The Pineapple

Botany, Production and Uses

 The Pineapple

Botany, Production and Uses

Edited by

D.P. Bartholomew

R.E. Paull

and

K.G. Rohrbach

University of Hawaii at Manoa

Honolulu
USA

CABI Publishing
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Library of Congress Cataloging-in-Publication Data

The pineapple : botany, production, and uses / edited by D.P.
Bartholomew, R.E. Paull, and K.G. Rohrbach.
p. cm.
Includes bibliographical references (p. ).
ISBN 0-85199-503-9
1. Pineapple. I. Bartholomew, D. P. (Duane Phillip), 1934- .
II. Paull, Robert E. III. Rohrbach, K. G. (Kenneth G.)
SB375.P47 2002
634.774--dc21

2002005741

ISBN 0 85199 503 9

Typeset by Columns Design Ltd, Reading

Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn
 Contents

Contributors vii

Preface ix

1. History, Distribution and World Production 1

Kenneth G. Rohrbach, Freddy Leal and Geo Coppens d’Eeckenbrugge
2. Morphology, Anatomy and Taxonomy 13
Geo Coppens d’Eeckenbrugge and Freddy Leal
3. Breeding and Variety Improvement 33
Y.K. Chan, G. Coppens d’Eeckenbrugge and G.M. Sanewski
4. Biotechnology 57
M.K. Smith, H.-L. Ko, S.D. Hamill, G.M. Sanewski and M.W. Graham
5. Crop Environment, Plant Growth and Physiology 69
Eric Malézieux, Francois Côte and Duane P. Bartholomew
6. Cultural System 109
Anthony Hepton
7. Plant Nutrition 143
Eric Malézieux and Duane P. Bartholomew
8. Inflorescence and Fruit Development and Yield 167
Duane P. Bartholomew, Eric Malézieux, Garth M. Sanewski and Eric Sinclair
9. Pests, Diseases and Weeds 203
Kenneth G. Rohrbach and Marshall W. Johnson
10. Postharvest Physiology, Handling and Storage of Pineapple 253
Robert E. Paull and Ching-Cheng Chen
11. Processing 281
Anthony Hepton and Aurora Saulo Hodgson

Index 291

The colour plates can be found following page 166

v
 Contributors

D.P. Bartholomew, Department of Tropical Plant and Soil Science, CTAHR, University of
Hawaii at Manoa, 3190 Maile Way, Honolulu, HI 96822, USA.
Y.K. Chan, MARDI, GPO Box 12301, Kuala Lumpur, 50774, Malaysia.
C.-C. Chen, Department of Horticulture, National Chung Hsing University, 250 Kuo Kuang
Road, Taichung, Taiwan, ROC.
G. Coppens d’Eeckenbrugge, CIRAD–FLHOR/IPGRI, AA 6713, Cali, Colombia.
F. Côte, CIRAD, BP 5035, 34032 Montpellier Cedex 01, France.
M.W. Graham, Queensland Agricultural Biotechnology Centre, Gehrmann Laboratories,
Level 4, The University of Queensland, St Lucia, Qld 4072, Australia.
S.D. Hamill, Maroochy Research Station, PO Box 5083, SCMC, Nambour, Qld 4560,
Australia.
A. Hepton (retired), Dole Food Company, 5795 Lindero Canyon Rd, Westlake Village, CA
91362, USA.
A.S. Hodgson, Tropical Plant and Soil Sciences, CTAHR, University of Hawaii at Manoa,
3190 Maile Way, Honolulu, HI 96822, USA.
M.W. Johnson, Plant and Environmental Protection Sciences, CTAHR, University of Hawaii
at Manoa, 3190 Maile Way, Honolulu, HI 96822, USA.
H.-L. Ko, Maroochy Research Station, PO Box 5083, SCMC, Nambour, Qld 4560, Australia.
F. Leal, Universidad Central de Venezuela, Facultad de Agronomía, Maracay, Aragua,
Venezuela, and IPGRI, AA 6713, Cali, Colombia.
E. Malézieux, Centre de Coopération Internationale en Recherche Agronomique pour le
Developpement (CIRAD), TA 179/01 Avenue Agropolis, 34398 Montpellier Cedex 5,
France.
R.E. Paull, Department of Tropical Plant and Soil Sciences, College of Tropical Agriculture
and Human Resources, University of Hawaii at Manoa, 3190 Maile Way, Honolulu, HI
96822-2279, USA.
K.G. Rohrbach, Plant and Environmental Protection Sciences, CTAHR, University of Hawaii
at Manoa, 3190 Maile Way, Honolulu, HI 96822, USA.
G.M. Sanewski, Queensland Horticulture Institute, PO Box 5083, SCMC, Nambour, Qld
4560, Australia.
E. Sinclair, Golden Circle Ltd, PO Box 150, Nundah, Qld 4012, Australia.
M.K. Smith, Maroochy Research Station, PO Box 5083, SCMC, Nambour, Qld 4560, Australia.

vii
 Preface

Pineapple is the third most important tropical fruit in world production after banana and cit-
rus. Seventy per cent of the pineapple produced in the world is consumed as fresh fruit in the
country of origin. This worldwide production has developed since the early 1500s when
pineapple was first taken to Europe and then distributed throughout the world’s tropics.
International trade is dominated by a few multinational companies that have developed the
infrastructure to process and market pineapple. The most famous variety in world trade is
Cayenne Lisse (‘Smooth Cayenne’), which was introduced to Europe from French Guiana.
Many clones of this cultivar are known and a number of other cultivars are only grown
regionally for consumption in the country of production or represent a minor component of
the fresh-fruit export market.
Drought tolerance and the ease of transport of vegetative propagules facilitated the wide
diffusion of pineapples around the world. However, the relatively short shelf-life of fresh
pineapple fruit limited early commercial trade to relatively short transportation routes or a
preserved form of the fruit. The commercial processing of pineapple started in Hawaii at the
end of the 19th century and Hawaii was a world leader in pineapple research and processing
until after the Second World War. The invention and refinement of the automatic peeling and
coring machine by Henry Ginaca in Hawaii, between 1911 and 1919, allowed the develop-
ment of a large-scale economically viable canning industry. This was paralleled by the major
expansion of pineapple production and expanded research on the crop.
This book is the outcome of discussions between the editors over the past 10 years. The
most recent comprehensive book on pineapple was published in French in 1983 and an
English version with minor additions was published in 1987. That book is now out of print.
The need for such a book was also seen as opportune, as many of the individuals involved in
the research associated with the international expansion of pineapple production and mar-
keting were retiring. This book is an attempt to put in one place a synopsis of the worldwide
research findings and commercial practices, mainly for ‘Smooth Cayenne’, but mention is
made of other cultivars where they differ from ‘Smooth Cayenne’ and where published infor-
mation was available.
The book integrates genetics, breeding, physiology, pathology, handling and processing

ix
x Preface

with commercial practices. Successful commercial production requires not only an integra-
tion of these findings, but their management in different biotic and abiotic environments and
with different market needs. The information contained in the book should be of use to all
interested in pineapple production and utilization.
Many have contributed to this text. The editors wish to thank the individual chapter
authors for their patience and understanding. Special thanks are due to Valent BioSciences
Corporation, California, for a generous donation that allowed us to include the eight pages of
colour plates. Tania Ishimaru worked for many hours typing and editing and checking the
individual chapters, and her help is sincerely appreciated by the editors. All errors and omis-
sions are the responsibility of the editors.
In closing, we would like to dedicate this book to the many scientists who spent much of
their life working on pineapple. Cristos Sideris, Walter Carter, J.L. Collins, Claude Py and
Eloys Giacommelli warrant special mention for their many contributions to the advancement
of pineapple science and culture. Many scientists at the Pineapple Research Institute of
Hawaii contributed much to our knowledge of pineapple, but their work was done for pri-
vate growers and so remains largely unpublished. We hope progress will continue to be
made in understanding this unique and fascinating crop.
Duane Bartholomew
Kenneth Rohrbach
Robert E. Paull
Honolulu, 2002
 1 History, Distribution and
World Production

Kenneth G. Rohrbach,1 Freddy Leal2 and

Geo Coppens d’Eeckenbrugge3
1Plant
and Environmental Protection Sciences, CTAHR, University of Hawaii at
Manoa, 3190 Maile Way, Honolulu, HI 96822, USA; 2Universidad Central de
Venezuela, Facultad de Agronomía, Maracay, Aragua, Venezuela; 3IPGRI/CIAT,
AA 6713, Cali, Colombia

Early History had done the same with pineapple long

before Colón. Other evidence points to the
Prior to the discovery of the pineapple fruit antiquity of pineapple cultivation. Thus, the
by Cristobal Colón (Christopher Columbus) names ‘nanas’ and ‘ananas’ were extensively
on 4 November 1493 (Morrison, 1963), the used throughout South America and the
fruit was already a stable component of the Caribbean. Early European explorers
vegetative-crop complex and in the diet of observed a high degree of domestication and
native Americans in the lowland tropics selection exhibited in the pineapples they
(Laufer, 1929). The European explorers were found. The Amerindians easily distinguished
impressed by this large and delicious fruit landraces from the wild types and had
and often mentioned and described it in developed a thorough knowledge of the crop
their chronicles. These early reports indicate agronomy, including its production cycle.
that domesticated pineapple was already Specifically adapted landraces (e.g. the
very widely distributed in the Americas Andean ‘Perolera’ and ‘Manzana’) were
(Orinoco, Amazon, coastal Brazil around Rio found with variation in fruit yield and qual-
de Janeiro) and the Caribbean prior to the ity. Five additional centuries of work by tal-
arrival of Columbus (Collins, 1960). In some ented horticulturists and modern plant
cases, the Europeans themselves could have breeders have not added significantly to the
contributed to pineapple dispersion in the variety of domesticated types (Leal and
continent. Indeed, the many plants (e.g. Coppens d’Eeckenbrugge, 1996; Coppens
lemon, orange, sugar cane, banana and plan- d’Eeckenbrugge et al., 1997).
tain) introduced by Colón from the Canary In addition to the fresh fruit, the native
Islands since his second voyage were distrib- Americans used pineapple for the prepara-
uted by the natives throughout tropical tion of alcoholic beverages (pineapple wine,
America in less than two decades, as evi- chicha and guarapo), for the production of
denced by banana and plantain cultivation fibre, and for medicinal purposes, as an
in Puerto Bello in 1503 (Colón, 1506). On the emmenagogue, abortifacient, antiamoebic
other hand, if the natives dispersed these and vermifuge and for the correction of
new crops so quickly, they very probably stomachal disorders, and for the poisoning

© CAB International 2003. The Pineapple: Botany, Production and Uses

(eds D.P. Bartholomew, R.E. Paull and K.G. Rohrbach) 1
2 K.G. Rohrbach et al.

of arrowheads. Most of these medicinal uses most important variety in world trade; many
are related to the proteolytic enzyme brome- others are only grown regionally for local
lain of the pineapple (Leal and Coppens consumption. Both Smooth Cayenne and
d’Eeckenbrugge, 1996). The native Singapore Spanish can be called true culti-
Americans also domesticated the curagua, a vars (see Coppens d’Eeckenbrugge and Leal,
smooth-leaved type with a higher yield of Chapter 2, this volume).
long and strong fibres, and used it for mak-
ing nautical and fishing-lines, fishing nets,
hammocks and loincloths (Leal and Amaya, Development of the Pineapple Trade
1991). There is still a small traditional indus-
try based on pineapple fibre in Brazil (Leme Drought tolerance and the ease of transport
and Marigo, 1993) and even in the of vegetative propagules facilitated the wide
Philippines, where ‘piña cloth’ was men- diffusion of pineapples around the world.
tioned as early as 1571 (Collins, 1960; However, the relatively short shelf-life of
Montinola, 1991). fresh pineapple fruit limited early commer-
From the early 1500s, the pineapple fasci- cial trade to relatively short transportation
nated the Europeans, who introduced and routes or some form of preservation. Jams
grew it in greenhouses. The first successful and sweets made in the West Indies, Brazil
greenhouse cultivation was by Le Cour, or and New Spain (Mexico) were the first com-
La Court, at the end of the 17th century near mercial products of pineapple (Thévet, 1557;
Leyden. He published a treatise on pineap- Acosta, 1590; Loudon, 1822). In the early
ple horticulture, including ‘forcing’ the 19th century, fresh pineapples were sent
plants to flower. Pineapple plants were dis- from the West Indies to Europe attached to
tributed from The Netherlands to English the entire plant, which lowered the price in
gardeners in 1719 and to France in 1730 the European markets and led to a decline in
(Gibault, 1912). As pineapple cultivation in European glasshouse production (Loudon,
European greenhouses expanded during the 1822). Commercialization during the mid-
18th and 19th centuries, many varieties were 19th century developed based on the short-
imported, mostly from the Antilles. Griffin est trade routes rather than an optimum
(1806) described ten of them and considered pineapple production environment.
most of the others as useless and their culti- Production in Florida, the Bahamas, Cuba
vation cumbersome. Others have described and Puerto Rico supplied the North
numerous varieties (Loudon, 1822; Munro, American market and the Azores the
1835; Beer, 1857). The now famous variety European market. The Azores maintained
Cayenne Lisse (‘Smooth Cayenne’) was their monopoly of the European fresh-fruit
introduced from French Guiana by Perrotet market until after the Second World War,
in 1819 (Perrotet, 1825). With the notable when production shifted to Africa (Py et al.,
exceptions of ‘Smooth Cayenne’ and 1987).
‘Queen’, most of these early varieties disap- Commercial processing of pineapple
peared as commercial cultivation in Europe started in Hawaii at the end of the 19th cen-
declined and pineapple fruit was imported tury. The invention and refinement of the
from the West Indies. automatic peeling and coring machine by
’Smooth Cayenne’ and ‘Queen’ were Henry Ginaca, a Hawaiian Pineapple
taken from Europe to all tropical and sub- Company (Dole) employee, between 1911
tropical regions (Fig. 1.1; Collins, 1951). The and 1919 allowed the development of a
Spaniards and Portuguese dispersed other large-scale economically viable canning
varieties, including ‘Singapore Spanish’, to industry. This was paralleled by a major
Africa and Asia during the great voyages of expansion of pineapple production. The 1919
the 16th and 17th centuries. However, the ginaca peeled and cored up to 65 pineapples
diversity of these varieties is still negligible min−1. A 1925 model, also developed by the
compared with the variation found in Hawaiian Pineapple Company, processed
America. ‘Smooth Cayenne’ is by far the 90–100 pineapples min−1. No additional sig-
 History, Distribution and World Production 3

England
Netherlands
and Belgium
1897
Hawaii
1860 1835
1923
Florida 1863
1885
1926 Azores ?
Taiwan
France
1947 Haiti 1820
Philippines 1946 Cuba
Mexico 1930
Ceylon
1893 1930
1858
French Africa
Fiji Guiana 1895

South
Africa
South
America
Australia

Fig. 1.1. Distribution of the ‘Smooth Cayenne’ cultivar (after Collins, 1951).

nificant improvements have been made on fresh fruit to Japan (Py et al., 1987). Today,
this machine since 1925 (Anon., 1993). Other the canned-product market remains very
important canning operations started around important but the value of the international
the same period in South-East Asia fresh-fruit market is rapidly increasing.
(Malaysia in 1888, Taiwan in 1902,
Philippines in 1920), Australia, South Africa,
the Caribbean (Martinique, Cuba and Puerto World Production and Trade
Rico) and Kenya. The Second World War
ruined the South-East Asian industry and Pineapple is now the third most important
destroyed international trade. Hawaii gained tropical fruit in world production after
a strong leadership position that lasted until banana and citrus. The processing of pineap-
the development of new competitors (Côte ple has made the fruit well known through-
d’Ivoire and the Philippines, followed by out the temperate developed world. Major
Thailand) between 1950 and the early 1960s. pineapple products of international trade are
In these same years, refrigerated sea trans- canned slices, chunks, crush (solid pack) and
portation developed and diminished the juice and fresh fruit (Fig. 1.2). International
importance of proximity to the market. trade is dominated by a few multinational
Hawaii, West Africa (mainly Côte d’Ivoire) companies that have developed the infra-
and Taiwan shifted part of their production structure to process and market pineapple.
to the fresh-fruit market, exporting, respec- Thailand and Indonesia are, to a degree,
tively, to the North American, European and exceptions, with small local processing oper-
Japanese markets. Philippine production ations. Despite the significance of canned
expanded greatly in the 1970s, exporting pineapple in international trade, approxi-
canned products and significant quantities of mately 70% of the pineapple produced in the
4 K.G. Rohrbach et al.

world is consumed as fresh fruit in the coun- (293,000 t) and Kenya (280,000 t). The value
try of origin (Loeillet, 1997). Important pro- of these statistics is relative to their accuracy.
ducing countries, such as Brazil, India, Thus, the joint use of statistics for production
China, Nigeria, Mexico and Colombia, pro- and planted areas allows yield estimations
duce fruit primarily for their own fresh-fruit ranging from a high (and unlikely) of 48 t
markets and canning is a minor industry. ha−1 for Cuba to a low of 7 t ha−1 for
Statistics on world pineapple production Indonesia (Table 1.1). Indeed, several coun-
are collected by the Food and Agriculture tries are thought to give rough estimates,
Organization of the United Nations (FAO). which may explain the surprising official
According to FAO statistics (Baker, 1990; production of countries such as Nigeria and
Anon., 2002), total pineapple production was the low correlation between yields and the
approximately constant in the 1999–2001 probable level of technology and inputs used
period, with a mean world production for in the production system. In addition, FAO
these 3 years of 13,527,149 metric tonnes (t). statistics do not separate fresh fruit from
World production has more than tripled dur- processed pineapple or export from local
ing the past 30 years (3,833,137 t in 1961 to consumption.
13,738,735 t in 2001). The leading pineapple- Pineapple produced in Thailand and the
producing countries are Thailand with Philippines dominates world trade. The for-
2,311,332 t, the Philippines with 1,520,715 t mer country processes approximately 1.6
and Brazil with 1,504,493 t (means 1999–2001). million t of its total production of 2 million t.
China (1,181,169 t), India (1.1 million t), and, Thailand’s pineapple is produced on almost
Nigeria (800,000 t) follow. Nigerian statistics 100,000 ha of small farms of 1–5 ha (Anupunt
announced year after year are a surprisingly et al., 2000). In contrast to Thailand, produc-
constant 800,000 t. Other producers exceeding tion and marketing in the Philippines is
250,000 t are Mexico (535,000 t), Costa Rica almost exclusively run by multinational cor-
(475,000 t), Colombia (360,000 t), Indonesia porations using large plantation production
(300,000 t), Venezuela (300,000 t), USA systems. Export and marketing from the

SPEARS
SUGAR AND
CATTLE FEED
Fruit
core Core CRUSH
Fruit hole
skin TITBITS

CHUNKS

FRUIT SLICE

Pineapple Longitudinal
fruit and fruit Fruit
crown section cylinder
JUICE
FRESH FRUIT
JUICE
CONCENTRATE

Fig. 1.2. Pineapple products.

 History, Distribution and World Production 5

Table 1.1. Pineapple production (t), ha harvested and yields (t ha−1) for 2001 by country
(Anon., 2002).

Country Production (t) Harvested (ha) Yield (t ha−1)

Thailand 2,300,000 97,300 24

Philippines 1,571,904 45,000 35
Brazil 1,442,300 59,238 24
China 1,284,000 57,700 22
India 1,100,000 80,000 14
Nigeria 881,000 115,000 8
Mexico 535,000 12,500 43
Costa Rica 475,000 12,000 40
Colombia 360,000 9,000 40
Indonesia 300,000 42,000 7
Venezuela 300,000 15,000 20
USA 293,000 8,130 36
Kenya 280,000 8,500 33
Côte d’Ivoire 225,675 5,200 43
South Africa 145,441 6,200 23
Australia 140,000 3,000 47
Dominican Republic 136,862 5,500 25
Malaysia 130,000 7,000 19
Guatemala 101,287 3,710 27
Honduras 70,000 3,900 18
Cameroon 42,000 4,000 11
Martinique 20,800 484 43
Swaziland 19,680 600 33
Cuba 19,000 400 48
Cambodia 16,500 1,600 10
Puerto Rico 15,000 500 30

Philippines of both processed and fresh fruit inflation has increased as production was
are frequently handled with other products, gradually shifted from processed to fresh
such as bananas. The large scale of produc- fruit for the domestic market (Rohrbach,
tion, high level of technology and low labour 2000). Taiwan (Lin and Chang, 2000), like
costs make competition with production, Hawaii, has shifted from a dominant pro-
processing and marketing of both canned cessing industry to a domestic and export
and fresh fruit from Thailand and the fresh-fruit market.
Philippines very difficult for the smaller pro- World trade in pineapple mainly consists
ducing countries. Australia and South Africa of processed products. World exports of
market canned and fresh fruit almost exclu- canned pineapple doubled between 1983 and
sively within the country and remain com- 1992, passing 1 million t and representing a
petitive because of efficient production and value of more than US$600 million. Asian
processing (Sanewski and Scott, 2000) and countries have been the primary suppliers,
international trade barriers. While Hawaii increasing their share from 69 to 85%, while
was the centre of world processing and tech- Africa’s share has decreased from 25 to 10%.
nology in the first half of the 20th century, its Leading countries are now Thailand (315,000
proportion of production has declined t), the Philippines (209,000 t), Indonesia
steadily as production and competition from (95,000 t), Kenya (84,000 t) and Malaysia
Thailand and the Philippines have risen (Fig. (44,000 t). The European Union imports
1.3). However, the value of the Hawaiian 450,000 t, a 2.4-fold increase between 1983
pineapple industry (US$) unadjusted for and 1993. The USA and Canada import
6 K.G. Rohrbach et al.

2500 Thailand
Production and processing (t) ( 1000)

Brazil
2000
Philippines

USA
1500

1000

500

1961 1971 1981 1991

Years

Fig. 1.3. Shifts in Hawaiian and multinational foreign production and processing industries (Anon., 1998a;
Baker, 1990).

380,000 t, representing a replacement for the USA, annual per capita consumption of
decreasing Hawaiian production (Loeillet, fresh pineapple fruit has gone from 0.3 to 0.9
1995; Anon., 1998a). kg. This is still very low when compared
The market for concentrated pineapple with the approximately 5 kg consumption of
juice, especially frozen concentrate, has also processed pineapple over the past 25 years
increased. Estimated at 40,000 t in 1983, it (Fig. 1.6) and with the consumption of other
increased to 167,000 t in 1993 (representing fruits such as bananas, apples and oranges
then up to US$400 million), to reach 215,000 t (Fig. 1.7). The European market is mainly
in 1993. Supply is dominated by Thailand supplied by Côte d’Ivoire, with significant
and the Philippines. The Philippines is also amounts transshipped through France to
largely dominant for the smaller market for several other European countries (Fig. 1.8;
single-strength juice (70,000 t). The USA and Aldrich, 1984; Anon., 1998b). European
Canada (90,000 t) are the major importers of countries such as the Netherlands and
concentrated juice, with Europe (118,000 t) Belgium obtain fresh pineapple from several
second (Loeillet, 1994). different countries, including Costa Rica, as
Per capita consumption of pineapple juice well as the Côte d’Ivoire through France. In
in the USA is essentially static at between 1 Europe, per capita consumption of fresh
and 1.3 l year−1 which contrasts with pineapple is highest in France and in 1984
increases of both orange- and apple-juice was approximately equal to the current US
consumption (Fig. 1.4). fresh-pineapple consumption (Aldrich,
The international fresh-pineapple market 1984). The principal South-East Asian fresh-
(about 670,000 t) is dominated by Costa Rica, fruit export market is Japan, which is domi-
the Philippines and the Côte d’Ivoire. The nated by the Philippines. Taiwan also
North American market is primarily sup- supplies significant amounts. China,
plied by Costa Rica and Hawaii (Fig. 1.5). In Indonesia and Hawaii occasionally supply
 History, Distribution and World Production 7

25

20
Juice consumption (l)

15
Orange

10 Apple

Pineapple
5

1975 1980 1985 1990 1995

Year

Fig. 1.4. Per capita consumption of orange, apple and pineapple juice in the USA (Putnam and Allshouse,
1999).

small amounts. Imports of fresh pineapple transported chilled at 0–1°C or shipped whole
into Japan have declined somewhat in recent without the crown to large metropolitan cen-
years (Fig. 1.9) (Anon., 1997). In contrast to tres and processed just before retail sales. The
Europe and the USA, Brazil’s consumption shelf-life of this product is limited to 1–3
of fresh pineapple is approximately 11 kg per weeks unless the product is actually frozen.
capita year−1 (Reinhardt and Souza, 2000). The chilled fresh-cut product addresses con-
Chilled fresh-cut fruit pineapple packed as sumer demand for ready-to-eat foods that do
spears or chunks in sealed plastic bags for not require any preparation time. Industry
retail sale is a relatively new product. Fruit sources estimate that the market for vacuum-
may be processed at the production site and packed fresh-cut pineapple in Japan will soon

Costa Rica Dom Republic Ecuador Honduras Mexico

250,000
US imports (t)

200,000

150,000

100,000

50,000

1981 1986 1991 1996

Year

Fig. 1.5. Countries supplying fresh pineapple to the US market (Anon., 1998b). Dom, Dominican.
8 K.G. Rohrbach et al.

Per capita consumption (kg) 8

4
Fresh
2

Processed
0

1970 1975 1981 1986 1991

Year

Fig. 1.6. Per capita consumption of fresh and processed pineapple in the USA (Putnam and Allshouse,
1999).

14.0

12.0
Per capita consumption (kg)

Bananas
10.0
Apples
8.0

Oranges
6.0

4.0
Pineapples
2.0

0.0
1970 1973 1976 1979 1982 1985 1988 1991 1994
Years
Fig. 1.7. Per capita consumption of bananas, apples, oranges and pineapple in the USA (Putnam and
Allshouse, 1999).

approach 20% of the total fresh-pineapple (about 95%), comes from the cultivar
fruit market for Japan (Anon., 1999). In ‘Smooth Cayenne’. ‘Queen’ is present in
Hawaii, it is estimated that chilled fresh-cut small specific niches of high-quality and
pineapple represents up to 10% of the total expensive fresh fruit (Loeillet, 1996). The
fresh-fruit market. High-pressure processing increasing importance of fresh pineapple in
has recently been used to extend the shelf-life the temperate markets is putting pressure on
of chilled fresh-cut fruit (Aleman et al., 1994). the multinational corporations to switch to
Commercialization of this process will be cultivars that are superior to ‘Smooth
dependent on the costs versus benefits of Cayenne’ as fresh fruit. As industries shift to
high-pressure processing equipment. domestic fresh-fruit markets because of com-
Most of the world production (about petition in processed fruit, ‘Smooth Cayenne’
70%), and most of the canned pineapple does not provide today’s consumer with the
 History, Distribution and World Production 9

180,000

160,000 Ivory Coast

Costa Rica
140,000
Honduras
Fruit supplied (t)

120,000
Ghana
100,000
Dominican Rep.
80,000

60,000

40,000

20,000

0
1970 1975 1980 1985 1990 1995
Year
Fig. 1.8. Major countries supplying fresh fruit to the European market (Anon., 1998b; Loeillet, 1994).

150,000
Japanese imports (t)

100,000

50,000

0
1988 1989 1990 1991 1992 1993 1994 1995 1996 1997
Year

Fig. 1.9. Metric tons of pineapple imported into Japan (Anon., 1997, 1999).

best fruit quality (Paull, 1993; Sanewski and production have been evaluated and used as
Scott, 2000). Taiwan, Hawaii, Malaysia, animal feeds (Wayman et al., 1976; Olbrich
Australia, Cuba, Brazil and the French are and Al, 1977; Stanley and Ishizaki, 1979). In
putting resources into developing cultivars many countries, feed tolerances must be
specifically for domestic fresh-fruit markets. established for pesticides used during pro-
A first result of these efforts has been the duction in order for by-products to be used
recent and successful introduction of a low- as animal feeds. The requirements for toler-
acid cultivar by Del Monte from Costa Rica ances affect the economic viability of by-
into the European and American markets product use for feeds, even though a
(Malezieux, 2000). disposal cost exists for cannery wastes.
Additionally, because of the low nutrient val-
ues of pineapple by-products, animal weight
By-products gains may not be economical when higher-
quality feeds are available.
Portions of the pineapple plant and process- Pineapple fibre is considered to be more
ing wastes, in the form of shell and core delicate in texture than any other vegetal
materials, and centrifuged solids from juice fibre. About 60 cm long, white and creamy
10 K.G. Rohrbach et al.

and lustrous as silk, it easily takes and Bromelain was originally only extracted
retains dyes. Numerous tests in Brazil, from Hawaiian pineapple stems but now is
Florida, India and the Philippines have manufactured in Taiwan, Thailand, Brazil
shown its exceptional resistance to salt, and Puerto Rico. The variability in the com-
vapour and traction (Correa, 1926; mercially produced product and its multiple
Montinola, 1991). However, while small cot- ingredients have limited successful develop-
tage industries exist for speciality uses of ment. Pineapple bromelain has been used
pineapple fibre from particular cultivars, commercially as a meat-tenderizing enzyme
numerous attempts at commercial produc- and as a nutraceutical. Attempts have been
tion as a subproduct of the fruit industry made to develop bromelain for pharmaceuti-
have failed. In fact, fibre quality and yield cal use. The complexity of the active compo-
are highly dependent on the cultivar, and nents of bromelain has limited
those of ‘Smooth Cayenne’ are among the pharmaceutical research. Bromelain has
weakest. In addition, cultural practices for shown the following activity: (i) interference
fruit production have detrimental effects on with the growth of malignant cells; (ii) inhi-
fibre characteristics. Pineapple fibre has also bition of platelet aggregation; (iii) fibrinolytic
been processed into paper, which shows action; (iv) anti-inflammatory processes; and
remarkable thinness, smoothness and plia- (v) skin débridement (Lotz-Winter, 1990).
bility (Collins, 1960; Montinola, 1991). Recent These biological properties of bromelain
studies have resulted in several patents on have potential therapeutic activity in: (i)
paper production and the development of tumour growth; (ii) blood coagulation; (iii)
low-density polyethylene composites inflammatory changes; (iv) débridement of
(Fujishige et al., 1977; Fujishige and Tsuboi, severe burns; and (v) enhancement of drug
1978; George et al., 1993, 1995). absorption (Taussig and Batkin, 1988).

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 2 Morphology, Anatomy and Taxonomy

Geo Coppens d’Eeckenbrugge1 and Freddy Leal2

1CIRAD–FLHOR/IPGRI, AA 6713, Cali, Colombia; 2Universidad Central de Venezuela,
Facultad de Agronomía, Maracay, Aragua, Venezuela, and IPGRI, AA 6713,
Cali, Colombia

Morphology structures to be distinguished are the stem,

the leaves, the peduncle, the multiple fruit or
Ananas comosus is a herbaceous perennial of syncarp, the crown, the shoots and the roots
the Liliopsidae (monocotyledonous), whose (Fig. 2.1). The following description is
terminal inflorescence gives origin to a mul- mainly focused on the cultivated pineapple.
tiple fruit (sorose). After maturation of the It is partly based on the anatomical studies
first fruit, the plant develops new shoots of Krauss (1948, 1949a,b) and Okimoto
from axillary buds, so producing new (1948).
growth axes capable of producing another
fruit. The same plant may thus give a
sequence of various production cycles. In Stem
most commercial plantings, the plants are
not allowed to produce more than two to The pineapple stem is club-shaped, with a
three crops, due to a reduction in fruit size length of 25–50 cm and a width of 2–5 cm at
and uniformity. Then a new plantation must the base and 5–8 cm at the top. Its aerial part
be regularly established. This may be done is straight and erect, while the shape of the
with the same lateral shoots of the preceding earthed part depends on the material used
crop, or with other vegetative propagules, for planting. It is markedly curved when
such as the fruit crown, or, in many cultivars, coming from a slip, as the stems of these
slips produced along the peduncle. This veg- propagules are comma-shaped, less curved
etative reproduction is also dominant in wild when coming from a stem shoot and erect
pineapples, where, in addition to lateral when coming from a crown. Nodes can be
shoots, the crown and slips contribute to visualized by the leaf scars left after strip-
propagation as they resume rapid growth at ping the leaves from the stem. Internodes are
fruit maturity. The long peduncle then bends short (1–10 mm according to their position),
because of this mass and the crowns and so the whole rosette appears dense and com-
slips reach the ground and may root. Thus pact. Flattened shoot buds, 3–5 mm high and
most natural populations appear to consist about 5 mm wide, occur in the leaf axils. In
of a single clone, expanding as if propagat- the central portion of the stem, they are
ing by stolons. larger because of an increase in the size of
The adult plant is 1–2 m high and 1–2 m their prophyll (the first leaf of the shoot,
wide, and it is inscribed in the general shape which encloses it). A striking feature of the
of a spinning top. The main morphological pineapple stem is the presence of adventi-
© CAB International 2003. The Pineapple: Botany, Production and Uses
(eds D.P. Bartholomew, R.E. Paull and K.G. Rohrbach) 13
14 G. Coppens d’Eeckenbrugge and F. Leal

Crown

Axillary
crownlets

Slip

Stem shoot

Ground shoot
or sucker

Fig. 2.1. Main morphological structures of the pineapple plant.

tious roots breaking through the epidermis, little phloem. In this tissue, areas of non-vas-
and growing flattened and distorted, tightly cular tissue, or leaf gaps, are disposed at
wound around the stem, between the leaves. intervals, allowing leaf-trace bundles to pass
Their older portion is suberized. These aerial from the cortex into the stele. This vascular
roots rarely produce laterals. They are elon- cylinder is thicker and suberized at the stem
gated from a few millimetres in the subapical base. On the cortical side, a narrow layer of
region to 10 cm or more near the stem base. long, thin-walled cells bound it. Vascular
Thus, the underground portion of the stem is bundles are very numerous throughout the
covered with a tuft of adventitious fibrous stem but less so in the cortex than in the
roots. stele. The latter is mainly constituted of a
The stem (Fig. 2.2) constitutes a central compact parenchyma with abundant
cylinder, or stele, and a cortex, separated by starch. It contains large cells with raphides
a thin layer of vascular bundles produced by of calcium oxalate crystals. The cortex is
the dome-shaped apical meristem. The dense composed of a parenchymatous tissue,
network of vascular tissue separating cortex crossed by the isolated vascular bundles
and stele consists chiefly of xylem, with very going to the leaves, of the adventitious roots
 Morphology, Anatomy and Taxonomy 15

originating at the boundary with the central width, depending on the cultivar and ecolog-
cylinder and of circumferential small bands ical conditions. The apical ones are short and
of vascular tissue lying just above the leaf erect. The leaves are ensiform and, except for
attachment to the stem. The inner the young apical ones, broader at their base,
parenchyma of this cortex is also rich in which forms a non-chlorophyllous sheath
starch and contains raphide cells. Limiting around the stem. The blades then taper pro-
the stem externally is the epidermis, with gressively to a sharply pointed indurated tip.
peltate trichomes in the nodal regions. The constriction between the sheath and the
blade is more marked in certain wild pineap-
ples. Temporary stress during leaf growth
Leaves may cause variations in width or spininess,
or both, along the blade. Leaves are semi-
The sessile leaves enclose the stem on two- rigid, thanks to their crescent-shaped section.
thirds of its circumference. The phyllotaxy As in other bromeliads, this allows the plant
varies, being 5/13 in large-fruited cultivated to collect water in the rosette, where it can be
pineapples and 3/8 in small-fruited wild absorbed by the aerial roots present along
pineapples (Kerns et al., 1936). Leaf number the stem or through the epidermis of the
is variable between cultivars but generally sheath. The concave adaxial face is green or
around 40–80. The lower ones, originating dark green, with some anthocyanins, to dark
from the planting material or produced soon red or purple, according to the cultivar and
after planting, are smaller (5–20 cm) com- conditions. The abaxial side is convex, with a
pared with the younger ones, which can surface corrugated by longitudinal grooves.
reach more than 1.6 m in length and 7 cm in Both sides are covered by peltate trichomes,

A B

lg iar
vn

cvt
ltb cvt
rb
lb
vn
lbb lg
lg ltb

ear arb
iar
iar
ls

Fig. 2.2. A. Central region of the ‘Smooth Cayenne’ pineapple stem cut to show transverse, radial and
tangential views of gross external and internal structure. Tangential section cut at boundary between cortex
and stele. iar, internal, and ear, external portions of adventitious roots; cvt, circumferential band of vascular
tissue; vn, vascular network at the boundary between cortex and stele; lg, leaf gap; ltb, leaf-trace bundle; ls,
leaf scar; lbb, lateral branch bud; rb, tip of emerging root, commonly called a root ‘bud’. B. Diagrammatic
representation, showing portion of the cortex cut away to expose the vascular network at the boundary
between cortex and stele, and xylem elements from roots. arb, adventitious root base; cvt, circumferential
band of vascular tissue; lb, leaf base; lg, leaf gap; ltb, leaf-trace bundle. (After Krauss, 1948. © The
University of Chicago, all rights reserved.)
16 G. Coppens d’Eeckenbrugge and F. Leal

particularly the abaxial one, which is densely presence of large numbers of mitochondria
furfuraceous and silvery (see Fig. 5.7). The in the stalk cells give indirect evidence for an
leaf margins are usually thorny; however, important role in uptake of dissolved nutri-
certain cultivars are partially or totally iner- ents (Sakai and Sanford, 1980). Another
mous. In some smooth cultivars, the lower important role is to protect the plant from
epidermis is folded over the leaf edge and excessive transpiration and intense sunlight.
extended over the upper surface, so produc- More generally, the thick cuticle, the water-
ing a narrow silvery stripe, a trait called ‘pip- storage tissue, the disposition of the stomata,
ing’ by Collins (1960). the trichomes and the crassulacean acid
A section across the leaf (Fig. 2.3) shows metabolism (CAM) all contribute to the
successively a thick and smooth cuticle; a remarkable water economy of pineapple.
particular upper epidermis, which consists
of a single layer of cells, each containing a
silica body, orientated perpendicularly to the Roots
leaf axis, and rigidified by thick and undu-
lated lateral and inner cell walls; the hypo- Primary roots are only found in very young
dermis; the water-storage tissue, consisting seedlings. They die soon after germination
of various layers of thin-walled cells, which and are replaced by the adventitious roots.
accounts for a quarter to half of the leaf thick- These form a short and compact system at
ness, depending on the water status of the the stem base, with numerous strong roots
plant; the chloroplast-rich mesophyll, with and limited branching. Under ideal condi-
the vascular bundles, fibre strands and aerat- tions, the soil root system may spread up to
ing canals; and the lower hypodermis and 1–2 m laterally and 0.85 m in depth. The
epidermis, with the stomata arranged longi- number of roots produced after planting is
tudinally along the characteristic grooves of positively correlated with shoot weight, and
the abaxial leaf side, covered by numerous crowns produce more roots than do shoots.
trichomes, giving it a silvery appearance and The root internal anatomy is typical of
increasing reflectance. The fibre strands con- monocots, with, towards the centre: the epi-
fer a high tear resistance to the pineapple dermis, with root hair cells; the cortex com-
leaves. Stomatal density is rather low, about prising the exodermis, the outer cortex, with
80 stomata mm−2. The trichomes, present in sclerenchyma and aerating canals, and the
almost all the known bromeliads, are flat and inner cortex, with a lagunar parenchyma; the
shield-shaped, parallel to the leaf surface. endodermis, pericycle, vessels and pith (Fig.
They consist of a central disc of live cells, an 2.4). Their most characteristic trait is the
outer ring of dead cells and a pluricellular medullar structure given by the aerating
stalk arising from the epidermis and subepi- canals, formed by the tip-to-tip junction of
dermis. Like the stomata, they are nested in raphide cells in the outer cortex, and by the
small cavities along the bottom of the under- air lacunae formed by the disappearance of
side furrows, with their broad heads spread- thin-walled cell groups. Branch roots origi-
ing out to virtually cover the entire leaf nate in the pericyclic region of the main
surface. In many bromeliad species, the dead roots.
cells can absorb water and nutrients and the
stalk can carry them into the inner leaf tis-
sues. The shield also acts as a plug, closing Inflorescence and fruit
the concavity where the stalk is inserted and
protecting it from drying out. These tri- The peduncle and inflorescence develop
chomes function as one-way valves, playing from the apical meristem, the diameter of
an important role in the capacity of bromeli- which is suddenly increased until the initia-
ads to improve and maintain their water sta- tion of the peduncle (Kerns et al., 1936). The
tus (Benzing, 1980). However, the pineapple stage of inflorescence emergence is called
trichomes appear to be hydrophobic and do ‘red heart’ because of the five to seven red-
not absorb water (Krauss, 1948, 1949a,b). The dish peduncle bracts at its base. These bracts
 Morphology, Anatomy and Taxonomy 17

 200
 650  200

cu u.w. chl c.a.c. p.c.

s.b. chl
s.c.

3 4

2 e ch
ws.t.
1
1 cm

v.b.
a.c. m s.a.c.
ss.r.
7 f.s.
5 ss.c.
xy
6
fl

chl
ch
l.a.c.

hy g.c.
ph s.b. p
 200  650
 200 (after B.H. Krauss)

Fig. 2.3. 1. Diagram of a transverse section of a ‘Smooth Cayenne’ leaf showing: a.c., aerating canal; f.s.,
fibre strand; e, epidermis; v.b., vascular bundle; m, mesophyll; ws.t., water-storage tissue. 2. Outer epidermis
showing: cu, cuticle; s.b., silica body; u.w., undulating wall. 3. Aerating canal showing: c.a.c., central
aerating canal; s.c., stellate cell; chl, chloroplast. 4. Lower boundary of water-storage tissue showing: ch,
chlorenchyma; chl, chloroplast; p.c., palisade cells of water-storage tissue. 5. Lower epidermis showing:
ss.r., substomatal ring; s.a.c., secondary aerating canal; l.a.c., lateral accessory cell; g.c., guard cell; ss.c.,
substomatal chamber; s.b., silica body; p, pore. 6. Hypodermis and mesophyll (trichomes not shown)
showing: ch, chlorenchyma; chl, chloroplast; fi, fibres; hy, hypodermis. 7. Vascular bundle (mesophyll
without chloroplasts) showing: ph, phloem; xy, xylem. (After Py et al., 1987, with permission.)

are shorter and narrower than the ordinary The inflorescence consists of fewer than
leaves. The peduncle elongates after flower 50 (in some wild clones) to more than 200 (in
formation. Its length varies widely with the some cultivars) individual flowers; it is
botanical varieties or even cultivars. In addi- capped by a crown, composed of numerous
tion to its bracts, it bears, in many cultivars, a short leaves (up to 150) on a short stem. The
variable number of slips (up to a dozen or flowers or individual fruits are disposed
more), which can be positioned more or less around the central axis according to an 8/21
regularly between the stem and the fruit, at phyllotaxy in large-fruited cultivated
the axis of the peduncle bracts, or grouped pineapples (M.B. Linford, cited by Kerns et
just beneath the fruit. These slips can be con- al., 1936) and a 5/13 phyllotaxy for small-
sidered as dwarfish ‘aborted’ fruits with a rel- fruited wild pineapples or for young culti-
atively large crown (Collins, 1960). They may vated pineapples flowering prematurely
constitute an appreciable source of planting (Kerns et al., 1936). The fibrous axis contain-
material in extensive cultivation systems. ing the many vascular bundles that supply
18 G. Coppens d’Eeckenbrugge and F. Leal

ex
oc rh

ic

ph
t

rh
v

pc
en
l

rc
e

Fig. 2.4. Transverse section of a mature root of ‘Smooth Cayenne’ pineapple showing: e, epidermis; ex,
exodermis; oc, outer cortex with raphide cells (rc), whose transverse walls are collapsed to form air
passages; ic, outer portion of inner cortex; t, tracheids; ph, phloem; p, pith; v, vessel; pc, pericycle; en,
endodermis; l, large lacunae formed by collapse of many cells of inner cortex; rh, root hairs. (After Krauss,
1949b. © The University of Chicago, all rights reserved.)

the flowers is continuous with the peduncle trilobed and trifid style is almost as long as
and with the short stem of the crown (Fig. the petals and equal to or longer than the
2.5). Between the uppermost flower and the stamens. At anthesis, each stylar canal is an
crown is a transition region with bracts but unobstructed open channel from the stigma
no flowers. The edible part of the fruit con- to the locule directly above the placenta.
sists chiefly of the ovaries, the bases of sepals Petals are ligulate and free, each bearing at
and bracts and the cortex of the axis. The its base two slender funnelform scales or,
fruit shell is composed chiefly of sepal and more rarely, lateral folds that overlap the fila-
bract tissues and the apices of the ovaries ments. Petals are white at their base to violet-
(Okimoto, 1948). Anthesis normally takes blue at their tip. They are so close together at
place within a day. Flowering lasts 10–15 their outer end that only small insects can
days and occurs in a more or less acropetal enter the flower. This narrow tubular flower
succession along the inflorescence axis, but and the abundant nectar production are par-
some cultivars flower in a very disorderly ticularly adapted to hummingbird pollina-
manner. tion. Indeed, the three large nectary glands
Flowers are hermaphroditic and trimer- are so productive that nectar often fills the
ous, with three sepals, three petals, six sta- corolla and seeps out. The sepals are deltoid
mens in two whorls of three and one and appear similar to the bracts in colour
tricarpellate pistil (Fig. 2.6). The anthers are and texture. Each flower is surrounded and
bilobed, introrse and dorsifixed. The hollow, subtended at its base by a pulpous and thick
 Morphology, Anatomy and Taxonomy 19

Base of the crown

Shell

Ovary

Sepal and bract bases

Inflorescence axis,
the core

Cortex of the core

Theoretical limit of
the cortex of the core

Peduncle cortex

Peduncle

Fig. 2.5. The morphology of a ‘Smooth Cayenne’ pineapple fruit. (After Okimoto, 1949. © The University of
Chicago, all rights reserved.)

bract, covered by trichomes, which becomes tion). Orthotropous ovules are much less fre-
pointed and papyraceous at its tip. Parts of quent than anatropous ovules and their pres-
three other bracts complete the enclosure of ence and numbers are a varietal
the flower. Bract spininess is correlated with characteristic (M.F. Duval and G. Coppens
leaf spininess. d’Eeckenbrugge, unpublished results).
In the syncarpic inflorescence resulting Pollen grains are prolate and spheroidal,
from the fusion of the basal part of the flow- biconvex, isopolar and bilaterally symmetri-
ers and their axis, ovaries of adjacent flowers cal and diaperturate, with circular to slightly
are separated by the parenchymatous tissue elongated apertures situated at the poles.
of the calyx and bract bases. The ovary is The equatorial (36–51 m) and polar (46–59
inferior, tricarpellate and trilocular, with the m) dimensions are variable. The exine is
three septa forming an inverted Y when seen reticulate, and the polar areas show finer
in tangential section of the inflorescence. The reticulation than the rest of the surface (Wee
placentae and ovules are located in the and Rao, 1979). Male and female gametogen-
upper part of three deep cavities, called esis and embryogenesis are revised by Chan
locules, which are separated by the nectary et al. (Chapter 3, this volume).
glands. The ovules are caudate and arranged There is no floral abscission, and, except
in two single or double rows. The number of for the withering of the style, stamens and
ovules per flower varies with the cultivars, petals, the entire blossom develops
from 16 to 71 (Coppens d’Eeckenbrugge et parthenocarpically into a berry-like fruitlet.
al., 1993). The occurrence of two types of In the cultivated pineapple, growth from
ovules (unitegmic orthotropous and blossoming inflorescence to mature fruit
bitegmic anatropous) within the same ovary results in a 20-fold increase in weight. The
is common and orthotropous ovules are fer- enlargement of the calyx results from contin-
tilized (Okimoto, 1948; Rao and Wee, 1979; F. ued growth by cellular division, in stages up
Van Miegroet, 1993, personal communica- to flowering, and cell enlargement, in the
20 G. Coppens d’Eeckenbrugge and F. Leal

P STY
S
P P ST
ST S
STY S BR
DU PL
S BR LO
GL
O

A B C D E

Fig. 2.6. Pineapple flower. A. Floral diagram. B. Flower and subtending bract. C. Cross-section of ovary at
placenta level. D. Longitudinal section. E. Petal with scales and opposite stamen. BR, subtending bract; DU,
nectary duct; GL, septal gland; LO, locule; O, ovary limit; P, petal; PL, placenta; S, sepal; ST, stamen; STY,
style. (After Okimoto, 1949. © The University of Chicago, all rights reserved.)

later stages. During this size increase, cell basal eyes of the fruit (collar of slips). Slips
walls get thinner. The bract, sepal and ovary are curved at their base. As they are numer-
tissues are prominent structures in the ous in most cultivars, they are useful for
mature fruit. The large, conspicuous bract is rapid propagation. The crown can also be
fleshy and widened at its base and bends used for planting when the pineapple fruit is
over the flattened calyx surface, covering processed. Some plants may lack a crown or,
half of the fruitlet. Its papery tip dries during on the contrary, produce multiple crowns.
maturation. Internally, the locules get longer Also, crownlets may grow at the base of the
but relatively narrower and less conspicuous main crown or from some of the upper
in the developed fruit because of the expan- fruitlets.
sion of adjacent tissues, especially of the
septa. Placentas show some enlargement but
far less than the septal tissues, unless they Taxonomy
bear mature seeds. The seeds are approxi-
mately 3–5 mm long and 1–2 mm wide, flat The Bromeliaceae
on one side and curved on the other, with a
pointed end. They contain a hard flinty Pineapple belongs to the order Bromeliales,
endosperm and a minute embryo enclosed in family Bromeliaceae, subfamily Bromelioideae.
a brown to black coat, extremely tough and With 2794 species among 56 genera, accord-
leathery and roughened by numerous longi- ing to Luther and Sieff (1998), this is the
tudinal ridges (Miles Thomas and Holmes, largest family whose natural distribution is
1930). In the mature fruit, the stylar canals restricted to the New World, with the
get completely closed, first by a mucilagi- exception of Pitcairnia feliciana (Aug. Chev.)
nous plug, soon after anthesis, and a week or Harms & Mildbr., which is native to Guinea.
two later by cellular occlusion. Their unified geographical distribution and
their strong adaptation towards an epiphytic
mode of life indicate that this is quite a young
Vegetative propagules family. On the other hand, morphologically
they seem older than Rapateaceae, which led
Vegetative propagules are classified accord- Smith (1934) to accept that their restriction to
ing to their position on the plant. Suckers the Americas does not indicate extreme
appear on the earthed part of the stem. Stem youth. Judd et al. (1999) consider that the
shoots, which appear on the aerial part, are family probably represents an early divergent
more frequent. Slips appear on the peduncle. clade within the superorder Commelinanae.
They are often grouped near the base of the The Bromeliaceae have adapted to a very wide
fruit. Sometimes, they are produced from the range of habitats, ranging from terrestrial to
 Morphology, Anatomy and Taxonomy 21

epiphytic, deep shade to full sun, mesic to sules containing naked or appendaged seeds
extremely xeric and sea level to alpine, and adapted to wind dispersal. Their monophyly
from the hot and humid tropics to the cold is now questioned and new subfamilies
and dry subtropics. They thus cover a wide could be defined (Ranker et al., 1990; Terry et
area, from the centre of the USA to the north- al., 1997). The subfamilies Tillandsioideae and
ern regions of Argentina and Chile (Smith, Bromelioideae are considered monophyletic.
1934). The Bromeliaceae are set apart from The former include mostly epiphytic species,
other monocots by the unique, stellate or with smooth leaf margins, flowers usually
scale-like multicellular hairs and the unusual hypogenous and dry dehiscent capsules con-
conduplicate, spiral stigmas (Gilmartin and taining many plumose seeds adapted to
Brown, 1987). They are also characterized by wind dispersal. The second is the most
a short stem, a rosette of narrow stiff leaves, numerous. Most Bromelioideae are epiphytic,
terminal inflorescences in the form of frequently spiny, with epigynous flowers
racemes or panicles, hermaphroditic and and fleshy or leathery berries containing
actinomorphic trimerous flowers with well- naked seeds as an adaptation to dispersion
differentiated calyx and corolla, six stamens by birds or mammals. All the species exam-
and superior to inferior trilocular ovary, with ined exhibit CAM, with the exception of
axile placentation and numerous ovules. those of the genus Greigia, a trait much less
Fruits are capsules or berries and contain frequent in the other subfamilies (Medina,
small naked, winged or plumose seeds, with 1990). They seem to have evolved from east-
a reduced endosperm and a small embryo. ern Brazil and the Amazon basin, and they
Most species are epiphytic or saxicolous, but show a tendency to fusion of parts, fusion of
some are terrestrial. They are particularly their carpels to make an indehiscent fruit,
adapted to water economy, based on: (i) formation of an inferior ovary and fusion of
rosette structure; (ii) ability to absorb water sepals, petals and filaments. As stated by
and nutrients through their waxy leaves and Smith (1934), ‘Ananas capped the fusion ten-
aerial roots; (iii) ability to store water in spe- dency by merging the whole inflorescence,
cialized aquiferous leaf tissue; (iv) multicellu- flowers, bracts and all into one massy com-
lar trichomes functioning as water valvulae pound fruit.’
and reflecting radiation; (v) a thick cuticle; Pineapple is by far the most important
(vi) location of stomates in furrows, limiting economic plant in the Bromeliaceae. However,
evapotranspiration; and (vii) CAM. Their in the same Bromelioideae subfamily, some
root system is not well developed and func- Aechmea and Bromelia species also yield edi-
tions mostly to anchor the plant. ble fruits, such as A. bracteata (Swartz)
The Bromeliaceae are divided into three Grisebach, A. kuntzeana Mez, A. longifolia
subfamilies, the Pitcarnioideae, the (Rudge) L.B. Smith & M.A. Spencer, A. nudi-
Tillandsioideae and the Bromelioideae. While caulis (L.) Grisebach, B. antiacantha Bertoloni,
this division is widely recognized, their phy- B. balansae Mez, B. chrysantha Jacquin, B.
logenetic study has been confounded by karatas L., B. hemisphaerica Lamarck, B. nidus-
high levels of homoplasy for morphological puellae (André) André ex. Mez, B. pinguin L.,
and ecological characters. Most experts agree B. plumieri (E. Morren) L.B. Smith and B. tri-
that certain attributes have arisen several anae Mez (Rios and Khan, 1998). The most
times independently as adaptations to their common are known and consumed locally,
extreme environment. Molecular studies under names like cardo or banana-do-mato
have given contradictory results, placing dif- (bush banana), piñuelas (small pineapple) or
ferent subfamilies at the base of the karatas, gravatá and croata (derived from
Bromeliaceae (Clark and Clegg, 1990; Givnish Amerindian names given to terrestrial
et al., 1990; Ranker et al., 1990; Terry et al., bromeliads). Many other bromeliads are cul-
1997). The Pitcarnioideae were long held to be tivated as ornamentals, gathered for fibre
the most archaic. They are almost terrestrial, extraction or used in traditional medicine
with armed leaf margins, hypogenous or (Corrêa, 1952; Purseglove, 1972; Reitz, 1983;
epygenous flowers and dry dehiscent cap- Rios and Khan, 1998).
22 G. Coppens d’Eeckenbrugge and F. Leal

Evolution of pineapple classification sion of species descriptions, a few authors,

such as Bentham and Hooker (cited by
From the first observations of the pineapple André, 1889), claimed that the genus Ananas
by European explorers to the present time, is monospecific, with multiple wild and cul-
pineapple taxonomy has varied consider- tivated forms. Thus, in his first attempt at
ably. The first botanical description of culti- simplication, published in the Flora
vated pineapples was by Charles Plumier at Brasiliensis, Mez (1892) recognized only one
the end of the 17th century (but only pub- species, Ananas sativus, with five botanical
lished in 1755), when he collected plants varieties. The variety lucidus included the
called karatas and ananas on the island of pineapples with smooth leaves and a large
Hispaniola. Following the native classifica- fruit, but also the pitte or pitta, a small-
tion, he created the genus Bromelia for the fruited pineapple only cultivated for fibre.
karatas, in honour of the Swedish physician A. debilis was downgraded to a second vari-
Olaf Bromel, and described the ananas, ety debilis, ‘only known from European
using polynomials such as Ananas aculeatus glasshouses’. The variety bracteatus
fructu ovato, carne albida. In his Species included A. bracteatus (Lindl.) Schult. f., but
Plantarum, Linnaeus (1753) designated the also A. macrodontes Morren. The variety
pineapple as Bromelia ananas and Bromelia muricatus was made from A. muricatus
comosa, while Miller (1754, 1768) maintained Schult. f., although Mez expressed doubts
the name Ananas, with six varieties, all cul- about its existence. The last variety,
tivated. In the following classifications of microstachys, corresponded to the wild
the 18th and 19th centuries, as pineapple pineapple Acanthostachys ananassoides Baker.
was mainly known from attractive large- In 1917, Merrill established the binomial
fruited types, these varieties and other culti- Ananas comosus. In 1919, Hassler divided
vars were easily confused with species, the genus Ananas in two sections Euananas
which resulted in an overwhelming number and Pseudoananas. The latter was raised to a
of different names (Leal et al., 1998). Thus, distinct monospecific genus by Harms
Lindley (1827) used such names as Ananassa (1930), with Pseudananas macrodontes
sativa for ordinary cultivars, Ananassa lucida (Morren) Harms. In his second classifica-
for smooth-leaved cultivars such as tion, Mez (1934) did not recognize this new
‘Smooth Cayenne’ (from the variety Ananas genus, and proposed three species: (i) A.
(Lucidus) of the eighth edition of Miller’s comosus, including the wild pineapples and
Gardener’s Dictionary, published in 1768), the cultivated forms sativus (spiny types),
Ananassa debilis for a particular cultivar lucidus (smooth-leaved pineapples) and
with undulated leaves and Ananassa debilis (with doubts); (ii) A. sagenaria, corre-
bracteata for a crowned pineapple with long sponding to A. bracteatus (Lindl.) Schult. f.;
bracts. Schultes and Schultes (1830) and (iii) A. macrodontes Morren.
returned to the original name Ananas, with Unfortunately, the second simple classifica-
A. sativus, A. debilis, A. semiserratus (instead tion of Mez was no more successful than the
of A. lucida) and A. bracteatus. Linden (1879) first. From 1934 on, pineapple taxonomy
described a Colombian smooth-leaved culti- was dominated by the views of L.B. Smith
var with the ‘piping’ character under A. and F. Camargo. These authors divided the
mordilona. Morren (1878) gave the first clear genus Ananas, renaming and multiplying
description of a distinct pineapple, the the species in a long series of publications,
yvira, characterized by long bracts, the without describing new variation
absence of a crown and propagation by (Camargo, 1939, 1943, 1956; Smith, 1939,
stolons, which he named A. macrodontes. In 1961, 1962; Camargo and Smith, 1968; Smith
1889, both Baker and André described a and Downs, 1979). The resulting overclassi-
wild pineapple, with a long scape and a fication into two genera and nine species
small crowned fruit, respectively, under the (Smith and Downs, 1979; Table 2.1) has been
names of Acanthostachys ananassoides and A. severely criticized on the basis of practica-
pancheanus. In contradiction to this profu- bility, as well as inconsistency with avail-
 Morphology, Anatomy and Taxonomy 23

able data on reproductive behaviour and biology, according to previous reviews (Leal
morphological, biochemical and molecular and Coppens d’Eeckenbrugge, 1996; Leal et
diversity (Leal, 1990; Loison-Cabot, 1992; al., 1998).
Leal and Coppens d’Eeckenbrugge, 1996;
Coppens d’Eeckenbrugge et al., 1997; Leal et
al., 1998). Indeed, there are no reproductive
Ananas: one genus and two species
barriers between the Ananas species defined
by Smith and Downs and no differences
In the present classification, all pineapples
either in their floral structure and cytology
are regrouped into one Ananas genus, char-
or in their chromosome number or breeding
acterized among Bromeliaceae by the unique
system. The studies on diversity of
feature of their inflorescence, which is fused
isozymes (García, 1988; Aradhya et al., 1994)
into a syncarp. The seven valid Ananas
and on ribosomal and total DNA (Noyer,
species proposed by Smith and Downs
1991; Noyer and Lanaud, 1992; Duval et al.,
(1979) are downgraded to the level of five
1998) all showed a limited differentiation
botanical varieties of A. comosus.
between the different Ananas taxa, a strong
geographical component of variation and a
close affinity between certain wild pineap-
Ananas Miller, Gard. Dict. Abr. Ed. 4. 1754
ples and those cultivated for the fruit or for
the fibre. The study by Duval et al. (1998) Synonyms: Bromelia L. Spec. Pl. ed. 1 (1753) –
justifies the distinction between the gener- Ananassa Lindley, Bot. Reg. 13:1081 (1827).
ally diploid crowned pineapples propagat- Leaves densely rosulate, blades linear,
ing by shoots and the tetraploid crownless scarcely enlarged at base, often spiny. Scape
yvira propagating by stolons, although both evident. Inflorescence densely strobiliform,
taxa show much more similarity with each flowers hermaphrodite, sessile. Sepals free,
other than with other bromeliads, suggest- obtuse, asymmetric, persistent; petals free,
ing that they belong to the same genus. erect, violet or purplish pink. Stamens
For these reasons, we return to a much included, pollen ellipsoid, biporate; ovaries
simpler and more consistent classification, coalescing with each other and with the
taking into account morphological, bio- bracts and axis to form a fleshy syncarpic
chemical and molecular variation, geo- fruit; epigynous tube short; placentae apical;
graphical distribution and reproductive ovules caudate. Fruit succulent.

Table 2.1. Correspondence between the classification of Smith and Downs (1979) and the present
classification.

Smith and Downs Present classification

Pseudananas sagenarius (Arruda da Camara) Camargo Ananas macrodontes Morren

Genus Ananas Miller including eight species Ananas comosus (L.) Merrill including five
botanical varieties
Ananas ananassoides (Baker) L.B. Smith Ananas comosus var. ananassoides
Ananas nanus (L.B. Smith) L.B. Smith (Baker) Coppens & Leal
Ananas lucidus Miller Ananas comosus var. erectifolius (L.B.
Smith) Coppens & Leal
Ananas parguazensis Camargo & L.B. Smith Ananas comosus var. parguazensis
(Camargo & L.B. Smith) Coppens & Leal
Ananas comosus (L.) Merrill Ananas comosus var. comosus
Ananas monstrosus Invalidated by Leal (1990)
Ananas bracteatus (Lindley) Schultes f. Ananas comosus var. bracteatus (Lindl.)
Ananas fritzmuelleri Camargo Coppens & Leal
24 G. Coppens d’Eeckenbrugge and F. Leal

Type: Bromelia ananas Linnaeus (1753). fibre, fishing-net bromeliad, fierce fence
bromeliad and strong spine pineapple, in ref-
KEY TO THE SPECIES AND BOTANICAL VARIETIES erence to the uses of the plant.
1a. Apical coma inconspicuous or lacking, Type: Ananas macrodontes E. Morren, Belg.
plant propagating by elongate stolons . . . . . . Hortic. 28:140 (1878).
. . . . . . . . . . . . . . . . . . . . . Ananas macrodontes Plant propagating by elongate basal
1b. Foliaceous coma usually crowning the stolons, very rarely by axillary shoots, up to
syncarp. Plant propagating by the crown and 2 m high. Leaves densely rosulate, generally
shoots from the stem or peduncle (slips) . . . . arching decurved, 2–3 m long, blades 2–7 cm
. . . . . . . . . . . . . . . . . . . . . . . . . Ananas comosus wide, coriaceous, lustrous above, densely
2a. Flower bracts conspicuous, imbricate lepidote beneath, laxly serrate with curved
and covering the ovary . . . . . . . . . . . . . . . . . . retrorse and antrorse spines 3–10 mm long.
. . . . . . . . . . . . Ananas comosus var. bracteatus Scape erect, 20–50 cm high, 15–25 mm wide,
2b. Flower bracts relatively inconspicuous, furfuraceous; scape bracts foliaceous,
soon exposing the ovary apices reduced, red or green. Inflorescence simple,
3a. Spiny leaves. Leaf spines both antrorse strobilate, globose to ovoid, 8–20 cm long
and retrorse, blades narrowed towards and 6–9 cm wide, 100–200-flowered. Floral
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bracts densely imbricate, wide at base, lance-
. . . . . . . . . Ananas comosus var. parguazensis olate, acuminate, 2–5 cm long, erect, spiny,
3b. Spiny or smooth leaves. Leaf spines all red, reddish brown or green, cinereous–lepi-
antrorse, blades not narrowed at base dote, persistent; flowers opening in upward
4a. Dense rosette, wide blades, scape succession. Sepals erect, asymmetric, ovate
wide, fruit medium to large . . . . . . . . . . . to lanceolate, narrowed at base, apiculate,
. . . . . . . . . . . Ananas comosus var. comosus 10–15 mm long, thick with membranous
4b. Sparse rosette, blades usually nar- margins, green to red orange, sparsely lepi-
row, scape thin and long, usually erect, dote; petals tubular–erect, 3–5 cm long, the
fruit small to medium claw oblong, white, the blade lanceolate,
5a. Blades recurved or erect, spiny. purplish; stamens included, adnate to the
Fruit small to medium . . . . . . . . . . . . . . petals, shorter than the petals. Syncarp up to
. . . . . . Ananas comosus var. ananassoides about 20 cm long and 10 cm in diameter, not
5b. Blades erect, usually narrow and growing appreciably after anthesis, succu-
unarmed with the exception of the api- lent, without leafy crown.
cal thorn. Abundant production of A. macrodontes lacks a crown at the top of
shoots . . . . . . . . . . . . . . . . . . . . . . . . . . . . the syncarp and reproduces vegetatively by
. . . . . . . . Ananas comosus var. erectifolius stolons. Exceptional genotypes may also pro-
duce shoots directly from the stem (e.g.
ANANAS MACRODONTES MORREN (Plate 1). accession CRF001 at the Venezuelan Centro
Synonyms: Ananas microcephalus (Bak.) Nacional de Conservación de Recursos
Bertoni, An. Cient. Parag. II. 4: 250 (1919) – Fitogenéticos) or produce a rudimentary
Pseudananas macrodontes (Morren) Harms, crown (as in specimen US-2622957). Its leaf
Engl. Prantl., Nat. Pflanzenfam. 2. 15a (1930) margins bear strong spines, which are
– Pseudananas sagenarius (Arruda da Câmara) retrorse at the leaf base. The fruits are low in
Camargo, Rev. Agr. 14. 7–8:15–17 (1939). acid. A. macrodontes has been reported as
Synonymy with Bromelia sagenaria Arruda da self-compatible, with 2n = 4x = 100 chromo-
Câmara and Bromelia sylvestris Vellozo is somes (Collins, 1960). Its natural distribution
dubious because of very incomplete descrip- corresponds to coastal and southern Brazil,
tions, not mentioning clearly the absence of a up to Pernambuco, and to the drainage of
crown and hence generating a confusion the Paraguay and Paraná rivers, from south-
with A. comosus var. bracteatus. eastern Paraguay and north-eastern
Local names: yvira (Paraguay), gravatá de Argentina up to Mato Grosso (Coppens
rede, gravatá de cerca brava, nana caçaba d’Eeckenbrugge et al., 1997). Its habitat is
(Brazil). These names, respectively, mean limited to forest areas, under semidense
 Morphology, Anatomy and Taxonomy 25

shade. It is subjected to a rainy season dur- compound fruit; epigynous tube short; pla-
ing most of the year or even to periods of centae apical; ovules caudate.
flooding. Populations of A. macrodontes are Instead of producing stolons, A. comosus
rare now because of a very strong reduction multiplies by stem shoots (terrestrial and
of its habitat. aerial), slips (from the peduncle) and crown.
Camargo (cited by Reyes-Zumeta, 1967) The syncarpic fruit is formed of 50–200
considered that this species includes at least berries. The spines are generally antrorse but
three botanical varieties, one corresponding some genotypes also exhibit a few retrorse
to the description by Morren and another to spines. As commonly found in Bromeliaceae,
A. microcephalus, described by Bertoni (1919) A. comosus is diploid, with 50 minute and
in Paraguay. On the contrary, Baker and almost spherical chromosomes (Collins and
Collins observed very little variation in those Kerns, 1931; Canpinpin and Rotor, 1937;
areas (Smith, 1939). However, the few types Marchant, 1967; Sharma and Ghosh, 1971;
observed by Ferreira et al. (1992) in Paraguay Lin et al., 1987; Brown and Gilmartin, 1989;
and South Brazil showed significant varia- Dujardin, 1991). Giant unreduced gametes
tion, which was confirmed by the restriction may appear and produce natural triploids
fragment length polymorphism (RFLP) and tetraploids (Collins, 1933, 1960). Most
study of Duval et al. (1998). genotypes present reduced self-fertility
because of the action of a self-incompatibility
ANANAS COMOSUS (LINNAEUS) MERRILL. system, with considerable variation in its
Synonyms: Bromelia ananas L., Sp. Pl. 285 expression (Coppens d’Eeckenbrugge et al.,
(1753) – Bromelia comosa L., Herb. Amboin. 1993). The natural distribution of A. comosus
(1754), based on Anassa domestica Rumphius corresponds to that of its variety
– Ananas sativus sensu Mez, Mart. Fl. Bras. ananassoides, extending east of the Andes
3(3):290 (1892), in part – Ananas comosus (L.) (with some presence in the foothills of the
Merril sensu Mez, Pflanzenreich IV. 32:102 central Andes of Colombia), from northern
(1934) – see also synonyms in the following South America to northern Argentina and
descriptions. Paraguay.
Common names: nanas or ananas (in many
Indian languages of South America, and in Ananas comosus var. ananassoides (Baker)
French and Portuguese), piña (Spanish), Coppens & Leal (Plate 2). Synonyms:
pineapple. Acantostachys ananassoides Baker, Handb.
Type: Anassa domestica Rumphius from Brom. 25 (1889) – Ananas microstachys
Amboina in the Moluccas, Belg. Hortic. Lindman, Sv. Vet-akad. Handl. III. 24(8):39
28:140 (1878). (1891; nom. Ill.) – Ananas sativus var.
Plant not producing stolons but axillary microstachys (Lindm.) Mez, Mart. Fl. Bras.
shoots from the stem and/or slips from the 3(3):294 (1892) – Ananas guaraniticus Bertoni,
scape, up to 1.8 m high. Leaves densely rosu- An. Cient. Parag. II. 4: 274 (1919) – Ananas
late, scarcely enlarged at base, coriaceous, comosus var. microstachys (Mez) L.B. Smith,
densely lepidote beneath. Scape evident, Contr. Gray Herb. 104:72 (1934) – Ananas
erect; scape bracts foliaceous, usually red- ananassoides (Baker) L.B. Smith, Bot. Mus.
dish at base. Inflorescence densely strobili- Leafl. Harvard 7:70 (1939) – Ananas nanus
form, usually crowned with one or several (L.B. Smith) L.B. Smith, Brom. Soc. Bull.
rosettes of foliaceous bracts; flowers sessile, 12:54 (1962) – Ananas ginesio-linsii Reitz,
highly variable in number, from less than 50 Brom. Soc. Bull. 18:109 (1968).
to more than 150. Floral bracts broadly ellip- Local names: ananaí or nanaí, ananas de
tic to lanceolate, acuminate, sublepidote or ramosa (Brazil, Pará), curibijul, maya piñon,
lepidote. Sepals free, obtuse, slightly asym- piñuela, ananas do indio.
metric; petals free, erect, violet or purplish Lectotype: Regnell III 1261 in part (holotype,
pink, stamens included, pollen biporate and P), Caldas, Minas Gerais.
ellipsoid; ovaries coalescing with each other Leaves up to 2 m long and narrow; blades
and with the bracts and axis to form a fleshy linear, usually less than 4 cm wide,
26 G. Coppens d’Eeckenbrugge and F. Leal

subdensely serrate with wholly antrorse Harvard 7:78 (1939) – Ananas lucidus Miller,
spines. Scape elongate, slender, usually less sensu Smith, Bromel. In: Flora de Venezuela
than 15 mm thick; scape bracts large, subfoli- 12(1):1–361 (1971).
aceous. Inflorescence small to medium, glo- Synonymy with the lucidus variety of Miller
bose to cylindrical. Fruit slightly enlarged (Gard. Dict. ed. 8) is not founded because the
after anthesis, flesh scant to abundant, latter was created for the smooth-leaved cul-
fibrous; coma relatively large at maturity. tivars giving a large fruit.
Ananas nanus (L.B. Smith) L.B. Smith is Common names: curagua, curauá, curaná,
considered here just a dwarf form of A. como- kulaiwat, pitte.
sus var. ananassoides. The constancy of the Leaves stiff and erect, 1 m or longer,
distinctions reported by Smith (1962) do not blades about 35 mm wide, entire or nearly
constitute any argument in a vegetatively so, except for the pungent apex. Scape erect,
propagated plant. Indeed, sexual hybrids elongate, slender, to 15 mm thick; scape
segregate for size and most other traits. bracts foliaceous, erect. Syncarp small, cylin-
Ananas ginesio-linsii Reitz (ananas do indio) is drical or subcylindrical; floral bracts small,
a triploid clone showing particular vegeta- entire. Fruit slightly larger than inflores-
tive vigour (Lin et al., 1987; Dujardin, 1991). cence, coma relatively large, sometimes sur-
A. comosus var. ananassoides is the most rounded by additional crownlets; flesh scant,
common wild pineapple and the botanical fibrous, unpalatable.
variety that shows the highest genetic diver- Ananas comosus var. erectifolius is essen-
sity; it is very probably at the origin of the tially distinguished from A. comosus var.
cultivated pineapples (Leal and Coppens ananassoides by its smooth leaves. It is culti-
d’Eeckenbrugge, 1996; Coppens d’Eecken- vated by the natives in the Guianas, includ-
brugge et al., 1997; Duval et al., 1998). A. como- ing the Orinoco basin, and in the north of the
sus var. ananassoides is found in all tropical Amazon basin, for its very strong and long
areas of South America east of the Andes, fibres, used to make hammocks and fishing
generally in savannahs or in low-shaded for- nets (Leal and Amaya, 1991). The dry fibres
est, growing well on soils with limited water- constitute 6% of the plant weight (Camargo,
holding capacity (sand, rocks) and forming 1943). A. comosus var. erectifolius has never
populations of very variable densities. been found in the wild. Plants are medium-
However, a few genotypes thrive in dense sized, with abundant shoots and numerous
rain forest (in the Guianas). Most populations erect leaves and with a small, very fibrous
are monoclonal, but some are polyclonal, (inedible) fruit. It is also discriminated from
with variation of recent sexual origin (Duval A. comosus var. ananassoides by the absence of
et al., 1997). The plant has long and generally spines along the leaf margin. However, spiny
narrow spiny leaves and bears a small to types have been observed under cultivation
medium, globular to cylindrical syncarp on a or as mutants in germplasm collections. The
long and thin peduncle. The fruit is often absence of spines, as well as its erect habit,
seedy, and its pulp is white or yellow, firm are probably the product of human selection
and fibrous, palatable, with a high sugar and for high yield of easily extractable fibres from
acidity content. Some clones producing a strains of A. comosus var. ananassoides.
fruit of intermediate size (as compared with According to the DNA diversity study of
the comosus variety) are found both wild and Duval et al. (1998), different clones of A. como-
cultivated in the Guianas. Some dwarf types sus var. erectifolius show more affinity with
have been recently cultivated intensively as different clones of A. comosus var. ananassoides
ornamentals. than with each other, which suggests that this
process may have taken place independently
Ananas comosus var. erectifolius (L.B. Smith) at different times or places, or both.
Coppens & Leal (Plate 3). Synonyms: Ananas
sativus var. Duckei Camargo, Rev. Agr. 14. Ananas comosus var. parguazensis (Camargo
7–8:13–14 (1939), without Latin diagnosis – & L.B. Smith) Coppens & Leal (Plate 4).
Ananas erectifolius L.B. Smith, Bot. Mus. Leafl. Synonyms: Ananas ananassoides sensu L.B.
 Morphology, Anatomy and Taxonomy 27

Smith in part, Contr. U.S. Natl. Herb. 33:300 Ananas comosus var. comosus (Plate 5).
(1957) as to Colombian material; Fl. Venez. Synonyms (not mentioning horticultural
12:341 (1971) as to Venezuelan material – forms described as cultivars): Bromelia ananas
Ananas parguazensis Camargo & L.B. Smith, L., Sp. Pl. 285 (1753) – Bromelia comosa L.,
Phytol. 16:464 (1968). Herb. Amboin. (1754), based on Anassa
Common names: piña montañera (Amazonas, domestica Rumphius – Ananas varieties
Venezuela), ananaí (Roraima, Brazil), described by Miller, Gard. Dict. ed. 8 (1768) –
kurupira-nana (Río Negro, Brazil), gravatá Ananassa sativa Lindley, Ananassa lucida
(Pará, Brazil). Lindley and Ananassa debilis Lindley, Bot.
Type: Camargo 3 (holotype, COL; isotype, Reg. 13:1068 (1827) – Ananas sativus Schultes
US), Angelita, Bolívar, Venezuela, 16 June f., Ananas debilis Schultes f. and Ananas semi-
1966. serratus Schultes f., in Roemer & Schultes,
Leaves up to 2 m long, blades wide (usu- Syst. 7(2):1283, 1287 (1830) – Ananas sativus
ally more than 3 cm), somewhat contracted var. lucidus Baker, Handb. Brom., 23 (1889) –
towards base, green to red, sinuate–serrate, Ananas mordilona Linden, Belg. Hort. 29, 302
spines strong, antrorse and retrorse. Scape (1889) – Ananas sativus var. lucidus Mill. and
slender, elongate, often decumbent (depend- Ananas sativus var. debilis Lindley, Mez, Mart.
ing on fruit size); scape bracts foliaceous, Fl. Bras. 3(3):292–293 (1892) – Ananas sativus
notably dilated above base. Inflorescence var. pyramidalis (Miller) Bertoni, Ananas
few-flowered, subglobose, 4–10 cm long in sativus var. viridis (Miller) Bertoni and
fruit; apical coma large (reaching 30 cm in Ananas sativus var. hispanorum Bertoni, An.
height), basally more or less retrorse serrate. Cient. Parag. II. 4: 271, 272 (1919) – Ananas
Fruit succulent, flesh palatable. comosus f. sativus (Lindl.) Mez, Ananas como-
As mentioned by Smith and Downs sus f. lucidus (Mill.) Mez and Ananas comosus
(1979), Ananas pancheanus André could also f. debilis (Lindl.) Mez, Pflanzenreich IV.
correspond to A. comosus var. parguazensis. 32:102–104 (1934) – Ananas comosus (L.)
Indeed, André (1889) mentions a forest habi- Merril, interpr. Rumph. Herb. Amb.
tat, wide leaves (6 cm), antrorse and retrorse Common names: ananas, nanas, piña, pineap-
spines and a small subspherical syncarp. On ple, abacaxi (Brazil), matzatli (Aztec).
the other hand, the short and thin spines, Leaves numerous and wide, spines
with a density increasing regularly from the antrorse if present. Scape stout, usually 2 cm
base to the top of the leaf, are not typical of or more in width and less than 40 cm in
the variety. The exsiccata observed in Kew is length. Inflorescence large, many-flowered.
too poor to offer an opinion on. Floral bracts inconspicuous, soon exposing
The wild A. comosus var. parguazensis is the apices of the ovaries, weakly serrulate or
very similar to A. comosus var. ananassoides, entire. Syncarp growing appreciably after
with a difference in the retrorse orientation anthesis, to reach a final size several times
of some spines and a wider leaf slightly con- superior to that of the inflorescence; fruit
stricted at its base. Its distribution is mostly fleshy; seeds usually rare.
limited to the Orinoco and Rio Negro basins, The highly variable A. comosus var. como-
with a few observations in eastern Colombia sus is characterized by its large fruit (up to
and in the north-eastern Amazon (Coppens several kilograms in some cultivars), borne
d’Eeckenbrugge et al., 1997) and it shows less on a wide and strong peduncle. Its leaves are
variability than A. comosus var. ananassoides numerous (40–80) and wide ( 5 cm). In
(Duval et al., 1998). While A. comosus var. spiny cultivars, spines are antrorse and gen-
ananassoides shows a higher water-use effi- erally denser than in other botanical vari-
ciency and grows in both partial shade and eties. Certain cultivars are partially spiny
full sun, A. comosus var. parguazensis is (e.g. the cultivar ‘Smooth Cayenne’) or even
restricted to shadier environments (Leal and completely smooth (as in the cultivars with
Medina, 1995). It grows in the lowland the ‘piping’ trait). Seeds are rare because
forests, under canopies of variable densities, most cultivars present reduced fertility com-
from clearings or river-banks to dense forest. bined with self-incompatibility (Coppens
28 G. Coppens d’Eeckenbrugge and F. Leal

d’Eeckenbrugge et al., 1993). Since the first its long bracts (longer than the fruitlet) of a
Spanish and Portuguese great voyages, culti- bright pink to red colour. The fruit and
vation of A. comosus var. comosus has become peduncle are medium-sized. There are two
pantropical. main forms of A. comosus var. bracteatus. The
The species A. monstrosus (Carrière) L.B. most common corresponds to A. bracteatus
Smith was invalidated by Leal (1990) sensu Smith & Downs, with a spectacular
because the crownless fruit characteristic is bright red inflorescence and antrorse spines,
not stable. In addition, the names A. monstro- of which a variegated cultivar has been
sus and A. lyman-smithii Camargo, on which derived and propagated in all the tropical
it is based, are illegitimate, the former regions as a garden ornamental. It has also
because Carrière never intended to describe been used for the cut-flower market and a
a new species and the latter because it is a further mutation suppressing spines has
nomen nudum. been selected recently. The second form cor-
responds to A. fritzmuelleri Camargo, with
Ananas comosus var. bracteatus (Lindl.) antrorse and retrorse spines. Both forms
Coppens & Leal (Plate 6). Synonyms: Ananassa show very little variation, suggesting that
bracteata Lindley, Bot. Reg. 13:1081 (1827) – they are very particular types, probably
Ananas bracteatus (Lindley) Schultes f., Syst. selected by humans and dispersed through
7(2) (1830) – Ananas sativus var. bracteatus clonal propagation and still capable of devel-
(Lindley) Mez, Mart. Fl. Bras. 3(3):293 (1892) oping subspontaneous populations. They
– Ananas sagenaria sensu Mez, Pflanzenreich share certain morphological, biochemical
IV. 32:104 (1934) – Ananas bracteatus var. albus and DNA characters with A. macrodontes. In
L.B. Smith, Bot. Mus. Leafl. Harvard 7:76 diversity studies, they appear uniform and
(1939) – Ananas fritzmuelleri Camargo, Bol. well grouped (García, 1988; Duval and
Téc. Inst. Agr. Norte, Pará. 1:16 (1943). Coppens d’Eeckenbrugge, 1993; Aradhya et
As mentioned in the description of A. al., 1994; Duval et al., 1998), at some distance
macrodontes Morren, there is a possible syn- from the other botanical varieties of A. como-
onymy with Bromelia sagenaria Arruda da sus, but closer to A. comosus than to A.
Câmara and Bromelia sylvestris Vellozo (and macrodontes (Duval et al., 1998). A. comosus
all subsequent names referring to these two var. bracteatus hybridizes easily with A.
descriptions). comosus.
Common names: ananas de cerca, ananas
bravo, ananas do mato, karaguata-ruha (south-
ern Brazil). Origin and Evolution
Type: Lyndley s n (holotype, CGE n v), or
original description and plate. After inventorying and describing the vari-
Leaf blades wide, coarsely and usually ability found in Paraguay, Bertoni (1919)
laxly spinose–serrate, spines antrorse or stated that the pineapple was domesticated
antrorse and retrorse. Scape thick and stout. by the Tupi-Guarani Indians from A.
Syncarp medium to large. Floral bracts con- guaraniticus (A. comosus var. ananassoides) and
spicuous, imbricate and covering the ovaries, then accompanied them in their northward
coarsely serrate, usually red or pink. Fruit migrations to the Antilles, northern Andes
succulent at maturity. and Central America. This hypothesis has
A. comosus var. bracteatus has the same been retained in many reviews on crop ori-
original southern distribution area as A. gins (e.g. Collins, 1948, 1949, 1960,
macrodontes. It is always found cultivated as Purseglove, 1972; Pickersgill, 1976; Sauer,
a living hedge, for fibre and fruit juice, or 1993). In his discussion of Vavilov’s theory,
abandoned in ancient settlements (Baker and Brücher (1971) underlined the presence of
Collins, 1939; Duval et al., 1997). The plant is wild forms and primitive cultivars in the
very vigorous, with wide and long leaves, north of South America and proposed that
large spines and abundant production of domestication occurred on the Guiana
shoots. The inflorescence is characterized by shield, along the river system, and, possibly,
 Morphology, Anatomy and Taxonomy 29

in north-eastern Brazil. This work, written in A. comosus: the cultivated A. comosus var.
German and presenting imperfect informa- comosus and A. comosus var. bracteatus (the
tion on pineapple taxonomy and distribu- latter with poor morphological variation)
tion, was largely ignored. and savannah types of A. comosus var.
The hypothesis of a northern origin of ananassoides (Leal and Coppens
pineapple came back with Leal and Antoni d’Eeckenbrugge, 1996; Coppens d’Eecken-
in 1981. These authors suggested that the brugge et al., 1997; Duval et al., 1997).
centre of origin of the genus should be Molecular studies tend to confirm this
located in an area within 10°N–10°S latitude hypothesis (Duval et al., 1998). Thus, it seems
and 55°–75°W longitude, because the flora of that A. comosus var. comosus and A. comosus
this region is endemic and included the var. erectifolius evolved from A. comosus var.
largest number of species considered valid at ananassoides and/or A. comosus var. par-
that time. They proposed that south-eastern guazensis, the first by a selection based on
Brazil could be a secondary centre of origin large fruit size, through an increase in the
and distribution. Extensive expeditions in fruitlet number and size, high quality (lower
Venezuela, Brazil and French Guiana (Leal et acidity) and reduced seediness, and the sec-
al., 1986; Ferreira et al., 1992; Duval et al., ond by a selection for long, fibrous and
1997) widened significantly the known smooth leaves. Other traits were necessarily
diversity of wild and cultivated forms and modified in the course of domestication and
reinforced the hypothesis of a northern cen- selection.
tre of origin for the cultivated pineapple. Increased fruit size in A. comosus var.
Indeed, a wider morphological variation was comosus implied increased size of other
observed both in the wild and under cultiva- organs, i.e. wider leaves, bigger stem, shorter
tion in the areas to the north of the Amazon and wider peduncle. Sensitivity to natural
River (Orinoco and Rio Negro basins, flowering induction was reduced, allowing a
Amapá, Guianas) than in the southern areas longer cycle and hence a larger fruit.
(Paraguay, south of Brazil). Cultivation based on asexual propagation
Four botanical varieties of A. comosus and artificial selection of seedless genotypes
occur in the area north of the Amazon River: reduced the natural selective pressure on fer-
the cultivated A. comosus var. comosus, with tility and reinforced self-incompatibility.
many cultivars, and A. comosus var. erecti- Domestication also influenced spininess. In
folius, and the wild A. comosus var. parguazen- some cases, spininess was suppressed by the
sis and A. comosus var. ananassoides, with a selection of rare dominant mutations. The
wide range of morphological and adaptive excellence of the selections obtained by the
variation, from forest types to dry savannah Amerindians, still unsurpassed, their knowl-
types, including medium-fruited types still edge of the crop and the wide pre-
submitted to domestication cycles. In the Columbian distribution of the pineapple all
southern areas are found A. macrodontes and indicate a very ancient domestication, prob-
representatives of three botanical varieties of ably several thousands of years before Christ.

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Phytologist 83, 485–497.
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32 G. Coppens d’Eeckenbrugge and F. Leal

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 3 Breeding and Variety Improvement

Y.K. Chan,1 G. Coppens d’Eeckenbrugge2 and G.M. Sanewski3

1MARDI, GPO Box 12301, Kuala Lumpur, 50774, Malaysia; 2CIRAD-FLHOR/IPGRI,
AA 6713, Cali, Columbia; 3Maroochy Research Station, PO Box 5083, SCMC,
Nambour, Qld 4560, Australia

The pineapple industry of the world is domi- of new, resistant cultivars seemed the right
nated by the cultivar ‘Smooth Cayenne’, strategy to redress this situation. Further,
which is used both for fresh fruit and for while ‘Smooth Cayenne’ is quite good for
processing. The near-total reliance on a sin- processing, the fresh-pineapple markets in
gle cultivar in crop cultivation is uncommon the world are more diversified and, where a
and is made even more unbelievable by the choice is offered, this cultivar is not always
fact that ‘Smooth Cayenne’ has been the preferred. These are primary justifications
backbone of the global pineapple industry for the commencement of breeding pro-
for more than a century. First collected by grammes for pineapple worldwide.
Perrotet in 1819 in French Guiana under its In 1914, the Pineapple Growers
local name ‘Maipuri’ (Perrotet, 1825), it Association of Hawaii started one of the ear-
rapidly spread to other geographical regions, liest and most concerted efforts in pineapple
has adapted well and is known by such improvement. The experimental station of
other synonyms as ‘Kew’, ‘Giant Kew’, this association later became the Pineapple
‘Champaka’ and ‘Sarawak’. The ‘Smooth Research Institute of Hawaii (PRI). The
Cayenne’ monopoly is undoubtedly due to works of K.R. Kerns and J.L. Collins from the
its high yield, adaptability and good charac- PRI are renowned and are still often cited by
teristics for canning. Further, highly special- present-day researchers. One of the main
ized systems of production and processing objectives of the PRI programmes was to
protocols have been developed almost exclu- develop pest and disease resistance in a
sively for this cultivar. These factors, ‘Smooth Cayenne’-type variety. Varieties
together with impending high costs and were developed which had better resistance
attendant inconveniences in changing to a to Phytophthora, mealybug wilt, nematodes,
new variety, account for the difficulty in pink disease and internal brown spot. Some
replacing ‘Smooth Cayenne’. selections also had higher vitamins A and C,
less acid in winter-ripened fruits, a better har-
vesting peak, higher yield and plant vigour
Global Pineapple Breeding Research and improved cannery recovery. Many vari-
eties were selected which were better than
The overdependence of the pineapple indus- ‘Smooth Cayenne’ in certain individual char-
try on a single cultivar with a narrow genetic acteristics, but eventually none could replace
base has made it extremely vulnerable to the ‘Smooth Cayenne’ because, after extensive
threats of pests and diseases. Development evaluation, each would reveal a fatal flaw

© CAB International 2003. The Pineapple: Botany, Production and Uses

(eds D.P. Bartholomew, R.E. Paull and K.G. Rohrbach) 33
34 Y.K. Chan et al.

(Williams and Fleisch, 1993). The PRI closed and Australia. In the Philippines, the recent
in 1975 and evaluation of progenies was com- breeding programme carried out by the
pleted at the Maui Pineapple Company in Institute of Plant Breeding (Los Baños) was
1985. A small number of the PRI varieties and based on crosses between ‘Smooth Cayenne’
the genetic collection were turned over to the and ‘Queen’, with the objective of develop-
US Department of Agriculture (USDA) ing spineless ‘Queen’ types. One selection
germplasm repository in 1986. was micropropagated (Villegas et al., 1995).
Many other pineapple-growing countries The research on pineapple in Côte d’Ivoire
have also started breeding programmes to was started in 1978 and conducted by the
develop high-yielding varieties with specific fruit department (Département des
adaptation to their own environments. Most Productions Fruitières et Horticoles) of the
current breeding programmes are focused on Centre de Coopération Internationale en
the development of cultivars for the fresh Recherche Agronomique pour le
market. It is important to note that hybrid- Développement (CIRAD-FLHOR). ‘Smooth
ization programmes attempting to replace Cayenne’ was crossed with ‘Perolera’ (in fact
‘Smooth Cayenne’ for processing have been ‘Manzana’) with the objectives of developing
rare since the termination of the PRI pro- high ascorbic acid content for resistance to
gramme in Hawaii. Most hybridization black-heart disorder (an internal fruit break-
programmes revolve around the ‘Smooth down), developing resistance to Phytophthora
Cayenne’ because of its all-round good and nematodes and prevention of green ripe
qualities and universal acceptance. In 1926, fruits (internal flesh colour turns yellow
Taiwan (then Formosa) started a breeding before the change of peel colour) (Loison-
programme using ‘Smooth Cayenne’ crossed Cabot, 1987). A new hybrid called ‘Scarlett’,
with several of its local varieties, ‘Ohi’, ‘Uhi’, with commercial potential as a fresh fruit,
‘Anpi’ and ‘Seihi’. A later hybridization was selected from this hybridization pro-
programme using ‘Smooth Cayenne’ and gramme (Coppens d’Eeckenbrugge and
‘Queen’ resulted in the selection of ‘Tainung Marie, 2000).
4’ (‘Easy Peeler’), the eyes (fruitlets) of which In Cuba, the hybridization programme
can be picked off for eating without having was started in 1991 using ‘Serrana’ (‘Smooth
to peel the fruit (Fitchet, 1989). This pro- Cayenne’) × ‘Perolera’ and three hybrids
gramme continued, with the release of were micropropagated for commercial fruit
‘Tainung 13’ and ‘Tainung 16’ in 1995 and production (Benega et al., 1998b). In Puerto
1996, respectively (Chang et al., 1997). Rico, a natural cross between ‘Smooth
In Malaysia, early pineapple improvement Cayenne’ and ‘Española Roja’ resulted in a
programmes focused on the selection of selection called PR1–67 (Ramírez et al., 1972).
promising variants in clonal fields. This has In Australia, the breeding programme uti-
resulted in the development of ‘Masmerah’, a lizes mainly ‘Smooth Cayenne’, ‘Queen’ and
higher-yielding variant of the ‘Singapore selected PRI hybrids. Sanewski (1998) found
Spanish’ (Wee, 1974). Later hybridization that the PRI hybrids ‘73–50’, ‘53–116’ and
programmes conducted by the Malaysian ‘59–656’ and the Philippine hybrid ‘24–80’
Agricultural Research and Development generally produced more hybrid progenies
Institute (MARDI) developed a hybrid called with commercially acceptable attributes for
‘Nanas Johor’ from a cross between ‘Smooth the fresh-fruit market.
Cayenne’ and ‘Singapore Spanish’ (Chan and In Brazil, the major cultivars are ’Pérola’
Lee, 1985). In 1996, another hybrid called and ‘Smooth Cayenne’ and the main objec-
‘Josapine’ suitable for table fruit, was devel- tive of the breeding programme conducted
oped from a cross between ‘Nanas Johor’ and by the Empresa Brasileira de Pesquisa
‘Sarawak’ (a variant of ‘Smooth Cayenne’) Agropecuaria (EMBRAPA) was to develop
(Chan and Lee, 1996). cultivars with resistance to fusariosis disease
The ‘Smooth Cayenne’ was again featured caused by Fusarium subglutinans. The resis-
in breeding programmes carried out in the tant parents used in the crosses with ‘Pérola’
Philippines, Côte d’Ivoire, Cuba, Puerto Rico and ‘Smooth Cayenne’ gave the highest pro-
 Breeding and Variety Improvement 35

portion of hybrid progenies with desirable ‘Española Roja’ in the Caribbean and
characters (Cabral et al., 1993). Fifty-seven ‘Perolera’ and ‘Manzana’ in the Andes of
hybrids have been selected for further Colombia and Venezuela. Germplasm move-
evaluation. Three ‘Perolera’ × ’Smooth ment has been accompanied by clonal differ-
Cayenne’ and two ‘Primavera’ × ’Smooth entiation and/or selection and the cultivars
Cayenne’ hybrids are being tested for even- have frequently been renamed, often with
tual release. the same or similar names (e.g. ‘Sugar Loaf’).
Breeding strategies used by most coun- Some authors have classified them into horti-
tries appeared to be hybridization and selec- cultural groups. The last of these classifica-
tion of the segregating hybrid progenies. It tions, by Py et al. (1987), recognizes five such
was evident that ‘Smooth Cayenne’, the stan- groups: ‘Cayenne’, ‘Queen’, ‘Spanish’,
dard benchmark cultivar worldwide, is the ‘Pernambuco’ and ‘Perolera’. Since then, a
favourite parent for the crosses. It seemed much wider diversity of cultivars has been
fairly strange that hitherto, only a handful of collected and characterized, making the ear-
hybrids have been released from these lier classifications inadequate, confusing and
hybridization programmes. As noted by inappropriate. Coppens d’Eeckenbrugge et
Nakasone (1976): ‘breeding work has been al. (1997) proposed abandoning such classifi-
done in Hawaii, Australia, Taiwan, Malaysia, cations and coming back to the international
South Africa and other areas, resulting in sig- nomenclature for cultivars (Anon., 1980).
nificant contributions to knowledge in cytol- Accordingly, a new name can only be attrib-
ogy and genetics, but not to the list of new uted to a new cultivar whose characteristics
cultivars’. This strongly emphasizes the are stable and clearly distinct, regardless of
quality and stability in performance of the whether the method of development is
singular cultivar ‘Smooth Cayenne’, whose through clonal selection or sexual recombi-
durability and ability to weather the advent nation. Thus, most minor variants and clonal
of new hybrids are really remarkable. selections should not be considered as new
cultivars, although a small difference in fruit
size may be economically very significant. To
Germplasm Resources take into account interclonal genetic varia-
tion and regional nomenclature, Coppens
Origin, diversity and description of cultivars d’Eeckenbrugge et al. (1997) proposed men-
tioning local names or specific clones
Early European pineapple experts recog- together with the cultivar name, which
nized as many as 48–68 pineapple cultivars would give, for instance, ‘Champaka’
and classified them on the basis of spininess, (‘Smooth Cayenne’) and ‘McGregor’
fruit shape and flower colour (Munro, 1835; (‘Queen’).
Beer, 1857). Most of them originated from
coastal regions, such as the Caribbean, from
which sea trade favoured their distribution Main cultivars
to other tropical countries and to the
European glasshouses. Others were grown ’Smooth Cayenne’ (Plate 7), or ‘Cayenne
from seeds in Europe. Nearly all disappeared Lisse’, is still common in the Guianas under
with European production and only ‘Queen’, the name ‘Maipuri’, meaning ‘tapir’. It has
‘Smooth Cayenne’, ‘Baronne de Rothschild’, been renamed ‘Kew’, ‘Sarawak’, ‘Esmeralda’,
‘Havannah’ (‘Española Roja’), ‘Black ‘Claire’, ‘Typhoon’ and ‘Saint Michel’. The
Antigua’ and ‘Montserrat’ are still known ovoid medium-sized fruit (1.5–2.5 kg) of
today. The first two have reached all tropical ‘Smooth Cayenne’ is held on a short and
and subtropical countries. ‘Singapore strong peduncle (Fig. 3.1). It ripens progres-
Spanish’, which was introduced very early, is sively, turning yellow from the base to the
still cultivated to some extent in Asia. Other top, which is reflected in a strong internal
cultivars are economically important in their maturity gradient too. The flesh is pale
region of origin, such as ‘Pérola’ in Brazil, yellow, soft and juicy, with considerable
36 Y.K. Chan et al.

variation in sugar (from 13 to 19°Brix) and sidered to be tolerant to Phytophthora sp. (Py
acidity, depending on environmental condi- et al., 1987) and resistant to fruit collapse
tions – mainly rain and temperature – and caused by Erwinia chrysanthemi Burkbolder
low ascorbic acid content. Despite the high (Lim and Lowings, 1979).
sugar content of ‘Smooth Cayenne’, its acid- The cultivar ‘Cayenne Baronne de
ity is often considered excessive among trop- Rothschild’, cultivated in West Africa, is the
ical consumers and this has largely spiny form of ‘Smooth Cayenne’. ‘Hilo’, a
contributed to the image of pineapple as an Hawaiian variant, might merit cultivar sta-
acidic fruit. ‘Smooth Cayenne’ juice is not of tus as it produces many more slips than the
good quality, because of poor colour, high standard ‘Smooth Cayenne’ (Collins, 1960).
sugar and turbidity. The plant is medium- Collins and Kerns (1938) and Collins (1960)
sized (80–100 cm), with 60–80 dark green have described more than 30 ‘Smooth
leaves (c. 100 cm long and 6 cm wide), which Cayenne’ mutants in Hawaii, the most com-
bear small spines at their base and tip. mon being spiny leaves, multiple crowns
Vegetative propagation is normally by shoots and fasciated crowns or fruits and collar of
and slips, which are quite limited in produc- slips. The most interesting are ‘seedy fruit’, a
tion. The production cycle of ‘Smooth series of mutations that breaks down self-
Cayenne’ is longer than that of most other incompatibility in pollen, and resistance to
cultivars and this is exacerbated in cool cli- mealybug wilt, which was also observed in
mates. ‘Smooth Cayenne’ is sensitive to Mexico (Torres Navarro et al., 1989).
many known pests (fruit borers, mites, sym- ’Singapore Spanish’ (Plate 8) is second in
phillids, nematodes) and diseases (mealybug importance for canning. Other names and
wilt, fusariosis, fruitlet core rot, butt rot) and particular strains are ‘Singapore Canning’,
to internal browning (Rohrbach and Schmitt, ‘Ruby’, ‘Red Pine’, ‘Nanas Merah’, ‘Nangka’,
1994; Bello et al., 1997a,b). However, it is con- ‘Gandul’, ‘Betek’ and ‘Masmerah’ (Wee,
1972). ‘Singapore Spanish’ is cultivated in
South Asia, particularly in Malaysia, because
of its good adaptation to peat soils and its
golden-yellow flesh colour. Fruits are small
(around 1 kg – heavier in ‘Masmerah’), cylin-
drical and dark purple, turning copper-
orange when ripening (Fig. 3.2). Sugar and
acidity are low (10–12°Brix) and the taste is
poor; however, the juice is of good colour
and quality. The plant is medium-sized
(80–100 cm), with 35–70 dark green leaves,
about 150 cm long and 5 cm wide. Leaf
spininess is highly variable from clone to
clone, from complete spininess to very few
spines. The bracts of the peduncle and inflo-
rescence are bright red. The plant is vigorous
and produces many slips (about two to six)
and shoots. Multiple crowns are frequent.
‘Singapore Spanish’ is adapted to high soil
pH, which is probably related to its tolerance
to Phytophthora, a fungus that develops bet-
ter in such conditions. It shows severe
chlorosis when exposed to high manganese
concentration in the soil. It is also sensitive to
fruit collapse, the most serious disease in
Malaysia (Lim and Lowings, 1979), and to
Fig. 3.1. ‘Smooth Cayenne’. nematodes (Collins and Hagan, 1932).
 Breeding and Variety Improvement 37

Fig. 3.2. ‘Gandul’ (cv. ‘Singapore Spanish’).

The cultivar ‘Selangor Green’, also named between clones. ‘Queen’ is a robust cultivar,
‘Green Pine’, ‘Green Spanish’, ‘Nanas Hijau’ showing more general tolerance to stress,
and ‘Selassie’, is derived from a mutation pests and diseases than ‘Smooth Cayenne’.
suppressing anthocyanins in all the organs of On the other hand, it is susceptible to
‘Singapore Spanish’. Thus the leaves and Phytophthora and fruit collapse and highly
inflorescence are uniformly green, the petals susceptible to chilling and internal brown-
pale yellow and the fruit yellow at maturity. ing, particularly if harvested before maturity,
Other traits have not been altered. Another and to fruitlet core rot and butt rot (Lim,
mutation produced a variegated red-fruited 1971; Winks et al., 1985; Swarts, 1990;
cultivar used as an ornamental. Lutchmeah, 1992).
The cultivar ‘Queen’ (Plate 9) is widely ’Española Roja’ or ‘Red Spanish’ (Plate
distributed, but it is more particularly culti- 10) is widely cultivated in Venezuela and the
vated in the southern hemisphere, in South Caribbean basin, under names such as ‘Black
Africa and Australia, for the fresh-fruit mar- Spanish’, ‘Key Largo’, Havannah’ or
ket. It has been called ‘Mauritius’, ‘Malacca’, ‘Habana’, ‘Cubana’, ‘Cowboy’, ‘Bull Head’,
‘Red Ceylon’, ‘Buitenzorg’, ‘Victoria’, ‘Ripley ‘Cumanesa’ and ‘Native Philippine Red’
Queen’ and ‘Alexandra’ (Mendiola et al., (Leal, 1990). The fruit is medium-sized (1.2–2
1951; Leal, 1990). In Malaysia, it is called kg) and orange, with a barrel shape. The
‘Moris’ and Chan and Lee (1995) have flesh is firm, pale, aromatic and sweet, with
described several variants. ‘McGregor’ is a moderate sugar content (around 12°Brix) but
vigorous Australian selection. The tetraploid low acidity. The plant is medium-sized, with
‘Z’ or ‘James Queen’ was found in South spiny or half-spiny dark green leaves.
Africa (Nyenhuis, 1974). The plant is small Smooth clones have been selected. Floral
(60–80 cm), with short and very spiny silvery bracts are an intense bright red colour. The
leaves, and gives a small fruit (0.5–1 kg), plant regularly produces some slips (about
with a full yellow shell and small prominent one to three) and suckers. ‘Española Roja’ is
eyes (Fig. 3.3). The golden-yellow pulp is vigorous and tolerant to high temperature,
crispy and sweet (14–18°Brix), with an excel- drought, internal browning, butt rot, wilt
lent flavour and long shelf-life. The produc- and Phytophthora (Py et al., 1987; Rohrbach
tion of slips and shoots varies widely and Schmitt, 1994), but not to high
38 Y.K. Chan et al.

Fig. 3.3. ‘Mauritius’ (cv. ‘Queen’).

manganese in the soil and nematodes (Ayala

et al., 1969). It is highly susceptible to the
South American lepidopter Strymon basilides
(Geyer).
’Pérola’ (Plate 11), ‘Pernambuco’ or ‘Branco
de Pernambuco’ is the main cultivar in the Fig. 3.4. ‘Pérola’.
Brazilian fresh market. ‘Jupi’ is a particular
strain. In Florida, it has also been named
‘Abacaxi’, ‘Abakka’ and ‘Eleuthera’. The important cultivar of the north-eastern
fruit is small to medium (0.9–1.6 kg), ovoid Andes of Colombia and Venezuela. The fruit
(when small) to conical and green, with a lit- is large (1.5–3 kg), yellow to orange, with an
tle yellow in the centre of the mature eyes irregular cylindrical shape. It is borne on a
(Fig. 3.4). The flesh is soft, white and juicy long peduncle, which makes it susceptible to
with an exquisite aroma. Its sugar fruit lodging and sunburn, particularly in
(13–16°Brix) and ascorbic acid contents are hot lowlands. The flesh is pale yellow to yel-
high. As with most conical fruits, it shows a low, firm and sweet (although only around
strong gradient of maturation from base to 12°Brix). Numerous crownlets protrude from
top. The plant is medium-sized and vigor- the base of the crown and the upper eyes.
ous, with dark green, erect and spiny leaves. Slips are also numerous (from four to 11).
In some strains, long and erect basal slips The plant is medium to high. The leaves are
surround the fruit. ‘Pérola’ is a very robust completely smooth because the lower epi-
cultivar, showing resistance to Phytophthora dermis is folded over the leaf edge, a trait
(Collins, 1960), probably related to its erect that was named ‘piping’ by Collins and
habit, and tolerance to drought, mealybug Kerns (1946). ‘Perolera’ is susceptible to
wilt and nematodes (Collins and Hagan, fruitlet core rot and to the fly Melanoloma
1932; Sipes and Schmitt, 1994; Sarah et al., canopilosum Hendel. It has been used in the
1997). However, it is threatened because of Brazilian breeding programme because of its
its high susceptibility to fusariosis, which resistance to fusariosis.
affects Brazilian production severely. The cultivar ‘Manzana’ (Plate 12), also
‘Perolera’, also named ‘Lebrija’, called ‘Bumanguesa’, is cultivated in the same
‘Motilona’, ‘Capachera’ or ‘Tachirense’, is an zone as ‘Perolera’, as well as in the western
 Breeding and Variety Improvement 39

is firm and delicious. The Peruvian ‘Samba’

and ‘Roja Trujillana’ produce red fruits and
show resistance to S. basilides (Bello et al.,
1997a). Certain Amazonian cultivars, such as
‘Gigante de Tarauacá’ or ‘Cabeça de Onça’,
yield enormous fruits up to 15 kg (Ritzinger,
1987). Others are distinguished by an origi-
nal red or purple colour of their fruit, such as
the Amazonian ‘Cabeça de Arara’ (meaning
‘Macaw Head’) and ‘Roxo de Tefé’, the
Trinidadian ‘Mundo Nuevo Red’ and the
Venezuelan ‘Morada’. These colours are very
probably determined by a single dominant
gene, as observed in the case of ‘Roxo de
Tefé’ (Cabral et al., 1997). Further evaluation
of the germplasm recently collected in the
basins of the Amazon and the Orinoco will
provide many new traits of interest for vari-
etal diversification.
The hybrid ‘MD-2’, ‘Golden Ripe’ or
‘Extra Sweet’ and its sibling ‘CO-2’ (US
Patent PP8863) (Plate 13) were developed by
Del Monte Fresh Produce Hawaii Inc. from a
cross made between the PRI hybrids 58–1184
and 59–443 for the fresh-fruit market. ‘MD-2’
gives a medium to large (1.3–2.5 kg) cylindri-
cal, square-shouldered fruit, with large flat
eyes and an intense orange-yellow colour.
Fig. 3.5. ‘Manzana’.
The clear yellow pulp is sweet, compact and
fibrous. It is high in sugar (15–17°Brix) and
Andes of Colombia. It is said to be a sport ascorbic acid but lower in total acid than
from ‘Perolera’, from which it differs in the ‘Smooth Cayenne’. The core is tender, edible
regular globular to cylindrical shape and and thinner than in ‘Smooth Cayenne’.
large flat eyes of its fruit, which is of an However, the crown is larger and must be
attractive dark to bright red colour when broken at harvest. The leaves are yellow-
produced in tropical highlands (Fig. 3.5). It is green with a reddish tip and are mostly
susceptible to the nematode Pratylenchus spineless. ‘MD-2’ is said to be more produc-
neglectus but tolerant to Meloidogyne incognita tive than ‘Smooth Cayenne’. It is resistant to
Kofoid & White (Redondo-Echeverri and internal browning, but susceptible to fruitlet
Varón, 1992). core rot and more sensitive to Phytophthora
Other traditional cultivars of lesser than ‘Smooth Cayenne’ (Anon., 1996). ‘CO-2’
regional importance are sometimes is said to resemble the parents, but is distin-
described. ‘Cabezona’ is a vigorous spiny guished from them in being sweeter and
triploid producing yellow to orange fruits of higher in ascorbic acid and in having a ‘dis-
more than 3 kg with white pulp. ‘Monte tinct tropical flavour’, greater pigmentation
Lirio’, also called ‘Cambray’ or ‘Milagreña’, and fibre content of the flesh, greater resis-
is a smooth (‘piping’) cultivar with a mid- tance to internal browning and a good over-
size fruit and sweet white pulp, found from all appearance after refrigerated storage.
Mexico to Ecuador. ‘Black Antigua’ is an old ’Josapine’ (Plate 14) is a hybrid between
spiny cultivar producing a small to medium ‘Johor’ (‘Spanish’) and ‘Sarawak’ (‘Smooth
fruit with a dark green shell turning yellow Cayenne’) released by MARDI in 1996 (Chan
to orange at maturity. Its golden-yellow pulp and Lee, 1996). It is recommended as a table
40 Y.K. Chan et al.

variety. It fruits very early and is cultivated and flat. The flesh is firm and golden-yellow
on an annual cycle in Malaysia. The plant is with a high sugar content (15–18°Brix).
vigorous and produces two to three shoots. Acidity is similar to that of ‘Smooth
Leaves are lightly purple-tinged, usually Cayenne’ but it is higher in ascorbic acid
with spineless margins except for the leaf tip. content. The core is thinner, the flesh less
Crowns are medium, occasionally with mul- fibrous and the crown is lighter and erect.
tiple proliferation. Fruits weigh between 1.1 The plant is compact, with erect, smooth
and 1.3 kg and are cylindrical-shaped with (‘piping’) leaves. The peduncle is long but
dark purple peel ripening to an attractive resistant to lodging. ‘Scarlett’ has a short
orange-red (Fig. 3.6). The flesh is deep production cycle and responds well to floral
golden-yellow and has a strong aroma and a induction. It is susceptible to fruitlet core rot
sugar content between 17°Brix on peat soil (Coppens d’Eeckenbrugge and Marie, 2000).
and 22°Brix on mineral soils. ‘Josapine’ fruits
have a good storage life and are resistant to
black-heart disorder or internal browning Genetic resources from other species and
caused by low temperatures. botanical varieties
’Scarlett’ is a hybrid between ‘Smooth
Cayenne’ and ‘Manzana’ developed by The wild pineapples, Ananas comosus var.
CIRAD-FLHOR. Fruits are 1.4–2 kg and ananassoides (Plate 2) and A. comosus var. par-
orange to red in colour, with a regular cylin- guazensis (Plate 4), constitute the main reser-
drical shape. The eyes are medium to large voir of genetic diversity. The former is
generally considered the most drought-
resistant pineapple. Its fruit is often seedy,
with white to yellow pulp, which is firm and
fibrous. Sugar and acid contents are high and
it has a good flavour and aroma. The core is
narrow. According to Collins (1960), crosses
between A. comosus var. comosus and A. como-
sus var. ananassoides yield hybrids that are
entirely comparable with crosses between
cultivars, with highly variable sugar and
acidity values. Mean acidity value is inter-
mediate between those of the parents, but
the average sugar content of the hybrids is
much closer to that of the A. comosus var.
ananassoides parent. A. comosus var. ananas-
soides is a potential source of resistance or
tolerance to wilt, nematodes (Meloidogyne
javanica, Rotylenchulus reniformis) and crown
and root rot (Collins and Hagan, 1932;
Hagan and Collins, 1932; Collins, 1960;
Ayala, 1961; Ayala et al., 1969; Sipes and
Schmitt, 1994). A. comosus var. parguazensis is
less variable. It has a globular fruit with rela-
tively wide flat eyes and a white and fibrous
pulp. Resistance to fusariosis is variable in A.
comosus var. ananassoides and A. comosus var.
parguazensis (de Matos et al., 1991).
A. comosus var. bracteatus (Plate 6), first
cultivated for the fruit and as a living fence,
is now widely used as a tropical garden
Fig. 3.6. ‘Josapine’. ornamental. Its variability is very limited. It
 Breeding and Variety Improvement 41

grows well on poor and dry soil and it is recombination. A. macrodontes also combines
well adapted to cool conditions and altitude. vegetative and sexual reproduction, but
The fruit is small to medium, with white apparently with a higher efficiency of the lat-
fibrous flesh, a small core and an average ter than in A. comosus, and it is not self-
sugar and acid content. It is resistant to heart incompatible.
rot, root rot and fusariosis (de Matos et al.,
1991). Studies on nematode resistance pro-
duced contradictory results (Collins, 1960; Cytogenetics
Sipes and Schmitt, 1994). Crosses with A.
comosus var. comosus yield hybrids with fruits Chromosome number
of good size and weight and a wide variation Like many other Bromeliaceae, all the botanical
in flavour (Collins, 1960). varieties of A. comosus have a diploid number
A. comosus var. erectifolius (Plate 3) has of 50 minute, almost spherical, chromosomes,
been cultivated for its long and strong fibres. with a length of 0.5–1.7 m. Among three
In the past decades, a dark red cultivar has pineapple accessions, Sharma and Ghosh
also been exploited as an ornamental for the (1971) identified four chromosome types and
European cut-flower market. This trait is observed a slight variation in their distribu-
controlled by a single dominant gene (Cabral tion and in chromatin content. They also
et al., 1997). The use of A. comosus var. erecti- observed secondary constrictions in two to
folius in breeding for dual-purpose (fruit and five pairs of chromosomes. Arumuganathan
fibre) cultivars has been envisaged in the and Earle (1991) reported values of 1.09 and
Philippines (V. Villegas, Fort-de-France, 1992, 0.92 pg (corresponding to c. 526 and 444 Mbp)
personal communication). A. comosus var. for the DNA content of 2C nuclei of A. como-
erectifolius is resistant to heart rot and root rot sus var. comosus and A. comosus var. bracteatus,
(Collins, 1960) and relatively tolerant to respectively. Triploids (e.g. cultivars
nematodes (Sipes and Schmitt, 1994). ‘Cabezona’, ‘Caicara’, ‘Monte Oscuro’),
Ananas macrodontes (Plate 1) has been tetraploids and even heteroploids (60 chromo-
mainly used as a source of fibre. As with all somes) may also occur (Collins and Kerns,
other pineapples, it is resistant to drought 1931; Canpinpin and Rotor, 1937; Collins,
but it also tolerates temporary flooding. It is 1960; Antoni, 1983; Lin et al., 1987; Dujardin,
resistant to heart rot, root rot, wilt and fusar- 1991). Collins and Kerns (1931) reported six
iosis (Collins, 1960; de Matos et al., 1991). cases of triploidy out of 8000 plants resulting
When selfed, A. macrodontes shows little vari- from a cross between ‘Smooth Cayenne’ and a
ation, indicating that it is relatively homozy- clone of A. comosus var. ananassoides. Collins
gous (Collins, 1960). Hybridization with A. (1933b) demonstrated that these triploids had
comosus var. comosus produces 5–10% fertile arisen from unreduced female egg cells fertil-
seeds, most of which are tetraploid. These ized by normal haploid pollen. Tetraploids
tetraploids are fully fertile. could then be obtained from unreduced male
and female gametes, although their occur-
rence is rarer than triploids. These tetraploids
Breeding System produce about 90% viable pollen, of a larger
size than pollen cells of diploid plants. By
The breeding system of A. comosus combines crossing them with diploids, hybrid
efficient vegetative reproduction with func- tetraploid seedlings were obtained, which
tional allogamous sexual reproduction. grew faster and larger than diploids. A.
Although the former is largely dominant, the macrodontes is a natural tetraploid with 100
latter is responsible for most of the present chromosomes (Collins, 1960).
genetic variation. Pineapple allogamy, as
well as its consequent heterozygosity, is
Ploidy level and morphology
related to the existence of self-incompatibil-
ity and pollination by hummingbirds. A high Collins (1933b, 1960) and Kerns and Collins
chromosome number also favours genetic (1947) compared triploid and diploid
42 Y.K. Chan et al.

hybrids and diploid and autotetraploid 1993). There is also variation in all the
plants of ‘Smooth Cayenne’. The polyploids species in the relative frequencies of anat-
were taller, with wider leaves, placed at a ropous and orthotropous ovules. The
greater internodal length, so the total leaf embryo sac develops according to the
area does not differ much. The cross-section Polygonum type. Shrinking embryo sacs have
of the tetraploid leaves is more closed. Their been commonly observed at the eight-nucle-
larger fibres increase leaf rigidity. The size of ate stage in ‘Masmerah’ (Rao and Wee, 1979)
cells, trichomes and stomata also increases and the six main cultivars of A. comosus var.
with the ploidy level, while the stomata den- comosus and in A. comosus var. ananassoides,
sity is reduced. Water content is higher in A. comosus var. erectifolius and A. comosus var.
polyploids. Triploid hybrids and tetraploid bracteatus (F. Van Miegroet, personal commu-
‘Smooth Cayenne’ have longer vegetative nication). The proportion of ovules with nor-
growth periods, mature more slowly and mal embryo sacs is correlated with the
have a lower fruit sugar content. Tetraploid proportion of stained pollen and with fertil-
fruits are smaller, with fewer but larger ity (F. Van Miegroet and Coppens
fruitlets. d’Eeckenbrugge, unpublished results).
Fertility rate (i.e. the percentage of ovules
producing a seed) is generally lower in A.
Micro- and megagametogenesis and fertility
comosus var. comosus than in the other botan-
Meiosis is primarily regular in diploid and ical varieties. In the latter, fertility ranged
tetraploid A. comosus, as well as in A. from 6% (0.85 seeds per flower) in A. como-
macrodontes, with only rare irregularities, sus var. ananassoides to 35% (18.4 seeds per
such as lagging chromosomes and some flower) in A. comosus var. bracteatus. Within
abnormal tetrads (Collins and Kerns, 1931; A. comosus var. comosus, the cultivars with
Canpinpin and Rotor, 1937; Collins, 1960). ‘piping’ leaves often exhibit a higher fertility
However, some irregular gametes are pro- (4–11%; two to five seeds per flower) than
duced. The proportion of giant pollen cells the common cultivars ‘Smooth Cayenne’,
may reach 6.5% and that of giant ovules 1%. ‘Española Roja’, ‘Singapore Spanish’, ‘Pérola’
Bhowmik (1977) observed large variations in and ‘Queen’ (less than 5%; zero to two seeds
size and fertility of pollen grains in ‘Kew’ per flower). However, the highest fertility
(‘Smooth Cayenne’) and ‘Queen’ (from 35 to rate (29%; 9.45 seeds per flower), which is
81 m and from 36 to 68 m, respectively) comparable to that of wild types, was
and correlated them with the frequencies of observed in a spiny landrace. These fertility
univalents and secondary associations of up rates were estimated under an open-pollina-
to three or seven bivalents in these cultivars. tion situation and higher values may be
Pollen size and fertility also vary between expected under assisted pollination. Fertility
cultivars and clones (Collins, 1960; Ramírez, is correlated with ovule fertility, pollen
1966; Wee and Rao, 1979; Nayar et al., 1981; stainability and the amount of pollen pro-
Antoni, 1983; Coppens d’Eeckenbrugge et al., duced per flower. It is not correlated with
1993). Triploid genotypes are sterile because ovule number, probably due to crowding or
of irregular meiosis. Tetraploids produce competition effects between fertilized
90% of good pollen grains, which are uni- ovules. Indeed, many genotypes of A. como-
form but larger than those of diploids sus var. comosus and A. comosus var. bractea-
(Collins, 1960). tus can develop only a small proportion of
The number of ovules varies widely in A. their numerous ovules into seeds (Coppens
comosus. While A. comosus var. bracteatus may d’Eeckenbrugge et al., 1993). Fertility may be
have 40–70 ovules per flower, this number affected by abnormal anthesis. Thus, in cer-
generally ranges from 14 to 30 in the wild tain clones of ‘Española Roja’ and
botanical varieties and in A. comosus var. erec- ‘Singapore Spanish’, a significant proportion
tifolius and from 16 to 71 in A. comosus var. of flowers do not open, hampering pollina-
comosus (Coppens d’Eeckenbrugge et al., tion.
 Breeding and Variety Improvement 43

Self-incompatibility another was pseudo-self-compatible. A.

macrodontes is highly self-fertile and pro-
In A. comosus, the self-incompatibility is
duces uniform progenies, suggesting that
brought about by the inhibition of pollen-
this species is autogamous.
tube growth in the upper third of the style
(Kerns, 1932; Majumder et al., 1964). It is
gametophytically controlled by a single Hybridization between botanical varieties and
locus with multiple alleles (Brewbaker and between species
Gorrez, 1967) and is generally stronger in A.
There is little difficulty in crossing cultivars
comosus var. comosus than in the other botani-
of A. comosus var. comosus with clones from
cal varieties. Later work, however, suggested the other botanical varieties. Instead, these
that more than one gene might be involved crosses often show high fertility, as a result
in the control of self-incompatibility (D.D.F. of higher gamete viability of the latter. When
Williams, Colorado, 2001, personal commu- A. comosus is crossed with A. macrodontes, a
nication). However, many cultivars exhibit few fertile seeds are produced, which usu-
pseudo-self-compatibility, expressed in the ally yield vigorous tetraploids and some-
variable production of self-seeds, although times self-sterile triploids. The tetraploid
the resulting self-fertility is always lower hybrids are highly fertile and self-fertile
than cross-fertility. For instance, pseudo-self- (Collins, 1960).
incompatibility is common in ‘Singapore
Spanish’, ‘Perolera’, ‘Manzana’, ‘Primavera’,
‘Rondon’, ‘Cambray’, ‘Amarelo de Uaupés’, Breeding and Selection Techniques
‘Roxo de Tefé’, ‘Morada’ and ‘Alto Turi’ and
in some clones of ‘Queen’ (DeWald et al., Clonal selection and roguing
1992; Coppens d’Eeckenbrugge et al., 1993,
1997; Muller, 1994). In a sample of 84 clones Many commercial pineapple cultivars are
from 51 cultivars and primitive landraces improved through selecting genetic variants
studied by Muller (1994), 55 appeared that occur occasionally in experimental plots
strongly self-incompatible and 21 pseudo- or production fields. Such variants are per-
self-compatible and eight exhibited a self-fer- petuated as clones whose genetic make-up is
tility comparable to their fertility, which identical and ‘fixed’ by vegetative propaga-
could be due to self-compatibility or a very tion until new variation arises. Clonal planti-
weak self-incompatibility. Indeed, a domi- ngs form the basis of modern pineapple
nant mutation suppresses the incompatibil- culture, and many different clones are used
ity reaction in the pollen of ‘Smooth throughout the world. All ‘Smooth Cayenne’
Cayenne’ and results in the production of clones in Hawaii, including ‘F200’ and the
selfed seeds (Collins, 1960). The self-incom- ‘Champaka’ clones were developed through
patibility of A. comosus var. comosus does not clonal selection (Py et al., 1987). In Australia,
appear to be affected by tetraploidy (Kerns clonal selection began in 1950 with the selec-
and Collins, 1947). tion of 100 plants from commercial fields.
Pseudo-self-compatibility is more fre- Four clones – ‘C8’, ‘C10’, ‘C13’ and ‘C30’ –
quent in the wild pineapples. In Muller’s were eventually released to the industry in
study, five out of 11 A. comosus var. ananas- 1975 (Groszmann, 1982) and, in conjunction
soides clones and four out of seven A. comosus with the Hawaiian clone ‘Champaka F180’,
var. parguazensis clones exhibited pseudo- form the basis of the present-day industry.
self-compatibility. One A. comosus var. par- Several clones of the cultivar ‘Queen’ have
guazensis clone appeared self-compatible. In also been established in Australia, including
A. comosus var. bracteatus, three clones that ‘Ripley Queen’, ‘Alexander’ and ‘McGregor’,
were phenotypically identical appeared to be through a process of clonal selection by pri-
self-compatible. A variegated clone of the vate growers (Lewcock, 1940). In Malaysia,
same species was self-incompatible, while an improved cultivar, ‘Masmerah’, with
44 Y.K. Chan et al.

higher vigour and yield, was obtained by (slips arise from the peduncle very close to
clonal selection from a commercial field of the base of the fruit) will be multiplied faster
‘Singapore Spanish’ (Wee, 1974). Clonal than normal types and hence the frequency
selection has also been used in the will increase. Therefore, while the genotype
Philippines to develop a new clone of is considered stable, some specific characters,
‘Smooth Cayenne’ less prone to the green- such as leaf spininess, are considered unsta-
ripe disorder (Obrero, 1995). ble and the frequency of mutation for these
characters is high (Collins and Kerns, 1938).
Spontaneous mutations
Roguing
At least 30 major mutations have been
recorded in ‘Smooth Cayenne’ pineapple As most mutations are regressive and con-
since the early 1920s. These include white sidered to be off-types, the population per-
flowers (as opposed to purple), foliar floret formance will gradually decline if no
proliferation, multiple sepals and bracts, selection is done to maintain its standards.
enlarged fruit segments, protruding fruit Inferior types may be difficult to detect visu-
segments, increased or decreased trichome ally, especially those that have slightly
density, seediness, multiple crowns, a lower reduced plant vigour or fruit size. With the
harvest index, misshapen fruits, smaller- large number of plants grown and the effi-
diameter fruits, leaf spininess, reduced cient propagation methods used, inferior
chlorophyll or anthocyanin levels, dwarfed types that escape detection can be quickly
habit, translucent fruits, collar of slips, multiplied. Roguing can rapidly reduce the
increased or decreased slip number and percentage of most off-types, where they are
fruits with an increased incidence of basal initially numerous, to low levels. Therefore,
knobs (Anon., 1925, 1940; Kerns, 1928; major pineapple producers routinely rogue
Collins and Kerns, 1938; Groszmann, 1939; inferior spontaneous mutants to maintain
Dalldorf, 1975; Sanewski et al., 1992). Other productivity and efficiency.
measurable differences between clones likely Roguing is sometimes called mass selec-
to have been caused by mutations are sucker tion, although, strictly speaking, it is a form
number, length of the fruit development of culling rather than selection. Roguing
period and peduncle length and width aims to minimize the number of undesirable
(Sanewski, 1997). Only two mutations – non- variants in a field by not replanting material
translucent fruit and resistance to mealybug from those plants. Roguing is, by necessity,
wilt – could be considered substantially pro- an ongoing process and is best performed on
gressive (Collins and Carter, 1954; Obrero, well-grown plant crops. To identify the infe-
1995). Mutations can also be induced with a rior plants the grower walks through the
range of techniques, including tissue culture field before harvest and marks the crown or
(Williams and Fleisch, 1993), exposure of in plant itself with paint. The exact process will
vitro leaf cultures to cobalt gamma rays depend on what type of planting material is
(Cisneros et al., 1998) and the treatment of used. In Australia, it is performed on sum-
seed and plants with chemical mutagens mer plant crops, where the expression of
(Singh and Iyer, 1997). defects such as collar of slips is greatest
Despite the large number of mutations (Groszmann, 1945). The main characteristics
identified in pineapple cultivars, they are not targeted in roguing are described below.
generally considered to be unstable geno-
types. The incidence of mutations is proba- FRUIT WEIGHT. Fruit weight is strongly
bly a function of the large number of plants affected by environment and cultural opera-
propagated and observed. The type of plant- tions. Also, subjective assessment of fruit
ing material used may also determine the weight in the field is difficult even where the
frequency of some off-types. As an example, differences are as much as 10%. For these
if slips are used as planting material, then reasons, roguing against decreased fruit
plants that produce the mutation near-collar weight is considered not to be very effective
 Breeding and Variety Improvement 45

(Kerns, 1928). There is, however, a tendency unless the parent plants have been checked
for the very heavy and very light types before harvest for the incidence of collar of
within a field to be genetically different. slips. Eliminating this off-type should mini-
Roguing can therefore be used to discrimi- mize its incidence. Collar of slips is one of
nate against very small-fruited types (Anon., the main characteristics targeted in a roguing
1925). Where more objective methods of programme (Py et al., 1987).
assessment of fruit weight are used, such as The number of slips produced lower
in a clonal selection programme, gains in down on the peduncle is not considered the
fruit weight are possible, provided measures same as collar of slips, as it is a quantitative
are taken to avoid inadvertent selection for trait whose expression is more strongly influ-
more translucent types. As an example, a enced by plant vigour and environment
10–15% increase in fruit weight was (Collins and Kerns, 1950). Vigorous plants on
achieved in the Australian clonal selection the outer edges of fields commonly produce
programme (Wassman, 1982). more slips. The clone ‘F200’, which is grown
widely in the Philippines, reputedly does not
MULTIPLE CROWNS. Multiple or fasciated produce slips. Roguing against increased or
crowns can be heritable but the trait is also decreased slip number is not likely to make
strongly influenced by environment and cul- much difference, because of the large envi-
tural operations (Dalldorf, 1975). Excessive ronmental and cultural influences. However,
nitrogen fertilization and high temperature a well-conceived clonal selection programme
during flower-cone development are known should make progress in this regard.
to increase the incidence of multiple crowns
(Anon., 1925; Linford and Spiegelberg, 1933). FRUIT CONFORMITY. There are several distinct
Linford and Spiegelberg (1933) observed that mutations that affect fruit shape or appear-
the condition can start at the top of the fruit ance. Long Tom or long, slender fruits is a
or, as evidenced by the change in eye pat- heritable condition where the fruits are long
tern, at a position in the fruit itself. with a small diameter. Other common muta-
Replanting only single crowns from vigor- tions in fruit conformity include dry fruit
ously growing plants that flowered during and bottleneck. With these off-types, the
the hotter periods of the year should reduce flowers are absent from either the upper por-
its incidence (Dalldorf, 1975). Roguing can- tion of the fruit or the entire fruit. The effect
not completely eliminate this disorder. is a small malformed fruit with obvious
bracts but poorly developed eyes. These
COLLAR OF SLIPS AND KNOBBINESS. Collar of slips genetic disorders should not be confused
is a condition where several slips emerge with the physiological disorder associated
from the base of the fruit. This severe defect with sun damage (Broadley et al., 1993).
is caused by a dominant mutation (Collins Roguing against these off-types should even-
and Kerns, 1938). The incidence of knobs on tually eliminate them.
the base of the fruit is closely associated with
collar of slips and is sometimes considered a SPINY LEAVES. Genetic studies have shown that
milder expression. Collar of slips is highly spiny leaf is recessive to smooth leaf and
heritable but influenced by environment. In ‘Smooth Cayenne’ carries the gene for spines
Australia, collar of slips is usually expressed in a recessive condition (Collins and Kerns,
in summer fruit but not winter fruit 1946). The mutation of the dominant gene
(Groszmann, 1940). Also, plants propagated for smooth leaves to the recessive gene for
from the collar-of-slips type will occasionally spines can occur at any time in the growth of
produce about 10% normal types (Collins, the plant. Depending on the developmental
1933a; Groszmann, 1945). Fruit from collar- stage of the plant when the mutation occurs
of-slips types is usually smaller and slip and the position in the plant of the mutating
number is more than double that of normal cell, it can result in a chimera with partially
plants (Collins, 1933a; Py et al., 1987). It is spiny leaves or a completely spiny plant
best to avoid using slips as planting material (Collins and Kerns, 1933).
46 Y.K. Chan et al.

The mutation to spiny leaves should not sorting several thousand fruit using a weight
be confused with the tendency of ‘Smooth grader appears more reliable than walking
Cayenne’ to develop spines as a result of through fields and selecting plants by subjec-
stressful conditions and the cessation of leaf tive assessment of fruit weight (G.M.
growth. As an example, a high night temper- Sanewski, unpublished results).
ature increases the development of spines
along the leaf (Friend, 1981). A genetic
change is not thought to have occurred in Hybridization
these instances and new leaves develop nor-
mally once the growing conditions are Cultivated pineapple is self-incompatible but
favourable. Roguing is very effective in sets seeds readily when cultivars from differ-
removing spiny-leaf off-types. ent groups are crossed. Pineapple cultivars
are heterozygous, and hybridization is a
valuable method in generating widely vari-
Clonal selection
able genotypes through gene recombination.
While roguing is a form of clonal mainte- With clear objectives and judicious selection
nance, clonal selection offers the possibility of the parents, the hybrid population would
of clonal improvement. Clonal selection aims present a spectrum of variation within which
to identify superior individual plants that the desired improved genotypes may be
are eventually multiplied as a new clone. found. It was believed that many important
Clonal selection, however, is a slow process characteristics, such as high carotene con-
and is usually beyond the resources of the tent, greater translucency and lower acidity,
commercial grower. that could not be achieved by clonal selec-
It has been demonstrated that apparently tion could be realized by hybridization
normal plants can carry some undesirable (Wortman and Kerns, 1959).
traits, such as multiple crown. If enough
plants of that genotype are observed over
Considerations in hybridization
several environments, these traits will even-
tually be expressed and that line or clone can FERTILITY, COMPATIBILITY AND DIRECTION OF CROSS.
be eliminated. Clonal selection is therefore Fertility and compatibility are important con-
more successful than roguing against defects siderations in deciding the parents to be used
such as multiple crowns that are strongly for hybridization and also the direction of the
influenced by environment. cross, i.e. deciding which parent should be
The initial population in a clonal-selection paternal or maternal. Chan (1986) found that,
programme is established usually by walking for crosses between ‘Smooth Cayenne’,
through fields of maturing pineapples and ‘Queen’, ‘Singapore Spanish’ and ‘Johor’, a
selecting the crown from superior types or hybrid between ‘Smooth Cayenne’ and
marking the plant with paint and a stake to ‘Singapore Spanish’ was compatible, but there
collect the entire plant later. These selections was differential seed set between the crosses
are then assessed and multiplied over several and their reciprocals. ‘Smooth Cayenne’, for
crops until only the most consistent clones instance, has low fertility and was a poor
remain. When sufficient plants are available, female parent because only a few seeds were
replicated trials can be established in differ- formed. However, reciprocal crosses using
ent environments. The method of obtaining ‘Smooth Cayenne’ as the pollen donor pro-
the initial population will, of course, vary vided satisfactory seed set in all crosses. In the
depending on the specific characters in which extreme case, the ‘Smooth Cayenne’ female
the breeder is most interested. Novel meth- crossed with ‘Johor’ was incompatible but the
ods of selection can be used for some traits. reciprocal produced about 1000 seeds in a
As an example, a different approach to select- well-pollinated fruit (Chan, 1986). ‘Smooth
ing for larger-fruited types has recently been Cayenne’ was also shown to be a poor female
used in the Australian clonal-selection pro- parent, bearing only about 200 seeds per fruit.
gramme. Selection of larger-fruited types by When a reciprocal cross was made with the
 Breeding and Variety Improvement 47

much more fertile ‘Perolera’ as the female example, the range of values showed trans-
parent, there were more than 2000 seeds per gressive segregation, i.e. the minimum and
fruit (Cabral et al., 1993). maximum values of the hybrid progenies
The direction of the cross was reported by exceeded the lowest and highest values of
Loison-Cabot and Lacoeuilhe (1990) to have both parents. Sanewski (1998) reported that
an influence on the performance of the the range of TSS from hybrids of the ‘Queen’
hybrids. In a cross between ‘Smooth × ‘Smooth Cayenne’ cross was 9–24%, while
Cayenne’ and ‘Manzana’, the latter used as the parents had a narrower range of 14–19%.
the female parent improved the yield of the The coefficients of variance in the hybrid
hybrid population compared with the recip- population for many quantitative traits quite
rocal. In the case of total soluble solids (TSS), often exceeded twice that of the parents
‘Smooth Cayenne’ as the female parent gen- (Chan, 1989, 1991). As a matter of fact, the
erated progenies with higher TSS content. variation generated through hybridization of
even closely related cultivars may be as large
SYNCHRONY IN FLOWERING. Synchrony in flow- as that obtained for some pineapple collec-
ering of parents is obviously an important tions (Chan, 1989). This wide variation will
consideration in hybridization. The time provide good opportunities for selection of
taken for appearance of red-heart from time improved recombinants.
of hormoning is variable between various
pineapple cultivars. Inflorescence appear-
Choosing the most suitable parents
ance in ‘Smooth Cayenne’ is 10–12 days later
than in ‘Singapore Spanish’ and ‘Queen’, In hybridization for improvement of pineap-
respectively. This means that, while ple, the choice of parents has usually been
‘Singapore Spanish’ and ‘Queen’ may be based on the strength of one in complement-
induced to flower simultaneously, ‘Smooth ing the weakness of the other. For example,
Cayenne’ should be induced about 2 weeks the ascorbic acid content of ‘Smooth
earlier in order that inflorescences of all vari- Cayenne’ is improved using ‘Perolera’
eties appear simultaneously for hybridiza- (‘Manzana’), which has higher ascorbic acid
tion. Where the time to anthesis is unknown (Loison-Cabot, 1987), and ‘Queen’ for
for each parent cultivar, a percentage of increasing TSS of ‘Smooth Cayenne’ winter
plants of each cultivar should be induced at, fruits (Winks et al., 1985). Where biparental
say, 5-day intervals. crosses are focused narrowly on the
improvement of a single character, there is
TIME OF DAY FOR HYBRIDIZATION. Dawn to 9 a.m. danger that the donor parent might inadver-
is probably the best time for pollination tently introduce other adverse characters.
(Cabral et al., 1993; Sanewski, 1998). In practice, breeders are always faced
However, pollination may be done during with selection of a set of economic traits and
the entire length of the day, although seed not just single characters. In this case, selec-
set may vary depending on the time of day tion of suitable parents may be complicated.
and appears to be temperature-related Chan (1991) computed breeding values for
(Chan, 1986). Higher seed set was obtained parents based on consideration of eight agro-
when crossing was done during the early nomic characters. The values were computed
morning or cooler evening and a signifi- by obtaining the product of the percentage of
cantly lower seed set when it was done dur- progenies that qualified for selection in each
ing the hotter midday or early afternoon. of the eight characters. ‘Queen’ and ‘Smooth
Cayenne’ were better all-round parents for
SEGREGATION IN HYBRID POPULATIONS. Evaluation hybridization because 1.1–1.2 % of the proge-
of hybrid populations from biparental nies qualified for selection when eight char-
hybridization indicated that there is wide acters were considered. This was twice as
segregation of progenies for most quantita- much compared with the ‘Singapore
tive characters (Chan, 1989, 1991; Loison- Spanish’ parent. Sanewski (1998) used a sim-
Cabot, 1990; Sanewski, 1998). For TSS, for ilar computation and found that advanced
48 Y.K. Chan et al.

hybrids, such as ‘73–50’ developed by PRI, Cayenne’ and ‘Perolera’ parents, recom-
had a high breeding value. Sanewski (1998) mended 30,000 progenies for each annual
concluded that these hybrids are valuable as selection cycle, while Cabral et al. (1993) and
parents because they have accumulated Benega et al. (1998b) maintained 23,000 and
desirable genes during earlier multiple cross- 34,000 progenies, respectively, to obtain
ings and subsequent selection. Breeding val- 120–169 first-cycle selections. The PRI breed-
ues are helpful for guidance in the choice of ing programme has been the most extensive
parents and in ascertaining the probability of to date and, at its peak, was producing over
obtaining useful progenies. They are also 1 million seedlings per year. Up to 100,000
useful for determining the population size of seedlings were developed from some of the
the hybrid progenies. Quite evidently, higher better parent combinations (Williams and
breeding values of parents will yield higher Fleisch, 1993). Sanewski (1998), using some
percentages of promising hybrids, thereby of these PRI selections, obtained 300 selec-
reducing the need for larger populations. tions from populations of 50,000 seedlings
The high level of heterozygosity of because the parents have very high breeding
pineapple cultivars reduces the chances of values.
quickly obtaining progenies with a large Some interest has been shown in the use of
complement of specific characteristics and it other species for imparting characteristics
is this heterozygosity that is largely responsi- such as disease resistance. Collins (1960) indi-
ble for the slow rate of progress in most cated that, where crosses with wild
breeding programmes. Strategies such as germplasm were performed, only 1000 proge-
selfing (Cabral et al., 2000), backcrossing and nies were necessary in the first generation but
sib-crossing need to be investigated more 15,000–25,000 per backcross were required for
thoroughly, especially considering the each of four backcross generations.
demonstrated inefficiencies of combining
heterozygous genotypes. Other strategies to
Crossing technique
circumvent the heterozygosity of potential
parents such as the development of homozy- Pollen is collected from flowers at anthesis
gous haploid parents by anther and ovule early in the morning. The flowers are
culture have now been reported (Benega et removed from the inflorescence by making
al., 1998a; Cabral et al., 2000). three deep triangular incisions into the base
of the flower and then gouging them out.
This method of collecting pollen from
Suitable hybrid population size
excised flowers is better than removing only
Hybridization produces recombinants, but the stamens. Pollen in excised flowers
potential selections that meet all the selection should stay fresh longer and dehydrates less
criteria of the breeder will appear very rapidly. There are no conclusive data on the
rarely. Therefore, very large populations are storage of pollen, but Kerns (1931) indicated
usually required to increase the chances in that ‘Smooth Cayenne’ pollen could be
the draw of this genetic lottery. With the stored in cool, dry air for 15 days with mini-
breeding values of parents known, the popu- mal loss of viability.
lation size that may be required can be esti- During hand-pollination, the petals of the
mated (Chan, 1991). The number of flower are removed to expose the freshly
selections in the second cycle that the dehisced anthers with an abundance of
breeder would want to evaluate must first be pollen. The anther is picked up with a fine
ascertained. At MARDI, it was put at 300 pair of forceps and gently brushed on to the
selections and hybrid populations of stigma of the flower to be pollinated. In most
23,000–45,000 progenies, depending on the cases, there is no necessity to emasculate the
parents’ breeding values, will be needed to flowers before pollination because most cul-
have a realistic chance that the desired num- tivars are self-incompatible. In cases where
ber of potential recombinants would be gen- fidelity of the pollen source is vital, as in
erated. Loison-Cabot (1987), using ‘Smooth genetic studies, or when natural pollinators
 Breeding and Variety Improvement 49

are present, the contamination of foreign In pseudo-self-compatible cultivars, a few

pollen can be prevented by using a fine mos- inflorescences would be sufficient to obtain
quito netting or paper bag placed over the the small number of self-seeds required. In
inflorescence (Leal and Coppens addition, selfing is much less labour-inten-
d’Eeckenbrugge, 1996). Depending on vari- sive than crossing since these few seeds can
eties, the number of flowers in an inflores- be obtained by simply bagging the inflores-
cence ranges between 75 and 150. About cence before anthesis (Coppens
three to seven flowers open daily and 2–3 d’Eeckenbrugge et al., 1993). However, for
weeks may be required to complete pollina- strongly self-incompatible cultivars, a high
tion of the whole inflorescence (Chan, 1986). number of selfings may be necessary.
Attempts have been made to overcome self-
incompatibility with X-rays (Marr, 1964) or
Inbreeding gamma rays (Subramanian et al., 1981).
Bhowmik and Bhagabati (1975) tested bud
The difficulty in finding a superior recombi- pollination, style-grafting and naphthalene
nant among the widely varying progenies acetic acid (NAA) application, but only the
obtained through direct hybridization of last technique produced a few self-seeds in
highly heterozygous pineapple cultivars is ‘Queen’. Others have proposed obtaining
the main cause of the failure of many breed- homozygous plants through the production
ing programmes (Collins, 1960; Williams and of doubled haploids. However, Collins’s
Fleisch, 1993; Coppens d’Eeckenbrugge and results throw some doubt on the viability of
Duval, 1995). This led some researchers to this approach. Highly depressed and sterile
evaluate the potential of inbreeding in homozygotes obtained through this method
pineapple breeding. Thus, Collins (1960) would be of little use for breeding.
selfed a mutant of ‘Smooth Cayenne’ and Coppens d’Eeckenbrugge et al. (1993)
observed in the resulting progenies a loss of observed similar seed abortion rates after
vigour and a wide range of variation for self- and open pollination of 71 cultivated
qualitative and quantitative traits. In the sec- and wild clones, which indicates no depres-
ond generation, inbreeding depression was sion effect at the zygotic or embryonic level.
so severe that only a few plants produced No difference was observed in germination
fruits and none of them set seed. Backcrosses of self- and cross-seeds or in the early devel-
of first-generation inbreds to ‘Smooth opment of the resulting seedlings from
Cayenne’ restored vigour. Collins (1960) sug- ‘Española Roja’, ‘Primavera’, two clones of
gested the possibility of using this method to A. comosus var. ananassoides and two clones
synthesize new cultivars very similar to of A. comosus var. bracteatus. When trans-
‘Smooth Cayenne’. Following a similar idea, planted to the field, these seedlings showed
Coppens d’Eeckenbrugge and Duval (1995) about 50% size reduction compared with
proposed the introduction of one cycle of progenies from open pollination. In contrast,
selfing before the final hybridization to pro- a strong growth depression was observed in
duce inbred genotypes from the parental cul- the early development of seedlings of self-
tivars. Crossing genotypes with 50% progenies from ‘Manzana’ and from a clone
homozygosity would considerably lower the of A. comosus var. ananassoides, causing their
variability of the hybrid progeny. In addi- death in the nursery.
tion, studies on segregation of selfed proge-
nies would provide specific information on
Progeny evaluation
the genetic nature of the parental cultivars,
allowing identification and selection of reces-
Seed handling, germination and nursery
sive favourable alleles and elimination of
management
codominant and recessive unfavourable alle-
les. The final hybridization between the best Seeds can be harvested when fruits are ripe.
inbreds from two different cultivars would Seeds develop in the ovarian cavities (eyes)
restore vigour. and the cavities can be exposed by cutting
50 Y.K. Chan et al.

the peel of the fruit to a depth of about 1.5 Field trials

cm. The exposed seeds in the cavities are
In hybrid or inbred populations, each indi-
scraped out and washed. Alternatively, seeds
vidual plant is unique and the segregation is
can be extracted from the pulp using a low-
very wide, but less so for inbred progenies.
speed blender, preferably with a plastic
Evaluation is carried out on each plant and
blade. Most non-viable seeds are empty and
this may be very tedious considering the
remain afloat and these can be eliminated
large populations that are normally grown.
with the rinse water. Clean seeds are disin-
For this reason, only minimal data, covering
fected with 2–3% sodium hypochlorite and a
mostly qualitative characters that can be
wetting agent, dried and treated with a suit-
able fungicide before storage. The use of a visually recorded, are given emphasis. At
fungicide is critical. Storage of seeds with sil- this level of evaluation, basic data on fruit
ica gel in sealed plastic bags in a refrigerator shape, fruit weight, flesh colour and proba-
at 4–5°C can maintain viability for 2 years bly TSS, quickly recorded with a hand refrac-
(Leal and Coppens d’Eeckenbrugge, 1996). tometer, are collected. Plants with obvious
Seed pretreatment with concentrated sul- defects, such as fasciated fruits, multiple
phuric acid for 30 s improves uniformity in crowns, collar of slips, conical fruits and long
germination (Collins, 1960) but is not essential. peduncles, likely to lead to fruit lodging, are
In the tropics, good germination can be eliminated. The individual selected plants
obtained by sowing seeds evenly in shallow are usually propagated to some extent,
trays filled with fine river-sand. The trays are depending on the resources available. One
covered with clear polyethylene sheets to approach is to use the leaf-bud technique to
retain high humidity (Chan, 1986). In coun- propagate from the crown, suckers, slips and
tries with cooler temperatures, germination in stem, with each of the selections kept as a
temperature-controlled glasshouses or in separate family. It should be possible to
heated mist-beds inside a shade-house may be obtain up to 100 plants if all materials are
required (Sanewski, 1998). A range of different utilized. A less intensive approach is to
temperatures, from 24 to 35°C, have been used replant the crown, slips, suckers and stem
to germinate seed (Iyer et al., 1978). Seeds can sections only.
also be sown on sterile agar medium in cul- In the second round of selection, the
ture rooms (Loison-Cabot, 1987; Benega et al., selected families (clones) are further evalu-
1997). Germination occurs normally after 2 ated in replicated trials and more agronomic
weeks. The germination percentage of healthy characters, such as ascorbic acid, fruit titrat-
bold seeds is generally 80% or more. able acid, incidence of diseases and flavour,
When the seedlings reach the sixth- to are recorded. In the Hawaiian programme,
eighth-leaf stage after 3 months, they are screening techniques for the most important
either potted or carefully transplanted to diseases were applied by the third asexual
peat beds in the nursery. During transplant- generation. Subsequent selections based on
ing, some preliminary selection may be car- clonal performance and not individual plant
ried out to cull the obviously undesirable performance are made. Obviously poor-
characters, such as poor vigour, albinism and performing clones are discarded and, depend-
spiny leaves. The seedlings are kept in the ing on management capabilities, up to ten
nursery for 6–12 months before they are clones will make the next important selection
transplanted in the field for evaluation. The round, which is the genotype × environment
simplest approach is to plant in blocks coded trial. At this stage of testing, more materials
by the respective parent combinations. are available from propagation of leaf buds
About 2% of the population are planted with obtained from plants in the selected clones.
check plants, usually the parents, inter- Tissue culture can also be used to great
spersed randomly within the block advantage at this stage. This penultimate
(Sanewski, 1998). More complex designs can stage of testing is carried out over a few
be used where parent combinations are diverse environments and the selected fami-
being compared. lies are tested with contemporary commer-
 Breeding and Variety Improvement 51

cial varieties. The results of these trials will produce less variable but more useful hybrid
indicate the performance and stability of the populations. Sanewski (1998) reported that
selections compared with standard cultivars the PRI hybrids, such as ‘73–50’, have high
(Chan, 1997). breeding values and make very good parents
In Malaysia, the hybridization pro- because they have accumulated desirable
gramme took about 12 years before release of genes in their previous selection. It is envis-
the new variety ‘Josapine’. Started in 1984 aged that progress in hybridization should
with the diallele cross of four cultivars, the increase dramatically in the future as breed-
progeny evaluation and selection went ing programmes move beyond the basic-
through four cycles before the potential type crosses and into interhybrid
selection was identified in 1993 (Chan, 1993). combinations. However, it can be concluded
It took another 3 years to build up sufficient that hybridization, with its attendant inva-
planting materials before the variety was sive arrangement in gene constitution of an
released in 1996 (Chan and Lee, 1996). established variety, is a long and tedious
Loison-Cabot (1987) estimated that it would method, whose success depends, to a large
require 15 years to develop hybrid varieties extent, on the genetic lottery of a favourable
for growers, the major part of the time being recombinant.
spent on evaluation trials and in vitro propa- Improvement of pineapple varieties
gation of planting materials. Collins (1960) should also involve techniques that are less
estimated that 20–25 years would be invasive in genetic changes to the established
required for the development of commer- cultivar. Spontaneous mutations (or ‘spots’)
cially acceptable hybrids involving wild in the field or deliberately caused somaclonal
germplasm because of the need to incorpo- variants during tissue culture are some ways
rate about four additional back-crosses. in which small, favourable changes may be
made to a standard cultivar. The prospects of
genetically engineering target modifications
Prospects to the host variety appear very promising.
This area of research is already in progress
A historical review of pineapple-breeding for developing resistance to black-heart dis-
research showed that many breeders are order and mealybug wilt, resolving preco-
quite content with working around the cious flowering and delaying ripening of
benchmark ‘Smooth Cayenne’ cultivar, fruit for extension of storage life. It would
focusing on improving specific weakness, also be useful to obtain herbicide resistance
such as disease and pest resistance, improv- in pineapple for ease of weed management.
ing winter-fruit quality, peel colour and Genetic engineering must be supported by
resistance to black-heart disorder. However, adequate gene mapping of the pineapple
in the process of introducing the desired for- chromosomes. Present understanding in this
eign gene from another variety through area is still inadequate in locating, isolating
hybridization, many other undesirable and transferring genes in pineapple. Even
attributes are introduced as well. This is par- with successful gene transfer, the task of field
ticularly true when the source of the resis- testing under biosafety regulations for geneti-
tance gene is a wild relative with poor cally modified organisms will take many
agronomic traits. Good selections in the seg- years to complete. It can be expected that
regating hybrid population in this case are there will still be non-governmental organi-
expected to be extremely rare and this zation (NGO) groups and consumers in the
explains the lack of success in hybridization European Union (EU) resistant to accepting
programmes. Better progenies are often genetically modified organisms. All these dif-
obtained in back-crosses or between crosses ficulties aside, genetic engineering is
involving inbreds or selected hybrids, as expected to be the tool to fine-tune the per-
these procedures add on favourable genes to formance of current pineapple cultivars.
52 Y.K. Chan et al.

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 4 Biotechnology

M.K. Smith,1 H.-L. Ko,1 S.D. Hamill,1 G.M. Sanewski1 and

M.W. Graham2
1Maroochy Research Station, PO Box 5083, SCMC, Nambour, Qld 4560, Australia;
2Queensland Agricultural Biotechnology Centre, Gehrmann Laboratories, Level 4, The
University of Queensland, St Lucia, Qld 4072, Australia

Plant biotechnology is a new and rapidly include: micropropagation via axillary-bud

developing discipline that offers substantial proliferation, adventitious-bud proliferation
opportunities in the field of plant identifica- and regeneration from callus cultures, in
tion, multiplication and genetic improve- vitro germplasm conservation, and culture of
ment. The new tools of biotechnology have excised protoplasts, ovules or anthers.
made it possible to excise precise lengths of
DNA, to isolate and recombine them with
other pieces of DNA, to move them around Micropropagation
in DNA plasmids or vectors and to transfer
the vectors into the same or other species in Most tissue culture research in pineapples
ways that permit the desired DNA to relates to the use of micropropagation for the
become part of the new organism. rapid multiplication of cultivars. This has
Plant tissue culture is an integral part of been stimulated by the relatively slow proce-
plant biotechnology and includes the culture dures currently used for clonal propagation
of plant cells, tissues or organs under aseptic and the large and regular demand for plant-
conditions. This term can also be extended to ing material. Traditional methods of pine-
include the culture of excised embryos and apple propagation usually produce up to ten
protoplasts. Transformation develops the plants using crowns, slips and suckers from
tissue culture process further by delivering a single plant in a year. Sectioning of these
recombinant DNA to plant cells and the components, including the stem, can deliver
regeneration of plants from these cells. The up to 100 plants. The chemical chlorfurenol
goal of transformation is to ‘engineer’ plants can be used in commercial situations to
that express the gene of interest without enhance slip production as much as 30-fold.
altering the remaining, and often unique, But, for sheer scale of multiplication, none
part of the genotype. compares with tissue culture, where, accord-
ing to Pannetier and Lanaud (1976), 1 million
plants could theoretically be achieved in 2
Tissue Culture years from a single axillary bud.
The first report of pineapple in vitro prop-
Smith and Drew (1990) have reviewed many agation comes from the work of Aghion and
of the applications of tissue culture for plant Beauchesne (1960), and many of the earlier
propagation and improvement. Applications studies (Mapes, 1973; Mathews et al., 1976;
for pineapple propagation and improvement Drew, 1980) concentrated on culture estab-
© CAB International 2003. The Pineapple: Botany, Production and Uses
(eds D.P. Bartholomew, R.E. Paull and K.G. Rohrbach) 57
58 M.K. Smith et al.

lishment and the ability to take plants suc- along the nodal segments when placed hori-
cessfully through the micropropagation zontally on the culture medium.
process. Subsequent studies have concen- Once shoots have been produced and
trated on methods to optimize multiplica- multiplied, they are usually transferred to an
tion, and today pineapple micropropagation MS solid medium containing an auxin, such
is being used commercially in the pineapple as indole butyric acid (IBA), at between 0.5
industry to rapidly multiply important culti- and 2.0 mg l−1, or to a hormone-free medium
vars (Smith and Drew, 1990). In practice, for root development. Plants can be success-
micropropagation is used for the establish- fully established in a soilless potting mix and
ment of multiplication blocks, which then ‘hardened off ’ in the glasshouse prior to field
provide conventional planting material for establishment. The use of Azobacter or
larger production blocks. This is because endomycorrhizal fungi has also been sug-
micropropagated plants are more expensive gested as a way of improving the growth of
and there are grower concerns about genetic micropropagated plants (Gonzales et al.,
off-types (somaclonal variants). The initial 1996; Guillemin et al., 1996; Matos et al.,
limited use of micropropagation in the mul- 1996).
tiplication of new cultivars allows screening Plants can also be regenerated from cell or
of variants on fruit characteristics before con- callus cultures. When the cytokinin is sup-
ventional methods of multiplication are plemented with an auxin, such as naphtha-
used. lene acetic acid (NAA), the development of
Table 4.1 summarizes some of the meth- ‘lumpy’ tissue with protocorm-like bodies,
ods developed for pineapple micropropaga- or callus tissue can result (Mathews and
tion. The most common explant used to Rangan, 1979, 1981; Wakasa, 1989; Devi et al.,
initiate cultures is the axillary bud dissected 1997). Proliferation of these types of tissue
from crown leaves. Fitchet (1990) also sug- often results in cultures with an enhanced
gests that the crowns of some cultivars, such regeneration capacity via either organogene-
as ‘Smooth Cayenne’, should first be desic- sis (Fitchet, 1990) or embryogenesis
cated for a short period to break bud dor- (Daquinta et al., 1996; Cisneros et al., 1998).
mancy. Even though contamination of the Regeneration of plants from cell and callus
initial cultures is common, owing to the cultures opens up possibilities for the genetic
closely packed whorl of leaves in the crown transformation of pineapples. However,
which traps water and airborne particles, these procedures must be viewed very cau-
enough buds can be obtained to ensure suc- tiously because of perceived problems with
cessful establishment of some explants fol- somaclonal variation. As Scowcroft (1984)
lowing surface disinfestation. ‘Clean’ buds pointed out, as regeneration proceeds from
are most commonly grown on a Murashige more organized structures (axillary buds) to
and Skoog (1962) (MS) solid medium supple- unorganized tissues (callus), the propensity
mented with a cytokinin, usually benzylade- of the cells to undergo genetic changes
nine (BA), at between 2 and 5 mg l−1. Using increases.
these concentrations with a cultivar such as
‘Smooth Cayenne’, ten to 15 plants per month
can be produced. Multiplication may be Somaclonal variation
enhanced two- to threefold by the use of agi-
tated liquid media (Mathews and Rangan, Even though pineapple is not generally con-
1979; DeWald et al., 1988; Moore et al., 1992), sidered to have an unstable genotype, at
and further refinements have been made by least 30 mutants have been recorded in
the use of temporary immersion systems ‘Smooth Cayenne’ since the early 1920s
(Firoozabady et al., 1995; Escalona et al., (Collins and Kerns, 1938). Spiny leaves are
1998). A novel micropropagation method commonly encountered during conventional
was developed by Kiss et al. (1995); this pineapple propagation of ‘Smooth Cayenne’,
involved the induction of etiolated shoots, and it is no surprise that spininess is also
with subsequent multiplication of shoots most commonly encountered during micro-
Table 4.1. Review of methods used for the micropropagation of pineapple.

Stage I Stage II Stage III Stage IV Reference

Explant Axillary and terminal buds Medium MS with 2.0 mg l−1 KIN, Medium MS with 0.1 mg l−1 NAA ‘Soil’ Mathews et al. (1976)
from crown 2.0 mg l−1 IBA and and 0.4 mg l−1 IBA
Medium Nitsch with 0.1 mg l−1 BA 1.8 mg l−1 NAA
and 0.1 mg l−1 NAA 8 plantlets per month

Explant Axillary buds from slips and Medium MS with 2.3 mg l−1 BA Not stated ‘Gro-pots’ Drew (1980)
suckers and 2.2 mg l−1 KIN
Medium MS, hormone-free 50 plantlets per month
(callus involved)

Explant Axillary bud from crown Medium MS with between 0.5 Medium  MS, hormone-free Not stated Zepeda and Sagawa
Medium MS with 25% coconut water and 1.0 mg l−1 BA (1981)
3 plantlets per month
Biotechnology

Explant Axillary bud from crown Medium As for stage I, but liquid No roots ‘Commercial DeWald et al. (1988)
Medium MS with 2 mg l−1BA and (shaken) cultures soil mix’ Moore et al. (1992)
2 mg l−1 NAA 17 plantlets per month Plantlets  2.5 cm
for ‘Smooth Cayenne’;
76 plantlets per month
for ‘Perolera’

Explant Axillary bud from crown Medium MS with 0.5 mg l−1 BA Medium MS, hormone-free ‘Peat/perlite mix’ Cote et al. (1991)
Medium MS with 0.5 mg l−1 BA and ‘Plantlets halved or Plantlets 2–3 cm
0.2 mg l−1 IAA quartered during
subculture’
10 plantlets per month

Continued
59
 60

Table 4.1. Continued

Stage I Stage II Stage III Stage IV Reference

Explant Axillary bud from crown Medium MT with 2.0 mg l−1 KIN Medium MT with 1.0 mg l−1 NAA ’Peat/perlite/sand’ Fitchet (1990)
Medium MT with 2.0 mg l−1 KIN, and NAA and 500 mg l−1 malt Fitchet-Purnell (1993)
IBA and NAA 14 plantlets per month extract

Explant Axillary bud from stem Medium As for stage I Medium MS with 0.3 mg l−1 IBA Not stated Osei-Kofi and Adachi
Medium MS with 2.3 mg l−1 BA 8 plantlets per month and 0.1 mg l−1 IAA (1993)
and 0.6 mg l−1 NAA

Explant In vitro plantlets Medium N6 with 5 mg l−1 KIN Medium MS, hormone-free ‘Soil’ Kiss et al. (1995)
Medium MS with 2.0 mg l−1 NAA in and 4.5 mg l−1 BA Plantlets 8 cm
the dark ‘Etiolated shoots placed
horizontally on medium’
M.K. Smith et al.

60 plantlets per month

Explant Apical bud from crown Medium As for stage I, except Medium MS with 2.0 mg l−1 IBA Not stated Devi et al. (1997)
Medium MS with 1.0 mg l−1 BA and 2.0 mg l−1 BA
0.1 mg l−1 NAA ‘Regeneration from pro-
liferating callus at base
of plantlets’
56 plantlets per month

BA, benzyladenine; IAA, indole acetic acid; IBA, indole butyric acid; KIN, kinetin; NAA, naphthalene acetic acid; MS, Murashige and Skoog (1962); MT,
Murashige and Tucker (1969); N6, Chu (1978); Nitsch, Nitsch (1951).
 Biotechnology 61

propagation (Wakasa, 1979, 1989; Smith and they also offer the convenience of medium-
Drew, 1990). Spiny variants have also been and long-term storage of germplasm and
observed in micropropagated ‘Red Spanish’ facilitate its safe distribution.
(Liu et al., 1987). A range of other somaclonal Lower temperatures (16–20°C, as opposed
variants have been described by Wakasa to 25–28°C for normal micropropagation)
(1989), including variants for leaf colour, leaf have been used successfully to extend sub-
shape, wax, foliage density and abnormal culture times for up to 4 years (Sugimoto et
phyllotaxy. Spininess and dense-foliage vari- al., 1991). Zee and Munekata (1992) found
ants were attributed to chimeras at the that reducing the nutrient salts of the MS
donor-plant level. However, most of the medium to one-fourth was successful for
other variations were attributed to regenera- medium-term (12 months), low-input main-
tion of plants from callus cultures, especially tenance of pineapple cultures. For the long
undifferentiated callus derived from the syn- term, cryopreservation offers the best solu-
carp. Further studies of this nature are tion, as the processes leading to somaclonal
needed to establish the explant sources and variation during cell division are halted.
culture conditions required to minimize Gonzalez-Arnao et al. (1998) obtained the
somaclonal variation during micropropaga- first positive results in pineapple. Optimal
tion and transformation. conditions included a 2-day preculture of
Smith (1988) and Damasco et al. (1998) apices on a semisolid MS medium with 0.3 M
outlined a number of strategies for minimiz- sucrose, loading treatment for 25 min with a
ing somaclonal variation in micropropagated cryoprotective solution containing glycerol
bananas, and central to these approaches (1M) and sucrose (0.75M) and dehydration
were regular initiations of cultures while with a vitrification solution (30% (w/v)
providing limits to the number of plants glycerol + 15% (w/v) DMSO + 0.4 M
multiplied from a single explant. While no sucrose) at 0°C for 7 h before immersion in
data exist on this aspect in pineapple, com- liquid nitrogen.
mercial tissue-culture laboratories usually
limit multiplication to 300–1000 plants per
explant. Roguing of off-types, particularly Anther and Ovule Culture
during nursery establishment, is also essen-
tial to reduce the percentage of off-types There is some interest in the development of
reaching the grower. Many of the somaclonal haploid cultivars as a first step to circumvent
variants recorded by Wakasa (1989) would the heterozygosity observed in segregating
lend themselves to easy identification and seedling populations. There is doubt
roguing while the plants are quite small. whether the development of doubled hap-
Other less obvious variants, such as those loids is likely to be a useful strategy, and
that produce small, malformed or cracked other approaches, such as those involving
fruit, require a fruiting cycle for identifica- some level of inbreeding, may be better. To
tion. It is particularly important to grow all date, limited success has been achieved with
plants arising from tissue culture through a ovule culture, but anther culture appears
cycle of fruiting to enable roguing of all more difficult (Benega et al., 1996).
somaclonal variants.

Protoplast isolation and fusion

In vitro germplasm conservation
Like anther and ovule culture, there is some
Advancements in breeding and genetic engi- doubt whether the development of tech-
neering require that the genetic diversity of niques for protoplast isolation and fusion are
pineapples and their close relatives not only likely to be worthwhile in pineapple.
be preserved but be put to use for pineapple Protoplasts have been used in some monocot
improvement. While tissue culture techniques species for transformation, but there are
offer the opportunity for rapid propagation, inherent problems associated with the use of
62 M.K. Smith et al.

protoplasts and other approaches appear pineapple. The maize ubiquitin (Ubi-1) and
more applicable in pineapple. Protoplast the cauliflower mosaic virus 35S (CaMV35S)
fusion is sometimes used where parent promoters have been shown to function well
incompatibility prevents hybridization by in transient assays in pineapple fruit, leaves
conventional means. This approach, how- and callus (Graham et al., 1998). The transfor-
ever, will result in the development of mation technique developed by DNA Plant
tetraploid lines. Tetraploid pineapple culti- Technology Inc. (DNAP) in the USA also uti-
vars studied to date have been inferior types lizes these two most commonly used pro-
(Collins and Kerns, 1947). Nevertheless, pro- moters (Firoozabady and Gutterson, 1998).
toplasts of the cultivar ‘Perolera’ have been However, confirmation of the genuine utility
successfully isolated (Guedes et al., 1996), but of these promoters in transgenic pineapple
plant regeneration was not achieved. will await the generation and analysis of
many stably transformed lines. Similarly, for
driving expression of the nptII selectable
Genetic Engineering marker gene, both the Ubi-1 and CaMV35S
promoters, and promoters derived from seg-
The pineapple industry around the world is ments of the subclover stunt circovirus, have
unique in its almost total reliance on a single been used successfully to create stably trans-
cultivar. This industry, through its consistent formed plants (Graham et al., 1998).
use of a single cultivar from the birth of
pineapple processing, has virtually fixed the
identity of canned pineapple as ‘Smooth Transformation
Cayenne’. New cultivars for this market
would need to be almost identical to ‘Smooth Transformation techniques, using methods
Cayenne’ in production, processing and such as infection with Agrobacterium, electro-
organoleptic qualities. Improvements to this poration or biolistics, enable us to introduce
cultivar would therefore be difficult using ‘engineered’ genes into plant cells. The
conventional hybridization techniques. Agrobacterium co-cultivation technique is
Genetic engineering, however, would have based on the infective activity of the bacteria
tremendous application for making small tar- to introduce the recombinant DNA into the
geted changes without changing the integrity plant cells and thus act as a natural vector.
of the cultivar we know as ‘Smooth Cayenne’. Agrobacterium co-cultivation was convention-
To successfully genetically engineer ally the transformation method used for
pineapple, genes of interest first have to be dicots. Monocots were found not to be sus-
identified and sequenced and a complemen- ceptible to Agrobacterium infection in early
tary DNA (cDNA) clone generated. A plas- studies, but more recent studies have indi-
mid construct, including promoter, cated that many are and that Agrobacterium
terminator regions, gene of interest and a can be successfully employed for pineapple
selectable marker gene, can then be designed transformation. Other transformation meth-
and constructed. Transformation techniques ods employed for monocots include the
are then employed to introduce the new gene introduction of novel genes into plant proto-
sequence into cells of the chosen cultivar. plasts (Hauptmann et al., 1987; Rathus and
Birch, 1992). However, protoplasts are easily
damaged and cultures do not always grow
Promoters into morphogenic callus, to produce trans-
genic plant material. Another more success-
Promoters are segments of DNA that drive ful transformation technique is biolistics, in
the expression of the new gene sequence which recombinant DNA coated on to gold
(transgene) and they may or may not be or tungsten particles is propelled under gas
tissue-specific. Early work indicates that pro- pressure in a vacuum chamber into the plant
moter elements and constructs developed for cells. However, with this method multiple
use in other species can be adapted for use in copies of the DNA can become incorporated
 Biotechnology 63

and there is less control over the precise loca- genetic engineering include nematode resis-
tion of the inserted DNA in the plant genome. tance, pineapple mealy-bug wilt-virus resis-
The transformation techniques that have tance, flowering and fruit-ripening control
achieved the most success to date are the and black heart resistance (Botella et al., 1998;
direct introduction of transgenes by micropro- Graham et al., 1998; Rohrbach et al., 1998;
jectile bombardment (biolistics) and the indi- Sanewski et al., 1999).
rect method by co-cultivation using an
Agrobacterium vector. Both methods have been
applied to pineapple; however, commercial- In vitro selection procedures
ization and patenting issues have in the past
prevented the results of the work from being In vitro selection of transgenic material is
published widely. Nan et al. (1996) reported achieved by cointroducing a resistance gene
the use of the biolistics technique on liquid with the gene of interest. The most com-
embryogenic protocorm cultures, obtaining monly used selective genes are those confer-
low levels of transgenic material. Graham et ring resistance to an antibiotic (Dekeyser et
al. (1998) have successfully employed the al., 1989; Caplan et al., 1992) or herbicide
biolistics technique to callus cultures, achiev- (Vasil et al., 1992; Cole, 1994; Dennehey et al.,
ing an average of 1% transformation effi- 1994). The transgenic material can be
ciency. On the other hand, Firoozabady and screened in vitro by survival on a culture
Gutterson (1998) have produced transgenic medium containing the selective agent.
pineapples using the Agrobacterium-mediated Graham et al. (1998) found that geneticin at
technique on embryogenic callus cultures, 50 g ml−1 and hygromycin at 20 g ml−1
while Isidron et al. (1998) have reported GUS were useful selective agents, but not
positive activity using a similar system. kanamycin, even at concentrations as high as
Graham et al. (1998) have successfully 500 g ml−1.
employed Agrobacterium co-cultivation to The choice of selective gene can have sub-
‘Smooth Cayenne’ pineapple leaf bases, stantial consequences on the consumer accep-
resulting in the production of organogenic tance of the final product. Consideration
callus cultures and transgenic plants, with should be given to the likely concerns of the
transformation efficiency levels of about 2%. public when choosing selective genes.
All methods require a good and reliable
regeneration system (Rangan, 1982) to pro-
duce transgenic plants from transformed Evaluation of transgenics
cells. Using Wakasa’s (1989) recipe for
‘Smooth Cayenne’, Graham et al. (1998) initi- Incorporation of the introduced gene in
ated callus on 90% of excised leaf bases from putative transgenic material can be deter-
well-developed micropropagated shoots mined in vitro by polymerase chain reaction
within 3–4 weeks. Callus was maintained on (PCR) analysis, in which specifically
the same medium for proliferation, produc- designed primers can detect the presence or
ing different types of callus, very similar to absence of the novel gene in the plant tissue.
those described by Taylor et al. (1992) for This early evidence can identify ‘escapes’ or
sugar cane. The majority of callus consisted non-cotransformed individuals when coin-
of compact, hard, globular, yellow-green, troduction of the novel gene and the selec-
smooth-surfaced structures. This type of cal- tive gene was employed.
lus was regenerated through organogenesis. To confirm insertion of the novel and
To a much lesser extent the second (yellow selectable marker genes in the transgenic
and friable) and third type (soft, mucilagi- lines, the PCR needs to be followed up by
nous, grey-yellow) of callus was also pro- Southern blot analysis, which detects the
duced. Only the first type of callus is actual presence of the introduced DNA frag-
believed to be morphogenic. ment after DNA extraction from the tissue.
Traits that have been targeted for This technique will also give an indication of
improvement in ‘Smooth Cayenne’ using copy numbers inserted in the genome.
64 M.K. Smith et al.

Intellectual property and licensing laws will then require identification and seg-
regation of fields of genetically modified
Intellectual property (IP) is another impor- plants. It is important for the pineapple
tant issue in the application of genetic engi- industry to embrace protocols at this early
neering for pineapple improvement. Many stage to enable identification and segregation
of the enabling technologies are patented of genetically modified plants and thus
and cannot be used commercially without ensure consumer confidence.
licence agreements. The Agrobacterium co-
cultivation technique for pineapples has
been patented, although patents tend to be Molecular Markers
written quite specifically to cover particular
sections of the protocol and are country- Genetic classification in the genus Ananas
specific with an expiry date. The promoters was originally based on quantitative, mor-
Ubi-1 and 35S are also patented. IP issues phological (Smith and Downs, 1979) and
may even impede the design of a certain qualitative variables. Such classifications
plasmid construct, e.g. patents on using were initially re-evaluated using enzymatic
an antisense or co-suppression construct markers (isozyme analysis). In more recent
design. years, several DNA-based marker tech-
It is generally prudent to be aware of all niques, such as random amplified polymor-
IP before commencing work that may ulti- phic DNA (RAPD) and restriction fragment
mately infringe the rights of others when length polymorphism (RFLP) analysis, have
commercialized. The issues associated with been used. There are no reports of these tech-
this are very complex, but must be resolved niques being used to directly assist
before transgenic pineapples are grown com- hybridization strategies in pineapple.
mercially.

Enzymatic markers
Environmental and health risk assessment
Isozymes were used in the first systematic
Biosafety and public acceptance of trans- studies of the pineapple by Garcia (1989),
genic crops represent at least two other fac- with eight enzyme systems involving ten
tors that must be considered before loci, and by Aradhya et al. (1994), with six
commercialization. The former does not systems involving seven loci. They found a
seem to be a major issue with pineapple, pri- high heterozygosity and polymorphism in A.
marily because there is little chance of gene comosus, with most variation between the
escape into the environment. botanical varieties. In some cases, clones
Public acceptance is currently a major from the same cultivar showed differences in
issue with all transgenic crops. The per- some systems. DeWald et al. (1992) tested
ceived safety of transgenic foods and the eight enzymatic systems and found variable,
ownership of genetic material are at the cen- well-resolved banding patterns for peroxi-
tre of these concerns. It is difficult to antici- dase (PER) and phosphoglucomutase
pate changes in acceptance, but advances in (PGM). Their study, as well as that of
transformation techniques, such as the Laempet and Saghuanrungsirikul (1998),
development of selectable markers to replace also showed incomplete correspondence
antibiotic resistance markers and the use of between cultivars and zymotypes, and hence
the technology for more consumer-orien- some unreliability for cultivar identification.
tated product improvements might improve Pérez et al. (1995) applied the isozyme analy-
acceptance. Irrespective of changes in accep- sis technique to detect mutant lines devel-
tance, it is evident that food-labelling laws in oped by cobalt-60 (60Co) gamma irradiation
many countries will demand the identifica- of callus, while Arias Valdes et al. (1998) used
tion of genetically modified contents. These the technique to evaluate ploidy levels.
 Biotechnology 65

DNA markers Conclusion

DNA-based markers are more sensitive than Pineapple tissue culture will continue to play a
enzymatic markers and are widely used for valuable role in clonal propagation schemes.
cultivar identification, for gene isolation and, Genetic engineering will also become an estab-
perhaps most significantly, to monitor the lished method for pineapple improvement,
segregation of chromosome regions known particularly in cultivars, such as ‘Smooth
to be involved in controlling agronomic Cayenne’, that do not lend themselves readily
characters (quantitative trait loci (QTLs)). In to improvement by conventional breeding
pineapple this technique has been used strategies. The main obstacle to their wide-
mainly to classify cultivars. spread adoption will be somaclonal variation.
Ruas et al. (1995) used RAPD analysis to Reliable methods will be required for the con-
estimate the relationships among four vari- trol and detection of deleterious off-types pro-
eties. Polymorphism was observed in agree- duced during in vitro culture, and certainly
ment with classification based on before plants are grown in the field.
morphological characters. Noyer et al. (1996) Molecular markers have become powerful
used RFLP analysis on ribosomal RNA and tools in pineapple taxonomy and will assist
found the genus Ananas to be very homoge- with more precise identification and classifi-
neous. The results suggested variation cation. They will also become extremely use-
within A. ananassoides that could constitute ful in hybridization programmes for the
the origin of A. comosus. The most compre- selection of parent combinations containing
hensive study has been by Duval et al. (1998). the most important traits, thus reducing the
Their results were consistent with the classi- number of generations required.
fication of pineapples in one genus and two At the annual meeting of the
species (see Coppens d’Eeckenbrugge and Biotechnology Industry Organization in June
Leal, Chapter 2, this volume). This study 1998, Charles Shapiro, the Chairman of
indicated that variability was generally con- Monsanto Agricultural Company, compared
tinuous between current species, and A. the stage of development of the biotechnol-
comosus and A. ananassoides appear to be the ogy industry today to that of the information
most diverse groups. technology industry in 1973. Although com-
In the long term, spectacular advances in puters had been known since the 1940s, most
mapping technologies and the study of of the rapid advances that have made the
whole genomes can be confidently predicted. information technology industry what it is
A major European Union (EU)-funded pro- today have taken place in the past 25 years.
ject in this field aims at characterization and If that is truly where the biotechnology
gene mapping for linking markers to mor- industry is today, then today may be just the
phological traits and disease resistance beginning and the next 25 years could be the
(Coppens d’Eeckenbrugge et al., 1998). real biotechnology revolution.

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 5 Crop Environment, Plant Growth and
Physiology

Eric Malézieux,1 Francois Côte1 and Duane P. Bartholomew2

1Centre
de Cooperation Internationale en Recherche Agronomique pour le
Developpement (CIRAD), TA 179/01 Avenue Agropolis, 34398 Montpellier Cedex 5,
France; 2Department of Tropical Plant and Soil Science, University of Hawaii at
Manoa, 3190 Maile Way, Honolulu, HI 96822, USA

Introduction ductivity is reduced in drought conditions.

This is partly due to the fact that pineapple,
Pineapple, a crop native to the tropical as with most Bromeliaceae, has the crassu-
Americas, is currently grown commercially lacean acid metabolism (CAM) photosyn-
over a wide range of latitudes from approxi- thetic pathway. This specificity of pineapples
mately 30°N in the northern hemisphere is unique among plants widely cultivated for
(30°45’N in India (Hayes, 1960) and 28°30’ in commercial purposes.
the Canary Islands (Galan Sauco et al., Pineapple-based cropping systems
1988b)) to 33°58’S in South Africa throughout the world vary from gathering
(Bartholomew and Kadzimin, 1977; Table fruit from wild plants under tree cover, as in
5.1). Pineapple seldom requires less than 12 some areas of Brazil, and intercropping sys-
months from planting to harvest and more tems that include pineapple and a wide
commonly 18–24 months, or even as much as variety of tree and herbaceous crops (Lee,
36 months in cool subtropical environments. 1972; Nair, 1977; Bavappa et al., 1986;
The induction of flowering of pineapple is Darwis, 1990; Oladokun, 1990; Cai and
almost universally initiated with a growth Zheng, 1991) to highly intensive monocul-
regulator (see Hepton, Chapter 6, and ture cropping systems on thousands of
Bartholomew et al., Chapter 8, this volume), hectares, as is the case in Hawaii, Indonesia,
which has made large-scale production of the Philippines and Thailand. As a result,
pineapple possible in areas where flowering yields are extremely variable and can range
would otherwise be too erratic for commer- from 30 to more than 100 Mt ha−1, depend-
cial production. Further, in many areas ing on the length of the crop cycle, environ-
where the crop is grown, fruiting may be mental conditions, cultural practices and the
forced in most or all months of the year, so it pineapple variety being grown (Py et al.,
is common for both vegetative and repro- 1987). High productivity of pineapple
ductive growth to occur during both warm depends on successfully managing control-
and cool seasons. lable factors, such as water and nutrient
Production at the more extreme latitudes supply and pests and diseases (see Hepton,
is mainly restricted to areas where the cli- Chapter 6, and Rohrbach and Johnson,
mate is moderated by the ocean, as the crop Chapter 9, this volume). Environmental fac-
will not tolerate freezing temperatures. One tors, such as temperature, irradiance and
of the main features of pineapple is its adap- rainfall, interacting with the plant’s CAM
tation to areas of low rainfall, although pro- and the unique morphological and anatomical
© CAB International 2003. The Pineapple: Botany, Production and Uses
(eds D.P. Bartholomew, R.E. Paull and K.G. Rohrbach) 69
 70

Table 5.1. Temperature and length of crop cycle at various pineapple-growing areas in the world.

Temperature (°C ) Months to plant crop

Elevation F to H
Location Latitude (m) Average Max. Min. Force (F) Harvest (H) (days) Reference

Nyombe, Cameroon 4.5°N 70 6 11 Gaillard, 1970

Mt Cameroun, Cameroon 0 26.2 30.2 22.3 10 150 Aubert et al., 1973
550 22.6 27.6 17.7 10 180
1000 19.9 25.0 14.9 10 210
Johore, Malaysia 1°22N 5 26.9 35.0 18.9 Wee, 1969
Pekan Nenas, Malaysia 10 15–17* Wee, 1969
Abidjan, Côte d’Ivoire 5°N 25 26.6 30.9 22.3 8–11* 13–15* 156–166* Lacoeuilhe, 1978
Bangkok, Thailand 32.2 23.3 Neild and Boshell, 1976
Ivoloina, Madagascar 18°03S 20 23.8 28.0 19.6 8–10* 144–221* Moreau and Moreuil, 1976
Buenos Aires, Costa Rica 9°10N 23.0 32.0 19.9 13 20 Romero et al., 1973
Cali, Colombia 31.1 18.1 16 Neild and Boshell, 1976
Fort de France, Martinique 14°30N 20 10 Lacoeuilhe and Gicquiaux, 1971
60 10
350 11
E. Malézieux et al.

Foulaya, Guinée 10°N 400 24.7 30.7 18.7 13 19 180 Py et al., 1957
Sape, Brazil 7°5S 124 25.9 31.2 20.6 Giacomelli and Py, 1982
Coracao de Maria, Brazil 12°14S 267 23.6 30.0 17.3
Serra, Brazil 21°S 76 24.1 27.6 20.6
Lagoa Santa, Brazil 19°38S 778 21.4 26.5 16.3
Monte Alegre, Brazil 18°52S 756 22.3 29.1 15.6
Osorio, Brazil 29°55S 38 19.6 23.8 15.4
Rio Pedras, Puerto Rico 18°23N 23 24.8 Pico, 1974
Rock Hampton, Australia 23°26S 11.3 22.7 27.2 16.7 Bartholomew and Kadzimin, 1977
Brisbane, Australia 27°28S 42 20.5 25.5 9.5 Tkatchenko, 1947
Nambour, Australia 26°38S 29–20.5† 19–6.0† 203–301* Wassman, 1990
Yeppoon, Australia 23°6S 30–23.0† 21–11.0†
Wahiawa, Hawaii 21°20N 200 22.6 30.1 14.3 14 23 Bartholomew and Kadzimin, 1977
Kunia, Hawaii 216 25–19.5† 9 175–223* Fleisch, 1988
Maui, Hawaii 91 23.5 27.9 19.1 12 181–207* Bartholomew and Malézieux, 1994
300 21.8 25.7 17.9 12 195–213*
792 19.5 24.3 14.7 12 230–255*
Touliu, Taiwan 23°44N 48 16 23 Su, 1969
Karenko, Taiwan 23°58N 19 22.2 27.1 17.3 Tkatchenko, 1947
Thika, Kenya 1°01S 1463 20.5  35 5.5 22–30* Lebedev, 1970
Kiamba, Kenya 1°12S 1731 19.0
Tenerife, Canary Islands 28°N 50 19.5 9–15 16–24* 210–270* Galan Sauco et al., 1988a, b
150 21.1
475 15.8
Zululand, South Africa 28°S 18‡ 210–240*‡ Lacoeuilhe and Sarah, 1990
East London, South Africa 33°02S 125 18.6 22.8 14.4 24–36* Strauss, 1960; Bouffin, 1991
Port Elizabeth, South Africa 33°58S 55 17.2 21.2 13.3 Tkatchenko, 1947
Malkerns, Swaziland 26°30S 16.8 28.0 4.0 22 32 Dodson, 1968

*Range for two or more plantings in different seasons.

†Range of the maximum and minimum mean monthly air temperature.
‡Clone of the Queen group.
Crop Environment, Plant Growth and Physiology
71
72 E. Malézieux et al.

features of pineapple ultimately determine Photosynthetic rhythm in CAM plants and in

productivity within a given environment. pineapple
Rhythmic patterns of net CO2 exchange are
well known in CAM plants, while photosyn-
Carbon Assimilation and Water Use
thetic O2 evolution is less well documented.
The diel rhythm of CO2 fixation and O2 evo-
Carbon fixation via the CAM photosynthetic
lution of an attached pineapple ‘D’ leaf (Fig.
pathway
5.1) displays the typical four-phase pattern
of net CO2 exchange in CAM plants
Three main types of photosynthetic path-
described by Osmond (1978).
way exist in higher plants. In the C3 type,
the most widely distributed, CO2 fixation • Phase I corresponds to nocturnal carboxy-
catalysed by the enzyme ribulosebisphos- lation of phosphoenolpyruvate (PEP) via
phate carboxylase/oxygenase (Rubisco) PEP-Case to form oxaloacetate (OAA).
results in the synthesis of a three-carbon OAA is reduced to malate and stored in
acid. In C4 plants – for crops, mainly the chlorenchyma cell vacuoles. The inter-
Graminaceae of tropical origin – CO2 is ini- nal CO2 partial pressure remains low dur-
tially fixed by the enzyme phospho- ing this time and stomatal conductance is
enolpyruvate carboxylase (PEP-Case) into a high. The substrate for PEP production in
four-carbon acid in leaf mesophyll cells. The pineapple is mainly soluble sugars
four-carbon acid is then transported to adja- (Borland and Griffiths, 1989; Carnal and
cent bundle-sheath cells where decarboxyla- Black, 1989). In terms of balance, internal
tion liberates CO2, which is fixed by CO2 from respiration is also fixed via
Rubisco. This decarboxylation is accompa- PEP-Case and contributes to the malate
nied by an increase in the intracellular CO2 pool. Night O2 uptake data (Fig. 5.1) indi-
concentration, which stimulates photosyn- cate that, in a ‘D’ leaf, CO2 produced by
thesis by inhibiting photorespiration and respiration represents approximately 10%
increases water-use efficiency (WUE) of the total CO2 fixed into malate at night
(Hatch, 1975; Wong et al., 1979). A third cate- (Côte, 1988). Night net CO2 fixation in
gory of plants, which includes pineapple, pineapple generally accounts for at least
also fix CO2 via PEP-Case and Rubisco. two-thirds of the total net fixation of the
However, in these plants, a temporal separa- day–night period (Neales et al., 1980;
tion occurs between the two carboxylation Bartholomew, 1982; Nose et al., 1986;
steps. Nocturnal fixation of CO2 occurs via Côte, 1988). Reported values for 21 stud-
PEP-Case, and the C4 acids synthesized, ies in controlled environments with vari-
mainly malic, are stored in the vacuole. The ous photo- and thermoperiods and light
following day, decarboxylation of malate levels ranged from 40 to 100%, with a
liberates CO2, which is fixed by Rubisco. mean of 80  17% (Bartholomew and
This type of photosynthesis, which results in Malézieux, 1994). The proportion of CO2
large daily variations in the concentration of fixed nocturnally by pineapple plants
malate and which is found in species of the propagated in vitro shifted progressively
Crassulaceae, is termed crassulacean acid from 0 to 70% as plant fresh weight
metabolism (CAM). The main feature of increased from 1 to 300 g (Côte et al.,
CAM plants is their high WUE. Malate 1993). These observations support the
decarboxylation during the day is associated idea that pineapple is an obligate CAM
with an increase in the internal CO2 concen- plant.
tration and a subsequent decrease in the • Phase II, at the beginning of the day, cor-
stomatal conductance, which reduces exter- responds to the transition from net CO2
nal CO2 uptake and transpiration (for fixation via PEP-Case to CO2 fixation via
reviews, see Kluge and Ting, 1978; Osmond, Rubisco. Malate decarboxylation begins
1978; Winter, 1985). Pineapple is the most during this phase, the internal CO2 partial
important crop species exhibiting CAM. pressure gradually increases and stomatal
 Crop Environment, Plant Growth and Physiology 73

18
Night Day
16
Gas exchange (mol m–2 s–1) 14 I II III IV
12
CO2
10
8 O2

6
4
2
0
–2
0 6 12 18 24
Time (h)

Fig. 5.1. Net CO2 and O2 exchange rates of an attached ‘Smooth Cayenne’ pineapple ‘D’ leaf throughout a
night/day cycle (Côte, 1988). Both CO2 fixation and O2 evolution are indicated as positive values. Positive
values for O2 evolution during the day result from photosynthesis, while negative values during the night
indicate O2 uptake due to respiration. The four phases of the CAM cycle as defined by Osmond (1978) are
indicated at the top of the figure. Environmental conditions include: photosynthetic photon flux density, 400
mol m−2 s−1; photoperiod, 12 h night/12 h day; night/day temperature 22°C/26°C. Data are the average for
4 consecutive days.

conductance declines. The overall contri- evolution during this phase, could be 1.3-
bution to carbon gain is small during to six-fold higher than the maximum rate
phase II and was 1–3% of the total of net CO2 fixation achieved by pineapple
day–night net CO2 uptake by pineapple during the light period (Côte, 1988). In
plants maintained in controlled condi- the data of Fig. 5.1, it is 2.4 times higher.
tions (Fig. 5.1). • Phase IV begins when the malate pool
• Phase III is a period when net CO2 fixa- becomes depleted, the internal CO2 par-
tion is negligible for up to 5 h, while CO2 tial pressure declines, stomatal conduc-
generated by malate decarboxylation is tance increases and net external CO2
fixed via Rubisco. The internal CO2 par- uptake resumes for the last hours of the
tial pressure rises well above atmospheric day. In pineapple, this fixation generally
levels, because the rate of malate decar- amounts to 15–25% of the total net CO2
boxylation exceeds the rate of internal uptake (Fig. 5.1). Early in the phase, there
CO2 fixation via Rubisco. Net CO2 release is a progressive shift from fixation of
sometimes occurs during phase III. This internally generated CO2 towards exter-
high internal CO2 partial pressure stimu- nal net CO2 uptake. Following the
lates photosynthesis and results in the assumption that 1.0 mol O2 is evolved per
low stomatal conductance, which limits mole of CO2 fixed, it was estimated that
transpiration during this phase. Although only 60–80% of the total CO2 fixed during
net CO2 uptake is nil during phase III, the the night by a pineapple plant was reas-
maximum rate of photosynthetic carbon similated via Rubisco when net CO2
reduction via Rubisco occurs during this uptake resumed in phase IV (Côte et al.,
phase. Based on the assumption that 1.0 1989). A value of 75% can be deduced
mol O2 is evolved for every mol CO2 from Fig. 5.1. PEP-Case has been reported
reduced (Kaplan and Björkman, 1980), the to be active during late phase IV in CAM
rate of CO2 fixation via Rubisco during plants (Kluge, 1969; Kluge et al., 1982) and
phase III, deduced from the rate of O2 measurement of malate concentration
74 E. Malézieux et al.

also suggested that CO2 uptake via PEP- Photosynthetic rate in pineapple
Case occurs in pineapple during late
In CAM plants, both net CO2 uptake and
phase IV (Black et al., 1982; Kenyon et al.,
malate-decarboxylation-dependent CO2
1985). The O2 and CO2 gas-exchange data
assimilation via Rubisco during phase III
suggest that CO2 fixation via PEP-Case
have to be considered to evaluate the photo-
during phase IV represents 3–15% of the
synthetic rate. Net CO2 fixation in the light
nocturnal net CO2 fixed by pineapple
(phase IV) ranged from 0.13 to 6.3 mol m−2
(Côte, 1988; Côte et al., 1989).
s−1 for pineapple plants or ‘D’ leaves main-
Gas exchange measured under controlled tained in various conditions (Connelly, 1972;
conditions (Fig. 5.2) is atypical of what Neales et al., 1980; Sale and Neales, 1980;
occurs in the field, where stochastic, rather Côte, 1988; Bartholomew and Malézieux,
than steady-state, environmental conditions 1994). Night net CO2-fixation values reported
prevail. Diurnal changes in weather proba- by the same authors ranged from 0.13 to 2.5
bly influence to some extent both the dura- mol m−2 s−1, while others reported values as
tion and the intensity of the four phases of high as 5.0 to 7.8 mol m−2 s−1 (Côte, 1988;
CAM. While extrapolation of results Borland and Griffiths, 1989; Fig 5.1). It can be
obtained in controlled conditions to plants in deduced from photosynthetic O2 evolution
natural environments must be done care- that CO2 reduction via Rubisco never
fully, in situ measurement of stomatal con- achieves a steady state in pineapple (Côte,
ductance and leaf titratable acidity confirm 1988; Côte et al., 1989; Fig. 5.1). Data for O2
the general conclusions derived from studies evolution from a pineapple ‘D’ leaf also
in controlled environments. show that the CO2 assimilation rate during

Fig. 5.2. Controlled-environment chambers used for gas-exchange measurements of ‘Smooth Cayenne’
pineapple plants from in vitro culture (Côte, 1988). Measurements were made in controlled atmosphere
automatic growth chambers (CEA, Commissariat à l’Energie Atomique, 13108 St Paul-lez-Durance, France).
 Crop Environment, Plant Growth and Physiology 75

phase III reaches 9.9 mol m−2 s−1, a value WUE is two times greater during phase I
much greater than the maximum value than during phase IV and reaches a mini-
observed in phase IV (Côte, 1988) (Fig. 5.1). mum during phase III (Fig. 5.3). The transpi-
The maximum CO2-fixation rates for ration rates of pineapple ranged from 0.05 to
pineapple are comparable to those for other 0.23 mmol m−2 s−1 (Ekern, 1965; Joshi et al.,
CAM plants, but are low compared with val- 1965; Neales et al., 1968; Nose et al., 1981),
ues of 8.33–25 mol m−2 s−1 reported for C3 while values for sunflower and tobacco were
plants (Black, 1973). The low rates for at least tenfold higher at about 3.1 mmol m−2
pineapple relative to those of C3 plants is in s−1 (Neales et al., 1968). Transpiration by
part due to low leaf conductances to gas dif- pineapple in the light in controlled condi-
fusion during phase IV (Côte, 1988), even tions was only 4% of that of sunflower leaves
when the stomata are wide open. The con- grown in a similar environment (Neales et
centration of Rubisco per unit area of leaf al., 1968) and only 6% of that for wheat, and
could also be low relative to C3 plants WUE was 3.3-fold greater (Côte et al., 1993).
(Winter et al., 1982). Leaf and canopy transpiration rates for
When CO2 fixation by pineapple is pineapple are lower than those for most cul-
expressed on a unit area of soil, fixation rates tivated crops. At midday, there was no mea-
are also much lower than those for C3 plants. surable water-vapour loss from a full
For example, in controlled conditions, a pineapple canopy with an LAI of 7 (Ekern,
pineapple plant with a leaf area index (LAI) 1965) and transpiration from pineapple
close to 4 fixed 20–25% of the CO2 fixed by plants in pots was 1.0 mm day−1 (monthly
wheat (Côte et al., 1993). However, dry-mat- average) (Shiroma, 1971). The transpiration
ter accumulation by a pineapple crop is high. ratio (TR) (units water lost per unit dry-mat-
Lacoeuilhe (1976) reported that pineapple ter gain; kg kg−1), a measure of the WUE, for
accumulated 41 t of dry matter ha−1 during a pineapple was about 50 (Sideris and Krauss,
crop cycle in Côte d’Ivoire. Bartholomew 1955; Joshi et al., 1965; Côte, 1988) to 116
(1982) reported 62 t of dry matter ha−1 in 24 (Neales et al., 1968), while the range for C3
months in Hawaii. By way of comparison, crops was 450–950 and that for C4 crops
dry-matter production of wheat was 250–350 (Kluge and Ting, 1978). A conse-
reported to be between 18 and 29 t ha−1 year−1 quence of the low rate of water use is that
(growing season), while that for sugar cane the crop can sustain an LAI of greater than 7
was 67 t ha−1 year−1 (Loomis and Gerakis, (Malézieux, 1991; Zhang, 1992) over long
1975). The large dry-matter accumulation by periods under low-rainfall conditions
pineapple is associated with a high LAI and (Ekern, 1965; Aubert, 1973).
the ability of leaves to maintain their photo-
synthetic capacity for long periods of time.
Origins of the high water use efficiency
related to CAM
Diffusion of water vapour from and CO2 into
Water-use efficiency related to CAM in
plant leaves can be described by the equa-
pineapple
tions:
Transpiration rhythm and transpiration rate in W = r T (1)
pineapple CO2 = 1.6 r P (2)
In pineapple, the diel rhythms of stomatal where W is the difference in water-vapour
conductance and transpiration are closely pressure between the stomatal cavity and the
linked to the net CO2-uptake rhythm (Neales atmosphere, r is the resistance to water-
et al., 1968; Nose et al., 1981; Bartholomew, vapour diffusion, T is the transpiration rate,
1982; Côte et al., 1993; Fig. 5.3). The lowest CO2 is the difference in CO2 partial pres-
rates of transpiration occurred during phases sure between the stomatal cavity and the
I, II and III and the highest during phase IV. atmosphere, 1.6 is the ratio of the diffusion
76 E. Malézieux et al.

2.5 0.25
Night Day

2.0 0.20
CO2 fixation
(mmol h–1 per plant)

(mmol h–1 per plant)

Transpiration
0.15
CO2 exchange

1.5

Transpiration
1.0 0.10

0.5 0.05

0 0
0 6 12 18 24
Time (h)

Fig. 5.3. Transpiration and net CO2 exchange during a night/day cycle by a ‘Smooth Cayenne’ pineapple
plant (Côte et al., 1993). Environmental conditions include: photosynthetic photon flux density, 650 mol
m−2 s−1; photoperiod, 12 h night/12 h day; night/day temperature, 22°C/28°C. The plant (fresh weight of
aerial parts = 120 g) was obtained from in vitro culture. Data are the averages for 3 consecutive days.

coefficients for water vapour and CO2 in air Effects of environmental factors
and P is the rate of net CO2 fixation.
Combining equations (1) and (2) gives: Effects on photosynthesis
T = 1.6 × W × P/CO2 (3) TEMPERATURE. The effects of temperature on
photosynthesis of pineapple are complex and
The low transpiration rate in CAM plants is
relatively few studies have been conducted.
related to morphological and physiological
While the results of these studies are often not
adaptations that influence W, P and CO2
of equation (3) to reduce T: directly comparable, the general trends show
that the intensity of CAM (phase I) is highest
• Net CO2 fixation (P) occurs mainly during when the light/dark temperature differential
the night when W is low. is approximately 10°C and the night tempera-
• During the day, when W is high, malate ture is cool, about 20°C. These results are con-
decarboxylation increases the internal sistent with other observations that the
CO2 concentration; consequently P is low optimum day/night temperature for growth
or even zero. also is about 30/20°C (Neild and Boshell,
• During phase IV CO2 is high (Côte et al., 1976; Bartholomew and Malézieux, 1994).
1993), reducing P and T.
The highest leaf conductances recorded INCREASING NIGHT TEMPERATURE. At a given day
for pineapple were about 3% of those temperature, an increase in the night temper-
observed for cotton (Neales et al., 1968). This ature decreases night net CO2 uptake
low conductance was attributed to a thick (Connelly, 1972; Neales et al., 1980; Zhu et al.,
mesophyll, low stomatal frequency and 1999), as well as the proportion of night fixa-
sunken stomata located beneath multicelled tion in a 24 h period (Connelly, 1972;
trichomes (Bartholomew and Kadzimin, Bartholomew, 1982; Bartholomew and
1977). Malézieux, 1994; Zhu et al., 1999). Neales et
al. (1980) and Zhu et al. (1999) reported a 50%
 Crop Environment, Plant Growth and Physiology 77

decrease in night uptake as the night temper- IRRADIANCE AND PHOTOPERIOD. Because CO2
ature was increased from 20 to 25°C at a con- assimilation in CAM plants, including
stant day temperature of 30°C. The decrease pineapple, occurs via the photosynthetic car-
could be due in part to an increase in respira- bon reduction cycle, it is assumed that pho-
tion-derived CO2 fixation, but leaf titratable tosynthesis will increase with increasing
acidity also decreases (Min, 1995), indicating irradiance, and the results of Shiroma (1977)
less total CO2 dark fixation. Some studies and Nose et al. (1981, 1985, 1986) support this
show little or no change in net CO2 uptake assumption. Nose et al. (1985, 1986) found
during the day with an increase in night that light saturation of pineapple plants
temperature (Neales et al., 1980; Zhu et al., occurred at a photosynthetic photon flux
1999), but Connelly (1972) reported a dra- (PPF) of about 500 mol m−2 s−1. In vitro-
matic increase when night temperature grown plants maintained under continuous
increased from 15 to 30°C. It is at present not illumination also saturated at a PPF of 500
possible to resolve this discrepancy. mol m−2 s−1 (Côte, 1988). However, it is
The relationship between net CO2 fixa- probable that variation in irradiance can
tion and dry-matter accumulation under alter the pattern of CO2 fixation. For exam-
various thermoperiods has also not been ple, high irradiance decreases the duration of
well studied. In one study, net CO2 fixation phase III and increases the duration of phase
of single attached leaves of plants grown for IV assimilation relative to what occurs at low
8 months at 30/20°C was 1.6 times higher irradiance (Nose et al., 1986). This is because
than that for leaves of comparable plants the rate of malate-decarboxylation-depen-
grown at 30/25°C, while whole-plant dry- dent CO2 assimilation is more rapid during
matter accumulation was only 1.1 times phase III (Sale and Neales, 1980), so the
higher in the cooler environment (Zhu et al., malate pool is depleted more rapidly.
1997a). However, assimilation was mea- Increasing irradiance probably also increases
sured only on ‘D’ leaves, while dry-matter the quantity of soluble sugar formed in
accumulation reflects the contribution of leaves during phase III and IV. An increased
leaves of all ages. supply of sugar available for the production
of PEP, the substrate required for CO2 fixa-
INCREASING DAY TEMPERATURE. At a given night tion into malate at night, would increase the
temperature, the fraction of total CO2 assimi- amount of CO2 fixed into malate during
lated in a 24 h period that is fixed at night phase I. Consistent with this hypothesis, an
increases with increasing day temperature increase in irradiance during the day
(Neales et al., 1980; Bartholomew, 1982; increased CO2 fixation the following night
Bartholomew and Malézieux, 1994; Zhu et al., (Nose et al., 1981, 1985, 1986; Shiroma et al.,
1999). This increase is due primarily to a 1977), and leaf titratable acidity also
decrease in net CO2 uptake in the day and to a increased with increasing irradiance (Sideris
lesser extent to an increase in net CO2 uptake et al., 1948; Connelly, 1969; Aubert, 1971; Sale
at night (Connelly, 1972; Neales et al., 1980; and Neales, 1980). However, increases in
Zhu et al., 1999). For example, Zhu et al. (1999) night CO2 fixation that follow an increase in
reported that CO2 uptake in the day decreased irradiance generally are smaller than the
50% as the day temperature was increased increase in CO2 assimilation in the light.
from 30/25°C to 35/25°C, while the increase Thus, the intensity of CAM tends to decrease
in CO2 uptake at night was much smaller. As a as irradiance increases, at least until saturat-
result, there was a 37% increase in the fraction ing levels are attained.
of CO2 fixed at night. The biochemical basis Little is known about the effect of pho-
for the changes in CO2 uptake with changing toperiod on pineapple photosynthesis. Nose
temperature is not well understood. It is et al. (1986) reported that as day length
hypothesized that the decline in net CO2 increased from 10 to 16 h, total uptake in the
uptake during phase IV with increasing day light increased while uptake at night
temperature is associated with increased respi- decreased. However, expressed on an hourly
ration, but further studies are needed. basis, the increase in CO2 uptake in the light
78 E. Malézieux et al.

as day length increased was small, while the When well watered, CO2 fixation during the
decrease at night was relatively large. A day and night was normal, with 45% of the
40–50% reduction in CO2 uptake at night can CO2 being fixed at night. Depending on plant
be deduced from the data of Nose et al. size and perhaps moisture supply, drought
(1986) as photoperiod decreased from 16 to 8 reduced CO2 uptake in the light rapidly (Fig.
h. In natural environments, e.g. Hawaii, the 5.4) or relatively slowly (Zhu, 1996). With small
decrease in photoperiod from summer to plants, uptake in the day was nil after 4 days
winter is only 21%, while the decrease in without water, while, for large plants, net
irradiance is about 50%. uptake in the day by attached ‘D’ leaves ceased
after about 15 days. For small plants (Fig. 5.4),
CO2 CONCENTRATION. The effect of elevated
transitory net CO2 evolution occurred as the
CO2 concentration on pineapple photosyn-
period of water stress lengthened. Nocturnal
thesis has not been well studied, but gener-
CO2 fixation rate was not affected at the begin-
ally increases in response to elevated CO2
ning of water stress, but decreased progres-
have been observed. At a concentration of
sively as the duration of water stress
1700 mol mol−1 CO2, net CO2 fixation
lengthened (Fig. 5.4; Zhu, 1996). After rewater-
increased two- to threefold during phase IV
ing, CO2 uptake in both phase I and phase IV
(Côte, 1988) and a CO2 concentration greater
resumed rapidly (Fig. 5.4). The reduced sensi-
than 1000 mol mol−1 was required to satu-
tivity of night CO2 uptake to water deficit, as
rate photosynthesis in that phase. Saturation
compared with day CO2 fixation, and the
at such a high CO2 level indicates that
reversibility of the effects of water deficit in
pineapple has a high resistance to CO2 diffu-
pineapple are consistent with observations for
sion during phase IV.
other CAM plants (Kluge and Ting, 1978).
Zhu et al. (1997b) reported that the dry
Water deficit also alters the pattern of CO2
mass of pineapple plants grown for 4 months
assimilation. At the onset of water deficit,
at 730 mol mol−1 CO2 was 1.2 times higher
there was a transient stimulation of net CO2
than that of plants grown at ambient CO2
uptake at the beginning of the night period.
(330 mol mol−1). In that study, ‘D’-leaf titrat-
As drought was prolonged, the maximum
able acidity at the end of phase I was more
rate of CO2 assimilation shifted progres-
than 1.3 times greater for plants grown at ele-
sively towards the end of the night period
vated than at ambient CO2. Net CO2 uptake
(Côte et al., 1993; Zhu, 1996), and the shift
in phase I was also enhanced, but only where
was more marked at 25°C night temperature
average temperature was above 25°C. Uptake
than at 20°C. Zhu (1996) also showed that
of CO2 during phase IV was also enhanced
the rate of decline of net CO2 assimilation
and the effect of enrichment was greater as
during drought was related to the day/night
the average temperature increased from 25 to
temperature regime. Fluctuations in leaf
30°C (Zhu et al., 1997b). Nocturnal CO2 fixa-
titratable acidity indicated that significant
tion has been reported to be insensitive to
assimilation – about 35% of the maximum –
high CO2 levels in other CAM plants, so
still occurred after 70 days of drought at
the origin of these different responses is
30/20°C, while at warmer temperatures
unknown. Further studies are required to
(35/25 and 30/25°C), titratable acidity was
understand the combined effect of thermo-
20% or less of the maximum after only 40
period regime and elevated CO2 partial
days of drought. While pineapple is highly
pressure on pineapple photosynthesis.
tolerant of drought, assimilation declines
WATER STRESS. When CAM plants are subjected fairly rapidly with drought, and warm tem-
to drought, CO2 fixation during phase IV soon peratures hasten the rate of decline.
ceases. If the period of water stress is extended,
net CO2 fixation ceases but CAM plants con-
Effects on plant water relations
tinue to refix respired CO2, a phenomenon
referred to as CAM idling (Kluge and Ting, Transpiration rate is closely related to net
1978). Pineapple plants subjected to water CO2 uptake and to the differences in water-
deficit exhibit a similar response (Fig. 5.4). vapour pressure and CO2 partial pressure
 Crop Environment, Plant Growth and Physiology 79

18 H2O H2O H2O

Night fixation

Day fixation
(mmol 12 h–1 per plant)
14
Net CO2 exchange

10

–2
–10 –5 0 5 10 15 20 25 30 35
Time after cessation of watering (days)

Fig. 5.4. Effect of water deficit on the daily photosynthetic CO2 exchange by a ‘Smooth Cayenne’ pineapple
plant (Côte et al., 1993). The plant was placed on an inert culture support aluminium silicate). Watering
with the nutrient solution was stopped on day 0 and was resumed on day 26. Environmental conditions
include: photosynthetic photon flux density, 650 mol m−2 s−1; photoperiod, 12 h night/12 h day; night/day
temperature, 22°C/28°C. The plant (fresh weight of aerial parts = 100 g) was obtained from in vitro culture.

between the stomatal cavity and the atmos- higher during the day (51 mmol m−2 s−1)
phere (equation 3). Thus the effects of envi- than at night (8.1 mmol m−2 s−1) (Neales et al.,
ronmental factors on plant water relations 1980), reversing the typical inverted pattern
are closely linked to those of environmental of stomatal opening.
factors on photosynthesis (Côte, 1988; Zhu, Few data on gs are available under field
1996; Zhu et al., 1997a). conditions. In Côte d’Ivoire, gs data indicated
that the stomata opened earlier in the after-
TEMPERATURE. For pineapple held at a constant noon when the day/night temperature differ-
day temperature, daytime transpiration rates ential was 9°C than when it was 14°C (J.C.
decreased as the night temperature increased, Combres, 1981, personal communication).
while transpiration rates at night remained Conversely, a dry wind, such as the
constant (Yoder and Bartholomew, 1969a,b). Harmattan in West Africa, induced pro-
These results are consistent with the results longed closing of the stomata, with opening
of CO2 assimilation studies, which show that occurring late in the night (Py et al., 1987).
larger day/night temperature differentials Based on these data, we predict that con-
decrease assimilation in the light. Mean val- sumptive use of water by pineapple would
ues for WUE were higher when night tem- be higher in tropical environments, with high
peratures were lowest (Zhu, 1996). For night temperatures and a small day/night
example, WUE at night was more than 1.3 temperature differential, than it would be in
times higher for pineapple plants maintained more subtropical environments, where larger
in a day/night temperature of 30/20°C com- day/night temperature differentials prevail.
pared with those maintained at 30/25°C. In
the light, WUE was more than 1.5 times IRRADIANCE. Because pineapple transpires lit-
greater at 30/25 than at 35/25°C (Zhu, 1996). tle or not at all during the day, it is expected
Stomatal conductance (gs) of pineapple is that irradiance would have a limited effect
typically lowest during the day and highest on transpiration. However, high irradiance
at night, referred to as an inverted pattern of tends to increase the rate and duration of
stomatal opening. However, in a 10/25°C CO2 fixation during phase IV because the
day/night temperature regime, gs was much stomata open earlier in the afternoon. This
80 E. Malézieux et al.

would be expected to increase transpiration the root, stem and leaf meristems. Pineapple
during the day, and Nose et al. (1981) varieties that have strong apical dominance,
reported an increase in transpiration with such as ‘Smooth Cayenne’, generally do not
increasing irradiance. produce shoots from stem axillary buds
prior to flower induction, and shoots are typ-
CO2 CONCENTRATION. Zhu et al. (1999) reported ically not produced by ‘Smooth Cayenne’ in
a 1.1–1.2-fold increase in WUE during the warm tropical environments until after fruit
night and a 2.3–2.7 fold increase in the day for harvest. ‘Queen Victoria’ (Maerere, 1997) and
pineapple plants grown at 700 mol mol−1 ‘Red Spanish’ have weaker apical dominance
CO2 compared with those grown at 350 and may initiate shoots prior to flower
mol mol−1. Depending on the day/night induction, even in tropical environments.
temperature regime, the 24 h WUE was Shoots of the same initial mass, whether
1.3–1.6 times higher at 700 mol mol−1 than suckers or crowns, have approximately the
at 350 mol mol−1. Zhu et al. (1999) clearly same leaf area despite quite large differences
demonstrated that higher WUE was associ- in leaf number and size. Crowns have
ated with a two- to threefold decrease in numerous short leaves, while suckers have
stomatal conductance during phases IV and fewer but longer leaves. In an experiment in
I. Côte d’Ivoire, the leaf appearance rate in the
month after planting was 8 for crowns and 3
WATER STRESS. The patterns of gs during the for suckers (Lacoeuilhe, 1976). Beyond 3
onset of water stress are similar to those of months, leaf emergence rate was similar for
net CO2 uptake. Zhu (1996) observed a rapid crowns and suckers. During the first few
decrease in gs for pineapple plants subjected months of growth, leaves produced by suck-
to drought during phase IV and a somewhat ers are longer and heavier than those pro-
slower decrease during phase I. After 15 duced by crowns. For planting material of
days of drought, gs was near zero during similar size, leaf emergence rate and leaf size
phase IV. In phase I, the time when gs are similar within 3–5 months after planting
reached a maximum shifted from early in the (Py et al., 1987).
light period to towards its end. In ambient
CO2, WUE decreased progressively as water
Leaves
stress progressed. However, plants main-
tained at 700 mol mol−1 CO2 exhibited a Leaves represent approximately 90% of aerial
steady increase in WUE (Zhu, 1996). In a plant fresh weight during vegetative growth
field study, J.C. Combres (1981, personal (Py, 1959). Leaves grow from the base, the
communication) observed that, after rainfall maximum leaf length is reached several
following a period of drought, the stomata months after initiation and leaves of a wide
open progressively earlier in the morning range of sizes and ages are present on the
and in the afternoon in the following days. plant at the same time (Fig. 5.5). The time
from initiation to full elongation depends on
temperature and is approximately 4 months
Plant Growth and the Environment in the nearly equatorial conditions of Côte
d’Ivoire, but considerably longer in cooler
Main characteristics of vegetative plant environments. Sideris and Krauss (1936) sep-
growth arated pineapple leaves into categories based
on their similarity in size and age. The ‘D’
Pineapple is grown from a variety of propag- group of leaves were the longest on the plant
ules (see Hepton, Chapter 6, this volume) and had ‘succulent–brittle’ leaf bases. Over
but all develop in a similar manner. If condi- time it became common practice to apply the
tions for growth are favourable after plant- term ‘D’ to a single leaf, usually the tallest
ing, root initiation begins, followed by the leaf on the plant. This ‘D’ leaf represents an
appearance of new leaves. Between planting easily identified standard leaf (Fig. 5.6) that is
and inflorescence initiation, growth occurs in commonly used to index growth and evalu-
 Crop Environment, Plant Growth and Physiology 81

Fig. 5.5. Array of leaves present on a fruiting pineapple plant. The oldest leaves are to the right of the stem
and its associated peduncle and fruit. (Photo of E. Malézieux.)

ate plant nutrient status. When growth is

rapid, the tallest leaf is almost always nearly
but not fully expanded (Py et al., 1987). When
growth is unrestricted by stress, the maxi-
mum length, width, and weight of individual
leaves increase with each successive leaf until
a maximum is reached (Fig. 5.5). Maximum
leaf length may differ among pineapple vari-
eties, but, for ‘Smooth Cayenne’, the maxi-
mum length can reach 100 cm and the
maximum width can reach 7 cm (Py et al.,
1987). Total leaf area per plant can reach 2.2
m2 for a plant having a fresh weight of 3.6 kg
(Py, 1959). The LAI of a pineapple canopy can Fig. 5.6. The pineapple ‘D’ leaf is most accurately
reach 12 at floral induction, although values identified by the more or less parallel margins of the
leaf base as contrasted with the flared base of the
of 6–8 are more common.
older ‘C’ leaf on the left and the tapered base of the
young ‘E’ leaf on the right. For mineral-nutrient
Roots analysis, the middle one-third of the basal white
tissue is sampled. (Modified from Nightingale, 1949.)
Few data are available on root growth.
Though roots of ‘Smooth Cayenne’ pineap-
ple can theoretically reach a length of 1.8 m that root growth decreases after flower
and grow to a depth of 85 cm (Sideris and induction and that maximum root mass is
Krauss, 1934), the sensitivity of pineapple reached at anthesis.
roots to soil compaction (Rafaillac et al., 1978;
Ikan, 1990) generally confines the root sys-
Stem and axillary buds
tem to the tilled area (Ekern, 1965; Ikan,
1990; J.J. Lacoeuilhe and E. Malézieux, Stem mass increases progressively after
unpublished results). This restricted soil vol- planting, with no unique morphological
ume prospected by the roots can limit the changes until reproductive development
amount of water available to the plant. After begins. Plants may accumulate starch
planting, crowns produce more roots than reserves in the stem, but such accumulation
suckers (Py et al., 1987). There is evidence varies with plant age and size and aerial
82 E. Malézieux et al.

environment. Soon after floral induction d’Eeckenbrugge and Leal, Chapter 2, this
occurs in ‘Smooth Cayenne’, axillary bud volume). This shape and orientation channel
development usually begins, but active light rains and dew to the base of the plant,
growth is highly dependent upon climatic making a significant contribution to the
conditions. Axillary shoot growth occurs dur- water economy of the plant (Ekern, 1965,
ing fruit development on relatively large but 1968).
not on small ‘Smooth Cayenne’ plants in sub- Large trichomes completely cover both
tropical conditions, but only after fruit har- adaxial and abaxial leaf surfaces (Fig. 5.7),
vest in tropical environments having warm and a highly cutinized upper epidermis and
night temperatures. This may be related to a multicelled hypodermis are two significant
the higher starch content of the stem in plants morphological and anatomical features of
growing in subtropical conditions, where the pineapple leaves that contribute to the
night temperature is cooler and the plant’s water economy (Krauss, 1930, 1949;
day/night temperature differential is larger. see Coppens d’Eeckenbrugge and Leal,
The growth of one to a few suckers makes Chapter 2, this volume). There are 30–35 tri-
possible the production of one or more sec- chomes mm−2 on the abaxial leaf surface
ond fruits on the mother plant (ratoon crop) (Aubert, 1973), which cover strips of stomata
and these shoots are also used as propagules. located in furrows between ridges devoid of
stomata (Fig. 5.7). There are relatively few
stomata per unit leaf area (70–85 mm2) and
Morphological, anatomical and physiological
the stomatal pore is small (Krauss, 1949;
features and adaptations to environmental
Bartholomew and Kadzimin, 1977).
conditions
The mature ‘Smooth Cayenne’ leaf cross-
section can be up to 4 mm thick (Krauss,
Water use
1949), with approximately half the volume of
Leaves are arranged spirally around the stem the plant leaf occupied by a water-storage
in a dense rosette pattern (see Coppens parenchyma. The balance of the leaf volume

Fig. 5.7. Scanning electron micrographs (SEM) showing the large multicellular trichomes on the adaxial and
abaxial surfaces of a ‘Smooth Cayenne’ pineapple leaf. Note that the trichomes are much more prominent
on the abaxial surface. In the right-hand SEM, the trichomes have been removed from a section of the
abaxial surface to show the rows of stomata located in furrows that parallel the longitudinal axis of the leaf.
(Scanning electron micrographs of D. Bartholomew.)
 Crop Environment, Plant Growth and Physiology 83

is chlorenchyma. This water-storage d’Ivoire, water content was less and specific
parenchyma functions as a reservoir of leaf area (SLA) (g m−2, fresh weight basis)
water, which is utilized during periods of was greater in unirrigated than in irrigated
drought. The depleted tissue is replenished crops (E. Malézieux, unpublished results).
after adequate rain (Krauss, 1949; Sanford, The symptoms of drought develop slowly,
1962). the earliest being reduced growth and wilt-
As noted previously, these morphological ing of the older leaves (Swete Kelly and
and anatomical features result in low evapo- Bartholomew, 1993). With severe and pro-
transpiration (ET) (Ekern, 1965) and extreme longed drought, leaf colour changes from
tolerance to drought. Transpiration values of dark to pale green, then to pale yellow and
1.3 mm day−1 on cloudy days and 2.7 mm finally to red. At the later stages, leaf mar-
day−1 on fine days, average 2.1 mm day−1, gins curl downward, leaves lose their turgid-
were obtained for 15–16-month-old fruiting ity and become limp and growth stops (Py et
pineapple plants (Shiroma, 1973). In a pot al., 1987; Swete Kelly and Bartholomew,
study in sandy soil, transpiration was maxi- 1993).
mum during the summer when temperature The effects of drought are reversible and,
and solar radiation were highest (respec- when water again becomes available, the
tively about 27°C and 10–12 MJ m−2 day−1) leaves rehydrate and normal growth
(Shiroma, 1971); a low value of 0.4 mm day−1 resumes. Leaves not yet fully expanded
was estimated during the coldest periods. resume their growth. Leaf width rapidly
Average ET values for ‘Smooth Cayenne’ increases, resulting in a constriction at the
pineapple in field experiments in Hawaii point where elongation resumed (Py et al.,
were 0.83 mm day−1 with a plant-trash 1987). Such leaves generally develop spines
mulch and 1.25 mm day−1 with plastic mulch on the margins at the point where growth
(Ekern, 1965). For a leaf canopy formed by was restricted. Symptoms of water stress
small plants (LAI = 4.2) during fruit develop- may appear more rapidly where soil water-
ment, daily maximum ET was 1.3 mm on holding capacity is low, if rooting depth is
cloudy days (irradiance = 10.7 MJ m−2) and restricted or if the root system has been dam-
2.7 mm on a sunny day (irradiance = 16.6 MJ aged by pests or disease (Swete Kelly and
m−2), with an average of 2.1 mm day−1 Bartholomew, 1993).
(Shiroma, 1973). Measurements of ET made
in Côte d’Ivoire (N’Guessan, 1985) showed
Temperature
that values decreased from 0.25 mm h−1 over
a 10 h period 5–6 months after planting to Leaf and plant temperatures of this relatively
0.11 mm h−1 per 10 h period 7–10 months non-transpiring crop reach values that are
after planting. However, ET in Côte d’Ivoire detrimental, perhaps even lethal, to meso-
can reach 4.5 mm day−1 when solar radiation phytic crops. The temperature of horizontal
is high (Combres and Perrier, 1976). With leaves of 6-month-old ‘Smooth Cayenne’
irrigation, ET averaged 3.0 mm day−1 over a plants reached 48°C between 1 and 3 p.m. in
2-month period (Combres, 1979). Hawaii (Aubert and Bartholomew, 1973).
The difference between the leaf middle and
base reached 18°C and the maximum differ-
Drought
ence between leaf and air temperatures was
Despite the high resistance of pineapple to 18.6°C at 1 p.m. Such extreme leaf tempera-
drought, effects of drought on plant mor- tures caused no permanent harm, but their
phology and growth are important. In a dry specific effects on physiological processes
season, the width of young leaves, the rate of have not been studied. As discussed previ-
leaf emergence and the weight of successive ously, the effects of temperature on stomatal
‘D’ leaves may be reduced (Py, 1965). In conductance are significant.
Hawaii, leaf elongation decreased when the The effect of temperature on growth is
soil moisture content declined below 30–35% quite complicated. The morphology of
(Ekern, 1964). During the dry season in Côte ‘Smooth Cayenne’ plants is markedly
84 E. Malézieux et al.

affected by temperature. Optimum nutrition Whole-vegetative-plant SLAs over time were

in environments having a controlled night consistently lower for plants grown at night
temperature greater than about 25°C and in temperatures of 18, 22 and 26°C than for
warm, humid, low-altitude climates near the those grown at 30°C (Fig. 5.9). The lowest
equator produces plants with numerous, SLA was about 3.8 m2 kg−1 dry weight at a
wide, flaccid leaves (Fig. 5.8; Py et al., 1987; day/night temperature of 22/18°C
Bartholomew and Malézieux, 1994). (Bartholomew, 1982). SLA declines gradually
Conversely, in controlled environments with as plant mass and age increase (Fig. 5.9),
cool night temperatures (Friend, 1981) and in especially at the cooler night temperatures,
cooler climates, leaves are erect, straight, so comparisons of SLA between plants in dif-
rigid, shorter and fewer in number (Fig. 5.8; ferent environments must be made using
Py et al., 1987; Bartholomew and Malézieux, plants having approximately the same mass.
1994). In Hawaii, leaves of plants grown in Growth slows when night temperatures
high, cool elevations are shorter and more are cool, and a lower SLA at such tempera-
rigid than those on plants grown at lower tures will further reduce the rate of increase
elevations. in LAI and prolong the time required to
Indices such as SLA (leaf area per unit of reach full canopy closure and complete inter-
leaf dry mass, m2 kg−1) and leaf area ratio ception of available light. Thus, the rate of
(LAR) (leaf area per unit of total plant dry leaf area expansion will probably decrease
mass, m2 kg−1) allow the effects of tempera- more rapidly as night temperature decreases
ture on plant morphology to be quantified than would be predicted just on the basis of
and extend our ability to predict the effects the effects of temperature on leaf elongation.
of environment on vegetative growth. Mitigating this effect of cool temperature, at

Fig. 5.8. Effects of warm (30°C) and cool (22°C) night temperatures on the leaf orientation of ‘Smooth
Cayenne’ pineapple. The plants were grown for 5 months at the indicated temperatures. (Photo of D.
Bartholomew.)
 Crop Environment, Plant Growth and Physiology 85

8
Specific leaf area (m2 kg–1)

6
22/18
5 30/18
34/18
4
22/22
3 26/22
30/26
2
30/30
1 34/30

0
0 100 200 300 400
Days after treatment

Fig. 5.9. Effects of day and night temperature (°C) on the specific leaf area (SLA) (m2 kg−1 dry weight) of the
green leaf tissue of ‘Smooth Cayenne’ pineapple grown in controlled environments. Pineapple crowns were
started in a 26/22°C day/night temperature environment and transferred to the various temperatures 100
days after planting. Notice the gradual decrease in SLA with increasing time after planting. At a given
harvest date, larger plants generally had a lower SLA than did smaller plants. Plants harvested after 400 or
more days had produced fruit at night temperatures of 18 and 22°C while plants at warmer night
temperatures remained vegetative. Data points are means of total green leaf area (white and pale green
basal tissue removed) and dry weight for two plants. (Redrawn from Bartholomew and Malézieux, 1994.)

least to some degree, is the fact that plants elevation decreased and average tempera-
growing in cooler environments have a ture increased (Fleisch, 1988). The SLA and
higher net assimilation rate, which would LAR, both on a dry-mass basis, were related
offset to some degree the benefit imparted by to average air temperature (T) by the equa-
a more rapid rate of leaf expansion that tions:
would occur in warm environments. Thus,
SLA = 1.93T + 3.5, R2 = 0.33 (4)
dry-matter gain by plants in cooler environ-
LAR = 1.07T + 64.6, R2 = 0.37 (5)
ments may be as high as that which occurs in
warmer environments, where leaves expand where SLA is expressed in cm2 g−1 and T in
more rapidly. Unfortunately, simultaneous °C. The relatively low R2 values for equa-
changes in other climatic factors as tempera- tions (4) and (5) are assumed to be due, at
ture changes, possible differences in the least in part, to the decrease in SLA and LAR
growth rate of ‘Smooth Cayenne’ clones and with increasing plant mass and age.
differences in quality of management make it In contrast to many crops (Biscoe and
very difficult to compare productivity across Gallagher, 1977) and regardless of the air
environments having different temperature temperature, the leaves of the pineapple
regimes. mother plant persist at least into the first
Consistent with the effect of temperature ratoon in Hawaii, a period of more than 24
on SLA, LAR values for the ‘Smooth months. However, the functionality of
Cayenne’ clone ‘Champaka F-153’ also gen- pineapple leaves of various ages has not
erally increased with increasing night tem- been adequately determined.
perature (Fig. 5.10) and decreased with Both the ‘Smooth Cayenne’ pineapple leaf
increasing plant age. An increase in LAR was and stem are important storage organs and
also observed for plants grown in the field as can accumulate large quantities of starch.
86 E. Malézieux et al.

7
Leaf area ratio (m2 kg–1)

5 22/18
34/18
4 26/22
3 30/22
30/26
2 30/30
1 34/30

0
0 100 200 300 400
Days after treatment

Fig. 5.10. Effects of day and night temperature (°C) on the leaf area ratio (m2 of leaf kg−1 dry weight of
plant) of ‘Smooth Cayenne’ pineapple grown in controlled environments. Pineapple crowns were started in
a 26/22°C day/night temperature environment and transferred to the various temperatures 100 days after
planting. Data are means for two plants. (Redrawn from Bartholomew and Malézieux, 1994.)

The stem of a crown at the time of fruit mat- 30°C, plants had lower dry-matter content
uration can contain more than 20% starch on and little starch in the stem, while plants
a dry basis. Stem fresh weight increases grown at night temperatures of 18 and 22°C
slowly during the first months after planting had significant amounts of starch
and during this time, starch and dry matter (Bartholomew and Paull, 1986). Plant dry-
content initially decline and remain at about matter and starch content are believed to be
10–12% for several to many months – about especially important after floral induction
9–11 months in Hawaii (Pineapple Research (see Bartholomew et al., Chapter 8, this
Institute of Hawaii, personal communica- volume).
tion). In Hawaii, dry-matter content and While stem starch levels at the time of flo-
stem starch then begin to increase and stem ral induction can be as high as 60% in
dry-matter content can reach 30% (Fig. 5.11) Hawaii, starch levels do not reach 20% at the
and starch can exceed 60%, dry basis, at the time of forcing in Côte d’Ivoire, where tem-
time of flower induction (Bartholomew and peratures are higher. Forcing occurs much
Paull, 1986). At a plant population density of sooner after planting in Côte d’Ivoire than in
4.3 plants m−2, at 458 days after planting Hawaii, because suckers are the primary
when flowering was forced, stem starch source of planting material and vegetative
accumulation reached about 325 g m−2 of growth is more rapid in the warm environ-
ground area (Pineapple Research Institute of ment. Because the LAR increases as night or
Hawaii, personal communication). About 90 average temperature or both increase, large
days later, starch mass per unit of ground differences exist between statistical models
area exceeded 800 g m−2. In environments of vegetative growth developed in Hawaii
warmer than Hawaii, there may be no (Fleisch, 1988; Zhang, 1992) and in Côte
increase in stem dry-matter content until d’Ivoire (Malézieux, 1991).
vegetative growth is interrupted by forced Leaf dry-matter content also gradually
induction. Data for controlled conditions increases with time during vegetative and
showed that, at night temperatures of 26 or early reproductive growth (Fig. 5.11) and,
 Crop Environment, Plant Growth and Physiology 87

35

SDM
30
LDM
Dry matter content (%)

25

20

15

10

0
0 4 8 12 16 20 24 28
Months after planting

Fig. 5.11. Dry-matter content of ‘Smooth Cayenne’ pineapple stem (SDM) and leaves (LDM) from 3 months
after planting in December 1932 until after plant-crop fruit harvest in July 1934. The arrows indicate the
approximate time of natural induction of flowering and of fruit harvest. (Redrawn from King, 1935.)

though such changes are relatively small, (Ravoof, 1973). Also, as root temperature
when total storage per unit of land area is increased, the percentage of dry matter allo-
computed, the accumulation is significant. In cated to leaves increased from 77 to 80%,
an unpublished study conducted in Hawaii while that allocated to stem decreased from
(Pineapple Research Institute of Hawaii, per- 17 to 13% (Ravoof, 1973). Plant LAR prob-
sonal communication), the starch content of ably increased as root temperature increased.
leaves sampled early in the morning aver-
aged about 0.25% for the first 400 days after
Light interception
planting. Starch then gradually increased to
1.27% at 570 days after planting (120 days The spectral properties of the adaxial surface
after forcing). At that time, plants had accu- of pineapple leaves over the wavelength
mulated an average of 254 g m−2, land-area range from 520 to 750 nm were not signifi-
basis, of starch in leaves. Both stem and leaf cantly different from those of wheat, olive,
starch declined sharply after floral induction, orange and peach (N’Guessan, 1985).
presumably as reserves were drawn upon to However, pineapple leaves have very low
support fruit and sucker growth. In tropical reflectances in the infrared region with min-
environments, little starch is accumulated in ima at 1440 and 1990 nm (water peaks).
plant tissues by the time of forcing, so growth Despite reflectances in the visible range com-
of fruit and propagules such as suckers, is parable to those of mesophytes, much of the
dependent on current photosynthesis. In radiant energy falling on a pineapple plant is
these environments, propagule development entrapped by multiple reflections within the
is delayed until after the fruit is harvested. rosette leaf array (Ekern, 1965). As a conse-
Changes in plant dry-matter content and quence, the total reflectance of the canopy is
partitioning with changing root temperature low (Ekern, 1965), resulting in canopy albe-
at ambient air temperature were similar to dos that ranged from 0.14 to 0.16 for
the effects of temperature observed in con- canopies ranging in age from 1 to 16 months
trolled environments. Leaf dry-matter con- (Shiroma, 1973; Combres, 1983). There was
tent decreased from 16.5 to 12% as the root no significant variation in albedo with
temperature increased from 15 to 30°C canopy height (N’Guessan, 1986).
88 E. Malézieux et al.

Estimated canopy light extinction coeffi- Effects of environment on vegetative growth

cients for pineapple are typical of canopies
with a relatively erect leaf orientation. Specific effects
Extinction coefficients (k) of 0.56–0.59 were
TEMPERATURE. The plant growth rate of
calculated for ‘Smooth Cayenne’ pineapple
pineapple is strongly influenced by tempera-
in Hawaii (Fleisch, 1988; Zhang, 1992),
ture but few studies have been made in con-
resulting in 95% light interception at an LAI
trolled environments and the results from
of about 5.0. Extinction coefficients for meso-
field studies can be difficult to interpret. Leaf
phytic crops typically range from 0.4 for and root elongation rates measured in con-
erectophile canopies to 0.8 or greater for trolled environments over a range of temper-
planophile ones (Russell et al., 1989). A sig- atures (Sanford, 1962) generally show good
nificantly lower k of 0.29 was found for correspondence with the results of later stud-
pineapple in Côte d’Ivoire by measuring ies. The optimum temperature for leaf
total radiation interception of pineapple growth was 29°C and that for roots was 32°C
crops with a somewhat wider row spacing (Sanford, 1962). Growth of both organs was
(Malézieux, 1991). Low k values account in greatly reduced at temperatures below 20°C
part for the ability of pineapple to sustain and ceased at about 10°C.
very high LAIs. Data on the long-term effects of tempera-
ture on dry-matter accumulation are sparse
Photoperiod and irradiance and such data are needed if the vegetative
growth responses in the diverse field envi-
The data on the effects of photoperiod and ronments where pineapple is grown are to be
irradiance are sparse and few useful general- interpreted. Pineapple plants grown in con-
izations can be made from them. Also, trolled environments having night tempera-
changes in these parameters in natural envi- tures of 26°C or less had greater weights
ronments are often confounded with temper- than those grown in environments having
ature. At low light, seedling leaf lengths 30°C night temperatures, and the differences
decreased as the PPF increased from 100 to increased over time (Fig. 5.12; Bartholomew,
325 mol m−2 s−1 (Aromose, 1989). Average 1982). After about 400 days, plants grown at
leaf length was greater in 16 than in 8 or 24 h cooler night temperatures accumulated two
photoperiods (Aromose, 1989). At an aver- to three times the dry mass of those grown at
age PPF of 400 mol m−2 s−1, the length of 30°C night temperatures (34/30 and
the longest leaf of plants grown for 692 days 30/30°C). In environments with warm days
in controlled conditions increased as pho- (26–34°C) and cool nights (18–22°C), plants
toperiod increased from 6 to 10 h, was also had higher dry-matter contents and
unchanged at 12 h and declined sharply at 16 greater plant relative growth rates and leaf
h (Friend and Lydon, 1979). The width of the growth rates than did plants grown in warm
longest leaf increased continuously with days and warm (30°C) nights (Bartholomew,
increasing photoperiod while leaf thickness 1982). Plant leaf area at the end of the study
(g cm−2, fresh weight basis) declined (Friend was also greatest for plants grown at 30°C
and Lydon, 1979). day and 26 or 22°C night temperatures (Fig.
In natural environments, leaves of plants 5.13). The more rapid growth of plants at the
grown in low irradiance are long, erect and cooler night temperatures resulted mainly
dark green, while those of plants grown in from higher net assimilation rates. Net
very high irradiance become red or even yel- assimilation rates on a unit leaf basis ranged
low (Py et al., 1987). Physical damage and from about 1.0 g m−2 day−1 for plants grown
death, typically referred to as sunburn, due at 30°C dark temperatures to 2.0 g m−2 day−1
to high irradiance was reported in India for plants grown at dark temperatures of 26
(Srivastava and Singh, 1971). Connelly (1969) or 22°C (Bartholomew, 1982). Friend (1981)
found that average ‘D’-leaf length was 52 cm also reported that plants grown at a 30°C
in full sun, 55 cm in 25% shade and 50 cm in night temperature were smaller than those
50% shade. grown at cooler night temperatures.
 Crop Environment, Plant Growth and Physiology 89

1000

900 34/18
26/22
800 30/22
30/26
700
30/30
Plant dry weight (g)

600 34/30

500

400

300

200

100

0
0 50 100 150 200 250 300 350

Days after treatment

Fig. 5.12. Effects of day and night temperature (°C) on growth in dry mass of ‘Smooth Cayenne’ pineapple
plants grown in controlled environments. Pineapple crowns were started in a 26/22°C day/night
temperature environment and transferred to the various temperatures 100 days after planting (day 0 in the
figure). Each data point is the mean for two plants. (Redrawn from Bartholomew and Malézieux, 1994.)

4.5
34/18
4 26/22
Green leaf area (m2 per plant)

30/22
3.5 30/26
30/30
3
34/30
2.5

1.5

0.5

0
0 50 100 150 200 250 300 350
Days after treatment

Fig. 5.13. Effects of day and night temperatures (°C) on growth in green leaf (white and pale green leaf basal
tissue excluded) area per plant of ‘Smooth Cayenne’ pineapple plants grown in controlled environments.
Pineapple crowns were started in a 26/22°C day/night temperature environment and transferred to the
various temperatures 100 days after planting (day 0 in the figure). Each data point is the mean for two plants.
(Redrawn from Bartholomew and Malézieux, 1994.)
90 E. Malézieux et al.

Under field conditions, the mother plant (Bartholomew, 1982). A relatively sharp tem-
relative growth rate for the period from perature optimum occurs for leaf elongation
about 2 months after planting to induction of rate (Sanford, 1962), but no such sharp opti-
flowering was reduced by the cool winter mum was observed for dry-weight gain, pre-
temperatures of south Queensland (Sinclair, sumably because of morphological and
1992). When data from Hawaii and Côte anatomical differences induced by tempera-
d’Ivoire are compared, plants grown from ture or the day/night temperature differen-
crowns in Hawaii have 0.7 m2 of leaf area 8 tial, or both.
months after planting and would be grown Leaf initiation in natural environments, as
another 4–6 months before forcing indicated from rates of leaf appearance, is
(Bartholomew and Kadzimin, 1977). In Côte sensitive to temperature. In Taiwan, the
d’Ivoire, plants of the same age grown from equation:
larger suckers would have 1.8 m2 of leaf area
NLEAF = 18.125 × log10 T/11.669 (6)
and would be ready to be forced (Malézieux,
1991). Some of the difference in size of plants where T is mean monthly temperature in °C
after 8 months of growth is almost certainly and NLEAF is number of leaves to appear in
due to the larger propagules used in Côte 1 month, indicates the extent to which leaf
d’Ivoire, but some of the difference is also appearance rate is influenced by temperature
probably due to the more rapid expansion of (Shiroma, 1972). Data on leaf emergence col-
leaf area in the tropical environment of Côte lected from fields at elevations of 91, 305 and
d’Ivoire. In Thailand, large suckers, some 792 m in Hawaii could be described by a
weighing up to 2.0 kg, are planted and these similar relationship (D.P. Bartholomew, 1988,
will be forced within 5–6 months after plant- unpublished results). In Côte d’Ivoire, which
ing. has a high mean air temperature and narrow
It seems appropriate to interject here that, diurnal variation, the number of leaves
while much has been written about the rela- appearing per 2 months was best described
tive advantage of suckers over smaller by a quadratic function and showed an opti-
propagules (shorter time from planting to mum temperature at about 26–27°C (E.
forcing; larger average size of fruit), only Malézieux, 1992, unpublished results).
fairly recently has it been recognized that the Similar results showing the existence of an
size of the propagule is the main factor optimum temperature near 28°C were
accounting for the difference. A large obtained in Hawaii (J. Zhang, 1992, personal
propagule produces a large plant more communication).
quickly than does a small propagule. Leaf number can be predicted using heat
In areas where there is considerable sea- accumulation or daily thermal time (DTT)
sonal variation in climate, time of planting (Zhang et al., 1997). An enhanced model was
can also affect time from planting to forcing. developed which also accounted for the
In Hawaii, crowns will be forced 12–14 reduction in growth that occurs at low
months after planting (A. Hepton, personal (below 16°C) and high temperatures (greater
communication). Propagules planted during than 35°C). Such temperatures would nor-
periods of cool weather and low irradiance mally be reached only during part of the day
establish and grow more slowly than do (Zhang et al., 1997). DTT was calculated for
those planted when weather conditions are each day by the equation:
more favourable for growth.
DTT = TM – BT (7)
Theoretically, growth in leaf area depends
on the rate of both leaf initiation and expan- where TM is the mean temperature and BT
sion. Leaf expansion, as indicated by rate of the base temperature. A BT of 16°C provided
elongation, increased with temperature up to the best fit for the data. For a minimum tem-
29°C (Sanford, 1962). Leaf expansion was perature BT or a maximum day tempera-
also greater where the day/night tempera- ture  35°C, DTT was calculated using a
ture differential was large than where it was procedure similar to that reported by
small and night temperature was high Malézieux et al. (1994) for the estimation of
 Crop Environment, Plant Growth and Physiology 91

daily fruit thermal time. The leaf-emergence This surprising result is assumed to be due
model was calibrated and validated with to the fact that researchers who studied the
data from Hawaii and Côte d’Ivoire. Based effects of plant population density on
on Celsius temperature, the phyllochron pineapple generally did not collect data on
(interval between the appearance of two suc- plant fresh or dry mass, did not report such
cessive leaves) was 50 degree-days per leaf. data or estimated plant mass from ‘D’-leaf
Interplant competition had a significant mass. ‘D’-leaf mass is a faulty estimator of
effect on leaf number, and it was assumed plant mass, because as plants get larger, ‘D’-
that this effect was due to a reduction in leaf mass reaches a maximum, while leaf
meristem temperature caused by the intense number and leaf and plant mass continue to
shading that occurs at high plant population increase (Py et al., 1987).
densities. This effect was simulated in the Interplant competition does occur during
model using interplant competition factors vegetative growth of pineapple, and growth
calculated from leaves m−2 of ground area is reduced at high plant population densities
(Zhang et al., 1997). (Malézieux, 1992a; Zhang, 1992). In a study
In addition to the effects of root tempera- in Hawaii, there was no reduction in plant
ture on dry-matter partitioning noted above, mass with increasing plant population den-
Ravoof (1973) also showed that the total sity for the first 200 days after planting, even
fresh mass of 40-day-old plants grown from at a density of 123,500 plants ha−1, and light
slips increased with increasing root tempera- interception did not reach 95% at that den-
ture from 15 to 30°C, though the increase in sity until about 350 days after planting
growth was not proportional to the increase (Zhang, 1992). Plant mass at forcing, which
in root temperature. In Okinawa, Japan, a occurred for plantings made in June, August
relatively cold climate for pineapple (mean and October at approximately 440, 380 and
temperature
17°C, minimum temperature 335 days, respectively, after planting,
for more than 1 month  6°C) and in Hawaii decreased significantly with increasing den-
during the winter (mean temperature sity at all planting dates. Similar results were

22.3°C), growth is enhanced by black poly- obtained in Côte d’Ivoire (Malézieux, 1992a).
ethylene-film mulch (Ekern, 1967; Ogura et In the latter experiment, plant mass 8
al., 1968). The 33% increase in growth in months after planting increased by 33 and
Hawaii resulted from a soil temperature 20%, in irrigated and unirrigated conditions,
increase of only 1.6°C. In the Canary Islands, respectively, as plant population density
where mean average air temperature is decreased from 100,000 to 20,000 plants ha−1.
about 20°C, both black polyethylene mulch It is clear that competition for light occurs
and a polyethylene cover over the plants during vegetative growth in pineapple fields
increased yields (Galan Sauco et al., 1988b). even at moderate planting densities.
However, it may not be detectable where
IRRADIANCE AND PLANT POPULATION DENSITY. Few growth is limited by water or other stresses.
studies have been conducted on the effects of Unless the relationship between ‘D’-leaf
irradiance on vegetative growth. Connelly weight and plant mass is known, ‘D’-leaf
(1969) found that estimated plant weight of weights should not be used as an indicator
‘Smooth Cayenne’ plants grown in pots was of plant mass beyond about 6 months after
unaffected by 25% shade but was reduced by planting, or earlier than that where large
20% under 50% shade. Less nitrogen was suckers are planted. The effects of plant pop-
required for optimum growth at low than at ulation density on fruiting are discussed in
high light. detail later (see Bartholomew et al., Chapter
In most crops average plant mass 8, this volume).
decreases with increasing plant population Lest the details obscure the important
density due to interplant competition for message, it is clear that total biomass and
light, but no data illustrating this effect on often total yield increase with increasing
pineapple were found (Bartholomew and plant population density. However, studies
Kadzimin, 1977; Py et al., 1987; Scott, 1992). often show that, at densities much above
92 E. Malézieux et al.

74,000 plants ha−1 and sometimes well below that, when interplant competition is intense,
that, the percentage of fruit recovered, as there is insufficient photosynthate for the
well as the quantity and quality of products, development of both a fruit and suckers. The
declines. fruit is a stronger sink than developing suck-
Allometric data help to explain the effects ers, since forced plants will always develop a
of plant population density on vegetative fruit but may not develop suckers until after
growth. Leaf appearance rate decreased with the fruit matures. In tropical regions, such as
increasing planting density (Norman, 1977; Côte d’Ivoire, sucker growth was increased
Malézieux, 1992; Zhang, 1992) and, in one up to 20% by cutting off the leaves that
study, leaves were longer and narrower at extend above the developing suckers imme-
the higher planting densities (Wee, 1969). diately after fruit harvest (Combres, 1983).
Specific leaf mass (SLM) (dry-mass basis) Leaf removal may improve sucker exposure,
was significantly decreased when planting thus promoting their growth.
density was higher than 6 plants m−2.
However, at the base of the leaf, fresh weight DROUGHT. Despite the xeromorphic charac-
per unit area increased at high densities teristics of pineapple, growth and yield are
(Malézieux, 1992a). Zhang (1992) reported significantly reduced when drought is pro-
that SLA at 300 days after planting was longed. One effect of a water deficit is to
related to planting density (PD) by the equa- reduce the number and length of the roots
tion: (Kadzimin, 1975; E. Malézieux and G. Godo,
1991, unpublished results). White root tips
SLA = 50.3 + 0.51PD (8)
visible in the soil, expressed as a percentage,
and the effect was highly significant. have been used as an indicator of root health
The production of vegetative propagules, and water deficit (Sanford, 1962; Fig. 5.14),
such as slips and suckers, requires an input and root elongation ceases and root tips
of carbohydrates, at least until they develop suberize when the soil moisture approaches
sufficient leaf area to become autotrophic. −1.5 MPa (Ekern, 1967). Suberization does
Competition for light at higher planting den- not protect roots indefinitely as they will die
sities is assumed to reduce the supply of if the soil moisture stress is severe and pro-
assimilates available for growth and for stor- longed (Krauss, 1959). In Côte d’Ivoire, root-
age reserves, because the number of slips ing depth and root numbers increase with
and suckers per plant decreases at higher irrigation or with polyethylene mulch (E.
plant populations (Lacoeuilhe, 1974; Py et al., Malézieux and F. Zampatti, 1991, unpub-
1987). For clones that produce slips, slip lished results).
numbers decrease at a faster rate with Most of the quantitative data on the
increasing plant population density than do effects of water stress on growth were
suckers (Py et al., 1987). Suckers per unit area obtained on plants grown in pots; few data
may increase or, as occurred in a recent were found on the effect of water deficit on
study, become normalized across planting vegetative growth rate. Sideris and Krauss
density (Scott, 1992). Scott (1992) showed (1928) found that plants did not grow in soils
that, as plants per hectare increased from containing 5% moisture and, relative to well-
46,112 to 80,650, suckers per plant decreased watered soil, growth decreased significantly
from 1.84 to 1.11. Thus, despite a wide range in soils containing 10 or 15% moisture.
of planting densities, suckers ha−1 only Growth was not different in soils maintained
ranged from 82,162 to 89,572 (Scott, 1992). at 20, 25 and 30% moisture. In another pot
In addition to reducing the number of study, growth in a loamy soil continued at a
suckers per plant, sucker development, soil moisture content of 10% but almost
which in Australia (Sinclair, 1992) and stopped when soil moisture fell to 5% or less
Hawaii is initiated at the time apical domi- (Shiroma, 1971). In a pot study where the
nance is broken by forcing fruit develop- irrigation interval was varied from twice
ment, may also be delayed at higher planting weekly to bimonthly, ‘D’-leaf weight, leaf
densities. Circumstantial evidence suggests area, dry weight of all plant parts and fresh
 Crop Environment, Plant Growth and Physiology 93

Fig. 5.14. Variation in the percentage of white root tips used to diagnose root health and water availability
(modified from Nightingale, 1949).

fruit weight all decreased significantly as the decrease was due to a reduction in meristem-
irrigation interval was extended (Chapman atic activity associated with the internal
et al., 1983). After an early establishment water deficit (Kadzimin, 1975). In the clay
period, Kadzimin (1975) grew pineapple in soils of Hawaii, where water-holding capac-
black polyethylene bags for 7 months at ity is high, irrigation begun when soil
average soil water potentials (soil) of −0.1, reached −0.4 MPa was equal to or better than
−0.5, −1.0 and −1.5 MPa and in an unirri- irrigating at a soil of −0.03 or −0.07 MPa
gated treatment. In that study, plant total dry (Thorne, 1953). However, where soils are
mass decreased 27.6, 32.4 and 46.9%, respec- sandy and soil water-holding capacity is low,
tively, in the −1.0 and −1.5 MPa and unirri- growth was reduced as soon as the soil fell
gated treatments. Only leaf and stem growth below −0.015 MPa at a 15 cm depth
were reduced at −1.0 MPa; in the −1.5 MPa (Combres, 1979). Combres (1979) concluded
and unirrigated treatments, leaf, stem and that a soil of −0.015 could be used as a
root growth were reduced. The root system threshold value to initiate irrigation
as a proportion of the whole plant decreased (Combres, 1983).
significantly in the −1.5 MPa and unirrigated Plant moisture content decreases with
treatments. It was suggested that the increasing soil moisture stress and the thick-
94 E. Malézieux et al.

ness of the leaf water-storage tissue (Fig. leaves due to translocation to the fruit were
5.15) decreases as the plant water deficit estimated at 18 and 26.8% of leaf weight for
increases (Fig. 5.16). However, leaf water the sun and shade plants, respectively.
content changes slowly and pineapple toler- Pineapple plants show a remarkable ability
ates long periods of drought with minimal to transfer water and other nutrients from
plant loss. Nose et al. (1981) observed only a the leaves to the fruits when water stress is
0.1% decrease in leaf water content as soil pF extreme.
– a measure of soil water status – decreased Though normal values of pineapple leaf
from 1.1 to 4.5. Chapman et al. (1983) water potential (L) were about −0.6 MPa at
reported that the water content of ‘D’ leaves a soil of −0.1 MPa, L reached −2.3 MPa at a
was not significantly reduced by any irriga- soil of −1.8 MPa (Kadzimin, 1975). George et
tion treatment until fruit harvest and, even al. (1984) found that L of pineapple leaves
then, differences in the ‘D’-leaf moisture con- varied from −0.2 to about −1.8 MPa after 12
tents for the high and low water treatments weeks without any water supply, while leaf
were small. Even uprooted plants lose water relative water content (RWC) decreased from
slowly. Fruiting suckers removed from their about 96% to 42% over the same time period.
mother plants at flowering and kept without Leaf RWC and L were linearly correlated
water for 3 months on a glasshouse bench in with a 10% change in RWC corresponding to
sunlight lost 44.4% of their fresh weight by a 0.28 MPa change in L (George et al., 1984).
transpiration, while comparable shaded ones Similarly, Kadzimin (1975) found that, as L
lost 41.6% (Sideris and Krauss, 1955). Despite decreased from −0.1 MPa for well-watered
the high degree of dehydration, significant plants to −2.2 MPa for unirrigated plants,
fruit growth occurred. Water losses from the leaf RWC decreased from 96% to 70%.

Fig. 5.15. Light and scanning electron microscope cross-sections of a mature pineapple leaf showing the
prominent water-storage tissue on the adaxial side of the leaf and the chlorenchyma on the abaxial half of
the leaf (photos of D. Bartholomew).
 Crop Environment, Plant Growth and Physiology 95

Fig. 5.16. Leaf water-storage tissue deficiency (%) at inflorescence emergence (red bud) with decreasing
availability of water (modified from Nightingale, 1949).

However, leaf RWC was concluded to be a Kc can vary significantly over the plant cycle.
rather insensitive indicator of water stress in Kc decreased significantly from 0.93 5
pineapple, especially under minimal stress months after planting (50–60% of soil cover)
situations (Kadzimin, 1975; George et al., to 0.41 by 11 months after planting, at which
1984). time there was 80–90% of soil cover by the
Pineapple plants subjected to water stress canopy (N’Guessan, 1985). Where pineapple
have significantly lower ET values compared was planted through a polyethylene mulch
with unstressed plants. In central Côte in Hawaii, a 50% decrease in the daily rate of
d’Ivoire, ET of complete pineapple leaf consumptive use of water occurred by the
canopies averaged 3.8 mm day−1 when irri- time of 60% canopy closure (Ekern, 1964). It
gated and 2.9 mm day−1 when unirrigated is a striking feature of pineapple that the ET
(Combres and Perrier, 1977). The ET/ETg rate of a pineapple crop decreases as the
ratio, where ETg is the potential ET of a grass plant develops and the canopy cover
cover, is about 0.45 for an irrigated pineapple increases, whether in mulched or unmulched
crop but the ratio decreases during drought conditions (Ekern, 1964). With mulch, 15% of
(Combres, 1983). the incident energy was used for ET after the
ET of a pineapple crop depends both on canopy closed. The very low water-use rate
the stage of development and climatic condi- of pineapple, which is due to the inverted
tions. When well watered, the ET of com- pattern of stomatal conductance and xero-
plete canopies ranged from 0.6 to 0.7 of the morphic anatomical features, is further
potential evaporation of a grass cover reduced by water stress.
(Combres and Perrier, 1977). The crop para-
meter Kc, in the equation: FLOODING. Excess water can reduce the
growth and yield of pineapple, mainly when
ETR = Kc × ETo (9)
waterlogging occurs during root initiation
where ETR is actual ET and ETo is standard and at fruit filling. As with most crop plants,
evaporation (Priestley and Taylor, 1972), gen- root growth and efficiency are restricted by
erally ranges between 0.8 and 1.3 for most inadequate aeration associated with excess
crops. An average value of 0.74 was soil moisture. In solution cultures, root
observed for pineapple (Combres, 1983), but growth was increased by aerating the
96 E. Malézieux et al.

solution (Iwaoka et al., 1935) or by removing absence of frost, the plant is quite productive
roots from the solution for 2 h day−1 (Tisseau, in both cool subtropical and tropical environ-
1971). The formation and persistence of root ments, though the crop cycle is prolonged in
hairs seem to depend upon an oxygen sup- such environments.
ply to the roots (Py et al., 1987). Root Large areas are planted to pineapple in
aerenchyma is also increased by a dimin- hot, wet intertropical regions, in low-altitude
ished oxygen supply and a comparable effect areas and, more specifically, along coastal
is seen with an increase in soil compaction plains, where the climate is moister and hot-
(Rafaillac et al., 1978). Excessive water pro- ter than in continental areas. In most of the
duces a leaf colour change similar to that tropics, and especially in hot and wet regions
seen with water stress, in that leaves first close to the equator, the annual range of
become pale yellow and then red, and leaves monthly temperature is very small, some-
are reduced in length and are more erect (Py times not more than 1°C. Thus, the diurnal
et al., 1987). temperature cycle is often more important
than the seasonal cycle (Monteith, 1977).
Optimum day and night temperatures for
General effects
vegetative growth of pineapple are near 30
TEMPERATURE. Although there are few specific and 20°C, respectively, with an optimum
data on the effects of temperature on the mean temperature of 23–24°C (Neild and
growth and development of pineapple, tem- Boshell, 1976). Plant growth decreases
perature appears to be one of the most rapidly at mean temperatures below 15°C or
important environmental factors determin- above 32°C (Neild and Boshell, 1976). In
ing pineapple distribution and productivity Hawaii, the slower vegetative growth rates
in the world. Pineapple survives in hot, dry observed from December to April are mainly
environments where other crops would be due to a decrease in the average tempera-
non-productive, but is also cultivated in the ture, especially soil temperature (Ekern,
cool subtropics, where freezing temperatures 1967). In Australia, low temperatures from
may occur (Table 5.1). The lowest annual May to October reduce or even stop plant
average temperature where pineapple is growth in the midwinter period (Glennie,
grown on a commercial scale appears to be 1981; Wassman, 1986).
17.2°C in Port Elizabeth, South Africa The large environmental variation among
(Bartholomew and Kadzimin, 1977). In areas where the crop is grown (Table 5.1)
southern Queensland the mean monthly accounts for much of the large variation in
maximum/minimum temperature ranges the time from planting to maturation of the
from 29/19°C in summer to 20.5/6°C in win- mother plant crop. Within a given environ-
ter (Wassman, 1990). The plant does not tol- ment, fruit weight at harvest is determined
erate frost, but temperatures have been in large part by plant weight at forcing (Py
reported to drop below 0°C for short periods and Lossois, 1962; Gaillard, 1969; Tan and
of time in the pineapple-growing areas of Wee, 1973; Malézieux, 1988; Malézieux and
south Queensland, Australia (Swete Kelly Sébillotte, 1990a). Consequently, an impor-
and Bartholomew, 1993), South Africa, São tant objective of growers is to obtain a given
Paulo, Brazil (Giacomelli and Py, 1982; Py et plant weight at the time plants are to be
al., 1987), and southern Florida. As was forced. In the absence of other stresses, plant
noted before, prolonged exposures to tem- growth is determined by temperature. For
peratures less than 0°C can destroy the that reason, the interval from planting to
canopy and lead to the loss of the crop, as forcing varies considerably over the wide
has happened in some areas where freezing range of latitudes and altitudes where
temperatures can occur (Giacomelli and Py, pineapple is grown (Table 5.1). In the East
1982; Swete Kelly and Bartholomew, 1993). London area (33°S, South Africa), where
Overhead irrigation has been used to protect average temperature is 18.8°C (mean mini-
against freezes in Florida (J. Tenbruggencate, mum temperature of 14.7°C, mean maxi-
1986, personal communication). In the mum temperature of 22.8°C), the average
 Crop Environment, Plant Growth and Physiology 97

period from planting to forcing is 24 months TEMPERATURE INJURY. High temperatures are
(Bouffin, 1991), whereas in areas near the generally of minor concern during vegeta-
equator, such as in West Africa, the vegeta- tive growth of pineapple. Although leaf and
tive growth period is only 6–8 months. plant temperature may reach very high val-
The planting-to-forcing interval also varies ues due to the low transpiration rate and
with date of planting, especially in areas poor air circulation around the leaves within
where the seasonal temperature variation is the plant canopy, leaves tolerate high tem-
large. In the Canary Islands, the vegetative peratures well. Leaf sunburn is sometimes
cycle ranges from 9 months when planting is seen, but it is not a serious problem in any
done in the spring and plants develop in sum- region where pineapple is grown.
mer, to 14 months when the crop is planted in As noted above, low-temperature injury
winter (Galan Sauco et al., 1988b); similar is of greater concern where freezing temper-
large variations are encountered in Australia atures are encountered. With mild radiation
and South Africa. A plant fresh mass of 2.5 kg frost, the upper surface of horizontal leaves
– a common plant weight for forcing – can be develops a red/white-flecked, scorched
reached within 8 months after planting in appearance. If the injury is severe, the whole
Côte d’Ivoire (5°N, mean temperature of leaf dies and the leaves become papery.
26.6°C), within 10–11 months in Hawaii Depending upon the age of the plant when
(21°N, mean temperature of 23°C) and within frost injury occurs, vegetative plants may
13–14 months in Queensland, Australia (26°S, recover and produce an acceptable crop
mean temperature of 19°C). As a result, the (Swete Kelly and Bartholomew, 1993).
time from planting to harvest ranges from 12 Without some sort of frost protection, pro-
months near the equator (Gaillard, 1969), longed exposure to temperatures less than
where the average annual temperature is 0°C can destroy the aerial parts of the plant,
26–27°C, to 32 months in Swaziland (26° 30’S), leading to loss of the crop. As noted above,
where mean annual temperature is 16–17°C sprinkler irrigation has been used to protect
(Dodson, 1968). While there is wide variabil- pineapple plants from freezing in Florida (J.
ity in the length of the vegetative growth Tenbruggencate, 1986, personal communica-
phase for ‘Smooth Cayenne’ in the pineapple- tion).
growing areas of the world, there is also con-
siderable variation in the length of the IRRADIANCE. The significance of irradiance as
reproductive phase (Table 5.1; see a factor affecting pineapple yield was recog-
Bartholomew et al., Chapter 8, this volume). nized at least by the mid-1930s in Hawaii
In some pineapple-growing regions, it is (Sideris et al., 1936). However, there is little
common to produce one or two ratoon crops evidence that irradiance limits pineapple
from suckers borne on the mother plant. In production in most areas where the crop is
equatorial climates, sucker development is grown. Sanford (1962) attributed to Sideris
commonly delayed until after the fruit an observation that yield decreases about
matures. The first ratoon crop takes about 1 10% for each 20% decrease in solar radiation.
year in both Hawaii and Thailand, even Shiroma (1977) concluded that irradiance
though growth is more rapid in the warm, could be limiting for ‘Smooth Cayenne’
humid climate of Thailand. The long period pineapple in Okinawa, but his conclusion
of time required for ratoon-crop develop- was based on lower average global radiation
ment in Thailand results from the fact that values for Okinawa than for Taichung,
suckers do not begin to develop until after Taiwan and Hawaii, rather than data
the plant-crop fruit is harvested, whereas, in obtained in situ. A linear relationship
Hawaii, development begins at the time of between plant weight 8 months after plant-
forcing. In general, the plant and one ratoon ing and radiation intercepted by the canopy
crop are harvested after about 2 years in was established in an experiment in Côte
Thailand, 3 years in Hawaii, 3–4 years in d’Ivoire, where plants were planted every
Australia (Vuillaume, 1986) and as many as 5 month for 6 years (E. Malézieux, 1992,
years in Swaziland (Dodson, 1968). unpublished results). In this experiment, an
98 E. Malézieux et al.

empirical relationship that associated aver- tively stable and irradiance fluctuates signifi-
age irradiance during the entire crop cycle cantly throughout the year, potential plant
and an indicator of drought from planting to weight at forcing was correlated with irradi-
harvest explained 64% of the variation in ance intercepted during vegetative growth
plant weight at harvest (72 monthly planti- (Malézieux, 1988). In these unirrigated con-
ngs with similar cultural practices). ditions, yield was depressed when drought
However, interpretation of the results of was prolonged.
such studies is very difficult because of the
long-term nature of the crop and the fact that DROUGHT AND WATER EXCESS. Because pine-
changes in irradiance are confounded with apple has a great capacity to survive drought
simultaneous changes in air temperature. In (Sideris and Krauss, 1928), it has been grown
areas where the variation in temperature is successfully on Molokai, Hawaii, in an envi-
small compared with variation in irradiance, ronment with less than 600 mm year−1 of
such as near the equator, irradiance may be rain (Noffsinger, 1961). Where the soil is
of importance in determining plant growth. well drained, pineapple has also been culti-
Where both irradiance and temperature vated in relatively wet areas, such as
change significantly with season, it is very Guadeloupe, where rain exceeds 3500 mm
difficult to evaluate the effects of irradiance year−1 (Py et al., 1968). Hence, depending on
on pineapple growth and productivity. The the location and season, the climatic water
quality and average size of the available balance over the crop cycle or some part of it
planting material can vary with time of year can be largely positive or negative. In
(Louis and Nightingale, 1937; Zhang, 1992) Wahiawa, Hawaii, an almost ideal location
and these factors are important because they for pineapple culture without irrigation,
determine the initial plant size. At a given pan evaporation averages 1850 mm year−1
planting density in the same environment, while rainfall averages 1000 mm year−1
plant mass at a given time after planting is (Noffsinger, 1961; Ekern, 1965).
determined by the mass of the propagule (Py Despite the capacity to survive drought,
et al., 1987). Even modest differences in growth and yield are reduced in all areas
propagule mass between plantings in differ- where water stress occurs (Foote, 1955;
ent seasons can obscure the effects of sea- Black, 1962; Ekern, 1964; Py, 1965; Huang
sonal changes. After a 2–3 month and Lee, 1969; Malézieux, 1988). Hence, irri-
establishment period, which can be further gation or polyethylene mulch or both are
prolonged by cool temperatures or lack of regularly used (Hawaii) or are used by some
rain, vegetative growth is the product of growers (Queensland, Côte d’Ivoire).
accumulated irradiance over a period of 6–15 Though polyethylene mulch has several ben-
months. Fruit development requires an addi- efits, its impact on water economy is one of
tional 5–8 months. Since plant mass at forc- the main reasons it is used in Côte d’Ivoire
ing is an important determinate of fruit mass (Combres, 1983). Several studies report no
at harvest (Py and Lossois, 1962; Gaillard, effect of irrigation on pineapple growth,
1969; Tan and Wee, 1973; Malézieux, 1988; even where drought occurs (Rao et al., 1974;
Malézieux and Sébillotte, 1990b), anything Tay, 1974; Senanayake, 1978; Htun, 1986).
that retards vegetative growth in one season Despite such results, the benefits of irrigation
relative to another will either delay harvest during prolonged drought (Medcalf, 1950;
or reduce yield. Py, 1965; Huang and Lee, 1969; Combres,
Seasonal variations in productivity can 1979, 1983; Kuruvilla et al., 1988; Malézieux,
also be due to the seasonal distribution of 1992) are sufficient to cause it to be a widely
rainfall and seasonal and day-to-day varia- adopted practice. In Guinea, growth of unir-
tions in forcing success (Wee and Ng, 1968; rigated plants was delayed by up to 3.5
Wassman, 1991) and in pest and disease months by the end of a 5-month dry season
pressure. While it is difficult to demonstrate (Py, 1965). Despite good evidence of the ben-
an effect of irradiance on productivity, in efit of irrigation, no quantitative studies
Côte d’Ivoire, where temperature is rela- were found that would provide the data
 Crop Environment, Plant Growth and Physiology 99

needed to predict growth or yield reduction the physical damage provides points of entry
in response to water stress. for fungi (see Rohrbach and Johnson,
In regions of high rainfall, water excess can Chapter 9, this volume), but neither the
also limit growth, as well as increasing suscep- physical nor the fungal damage is consid-
tibility to disease (see Rohrbach and Johnson, ered economically significant in most
Chapter 9, this volume). Waterlogging instances (Py et al., 1987). Although wind
impedes gas exchange, resulting in elevated damage is generally minor, wind-breaks of
CO2 and depleted O2 levels in the soil, which napier grass or sugar cane have been used in
lead to decreased growth by inducing root some regions of South Africa (Anon., 1956).
anoxia. In the Mekong delta, for instance, beds In coastal areas, wind-borne salt spray can
may become submerged (Le Van Thuong, cause burns, leading to blackish spots near
1991), and pineapple plants died after 15 days the tips of the leaves (Sideris, 1955).
of submersion of the bed beneath 10–15 cm of Exceptionally strong winds caused by hurri-
water. Where rainfall is high, as in the humid canes can severely damage all parts of the
tropics, soils used for pineapple cultivation are plant or uproot it, leading to significant
generally lower in clay and higher in sand to reductions in yield (Py et al., 1987).
ensure good drainage. Such soils have low Hail is rare is most areas where pineapple
water-holding capacity and are susceptible to is grown but occurs in southern Queensland
erosion (Ducreux et al., 1980; Ciesiolka et al., (Swete Kelly and Bartholomew, 1993) and
1993; El-Swaify et al., 1993; Ciesiolka, 1994). southern Africa (Anon., 1956). Hail can seri-
Ridging practices, widely used in South and ously damage the rigid and brittle pineapple
West Africa and some areas of Australia, also leaf, significantly reducing leaf area. After a
decrease the risk of waterlogging during rainy severe hailstorm in Swaziland, most of the
periods. leaves wilted and died off, leaving only a
Plant growth and fruit weight are small growing point and leading to a loss of
decreased by waterlogging that occurs in 1 year’s growth (Anon., 1956).
small field depressions in Côte d’Ivoire. Few data on the effects of salt on pineap-
While few quantitative data on the effects of ple were found (see Malézieux and
water excess on growth under field condi- Bartholomew, Chapter 7, this volume), but
tions were found, until relatively recently, pineapple is grown along windward sea
when effective fungicides became available coasts in Hawaii with minimal adverse
to control Phytophthora spp., loss of plants to effects, even though the north-easterly trade
root and plant rots was a greater concern winds can be both persistent and strong.
than reduced growth due to waterlogging
(Rohrbach and Apt, 1986; see Rohrbach and
Johnson, Chapter 9, this volume). Conclusions

MISCELLANEOUS FACTORS. Wind is a minor con- Potential crop dry-matter production

cern for pineapple growers, though pro- depends on the various interacting effects of
longed winds have been reported to reduce the physical environment. The complex
plant size by 25% (Nightingale, 1942). It is physiological responses of this CAM plant to
not known if the reduction was due to the several simultaneously varying weather fac-
physical effect of wind, to an altered water tors (Fig. 5.17) make it difficult to link
balance or to reduced temperature within weather through its effects on physiological
the canopy. Though wind has little effect on processes to dry-matter accumulation, dry-
ET when the plant cover is complete, hot, matter partitioning or yield. Even where
dry winds, such as the Norte in Mexico or controlled-environment studies have in-
the Harmattan in Côte d’Ivoire, can lead to creased our understanding of physiological
leaf-tip drying (Py et al., 1987). The influence responses to one factor, it is difficult to
of wind on thermal exchanges in the canopy extrapolate such results to the field. These
is significant (Py et al., 1987). Strong winds considerations apply to many crops but are a
cause leaves to rub against each other and particular problem for pineapple because of
 100

Temperature Moderating factors

Latitude Commercial requirements
Irradiance
Elevation Grower’s decisions
Daily thermoperiod Techniques (houses, mulches)
Water supply Climate, geography
Propagule mass
Quantity of CO2 assimilated
Proportion of CO2 fixed by CAM Plant growth rate
Planting to forcing
interval
Plant mass at forcing
Transpiration
Stomatal opening pattern Consumptive use of water

Plant morphology Fruit weight

Specific leaf area
Leaf weight ratio
Dry-matter content

Susceptibility to forcing
Forcing success
Natural floral induction
E. Malézieux et al.

Fruit development rate Forcing to harvest interval

Yield
Stem starch accumulation

Sucker development Cycle duration

Sucker growth Possibility of one or more ratoon crops
Sucker position Planting to first ratoon harvest interval

Total soluble solids Skin colour Multiple crowns

Titratable acidity Flesh colour Collar of slips
Aroma Internal browning Fungal diseases Fruit quality
Green ripe fruit Storage life

Fig. 5.17. Flow diagram summarizing the effects of environmental factors, but mainly temperature, on pineapple growth and development.
 Crop Environment, Plant Growth and Physiology 101

the complex interactions between weather There is much opportunity for basic research
and CAM in carbon assimilation, water on pineapple to characterize the effects of
economy and growth processes. The effect of irradiance and temperature on carbon assimi-
temperature on leaf photosynthesis is rea- lation, dry-matter partitioning, reproductive
sonably well understood, but it is still haz- physiology, yield and fruit quality. Only
ardous to extrapolate this knowledge to the when some of these important issues are
effect of temperature on crop growth. Leaf resolved will it be possible to understand the
photosynthesis is apparently saturated at complex interactions between weather and
low irradiance, but crop productivity never- physiology. Although prediction of pineapple
theless seems to respond to total irradiance, growth and development as a response to the
as well as to the fraction intercepted by the main environmental factors is now possible
canopy. This is due in part to the high LAIs in some conditions (Zhang et al., 1997), many
sustained at high planting densities. Also, factors affect the accuracy of such predic-
although pineapple has one of the highest tions. A comprehensive understanding of the
WUEs among cultivated crops and can sur- effects of environmental variables on pineap-
vive severe drought, the reduction in crop ple growth and yield is essential if the accu-
growth due to water stress makes irrigation racy of growth models is to reach the point
profitable in many areas. where it will be possible to satisfactorily pre-
We show here that, despite a significant dict harvest date and yield. When sufficient
amount of research on pineapple, it is still data are collected, it may be possible to use
difficult to understand and simulate the growth models to make crop-loss assess-
effects of weather on yield by linking ments for the several important pest and dis-
weather variables to physiological processes. ease problems that reduce crop productivity.

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