Block 1 Comparative Anatomy of Vertebrates-I

UNIT 5

CIRCULATORY SYSTEM

Structure
5.1 Introduction 5.4 Venous System
Objectives Fishes

5.2 Heart Amphibians

Protochordate Heart Reptiles

Early Vertebrate Heart Birds

Early Tetrapod Heart Mammals

Late Ectotherm Heart 5.5 Blood

Endotherm Heart Composition of Blood

5.3 Arterial System Respiratory Pigment

Aortic Arches-Overview 5.6 Lymphatic System

Fishes Fishes

Amphibians Amphibians

Reptiles Reptiles

Birds Birds

Mammals Mammals

Evolution of Aortic Arches Other Lymphatic Organs

Types of Arteries 5.7 Summary

5.8 Terminal Questions
5.9 Answers

5.1 INTRODUCTION
In the previous unit on Respiratory system you learnt about the various
respiratory surfaces and organs that are involved in gas exchange in aquatic
and terrestrial vertebrates. In this unit you will learn that the circulatory system
of vertebrates consists of a heart, arteries, veins, capillaries and blood (the
138 blood vascular system) and of lymph channels and lymph (lymphatic system).
Unit 5 Circulatory System
Blood carries oxygen collected in respiratory organs, nutrients from extra
embryonic membranes of embryos and from the adult digestive tract. It also
carries hormones from endocrine tissues, substances associated with
maintaining, immunity and disease. While flowing, it removes waste products
of metabolism from the excretory organs. Lymph channels collect interstitial
tissue fluids not taken up by the blood stream and emulsified fats absorbed in
the small intestine. These lymph channels terminate in one or more of the
large venous channels of the blood vascular system. Pressure differences that
drive this flow of blood through the circulatory system are created by the
pumping action of the heart.

Arteries carry blood away from heart. They have muscular elastic and fibrous
walls capable of swelling with each intrusion of blood and of active constriction
and dilation in response to nerve impulses. Arteries, thereby assist in
regulating blood pressure. They terminate in capillary beds. Veins have
proportionately less muscle and elastic tissue and more fibrous tissue and are
therefore, capable of less dilation or constriction. They carry blood towards
heart from capillary beds. Capillaries consist of endothelium only, with a lumen
just large enough to accommodate red blood cells. In fact, the red cells must
squeeze through and in so doing, become deformed. In vertebrates respiring
by gills, blood is pumped from the heart to the gills, where external respiration
takes place. From the gills it typically flows via arteries to capillaries
throughout the body. Thus we see that the respiratory and circulatory systems
work in tandem.

We begin by discussing the evolution of the vertebrate heart and aortic arches
and see how the vertebrate circulatory system exhibits diverse adaptations
across the radiation of vertebrates culminating in the highly specialized
mammalian heart. Along with changes seen in the structure of the heart you
can see the evolution of double circulation and its importance. You will realize
that the structure and function of the circulatory system of each group of
vertebrates has a vital role that benefits that particular group. We then go on to
discuss the composition of blood and describe the lymphatic system.

Objectives
After studying this unit you should be able to:

 trace origin and evolution of the heart of vertebrates,

 trace origin and evolution of aortic arches,

 describe arterial and venous systems of vertebrates,

 describe composition of blood,

 describe lymphatic system, and

 explain the phenomenon of double circulation.

5.2 HEART
139
Block 1 Comparative Anatomy of Vertebrates-I
The vertebrate heart is really a modified blood vessel with highly muscular
walls. There are valves to prevent backflow of blood. The heart consists of
three layers:

1) Endocardium - inner lining (endothelium and elastic tissue)

2) Myocardium - between endocardium and epicardium

(muscular layer)

3) Epicardium - outer fibrous tunica, covered by visceral

pericardium (subdivision of coelom).

Heart pulsates as a result of response of the muscle cells to electrolytes that

infuse it. Evolutionary changes of the heart lead from a primitive, simple,
straight tube in the protochordates through the sinuous multi chambered organ
seen in fishes and the partially subdivided, but otherwise simple structure of
early tetrapods, to the compact, highly efficient structure of birds and
mammals. The evolution is summarized as follows:

i) the protochordate stage with alimentary pharynx,

ii) the piscine stage with branchial pharynx,

iii) the early tetrapod stage with primitive lungs,

iv) the late tetrapod stage of higher ectotherms and

v) the stage of endothermic tetrapods.

As implied by the names designating these stages, evolution of heart reflects

changes in other structures, particularly respiratory organs.

5.2.1 Protochordate Heart

Protochordates (tunicates and Amphioxus) possess a simple, tubular heart
that beats by peristaltic waves. Since arteries are more muscularised than
veins, they may be taken as the primitive heart. Therefore, extensive
muscularization of arteries has its functional significance in forcing blood into
the smaller vessels (capillaries) from which blood may return back without
special means of propulsion since it is passing from smaller into larger
vessels.

The protochordate stage does not need, a more specialized heart, this can be
explained on the basis of lack of proper respiratory function of the pharyngeal
clefts. The pharyngeal clefts in protochordates primarily serve as a sieve to aid
in capturing the food.

5.2.2 Early Vertebrate Heart

When pharynx is provided with capillary network, then in order to facilitate the
gaseous exchange, a more efficient and specialized pump is required to force
blood through to the dorsal aorta. The most primitive vertebrate heart is seen
140 in agnathans. They have a circulatory system consisting of a main systemic
Unit 5 Circulatory System
heart and 3 accessory hearts (Fig.5.1). In the figure you can see the general
arrangement of the circulatory system, the portal heart pumps blood from the
intestines to the liver, the cardinal heart pumps blood from the head to the
body, caudal heart pumps blood from the trunk and kidneys to the rest of the
body. The agnathans have a different sinus venosus that is attached to the left
side of the atrium. The circulatory system of hagfish has not changed in
millions of years and since they are bottom feeders and dwellers they have
adapted to a low arterial blood pressure and low cardiac output is sufficient.

Fig. 5.1: Simplified view of the relationship between the main ‘systemic (or
brachial)’ heart and the three accessory hearts of the hagfish (adapted
from Jensen, 1965 and Jorgensen et al. 1998).

Thus a true heart is seen for the first time in fishes; a heart consisting of a
series of specialized chambers to which the function of forcible propulsion of
blood in arterial channels is restricted. The heart of fishes is known as the
branchial heart as its main function is to provide deoxygenated blood through
the ventral aorta into the gills. Basically there are two main chambers : the
atrium and the ventricle; two other chambers the sinus venosus (SA) and
conus arteriosus (CA) in lamprey, elasmobranchs and holosteans (Fig.5.2 &
5.3a). The same chamber in teleosts is known as bulbous arteriosus which is
elastic and continues to form the ventral aorta (Fig.5.3b).

Actually two operations must be performed to carry out the cardiac function
efficiently in a gill-respiring vertebrate. The blood must be first collected and
then pushed along. Collecting chambers thus appear at the rear and
propulsive chambers at the front. Fishes are provided with a posterior sinus
venosus, followed by an atrium, then a ventricle, and finally a conus arteriosus
in the anterior arranged in series (Fig. 5.2). 141
Block 1 Comparative Anatomy of Vertebrates-I
The sequence of beat is the peristaltic sequence; from the rear towards the
front so the blood flows from posterior chambers to anterior chambers.

The sinus venosus is little more than an expansion of the junction of the
primary somatic (common cardinal) and visceral (hepatic) veins. The walls are
very thin and contain the pacemaker cells that initiate contraction. A slight
negative pressure actually exists in this chamber, resulting from the integration
of atrial contraction and action of the sinoatrial valve. The venous blood from
the body fills the sinus venosus and through the sinoatrial valve blood passes
with minimum cardiac assistance into the atrium. Because the S-shaped
arrangement of the chambers in the fish heart (Fig. 5.2) places the thin-walled
sinus venosus and atrium dorsal to the ventricle, the blood is forced by gravity
and atrial contraction through the atrioventricular valve into the ventricle. The
sinoatrial and atrioventricular valves prevent the back flow of the blood. The
thickened walls of ventricle with cardiac muscle provide energy to move blood
into the ventral aorta and gills.

Once received in the ventricle, blood passes through one or more series of
semilunar valves (conal valves) in reaching the fairly rigid conus arteriosus,
which adds still further push to the blood and smooths its flow on its way to the
ventral aorta. The entrance into the latter is guarded by several rows of other
semilunar valves. The semilunar valves are folds in the wall which prevent the
back flow of blood and possibly help in distributing blood to the aortic arches
(Fig. 5.2 & Fig. 5.3a).

Fig. 5.2: Lamprey heart showing the chambers characteristic of most fishes.

In teleosts the blood flows from the ventricle into the bulbous arteriosus which
is a unique structure, it is thin walled and elastic and expands each time the
blood is pumped from the ventricle and connects to the ventral aorta. The
blood pressure for this is generated in the ventricle which is much higher than
what is required in the thin walled gills so the bulbous arteriosus regulates the
pressure as it enters the gills. In fact the teleosts synchronize the opercular
movement with the pulsation of the heart so that a steady flow of blood is
maintained.

142
Unit 5 Circulatory System

(a)

(b)

Fig. 5.3: Fish hearts, (a) Shark (b) Teleost. Blood leaves the shark heart through
the muscular conus arteriosus, a chamber that is absent in teleost
fishes. In the teleost heart, the base of the ventral aorta is swollen,
creating the elastic, non contractile bulbus arteriosus. A single pair of
bulbus valves prevent the back flow of blood into the ventricle.

5.2.3 Early Tetrapod Heart

This stage was initiated by the development of air bladder in pretetrapod

ancestors for the perfection of aerial respiratory function in tetrapods.
Gradually, the air bladder became specialised to serve the role in external
respiration as seen in the present day lung fishes, the sarcopterygians.
Lungfishes primarily use their gills for respiration but when they are outside
water they adopt a vascularized lung supplied by two pulmonary arteries to
assist in breathing air. The aerated blood does not supply the body directly
but returns to the heart via a single pulmonary vein which opens on the left
side of the atrium from where it is pumped into systemic circulation. The blood
returning after circulation in the body comes back via the sinus venosus which 143
Block 1 Comparative Anatomy of Vertebrates-I
has shifted to the right side of the heart. This modification forms the basis for
dual circulation first seen in vertebrates which chiefly involved changes in the
heart. First the atrium was subdivided into two chambers; a systemic chamber
(right), receiving non-aerated blood from the sinus venosus, and a pulmonary
chamber (left), receiving aerated blood from the pulmonary veins (Fig. 5.4).

Fig. 5.4: The early tetrapod heart, ventral view with parts drawn to side instead
of superimposed as in life.

A second related alteration involved a partial separation of two chambers in

the conus arteriosus, serving apparently to direct the aerated blood (from the
left side) chiefly into the anterior vessels of the ventral aorta, whereas the non-
aerated blood (from the right side) was directed chiefly into the posterior pair
of vessels bearing the pulmonary arteries. Thus we can recognize a
pulmonary and a systemic chamber in the conus arteriosus. In amphibians
that have functional lungs; the heart consists of 3 chambers:, atrium
completely divided into right and left atria, a ventricle and a sinus venosus and
conus arteriosus which is partially divided with the help of a spiral valve. In
frogs, whose cardiovascular system is best studied, the conus arteriosus
arises from the single trabeculate ventricle. Trabeculae are the projecting
cones of the muscles that arise from the inner wall of myocardium of the
ventricle creating deep recesses or compartments in the wall. As shown in Fig.
5.5 semilunar valves lie at the base of conus arteriosus and prevent the
backflow of the blood into the ventricle. A spiral valve twisting almost through a
complete rotation establishes chambers within the conus arteriosus that target
blood to the systemic and pulmocutaneous arches, both of which arise from
truncus arteriosus, a remnant of ventral aorta. The sources of oxygenated
blood vary in amphibians; as for respiration they depend on the skin, on gills,
on lungs or on all the three modes. It is for this reason the heart structure
varies in different amphibians.

In lungless salamanders or the ones with reduced lung function the septum
dividing the atrium as well as the spiral valve may be much reduced or entirely
absent. In amphibians such as Necturus where gills dominate lungs as
respiratory organs, the interatrial septum is reduced or perforated. The atrium
is completely divided in modern anurans and in all living reptiles, birds and
144 mammals.
Unit 5 Circulatory System

Fig. 5.5: Structure of the heart of a bull frog.

Another alteration in amphibian heart is the reduction of sinus venosus both in

size and importance as a blood-gathering chamber. A large thin-walled heart
chamber (sinus venosus), into which the blood can flow against an absolute
minimum of resistance is essential for fishes. In land-dwelling vertebrates,
subjected to the markedly lesser atmospheric pressure, such an elaborate
collecting device is not essential; in later stages it is discarded completely.

In amphibians the deoxygenated and oxygenated blood streams returning

from systemic and pulmonary circuits are kept separate as they pass through
the heart. The deoxygenated blood is selectively directed to the lung via
pulmonary artery and the oxygenated blood is directed to the systemic tissue
via aortic arches. In frogs during the time of air breathing, the trabeculae in the
ventricle separate the two different streams of blood in the heart. It is thought
that as one stream enters the ventricle, it fills the compartments between the
trabeculae, and then the second stream occupies the centre of the ventricle.
Because of their different positions, the oxygenated and deoxygenated
streams depart by different exits to reach appropriate set of arteries.
Whenever a frog dives in the water, the lung collapses from the water
pressure on the body wall. The blood flow is reduced in the lungs and
increased in the skin. Thus, the loss of pulmonary respiration is somewhat
compensated by increased cutaneous respiration in submerged frogs. Before
we proceed further, try the following SAQ.

SAQ 1
Fill in the blanks with appropriate words.
i) The tubular heart of protochordates beats by ………………. .
ii) In the early tetrapod heart the …………… is divided into two chambers.
iii) Ventricle is ………………. in frogs.
iv) In Necturus the …………… ……….……… is reduced or perforated.
v) In amphibians during diving the ……………… respiration is increased.
145
Block 1 Comparative Anatomy of Vertebrates-I
5.2.4 Late Ectotherm Heart
Living reptiles varied in cardiac structure though they exhibit uniformly a
number of improvements over the early tetrapod heart (Fig.5.6). Since most
reptiles are adapted to a fully terrestrial environment and have more active
lifestyle, their cardiovascular system supports the accompanying higher
metabolic rates and elevated levels of oxygen and carbon-dioxide transport. It
is also capable of generating higher cardiac out-put, elevated blood pressures
and more efficient separation of oxygenated and deoxygenated blood streams
as compared to amphibians. Basically two types of reptilian hearts (one in
chelonians and squamates and other in crocodiles) have been recognised.

Fig. 5.6: Late ectothermic heart, ventral view with parts drawn to side instead of
superimposed as in actual structure.

In chelonian/squamate heart, the sinus venosus is reduced in size as

compared to that in amphibians and in advanced types ancestral to mammals,
it has disappeared completely as a chamber, although it is present in all living
reptiles. From the very first vertebrates, however, the sinus venosus served a
function not only as a collecting chamber but also as a site of origin of
heartbeat. Although its identity as a chamber may have been lost in at least
some reptiles and certainly in birds and mammals, the function it formerly
performed for initiation of heart beat still remains. The excitatory tissue
remained embedded in the wall of the right atrium near the point of the entry of
veins, that now with the loss of the sinus as a chamber, empty directly into the
atrium. This myogenic center is the sinoatrial node, which serves in all
amniotes lacking sinus venosus as the originator of each heartbeat. The conus
arteriosus also disappeared, though it appears during early embryonic
development, in adults it was subdivided to form the bases (trunks) of the
three large arteries leaving the ventricle: the pulmonary trunk and the right and
left aortic trunks (systemic trunks). This pairing of systemic trunk is seen in
many reptiles, the right trunk connected with the left side of the heart, the left
with the right side. Despite the loss of the role of the conus arteriosus as a
cardiac chamber in reptiles, the semilunar valves that conus formerly
possessed remained intact and unchanged. These persist at the bases of the
pulmonary and systemic trunks in all amniotes, but are reduced into three
valves in each vessel. Atrium is divided into right and left atria. Entry to the
146 ventricle is guarded by the atrioventricular valves. Ventricle is partially divided
Unit 5 Circulatory System
internally resulting in a rather effective separation of venous and arterial blood.
As you can see in Fig. 5.7, it has three interconnected compartments: cavum
venosum, cavum pulmonale, cavum arteriosum. Cavum venosum and cavum
pulmonale are separated from each other by a muscular ridge and cavum
arteriosum is connected to the cavum venosum through an interventricular
canal.

(a)

(b)

Fig. 5.7: Ventral view of the lizard heart. (a) Small part of the ventral wall has
been removed. The arrow shows the direction of blood flow from
cavum arteriosum to cavum venosum through interventricular canal.
The blood goes to systemic arches from cavum venosum. (b) Structure
of the heart after some more part of the ventral wall has been removed. 147
Block 1 Comparative Anatomy of Vertebrates-I
As you can see in Fig. 5.8, the pattern of blood flow through the hearts of
chelonian and squamates varies depending on whether they breathe air or
hold their breath (a condition called apnea). For example, when turtle is air
breathing on the land most of the deoxygenated blood returning from systemic
tissues is directed to the lungs and most of the oxygenated blood from the
lungs is directed to the systemic tissues via the aortic trunk. When the turtle
Apnea, a condition of
dives in the water, the heart responds with a right to left cardiac shunt. Blood
holding breathe flowing in the cavum venosum is directed to the aortae rather than the
occurs not only during pulmonary circuit. This shunting is controlled by the differences in the
diving. Most reptiles,
resistance of systemic and pulmonary circuits.
while resting on land
go for long intervals
without taking a
breath. As apnea
continues, oxygen
from the lungs
becomes depleted
and pulmonary
perfusion declines
until just before
another breath.

Fig. 5.8: The path of blood flow in the squamate heart. (a) In the squamates
breathing air, venous blood from the right atrium enters the cavum
venosum, crosses the muscular ridge and fills the cavum pulmonale.
When ventricle contracts, most of this blood flows in the pulmonary
artery. Simultaneously, blood from the left atrium enters the deep
cavum arteriosum. Upon contraction of the ventricle this blood passes
through the interventricular canal to the left and right aortic arches. (b)
On diving, due to the resistance to the pulmonary blood flow, the blood
moves across the muscular ridge and departs mainly through the left
148 aortic arch.
Unit 5 Circulatory System
The heart of crocodiles shows structural variations in certain aspects. The
ventricle is completely divided into right and left chambers by a complete
interventricular septum. The pulmonary trunk and the left aortic arch opens off
the right ventricle and the right aortic arch opens off the left ventricle.

Curiously, the left arch does not receive blood directly from its own (right)
ventricle, since the semilunar valves actually prevent flow from ventricle to
aorta except under unusual stress situation. The left arch receives blood,
through the foramen of Panizza that connects the right and left arches where
they cross a short distance from the heart (Fig. 5.9). You can see in Fig. 5.10
the comparison of the path of blood flow in the hearts of chelonians /
squamates and crocodiles.

Fig. 5.9: Blood flow through the crocodile heart. (a) Systemic and pulmonary
blood flow during air breathing period. (b) Internal changes that result
in decreased pulmonary flow when the crocodile dives. 149
Block 1 Comparative Anatomy of Vertebrates-I

Fig. 5.10: The flowchart comparing the blood flow patterns in the hearts of (a)
chelonian and squamate, and (b) crocodiles. Dashed lines indicate the
cardiac shunts that divert blood flow from pulmonary circuit to
systemic circuit while diving. During this cardiac shunt resistance to
pulmonary flow increases due to contraction of the sphincter at the
base of pulmonary artery. In crocodiles the vasoconstriction of the
vascular supply to the lungs also aids in this resistance.

So we can summarise that in the later ectothermic stage, out of the six
occurring in the, preceeding stage, only three cardiac chambers remain. One
of these three (the ventricle) is further subdivided so that in this stage there
are four cardiac chambers at least partly separated from each other.

5.2.5 Endotherm Heart

Only a few, relatively slight alterations have occurred in endotherms to effect
150 the maximum efficiency of the heart (Fig. 5.11).
Unit 5 Circulatory System

Fig. 5.11: Endothermic heart, ventral view with parts drawn to side.
Most important is the complete closure of the ventricular wall, rendering
absolutely impossible any mixture of aerated and non-aerated blood. The
trend towards double circulation in the heart was initiated certainly in the
sarcopterygians. It attained perfection in endotherms. Closure of the ventricles
along with simplification of the systemic aorta is the last step in reaching the
greatest possible perfection in the cardiac link of the "forced draft" respiratory
mechanism unique to birds and mammals and chiefly responsible for their
endothermic condition.

In birds, though the sinus venosus is reduced, it still remains a small discrete
area. The conus arteriosus is only transiently present in embryonic condition,
that gives rise to the pulmonary trunk and a single aortic trunk (Fig. 5.12).

Fig. 5.12: The heart of a bird (ventral view). 151

Block 1 Comparative Anatomy of Vertebrates-I
In mammals the sinus venosus is reduced to a patch of Purkinje fibers (also
called sinoatrial node) in the wall of right atrium. This node acts as a
pacemaker, initiating the wave of contraction i.e. rhythmic heart beat that
spreads across the heart like in all other vertebrates. Another mass of typical
muscle fibers called as the atrioventricular node in the waIl of right atrium in
the four chambered heart, also acts as pacemaker under experimental
conditions when the sinoatrial node is destroyed or prevented from
functioning. Similar to that in birds the conus arteriosus during embryonic
development splits to produce the pulmonary trunk and single aortic trunk of
the adult (Fig. 5.13). The mammalian heart possesses only two of the three
sets of valves present in piscine ancestors; the semilunar valves and the
atrioventricular valves. The latter set is now divided into two, to which the
names tricuspid valve and bicuspid or mitral valve are applied in mammals.

Fig. 5.13: Ventral view of the mammalian heart.

Although the structure of avian and mammalian heart is similar they both have
evolved from different reptilian groups. This difference is reflected in their
embryonic development. Both these hearts also function similarly as both
consist of parallel pumps with double circulation circuits. The right side of the
heart gathers deoxygenated blood from the systemic tissues and pumps it in
pulmonary circuit. The left side of the heart pumps oxygenated blood from the
lungs through the systemic circuit. There is no cardiac shunting as hearts of
birds and mammals are divided into right and left chambers.

In single circulation pattern the blood passes only once through the heart
during each complete circle as seen in most of the fishes (Fig. 5.14a).
Amniotes have double circulation pattern in which blood passes twice through
the heart in every circuit (Fig. 5.14b). From the heart the blood goes to the
lungs, back to heart out to systemic tissues and again back to the heart. The
major evolutionary event was addition of pulmonary circuit in the circulatory
pattern. The intermediates that have characteristics of both the conditions
include lung fishes, amphibians and reptiles. The evolution of this type of
circulatory system design is the adaptive advantage of the transitional forms
152 that came onto land.
Unit 5 Circulatory System

Fig. 5.14: Diagrammatical representation of single circulation (a) and double

circulation (b).

You have finished studying about the evolution and structure of heart. In order
to judge your progress attempt the following SAQ.

SAQ 2
i) Fill in the blanks with appropriate words.

a. Sinus venosus is ……………. in size in chelonian heart.

b. Pacemaker or sinoatrial node ……………. the heart beat.

c. The ventricle is …………….divided in squamates and is

……………. divided in crocodiles.

d. The systemic trunk is paired in ……………. and ……………. .

e. Double circulation in heart was initiated in ……………. .

f. Tricuspid and bicuspid valves are present in the ……………. heart.

ii) What is rhythmicity of heart beat in mammals?

5.3 ARTERIAL SYSTEM

Although arterial system in different vertebrates appears to be different in
arrangement, a study of development reveals that all are built upon the same
fundamental plan. The increasing complexity of the heart from the simple two
chambered structure of the lower forms to the four-chambered organ of
crocodilians, birds and mammals is associated with certain variations to be
found in the blood vascular system. Modifications of the embryonic pattern,
which occur during later ontogeny are of such a nature that they adapt the
aortic arches for respiration either by gills or by lungs.

5.3.1 Aortic Arches-Overview

Early chordates evolved blood vessels that were retained as aortic arches in
vertebrates. During embryonic development in all vertebrates the anterior end
of the ventral aorta divides into two arches called aortic arches, which course
dorsally into the mandibular region. Dorsal to the pharynx these are continued 153
Block 1 Comparative Anatomy of Vertebrates-I
posteriorly where they are known as paired dorsal aortae. Additional pairs of
aortic arches then appear in an anterio-posterior sequence, forming
connections between ventral and dorsal aortae on each side (Fig5.15). Each
travels through the tissue between the adjacent pharyngeal pouches. The
typical number of aortic arches that are taken as basic embryonic pattern in
vertebrates is six pairs, although there are certain exceptions among lower
forms, such as, eight in lamprey, fifteen in hagfishes and ten or twelve in some
species of shark. The first is known as mandibular aortic arch and the second
is hyoid aortic arch. The remainder are referred to as the third, fourth, fifth and
sixth aortic arches, respectively and all are designated by roman numerals
(Fig. 5.15). Each aortic arch lies anterior to the visceral cleft bearing the
corresponding number. The two dorsal aortae soon fuse posterior to the
pharyngeal region, so that ultimately only a single dorsal aorta is present. It is
continued in the tail region as a caudal artery. Blood which is pumped
anteriorly by the heart, passes through the ventral aorta to the aortic arches.
These vessels carry blood to the paired dorsal aortae, from which it goes
anteriorly to the head and posteriorly to the single dorsal aorta which
distributes it to the remainder of the body. Veins return blood to the sinus
venosus, atleast during early stages of development. As we go up the
vertebrate series we find that the number of aortic arches decreases from 6 in
fishes to 3 in amniotes. As you read why this decrease happens you will be
able to correlate it with the complexity in heart structure seen from fishes to
amniotes and also the transition from an aquatic life to terrestrial seen in
vertebrates. Let us learn about these changes in the aortic arches in different
groups of animals.

Fig. 5.15: Diagram illustrating the basic pattern of aortic arches in vertebrate
embryos. Ventral view. Six pairs of arches connect ventral and dorsal
154 aortae.
Unit 5 Circulatory System
5.3.2 Fishes

The aortic arches of gill bearing vertebrates are primarily for bringing blood
from the heart via the ventral aorta and afferent arteries through the gills
where the blood is oxygeneated in a capillary bed and drained via the efferent
arteries into the dorsal aorta for circulation in the body. Fish embryos have six
pair of arteries corresponding to the six arches but in the adult fishes there is a
reduction in the number of aortic arches within the superclass. The greatest
number occurs in certain primitive sharks, where it is directly related to the
number of gill pouches. The first pair is generally lost or modified. The IInd
arch is present in elasmobranchs but lost in many other fish (Fig. 5.16). The
anterior continuation of the paired dorsal aortae gives rise to the internal
carotid arteries supplying to the brain; those arising from the ventral aorta form
the external carotid arteries which supply blood to jaws and face.

Fig. 5.16: Diagram of aortic arch region as found in most teleost fishes : ventral
view. Arches I and II have degenerated. Each remaining arch is divided
into afferent and efferent branchial arteries that are connected by gill
capillaries.

Lungfishes have both gills and lungs and depend on their lungs for respiration,
thus they have lost the capillary beds associated with arches III and IV (Fig.
5.17).The corresponding arches (III and IV) are uninterrupted vessels. When
arches I and II are lost then the anterior extension of dorsal aorta from arch III
continues and the internal carotid artery to the head. Fifth and VI have
capillary networks and pulmonary arteries arise from the VI arch and go to the
lungs of lungfishes and to the swim bladder of coelacanths.

Pulmonary veins bring back blood from the lungs to the heart via the left
atrium. This is not a very efficient system but has some ability to shunt blood
to regions of the body where oxygenated blood is necessary.
155
Block 1 Comparative Anatomy of Vertebrates-I

Fig. 5.17: Aortic arches of Protopterus (lung fish).

5.3.3 Amphibians
Since amphibians rely on lungs for respiration, the gills are no longer important
for adults. Amphibians possess external gill filaments, atleast during early
development but they are not homologous with the internal gill lamellae of
fishes, nor are they supplied with blood in the same manner. In amphibians,
generally the first two aortic arches (I, II) disappear early in the development.
The pattern of remaining arches differs between larvae and metamorphosed
adults. The frog larva has last four aortic arches (III-VI) that deliver blood to
the internal gills. The embryonic pulmonary artery buds from the arch VI.
During metamorphosis the gills and arch V are lost in anurans but is present in
some urodeles (salamander) leaving arches III, IV and VI intact (Fig. 5.18).
Dorsal link between III and IV becomes thin in urodeles but is lost in anurans.

Fig. 5.18: Diagram showing modification of aortic arches as found in adult frog.

The anterior continuation of the ventral aorta becomes external carotid artery.
The third arch becomes the internal carotid artery, and both of these carotid
arteries arise from the common carotid which is the section of ventral aorta
that lies between III and IV arches. At the base of the internal carotid lies the
carotid body which is an enlarged portion of the carotid arteries and is formed
at the point of branching of carotid arteries.

The fourth aortic arch persists to become the systemic arches which unite
posteriorly to form the dorsal aorta. Arch VI on each side sends a branch to
the developing lung and another branch to the skin, thus becoming the
156 pulmocutaneous artery. Fig. 5.19 shows the arterial system of the adult frog.
Unit 5 Circulatory System

Fig. 5.19: Diagram of arterial system of adult frog (ventral view).

In salamanders (Caudates) the fifth arch may persist in a very reduced form.
Frequently the radix or the lateral aorta between arches III and IV fails to
degenerate completely. The ductus arteriosus also persists in caudates.

The external gills of larval caudates are supplied with vascular loops
connected to aortic arches. In caudates, at the end of metamorphosis gills
degenerate and atrophy of the vascular loops i.e. capillary net work
surrounding the gill lamellae and collecting loop occurs (Fig. 5.20). 157
Block 1 Comparative Anatomy of Vertebrates-I

(a)

(b)

Fig. 5.20: Diagram showing modification of aortic arches as found in most

caudate amphibians, (a) Salamander (larval); (b) Salamander (adult).

Certain caudates, such as neotenic salamander Necturus are called perenni

branchiates since they retain gills throughout life and fail to metamorphose. In
this amphibian, the fifth aortic arch persists and the pulmonary artery arises
from the fifth arch rather than the sixth, the ventral portion of which is lacking.
Blood going to the lungs in Necturus has already been oxygenated. The lungs,
therefore, under normal conditions are unable to function as respiratory
organs, and the gills are retained. Now try the following SAQ to assess your
understanding.

SAQ 3
Match the items given in column I with those in column II.
Column I Column II
i) Basic embryonic pattern in a) Supply blood to jaws and face
vertebrates
ii) External carotid artery b) Generally in amphibians
iii) Reduction/modification of I and c) Occurs in caudates
II aortic arches
iv) Disappearance of I and II aortic d) Characteristic of teleosts
arches during early
development
v) Atrophy of vascular loops at the e) Six pairs of aortic arches
end of metamorphosis
158
Unit 5 Circulatory System
5.3.4 Reptiles
Since reptiles are fully terrestrial, gills are totally absent and replaced by lungs.
Just as in amphibians, reptiles retain aortic arches III, IV and VI. The fifth arch
may also be retained in reduced form in certain lizards, and a remnant of the
radix between arches III and IV may persist in some snakes.

In most reptiles, further modifications occur in the aortic arches (Fig. 5.21).
Arches I, II and V are lost. Dorsal connection between III and IV disappears.
Arch III forms a part of the internal carotid arteries and forward extensions of
the ventral aorta form the external carotid arteries. Other important changes
are splitting of the distal portion of the conus arteriosus and part of the ventral
aorta into three vessels - left systemic arch, right systemic arch derived from
the IV arch and the pulmonary trunk.

Fig. 5.21: Diagram showing modification of the aortic arches as found in

reptiles. The ventral aorta has split into three vessels.

The right systemic arch arises from the left side of the ventricle carrying
oxygenated blood to the carotid arch (IIIrd) to be sent to the head. Left
systemic arch arises from the right side of the ventricle carrying mixed blood to
the body through the dorsal aorta some mixing may also occur through the
foramen of Panizza. Mixing of oxygenated and deoxygenated blood seems to
be associated with the poikilothermal mode of life. (Poikilotherms are those
animals which cannot adapt themselves to the changing temperature e.g.
fishes and amphibians). The sixth arch on each side gives off a pulmonary
artery carrying deoxygenated blood to the lung. The two pulmonary arteries,
therefore, arise from a common trunk, the pulmonary aorta, coming from the
right side of the ventricle. Thus, the modifications of the aortic arches give rise
to one pulmonary circuit and two systemic circuits each of which arise
independently from the heart.

5.3.5 Birds

Though 6 arches are present in the embryo; only II, IV and VI are retained in
the adults. The ventral aorta splits into two portions, a systemic aorta and a
pulmonary trunk or aorta. The systemic aorta is connected to tile left ventricle,
and the pulmonary aorta to the right. The fourth aortic arch on the right side
leaves the right systemic aorta and together with the radix, leads to the dorsal
aorta proper, which supplies the entire body with oxygenated blood. The left
systemic aorta does not develop fully. The pulmonary trunk leading from the
right ventricle gives off the pulmonary arteries, which are actually outgrowths
of the sixth aortic arch. 159
Block 1 Comparative Anatomy of Vertebrates-I

Fig. 5.22: Diagram showing modification of aortic arches as found in birds.

Until the time of hatching there is a ductus arteriosus on each side,

representing the portion of sixth aortic arch between pulmonary artery and
radix (Fig. 5.22). These serve as shunts from the right ventricle to the dorsal
aorta until the lungs become functional. They close at the time of hatching,
and the blood from the right ventricle is then sent to the lungs for aeration.

5.3.6 Mammals
Changes in the aortic arches of mammals are rather similar to those of birds.
Only III, IV and VI arches are retained in the adult. The IV aortic arch on the
left side together with its radix becomes the arch of the definitive aorta and
therefore, is the left systemic arch in mammals. The IV arch on the right and a
portion of the right radix together become the right subclavian artery. The left
subclavian develops as an enlargement of one of the intersegmental arteries
coming off the aorta (left systemic arch) in this region (Fig. 5.23). From the
right subclavian emerges the right common carotid going to the neck and
head. The VI arch gives rise to the pulmonary aorta that divides in right and
left pulmonary arteries going to the lungs. In mammalian embryos there is at
first a ductus arteriosus on each side, but the one on the right persists for only
a short time. The left one, which serves as a shunt between pulmonary and
systemic aortae, persists until birth, when it finally becomes occluded. The
carotid arteries are formed from the paired third arch.

Fig. 5.23: Diagram showing modification of aortic arches as found in mammals.

5.3.7 Evolution of Aortic Arches

Till now you have studied about structure of aortic arches in different
vertebrates. Let us now summarise their evolution in vertebrates.
160
Unit 5 Circulatory System
The aortic arches in most of the fishes deliver the deoxygenated blood to the
respiratory surfaces of the gills and from there distribute the oxygenated blood
to the head region through carotid arteries and to the rest of the body through
the dorsal aorta. In lung fishes and tetrapods the aortic arches form two types
of circuits – arterial circuit to lungs through pulmonary arch and arterial circuit
to rest of the body through systemic arches. Blood to the head in tetrapods is
supplied by carotid arteries that arise from the systemic arch. The left and right
arches are present in reptiles also. However, these are reduced to single
systemic arch, the right arch in birds and left arch in mammals. You can see
the evolution of aortic arches in Fig. 5.24.

Fig. 5.24: The evolution of aortic arches in vertebrates. Hypothetical ancestor

with 6 aortic arches corresponding to 6 pairs of gill pouches. Primitive
fishes, represented by sharks, also have six paired gill arches. In
st nd
teleosts, the 1 and 2 gill arch arteries are absent and only four pairs
are seen in the caudal branchial arches. Lungfish have both gills and a
pulmonary circulation with the gill arches corresponding to arches
two, five, and six. During air respiration, the blood is shunted through
arches three and four, while the ductus arteriosus in arch six shunts
oxygen-poor blood away from the gills and to the lungs. In adult
amphibians, the gill arches are lost and the aortic arch vasculature
remains bilaterally symmetrical. Oxygenated and de-oxygenated blood
enter the ventricle through the right and left atrium and leaves the
heart through a single outflow tract containing a spiral valve. In
reptiles such as the turtle, aortic arch artery four remains bilateral but
is divided at the base of the outflow tract. The outflow tract is divided
into three arteries: right and left aortic arch arteries and the pulmonary
artery. In mammals, the fourth aortic arch arteries become bilaterally
asymmetrical and the outflow tract separates into two distinct outflow
vessels. Birds also have a completely divided outflow tract with
asymmetrical aortic arch arteries. 161
Block 1 Comparative Anatomy of Vertebrates-I
5.3.8 Types of Arteries
Most of the vertebrate body is supplied with blood through branches of the
aorta which for convenience sake may be grouped under two divisions:
somatic arteries supplying the body proper, and visceral arteries, distribute
blood to various portions of the digestive tract and associated structures,

1) Somatic arteries: They supply portions of the body derived from the
embryonic epimere, being distributed to the dorsal musculature and
vertebral column, where they are referred to as parietal, or segmental
arteries. In higher forms in which the body is divided into more or less
definite regions, terms such as intercostal, dorsolumbar, and sacral are
used. The vessels going to the pectoral appendage are called
subclavians and those to the pelvic are iliacs, which may be composed
of a union of several segmental arteries.

2) Visceral arteries: The arteries supplying the viscera are of two kinds:
paired and unpaired. The paired arteries are segmentally arranged and
supply the body parts that are derived from the embryonic mesomere
from which the urinogenital organs and their ducts arise. These are
termed as renal, genital, ovarian, spermatic and urinogenital arteries.
Renal and genital arteries are numerous in the lower vertebrates, but the
number is much reduced in higher forms. The unpaired visceral arteries
supplying spleen, and the digestive tract with its derivatives, are vessels
which course through the dorsal mesentery of the gut. There are usually
three unpaired visceral arteries in vertebrates. The most anterior of
these is the celiac artery, supplying the anterior viscera, including
stomach (gastric), spleen (splenic) pancreas (pancreatic), liver (hepatic),
and duodenum (duodenal). The second unpaired visceral artery is the
superior mesenteric, which supplies the entire length of the small
intestine, with the exception of the pyloric end of the duodenum, which is
taken care of by the celiac artery. The third unpaired artery is the inferior
mesenteric, supplying to the posterior part of the large intestine and
rectum.

SAQ 4
Give short answers to the following :

i) What is the major modification that occurs in reptilian aortic arches?

ii) What is the role of ductus arteriosus in bird's circulatory system'?

iii) How does right subclavian develop in mammals?

iv) Write difference between somatic and visceral arteries.

5.4 VENOUS SYSTEM

As in the case of the arterial system, a comparative account of veins in various
vertebrate groups shows that they are arranged according to the same
162 fundamental plan and that the variations form a logical sequence as the
Unit 5 Circulatory System
vertebrate scale is ascended. In its development, venous system of higher
forms passes through certain stages common to the embryos of lower forms
(Fig.5.25). Basically three major sets of paired veins are present in the early
developmental stage – vitelline veins from the yolk sac, cardinal veins from
the body of the embryo and lateral abdominal veins from the pelvic region.

Fig. 5.25: Diagram of basic embryonic venous complex from which the venous
system of vertebrates are derived.

Vitelline veins include the hepatic sinusoids and hepatic veins. Hepatic veins
collect the blood from hepatic sinusoid network and enter the sinus venosus.
The cardinal veins consist of anterior cardinal veins that collect the blood from
the head region and, posterior cardinal vein that collects blood from the rest of
the body of the embryo. Both these veins meet at common cardinal veins also
called as duct of Cuvier that opens into sinus venosus. The lateral abdominal
veins are present in fishes but are normally absent or merged in tetrapods. We
will discuss about these veins while discussing the venous system of fishes.
The vein that carries the absorbed end products of digestion from the digestive
tract to the vascular sinusoids in liver is called the hepatic portal vein. It is
present in all vertebrates and is developed from the embryonic subintestinal
vein originating in the caudal vein. The subintestinal vein loops around the 163
Block 1 Comparative Anatomy of Vertebrates-I
anus, extends along the ventral wall of intestine, passes through the liver and
finally joins the left vitelline vein. The structural variations in the venous system
of different vertebrates are described in further sub-sections.

5.4.1 Fishes
Many primitive features are retained by the venous system of fishes, but
important advances over conditions in primitive chordates are evident
(Fig. 5.26).

Fig. 5.26: Diagrams, a) shark and b) lungfish, illustrating the changes over the
primitive condition which occur in the venous system of fishes.

The sinus venosus receives a duct of Cuvier on each side. An anterior

cardinal, or jugular vein brings blood to the duct of Cuvier from the dorsal side
of the head region. In many fishes a pair of inferior jugular veins from the
ventrolateral part of the head also enters the common cardinal veins (Fig
5.26a). These are lacking in Polypterus, fused in Lepisosteus. Each common
cardinal also receives a postcardinal vein from the posterior end of the body.
Since fishes are the first vertebrates to possess paired appendages,
subclavian and iliac veins, bringing blood from the pectoral and pelvic fins,
respectively, make their appearance first in this group. The iliac veins join the
lateral abdominals which course in the body wall to join the common cardinal.
The posterior wall of the sinus venosus usually receives two hepatic veins,
which return blood from the liver to the heart. The subintestinal vein has, in
most cases, lost its connection with the caudal vein.

With the development of the opisthonephric kidneys (The adult kidney formed
from the mesonephros and additional tubules from the posterior region of the
164 nephric ridge. You will study more about this in the unit on Excretory System)
Unit 5 Circulatory System
and their posterior elongation, the postcardinals grow backward dorsal to the
kidneys, ultimately to unite with the caudal vein. A pair of subcardinal veins
develop ventral to the kidney, i.e. between opisthonephric kidneys. Small
blood vessels connect the postcardinal and subcardinal veins. They are not
associated with glomeruli, which receive arterial blood. Blood in the caudal
vein has now two alternative routes through which it may pass on its way to
the heart. It may either go into the postcardinals directly or indirectly through
the subcardinals renal veins. The next advance seen in teleosts involves an
interruption in the course of each postcardinal vein so that anterior and
posterior portions are no longer continuous. Blood from the tail now travels
only by one route. It passes through the kidneys, i.e. the renal portal veins to
enter the subcardinal veins thus, establishing the renal portal system. The
blood courses anteriorly via the postcardinals, which join the duct of Cuvier.
The postcardinals usually receive veins from the gonads. The blood passing
through the renal veins, flows in a direction reverse of that occurring during an
early stage of development, after the renal portal circulation is established.
These changes are evident upto elasmobranch fish stage.

In teleosts, lateral abdominal veins are not present. The subclavians enter the
common cardinals, and the iliacs join the postcardinal. Veins from the swim
bladder (derivative of the gut) usually join the hepatic portal vein. However, in
certain forms the connection is with the postcardinal veins. In Polypterus they
join the hepatic veins directly. In dipnoans, the pulmonary veins from the swim
bladder enter the newly formed left atrium of the heart, and the double type of
circulatory system appears for the first time.

Lung fishes show the characters that are connecting links between
amphibians and fishes. In Epiceratodus a single, midventral; anterior
abdominal vein, similar to that of amphibians, makes its appearance. The
lateral abdominal veins have fused to form the anterior abdominal vein, which
courses forward to enter the sinus venosus iliac joins with renal portal and the
pelvic vein is a branch of iliac which joins the anterior abdominal. The right
postcardinal, including its posterior portion, has become much larger than its
counterpart on the left side. The connection with the caudal vein on both sides
is retained. The larger vessel on the right side is now called the postcaval vein.
It passes through the liver and enters the left duct of Cuvier. The venous
system of Protopterus, except for the lack of an anterior abdominal vein,
resembles more closely with that of amphibians than that of Epiceratodus.

5.4.2 Amphibians

In amphibians, the renal portal and hepatic portal systems are brought into
close association, since blood from the hind limbs must pass through one or
the other.

Pulmonary veins from the lungs enter the left atrium, as in Dipnoi. In lungless
salamanders, of course, pulmonary veins are absent and the left atrium is
reduced in size.

The ducts of Cuvier, which originally received the subclavian, jugular, and
postcardinal veins, are further consolidated in amphibians and are now called 165
Block 1 Comparative Anatomy of Vertebrates-I
the precaval veins. These enter sinus venosus on each side. The jugular vein
has external and internal tributaries. Since cutaneous respiration has
developed to a high degree in amphibians, exceptionally large cutaneous
veins are present which join the subclavians to enter the sinus venosus.

Although both urodele (caudates) and anurans (salientians) are similar in the
above respects, they exhibit certain differences in regard to the arrangement
of the postcaval postcardinal complex. In most caudates and a few salientians
the anterior portions of the postcardinals persist in reduced form connecting
the middle portion of the postcava with the duct of Cuvier on each side. In
most adult anurans however, the anterior portions of the postcardinals usually
disappear and furnishes the only route through which blood from the kidneys
and gonads may return to heart (Fig. 5.27).

Fig. 5.27: Diagram of venous system of a typical anuran amphibian: ventral

view.

SAQ 5
Answer the following questions in short.

i) What are the three sets of paired veins present in the early embryonic
stage of vertebrates?

ii) What are the veins in fishes that bring blood from the fins?

iii) What is the fate of lateral abdominal veins in teleosts and lung fishes?

iv) What are precaval veins?

v) Why are large cutaneous veins present in amphibians?

166
Unit 5 Circulatory System
5.4.3. Reptiles
In this group following further partitioning of the heart the large systemic veins
entering heart have shifted more to right (Fig. 5.28). Two precavae are the
original ducts of Cuvier, which receive jugular, subclavian, and postcardinal
veins. The anterior portion of the jugular, subclavian, and postcardinal have
degenerated into two small vertebral veins. In snakes the subclavians are
lacking. More blood from the posterior part of the body now courses through
the anterior abdominal vein which joins the hepatic portal veins anteriorly. The
importance of the renal portal vein has diminished, and in some forms direct
channels may even pass through the kidneys, connecting renal portal and
postcaval veins. Since cutaneous respiration does not exist in reptiles,
pulmonary circulation has assumed greater importance.

(a) Turtle (b) Crocodile

Fig. 5.28: Venous channels in reptiles a) turtle, b) crocodile. A strong branch of

the renal portal vein in crocodilians continues directly to the postcava.
The pelvic vein occurs in turtles but has been omitted to facilitate
identification of the basic pattern.

Depending upon the degree of asymmetrical development of the respiratory

organs, pulmonary veins from the two lungs show discrepancies in size in
various forms. One pulmonary vein may even be entirely absent in certain
snakes in which the left lobe of the lung is absent. The reptilian postcava, as in
amphibians, is derived partly from the subcardinals and partly from the vitelline
veins. The postcardinals have practically disappeared.

5.4.4 Birds
The sinus venosus has been incorporated in the wall of the right atrium, so
that the two precavae and single postcava enter the right atrium directly. Each 167
Block 1 Comparative Anatomy of Vertebrates-I
precava is formed by the confluence of subclavian and jugular veins. The
original postcardinal connection is no longer in existence. The postcava is the
chief pathway for the return of blood from the posterior part of the body. The
posterior end of the postcava receives blood directly from the limbs via the
renal portal veins. The hepatic veins of birds join the postcava as it nears
heart. The caudal vein in birds is greatly reduced. A vein variously known as
the inferior mesenteric, coccygeomesenteric, and caudal mesenteric, connects
the caudal vein with the hepatic portal vein. A small epigastric vein carries
blood from the great omentum to one of the hepatic veins (Fig. 5.29).

Fig 5.29: Diagram of venous system of bird, ventral view.

5.4.5 Mammals

The shifting of the main venous channels to right side is more clearly indicated
in mammals than in other vertebrates. In some mammals two precavae are
present, but in others they are joined. The vessel on each side which receives
jugular and subclavian vein is then called the brachiocephalic (innominate)
vein. In mammals a portion of the anterior end of the right postcardinal persists
168 as the azygos vein. This drains the intercostal muscles and enters the
Unit 5 Circulatory System
precava. The greatest change in the venous system of mammals occurs in the
postcava, which has become considerably simplified. No trace of a renal portal
system is found in adults and all blood from the posterior end of the body is
collected by the postcava. The anterior abdominal vein has disappeared in
mammals, being found only in the monotreme, Echidna. Usually only the left
umbilical vein persists, passing through the liver as the ductus venosus to join
the postcava before it enters the heart. The hepatic portal vein usually referred
to as the portal vein, is similar to the hepatic portal vein of lower forms
(Fig. 5.30).

Fig. 5.30: Diagram of venous system of cat, ventral view. 169

Block 1 Comparative Anatomy of Vertebrates-I

SAQ 6
Fill in the blanks with appropriate words.

i) Subclavians are absent in ____________.

ii) The reptilian postcava is derived partly from ____________ and partly

from ____________.

iii) The ____________ brings the blood from the posterior part of the body.

iv) The caudal vein is greatly reduced in ____________.

v) The ____________, ____________ transports blood from great

omentum to hepatic vein.

vi) In ____________ the azygos vein brings the blood from intercostal

muscles.

vii) The anterior abdominal vein has disappeared in mammals, except in

____________.

viii) ____________ is the process of formation of blood vessels.

ix) The basic four chambered heart in most tetrapods is formed from
____________.

5.5 BLOOD
Blood may be described as a specialized connective tissue in which there is
liquid intercellular substance known as plasma, and formed elements, the red
blood cells, the white blood cells and the platelets suspended in the plasma.

5.5.1 Composition of Blood

The general composition of the whole blood is summarized below. Among the
cellular components the red blood cells or erythrocytes are the containers of
haemoglobin, the respiratory pigment. The erythrocytes in all the groups of
animals, except for mammals have nuclei. Red blood cells vary in size for
example, 8 m in humans, 9 m in elephants and 80 m in some
salamanders. The life of the erythrocytes is three to four months in the
circulating blood before being broken down and replaced. White blood cells,
the second major cellular component of the blood defend the body from
infection and disease. The third type of blood cells are the platelets that help in
the formation of clot at the site of tissue damage. Blood has important roles to
play in body processes such as respiration, protection against diseases,
nutrition, excretion, regulation of body temperature, maintenance of water
170 balance, and transport of hormones.
Unit 5 Circulatory System
Blood

Cells Plasma

- Red blood cells

- White blood
cells
- Platelets

Water 91-92% Solid 8-9%

%

Inorganic Organic
constituents constituents 7.5%
0.9%
- Sodium
- Potassium
- Calcium
- Magnesium
- Phosphorus
- Iron
- Copper

Proteins Non protein Fats Carbohydr Others Colouring

- Serum substances ates - Internal matters
- Neutral fats
albumin - Urea - Phospholipis - Glucose secretions - Bilirubin
- Serum - Uric acid - Cholesterol - Antibodies - Careteus
globulin - Xanthine - Cholesteroid - Enzymes - Xantho
- Fibrinogen - Hypoxanthine - Amylase phyllin
- Prothrombin - Creatin - Protease
- Ammonia
- Acids

5.5.2 Respiratory Pigment

Haemoglobin is the only respiratory pigment among vertebrates. This

combines with far greater amounts of oxygen than any of the other respiratory
pigment.

Haemoglobin is made up of an iron porphyrin compound associated with a

protein, globin. This heme component of the molecule is a constant feature of
all haemoglobins, but the globin portion varies in different vertebrates. The
oxygen carrying capacity of haemoglobin in various vertebrates is given in
Table 5.1. 171
Block 1 Comparative Anatomy of Vertebrates-I
Table 5.1: Haemoglobin and its oxygen carrying capacity in different vertebrate
groups.

Pigment Colour Site Animals Oxygen carrying

capacity in volume per
cent

Haemoglobin Red Corpuscles Mammals 15-30

Birds 20-25

Reptiles 7-12

Amphibians 3-10

fishes 4-20

5.6 LYMPHATIC SYSTEM

Lymphatic system resembles the blood vascular system in that it consists of
vessels, fluids in transit, and associated organs. A major difference is that
lymph flows in only one direction that is towards the heart. The lymphatic
system of all vertebrates consists of thin walled vessels called lymphatics or
lymph vessels. The walls of lymphatics are like that of veins and like veins
they also contain one way valves. The lymphatics penetrate nearly all the soft
tissue of the body and commence as blind-end lymph capillaries that collect
interstitial fluids. Once inside the lymph capillaries, the fluid is called lymph, a
colourless or pale-yellow fluid containing metabolites and secretions, which
constantly collect in the intercellular spaces. The lymph from any area mirrors
the metabolic activities of that area from moment to moment.

Lymphatic system also includes lymphatic tissue which consists of connective

tissue and free cells, the leucocytes, plasma cells and macrophages.
Lymphatic tissue can be found almost anywhere in the body as diffusely
distributed tissue, in patches or encapsulated in lymph nodes. A lymph node is
a collection of lymphatic anastomosing lymph capillaries, macrophages,
reticular cells and lymphocytes wrapped in capsule of fibrous connective
tissue. Lymph nodes are located within channels of lymphatic vessels and
occur in mammals and some water birds only. Lymph nodes have a crucial
role in the immune system of mammals as they form check points for
infections, bacteria and also intercept the cancer cells that migrate through
the lymph nodes (the rapidly dividing cancer cells fill up the lymph nodes. If
such a case is detected in the tests all the cancer affected nodes should be
removed).

The lymphatics in the villi of the small intestine collect globules of fat absorbed
from the intestine after a meal. If the meal has been particularly fatty, the
lymph in these vessels is milky. For this reason, the lymphatics of the intestinal
villi are called lacteals, and the lymph therein is called chyle.

Several factors that are instrumental in propelling the slow moving lymph
172 through lymphatic vessels and nodes are as follows.
Unit 5 Circulatory System
1) Muscular activity of various parts of the body.

2) Pressure built up in the smaller vessels by osmosis and absorption of

tissue fluid.

3) The action of pulsating lymph hearts which consists of enlargements in

lymphatic vessels with contractile walls, usually situated near a point
where lymph enter the venous system. Lymph hearts are not true hearts
because they lack cardiac muscles. The striated muscles in their walls
slowly develop pulses of pressure to drive the lymph.

Let us discuss the lymphatic system in different vertebrates.

5.6.1 Fishes

Lymphatic vessels in fishes are extensively developed, peripherally located

and extend into head, tail and fins. Deeper channels follow the course of some
of the larger veins.

Lymph hearts are usually not present, but in some forms they are present near
the point of junction of lymphatic vessels and veins. Eel has a lymph heart in
the tall. The European catfish has two caudal lymph hearts. Lymph nodes are
lacking in fishes.

5.6.2 Amphibians

Two main sets of lymphatic vessels are present in caudate amphibians.

Superficial vessels beneath the skin carry lymph to cutaneous and
postcardinal veins. Between 14 and 20 lymph hearts have been observed
along their course in various forms. Deeper channels follow the dorsal aorta
on each side and enter the subclavian artery. In anurans, with large lymph
sacs, most of the lymph flows towards heart. Two pairs of lymph hearts are
usually present in adult animals. First pair near the third vertebra, pumping
lymph into the vertebral vein, and a second posterior pair, located near the
end of the urostyle pumps blood into the transverse iliac vein. Lymph hearts
are more numerous in larval and tadpole stages. More than 200 lymph hearts
are present in caecilians, lying beneath the skin along the intersegmental
veins.

5.6.3 Reptiles

The lymphatic system is well developed. A large subvertebral trunk divides

anteriorly to enter the precaval veins. In snakes, lymphatic vessels and
sinuses are exceptionally large and numerous. A posterior pair of lymph hearts
pumps lymph into the iliac veins.

5.6.4 Birds

The lymphatic vessels of birds ultimately enter two thoracic lymph ducts which
join the precaval veins. Transitory lymph hearts, not found in adult birds, may
be observed in the pelvic region during embryonic development. 173
Block 1 Comparative Anatomy of Vertebrates-I
5.6.5 Mammals
Lymph hearts are altogether lacking in mammals. A main trunk, the thoracic
duct drains all the lymphatic vessels of the posterior part of the body, as well
as those coming from the left side of the head, neck and thoracic regions. In
mammals nodes are numerous in superficial regions of the head and neck,
axillae, and groin. Many lie within the body cavity, large and numerous in the
mesentry of the intestine. In all those regions they serve to prevent the
invasion of the body by bacteria (Fig. 5.31).

(a) (b)

Fig. 5.31: Lymphatic circulation and lymph nodes. (a) Lymphatic vessels
returning from all parts of the body join to form major lymphatic
vessels, the largest being the thoracic duct, which empties lymph into
the postcaval or subclavian, veins, (b) Cross section of a lymph node.
These lymph nodes house lymphatic tissue, which functions to
remove foreign materials from the lymph circulating through them.
Lymph nodes have a cortex and medulla bound by a fibrous
connective tissue capsule. Notice the entering and departing
lymphatic vessels.

5.6.6 Other Lymphatic Organs

Some organs of pharyngeal origin usually considered to belong to lymphatic
system include tonsils, adenoids and thymus glands. Tonsils and adenoids
guard the body by forming antibodies against antigenic substances. They filter
the tissue fluid and produce lymphocytes. Thymus gland has a role to play in
some of the immunological reactions in the body. Peyer’s patches are nodules
of lymphoid tissue in the small intestine especially numerous in the region of
the ileum. Some other organs related to circulatory system in the body are as
174 follows.
Unit 5 Circulatory System
Hemolymph glands: These glands of the body closely resemble lymph
glands except that they enclose blood vessels rather than lymphatic. Blood,
rather than lymph, filters through them. They are also called hemal nodes.
They are believed to play a role in the destruction of old and worn-out
corpuscles and also erythrocyte formation. Ruminating mammals have
numerous hemal-nodes but their occurrence in man is doubtful.

Spleen: The largest lymphoid organ in body is the spleen. It is sometimes Spleen enlargement
considered to be a hemolymph gland since it is interposed in the blood stream is feature of malaria
and in some other
rather than in lymphatic vessels. The blood vessels are peculiarly arranged in
conditions in which
the spleen so that the, blood comes in contact with the phagocytes the organ may
(macrophages) that engulf the fragments of disintegrating red corpuscles. assume relatively
Lymphocytes and plasma cells originating in thymus are later formed in large much larger size.
number in spleen. The spleen also serves as a storehouse for erythrocytes.
Spleen also produces antibodies serving in the defence mechanism of the
body.

SAQ 7
i) a) Name two blood proteins .

b) Name two non-protein substances of blood.

c) Name three inorganic constituents of blood.

d) Describe the composition of respiratory pigment present in

vertebrates.

ii) Match the items given in column I with those given in column II.

Column I Column II

a) Arteries i) Lymph

b) Unidirectional flow ii) Blood

c) Collection of interstitial fluid iii) Lacteals

d) Chyle iv) Lymph capillaries

e) Absence of lymph nodes v) Birds

f) Transitory lymph heart vi) Mammals

g) Absence of lymph heart vii) Fishes

h) Production of lymphocytes viii) Tonsils

i) Lymphoid nodules in ileum ix) Spleen

j) Phagocytosis of disintegrating x) Peyer’s patches

red corpuscles

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Block 1 Comparative Anatomy of Vertebrates-I

5.7 SUMMARY
Let us symmarise whatever you learnt in this Unit:

 The circulatory system includes blood vascular and lymphatic systems.

Blood vascular system consists of heart, arteries, capillaries, and veins.
Lymphatic system is composed of lymphatics (including lacteals), lymph
capillaries, lymph sinuses, lymph nodes in mammals and birds. Lymph
hearts are present in lower vertebrates, but absent in birds and
mammals. Lymph is transported from tissue spaces, and chyle, which is
also a form of lymph, is transported from intestinal villi to certain major
venous channels.

 An aortic arch is a blood vessel connecting the ventral and dorsal aortae
and located, in the embryo at least, in a visceral arch. Typically, six pairs
of aortic arches develop in each vertebrate embryo. During ontogeny the
aortic arches are reduced in number, the most highly evolved
vertebrates retaining fewest arches.

 A sinus venous occurs in fishes, amphibians, and reptiles. It is absent in

adult birds and mammals, having been incorporated in the right atrial
wall during embryonic development.

 There is no mixing of oxygenated and deoxygenated blood in fishes.

Considerable mixing occurs in tailed amphibians, and less occurs in
frogs and reptiles. Mixing occurs in fetal birds and mammals but not in
adults.

 Major venous channels in the basic pattern of vertebrate circulation are

anterior cardinal (internal jugular) veins from the head, postcardinal
veins from the trunk and kidneys, and subclavian vein from the anterior
appendages – all flow into common cardinal veins; abdominal veins from
the hind limbs; renal portal system from the tail; and hepatic portal
system from the chief digestive organs. Hepatic sinuses drain the liver,
coronary veins drain the musculature of the heart, and pulmonary and
postcaval veins are added in lung-breathing forms.

 The renal portal system drains only the tail in fishes. It acquires a
connection (external iliac) with the hind limb drainage in amphibians and
reptiles. In crocodilians and birds the connection may bypass the
kidneys and go directly to the postcava. The renal portal system is
absent in adult mammals, except monotremes, and the hind limbs and
tail are drained solely by the postcava.

 Blood may be described as a specialized connective tissue in which

there is intercellular substance known as plasma, and formed elements,
the red blood cells, the white blood cells and the platelets suspended in
the plasma. Specific gravity varies from 1.055 to 1.060.

 The lymphatic system of all vertebrates consists of thin walled

lymphatics. In birds and mammals, the lymph nodes are interposed
along the course of the lymphatics. The lymphatics penetrate nearly all
the soft tissue of the body and commence as blind-end lymph capillaries
that collect interstitial fluid which is pale yellow fluid containing
176 metabolites and secretions which collect in the intercellular spaces.
Unit 5 Circulatory System
 Lymphatic vessels in fishes are extensively developed. Peripherally
located channels extend into head, tail and fins. Lymph hearts are
present in amphibians in which two main sets of lymph vessels are
present. In reptiles and birds, lymphatic system is well-developed. In
mammals, there are no lymph hearts; but several lymph nodes are
present. The largest lymphoid organ in body is spleen.

5.9 TERMINAL QUESTIONS

1. Write short note on the general structure of heart wall.

2. Name different stages in the evolution of heart.

3. What is pace-maker?

4. Write short note on foramen of Panizza.

5. Discuss lymphatic system in different vertebrates.

5.10 ANSWERS
Self-Assessment Questions
1. i) peristaltic waves

ii) atrium

iii) tuberculate

iv) interatrial septum

v) cutaneous

2. i) a) reduced

b) initiates

c) partially, completely

d) squamates and crocodiles

e) chondrichthyes

f) mammalian

ii) Heart beat originates in a bundle of typical muscle fibers located in

the wall of the sinus. This is called as ‘Sinoauricular node’ (SA
node). The heart beat further spreads to another as of a typical
muscle fibers, in the wall of right atrium.

3. i -e

ii - a

iii - d

iv - b

v-c 177
Block 1 Comparative Anatomy of Vertebrates-I
4. i) Formation of right and left aortic arches, pulmonary trunk and a
part of ventral aorta.

ii) It serves as shunt from right ventricle to the dorsal aorta until the
lungs become functional.

iii) The fourth arch on the right and a portion of the right radix together
become the subclavian artery.

iv) Somatic arteries supply blood to the body parts derived from the
embryonic epimere and visceral arteries supply to digestive tract
and other organs derived from embryonic mesomere.

5. i) Vitelline veins from yolk sac, cardinal veins from the body of
embryo and lateral veins from pelvic region.

ii) Subclavian and iliac veins bring the blood from pectoral and pelvic
fins, respectively.

iii) Lateral abdominal veins are absent in teleosts but in lung fishes
they fuse to form the anterior abdominal veins which enter the
sinus venosus.

iv) The ducts of Cuvier which originally received the subclavian,

jugular and postcardinal veins are further consolidated in
amphibians and are called precaval veins.

v) It is because the cutaneous respiration has developed to the

higher degree in amphibians.

6. i) snakes

ii) subcardinals, vitelline veins

iii) postcava

iv) birds

v) epigastric veins

vi) mammals

vii) Echidna

viii) angiogenesis

ix) splanchnic mesoderm

7. i) a) Fibrinogen, prothrombin

b) Creatine, Urea

c) Sodium, potassium, calcium

d) Haemoglobin is made up of an iron porphyrin compound,

hence associated with a protein, globin. The heme
component is a constant feature of all haemoglobins, but the
globin portion varies in different vertebrates.

178 ii) a) ii; b) i; c) iv; d) iii; e) vii; f) v; g) vi; h) viii; i) x; j) ix.

Unit 5 Circulatory System
Terminal Questions
1) Heart wall consists of 3 layers viz.

a) Endocardium - the inner wall consisting of endothelium and

elastic tissue.

b) Myocardium - muscular portion in between endo- and

epicardium.

c) Epicardium - outer fibrous tunica, covered by visceral

pericardium (subdivision of coelom).

2) Following are the different stages in the evolution of heart.

a) The protochordate stage without alimentary pharynx.

b) The piscine stage with branchial pharynx.

c) The early tetrapod stage with primitive lungs.

d) The later tetrapod stage of higher ectotherms.

e) The stage of endothermic tetrapods.

3) The excitation tissue which is responsible for the initiation of the

heartbeat and embedded in the wall of the right atrium is called
sinoauricular (SA) node or sinuatrial node. This is also called the
pacemaker (myogenic center) of the heart.

4) The crocodiles, which have a left aortic arch, do not receive blood from
its own (right) ventricle, since the semilunar valves actually prevent flow
from ventricle to aorta except under unusual stress situation. The left
arch receives blood through a foramen of Panizza, connecting the right
and left arches where they cross a short distance.

5) Refer to Sub Section 5.6.1 to 5.6.5.

Acknowledgement of Figures
Fig. 5.24: Source: modified from Keyte, Anna & Alonzo, Martha & Hutson,
Mary. (2014). Evolutionary and Developmental Origins of the
Cardiac Neural Crest: Building a Divided Outflow Tract. Birth
Defects Research Part C: Embryo Today: Reviews. 102.
10.1002/bdrc.21076)

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Block 1 Comparative Anatomy of Vertebrates-I

180