dwarf individual with the genotype hhdd then the resulting F1 progeny will be phenotypically polled and non-dwarf but genotypically HhDd. If the Fi progeny are then intermated the resulting offspring (Fz) will exhibit four different phenotypes in the ratio of nine polled non-dwarf, to three polled dwarf, to three horned non-dwarf, to one horned dwarf individual. There will be, however, nine different genotypes (Fig. 4.9). The phenotypic ratio of 9 : 3 : 3 : 1 is mathematically the square of the ratio of 3: 1, which is the phenotypic ratio when one characteristic is considered. If three pairs of characteristics that all exhibit dominance are considered then a phenotypic ratio of 27 :9 :9 :9 : 3 : 3 : 3 : 1 will result. The number of possible combinations increasesrapidly with an increase in the number of gene pairs. This can be seen from Table 4.5. Thus, in domestic animals where the number of heterozygous gene pairs is very large it is not surprising that no two individuals, with the
Table 4.5
Comhirzation possibilities in the FZ generatiorz when F1 irzdividlrals are hetero~ygous~for a specific number of gerle pairs No. of gametes Combination possibilities No. qf genotypes No. of homozygorts combinations

Pairs of
genes

1 2 3 4 10
n

2 4 8 16 1,024 2

4 16 64 256 1,048,576 4 .

3 9 27 81 59,049 3

2 4 8 16 1,024 2

Source: Johannsson and Rendel(1968).

exception of identical twins, are genotypically or phenotypically completely alike. Where dominance exists in one pair of genesbut is lacking in the other the situation is different. For example, if we consider Shorthorn cattle where the polled trait is dominant and the coat colour trait lacks dominance, then if a homozygous polled red Shorthorn is mated with a homozygous homed white Shorthorn all the F1 progeny will be heterozygous polled and roan in colour (Fig. 4.10). When the F1 generation are intermated the following ratio of different phenotypic types of cattle would be obtained in the FZ generation: three polled and red, six polled and roan, three polled and white, one horned and red, two homed and roan and one homed and white (Fig. 4.10). From the examples that have been discussed above it will be appreciated that if the mode of inheritance of traits is known then expected ratios of different types of offspring from specific matings can be calculated, as can the probability of obtaining a particular type of

168

Reproduction

and Breeding

HHRR polled : red coat

hhrl horned : white coat

FI generation -------------------------------------------------

I HhRr polled : roan coat HhRr polled : roan coat

HhRr polled : roan coat Fz generation HHRR polled : red HHRr polled : roan HhRR polled : red HhRr polled : roan HHRr polled : roan HHrr polled : white HhRr polled : roan Hhrr polled : white

HhRR polled : red HhRr polled : roan hhRR horned : red hhRr horned : roan

HhRr polled : roan Hhrr polled : white hhRr horned : roan hhrr horned : white

Ratio of phenotypes in the F2 generation (polled : red) 3:(polled : roan) 6: (polled: white) 3: (horned : red) 1: (horned : roan) 2 : (horned : white)] Fig. 4.10 Schematic representation of the inheritance of the polled trait and coat colour in Shorthorn cattle. Note: H represents the polled trait, h the horned, R red coat colour and r white coat colour.

individual from a specific mating. There are, however, a number of important exceptions to these general rules. Modifications of the basic Mendelian law may be necessary for a number of reasons. These include, among others, an additive effect of genes, the effect of duplicate recessivegenes and the non-linear interaction effects between genesat different loci or epistasis. Also important are the effects of linkage, crossing-over, multiple alleles and disturbances in the chromosome mechanism. In general, no sharp distinction can be made between qualitative and quantitative characteristics. A characteristic may be determined by a major gene or oligogene, but one or more other minor genes or polygenes may cause some variation in the manifestation of the characteristic.

The inheritance of characteristics

169

Where geneinteraction is between alleles there may be overdominance if the heterozygotes are superior in one way or another to both the homozygotes. The occurrence of overdominance is assumed to play an iImportant role in such qualitative characteristics as viability and fertility and will be further discussedin the section on crossbreeding. The effect may be due to one genehaving a more advantageouseffect than two genes of the same type- i.e. a level of dosage effect-or to the alleles complementing each other in some way. In epistaais,the effect of a single gene depends upon with which other genes it interacts. The effect on the normal additive situation can be either negative or positive. An example of linkage is two characteristics that are somewhat related and not completely independent becausethey are controlled by genes that are located on the same chromosome rather than on different chromosomes. Under thesecircumstances independent assortment does not occur and the ratios of the phenotypes may be different from those stated above, where it has been assumed that the genes are located on separate chromosomes. Crossing-over is said to occur when - although the genesare known to be located on different chromosomes-i:1 some way they stay together. It happens if chromosomes do not completely sort themselves independently but somehow a part of one chromosome joins up with a part of its homologue or vice versa. It has been assumed in the examples of segregation and independent assortment given above that at any given location in the chromosomes there would be two alleles.More than two alleles can, however, be present and this also modifies the normal Mendelian ratios. One of the bestknown examples is concerned with the inheritance of blood type in man. In this case there are four blood types derived from combinations of three allelic genes. The livestock breeder is not of course usually concerned with the inheritance of one or two single traits. What is of major interest is the inheritance of quantitative characteristics such as body size, milk yield, the fat content of milk and the egg production of hens. It is therefore ii portant to realize that although the fundamental laws of Mendelian inheritance still apply, characteristics of this type are strongly modified by environmental variation and that a very large number of genes are involved that may have different degreesof effect and interact in a complicated way with each other. The inheritance of sex In the examples of inheritance that have been so far described it has been assumedthat the geneswere located in the autosomes and not on the sex chromosomes.The la&i ore different from the autosomes in that they are the primary determinants of sex in the offspring. In mammals

170

Reproduction

and Breeding

the femalepossesses pair of homologues known as the X-chromosomes, a while the male possesses X-chromosome and one that is very different one called the Y-chromosome. The inheritance of sex can be shown schematically as fo!lows:
XX female xx female XY male 1 xx female XY male XY male

Thus the sex ratio should theoretically be one to one. In farming practice the male: female ratio has been calculated to be 515, 52.3 and 49.2 for cattle, pigs and poultry, respectively (Johansson and Rendel, 1968).It is generally assumedthat in mammals the male:female ratio at fertilization is considerably higher than 50 per cent, but that the ratio is reduced during the gestation period becausemales have a higher foetal death rate. The reason why the sex ratio is higher at fertilization could be because some reasonspermscarrying the Y-chromosome are more for viable. If there are differences in viability between the sperms carrying the X- and the Y-chromosomes then it might be possible to separate them. This could be the basis for some future method of sex determination. In birds the male is the homogametic (XX) sex and the female the heterogametic (XY) sex. Other geneswhich are located on the sex chromosomes will obviously be linked or associatedwith sexand will be transmitted to the next generation in combination with sex.The fewer the chromosomes that the species possesses the larger the sex chromosomes, the more traits are likely and to be associated with sex and inherited in a sex-linked manner. To date, no sex-linked characteristics of economic importance have been recorded in mammalian livestock, but there are several in poultry. These include barred compared with dark heads, silver compared with gold down colour and curly compared with late feathering in chicks. Sex-linked inheritance of a colour factor has also been demonstrated in turkeys and geese.Man also demonstrates a number of well-known sex-linked characteristics, including haemophilia and red-green colour blindness. Haemophilia is also sex-linked in dogs. It was thought at one time that the colour-marking genes could be usedfor the rapid determination of sex in chicks, but as poultry are bred strictly for their production characteristics breeders could not afford to restrict their breeding programmes to the breeds exhibiting sex-linked colour traits. In any casesexing by examination of the external genitalia of day-old chicks is now commonplace so that the economic need for a colour determinant of sex in the chick has declined. A schematic diagram of the sex-linked inheritance of barring is shown in Fig. 4.11. When a

The inheritance of characteristics

171

black cock is mated with a barred female, in the F1 generation all the males are barred and the fem.ales black. After hatching the difference are between the sexes can be seen immediately as the male chicks have a light patch on the back of their head and are relatively light coloured on the other parts of the body while the female chicks possessdown of an even colour.

Parents

Black cockerel

e-----v
----- -Barred hen

Chromosomes of parent stock

Chromosomes of offspring

Or?

----_--e-ev
----- Barred cockerel

Offspring

Black hen

Fig. 4.11 Illustration of the sex-linked inheritance of the barring trait in poultry (tiammond et al.. 1971). Note: B represents barred and b self-colour (black).

The White Leghorn breed possesses sex-linked gene for early feathera ing, an economically desirable characteristic as it is linked with earlier maturity and the chicks are lessliable to chill, whereas someaf the heavier breeds have a corresponding dominant allele for late feathering. If White Leghorn cocks are bred to the heavier breedhens the femalechicks exhibit the early feathering trait.

172

Reproduction

and Breeding

Autosexing breedssuch as the Legbar and the Cambar have been produced, but they have not been particularly popular as they have not been so productive as some other non-autosexing breeds.

Mutation
There are many examplesof the suddenappearance of variants or sports in livestock. The processis known as mutation. It may be defined as every change in the heritable sensewhich is not due to segregation or to a recombination of previously existing genes. This spontaneous change may be due to a variety of reasons, and there is presumably a change in the chemical composition of the gene. Mutation can take place in both somatic and germ cells; it occurs naturally at all times, but its frequency can be increased by the effect of radiation and certain chemicals. Natural mutations apparently occur at rates ranging from one in 100,000 generations to one in 10 million generations. The majority of visible mutations appear to be generally undesirable as far as the livestock breeder is concerned, but more of them may be advantageousfor the breeders of fancy pet animals. It is believed that the majority of the fancy breeds of dogs, cats, rabbits and various speciesof birds owe their origin to a mutation. If the mutant is a dominant gene the trait that it controls should be exhibited immediately, but if it is a recessive gene the trait may be hidden for generations. It must be presumed that animals gradually acclimatize themselves generation by generation to a new environment partly on account of mutations that are favourable to them in their new environment. If a single gene has a multiple effect - a phenomenon known as pleiotropy - even mutations that could be considered to be desirable may have other effects that are very undesirable. For example, double muscling in beef cattle might be considered economically desirable, but when the trait is inherited in the homozygous form it is also the cause of unthriftiliess and infertility. Another interesting example is the link between polledness and intersexuality of female goats. The polled condition can appear due to a mutation in horned goats, and as it is a dominant character it would be expected that it should be possible to breed hornless goats. This is not the caseas homozygous polled females are intersexual and therefore sterile, and homozygotes are required in order to be able to breed for total homlessness. Many of the abnormalities common in livestock are due to mutations. Those that cause the death of the foetus are known as lethal factors. Others such as cleft palate, hernia, inverted nipples and atresia ani in pigs and cryptorchidism in cattle, sheep,goats and pigs cause difficulties after birth. The many types of coat colour in domestic livestock have all been derived by mutation from a more limited number of coat colours

Genetic-environmental

interactions

173

in wild animals. Abnormalities are often associated with white colour such as the underdevelopment of the vagina and uterus in white Shorthorn heifers. Details of known lethal factors in livestock in the temperate zone have been published by Lemer (1944). The majority of abnormalities noted in temperate-type stock are also known to occur in tropical-type livestock. Domestic livestock can and should be purged of unwanted mutations that causeabnormalities, particularly if male animals are going to be used to mate with a large number of females either naturally or by using AI. If the male is homozygous for the abnormality, then he will normally exhibit it so that his elimination from a breeding programme is simple. If he is heterozygous he may not exhibit the abnormality. He should be mated with two or three females of the same breed that exhibit the abnormality. If he is heterozygous for the trait half the resulting offspring will exhibit the trait; if he does not possessthe trait then none of the offspring will exhibit it. Heterozygous males can thus be identified and culled.

Changes in the number of chromosomes

When the chromosome number of a type or speciesis a multiple of the haploid chromosome number (n) and larger than the diploid number (211) is usual to refer to it as a polypbid. Changes in the chromosome it number have been mainly studied in plants and lower animals, and although polyploidy may have played a role in the evolution of animal species little is at present known of its incidence in livestock. In this context it is interesting to note than in man possessionof an additional X-chromosome, i.e. the male possessingXXY-chromosomes instead of the normal XY, may be one cause of intersexuality.

Genetic-environmental interactions
The suggestionby Hammond (1947),that livestock should be reared and bred in the most favourable environment if maximum improvements are to be made through selection, has been generally accepted in the temperate zone. However, this theory may not be sound if the differences between the environments in which the livestock are bred and used are very great, as they are between temperate and tropical environments. Genetic-environmental interactions may in fact be very important for the livestock breeder in the tropics. They may be defined as the differential response of a specific genotype in different environments.

174

Reproduction

and Breeding

One obvious effect of the en.vironment on the genotype is on the mature size of domestic livestock. Under natural grazing conditions in high-rainfall areas, where minerals have been leached from the soil, domestic livestock are generally smaller than genetically similar animals managed in drier areas where there is little leaching. An example occurs in East Africa where the semi-arid-area cattle such as the Boran and the Karamajong are on averagemuch larger than the East African Shorthorn Zebu that are found in the more humid areas, although it is highly probable that all these Zebu breedshad a common ancestry. In India virtually all breeds of Zebu hill cattle are on average smaller in size than the Zebu cattle found in the adjacent plains although they almost certainly possessa common ancestry (seeCh. 5, Pt II). The reasons for geneticenvironmental interactions are probably varied. One reason that has been suggestedis that the genes affecting a particular trait in an animal may not be the same in two very different environments. It is also possible that the effect of a specific mutation might be very different in different environments. For example, a mutation for additional or longer body hair could be very advantageous to an animal in a cold, temperate climate and very disadvantageous in a hot, humid tropical climate. What is certain is that the planning of breeding programmes in the tropics depends as much on the magnitude of geneticenvironmental interactions as it does on other genetic parameters. Readers who are particularly interested in interactions with regard to specific livestock should consult Pani and Lasley (1972).

Maternal influence
The mother has a very considerable influence on the development of her young during the gestation and suckling periods. Usually the birth weight decreaseswith an increase in the number of young born. In cattle and sheep the birth weights of individual twins are 25 to 30 per cent lower than the birth weights of singles. When reciprocal crosses are made between large and small breeds there is no experimental evidence that under most circumstances the maternal influence during the gestation period has any influence on body size at maturity of the offspring. Differences have, however, been noted where the disparities in size are extreme, i.e. between the Shire horse and the Shetland pony and between the Flemish Giant and the small Polish rabbit. Experimental work does not support the idea that the mother has, in the long term, any special influence on the offspring or that the offspring inherit more traits from their dam than from their sire. Differences that occur in the mature offspring of reciprocal crosses of extreme animal types must be considered to be due to carry-over effects of differences in the foetal and suckling environments.

Animal breeding practices

175

To what extent the age of the mother influences the frequency of congenital defects in domestic livestock is unknown. It has been shown that there are many instances of congenital malformation in domestic livestock where a change in the environment can have the same effect as a change in the genetic constitution of the animal. This phenomenon is known as phenocopy. Phenocopies of a number of genetically determined deformities in the offspring can be obtained by injecting the mother with specific chemical substancesduring the first part of her gestation period. Deficiencies of vitamins can also induce deformities in the foetus. In practice, it is suggestedthat deformities in young livestock should be scrutinized very carefully before they are necessarily attributed to the effect of heredity.

Animal breeding practices

Current animal breeding methods, reviewed by Mahadevan (1970) will be briefly discussed and the ways in which these practices can best be used in tropical environments will be considered. Selection In order to improve the average level of a livestock population for any trait by genetic means the population must be subjected to selection for the specific trait or combination of traits required. Some traits are strongly inherited, while others are weakly inherited as their development in the animal is more dependent upon environmental conditions. The intensity of inheritance of a specific trait can be measured. It is known as the heritability of the trait, is usually symbolized as h2, and it may be defined as those phenotypic differences in a trait that can be attributed to inheritance. It has been found that those parts of the animal that develop early in life tend to possesshigher heritabilities than those parts that develop later, and that heritability estimates are least variable if they are determined under standardized environmental conditions. Approximate estimates of the heritabilities of a number of selected productive traits in domestic livestock are shown in Table 4.6. It will be seen from this table that carcase traits are generally highly inherited, that liveweight gain and efficiency of feed conversion are moderately highly inherited, that milk and egg production are relatively poorly inherited and that such traits as fertility and viability are very poorly inherited. Heritability provides an index of the probable efficiency of selection. Where heritabilities are high the most effective programme for genetic improvement of the trait would be mass selection of those individuals exhibiting the desirable trait with little attention being given to ancestry, sibs and other collateral relatives and to progeny tests. Where heritabili-

176

Reproduction

and Breeding heritability

of selected traits in some domestic livestock

Table 4.6

The approximate Trait

Type of livestock

Approximate heritability (/, h*)

Dairy cattle

Butterfat percentage in milk Protein percentage in milk Butterfat production Milk production Reproductive performance Score for tenderness of meat Dressing percentage of carcase Bone percentage in carcase Heart girth measurement Daily liveweight gain Efficiency of feed conversicn Birth weight Weaning weight Daily liveweight gain Staple length of wool No. crimps per unit length in wool Birth weight Weaning weight _______ Score for leanness of meat Body length Back-fat thickness Daily liveweight gain Efficiency of feed conversion Thickness of eggshell Egg production Age at first lay Hatchability of eggs Viability of eggs Fertility

50-60 50-60 20-30 20-30 O-10 55-65 55-65 45-55 45-55 40-50 35-45 35545 20-25 50-60 40-50 40-50 25-35 25-35 65-75 45-55 45-55 40-50 35-40 35-45 20-30 15-25 IO-15 lo-15 O-5

Beef cattle

Sheep

Pigs

Poultry

Note: The majority of the estimates have been made in temperate-zone countries.

ties are low, selection should include some form of progeny test and be based on ancestry and the performance of close relatives. Selection acts by allowing selected individuals to contribute more traits to the next generation than other individuals in the same population. In fact the unselectedindividuals will have no influence on the next generation as they will not be allowed to breed. The rate at which selection can improve a population depends upon its overall accuracy, its level of intensity and the interval between generations. Accuracy in selection dependsupon how well the phenotype reflects the genotype and this can be improved by keeping as many records as is economically

Animal breeding practices

177

practicable. For example, the heritability of milk yield estimated from a single lactation record of a milking cow may be 0.25, whereas when two, three and four lactations are included in the estimate the heritabilities may be 0.33,0*37 and 040, respectively. Stressmust, however, be placed on the economic practicability of record-keeping. Multiple and complicated records do not necessarilyprovide additional information as to how accurately the phenotype reflects the genotype. The intensity of selection dependsto a very large extent on suitable replacement animals being available. Thus improvements in overall fertility and mortality improve the possibilities for intensifying selection. The interval between generationsdependsupon the interval betweenbirth and sexual maturity and the fertility rate. Thus any improvements in the age at first parturition and in overall fertility will help to decrease the interval between generations and thus increase selection pressure. Four major methods of selection are used in practice: individual or mass selection, pedigree selection, selection on the basis of collateral relatives and progeny testing. This is a most valuable method where a trait is highly inherited and where it can be observed in both sexes.In order to compare animals from different herds and therefore from slightly different environments the animals to be tested must be brought together under the sameconditions of feeding and management. An arrangement of this type is called a performance test. It can, of course, only be used for selecting for traits that can be measured in the live animal, such as growth rate and efficiency of feed conversion. However, the recent development of ultrasonic carcase measurementshas meant that some carcasemeasurementscan also be made, particularly in pigs. The major use of the performance test is for the selection of suitable breeding males.
Individual or mass selection and performance testing.

This is a method of selection based on the performance of ancestors. It was, until recent times, the major method by which selection was practised and it spawneda network of pedigree herd societies in the temperate zone. The methods can be of value only if the pedigree information is complete. Today, pedigree information is most useful when no data are available for the individual animal, either because it is too young or becausethe expression of a trait is sex-linked. The principal use of a pedigree in animal-breeding practice is to avoid too close an inbreeding ratio.
Pedigree selection.

This is most useful when family size is large, when traits are highly inherited, when there is a close genetic relationship between members of the family and when the mean generation interval is short. It is therefore obviously of more use in selecting for productive traits in poultry than in cattle.
Selection on the basis of collateral relatives.

178

Reproduction

and Breeding

Progeny testing.

The assessment the breeding value of an animal on of the basis of the performance of its offspring is known as a progeny test. As in most livestock speciesmales produce many more offspring during their lives than females,progeny tests are usually applied to males. It is a particularly valuable method to employ where a trait such as milk production is not measurablein mature animals of both sexes,where the heritability of a trait is low, where the breeding unit is large and where the increase in the generation interval, implicit in the method, is not too pronounced. Selection accuracy is increased at the expense of selection intensity and an increase in the interval between generations. Progeny testing is widely used in order to improve milk production in the temperate zone, but so far this method of selection has not been used to any major extent in the tropics. Aids to selection This method of breeding is now used for every speciesof domestic livestock. Nevertheless,its usefulnessis not unlimited and it can be practised successfully only under quite specific practical conditions. It is generally acceptedthat the benefits to be derived from AI are that it allows the maximum exploitation of the best sires, the fullest use of a selectedsire and a consequentreduction in the total number of sires that have to be maintained. Thus when AI is properly organized there can be a real reduction in breeding costs; AI also minimizes the spread of venerealand other diseases. AI demandsthat the farmer should closely As observehis female stock, it also probably improves general standards of management. Certainly it is a method by which livestock owners and livestock extension agents are brought into closer contact with each other and it probably stimulates a general increasing interest in livestock improvement. It is a method that is of particular utility to livestock farmers who wish to use different breeds of sires simultaneously, and is likely to bz economic in areas where there are a large number of smallholders, the majority owning one male and a limited number of female stock. It can also be of the greatest use where it is desired to import exotic livestock for crossbreeding and/or upgrading purposes, and where it is doubtful whether exotic sires will thrive. As AI involves the close handling of animals it cannot be undertaken on an economic scalewhere the livestock are too wild or too dispersed on the holding. Nor is it normally economic to service widely separated farms, particularly where roads and telephone communications are poor or non-existent. There are also some special technical problems for AI in the tropics. In most countries the low level of farm recording has been a major handicap in the testing of AI bulls. Obviously little is to be gained and considerable harm may be caused if the sires used in an AI programme
Artificial insemination.

Animal breeding practices

179

are unproven and not known to be superior to most local sires. If they are not proven they could in fact be inferior to some local bulls that would otherwise be used for natural service. Another problem is that many livestock-owning peoplesin the tropics rear all their male animals to maturity for meat purposes,so that the useof AI does not reduce their costs in bull maintenance. Perhaps the most serious technical difficulty is that in the females of many tropical breeds-in particular in the females of the Zebu or humped cattle breeds- their heat period is short in duration and often difficult to identify as it usually occurs at night. Under these circumstances many heat periods may be missed and this seriously reducesthe efficiency of the AI operation. Short and silent heats are also very frequent in the females of exotic dairy breeds managed in the tropics, and in consequenceAI in many exotic dairy herds has tended to become an uneconomic operation. It is apparent that the organization of an AI service is not necessarily the ultimate answer to livestock-breeding problems in any tropical country. In fact it may be an inefficient method of using scarce resources. Technological improvements in AI, such as the introduction of deepfrozen semen,may have improved the possibilities of importing the semen of exotic breeds into the tropics and may have solved some of the problems posed by scattered holdings and poor communications, but these new techniques require even more equipment and highly skilled manpower than the old and due consideration should be given to all the relevant factors before they are introduced.
Contemporary comparison test. This is probably the most effective selec-

tion method for dairy cattle that has been evolved to date. A sample of the averagemilk yield of unselected heifers in a number of herds under test is compared with that of contemporaries sired by other bulls. The comparisons are between heifers of approximately the same age and sexual m