MZO-003

COMPARATIVE ANIMAL
Indira Gandhi PHYSIOLOGY AND
National Open University
School of Sciences BIOCHEMISTRY

VOL

1
COMPARATIVE ANIMAL PHYSIOLOGY
BLOCK 1
FUNDAMENTALS OF ANIMAL PHYSIOLOGY 5
BLOCK 2
SENSING THE ENVIRONMENT AND COORDINATION 171
 COURSE: COMPARATIVE ANIMAL
PHYSIOLOGY AND BIOCHEMISTRY
Throughout history, physiological advances have been made because of detailed
observations of living and dead animals, united with carefully planned experiments to
elucidate how animals work. Advances in physiology have followed in step with advances in
physics, chemistry, and molecular biology, which have allowed physiologists to gain an ever-
increasing understanding of animal structure and function.

A number of important unifying themes apply across all of the subdisciplines of physiology.
(1) Physiological processes obey physical and chemical laws. (2) Physiological processes
are often regulated. (3) Genotype and phenotype are linked. (4) Phenotypes are the product
of evolution.

This course, entitled, “Physiology and Biochemistry” is organized in 2 volumes. Volume 1

deals with Physiology and Volume 2 concentrates on Biochemistry.

Volume 1 of the course on ‘Physiology’ consists of two blocks and will introduce you to the
various life supporting properties, functions and processes of animals of their parts. These
processes can be studied at various levels of organization from membranes through to
organelles, cells, organs, organ systems and to the whole animal. You have studied animal
physiology at your undergraduate level also. We have dealt with more complex principles
and process of physiological phenomena in this course material, so try to recapitulate your
knowledge acquired at undergraduate level before studying this course.

Volume 1 comprises two blocks with five units each. In the Block 1 of Volume 1 you will study
about transport of nutrients, waste products and respiratory gases between the blood and
tissue in cardiovascular and Respiratory Physiology. Regulation of total body balance of
any substance and reflexes that alter excretion via urine are discussed in Renal Physiology.
Physiology of movement, the characteristic property of animals will be discussed in Muscle
Physiology. Energy metabolism in animals in relation to oxygen consumption will be dealt in
Digestive System. Block 2 comprises of five units. Conceptual view of organisation of
nervous system and integrated function of the same will be discussed in Nervous System.
Specific sensory receptors that respond to environment in a well coordinated mechanism are
discussed in Sensory Receptors. Heat balance, an important mechanism in animals and
how do they adapt themselves in extreme temperature are mentioned in Thermoregulation
Phenomenon dynamics and applications of bioluminescence are discussed in Unit on
Bioluminescence. You will be introduced to the mechanism of electric discharge, and
electric organs in different organisms in the last Unit Electric Organs of the Volume 1.

Volume 2 of the course “Biochemistry” consists of two blocks. The ultimate goal of
biochemistry is to explain all life processes in molecular detail. As life processes are
performed by organic molecules, the discipline of biochemistry relies heavily on fundamental
principles of organic chemistry and other basic sciences.

This volume will speak about enzymes at length. Basic characters of enzymes and overview
of enzyme kinetics that includes kinetics of multi substrate enzyme catalyzed reactions and
allosteric enzymes will be provided. Various types of enzyme inhibition and regulation of
enzyme activity will also be explained.

Energetics and Design of living system are dealt in detail in this volume. You will appreciate
the metabolic network and integration of metabolism that are discussed in a fascinating may
in Volume 2. 3
 Objectives
After studying this course, you should be able to:

• describe the structure of the human heart, cardiac cycle and calculate the cardiac
output,

• explain the process of gas transport between the blood and tissues in mammals,

• describe the anatomical diversity and functional principles of excretory organs in

different groups of animals,

• identify the components of the neuromuscular junction, and summarize the events
involved in the neural control of skeletal muscle contraction and relaxation,

• describe the digestion process in humans as an example of mammalian digestion,

• explain energy metabolism in animals in relation to oxygen consumption,

• describe the conduction of action potentials along myelinated and non myelinated
axons,

• explain the effect of temperature on animals and explain how animals tolerate the
extreme temperatures,

• explain general properties of sensory receptors and mechanism of stimulus detection,

amplification, transmission and transduction,

• discuss the presence and control of chromatophores in different groups of invertebrate

and vertebrate animals,

• discuss the phenomenon of bioluminescence and its application for mankind,

• describe the role of enzymes in lowering the active activation energy and in coupled
reactions,

• list the types of enzymes and cofactors and discuss the mechanism of enzyme action,

• explain the effects of temperature, pH and enzyme concentration on the rate of enzyme
action,

• draw the curve for Michaelis-Menten equation,

• explain enzyme inhibition and regulation of enzyme activity,

• describe molecular approach to understand life forms and living processes,

• explain structure and function of ETS,

• describe various metabolic pathways and their linkages, and

• appreciate the integration of metabolism in human body.

4
 MZO-003
COMPARATIVE ANIMAL
Indira Gandhi PHYSIOLOGY AND
National Open University
School of Sciences BIOCHEMISTRY

Block

1
FUNDAMENTALS OF ANIMAL PHYSIOLOGY
UNIT 1
Circulatory System 9
UNIT 2
Respiratory System 33
UNIT 3
Osmoregulation and Excretion 56
UNIT 4
Muscle Physiology 93
UNIT 5
Feeding Systems and Digestion 125
COURSE NAME: COMPARATIVE ANIMAL PHYSIOLOGY COURSE CODE: MZO-003
AND BIOCHEMISTRY

Course Design Committee

Prof. Sujatha Varma Prof. Bharati Chauhan Prof. Amrita Nigam (Retd.)
Former Director, School of Department of Zoology School of Sciences, IGNOU
Sciences, IGNOU, Maidan Garhi University of Rajasthan Maidan Garhi, New Delhi-110068
New Delhi-110068 Jaipur-302004
Prof. Neera Kapoor
Prof. Neeta Sehgal Prof. Devinder Kaur Kochar School of Sciences, IGNOU
Department of Zoology Department of Zoology Maidan Garhi, New Delhi-110068
University of Delhi, Delhi-110007 Punjab Agricultural University
Ludhiana-141004 Dr. Siya Ram
Prof. Sarita Kumar School of Sciences, IGNOU
Department of Zoology, Acharya Prof. Varsha Wankhade Maidan Garhi, New Delhi-110068
Narender Dev College, University Department of Zoology
of Delhi, Delhi-110007 Savitribai Phule University Dr. Ravi Rajwanshi
Pune-411007 School of Sciences, IGNOU
Prof. S. Dinakaran Maidan Garhi, New Delhi-110068
Department of Zoology Prof. Sarita Sachdeva
Madurai Kamaraj University Department of Biotechnology
Tamil Nadu-625021 Manav Rachna International
University, Sector-43 Faridabad
Haryana-121004

Block Preparation Team

Prof. Neera Kapoor (Units 1, 2, 3, 4 and 5)
School of Sciences, IGNOU
Maidan Garhi, New Delhi-110068

Programme Coordinators : Prof. Neera Kapoor and Dr. Siya Ram

Course Coordinator : Prof. Neera Kapoor

Course Editor : Dr. Anjali Nawani

Sri Venkateswara College, University of Delhi
Delhi-110021

Production : Mr. Hemant Kumar

SO(P), MPDD, IGNOU

Acknowledgement:
• Prof. Neera Kapoor and Mr. Ajit Kumar, Suggestions for figures and Cover Design.
• Mr. Vikas Kumar, JAT for word processing and CRC preparation.
• Some of the units material has been taken from IGNOU course material.
August, 2023
Indira Gandhi National Open University, 2023
ISBN: ..................................
All rights reserved. No part of this work may be reproduced in any form, by mimeograph or any other
means, without permission in writing from Indira Gandhi National Open University.
Further information on Indira Gandhi National Open University courses may be obtained from the
University’s office at Maidan Garhi, New Delhi-110 068 or IGNOU website www.ignou.ac.in.
Printed and published on behalf of Indira Gandhi National Open University, New Delhi by the Registrar,
MPDD, IGNOU.
Printed at:
 BLOCK 1: FUNDAMENTALS OF ANIMAL
PHYSIOLOGY
Human Physiology is about how a human body functions. It describes the function performed
by animals in physical and chemical terms from the sub-cellular level to the level of an
integrated individual. However, none of these functions work in isolation. They are
interconnected and interdependent with common principles underlying each function at the
cellular and molecular levels in all animals and humans. At the cellular and sub cellular level
we find that certain principles keep on occurring throughout the animal kingdom.

Block 1 consists of five units, Block 1 “Circulatory System” speaks about the general plan of
the cardiovascular system (CVS) and principles associated with the circulation blood. You
will also study about the response of CVS to extreme conditions. The unit explains about
single and double systems of circulation. Unit further speaks about origin of heart beat, blood
pressure and their regulation.

In Unit 2, you will learn about processes involved in transport and exchange of gases and the
role of respiration in maintaining acid-base balance and blood pH. You will appreciate the
interrelationship of respiration and circulation mechanisms of human body. The unit reviews
the transport of oxygen and carbon dioxide in the blood and transfer of these gases between
blood and tissues.

In Unit 3 “Osmoregulation and Excretion” you will be amazed to know the anatomical
diversity and functional principles of excretory organs in different groups of animals and
appreciate the adaptive principles of osmoregulators and osmoconfomers to the
environment. Patterns of nitrogen exertion are discussed in the Unit. Osmoregulation in
aquatic and terrestrial environments are also explained. Extra renal osmoregulatory organs
that play important role in metabolism and are also dealt with.

Unit 4 entitled “Muscle Physiology” speaks about the events involved in the neural control of
skeletal muscle contraction and relaxation. Muscle contraction energetics are also discussed
in the Unit. Various types of muscles, their structure, properties and control are explained in
this Unit.

All the physiological processes in the body require energy which further needs nutrients
which are supplied by food. Various feeding strategies, process of digestion and absorption
of food are explained in the last unit of this block “Feeding Systems and Digestion”. Effects of
starvation are also briefly explained.

Objectives
After studying this course, you should be able to:

• describe the structure of the human heart, cardiac cycle and calculate the cardiac
output,

• discuss how cardiovascular system responds to extreme conditions,

• explain the process of gas transport between the blood and tissues in mammals,

• appreciate how respiratory system regulates blood pH and maintain acid-base balance, 7
 • describe the anatomical diversity and functional principles of excretory organs in
different groups of animals,

• identify the components of the neuromuscular junction, and summarize the events
involved in the neural control of skeletal muscle contraction and relaxation,

• describe the digestion process in humans as an example of mammalian digestion, and

• explain energy metabolism in animals in relation to oxygen consumption.

8
 UNIT 1
CIRCULATORY SYSTEM

Structure
1.1 Introduction 1.4 Peripheral Blood Circulation

Objectives Arteries

1.2 General Plan of Circulatory Veins

System
Capillaries
Open Circulation
Blood Pressure and its
Closed Circulation Regulation

1.3 Mammalian Heart 1.5 Cardiovascular Response to

Extreme Conditions
Location of the Heart
Exercise
Layers in the Heart
Diving
Internal Structure of the Heart
Hemorrhage
Blood Flow in the Heart
1.6 Summary
Initiation of Heartbeat and the
Cardiac Cycle 1.7 Terminal Questions

Cardiac Output 1.8 Answer

1.1 INTRODUCTION
You have learnt in your undergraduate course on Animal Physiology that for
cellular respiration to take place there must be a steady supply of nutrients
and oxygen and continual removal of carbon dioxide. In single celled Blood has RBCs,
WBCs, platelets, and
organisms, the supply of nutrients and removal of waste products is easier as
a fluid called plasma,
their cell surface is in direct contact with their environment which facilitates whereas lymph has
easy diffusion. In small organisms (diameter < 1mm) gas exchange can take WBCs and watery
place by simple diffusion across the cell membrane. Such small organisms do fluid. Blood flows
not need any internal system of transport, However, most multicellular though blood vessels
organisms (except sponges, cnidarians and flatworms) require a system for and lymph through
lymphatic vessels.
transport of nutrients, waste products and respiratory gases. Such a transport
system is provided by the cardiovascular system or the circulatory system.
The body fluids which transport the nutrients, waste products and respiratory
gases are blood and lymph.
Block 1 Fundamentals of Animal Physiology
In this unit, we will study about general plan of the cardiovascular system
(heart as a pump with its network of arteries, veins and capillaries which are
connected vessels) and the principles associated with the circulation of blood
will also be discussed. You will also study how the cardiovascular system
responds under extreme conditions. The cardiovascular system of humans
which is best known is taken as a representative model and discussed in detail
in this unit.

All circulatory systems, however, comprise the following basic parts, which
have similar functions in different animals:

1. A main propulsive organ, usually a heart, which forces blood around in

the body

2. An arterial system, which can act both to distribute blood and as a

pressure reservoir

3. Capillaries, in which transfer of materials occurs between blood and

tissues

4. A venous system, which acts as a blood (volume) reservoir and as a

system for returning blood to the heart

The arteries, capillaries, and veins constitute the peripheral circulation.

Objectives
After study this Unit, you should be able to:

 describe the difference between open and closed circulatory system,

 describe the structure of the human heart, cardiac cycle and calculate
the cardiac output,

 explain the flow of blood and blood pressure in arteries and veins, its
regulation,

 describe the process of microcirculation and exchange of fluid across

capillary beds, and

 discuss how cardiovascular system responds to extreme conditions.

1.2 GENERAL PLAN OF THE CIRCULATORY

SYSTEM
The movement of blood through the body results from any or all of the
following mechanisms:
• Forces imparted by rhythmic contractions of the heart
• Elastic recoil of arteries following filling by cardiac contraction
• Squeezing of blood vessels during body movements
• Peristaltic contractions of smooth muscle surrounding blood vessels
The relative importance of each of these mechanisms in generating flow varies
10 among animals. In vertebrates, the heart plays the major role in blood
Unit 1 Circulatory System
circulation; in arthropods, movements of the limbs and contractions of the
dorsal heart are equally important in generating blood flow; in the giant
earthworm, Megascolides australis, peristaltic contractions of the dorsal vessel
are responsible for moving blood in an anterior direction and filling the lateral
hearts, which pump blood into the ventral vessel for distribution to the body
(Fig. 1.1b).This worm, which can be up to 6 m in length, is divided into
segments separated by membranous structures (septa).Tracer studies have
shown that the anterior 13 segments, each of which contains two lateral
hearts, have a rapid circulation, but the remaining segments, which lack lateral
hearts, have a very sluggish circulation. Because of the peristaltic contractions
of the dorsal vessel, the blood pressure is considerably higher in the dorsal
vessel than in the ventral vessel. In all animals, valves and/or septa determine
the direction of flow, and smooth muscle surrounding blood vessels alters
vessel diameter, thereby regulating the amount of blood that flows through a
particular pathway and controlling the distribution of blood within the body.

1.2.1 Open Circulation

Many invertebrates have an open circulation, that is, a system in which blood
pumped by the heart empties via an artery into an open fluid space, the
hemocoel, which lies between the ectoderm and endoderm. The fluid
contained within the hemocoel, referred to as hemolymph, or blood, is not
circulated through capillaries but bathes the tissues directly. The hemocoel is
often large and may constitute 20%-40% of body volume. In some crabs, for
instance, blood volume is about 30% of body volume. Open circulatory
systems have low pressures, with arterial pressures seldom exceeding 0.6-
1.3 kilopascals (kPa),or 4.5-9.7 rnm Hg (1kPa = 7.5 rnrn Hg). Higher
pressures have been recorded in portions of the open circulation of the
terrestrial snail Helix, but these are exceptional. In snails, these high pressures
are generated by contractions of the heart, whereas in some bivalve molluscs
high pressures in the foot are generated by contractions of surrounding
muscles rather than of the heart.

Animals with an open circulation generally have a limited ability to alter the
velocity and distribution of blood flow. As a result, in bivalve molluscs and
other animals that have an open circulation and use blood for gas transport,
changes in oxygen uptake are usually slow and maximal rates of oxygen
transfer low per unit weight. Nonetheless, such animals exert some control
over both the flow and distribution of hemolymph; moreover, the blood is
distributed throughout the tissues in many small channels in animals with an
open circulation. In the absence of such features, even moderate rates of
oxygen consumption would be impossible because of the large diffusion
distances for oxygen between the hemolymph and the active tissue.

Insects have an open circulation but do not depend on it for oxygen

transport and thus can achieve high rates of oxygen transfer. They have
evolved a tracheal system in which direct gas transport to tissues occurs
through air-filled tubes that bypass the blood, which plays a negligible role in
oxygen transport. Consequently, although insects have an open circulation,
they have a large capacity for aerobic metabolism (Fig. 1.1a)

1.2.2 Closed Circulation

In a closed circulation, blood flows in a continuous circuit of tubes from arteries
to veins through capillaries. All vertebrates and some invertebrates, such as 11
Block 1 Fundamentals of Animal Physiology
cephalopods (octopuses, squids), earthworm, have this type of circulation (Fig.
1.1b). In general, there is a more complete separation of functions in closed
circulatory systems than in open ones. The blood volume in the closed
circulation of vertebrates typically is about 5%- 10% of body volume, much
smaller than that of open-circulation invertebrates. The extracellular volume
in vertebrates, expressed as a percentage of body volume is similar to
the hemocoel volume in invertebrates. The closed circulatory system of
vertebrates is a specialized portion of their extracellular space.

In a closed circulation, the heart is the main propulsive organ, pumping blood
into the arterial system and maintaining a high blood pressure in the arteries.
The arterial system, in turn, acts as a pressure reservoir forcing blood through
the capillaries. The capillary walls are thin, thus allowing high rates of transfer
of material between blood and tissues by diffusion, transport, or filtration. Each
tissue has many capillaries, so that each cell is no more than two or three cells
away from a capillary. Capillary networks are in parallel, allowing fine control
of blood distribution and, therefore, oxygen delivery to tissues. Animals with a
closed circulation can increase oxygen delivery to a tissue very rapidly.
For this reason, squid, unlike many other invertebrates, can swim rapidly
and maintain high rates of oxygen uptake; that is, their closed circulation
permits sufficient flow and efficient distribution of hemolymph to the muscles to
support short bursts of high- level activity.

Also, the blood is under sufficiently high pressure in a closed circulation to

permit the ultra filtration of blood in the tissues, especially the kidneys. Ultra
filtration refers to the separation of an ultra filtrate, devoid of colloidal
particles, from plasma by filtration though a semi permeable membrane
(capillary wall) using pressure (blood pressure) to force the fluid through
the membrane. Ultra filtration occurs in most vertebrate kidneys, resulting in
the net movement of a protein-free plasma from the blood into the kidney
tubule. In general, all capillary walls are permeable and as pressures are high,
so fluid slowly filters across the walls and into the space between cells. A
lymphatic system has evolved in conjunction with the high-pressure, closed
circulatory system of vertebrates to recover fluid lost to tissues from the blood.
The extent of filtration depends largely on the blood pressure and the
permeability of the capillary wall. Filtration across capillary walls can be
decreased either by a reduction in the permeability of the capillary walls or in
the pressure of the blood. For example, the vessels in some tissues have less
permeable walls than those found in other tissues. And in the liver and lung,
where permeability is high for other reasons, pressures are lower than those in
the rest of the body.

Different pressures can be maintained in the systemic (body) and pulmonary

(lung) circulations of mammals because the mammalian circulatory system is
equipped with a completely divided heart. The right side of the heart pumps
blood in the pulmonary circulation, and the left side pumps blood in the
systemic circulation. This means, however, that the flows in the pulmonary
and systemic circuits must be equal because blood returning from the lung is
pumped around the body. In other vertebrates the heart is not completely
12 divided and flow to the lung can be varied independently of body blood flow.
Unit 1 Circulatory System
The venous system collects blood from the capillaries and delivers it to the
heart via the veins, which are typically low-pressure, compliant structures in
which large changes in volume have little effect on venous pressure. Thus, the
venous system contains most of the blood and acts as a large-volume
reservoir. Blood donors give blood from this reservoir, and since there is little
change in pressure as the venous volume decreases, the volumes and flow in
other regions of the circulation are not markedly altered.

In the next section we will study the structure and function of the human heart
as the example of a highly developed pump which plays a vital role in
vertebrate circulation.

Fig.1.1: Diagramatic representation of a) Open circulatory system seen in

invertebrates like arthropods, molluscs. The haemolymph returns to the
heart by ostia; b) closed circulatory system seen in all vertebrates and
some invertebrates like annelids, echinoderms and cephalopods.

SAQ 1
Chose the correct answer.
a) A closed circulatory system is required by all animals that have
i) high metabolism
ii) interstitial fluids
iii) intracellular digestion
iv) low energy requirement
b) In the open circulatory system haemolymph is not segregated from:
i) interstitial fluid
ii) cytoplasm
iii) digestive system
iv) urine
13
Block 1 Fundamentals of Animal Physiology
1.3 MAMMALIAN HEART
You all must have felt or heard your heart beating. Did you know that on an
average an individual’s heart beats at about 80 beats per minute that is
4,800 beats per hour, 115, 200 beats per day, and more than 42 million
beats per year, which calculates to roughly 3 billion beats if you live to the
age of 72. This signifies the vital role of the heart in our survival. To
comprehend the significance of the heart, you must remember that our body is
made up of over 10 trillion cells and each cell has essential needs: oxygen,
nutrients, fluid environment and removal of wastes. To ensure this need for
each and every cell of the body a very efficient transport system is needed.
You already know that O2 is transported through blood which flows in a system
of arteries, veins and capillaries. To maintain the flow of blood in the arteries,
veins and capillaries a pump is needed. The heart acts as a muscular pump
which maintains the rhythmic blood flow. You have already studied in your
graduation course the structure of the heart of various vertebrates. In the
present unit we will be describing in greater detail the structure and function of
the mammalian heart in particular the human heart.

1.3.1 Location of the Heart

Figures 1.2 (a) and (b) show the position of a human heart in the chest or
thorax. The heart lies in the upper left part of the chest under the sternum
(breast bone) and is protected by the ribcage. It lies in between the lungs
forming a cardiac notch in the left lung.

Fig. 1.2: Position of a human (mammal) heart in the thorax: a) heart is protected
by the rib cage; b) heart lies in between the lungs forming a cardiac
14 notch in the left lung.
Unit 1 Circulatory System
1.3.2 Layers of the Heart
The mammalian heart as we mentioned earlier is a muscular organ and is
located within the pericardial space or cavity which is within the mediastinum
(central compartment of the thoracic cavity).The heart is covered by a tough
fibrous sheath, the pericardium which separates the heart from the rest of the
organs of the thoracic region. The pericardium has two layers– an inner
visceral layer that covers the heart from outside and external to it a parietal
layer that forms a sac around the outside of pericardial cavity (see Fig. 1.3).
The pericardium has a pericardial cavity which contains fluid that lubricates the
heart and permits it to contract with minimal friction.

Fig. 1.3: Layers of heart - epicardium, myocardium and endocardium

surrounded by the pericardial cavity that contains the pericardial fluid
and enclosed in the outmost layer of pericardium.

The wall of the heart is made up of three layers (Fig 1.3) and its thickness
varies in different parts of the heart. Just internal to the pericardial cavity is
the: i) epicardium, also called the visceral layer of pericardium. It forms the
outermost protective layer of the heart; ii) inner to the epicardium, is the
middle layer of the heart called the myocardium which consists of cardiac
muscles. The myocardium is the muscular part of the heart and makes up for
most of the thickness and mass of heart wall. It is responsible for the pumping
of blood; iii) inner to the myocardium is the innermost layer of the heart which
is called the endocardium. The endocardium is a thin layer which is made up
of an endothelial layer that lines the internal surface of the heart.

1.3.3 Internal Structure of the Heart

The mammalian heart has four chambers, which consist of two upper
chambers and two lower chambers. The upper chambers are called atria
(singular atrium) or auricles and the lower chambers are called
ventricles. Each half of the heart as shown in Figure 1.4 is made up of an
upper, thin walled atrium and a posterior, thick walled ventricle. The thin 15
Block 1 Fundamentals of Animal Physiology
walled atrium serves as a filling reservoir for blood and the thick walled
ventricle pushes the blood into circulation. The right atrium and the right
ventricle are connected by a tricuspid valve and the left atrium and the left
ventricle are connected by a bicuspid valve also called the mitral valve.
These two sets of valves are collectively called the atrioventricular valves.
Another set of valves called the semilunar valves are also present in the heart.
They are present at the region where the pulmonary artery leaves the right
ventricle (pulmonary semilunar valves) and at the region where the aorta
leaves the left ventricle (aortic semilunar valves). Both the atrioventricular
valves and the semilunar valves prevent the backflow of blood.
Cardiac muscle cells
are rectangular
shaped cells that are
connected by regions
called intercalated
discs. Intercalated
discs contain gap
junctions and
desmosomes. The
gap junctions, are
protein-lined tunnels,
which allow direct
transmission of the
depolarizing current Tricuspid valve
from cell to cell,
across the chambers
of the heart, so that
the cells contract in
unison. The
desmosomes hold the
cardiac muscle cells
together during
contraction.
Depolarizations of
myocardial cells Fig. 1.4: Internal structure of heart showing the valves.
occur, when they are
stimulated by the 1.3.4 Blood Flow in the Heart
electrical system of
the heart which You are already familiar of the fact that oxygenated blood from the heart flows
results in their in arteries and releases the oxygen bound to haemoglobin in the tissues and
contraction. Cardiac collects the carbon dioxide released from the tissues and so as a result
cells at rest are becomes deoxygenated as it returns to the heart. This is known as systemic
considered polarized. circulation.
Repolarization is the
process in which the Deoxygenated blood enters the heart from the systemic circulation via the
cardiac cells return to large veins that bring blood back from the top and bottom of the body namely
the resting state
via the superior vena cava and the inferior vena cava respectively. The
before they can be
electrically stimulated
deoxygenated blood (Blue in colour in Fig. 1.5) is received in the right atrium
again. which is a transit chamber from where the blood goes through the tricuspid
valve to the right ventricle. The valves make sure that blood only goes one
way i.e., from right atrium to right ventricle and prevents backflow. The right
ventricle (pump-1) has muscular walls that constrict and gently push the
deoxygenated blood through the tricuspid pulmonary semilunar valves into the
pulmonary artery. The pulmonary artery is the only artery that carries
16 deoxygenated blood. The tricuspid pulmonary semilunar valves ensure that
Unit 1 Circulatory System
the blood only goes one way. The pulmonary artery carries the deoxygenated
blood to the lungs where oxygen from an area of high oxygen concentration
(alveoli) diffuses to an area of low concentration (blood) so that the blood
becomes oxygenated. This oxygenated blood (red in colour in Fig. 1.5) then
returns through pulmonary vein to the left atrium of the heart. This part of
circulation is known as pulmonary circulation. Pulmonary vein is the only
vein that carries oxygenated blood.

Fig.1.5: a) Cross-section of the human heart, with the right ventricle labeled
“pump 1” and the left ventricle labeled “pump 2”. (Adapted from
Wikipedia image from Zoofari); b) Systemic and Pulmonary circulation.

The left atrium just like the right atrium can be thought of as a transit chamber
from where the blood passes through the bicuspid valve (mitral valve) to the
left ventricle. The left ventricle is stronger, thicker, and more muscular than the
right ventricle. As a result, the left ventricle (pump-2) forcefully pushes the 17
Block 1 Fundamentals of Animal Physiology
blood through the aortic semilunar valves into the aorta for the systemic
circulation. The muscle wall of the left ventricle is thicker than that of the right
ventricle because the left ventricle has to apply greater force for pumping the
blood through the systemic circulation. The aorta forks and the blood is divided
between major arteries which supply the upper and lower body. The major
artery divides into smaller arteries which further divide into arterioles and then
into capillaries of the body in order to supply oxygen to trillions of cells and to
take up their carbon dioxide. The relatively deoxygenated blood then travels to
the venules, which are the smallest types of veins. The venules coalesce into
veins, which are larger vessels. The veins then join to form the inferior and
superior venae cavae which finally enter back into the right atrium carrying
deoxygenated blood where the process began (Fig. 1.5 b).

The contraction of the heart is known as systole and its relaxation is known as
diastole. The rhythmic contraction and relaxation of the heart makes up
one heart beat or one cardiac cycle. Let us now see how this heart beat is
generated.

1.3.5 Initiation of Heartbeat and the Cardiac Cycle

Do you know that the heart pumps or beats continuously because of a small
electrical current that is generated by the cardiac conduction system of the
heart without any external stimulus? The normal human heart beats
continuously due to its own inherent and rhythmical electrical activity.

The electrical activity begins in a region of the heart known as the pacemaker.
The cells of the pacemaker are capable of spontaneous activity and maybe
either composed of neurons or muscle cells. If the heartbeat is initiated in a
pacemaker consisting of neurons, then the pacemaker is called a neurogenic
pacemaker as found in animals with open circulation and if the heartbeat is
initiated in a pacemaker composed of muscle cells then the pacemaker is
known as a myogenic pacemaker as seen in animals with closed circulatory
system. Based on the type of pacemaker, hearts are classified as neurogenic
or myogenic.

In vertebrates the heart is myogenic which means that the source of

stimulation for the beating of the heart is a network of specialized self-
excitable cardiac muscle cells called the autorhythmic cells. Proof of
autorhythmicity of cardiac muscle cells can be observed when we see that
the heart in a developing chick embryo begins to beat even before any nerve
has reached it and also when heart muscle cells grown in a culture contract
rhythmically without any external stimulus. These autorhythmic cells have two
important functions: (i) They act as you are aware as a pacemaker and set the
rhythm for the entire heart, and (ii) they form the conduction system, which is
the route for propagating action potentials or electric stimulus throughout the
heart muscle for it to contract or relax.

The conduction system ensures that the four heart chambers namely the two
atria and two ventricles become stimulated to function as effective pumps. The
main components of the cardiac conduction system as shown in Fig 1.6 a and
b are the sinoatrial node (SA node), atrioventricular node (AV node),
18 bundle of His, left and right bundle of His branches, and Purkinje fibers.
Unit 1 Circulatory System
Sinoatrial node (SA node) is the pacemaker of the heart which initiates the
electric impulse (Fig. 1.6 a and b). The spontaneous electric impulse or wave
of excitation produced by the SA node spreads across both atria via gap
junctions, causing both atria to contract (atrial systole). Contraction of atria
causes the flow of blood from the atria into the ventricle. After the electrical
impulses spread across the atria, they converge at the atrioventricular node
(AV node) - located within the atrioventricular septum. The atrioventricular
septum is the membranous part of the interventricular septum, and is present,
near the opening of the coronary sinus which is located along the heart's
posterior (rear) surface between the left ventricle and left atrium. AV node after
a brief delay sends the electrical impulse forward which ensures that the atria
have enough time to fully eject blood into the ventricles before ventricular
systole. The delay between atrial and ventricular excitation corresponds to the
PR interval of the electrocardiogram and it is 0.12-0.20 seconds or 120-200
milliseconds (ms) in humans. The impulse from the AV node is conducted
through the bundle of His (atrioventricular bundle) to the Purkinje fibres of
the ventricles.

Fig. 1.6: a) The individual components of the human heart (mammal) in the
electrical conduction pathway; b) The pathway of the conducting
system of the human heart. 19
Block 1 Fundamentals of Animal Physiology
From the AV node the impulse descends down the membranous part of the
interventricular septum, before dividing into two main bundles: Right bundle
of His branch which conducts the impulse to the Purkinje fibres of the right
ventricle and Left bundle of His branch which conducts the impulse to the
Purkinje fibres of the left ventricle.

Purkinje fibres are a network of specialised cells which are connected together
by desmosomes and gap junctions, but not by intercalated discs. The
Purkinje cells have extensive gap junctions that help in rapid transmission of
cardiac action potentials from the atrioventricular bundle to the myocardium of
the ventricles. This rapid conduction allows coordinated ventricular
contraction (ventricular systole) and blood is moved from the right and left
ventricles to the pulmonary artery and aorta respectively. This arrangement
allows the contraction to begin at the tip of the ventricles and to spread
upwards in order to squeeze out blood in the most efficient way. Thus, cardiac
conduction is the driving force behind the cardiac cycle. This cycle is the
sequence of events of: systole and diastole that occurs when the heart
beats. Cardiac muscles contracts when electric impulse reaches them.
Contraction of heart muscle means that the heart pushes in; making the heart
chambers smaller which results in pushing the blood out of the heart and into
the blood vessels. Once the heart contracts and pushes out the blood, its
muscles then relax, due to which the chambers of the heart get bigger and the
blood comes back into the heart and fills the chambers. Therefore, the
sequence of events are: atrial systole → ventricular systole → cardiac diastole
(Fig. 1.7). When this sequence of events happens one time, it is called
a cardiac cycle.

Fig. 1.7: Cardiac Cycle: a) During diastole the heart relaxes and blood is sucked
into the heart. The semilunar valves are closed; b) while during systole
heart contracts and blood is prevented from going back to atria by the
atrioventricular valves. Blood is pumped out from the ventricles
through the open semilunar valves.

1.3.6 Cardiac Output

Now that you know how blood is pumped out by the heart, let us see how
much blood is pumped out with each heartbeat and whether it is related to the
20 oxygen requirements of the animal. Heartbeat frequency/pulse rate/heart rate
Unit 1 Circulatory System
is usually given as the number of heart beats per minute. The pulse rate for an
adult human being is 72 per minute at rest. The volume of blood ejected from
the heart per minute is called cardiac output. In hearts with complete
separation of ventricles the cardiac output refers to output of only one
side of the ventricle and not to that of both ventricles.

The volume of blood ejected from the left ventricle in one contraction or with
each beat is called stroke volume (SV). So we can calculate cardiac output
as the product of the heart rate, which is the number of beats per minute, and
the stroke volume, which is amount of blood pumped by the left ventricle per
beat.

CO = HR × SV

(Cardiac out put = Heart rate × stroke volume)

The cardiac output is usually expressed in liters/minute. For example for

someone weighing about 70 kg, whose stroke volume at rest is about 70
ml/beat and the heart rate is 72 beats/min the cardiac output (70×72) would be
a little more than 5 liters/minute.

Heart rate is affected by autonomous innervations, hormones, age and the

fitness level of an individual. A nerve branch of parasympathetic origin slows
the heart rate by releasing acetylcholine while another nerve of sympathetic
origin releases epinephrine and norepinephnine at the pacemaker region to
increase the heart rate. Apart from these the pacemaker is influenced by
temperature and electrolytes.

SAQ 2
Tick mark the correct answer.

i) Stroke volume is the:

a) volume of blood pumped from the heart

b) volume of blood pumped into the heart

c) volume of blood pumped by the left ventricle during a single

heartbeat

d) number of heart beats per minute

ii) Cardiac output equals:

a) stroke volume multiplied by heart rate

b) stroke volume divided by heart rate

c) stroke volume added to the heart rate

b) Fill in the blanks with appropriate words from the text.

i) Sinoatrial node is the .................... of the heart. The electrical

impulse starts at this small piece of muscle and spreads
across both the .................... .
ii) In the heart the impulse from the AV node is conducted
through the.................... to the.................... of the ventricles.
21
Block 1 Fundamentals of Animal Physiology
1.4 PERIPHERAL BLOOD CIRCULATION
There are two main types of blood vessels in the circulatory system - arteries
and veins with characteristic differences among them. Arteries have relatively
thick walls that consist of heavy, strong layers of elastic fibres and smooth
muscles. As the arteries branch, their diameter becomes smaller and the
relative amount of muscle tissue increases in proportion to the elastic tissue.
The smallest units of the arteries are the capillaries which are single cell thick
and play a major role in all the exchanges between tissue and blood which
take place through the walls of the capillaries. The veins also contain smooth
muscles and fibres.
The walls of the arteries and veins are largely composed of living cells and
need to be supplied with nutrition. The walls of the arteries and veins also
produce wastes. For this reason these blood vessels have their own blood
supply in the form of a capillary network known as “vasa vasorum” or
“vessels within vessels”. Minute nerves are also present within the walls of
the capillaries of the vasa vasorum which control and coordinate the
constriction and relaxation of these capillaries.

1.4.1 Arteries
The arteries except for the pulmonary artery carry the oxygenated blood
from the heart to various cells and tissues. Figure 1.8 a shows the
structure of arteries and the different layers of the vessel wall.
The arteries have four main functions:
1. to act as a conduit for oxygenated blood(except for pulmonary artery
which carries deoxygenated blood) between heart and capillaries,
2. to act as a pressure reservoir for forcing blood into small diameter
arterioles,
3. to produce a more or less even flow of blood by means of the capillaries,
4. to control distribution of blood to different capillary networks via selective
constriction of the terminal branches.
There is a precise control on arterial blood pressure. The nature of the arterial
wall and the volume of blood pumped into the arteries determines the
pressure. If any of these are changed, the pressure will also change. Normally,
arterial blood pressure varies very little as the cardiac output and the capillary
flow are evenly matched.
The elastic properties of arterial walls vary. Close to the heart the arteries are
elastic and dampen (reduce) the oscillations in pressure and blood flow,
generated by the contractions of the heart.

1.4.2 Veins
The veins bring back the deoxygenated blood (except for pulmonary vein
which carries oxygenated blood) to the heart from the capillaries via the
venules. They form a large volume, low pressure system. The vessels have a
larger internal diameter and pocket valves (Fig.1.8b). In mammals 50% of the
total blood volume is present in the veins and the pressure in the veins is
between 8 and 10 mm Hg. If there is any blood loss, the venous volume is
decreased and not the arterial volume. So that arterial blood pressure and
22 capillary blood flow is maintained.
Unit 1 Circulatory System
The flow of blood in veins is affected by several factors. Pressure exerted by
the diaphragm on the gut and the activity of the limbs both help to squeeze the
veins of those regions. This squeezing and also the action of the pocket valves
(that prevent back flow) help the flow of blood towards the heart. Breathing in
mammals also helps in drawing the blood from the veins in the head and
abdominal cavity.
Smooth muscles in veins also help in regulating blood supply in the venous
system. When a person changes his position from sitting to standing the
change in the relative position of heart and brain with respect to gravity
activates the nerve fibres that are present in the veins of the limb. This causes
a contraction of the smooth muscles of the veins and results in the pooled
blood to be, redistributed.

Fig. 1.8: Characteristic features of a) artery; b) vein and c) capillary.

1.4.3 Capillaries
We had said earlier that most tissues have such an extensive network of
capillaries that any single cell is hardly 2-3 cells away from any capillary. The
small terminal arteries subdivide to form arterioles which divide to form
metarterioles and then capillaries (Fig. 1.8 c). The capillaries join the venules
which contain the venous blood of the capillaries. The smooth muscles of the
arterioles become discontinuous in the metarterioles and end in a muscle ring,
the precapillary sphincter that controls the blood supply to each capillary
bed. Through the precapillary sphincters, fresh blood supply to the capillary
bed can be bypassed altogether and blood can be diverted to areas of greater
demand. All capillaries of an animal have the potential to hold 14% of the total
blood volume. However, only 30% to 50% of all capillaries are open at a time
and thus only 5-7% of the total volume is contained in them. Several venules
join together to form the veins. 23
Block 1 Fundamentals of Animal Physiology
Capillaries are made up of a single layer of endothelial cells (Fig. 1.8 c)
surrounded by a basal membrane. The walls are thin and fragile but because
of their small diameter can resist stretching in response to capillary blood
pressure. Water and dissolved substances of small molecular weight (gases,
salts, sugars, amino acids etc.) can diffuse easily in and out of the capillaries
(Fig. 1.9). In addition, fluid is forced out through the capillary walls.
Substances of molecular weight more than 70,000 dalton (mostly proteins) do
not pass in or out of the capillary walls. These proteins of molecular weight
more than 70,000 dalton when present in the capillaries exert an osmotic
pressure called the colloidal osmotic pressure, which tends to draw water
back into the capillary from the surrounding tissue fluid (reabsorption) (Fig.
1.9). Another force, known as hydrostatic pressure of blood tends to push the
water across the endothelial cell layer and so out of the capillaries (filtration)
(Fig. 1.9). When the hydrostatic pressure within the capillary exceeds colloidal
osmotic pressure, fluid is passed out through the capillary wall into the tissues.
However, when the hydrostatic pressure in the capillary falls below the
colloidal osmotic pressure then fluid is drawn in from the tissues. At the arterial
end of the capillary the hydrostatic pressure is higher than the colloidal
osmotic pressure while at the venous end it is often lower. Therefore, fluid is
filtered out at the arterial end and taken at the venous end (see Fig.1.9).

The amount of fluid which is forced out and the amount of fluid re-entering the
capillary varies greatly. Usually outflow exceeds inflow and excess fluid
remains in the interstitial spaces. This as you already know forms the lymph.

Fig 1.17: a) Valves in

veins open for one
way flow of blood
vein towards the
heart; b) Valves of
Fig. 1.9: Fluid exchange across the capillary wall, showing differences in blood
vein closed to
prevent backflow of pressure and colloidal osmotic pressure. Net filtration occurs near the
the blood. arterial end of the capillary since the capillary’s hydrostatic pressure
(35mmHg) is greater than the blood colloidal osmotic pressure
(25mmHg). There is no net movement of fluid near the midpoint of the
capillary. At the venous end, fluid (water) is reabsorbed as the capillary
hydrostatic pressure is less (18mm Hg) than the colloidal blood
pressure (25mm Hg).

1.4.4 Blood Pressure and Its Regulation

We are all familiar with the term high blood pressure. What exactly does this
term mean? By this term we really mean only arterial blood pressure. When
systemic blood pressure is measured it is recorded as a ratio of systolic and
diastolic pressure. In a normal human adult individual blood pressure is
120/80mm Hg. The higher value of 120mmHg is the arterial pressure resulting
from the ejection of blood during ventricular contraction and is termed systolic
pressure. The lower value of 80mmHg is the arterial blood pressure during the
24 relaxation of the heart and is termed as diastolic blood pressure. The
Unit 1 Circulatory System
difference between the two is referred to as pulse pressure which is normally
25 % of the systolic pressure. Blood pressure falls as blood flows from the
aorta to the vena cava. The greatest pressure drop takes place in the smallest
arterioles which by changing their diameter can regulate the flow of blood to
various body organs. Persistently lower values of pulse pressure could
indicate lower stroke volume and possibly congestive heart failure. Higher
values can temporarily be attained during exercise but may indicate a
diseased condition.

The mechanisms that regulate systemic blood pressure may be divided into
two types: intrinsic mechanisms (long term regulation) and nervous
mechanisms (short term regulation). The nervous mechanisms involve the
nervous system, and the intrinsic mechanisms do not require nerve impulses.
INTRINSIC MECHANISMS
The term intrinsic means “within.” Intrinsic mechanisms work because of the
The Frank Starling
internal characteristics of certain organs. The first such organ is the heart.
Law is the description
When venous return increases, cardiac muscle fibers are stretched, and the
of cardiac
ventricles pump more forcefully (Starling’s law). Thus, cardiac output and
hemodynamics as it
blood pressure increase. This is what happens during exercise, when a higher relates to myocyte
blood pressure is needed. When exercise ends and venous return decreases, stretch and
the heart pumps less forcefully, which helps return blood pressure to a normal contractility. The Law
resting level. states that the stroke
volume of the left
The second intrinsic mechanism involves the kidneys. When blood flow
ventricle will increase
through the kidneys decreases, the process of filtration decreases and less as the left ventricular
urine is formed. This decrease in urinary output preserves blood volume so volume increases due
that it does not decrease further. Following severe hemorrhage or any other to the myocyte stretch
type of dehydration, this is very important to maintain blood pressure. causing a more
forceful systolic
The kidneys are also involved in the renin- angiotensin mechanism. When
contraction.
blood pressure decreases, the kidneys secrete the enzyme renin, which
initiates a series of reactions that result in the formation of angiotensin II.
These reactions are depicted in Fig. 1.10. Angiotensin II causes
vasoconstriction and stimulates secretion of aldosterone by the adrenal cortex,
both of which will increase blood pressure.

Fig. 1.10: The renin-angiotensin mechanism. 25

Block 1 Fundamentals of Animal Physiology
NERVOUS MECHANISMS (Short term mechanisms)

The medulla and the autonomic nervous system are directly involved in the
regulation of blood pressure. The first of these nervous mechanisms concerns
the heart, stroke volume and cardiac output; this was described previously.

The second nervous mechanism involves peripheral resistance, that is, the
degree of constriction of the arteries and arterioles and, to a lesser extent, the
veins (see Fig. 1.11). The medulla contains the vasomotor center, which
consists of a vasoconstrictor area and a vasodilator area. The vasodilator area
may depress the vasoconstrictor area to bring about vasodilation, which will
decrease blood pressure. The vasoconstrictor area may bring about more
vasoconstriction by way of the sympathetic division of the autonomic nervous
system.

Sympathetic vasoconstrictor fibers innervate the smooth muscle of all arteries

and veins, and several impulses per second along these fibers maintain
normal vasoconstriction. More impulses per second bring about greater
vasoconstriction, and fewer impulses per second cause vasodilation. The
medulla receives the information to make such changes from the presso-
receptors in the carotid sinuses and the aortic sinus.

26 Fig. 1.11: Nervous mechanism that regulates blood pressure.

Unit 1 Circulatory System

SAQ 3
Select the four true statements:

a) Arteries generally have a larger diameter than veins.

b) Capillaries are made up of a single layer of endothelial cells surrounded

by a basal membrane.

c) The arteries near the heart are more elastic and dampen the oscillation
in blood flow.

d) Whole blood is more viscous than plasma because of the presence of

blood cells.

e) The maximum pressure during a heart beat is systolic pressure.

f) The maximum pressure during a heartbeat is known as diastolic

pressure.

1.5 CARDIOVASCULAR RESPONSE TO

EXTREME CONDITIONS
In the previous sections, you have studied the general organization of the
circulation and its regulation under usual conditions. The cardiovascular
system responds in characteristic ways during exercise, diving, and
hemorrhage to meet the physiological challenge of these extreme conditions
(Table 1.1).

1.5.1 Exercise
Regulation of the cardiovascular system during exercise is clearly a complex
process involving central neural control mechanisms, peripheral neural reflex
mechanisms (especially those involving skeletal muscle afferent fibers) and
local control.

During exercise, blood flow to skeletal muscle is increased in proportion to the

level of activity of the muscle. The increase in flow to a muscle may be as
much as twenty times; at the same time, transfer of oxygen from the blood to
muscle may increase threefold, resulting in a sixtyfold increase in oxygen
utilization by the muscle. At the same time, there is a reduction in flow to the
gut, kidney etc. and, at high levels of exercise, the skin. Cardiac output can
increase up to ten times above the resting level owing to large increases in
heart rate and small changes in stroke volume. Much of the increase in
cardiac output can be accounted for by a decrease in peripheral resistance to
about 50% of the resting value and by an increase in venous return to the
heart due to both the pumping action of skeletal muscles on veins and the
increase in breathing associated with exercise.

The increased sympathetic, but decreased parasympathetic activity in nerves

innervating the heart has the effect of increasing both heart rate and the force
of contraction, so as to maintain stroke volume at a relatively constant level. In 27
Block 1 Fundamentals of Animal Physiology
fact, stroke volume increases by about 1.5 times during exercise in mammals,
despite the large increase in heart rate and the associated reduced time
available for filling and emptying. Following sympathetic stimulation, however,
blood is ejected more rapidly from the ventricles with each beat, maintaining
stroke volume at higher heart rates. The relative role of changes in stroke
volume and heart rate in generating the increase in cardiac output with
exercise varies among animals. In fish, for example, the changes in stroke
volume are much greater than the changes in heart rate, whereas in birds
there are very large changes in heart rate and little change in stroke volume
during exercise.

1.5.2 Diving
Many air-breathing vertebrates can remain submerged for prolonged periods.
During submersion for any period, all air-breathing vertebrates stop breathing,
so the animal must rely on available oxygen stores in the blood. The
cardiovascular system is adjusted to meter out the limited oxygen store to
those organs-brain, heart, and some endocrine structures-that can least
withstand anoxia.

In mammals, but not in other vertebrates, stimulation of the facial receptors

that inhibit breathing cause a marked bradycardia. Although the initial
pressurization of the lung can lead to a transient increase in blood oxygen and
CO2, levels, the continued utilization of O2, during the dive results in a gradual
fall in blood O2, and rise in blood CO2, levels. This fall in blood O2, stimulates
the arterial chemoreceptors and, in the absence of lung stretch-receptor
activity, causes peripheral vasoconstriction and a reduction in heart rate and
cardiac output; thus blood flow to many tissues is reduced so as to maintain
flow to the brain, heart, and some endocrine organs. In some instances, blood
flow to muscle decreases, but this depends on the level of exercise associated
with the dive and the species.

It has been shown in the seal that the generation of the diving bradycardia
(slow heart rate) can involve some form of associative learning. In some
trained seals, bradycardia occurs before the onset of the dive and, therefore,
before the stimulation of any peripheral receptors. This psychogenic influence
on heart rate can have a marked effect on the change in heart rate during a
dive in many animals. In general, if heart rate is low before a dive, there may
be little or no change in heart rate during the dive. If the heart rate is high, then
there may be a marked bradycardia due to wetting the face and a decrease in
lung stretch-receptor activity.

The "water" receptors present in birds are not directly involved in the cardio-
vascular changes associated with submersion. A decrease in heart rate is not
observed either in submerged ducks breathing air through a tracheal cannula
or in submerged ducks following carotid body denervation. Thus, activation of
the "water" receptors causes suspension of breathing (apnea); the subsequent
drop in blood pO2 and pH and the rise in pCO2result in stimulation of
28 chemoreceptors, which then reflexly cause the cardiovascular changes.
Unit 1 Circulatory System
1.5.3 Hemorrhage
Normally stimulation of arterial and baroreceptors inhibits vasopressin release,
as well as sympathetic out flow to the peripheral circulation. Hemorrhage
reduces both venous and arterial blood pressure, reducing the discharge Baroreceptors are a
frequency of both a trial and arterial baroreceptors. This releases the type of
mechanoreceptors
baroreceptive inhibition of sympathetic outflow causing constriction of both
allowing for relaying
arteries (vaso constriction) and veins (veno constriction), and an increase in information derived
cardiac output. The peripheral vasoconstriction and increased cardiac output from blood pressure
raises arterial blood pressure, while the veno-constriction maintains venous within the autonomic
return to the heart. nervous system.
Erythropoietin (EPO)
As discussed in previous section, hemorrhage-induced reduction in baro
is a glycoprotein
receptor inhibition also promotes vasopressin (ADH) release. In addition, there hormone, produced
is an increase in renin/angiotensin/aldosterone activity, resulting from the fall by peritubular cells of
in blood pressure and the associated decreased renal blood flow. Both the kidney that
vasopressin and aldosterone reduce urine formation, thereby conserving stimulates RBCs
plasma volume. There is a marked stimulation of thirst and this helps to productions.
restore plasma volume. The reduced renal blood flow promotes kidney
production of erythropoietin, which stimulates red blood cell production by the
bone marrow. Thus lost red blood cells are replaced by increased production
in the days (week) following the hemorrhage. The liver is also stimulated to
increase the production of plasma proteins. The increase in production of
erythrocytes and plasma proteins, along with the reduction in urine production
and increased drinking rate, restores the blood to its original state.

Table 1.1: Cardiovascular response to extreme conditions.

Condition Cardiovascular Response

Exercise Increase in blood flow and oxygen transfer towards

muscles. Associated with decreased blood flow towards
other body organs like gut, kidney and skin.

Increase in heart rate, decrease in peripheral resistance,

increase in venous return leads to increase in cardiac
output.

Stroke volume is maintained constant.

Diving Peripheral vasoconstriction, reduction in heart rate and

cardiac output results in more blood flow towards brain,
heart and other endocrine organs.

Haemorrhage Vasoconstriction and venoconstriction leading to an

increase in cardiac output.

Release of ADH (Vasopressin). Increase in

renin/angiotensin/aldosterone activity to maintain blood
pressure.

Erythropoietin release from kidney, increased plasma

protein synthesis from liver to maintain blood
composition. 29
Block 1 Fundamentals of Animal Physiology

SAQ 4
State True and False:

a) Hemorrhage reduces both venous and arterial blood pressure.

b) Water stimulates the facial receptors that inhibit the breathing and cause
a marked bradycardia.

c) During exercise cardiac output and heart rate increases up to ten times
above the resting level however, small changes takes place in stroke
volume.

d) During exercise, blood flow to brain and heart is increased.

1.9 SUMMARY
In this unit you have studied that:

• Circulatory system can be divided into two broad categories: Open and
Closed.

• In open circulation the blood usually called haemolymph is pumped by

the heart into spaces where the organs are directly bathed by it. Blood
pressure is low in such a system.

• In closed circulation, blood is pumped by the heart into enclosed vessels

and passes from arteries into veins via capillaries. Blood pressure is high
in the closed circulatory system and the fluid that leaks across the
capillary wall into the interstitial spaces is returned to the circulation by
special vessels forming the lymphatic system.

• The vertebrate heart is a myogenic, muscular pump that ejects blood

into the arterial system. Excitation of the heart starts in a patch of muscle
tissue, the pacemaker and the contraction spreads to the rest of muscle
tissue of the heart. The heartbeat consists of rhythmic contraction
(systole) and relaxation (diastole) of the whole muscle mass of the heart.

• Cardiac output is dependent on the heart rate and stroke volume.

• The arterial system is the pressure reservoir and conduit for blood
between heart and capillaries. The elastic arteries dampen the
oscillations in pressure and flow caused by the contractions of the heart.
Blood to the capillaries is controlled by muscular sphincters at the
arteriole end. The velocity is maximum in arteries and minimum in
capillaries.

• The venous system acts as a conduit for blood between capillaries and
the heart and as a blood reservoir. In mammals 50% of the total blood is
contained in veins.

• Exchange of materials between blood and tissue takes place through the
30 capillary walls. Changes in the composition of extracellular fluid and
Unit 1 Circulatory System
blood in capillaries cause the fluids to leave the capillaries at the arterial
end and to be reabsorbed at the venous end.

• Regulation of the blood pressure could be intrinsic or mediated by

nervous system.

• Intrinsic mechanism is mediated in two ways: 1) The heart—responds to

increased venous return by pumping more forcefully (Starling’s law),
which increases cardiac output and BP. 2) The kidneys-decreased blood
flow decreases filtration, which decreases urinary output to preserve
blood volume. Decreased BP stimulates the kidneys to secrete renin,
which initiates the renin-angiotensin mechanism that results in the
formation of angiotensin II, which causes vasoconstriction and stimulates
secretion of aldosterone.

• The nervous mechanism is also mediated in two ways: 1) Regulation of

Heart rate and 2) By maintaining Peripheral resistance-the medulla
contains the vasomotor centre, which consists of a vasoconstrictor area
and a vasodilator area. The vasodilator area brings about vasodilation by
suppressing the vasoconstrictor area. The vasoconstrictor area
maintains normal vasoconstriction by generating several impulses per
second along sympathetic vasoconstrictor fibers to all arteries and veins.
More impulses per second increase vasoconstriction and raise BP; fewer
impulses per second bring about vasodilation and a drop in BP.

• During exercise, blood flow to skeletal muscle is increased in proportion

to the level of activity of the muscle. Cardiac output can increase up to
ten times above the resting level owing to large increases in heart rate
and small changes in stroke volume.

• In mammals, but not in other vertebrates, stimulation of the facial

receptors that inhibit breathing cause a marked bradycardia.

• Hemorrhage reduces both venous and arterial blood pressure, reducing

the discharge frequency of both atrial and arterial baroreceptors. This
releases the baroreceptive inhibition of sympathetic outflow causing
constriction of both arteries (vasoconstriction) and veins
(venoconstriction), and an increase in cardiac output.

1.10 TERMINAL QUESTIONS

1. What are the advantages of a closed circulatory system? In your opinion
which is superior - closed or open circulatory system?

2. Describe in brief the cardiac conduction pathway and cardiac cycle.

3. Differentiate between artery and vein.

4. Explain with help of a diagram fluid exchange across the capillary wall.

5. Describe the mechanism of blood pressure regulation under three

extreme conditions. 31
Block 1 Fundamentals of Animal Physiology
1.11 ANSWERS
Self-Assessment Questions
1. a) high metabolism

b) Interstitial fluid
2. a) i) volume of blood pumped by the left ventricle during a single
heart beat
ii) stroke volume multiplied by heart rate
b) i) Pacemaker, atria
ii) Bundle of His, Purkinje fibres.
3. b) Capillaries are made up of a single layer of endothelial cells
surrounded by a basal membrane.
c) The arteries near the heart are more elastic and dampen the
oscillation in blood flow.
d) Whole blood is more viscous than plasma because of the
presence of blood cells.
e) The maximum pressure during a heart beat is systolic pressure.
4. a) True, b) True, c) True, d) False.

Terminal Questions
1. Refer to Subsection 1.5.3.
2. Refer to Subsection 1.7.3.
3. Refer to Subsections 1.8.1 and 1.8.2.
4.

Fluid exchange across the capillary wall, occurs near the arterial end of the
capillary since capillary hydrostatic pressure (35mmHg) is greater than the
blood colloidal osmotic pressure (25mmHg). There is no net movement of fluid
near the midpoint of the capillary. At the venous end, fluid is reabsorbed as
capillary hydrostatic pressure is less (18mm Hg) than colloidal blood pressure
(25mm Hg). Draw a figure with help of Fig. 1.9.

5. Refer to Sections 1.4 and 1.5.

32
 UNIT 2
RESPIRATORY
RESPIRATORY SYSTEM

Structure
2.1 Introduction Chemical Regulation of
Respiration
Objectives
2.2 Gas Transfer in Air and Respiration and Acid Base
Water Balance

2.3 Transport of Gases in Blood 2.5 Respiratory Responses to

Extreme Condition
Hemoglobin
2.6 Summary
Oxygen Transport in Blood
2.7 Terminal Questions
Carbon Dioxide Transport in
Blood
2.8 Answers

2.4 Regulation of Respiratory

System
Neural Regulation of
Respiration

2.1 INTRODUCTION
You are aware that all animals utilize oxygen and produce carbon dioxide
during the course of their metabolism. The process of oxygen uptake and
release of carbon dioxide is called respiration. You have already learnt many
aspects of respiration in your graduation course on “Animal Physiology”.
Animals as you are aware take up oxygen from the medium they live in.
Aquatic animals take up oxygen from water and terrestrial animals take in air
from the atmosphere. In most multicellular animals respiration is more
complex as all the cells of the multicellular organisms are not in direct contact
with the respiratory medium (air or water). Therefore, respiratory strategy of
animals depends on the medium in which the animal lives (aquatic or
terrestrial). The present unit focuses on the acquisition of oxygen by animals
from the environment and its distribution to the cells where it is needed for
cellular respiration.

In this unit, the processes involved in the transport and exchange of gases will
be discussed in detail. You will also learn details of gas exchangers also called
respiratory pigment (haemoglobin) that aid in the transport of oxygen. You will
Block 1 Fundamentals of Animal Physiology
see how respiration helps to maintain acid-base balance and pH of the blood.
You will study the neural and chemical regulation of respiratory system and
how respiratory system responds to extreme conditions like hypoxia,
hypercapnia, diving and exercise.

As you already know that in higher animals all systems function in an

integrated manner and no system is able to function alone. So, in this unit you
will note that the circulatory system has been referred to quite often as it is
somewhat difficult to discuss one system with the exclusion of the other.

Objectives
After studying this Unit, you should be able to:

 describe gas transfer mechanisms in air and water,

 explain the process of gas transport between the blood and tissues in
mammals,

 discuss the role of haemoglobin in the transfer of oxygen and compare it

to the transport of carbon dioxide in blood,

 appreciate how respiratory system regulates blood pH and maintain

acid-base balance,

 explain neural and chemical regulation of respiratory system, and

 discuss respiratory response to extreme conditions.

2.2 GAS TRANSFER IN AIR AND WATER

Jelly fish can be very

Many small organisms obtain oxygen by diffusion through their body surfaces.
large. Since it has They do not have any specialized respiratory organs nor do they have blood
less than 1% organic circulation. Larger and more complex organisms, however, need specialized
matter and the rest surfaces for gas exchange and a circulatory system that transports oxygen
99% consists of salt more readily than that possible by simple diffusion across the body surface.
and water, it has a
very low average Calculations based on metabolic demands and rate of diffusion in protoplasm
oxygen consumption show that simple diffusion is sufficient only to meet the demands of organisms
rate. The actively not larger than 1 mm in diameter. These calculations appear reasonable when
metabolizing cells are
we see that animals like protists (protozoans) and flatworms that meet their
located near the
surface therefore,
respiratory requirements through diffusion are either quite small or have very
diffusion alone is low metabolic rates (Fig. 2.1 a). Giant land planarians which are 50 cm long
sufficient for gas are exceptions as they are flat with very large surfaces in relation to mass,
exchange. therefore, diffusion is sufficient to meet their oxygen demand. Coelentrates like
corals and sponges often reach very large sizes but have very modest
metabolic demands. They maintain a circulation of water over the surfaces of
cells by means of cilia which line their canal systems. Thus, sufficient gas
exchange takes place without the aid of a circulatory system or respiratory
pigments.

In animals that have inefficient circulatory system and readily permeable

vascular skins, gas exchange occurs through the integument. Thus we find
34 that animals like earthworms, flatworms (planarians) (Fig. 2.1 a), leeches and
Unit 2 Respiratory System
many larval fishes are among the many animals that obtain the oxygen they
need across their general body surface. Even larger animals such as many
amphibians and fish may rely on cutaneous respiration during emergencies
or use it as a supplement to gills or lungs.

The ability to respire through the integument is most developed in eels, and
amphibians that have moist and highly vascularized skins. The integumentary
contribution to respiration may be as high as 90 per cent in the giant
salamander (Cryptobranchus alleganiensis) and as low as 20 per cent in dry-
skinned toads. The giant salamander is 25 to 60 cm in length and may weigh
over 1 kg. It lacks gills and the unspecialized lungs play little or no part in
respiration. This aquatic amphibian is the largest animal relying exclusively on
cutaneous breathing (breathing by skin).

Animals that are large and have higher metabolic rates have specialized
respiratory organs. These organs have a thin respiratory surface to help in gas
exchange.

Fig. 2.1: Respiratory System of a) flatworm; b) insect; c) fish; and d) humans.

Insects have a special respiratory system (Fig.2.1b). Small openings present

on the insect's body surface are connected to small tubes or trachea that
branch and spread throughout the body. The gases diffuse through these
branches directly into the cells.

Generally gills are for aquatic breathing (Fig. 2.1 c) and lungs (Fig. 2.1 d) for
breathing air. Gills are highly vascularised extensions of gas exchange
membranes. Gill surface area must be large enough to provide adequate
exchange of gases. Therefore, highly active fish have the largest relative gill
area. 35
Block 1 Fundamentals of Animal Physiology
Gas exchange in fishes takes place in the gill lamellae as water flows between
them in one direction and blood within them in the other direction. This is
known as countercurrent flow. This type of flow permits the blood leaving the
fish gills to have the highest possible oxygen levels.

We know that air has more oxygen than water. The atmosphere provides a
constant supply of oxygen almost everywhere. The greatest disadvantage
however, is that air tends to dry out membranes involved in gas exchange. To
overcome this, terrestrial animals have to live in extremely moist environments
or find some ways to keep their respiratory surfaces moist. In order to
accomplish this, water loss in terrestrial animals has been minimized
evolutionarily by turning the respiratory membranes into lungs located inside
the body as seen in amphibians, reptiles, birds and mammals, and into
trachea as in insects. These structures which are contained within the body
are bathed by body fluid both internally and externally and so are kept moist.
Table 2.1 compares oxygen availability in respiration using air and water.

Table 2.1: Major advantages of breathing air.

Properties Air Water

Solubility of oxygen Air contains 209 ml of 7 ml of oxygen per litre

at 15°C oxygen per litre

Energy needed to Less energy needed More energy needed to

move medium (air or to move air over the move water over the
water) over the respiratory surface respiratory surface as
respiratory surface water is much more
viscous than air,

Diffusion of Oxygen Oxygen diffuses some Oxygen diffusion is slow

10,000 times more from water to the
rapidly in air than in respiratory surface in fish
water and so can gill as it takes place only in
diffuse in lungs over a small area constituting a
several millimeters tiny fraction of a millimeter.

As you must have studied in your graduation course that lungs are of two
types i) diffusion lungs which are characterized by air exchange with the
surrounding environment by means of only diffusion, are present in small
animals such as pulmonate snails, small scorpions, some spiders and some
isopods; ii) ventilation lungs which are typical of vertebrates. The air passes
through a tube into inflatable lungs where gas exchange takes place and not
the oxygen poor air, but carbon dioxide rich air is then forced out usually
through the same tube. This is known as tidal flow of air.

You have already studied details of mammalian lungs as they are the best
representatives of a respiratory surface adapted for terrestrial respiration. In
the next section of this unit you will study about gas exchangers and transport
36 of gases.
Unit 2 Respiratory System

SAQ 1
Why does diffusion alone suffice to supply oxygen in both protists (protozoans)
and flatworms?

2.3 TRANSPORT OF GASES IN BLOOD

We know that two types of gas exchanges are constantly occurring in the
animal body-one at the interphase of the respiratory membrane and external
environment (which you have studied in the previous section) and the other in
the rest of the animal tissues. The underlying principle is the same at both
places - passive diffusion occurs along a pressure gradient.

In many invertebrates, oxygen and carbon dioxide are carried dissolved in

blood or haemolymph. Since the amount of oxygen carried in a simple solution
is small, therefore, in highly organised animals (many invertebrates and all
vertebrates) oxygen is transported by being bound reversibly to proteins that
are oxygen carriers. These proteins contain a metal, commonly iron or copper
and are coloured. These proteins are thus, metalloproteins and are known as
respiratory pigments. There are four major classes of respiratory pigments in
animals. They are all metalloproteins that bind reversibly to oxygen at specific
sites on the metal ion in their molecular structure. The four classes of
respiratory pigments are haemocynins (copper containing), hemerythrins,
chlorocrurins and haemoglobins (all three have iron). Out of all these,
haemoglobins are the most widespread respiratory pigment in most
vertebrates.

All four respiratory pigments are adapted to load and unload oxygen
effectively in the habitats where they have evolved, whether animals live on
land where the air contains 210 ml of oxygen per litre or in fresh water
containing 8.0 ml of oxygen/litre or in the sea containing 6.4 ml of oxygen/litre.
The loading unloading reaction of oxygen can be written as follows:
respiratory surface
(haemoglobin) Hb + O2 Hb O2 (oxyhaemoglobin)
tissue

2.3.1 Haemoglobin
Among the respiratory pigments we shall consider haemoglobin in some detail
as this is the most familiar, and widespread and is present in humans. It is also
the most efficient respiratory pigment. In vertebrates haemoglobins are packed
into the red blood cells and combine with far greater amounts of oxygen than
other respiratory pigments that are found dissolved in the plasma. If the
amount of haemoglobin that is packed in the cells was to be free in the plasma
then, the viscosity of blood would be viscous (thick) like syrup and blood would
not be able to flow in the blood vessels as it does. Apart from this the red
blood cells enclosing the haemoglobin provide a stable environment as
reaction of oxygen and haemoglobin is affected by ion concentration and
organic compounds in blood. 37
Block 1 Fundamentals of Animal Physiology
Let us now consider the structure of haemoglobin in more detail so as to know
why haemoglobin has better oxygen carrying capacity. In Fig 2.1 a you can
Haemoglobin has see that hemoglobin (Hb) molecule is made up of four polypeptide subunits 2
even a stronger alpha (α) and 2(β) beta subunits. Each haemoglobin forms a tetrahedral
affinity for carbon structure. Each subunit of haemoglobin has a heme group (an iron-containing
monoxide than compound of the porphyrin class) embedded in it. The heme group which
oxygen. The bond is accounts for only 4 percent of the weight of the molecule, is composed of a
210 times stronger. ring like organic compound known as a porphyrin to which an iron atom is
Carbon monoxide attached (Fig. 2.1 b). It is the iron atom that binds oxygen as the blood travels
tends to displace
between the lungs and the tissues. Thus as there are four iron atoms in each
oxygen in
molecule of haemoglobin, each haemoglobin can accordingly bind to four
haemoglobin and
atoms of oxygen forming oxyhaeomoglobin in a reversible reaction. The
remains attached as
the blood passes unoxygenated haemoglobin compound is called deoxyhaemoglobin. Another
through the tissues. molecule in the body that has the ability to bind oxygen is myoglobin. It is a
The transport of single unit or monomeric form of haemoglobin and consists of a single
oxygen is impaired polypeptide chain the globin in which the heme group is embedded. Myoglobin
leading to dangerous is found in striated muscles of vertebrates and combines with one molecule of
consequences and oxygen.
even death.

Fig. 2.1: a) Chemical structure of heme group; b) Schematic representative of a

single subunit of haemoglobin.

2.3.2 Oxygen Transport in Blood

When no oxygen is bound to the haemoglobin molecule it is said to be in a
tense state (T-state). Haemoglobin molecule has the ability to change its
shape when it binds to the first molecule of oxygen. As a result of the first
binding of oxygen the affinity for oxygen of the haemoglobin increases and the
molecule is said to be in a relaxed state (R-state). This change in the state of
the haemoglobin molecule allows subsequent binding of oxygen molecules to
the rest of the 3 hemes easier and rapid. This type of binding is known as
cooperative binding between active sites.

The transport of oxygen in the blood takes place in two ways:1) the first and
major one is through formation of oxyhaeomoglobin (HbO2) and 2) the second
is through oxygen dissolved in plasma. In haemoglobin the amount of oxygen
bound as oxyhaeomoglobin is 20 ml oxygen/100 ml of blood while oxygen
38 transported through plasma is only 0.2ml of oxygen /100ml of blood.
Unit 2 Respiratory System
The transport of oxygen in blood depends on: i) the pO2 (partial pressure of
oxygen) of the environment and ii) the bond strength or affinity between
haemoglobin and oxygen. In the lungs where Po2 in the inhaled air is high
almost all deoxyhaemoglobin molecules bind to oxygen (Fig. 2.2). Low pO2 in
the systemic capillaries promotes unloading of oxygen. Haemoglobin has bond
strength (affinity) which permits 97% of haemoglobin to combine with oxygen
when leaving the lung. However, it has a stronger affinity for H+ ions than
oxygen. Therefore, in the presence of H+ ions the oxyhaemoglobin molecule
has the ability to unload or disassociate from its oxygen into the systemic
capillaries. We will explain this as we study the oxygen disassociation curves.

Fig. 2.2: Oxygen from the lungs binds to hemoglobin molecules and is carried
through blood vessels to the rest of the body.

Oxygen Dissociation Curves

Recall we had said earlier in the section that in humans the oxygen carrying
capacity is 20 ml oxygen per 100 ml blood. The relationship of oxygen carrying
capacity to the surrounding oxygen concentration can be shown graphically by
the oxygen dissociation curves (ODC) (Fig. 2.3) which give us information
about the most important function of red blood cells and the haemoglobin
contained within them - that is, the affinity of haemoglobin for oxygen and its
delivery to the tissues. These curves are obtained by subjecting blood
samples to different partial pressures of oxygen. The percentage of
oxyhaemoglobin saturation at different partial pressures of oxygen is plotted.

The oxygen dissociation curve is S-Shaped or a sigmoid curve with the

percentage of oxygen saturation of Hb shown on the Y (vertical) axis and the
partial pressure of oxygen depicted on the X (horizontal) axis. You must
remember that the amount of oxygen bound to hemoglobin i.e. the percentage
O2 saturation of hemoglobin at any time is related to partial pressure of O2 that
surrounds it. In the oxygen dissociation curve in Figure 2.3 we can see that
that as PO2 increases the oxygen saturation of haemoglobin also increases.
We can see that total saturation of Hb occurs in the lungs where partial
pressure of oxygen is above 95 mm Hg and the oxygen is unloaded at low
PO2 found in tissues (about 40 mm Hg). We can also see that at 60 mm Hg
partial pressure of oxygen the curve begins to flatten out even if PO2 levels are
increased sharply. The steep portion of the curve indicates that when Po2
drops (in systemic capillaries of tissues) the oxygen is dissociated from Hb
and is released to the tissue. The curve shows that changes in Po2 values
from arterial to venous blood result in 97-75 = 22% unloading when resting. 39
Block 1 Fundamentals of Animal Physiology

Fig. 2.3: Oxygen dissociation curve shows how haemoglobin's oxygen binding
capacity depends on partial pressure of oxygen. Note that there is a
22% decrease in the percent of oxygen as blood passes from arteries to
veins in the tissue. This results in unloading of approximately 5 ml of
The importance of oxygen per 100 ml of blood.
2.3-DPG within the
red blood cells is now The PO2 of oxygen when Hb is 50% saturated is about 28 mm Hg for healthy
recognized in blood person and this point in graph is called P50. In disease, the affinity of O2
banking. Old stored changes and the graph shifts to right or left.
red cells lose their
ability to produce 2.3- Oxygen dissociation curve for a sample of blood is affected by several
DPG which means factors. The most important of them are:
that such cells will not
unload their oxygen 1. Temperature
easily. Modern
techniques for 2. pH
storage of blood,
therefore, include the
3. CO2
addition of energy
4. 2, 3 - Diphosphoglycerate (DPG) - Organic phosphates
substrates for
respiration and 1. Temperature: Haemoglobin, at higher temperature gives up oxygen
phosphate sources
more readily and thus the dissociation curve shifts to the right (Fig 2.4).
needed for production
of 2, 3-DPG.
This is of physiological importance because increased temperature
means higher metabolic rate or higher oxygen requirement by the
organism.

2. pH: Another important factor that influences the oxygen dissociation

curve is pH. An increase in carbon dioxide or other acids in the blood
lowers the pH of the plasma and shifts the dissociation curve to the right
(Fig. 2.4). At high carbon dioxide concentration more oxygen is unloaded
at any given oxygen pressure. This effect is known as Bohr effect, after
40 the Danish scientist who first described it. Therefore, as carbon dioxide
Unit 2 Respiratory System
enters the blood from the respiring tissues, it encourages the release of
more oxygen from the haemoglobin. This is an important characteristic
because it allows more oxygen to be released to the tissues which need
it the most. More oxygen is loaded in the lungs and more is unloaded at
the tissues as a result of Bohr Effect, than would be the case if only
diffusion along the concentration gradient was responsible.

3. Carbon dioxide: Carbon dioxide lowers the oxygen affinity of

haemoglobin even if the pH is kept constant. This effect is due to the
binding of carbon dioxide to the terminal amino groups of haemoglobin
molecule forming carbaminohaemoglobin. This site is not the same site
on the molecule where oxygen is bound.

4. 2, 3-diphosphoglycerate (DPG): The presence of organic phosphates

especially 2,3-diphosphoglycerate (DPG), in the red blood cells helps to
explain many peculiarities of the oxygen dissociation curve. Previously
the red blood corpuscle was considered to be a bag full of haemoglobin
with no metabolism of its own because of the absence of a nucleus. Now
we know that it has an active carbohydrate metabolism and the RBC has
a high content of ATP and 2, 3-diphosphoglycerate (DPG). 2, 3-
diphosphoglycerate is a product of glycolysis and binds to the beta (β)
chain of the globin and reduces oxygen affinity (Fig. 2.4). Experimentally
it has been shown that pure haemoglobin has greater oxygen affinity
than whole blood (the dissociation curve for pure haemoglobin is far to
the left of the curve as compared to that for the whole blood). If 2, 3-DPG
is added to pure haemoglobin solution the oxygen affinity decreases and
approaches that of whole blood.

Fig. 2.4: The oxygen dissociation curve shifts to the right or left depending on
temperature, carbon dioxide, pH, or 2,3-DPG levels . Due to Bohr effect
oxyhaemoglobin surrenders its oxygen more readily in the presence of
increasing acidity which is the case in metabolically active cells where
more CO2 is released. 41
Block 1 Fundamentals of Animal Physiology
We have learnt about the S shaped curve for haemoglobin. Let us now
see how the oxygen dissociation curve of haemoglobin is different from
the curve showing oxygen affinity in myoglobin.

Fig. 2.5: A comparison of the dissociation curves for haemoglobin and for
myoglobin. At the PO2 of venous blood, the myoglobin retains almost
all of its oxygen, indicating a higher affinity than haemoglobin for
oxygen. The myoglobin does, however, release its oxygen at the very
low pO2 values found inside the mitochondria.

In myoglobin the heme reacts to oxygen independently. It does not show

cooperativity binding as in the case of haemoglobin. Fig. 2.5 shows the
dissociation curve for myoglobin which in contrast to the curve for
haemoglobin is rectangular. Myoglobin is described as middleman in the
transfer of oxygen from blood to mitochondria within muscle cells. Myoglobin
has a much stronger affinity for oxygen and stores oxygen in the muscles. It
binds to oxygen at a much lower partial pressure than haemoglobin and
dissociates from its bound oxygen only when the partial pressure of oxygen is
very low as found in the mitochondria.
Fig. 2.15 also indicates that the myoglobin remains oxygenated until quite low
levels of PO2 in the surrounding fluids is reached. You can see from the Figure
that at 20 mm Hg the haemoglobin in the blood is about 30% saturated but the
myoglobin in the muscles is saturated above 80%.

SAQ 2
Pick out the True statements from those given below:
i) The haemoglobin of human adults contains four identical subunits and a
haem group.
ii) When haem and globin fractions are separated the haem group binds,
reversibly with oxygen.
iii) Bohr's effect facilitates transfer of oxygen to the tissues because of
increased CO2 levels in blood.
iv) Haemoglobin molecule has the ability to change its shape when it binds
to the first molecule of oxygen due to which its affinity for oxygen
increases.

42
Unit 2 Respiratory System
2.3.3 Carbon Dioxide Transport in Blood
The same transport system that brings oxygen to the tissues must take back
carbon dioxide to the environment across the respiratory surface. However,
unlike oxygen that is transported exclusively by haemoglobin, carbon dioxide
is transported in three ways:

1. as dissolved carbon dioxide in plasma (about 8%),

2. in red blood cell as carbaminohaemoglobin. About 25% of the total

blood carbon dioxide is carried attached to the amino groups in
haemoglobin.

Haemoglobin − < N H+ CO → H+ + Haemoglobin − NHCOO−

H 2
Carbamino haemoglobin is carried to the lungs where haemoglobin
releases the carbon dioxide in exchange for oxygen,

3. as carbonic acid (H2CO3) and bicarbonate/bicarbonate ion (HCO -3 )

which accounts for most of the carbon dioxide (67%)carried by blood.

Carbon dioxide is generated in the tissues and diffuses freely into the venous
plasma and then into the RBCs. In the RBCs, CO2 combines with H2O to form
carbonic acid (H2CO3).

CO2 + H2O → H2CO3

This reaction occurs spontaneously in the plasma at a very slow rate but much
more rapidly within the blood cell due to the catalytic reaction of an enzyme,
carbonic anhydrase. The formation of carbonic acid is favoured by high
PCO2 in the capillaries of tissues. Carbonic acid dissociates rapidly in the red
blood cells into hydrogen ion (H+) and bicarbonate ion (HCO -3 ) .

H2CO3 → H+ + (HCO -3 )

The H+ (hydrogen ions) released are buffered by combining with haemoglobin

and HCO 3- moves out of the cells. The inside of the cell thus gains a net
positive charge. This attracts the chloride ion (CI-) which moves inside the red
blood cells. This exchange of anions as the blood moves through capillaries in
tissue is known as chloride shift (Fig. 2.6 a). The red blood cells are very
-
permeable to both Cl and HCO 3- ; because the membrane has a high
concentration of a special anion carrier protein called band III protein that
binds CI- and HCO 3- and transfers them in opposite direction through the RBC
membrane.

The formation of carbonic acid enhances oxygen unloading (Bohr Effect). The
oxygen unloading in turn improves the ability of blood to form carbonic acid
and transport carbon dioxide.

When blood reaches the pulmonary capillaries deoxyhaemoglobin is

converted to oxyhaeomoglobin which has a lower affinity for H+. The H+ is
released in the red blood cells. This attracts HCO 3- from plasma which
combines with H+ to form H2CO3 43
Block 1 Fundamentals of Animal Physiology
H + HCO -3 → H2CO3
+

Under low PCO2 of pulmonary vessels the carbonic anhydrase catalyses the
formation of carbonic acid to carbon dioxide and water

H2CO3 Carbonic anhydrase H2O + CO2

Low PCo2

Therefore, a reverse chloride shift occurs in the pulmonary capillaries to

convert carbonic acid, and biocarbonate to CO2 gas which is eliminated in the
expired breath (Fig. 2.6 b).

After having discussed the properties of blood and its role in the transport of
respiratory gases in the present unit we can now proceed to discuss the
regulation of respiration.

(a)

(b)
Fig. 2.6: Carbon dioxide transport in blood. a) carbon dioxide is transported in
three forms: (i) as dissolved CO2 gas in plasma; attached to
haemoglobin as carbaminohaemoglobin, and as carbonic acid and
bicarbonate in plasma due to chloride shift; b) carbon dioxide is
released from the blood as it travels through pulomonary capillaries. A
reverse chloride shift occurs and carbonic acid is transformed into CO2
44 and H2O releasing CO2 in the alveolus.
Unit 2 Respiratory System

SAQ 3
In how many ways does carbon dioxide transported in blood?

2.4 REGULATION OF RESPIRATION

In mammals, ventilation during respiration is operated by separate groups of
neurons that work alternatively. Ventilation in mammals and birds is regulated
primarily by the amount of carbon dioxide present in the lung air. Even a slight
increase in CO2 level in the inspired air increases the rate of ventilation. CO2
becomes dangerous in higher concentrations. Oxygen on the other hand has a
much smaller effect on ventilation. If the O2 concentration is decreased from
21% to 18.5% there is virtually no effect on ventilation. It is peculiar that even
though metabolism requires oxygen yet the respiratory centre is not very
sensitive to decreased oxygen levels.

2.4.1 Neural Regulation of Respiration

The rhythmic contractions of the diaphragm and of the intercostal muscles are
controlled by respiratory centre which consists of clusters of neurons located
in the area of the medulla oblongata and pons of the brain.

There are four respiratory groups that regulate respiration in the body - two in
the medulla and two in the pons. The two groups in the pons are known as
the pontine respiratory group. The respiratory group are as follows:

1. Dorsal respiratory group - in the medulla

2. Ventral respiratory group - in the medulla

3. Pneumotaxic center - various nuclei of the pons

4. Apneustic center - nucleus of the pons

Chemoreceptors that are located in the respiratory centers are sensitive to

increased carbon dioxide levels and increased acidity of the blood and the
cerebrospinal fluid and respond by increasing the lung ventilation so that more
CO2 is able to leave the lungs and its levels return to normal. Chemoreceptors
that are sensitive to changes in arterial carbon dioxide, oxygen, and pH
(resulting from changes in the levels of carbon dioxide) transmit the
information of these changes to the centres in the brain by nerve impulses
which transmit the information to respiratory centres so that they can respond
appropriately by sending impulses to increase or decrease ventilation (Fig.
2.11). Whenever the need for oxygen in the body increases, ventilation of the
respiratory organs also increases. The respiratory muscles in particular the
diaphragm and intercostal muscles as a result become activated and cause air
to move in and out of the lungs.

Central chemoreceptors are located on the medulla oblongata, near

the medullary respiratory groups of the respiratory center. They are mainly
sensitive to changes in the pH in the blood, (resulting from changes in the
levels of carbon dioxide). 45
Block 1 Fundamentals of Animal Physiology
The peripheral chemoreceptors are present in the carotid bodies and aortic
arch near the heart and are sensitive to changes in arterial carbon dioxide,
oxygen, and pH. Chemoreceptors located in the carotid bodies and aortic
arches respond to decreased PO2 levels by increasing the depth and
frequency of breathing. The carotid bodies in general are more important in
mediating this response and provide the principal mechanism by which
mammals sense decreased levels of oxygen. Information from the peripheral
chemoreceptors is conveyed along nerves to the respiratory groups of the
respiratory center.

Fig. 2.7: Respiratory centre in brain.

2.4.2 Chemical Regulation of Respiration

Chemical regulation refers to the effect of blood pH and blood levels of oxygen
and carbon dioxide on breathing. This is shown in Fig. 2.8. Chemoreceptors
that detect changes in blood gases and pH are located in the carotid and
aortic bodies and in the medulla itself.

A decrease in the blood level of oxygen (hypoxia) is detected by the

chemoreceptors in the carotid and aortic bodies. The sensory impulses
generated by these receptors travel along the glossopharyngeal and vagus
nerves to the medulla, which responds by increasing respiratory rate or depth
(or both). This response will bring more air into the lungs so that more oxygen
can diffuse into the blood to correct the hypoxic state.

Carbon dioxide becomes a problem when it is present in excess in the blood,

because excess CO2 (hypercapnia) lowers the pH when it reacts with water
to form carbonic acid (a source of H+ ions). That is, excess CO2 makes the
blood or other body fluids less alkaline (or more acidic). The medulla contains
chemoreceptors that are very sensitive to changes in pH, especially
decreases. If accumulating CO2 lowers blood pH, the medulla responds by
increasing respiration. This is not for the purpose of inhaling, but rather to
46 exhale more CO2 to raise the pH back to normal.
Unit 2 Respiratory System
Of the two respiratory gases, which is the more important as a regulator of
respiration? Our guess might be oxygen, because it is essential for energy
production in cell respiration. However, the respiratory system can maintain a
normal blood level of oxygen even if breathing decreases to half the normal
rate or stops for a few moments. This oxygen did not enter the blood but was
available to do so if needed. Also, the residual air in the lungs supplies oxygen
to the blood even if breathing rate slows.

Therefore, carbon dioxide must be the major regulator of respiration, and the
reason is that carbon dioxide affects the pH of the blood. As was just
mentioned, an excess of CO2 causes the blood pH to decrease, a process that
must not be allowed to continue. Therefore, any increase in the blood CO2
level is quickly compensated for by increased breathing to exhale more CO2.
If, for example, you hold your breath, what is it that makes you breathe again?
Have you run out of oxygen? Probably not, for the reasons mentioned. What
has happened is that accumulating CO2 has lowered blood pH enough to
stimulate the medulla to start the breathing cycle again.

In some situations, oxygen does become the major regulator of respiration.

People with severe, chronic pulmonary diseases such as emphysema have
decreased exchange of both oxygen and carbon dioxide in the lungs. The
decrease in pH caused by accumulating CO2 is corrected by the kidneys, but
the blood oxygen level keeps decreasing. Eventually, the oxygen level may fall
so low that it does provide a very strong stimulus to increase the rate and
depth of respiration.

Fig. 2.8: Chemical regulation of respiration.

2.4.3 Respiration and Acid Base Balance

As you have already studied in previous sections that respiration affects the
pH of body fluids because it regulates the amount of carbon dioxide in these
fluids. Remember that CO2 reacts with water to form carbonic acid (H2CO3),
which ionizes into H+ ions and HCO– ions. The more hydrogen ions present in
a body fluid, the lower the pH, and the fewer hydrogen ions present, the higher
the pH.

The respiratory system may be the cause of a pH imbalance, or it may help

correct a pH imbalance created by some other cause.

Respiratory Acidosis and Alkosis

Respiratory acidosis occurs when the rate or efficiency of respiration
decreases, permitting carbon dioxide to accumulate in body fluids. The excess
CO2 results in the formation of more H+ ions, which decrease the pH. Holding
one’s breath can bring about a mild respiratory acidosis, which will soon 47
Block 1 Fundamentals of Animal Physiology
stimulate the medulla to initiate breathing again. More serious causes of
respiratory acidosis are pulmonary diseases such as pneumonia and
emphysema, or severe asthma. Each of these impairs gas exchange and
allows excess CO2 to remain in body fluids.

Respiratory alkalosis occurs when the rate of respiration increases, and CO2
is very rapidly exhaled. Less CO2 decreases H+ ion formation, which
increases the pH. Breathing faster for a few minutes can bring about a mild
state of respiratory alkalosis. Babies who cry for extended periods (crying is a
noisy exhalation) put themselves in this condition. In general, however,
respiratory alkalosis is not a common occurrence. Severe physical trauma and
shock, or certain states of mental or emotional anxiety, may be accompanied
by hyperventilation and also result in respiratory alkalosis. In addition, traveling
to a higher altitude (less oxygen in the atmosphere) may cause a temporary
increase in breathing rate before compensation occurs.

Respiratory Compensation

If a pH imbalance is caused by something other than a change in respiration, it

is called metabolic acidosis or alkalosis. In either case, the change in pH
stimulates a change in respiration that may help restore the pH of body fluids
to normal.

Metabolic acidosis may be caused by untreated diabetes mellitus

(ketoacidosis), kidney disease, or severe diarrhea. In such situations, the H+
ion concentration of body fluids is increased. Respiratory compensation
involves an increase in the rate and depth of respiration to exhale more CO to
decrease H+ ion formation, which will raise the pH toward the normal range.

Metabolic alkalosis is not a common occurrence but may be caused by

ingestion of excessive amounts of alkaline medications such as those used to
relieve gastric disturbances. Another possible cause is vomiting of stomach
contents only. In such situations, the H+ ion concentration of body fluids is
decreased. Respiratory compensation involves a decrease in respiration to
retain CO2 in the body to increase H+ ion formation, which will lower the pH
toward the normal range.

Respiratory compensation for an ongoing metabolic pH imbalance cannot be

complete, because there are limits to the amounts of CO2 that may be exhaled
or retained. At most, respiratory compensation is only about 75% effective.

SAQ 4
Fill in the blanks:

a) An increase in the rate and depth of respiration to exhale more CO to

decrease H+ ion formation, which will raise the pH toward the normal
range is called………………..

b) When the rate of respiration increases, and CO2 is very rapidly exhaled
leads to decreases H+ ion formation, which increases the pH. This
condition is called …………………

48
Unit 2 Respiratory System
2.5 RESPIRATORY RESPONSES TO EXTREME
CONDITIONS
Variations in the levels of respiratory gases, diving by air- breathing animals,
and exercise all induce respiratory responses. Let's see how animals adjust to
these extreme conditions.

Reduced Oxygen Levels (Hypoxia) in Water

Aquatic animals are subjected to more frequent and rapid changes in oxygen
levels than are air-breathing animals. Both mixing and diffusion are more rapid
in air compared with water, so regions of local hypoxia develop more often in
aquatic environments. Although photosynthesis can cause very high oxygen
levels during the day in some aquatic environments, oxygen consumption by
both biological and chemical processes can produce localized hypoxic
regions. The changes in oxygen levels in water may or may not be
accompanied by changes in carbon dioxide.

Many aquatic animals can withstand very long periods of hypoxia. Some
fishes (e.g., carp) overwinter in the bottom mud of lakes where the PO2 is very
low. Many invertebrates also bury themselves in mud with a low PO2 but high
nutritive content. Some parasites live in hypoxic regions, such as the gut,
during one or more phases of their life cycle. Limpets and bivalve molluscs
close their shells during exposure at low tide to avoid desiccation, but as a
consequence are subject to a period of hypoxia. Many of these animals utilize
a variety of anaerobic metabolic pathways to survive the periodofre
duceoxygen availability. Others also adjust the respiratory and cardiovascular
systems to maintain oxygen delivery in the face of reduced oxygen availability.
For instance, aquatic hypoxia causes an increase in gill ventilation in many
fish, as a result of stimulation of chemoreceptors on the gills. The increase in
water flow offsets the reduction in oxygen content and maintains delivery of
oxygen to the fish. Fishes, such as tuna, ventilate their gills by swimming
forward with their mouth open, the size of the gap increases with hypoxia to
increase water flow over the gills.

Reduced Oxygen Levels (Hypoxia) in Air

Compared with aquatic environments, oxygen and carbon dioxide levels are
relatively stable in air, and local regions of low oxygen or high carbon dioxide
are rare and easily avoided. There is, of course, a gradual reduction in PO2
with altitude, and animals vary in their capacity to climb to high altitudes and
withstand the accompanying reduction in ambient oxygen levels. The highest
permanent human habitation is at about 5800 m, where the PO2, is 80 mm Hg
compared to about 155 mm Hg at sea level. Many birds migrate over long
distances at altitudes above 6000 m, where atmospheric pressures would
cause severe respiratory distress in many mammals. High altitudes are
associated with low temperatures as well as low pressures, and this also has a
marked effect on animal distribution.

A reduction in the PO2, of ambient air results in a decrease in blood pressure,

which in turn stimulates the carotid and aortic body chemoreceptors, causing
an increase in lung ventilation in mammals. The rise in lung ventilation then 49
Block 1 Fundamentals of Animal Physiology
leads to an increase in CO2, elimination and a decrease in blood PCO2. The
decrease in blood PCO2 causes an increase in pH of the CSF. Decrease in
blood PCO2 and increases in CSF pH tend to reduce ventilation, thereby
attenuating the hypoxia-induced increase in lung ventilation. If, however,
hypoxic conditions are maintained, as occurs when animals move to high
altitude, both blood and CSF pH are returned to normal levels by the excretion
of bicarbonate. This process takes about one week in humans. Thus, as CSF
pH returns to normal, the reflex effects of hypoxia on ventilation predominate;
the result is a gradual increase in ventilation as the animal acclimatizes to
altitude. The response to prolonged hypoxia may also involve modulation of
the effects of CO2, on the carotid and aortic bodies to reset these
chemoreceptors to the new lower CO2, level at high altitude.

As mentioned earlier, low oxygen levels cause a local vasoconstriction in the

pulmonary capillaries in mammals, producing a rise in pulmonary arterial blood
pressure. This response normally has some importance in redistributing blood
away from poorly ventilated, and therefore hypoxic, portions of the lung. When
animals are subjected to a general hypoxic environment, however, the
increase in the resistance to flow through the whole lung can have detrimental
effects. Some mammals that live at high altitudes exhibit a reduced local
pulmonary vasoconstriction in response to hypoxia; this is probably a
genetically determined acclimation. Humans residing at high altitudes are
usually small and barrel chested, and have large lung volumes. Lung
development is oxygen insensitive, but the growth of limbs is reduced under
hypoxic conditions. The high lung to body ratio enables these people to
maintain oxygen uptake under hypoxic conditions. Pulmonary blood pressures
are high, and there is often hypertrophy of the right ventricle. High pulmonary
pressures produce more even distribution of blood in the lung, and so
augment the diffusing capacity for oxygen.

Increased Carbon Dioxide Levels (Hypercapnia)

In many animals, an increase in blood PCO2 results in an increase in
ventilation. In mammals the increase is proportional to the rise in the CO, level
in the blood. The effect is mediated by modulation of the activity of several
receptors that send messages to the medullary respiratory center. These
receptors include the chemoreceptors of the aortic and carotid bodies and the
mechanoreceptors in the lungs, but the response is dominated by the central
H+ receptors. Correction of CSF pH, in the face of altered PCO2 levels, is very
important in the return of ventilation to normal.

A marked increase in ventilation occurs almost immediately in response to a

rise in CO. However, ventilation eventually returns to a level slightly above the
volume that prevailed before hypercapnia. This return to a value only slightly
greater than the initial ventilation level is related to increase in levels of plasma
bicarbonate and CSF bicarbonate, with the result that pH returns to normal
even though the raised CO levels are maintained.

Diving by Air-Breathing Animals

Many air-breathing vertebrates live in water and dive for varying periods of
time such as dolphins and whales rise to the surface to breathe but spend
50 most of their life submerged.
Unit 2 Respiratory System
Diving mammals and birds are, of course, subjected to periods of hypoxia
during submergence. The mammalian central nervous system (CNS) cannot
withstand anoxia and must be supplied with oxygen throughout the dive.
Diving animals solve the problem by utilizing oxygen stores in the lungs, blood,
and tissues. Many diving animals have high hemoglobin and myoglobin levels,
and their total oxygen stores generally are larger than those in non- diving
animals. To minimize depletion of available stores, oxygen is preferentially
delivered to the brain and the heart during a dive; blood flow to other organs
may be reduced, and these tissues may adopt anaerobic metabolic pathways.
Air-breathing animals that spend prolonged periods submerged at sea must
have sufficient oxygen stores to sustain aerobic metabolism, because they
cannot tolerate the high accumulation of lactic acid that results from anaerobic
metabolism. During prolonged dives, metabolic rates and thus oxygen needs
often are reduced in such animals (e.g., elephant seals).

Some diving animals, such as the Weddell seal, exhale before diving, thus
reducing the oxygen store in their lungs. During a deep dive, the increase in
hydrostatic pressure results in lung compression. In those animals that reduce
lung volume before a dive, air is forced out of the alveoli as the lungs collapse
and is contained within the trachea and bronchi, which are more rigid but less
permeable to gases. If gases remained in the alveoli, they would diffuse into
the blood as pressure increased. At the end of the dive, the partial pressure of
nitrogen in the blood would be high, and a rapid ascent would result in the
formation of bubbles in the blood the equivalent of decompression sickness, or
the "bends," in humans. Thus exhalation before diving reduces the chances of
the bends occurring. Since only about 7% of a Weddell seal's total oxygen
stores are in the lungs, pre-dive exhalation appears to be a reasonable trade-
off.

Receptors that detect the presence of water and that inhibit inspiration during
a dive are situated near the glottis and near the mouth and nose (depending
on the species). The decrease in blood oxygen levels and increase in CO,
levels that occur during a dive do not stimulate ventilation because inputs from
the chemoreceptors of the carotid and aortic bodies are ignored by the
respiratory neurons while the animal is submerged.

During birth, a mammal emerges from an aqueous environment into air and
survives a short period of anoxia between the time the placental circulation
stops and the time air is first inhaled. The respiratory responses of the fetus
during this period are similar in several respects to those of a diving mammal.

Exercise

Exercise increases oxygen utilization, CO2 production, and metabolic acid

production. Cardiac output increases to meet the higher demands of the
tissues. Even though the transit time for blood through the lung capillaries is
reduced, nearly complete gas transfer still occurs. Ventilation volume
increases in order to maintain gas tensions in arterial blood in the face of
increased blood flow. The increase in ventilation in mammals is rapid,
coinciding with the onset of exercise. This initial sudden increase in ventilation
volume is followed by a more gradual rise until a steady state is obtained both
for ventilation volume and oxygen uptake. 51
Block 1 Fundamentals of Animal Physiology
When exercise is terminated, there is a sudden decrease in breathing,
followed by a gradual decline in ventilation volume. During exercise, oxygen
levels are reduced and CO, and H+ levels raised in venous blood, but the
mean PO2 and PCO2 in arterial blood do not vary markedly, except during
severe exercise. The oscillations in arterial blood PO2 and PCO2 associated
with each breath increase in magnitude, although the mean level is unaltered.

Exercise covers a range from slow movements up to maximum exercise

capacity. The phrase moderate exercise refers to exercise above resting
levels that is aerobic, with only minor energy supplies derived from anaerobic
glycolysis. Severe exercise refers to exercise in which oxygen uptake is
maximal and further energy supplies are derived from anaerobic metabolism.
Heavy exercise is a term sometimes used to denote the exercise level
between moderate and severe exercise.

The onset of exercise involves many changes in lung ventilation and the
cardiovascular system, as well as muscle contraction. In the initial stages,
during the transition from rest to exercise, the animal is not in a steady state
and part of the energy supply is derived from anaerobic processes. If the
exercise level is moderate and sustained, the animal moves into a new steady
state typical of that exercise level, with increased lung ventilation, cardiac
output, blood flow to the exercising muscles, and oxygen uptake. The
relationship between lung ventilation and oxygen uptake is linear during
moderate exercise, the slope of the relationship varies with the type of
exercise.

Table 2.2: Respirarory response to extreme condition.

Condition Respiratory Response

Reduced Oxygen Many invertebrates switch to anaerobic

Levels (Hypoxia) in metabolic pathways.
water
Many fish Increase in gill ventilation.

Reduced Oxygen Many mammals show increase in lung

Levels (Hypoxia) in air ventilation, which leads to an increase in carbon
dioxide elimination and a decrease in blood CO
levels.

Long term response involves excretion of

bicarbonate ions, increase in RBC number and
blood haemoglobin content, reduction in
oxygen-haemoglobin affinity.

People living in high altitudes have high lung to

body ratio.

Increased Carbon Initial increase in ventilation which eventually

Dioxide Levels returns to normal level maintaining blood pH.
(Hypercapnia)

Diving by Air Breathing Utilize oxygen stores in the lungs, blood and
Animals tissues.
52
Unit 2 Respiratory System
Oxygen is preferentially delivered to brain and
heart; blood flow to other organs is reduced.

No increase in ventilation as receptors located

near mouth and nose detect the presence of
water.

Exercise (Increase in For complete gas exchange between blood and

oxygen utilization, carbon alveoli the transit time for blood through lung
dioxide and metabolic capillaries is reduced.
acid production)
Increase in ventilation volume and ventilation
speed.

SAQ 5
State ‘True’ or ‘False’.

a) The relationship between lung ventilation and oxygen uptake is linear

during moderate exercise.

b) The respiratory responses of the fetus during this period are similar in
several respects to those of a diving mammal.

c) Some diving animals, such as the Weddell seal, exhale before diving,
thus reducing the oxygen store in their lungs.

d) Compared with aquatic environments, oxygen and carbon dioxide levels

are relatively unstable in air.

2.6 SUMMARY
In this Unit you have studied that:

• Most aquatic animals exchange gases through gills that are evaginations
of respiratory surfaces. The air breathing vertebrate lung is a highly
branched inpocketing of respiratory surface containing numerous air
sacs or alveoli, intimately associated with capillaries and ventilated by a
tidal flow of air.

• Respiratory gases in vertebrates including human are transported mainly

by respiratory pigments in blood. The most well-known pigment is
haemoglobin which combines with oxygen to form oxyhaemoglobin.
Oxygen dissociation curve depicts percent oxyhaemoglobin saturation at
different values of PO2. Carbon dioxide, pH, temperature and presence
of organic compounds in blood influence this curve. A fall in pH
decreases the oxygen affinity of haemoglobin for oxygen. This is called
the Bohr’s effect. Decreased affinity causes unloading of oxygen to the
tissue. Oxygen affinity also decreases in the presence of 2, 3-DPG and
in conditions of increased blood temperature.

• Carbon dioxide from the tissues is transported to the lungs mainly by the
formation of carbonic acid in red blood cells; as carboxyhaemoglobin
and due to dissolved CO2 in plasma. Carbonic acid ionizes into H+ and 53
Block 1 Fundamentals of Animal Physiology
HCO 3- . Hydrogen ion is buffered by haemoglobin but anion balance is
maintained by chloride shift. Reverse chloride shift occurs in the
pulmonary capillaries and carbon dioxide is formed from carbonic acid
dissociation into water and carbon dioxide and hence carbon dioxide in
gas form is exhaled outside.

• The medulla contains the inspiration center and expiration center.In the
pons: the apneustic center prolongs inhalation, and the pneumotaxic
center helps bring about exhalation. These centers work with the
inspiration center in the medulla to produce a normal breathing rhythm.

• Decreased blood O2 is detected by chemoreceptors in the carotid body

and aortic body. Response: increased respiration to take more air into
the lungs. Increased blood CO2 level is detected by chemo- receptors in
the medulla. Response: increased respiration to exhale more CO2.

• Breathing also regulates the level of CO2 within the body, and this
contributes to the maintenance of the acid–base balance of body fluids.

• Variations in the levels of respiratory gases, diving by air- breathing

animals, and exercise all induce respiratory responses.

2.7 TERMINAL QUESTIONS

1. Write a formula representing the reaction between haemoglobin and
oxygen. What factors influence the rate and direction of the reaction?

2. How is carbon dioxide transported when it is released by the tissues into

the blood in mammals? What is the role of carbonic anhydrase?

3. Discuss respiratory response to extreme conditions.

2.8 ANSWERS
Self-Assessment Questions
1. Calculations by scientists, based on metabolic demands and rate of
diffusion in protoplasm show that simple diffusion is sufficient to meet the
demands of organisms not larger than 1 mm in diameter and since
protists (protozoans) and flatworms are less than 1 mm in diameter their
respiratory requirements are met through diffusion.

2. i) False; ii) False; iii) True; iv) True

3. Carbon dioxide is transported in three forms

a) as dissolved carbon dioxide gas in blood plasma

b) attached to haemoglobin as carbaminohaemoglobin

c) as carbonic acid and bicarbonate in plasma

4. a) Respiratory compensation

54 b) Respiratory alkalosis
Unit 2 Respiratory System
5. a) True

b) True

c) True

d) False

Terminal Questions
1. Hb + O2 ⇌ HbO2.

Hint: Temperature, CO2, pH and organic phosphates.

2. Refer to Subsection 2.3.

3. Refer to Subsection 2.5.

55
 UNIT 3
OSMOREGULATION AND
EXCRETI
EXCRETION

Structure
3.1 Introduction 3.6 Process of Urine Formation
Objectives Formation of Primary Urine

3.2 Principles of Osmoregulation Modification of Urine

Transport of Electrolytes Across 3.7 Regulation of Kidney

Cell Membranes Function
Ion Channels in Osmoregulation Juxtaglomerular Apparatus

3.3 Osmoregulation in Aqueous Renin-Angiotensin-Aldosterone

and Terrestrial Environments System (RAAS)

3.4 Extra Renal Osmoregulatory 3.8 Patterns of Nitrogen

Organs in Invertebrates Excretion
Contractile Vacuoles 3.9 Summary
Nephridia of Invertebrates 3.10 Terminal Questions
Molluscan Kidney 3.11 Answers
Green Glands

Malpighian Tubules

3.5 Structure and Function of the

Vertebrate Kidney

3.1 INTRODUCTION
You already know that water is an essential component of the cytoplasm and
of the body of organisms. Distribution of gases, transportation of nutrients and
waste products inside the body, all depend on water as a diffusion medium. If
the water content of the body changes, all cellular activities are disrupted and
life is jeopardized as all chemical reactions inside the body of an organism
occur in an aqueous environment. Thus maintaining water balance or
homeostasis inside the body is of paramount importance. This water balance
in the body is maintained by ensuring that the intake of water is equal to water
Unit 3 Osmoregulation and Excretion
loss. Water balance is also linked to the balance of solutes present throughout
the body. To survive, an animal must maintain normal volume and composition
of the extracellular fluids as well the intracellular fluids, which means that the
ionic (electrolyte) concentrations and pH of the fluids have to be maintained
along with their volume. The process of maintaining osmotic pressure by
maintaining the concentration of ions and water in the body fluids is
known as osmoregulation.

How this water and electrolyte balance is maintained in the body is the subject
of the first part of this unit. We will first look into the processes by which water Homeostasis: refers
moves in and out of the cells. We will then investigate how animals regulate to the body’s ability to
the volume of water and ionic concentration in the body in accordance with maintain a stable
internal environment.
their external environment. You will learn that even though the environments
Animal organs and
are different, the molecular mechanisms for facing these challenges by the organ systems
organisms are the same in many cases. You will also understand how organs constantly adjust to
such as gills and salt glands all work to achieve the ionic and water balance the internal and
in the body of different animals. external changes in
order to maintain this
Osmoregulation and excretion are intimately related as the ultimate aim of steady state.
these two processes is to maintain homeostasis. Excretion is the removal of
toxic waste products of metabolism from the body. The end products of
metabolism are either eliminated or conserved by an animal depending on its In humans, electrolyte
physiology or must be converted into a less toxic form before being excreted. imbalances lead to
muscle spasms,
The process of excretion and osmoregulation are performed by the same set confusion, irregular
of organs. In the later half of this unit we will discuss the structure and function heart beats, fatigue,
of extra renal osmoregulatory organs in different animal groups. paralysis and even
death.
Objectives
After studying this Unit, you should be able to:

 explain the meaning of osmoregulation and excretion,

 discuss how osmoregulators and osmoconformers adapt to their

environment,

 describe the anatomical diversity and functional principles of excretory

organs in different groups of animals, and

 explain different patterns of nitrogen excretion.

3.2 PRINCIPLES OF OSMOREGULATION

You are familiar with the processes of diffusion and osmosis. After studying
Unit 2, “Respiratory system”, of this course, you already know that diffusion
involves the movement of substances from a region of higher concentration to
a region of lower concentration. In living organisms solutes move between
cells and extracellular spaces along their concentration gradient across a
selectively permeable membrane. When the solutes are equally distributed
throughout the solutions on both sides of the membrane then they are said to
be in equilibrium and molecules can move back and forth at equal rates. The
concentration of solutes in the solution is known as the solution’s osmolarity 57
Block 1 Fundamentals of Animal Physiology
(Refer to Box 3.1). However, when solutes cannot cross the selectively
permeable membrane then water moves from the region where solutes are
less to the region where the solutes are more concentrated. We can say that
water moves across a semi permeable membrane from regions of lower
osmolarity to regions of higher osmolarity. This movement of water is called
osmosis and the pressure that would have to be applied to the water to
prevent it from passing across a semi permeable membrane is the osmotic
pressure (Refer to Box 3.1).

(a) (b) (c)

Fig. 3.1: a) Cells placed in a hypertonic environment tend to shrink due to loss of
water; b) the blood maintains an isotonic environment so that cells
neither shrink nor swell In a hypotonic environment ; c) cells tend to
swell due to intake of water (credit: Mariana Ruiz Villareal).

Recall we had defined osmoregulation as the process of maintenance of salt

and water balance (osmotic balance) across membranes within the body’s
fluids, which are composed of water, plus electrolytes and non-electrolytes. An
electrolyte is a solute that dissociates into ions when dissolved in water. In
most animals the most abundant solutes or electrolytes are sodium, potassium
chlorine and calcium ions. A non-electrolyte, in contrast, doesn’t dissociate
into ions during water dissolution. Both electrolytes and non-electrolytes
contribute to the osmotic balance. The body’s fluids which you all know include
the blood plasma, the cytosol within cells, and the interstitial fluid (the fluid that
exists in the spaces between cells and tissues of the body). The membranes
of the body (such as the pleural, serous, and cell membranes) are semi-
permeable membranes. Semi-permeable membranes are permeable (or
permissive) to certain types of solutes and water. Solutions on both sides of a
semi permeable membrane tend to equalize the solute concentration by
movement of solutes and/or water across the membrane. As seen in Figure
3.1 c, a cell placed in water tends to swell due to gain of water from the
hypotonic or “low salt” environment of water. A cell placed in a solution with
higher salt concentration (hypertonic), on the other hand, tends to shrink and
the membrane shrivels up due to loss of water into the hypertonic or “high salt”
environment (Fig. 3.1 a). Isotonic cells are those that have an equal
concentration of solutes inside and outside the cell; this equalizes the osmotic
pressure on either side of the cell membrane which is a semi-permeable
58 membrane (Fig. 3.1 b).
Unit 3 Osmoregulation and Excretion
Box 3.1

It is important to be clear about the basic concepts of osmolarity and

membrane permeability. The presence of dissolved solute confers on a
solution the property of osmotic pressure also called osmotic concentration.
Osmotic pressure is one out of the four colligative properties [The colligative
property of a solution is that which is dependent on the ratio between the total
number of solute particles (in the solution) to the total number of solvent
particles]. Colligative properties are not dependent on the chemical nature of
the solution’s component of a solution. The remaining three colligative
properties of a solution are boiling point elevation, freezing point depression
and relative lowering of the vapour pressure. Osmotic pressure of a
solution depends on the number of dissolved particles present in it per
unit volume. While the chemical concentration of a solution is expressed in
molarity (moles/litre), the osmotic concentration of a solution is
expressed in osmolarity (osmoles/litre). For ideal non-electrolytes (e.g.,
sucrose), a one molar solution is one osmolar. An electrolyte solution, on the
other hand, has a higher osmolarity than its molarity. For example, NaCl in
the solution dissociates into Na+and Cl-- ions. Thus, for every molecule of
NaCl, one gets two ionic particles in the solution, one of Na+ and one of Cl--
.Hence, one molar NaCl solution is nearly two osmolar. In the same way, one
molar CaCl2 solution is nearly three osmolar. An osmole is defined as that
amount of a solute which when dissolved in one litre of water has the same
osmotic pressure as one mole of an ideal non-electrolyte in one litre of water.
If two solutions (solution A and solution B) have the same osmotic
concentration, they are said to be isosmotic to each other. If solution A has
a higher osmolarity than solution B then A is hyperosmotic to B or B is
hypoosmotic to A.

3.2.1 Transport of Electrolytes Across Cell Membranes

Electrolytes, such as sodium chloride, ionize in water, meaning that they
dissociate into their component ions. In water, sodium chloride (NaCl),
dissociates into the sodium ion (Na+) and the chloride ion (Cl–). The most
important ions, whose concentrations are very closely regulated in body fluids,
are the cations sodium (Na+), potassium (K+), calcium (Ca+2), magnesium
(Mg+2), and the anions chloride (Cl–), carbonate (CO3-2), bicarbonate (HCO3–),
and phosphate (PO3–). Electrolytes are lost from the body during urination and
perspiration. For this reason, in very hot weather and during strenuous
exercise people are encouraged to replace electrolytes and fluids. Water can
pass through membranes by passive diffusion. If electrolyte ions could
passively diffuse across membranes, it would be impossible to maintain the
specific concentrations of ions in the body therefore ion channels present on
the cell membranes have a vital role to play in osmoregulation.

3.2.2 Ion Channels in Osmoregulation

Ion channels are protein molecules that span across the cell membrane
allowing the passage of ions from one side of the membrane to the other (see
Box 3.2). The mechanism of action of some of the ions channels and pumps
and the important role they play in osmoregulation are as given below: 59
Block 1 Fundamentals of Animal Physiology
+ +
The Na /K ATPase pump which is present in the cell (plasma) membranes is
powered by ATP and pumps to move sodium and potassium ions in opposite
directions, each against its concentration gradient. In a single cycle of the
pump, three sodium ions are taken out from the cell when the ATP molecule
provides it energy. Then there is a conformational change in the ion channel
protein and two potassium ions are imported into the cell and the phosphate of
the ATP is dissociated from the pump (Fig. 3.2). Na+/K+ ATPase pump
produces and helps to maintain osmotic equilibrium and a membrane potential
(difference in charge) across cell membranes.

+ +
Fig.3.2: Na /K ATPase pump functions to maintain osmotic balance and electric
+
gradient across membranes by taking 3 Na out of the cell and at the
+
same time bringing back 2 K into the cell. The energy required for this
comes from the phosphorylation of this protein ATPase pump by ATP.

• A Na+/Cl-/K+ cotransporter is a protein which uses the electric gradient

produced by Na+/K+ ATPase pump to bring three ions: sodium,
potassium and chloride into the cell across the membrane. The
cotransporter brings in 1 Na+, 2 Cl- and 1K+ together into the cell. There
is an abundance of these cotransporters in the nephrons of the kidney
where Na, K and Cl ions are extracted from the urine so that they can be
reabsorbed into the blood. In such a case the Na + ion is again taken out
60 of the cell by the Na+/K+ ATPase pumps into the extracellular fluid but the
Unit 3 Osmoregulation and Excretion
- +
concentration of Cl and K builds up in the cell. Selective ion channels
present in the cell membrane allow passive diffusion of the potassium
and chlorine ions. The potassium ion (K+) passively diffuses out of the
cell into the extracellular fluid through potassium channels and chlorine
ion (Cl-) diffuses down the concentration gradient via chloride
channels.

• Some sodium ions also diffuse down the concentration gradient through
sodium channels.

• Aquaporins (AQP) are integral membrane proteins that serve as

channels in the transfer of water, and in some cases, small solutes
across the membrane.

Box 3.2

Passive transport: Diffusion along an electrochemical gradient without the

use of energy involves the passive movement of ions across the membrane
through protein channels present in the plasma/cell membrane. In passive
transport specific channels for particular ions may be present. Furthermore,
there could also be carriers too that bind to specific proteins and undergo
shape changes in order to deposit the bound protein inside the cell, this is
known as facilitated diffusion.

Active transport is powered by ATP and functions in the movement of ions.

In this transport ions are brought inside the cell against the electrochemical
gradient through special membrane bound proteins. Once an
electrochemical gradient has been established inside the cell a
cotransporter or an antitransporter uses the same energy to bring in a
different ion against the concentration gradient.

SAQ 1
a) Fill in the blanks with the relevant word/words:

i) Osmolarity of an organism depends on the concentration of

…………… in their body fluid.

ii) The marine environment is ………………… to invertebrates such

as sponges and cnidarians occupying that habitat.

iii) The body fluids of marine teleost fish are …………………. to their
environment while the body fluids of elasmobranches are
…………………. to the surrounding sea water

iv) Aquaporins (AQP) are integral membrane proteins that serve as

channels in the transfer of …………………., and in some cases,
small …………………. across the membrane.

b) What is the the role of Na+/K+ATPase pump in osmoregulation?

61
Block 1 Fundamentals of Animal Physiology
3.3 OSMOREGULATION IN AQUEOUS AND
TERRESTRIAL ENVIRONMENTS
Animals face quite distinct osmotic problems in aqueous and terrestrial
environments. In this section, we first discuss osmoregulation by water-
breathing animals and then consider air-breathing animals.

A) Water Breathing Animals

Many aquatic animals find themselves and all their respiratory surfaces
immersed in water. The osmolarities of aqueous environments range from a
few milliosmoles per liter in fresh water lakes to about 1000 mosm per litre in
ordinary seawater, or even more in landlocked salt seas. Intermediate
environments, such as brackish bogs, marshes, and estuaries, have salinities
ranging between these extremes. As a rule, the body fluids (i.e., interstitial
fluids and blood) tend away from the environmental osmotic extremes.
Euryhaline aquatic animals can tolerate a wide range of salinities, whereas
stenohaline animals can tolerate only a narrow osmotic range. In this section,
we consider the nature of the osmotic problems faced by freshwater and
marine animals and their mechanisms for dealing with them.

Freshwater Animals

As you know the body fluids of freshwater animals, including invertebrates,

fishes, amphibians, reptiles, and mammals, are generally hyperosmotic to their
aqueous surroundings (see Table 3.1). Freshwater vertebrates have blood
osmolarities in the range of 200 to 300 mosm. L-I, while the osmolarity of
freshwater is generally much less than 50 mosm-L-l. Because they are
hyperosmotic to their aqueous surroundings, freshwater animals face two
kinds of osmoregulatory problems:

• They are subject to swelling by movement of water into their bodies

owing to the osmotic gradient

• They are subject to the continual loss of body salts to the surrounding
medium, which has a low salt content.

The freshwater animals must prevent the net gain of water and net loss of
salts, which they accomplish by several means.

One way to avoid a net gain of water is production of a dilute urine. Among
closely related fishes, for example, those that live in freshwater produce a
more copious (i.e., plentiful and hence dilute) urine than their saltwater rela-
tives (see Fig. 3.3).The useful salts are largely retained by reabsorption into
the blood from the ultrafiltrate in the tubules of the kidney, and thus a dilute
urine is excreted. Nonetheless, some salts pass out in the urine, so there is a
potential problem of gradually washing out biologically important salts such as
KCI, NaCl, and CaCI,. Lost salts are replaced, in part, from ingested food. In
addition, freshwater animals have remarkable abilities to take up salts from
their dilute environment. Freshwater fishes are able, for example, to extract
Na+ and Cl- ions with their gills from water containing less than 1mM NaCl,
even though the plasma concentration of the NaCl exceeds 100 mM (Fig.
62 3.3b). The mechanism of sodium reabsorption appears to be similar in the gills
Unit 3 Osmoregulation and Excretion
of freshwater fishes, frog skin, turtle bladder, and the mammalian kidney. In all
cases the cells of these epithelia are joined together by tight junctions.
Transport of Na+ into these cells is dependent on an electrogenic proton
ATPase, which actively transports protons out of the cells into the
environment. The mechanism of sodium reabsorption is discussed in detail
later.

In some freshwater animals, including fishes, reptiles, birds, and mammals,

water uptake and salt loss are minimized by an integument having low
permeabilities to both salts and water. As a general rule, animals living in
fresh- water refrain from drinking fresh water, reducing the need to expel
excess water.

Marine animals

In general, the intracellular and extracellular body fluids of marine

invertebrates and the ascidians (primitive chordates) are close to seawater
both in osmolarity (isosmotic) and in the plasma concentrations of the
individual major inorganic salts (see Table 3.1). Such animals therefore need
not expend much energy in regulating the osmolarity of their body fluids. A
rare example of a vertebrate whose plasma is also isosmotic to the
environment is the hagfish. It differs from most marine invertebrates, however,
in that it does regulate the concentrations of individual ions. In particular, blood
Ca2+,Mg2+,and SO-: are maintained at significantly lower concentrations than
they are in seawater, whereas Naf and Cl- are higher. Since various func-
tions of excitable tissues such as nerve and muscle of vertebrates are
especially sensitive to the concentrations of Ca2+and Mg2+,the regulation of
these divalent cations may have evolved to accommodate the requirements of
neuromuscular function.

Like the hagfish, the cartilaginous fishes such as sharks, rays, and skates, as
well as the primitive coelacanth Latimeria, have plasma that is approximately
isosmotic to seawater. They differ, however, in that they maintain far lower
concentrations of electrolytes (i.e., inorganic ions), making up the difference
with organic osmolytes such as urea and trimethylamine oxide (TMAO).High
urea concentrations tend to cause the breakup of proteins into constituent
subunits, whereas TMAO has the opposite effect, cancelling the effect, of urea
and stabilizing protein structure in the face of high urea levels. In the
elasmobranchs and coelacanths, excess inorganic electrolytes such as NaCl
are excreted via the kidneys and also by means of a special excretory organ,
the rectal gland, located at the end of the alimentary canal.

The body fluids of marine teleosts (modern bony fishes),like those of most
higher vertebrates, are hypotonic to seawater, so there is a tendency for these
fishes to lose water to the environment, especially across the gill epithelium.
To replace the lost volume of water, they drink salt- water. Most of the net salt
uptake is due to drinking sea- water rather than salt uptake across the body
surface or gills. By absorption across the intestinal epithelium, 70% to 80% of
the ingested water enters the bloodstream, along with most of the NaCl and
KCl in the seawater. Initially the ingested seawater is diluted by about 50% by
diffusional uptake of salts across the esophagus. Active salt uptake occurs in
the small intestine, via a Na/2Cl/K cotransporter across the apical membrane 63
Block 1 Fundamentals of Animal Physiology
and then by active transport via a Na+/K+ATPase across the basolateral
membrane. Left behind in the gut and expelled through the anus are most of
the divalent cations such as Ca2+,Mg2-, and SO:-. The excess salt absorbed
along with the water is subsequently eliminated from the blood by active
transport of Na+, Cl-, and some K+ across the gill epithelium into the
seawater, and by secretion of divalent salts by the kidney (see Fig. 3.3a).The
urine is isotonic to the blood, but rich in those salts (especially Ca2+,Mg2+,and
SO:-) that are not secreted by the gills. The net result of the combined osmotic
work of gills and kidneys in the marine teleost is a net retention of water.

The gills of marine teleosts, as might be expected, are organized differently

from those of freshwater fish. In marine teleosts, the gill epithelium contains
specialized cells, called chloride cells, that mediate transport of NaCl from
the blood into the surrounding water. The mechanism of this transport, which
makes it possible for these fishes to live in saltwater; is described in a later
section.

Fig.3.3: Saltwater and freshwater face different osmotic challenges. a) In marine

fish water is lost by osmosis and electrolytes are gained by diffusion; b)
In freshwater fish, water is gained by the fish body through diffusion by
osmosis and electrolytes are gained by the fish by active transport via
the gills. Freshwater fish excrete very dilute urine. Red arrows show
64 direction of ion movement and blue arrows show water movement.
Unit 3 Osmoregulation and Excretion
B) Air Breathing Animals

Animals in a terrestrial environment can be thought of as submerged in an

ocean of air rather than water. Unless the humidity of the air is high, animals
having a water- permeable epithelium will be subject to dehydration very much
as if they were submerged in a hypertonic medium such as seawater.
Dehydration would be avoided if all epithelial surfaces exposed to air were
totally impermeable to water. The evolutionary process has not found this to
be a feasible solution to the problem of desiccation, since an epithelium that is
impermeable to water (and thus dry) will have limited permeability to oxygen
and carbon dioxide, and will thus be unsuited for the respiratory needs of a
terrestrial animal. As a consequence, air-breathing animals are subject to
dehydration through their respiratory epithelia. Air-breathing animals utilize
various means to minimize water loss into the air by this route and others.

Marine reptiles (e.g., iguanas, estuarine sea turtles, crocodiles, sea snakes)
and marine birds drink seawater to obtain a supply of water but, like marine
teleosts, are unable to produce a concentrated urine that is significantly
hyperosmotic to their body fluids. Instead, they are endowed with glands
specialized for the secretion of salts in a strong hyperosmotic fluid. These salt
glands are generally located above the orbit of the eye in birds (Figure 3.4a)
and near the nose or eyes in lizards. Brackish-water crocodiles were long
suspected of using extrarenal means of excreting salts, and eventually salt
glands were discovered in the tongue of this reptile (Fig. 3.4b). Marine
mammals, which lack salt glands or similar specializations, avoid drinking
seawater, get their water entirely from their food intake and its subsequent
metabolism, and depend primarily on their kidneys for maintaining osmotic
balance.

Human beings, like other mammals, are not equipped to drink seawater. This
is so because to excrete the salt ingested with a given volume of seawater, the
human kidney must pass more water than is contained in that volume. This, of
course, will lead rapidly to dehydration.

Fig. 3.4: Salt glands in a) bird; b) iguana, a reptile.

Desert Living Mammals

Camels, desert mice, kangaroo rat and many other mammals can survive in
the arid conditions of the desert without drinking water. 65
Block 1 Fundamentals of Animal Physiology
The kangaroo rat and other desert mammals are faced with a physiological
double jeopardy-excessive heat and near absence of free freshwater. The
kangaroo rat, like many desert mammals, avoids much of the daytime heat by
remaining in a burrow during daylight hours and coming out only at night. This
nocturnal lifestyle is an important and widespread behavioural adaptation to
desert life. In addition, with the ambient temperature in the burrow being
significantly lower than the core temperatures of birds and mammals the nasal
counter- current mechanism for conserving respiratory moisture works well.
Because of its efficient kidneys, the kangaroo rat excretes a highly
concentrated urine, and rectal absorption of water from the feces results in
essentially dry fecal pellets. When deprived of drinking water the primary
source of water for kangaroo rat is the water produced during the oxidation of
foods.

Unlike the kangaroo rat, camels are too large to hide from the hot desert sun
in burrows. When deprived of drinking water, camels do not sweat but allow
their body temperature to rise rather than losing water by evaporative cooling
during the heat of the day. During the cooler night, the camel's body
temperature drops and increases only slowly the next day because of the
animal's large body mass and thick fur, which acts as a heat shield.
Nevertheless, the body temperature of a dehydrated camel may vary from
35°C at night to 41°C during the day. This strategy of heating during the day
and cooling at night is impossible in small rodents, whose body temperatures
oscillate much more rapidly than in the larger camel. Because of their small
size dessert rodents heat up rapidly in the sun and must return to their burrow
to cool down. The camel also reduces heating by orienting to give minimal
surface exposure to direct sunlight. The camel, like other desert animals,
produces dry feces and concentrated urine. When water is not available the
camel does not produce urine but stores urea in the tissues. The camel can
tolerate not only dehydration but also high urea levels in the body. When water
becomes available, these ships of the desert can rehydrate by drinking 80
liters in 10 minutes.

Marine Mammals

Marine mammals face problems similar to those of desert animals because

they live in an environment without available drinking water.

The physiological responses of marine mammals, although different in detail,

are generally similar to those of desert mammals. The emphasis is on water
conservation. They are endowed, as are other mammals, with highly efficient
kidneys capable of producing a very hypertonic urine. Seals have a
characteristic labyrinth-like proliferation of epithelial surfaces in the nasal
passages which reduces water loss via breathing. Whales and dolphins have
a blow hole rather than the typical mammalian nose. These animals have
large lung tidal volumes. The velocity of air flow through the blow hole is high
because both inspiration and expiration are rapid, and large volumes of gas
are moved with each breath. It is possible that the expansion of air passing
through the blow hole of a whale also cools the air, resulting in water
condensation in the region of the blow hole that can be used to wet inspired
66 air. This would reduce water loss via ventilation.
Unit 3 Osmoregulation and Excretion
Table 3.1: Animals living in different environments exhibit different
osmoregulatory mechanisms. (Reproduced from Eckert
Animal Physiology)

Terrestrial Arthropods

Certain terrestrial arthropods (like arachnids -ticks, mites and in a number of

wingless forms of insects, primarily larvae) have the ability to extract water
vapor directly from the air, even when the relative humidity is as low as 50%
(Table 3.1). 67
Block 1 Fundamentals of Animal Physiology
The water vapor pressure associated with a solution decreases with
increasing ionic content, so highly concentrated salt solutions will absorb water
from air. Insects take advantage of this by creating very concentrated solutions
that can absorb water from air. In insects that extract water from air, the site of
entry is often the rectum, which reduces the water content of fecal matter to
remarkably low levels. As water is removed from the feces, the latter can take
on new water from the air, if the water vapor pressure is high enough and if
the air has access to the rectal lumen. In ticks, tissues in the mouth have been
implicated in the uptake of water vapor. Here it appears that the salivary
glands secrete a highly concentrated solution of KCl that in turn absorbs water
from the air.

SAQ 2
State ‘True’ of ‘False’ for the following statements.

a) Certain terrestrial arthropods (like arachnids -ticks, mites and in a

number of wingless forms of insects, primarily larvae) have the ability to
extract water vapor directly from the air.

b) The nocturnal lifestyle is an important and widespread behavioural

adaptation to desert life.

c) Urea and TMAO have opposite effect on protein conformation.

d) Sea birds and marine reptiles have salt glands to remove extra salts
from their body.

e) Fresh water fish does not drink water and actively absorb salt ions from
surrounding water.

f) Marine fish secrete excess salt through their gills.

3.4 EXTRA RENAL OSMOREGULATORY

ORGANS IN INVERTEBRATES
In your graduation course on “Animal Physiology” you have already studied
about the anatomy and functional details of renal organs. Also, in the previous
section we have discussed about various mechanisms and organs (besides
kidney) present in vertebrates for osmoregulation. In the present section we
will study the extra renal osmoregulatory organs present in various
invertebrates.

In general, invertebrate osmoregulatory organs employ mechanisms of

filtration, reabsorption, and secretion similar in principle to those of the
vertebrate kidney to produce a urine that is significantly different in osmolarity
and composition from the body fluids. Insects and possibly some spiders are
the only invertebrates known to produce a concentrated urine. These
mechanisms are used to differing extents in various organs in different groups
of animals. That there has been convergent evolution of physiological
mechanisms in nonhomologous organs underscores the utility of these
68 mechanisms.
Unit 3 Osmoregulation and Excretion
3.4.1 Contractile Vacuoles
Contractile vacuoles are present in fresh water protists (protozoans) and
sponges, but are frequently absent from their marine counterparts. The
contractile vacuole complex (CVC) in protists is an osmoregulatory organelle
composed of cisternae and interconnecting ducts. Large cisternae act as
bladders that periodically fuse with the plasma membrane, forming pores to
expel water.

Since freshwater animals are hyperosmotic to their surrounding water and

their surface is permeable to water they gain water by diffusion and need to
pump it out continuously. Microscopic examination has revealed that in protists
the contractile vacuoles exhibit a cyclic function (Fig. 3.5). A vacuole gets filled
with fluid and increases in volume (diastole) till it reaches a critical size and
then it moves to the cell boundary, attaches to it and suddenly expels its
contents to the exterior of the cell and decreases in size (systole). Contractile
vacuoles are the primary organs of osmoregulation in protists and any role
they may have in excretion of ammonia is considered secondary.

Fig. 3.5: Cyclic function of osmoregulation in Amoeba.

3.4.2 Nephridia of Invertebtates

Nephridia are the excretory tubules in many invertebrates. While all nephridia
open to the exterior by an excretory pore known as nephridiopore, some
have their internal end closed and others open into the body cavity by a
ciliated funnel known as nephridiostome. The former type of nephridia are
known as protonephridia and the latter type as metanephridia. 69
Block 1 Fundamentals of Animal Physiology

Fig. 3.6: Excretory system of a) a planaria which has the protonephridium type
of excretory system that contains flame bulbs; b) an earthworm has
metanephridium type of excretory system with nephrostome and is
closely associated with the circulatory system.

Protonephridia are more primitive and occur in acoelomates and

pseudocoelomates. An animal may have one or two more extensively
branched protonephridia each terminating in a bulb like flame cells also
called flame bulb. Inside each flame cell is a tuft of cilia that moves to drive
the fluid through the tubules (Fig 3.6a). Metanephridia are found in coelomates
(e.g., annelids).The metanephridia of annelids (Fig. 3.6b) are closely
associated with their circulatory system resulting in the direct exchange of
material between the two systems. In these invertebrates the isoosmotic fluid
is filtered from the coelomic fluid into the nephridium through the ciliary mesh
of the nephrostome. Later salt is reabsorbed and a dilute urine is discharged.

3.4.3 Molluscan Kidney

In molluscs the coelom is reduced to the pericardial cavity and the cavities
enclosing the kidneys and gonads. The pericardial coelom opens into the renal
cavity via the reno-pericardial duct (Fig.3.7). Thus the molluscan kidney is a
modified metanephridial coelomo duct with one end opening into the
pericardial coelom and the other end opening into the mantle cavity. The initial
urine (pre - urine) of the molluscs is formed by ultra filtration of the blood
through the wall of the heart and enters the renal cavity through the reno-
pericardial duct. Reabsorption of valuable materials like glucose and other
ions and secretion of unwanted salts and molecules takes place in the kidney
sac and ureter resulting in the formation of the final urine. Most freshwater
snails are able to maintain their body fluids hyperosmotic to the exterior
environment in a varying range of osmolarities.

70 Fig. 3.7: Kidney and its relationship to the pericardium in freshwater snail.
Unit 3 Osmoregulation and Excretion
3.4.4 Green Glands
Crustaceans have a pair of renal organs, each of which consists of a closed
end sac with a labyrinth and a long coiled nephridial canal ending in a bladder
that opens to the exterior through an excretory pore located at the base of the
antennae in which case they are known as the antennary glands or opening
at the base of the maxillae, in which case they are known as maxillary
glands. The end sac in many cases is green hence these renal organs are
also known as the green glands (Fig. 3.8a).

Urine formation in crustaceans involves filtration, reabsorption and secretion.

In the antennal gland of the freshwater crayfish (Astacus), fluid is filtered
from the blood into the end sac and passes through a tubular labyrinth.
Chloride ion (Cl-) is reabsorbed from the isoosmotic ultrafiltrate during its
passage in the nephridial canal so that the final urine is hypoosmotic (Fig.
3.8b).Urine concentration is known to vary in crustaceans depending on
whether they are fresh water or marine. Like in other freshwater invertebrates
the urine in fresh water crustaceans is hypoosmotic and very dilute while in
brackish water crustaceans the urine is closer to blood osomolarity. This may
be due to the absence of the salt reabsorbing segment in their nephridial
tubule. The hydrostatic pressure of the blood is the driving force for filtration.

Fig. 3.8: a) Location of green gland in a crustacean b) Structure of the renal

organ (green gland) involved in urine formation in crayfish.

3.4.5 Malpighian Tubules

The osmoregulatory and excretory organs of insects are the Malpighian
tubules.

The production of insect excreta containing urine or pellets is a result of two

related processes: excretion and osmoregulation (maintenance of
favourable osmotic pressure and ionic concentration of body fluid). The
system responsible for excretion and osmoregulation is referred to as
excretory system and its activities are performed largely by the Malpighian
tubules and hindgut. The osmoregulatory excretory organs of insects are
the Malpighian tubules that may be two to several hundred in number and
also the hind gut (ileum, rectum and colon). Malpighian tubules are long,
tubular structures, usually arising at the junction of the mid- and hindgut. Their
lumen is continuous with the lumen of the gut at one end, while the other end
terminates blindly in the haemocoel and so these Malpighian tubules are
similar to protonephridia. The Malpighian tubules are bathed in the 71
Block 1 Fundamentals of Animal Physiology
haemolymph and have numerous tracheal connections (Fig. 3.9a). The tubule
walls of the malphighian tubules consist of a single cell layer of epithelial cells
and are differentiated by structure and function along the length of the tubule.

The mechanism of primary urine formation in the Malpighian tubules of insects

is however, different compared to other invertebrates. The tracheal system of
insects eliminates the necessity of a pressurized arterial circulatory system.
The formation of urine takes place by secretion rather than ultrafiltration.

Fig. 3.9: a) Location of malpighian tubule; b) Formation of urine in the

Malpighian tubules and gut in insect (locust). The Malpighian tubules
system and the hindgut in insects carry out osmoregulation and
removal of nitrogenous wastes. Note - 1. Nitrogenous wastes in insects
is uric acid; Uric acid and ions are moved into malpighian tubules by
active transport and water follows by osmosis. 2. Ions are actively
transported back to the coelom fluid and water follows. 3. Uric acid is
excreted with faeces.

Malpighian tubules work cooperatively with specialized glands present in the

wall of the rectum of the hindgut. Body fluids are not filtered, as in the case of
nephridial, instead, urine is produced by tubular secretion mechanisms by the
72 cells lining the Malpighian tubules. Metabolic wastes, such as urea and amino
Unit 3 Osmoregulation and Excretion
acids, freely diffuse into the tubules, while ions are transported through active
pump mechanisms. There are exchange pumps lining the tubules among
which the apical cation (K+) actively pumps(secretes) K+ or Na+ ions out into
the lumen of Malpighian tubules The secretion of these ions generates an
osmotic pressure gradiant due to which water passively enters the lumen of
Malpighian tubules. Electrolytes and nitrogenous waste (uric acid) also
passively enter into the lumen of Malpighian tubules. Before excretion the
essential ion molecules are reabsorbed in the hind gut. Reabsorption is highly
selective. The whole process of reabsorption is tightly regulated, allowing
precise control over water and ionic balance. Figure 3.9b summarizes the
process of urine formation in gut of insect locust. The proton pump called the
V-ATPase (vacuolar-type ATPase) actively pumps, and transports H+ ions
back into the cells of the hind gut. The H+ ions and other electrolytes useful for
the insect as well as water are reabsorbed into the haemolymph. The
excretory matter present in the Malpighian tubule predominantly in the form of
uric acid is precipitated and excreted as a thick paste or powder. Figure 3.9b
summarizes the process of urine formation in gut of locust. By not dissolving
wastes in water, these organisms are able to conserve water; this is especially
important for life in dry environments.

SAQ 3
a) Match the excretory organ given in column A with the animal in which it
occurs given in column B.

Column A Column B

i) Malphigian tubule a protists (amoeba)

ii) contractile vacuole b crustacean

iii) nephridia c insects

iv) green gland d worm (annelids)

b) Describe with the help of suitable diagram the osmoregulation in insects.

3.5 STRUCTURE AND FUNCTION OF THE

VERTEBRATE KIDNEY
Typically all vertebrates have a pair of kidneys, which function on the principle
of filtration, reabsorption and secretion. Only in a few teleost fish the kidney
is aglomerular (without glomerulus) and functions on the absorption-
reabsorption-secretion principle similar to the Malpighian tubules of insects. In
this unit for studying the vertebrate excretory system we will describe the
mammalian kidney and its functions though it is not representative of all
vertebrates as it performs several functions which in lower vertebrates are
performed or shared by other organs such as skin and bladder in amphibians,
gills in fish and salt glands in reptiles and birds.

You all know that mammals (Fig. 3.10) have paired kidneys which are bean
shaped organs. A large blood vessel, the renal artery brings in blood 73
Block 1 Fundamentals of Animal Physiology
containing nitrogenous wastes to each kidney and a renal vein takes away
the filtered blood. The urine that is formed is collected in the ureters leading to
a storage organ the bladder from where it is transported to the body surface
by a tube called the urethra and excreted outside the body. Figure 3.10 shows
the human urinary system and a longitudinal section of a kidney with its
internal parts. The functional unit of the vertebrate kidney is the nephron or
the uriniferous tubule.

Fig. 3.10: Anatomy of the human urinary system. a) A pair of bean-shaped

kidneys are located near the spinal column in humans; b) L S of a
human kidney showing the : cortex forming the outer part of the
kidney; medulla that forms the inner portion of the kidney; renal
pyramids which are the triangular-shaped divisions of the medulla of
the kidney; papilla which are the narrow, innermost tips of the
pyramids; pelvis also called the kidney or renal pelvis which is an
extension of the upper end of the ureter (the tube that drains urine into
the bladder); calyx (each calyx is a division of the renal pelvis and
Jacob Henle opens into the papilla of a pyramid).
described the loop of
Henle part of the The kidney of a small fish may be made up of just a few nephrons while a
nephron first in 1860s large mammal may have several million nephrons organized within the bean
hence it has been shape structure that we associate with the kidney. Most of the nephrons in a
named loop of Henle. human kidney (approximately one million) are located in the outer layer called
the renal cortex. However, some nephrons extend into the inner region known
as the renal medulla. As you can see in Figure 3.11 a and b that each
nephron begins as a closed tube in the shape of a double walled cup called
the Bowman’s capsule (imagine a balloon that has been pushed in from one
end forming a cup) (Fig. 3.11 b) and extends into a narrow tube that empties
into a collecting duct along with other nephrons (Fig. 3.11a). Several such
collecting ducts combine to form papillary ducts that finally empty into the
ureter. The Bowman’s capsule in each nephron encloses a knot of fine
74 capillaries forming the glomerulus (Fig. 3.11 a & b). Together the Bowman’s
Unit 3 Osmoregulation and Excretion
capsule and glomerulus form the renal capsule. The Bowman’s capsule of
each nephron continues posteriorly to form the long convoluted tubule that can
be distinguished into the - proximal and distal convoluted tubules
respectively. In mammals and birds there is a‘U’ shaped region of the nephron
that is present between the proximal and distal tubules which is called the
loop of Henle.

Fig. 3.11: a) Structure of nephrons in the human kidney; b) Bowman’s capsule

surrounding the glomerulus.

Vertebrates that lack a loop of Henle are incapable of producing urine

hypertonic to blood and so the concentration of the final uretal urine in them
maybe hypoosmotic or at best isoomotic. In mammals the nephrons are so
arranged in the kidney that their collecting tubules are arranged parallel to
each other giving the characteristic striated appearance to the interior of the
kidney. The glomeruli are located in the renal cortex of the kidney and the
loops of Henle are located in the renal medullary region. Two types of
nephrons are found in the mammalian and avian kidneys:

• Juxtamedullary nephrons – that have their glomeruli located deep

within the cortex near the cortica medullary border and their long loops
of Henle extend deep into the medulla.

• Cortical nephrons – that have their glomeruli located nearer to the

outer part of the cortex and their short loops of Henle donot extend into
the medulla.

The arrangement of the blood vessels in the kidney is suited to the mechanism
of urine formation. The blood enters via the afferent arteriole in the
glomerular capillaries and leaves via the efferent arteriole rather than via a
vein as is the case with other capillary networks. After leaving the Bowman 75
Block 1 Fundamentals of Animal Physiology
capsule and before leaving the kidney the efferent arteriole form another
interconnecting capillary network around the loop of Henle known as vasa
recta. The vasa recta lies parallel to the Loop of Henle. The flow of blood is
about 10% less in the efferent arteriole than in the afferent arteriole because
the glomerulus filters about 10% of the fluid into the Bowman’s capsule. Let us
now see how urine formation takes place in vertebrates using the example of
human (mammals).

SAQ 4
Correct the following statements.

a) Cortical nephrons- that have their glomeruli located deep within the
cortex and loops of Henle reaching into the renal medulla.

b) Juxtamedullary nephrons- that have their glomeruli located nearer to the

outer part of the cortex and their short loops of Henle donot extend deep
into the renal medulla.

c) The blood enters via the efferent arteriole in the glomerular capillaries.

d) The blood leaves via the afferent arteriole rather than via a vein as is the
case with other capillary networks.

3.6 PROCESS OF URINE FORMATION

As mentioned earlier three main processes take place in the formation of final
urine that is excreted as summarized in Figure 3.12:

Fig. 3.12: Schematic figure showing the movement and process of urine
76 formation in a mammalian nephron.
Unit 3 Osmoregulation and Excretion
As you can seen in Fig. 3.12, the following activites result in the formation of
urine.

• Filtration of blood plasma forms an ultrafiltrate in the Bowman’s capsule

• Tubular reabsorption of about 99% of water and the reabsorption of

other salts takes place from the ultrafiltrate. The ultra filterate is thus, left
behind with waste products such as urea.

• Tubular secretion of substances via active transport.

In addition, some substances are formed in the tubular cells and in the lumen
that assist in the formation of nitrogenous waste.

3.6.1 Formation of Primary Urine

You know that in the human kidneys each nephron has a glomerulus which
functions as a microscopic filter, that constantly filters blood (Fig. 3.13a). Blood
which is to be filtered enters the glomerulus, which consists of fenestrated
capillaries, as they have openings or pores (Fig. 3.13 b and c). These
fenestrations are 100 times more permeable than the continuous capillaries
(capillaries without fenestrations) that are found in other parts of the body. The
physical characteristics of the glomerular capillary wall determine what is
filtered and how much is filtered into the Bowman’s capsule. Working from the
inside towards outside, the capillary walls are made up of three layers. These
three layers of the capillary walls make the filteration membrane. The three
members are as follows:

i) Endothelium layer (inner most layer) has relatively large pores (70-100
nanometers in diameter) (Fig. 3.13a). Solutes, and fluid can pass
through the endothelium but not plasma proteins and blood cells. The
proteins associated with the pores of the endothelium are negatively
charged, so they tend to repel negatively charged substances and allow
only positively charged substances to pass more readily.

ii) Glomerular Basement membrane (middle layer) is made up of a

complex mesh of intracellular proteins and is fused to the endothelial
layer. Its job is to prevent plasma proteins from being filtered out of the
bloodstream.

iii) Epithelium layer (outermost layer) consists of specialized cells called

podocytes with extending finger-like arms (pedicels) (Fig. 3.13 b & c) to
cover the glomerular capillaries. They wrap around the capillaries, but
leave gaps/ slits between the pedicels, known as filtration slits. A thin
diaphragm between the slits acts as a final filtration barrier before the
fluid containing inorganic ions and small organic molecules such as
glucose, urea and amino acids pass from the blood plasma into the
lumen of the Bowman’s capsule. Urine formation starts with a size
dependent filtration step. The fluid filtered from the blood of the
glomerulus consists of water and small solutes that are pushed out from
the pores in the capillaries and through the filtration slits into the fluid
filled space inside the Bowman’s capsule. The force required for
filteration is supplied by blood pressure. 77
Block 1 Fundamentals of Animal Physiology
The glomerulus, as you will recall consist of two arterioles - afferent arterioles
which deliver blood to the glomerulus, and efferent arterioles which carry it
away. Constriction of efferent arterioles as blood exits the glomerulus provides
resistance to blood flow in the capillary, preventing a pressure drop. The two
arterioles change in size to increase or decrease the blood pressure in the
glomerulus. In addition, efferent arterioles are smaller in diameter than afferent
arterioles. As a result, the pressurized blood enters the glomerulus through a
relatively wide tube, but is forced to exit through a narrower tube. Together,
these unique features plus the fact that the heart supplies the kidneys with
over a liter of blood per minute (around 20% of its output) helps maintain a
high glomerular capillary pressure so that the filtration function of the kidney
continues, regardless of fluctuations in blood flow.

It is interesting to note that during the formation of this filtrate upto 25% of the
water and solutes are removed from the blood. The renal corpuscles
(glomerulus+ Bowman’s capsule) of the human kidney are capable of
producing 180 L of pre-urine a day but we produce only about 1-2 L of urine
per day which means that about 98% of the fluid volume of pre-urine is
reabsorbed and returned to the bloodstream.

Fig. 3.13: a) Structure of the renal capsule in humans showing Bowman’s

capsule containing the glomerulus; b) Cross section of glomerulus
showing its three filteration membranes; c) structure of a fenestrated
capillary (adapted from The Glomerulus: The Sphere of Influence: Martin
R. Pollak, et al CJASN May 2014, CJN.09400913; DOI:
https://doi.org/10.2215/CJN.09400913).

SAQ 5
a) Name the three layers of the capillary walls which make the filteration
membrane.

b) Name the three main processes which take place in the formation of the
final urine which is excreted.

78
Unit 3 Osmoregulation and Excretion
3.6.2 Modification of Urine
The pre-urine which is the glomerular filterate that enters the Bowman’s
capsule is isosmotic to plasma. This pre-urine leaves the Bowman’s capsule
and enters the proximal convoluted tubule (PCT) where it begins to be
modified and its volume is reduced by about 80%.The fluids inside the tubule
contain water, small solutes such as glucose, urea, amino acids, vitamins and
electrolytes. It is important to remember that some of these molecules are
wastes while the others are valuable nutrients.

Modification of Urine in the Proximal Convoluted Tubule

The epithelial cells of the PCT bear a series of projections termed microvilli -
that face the lumen of the uriniferous tubule (nephron). These microvilli
increase the surface area of these cells providing more space for membrane
proteins that act as pumps, cotransporters and channels. The epithelial cells in
the proximal tubule have a greater number of mitochondria as movement of
substances across the epithelium takes place through active transport
mechanisms. The proximal tubule is the region of the nephron where
reabsorption of water and varying amounts of salts and other solutes from the
pre-urine takes place. Figure 3.14 summarizes the molecular mechanism
believed to be responsible for this selective reabsorption. You can see in the
figure that the basolateral membrane of the cells in the proximal tubule
contains the Na+/K+ -ATPase pump and that the apical membrane has various
protein channels and cotransporters. The Na+/K+-ATPase pumps out sodium
ions from the cells and this creates a gradient for the Na ions to enter into the
cells from the lumen of the tubule. In the apical membrane adjacent to the
lumen of the uriniferous tubule, Na+ - dependent cotransporters use this
gradient to remove valuable ions and nutrients selectively from the pre-urine.
Thus, along with the Na+, another solute like glucose, or amino acids or Cl- or
vitamins is transported into the cell against its gradient.

Fig. 3.14: Water and electrolyte are reabsorbed in the proximal convoluted
tubule. Brush border on the apical membrane of the epithelial cells of
the tubule increases the surface area, facilitating the entry of solutes
through cotransporter molecules and water through aquaporins. The
basolateral plasma membrane has sodium potassium ATPase pump to
transport the sodium ions and other electrolytes and glucose back in
to the blood through the protein channels. Water as a result follows,
by osmosis. 79
Block 1 Fundamentals of Animal Physiology
As these solutes are removed, the tubular fluid becomes hypoosmotic and the
interstitial fluid surrounding the tubular wall becomes hyperosmotic. This
causes passive diffusion of water out of the tubule to restore isoosmotic
condition between the tubular fluid and the surrounding cortical interstitium.
Water from the tubule lumen enters the cells through special channels known
as aquaporins. Aquaporins can bring in 3 billion water molecules per second.
Aquaporins are located in the basolateral membrane of the epithelial cells and
transport back the water into the blood from the interior of the epithelial cells.
This reabsorption of water is secondary to solute transport and occurs
inevitably irrespective of the water requirements of the body. Hence, it is called
the obligatory reabsorption of water. It has been determined that two thirds
of the NaCl and water that is filtered through the glomerulus is reabsorbed in
the proximal tubule. However the osmolarity of the fluid in the tubule does not
change as the water reabsorption is proportional to the solute reabsorption.

Thus, at this stage the pre-urine that enters the proximal tubule is dilute but
the fluid that leaves the proximal tubule is isotonic though reduced in volume.
As the pre-urine enters the next part of the tubule it contains a high
concentration of waste molecules and low concentration of nutrients.

Countercurrent Multiplication in Loop of Henle

In mammals and birds the fluid from the PCT enters a hairpin shaped loop of
Henle. In most nephrons the loop is short and does not enter deeply into the
medulla or leave the cortex but in about 20% nephrons the loop is long and
enters deep into the medulla. As shown in Figure 3.12 you can see that the
loop of Henle consists of three regions the thin descending limb, the thin
middle loop, and the ascending limb which is composed of the thin ascending
limb and the thick ascending limb. The thin and thick ascending limbs differ in
the thickness of their walls. Ion transport is different in each of these
segments as the descending limb is permeable to water and the
ascending limb is permeable to solutes. It took almost a century, before an
explanation was offered for the function of the loop of Henle function. In 1951
B Hargitay and Werner Kuhn proposed that the Loop of Henle acts as a
countercurrent multiplier system of urine-concentrating mechanism in
mammalian kidneys. This means that the loop of Henle sets up an osmotic
gradient which results in exchange of water and solutes between cell of
different regions of the loop as the urine goes down from the cortex to the
medulla. As a results water is reabsorbed from the tubular fluid and
concentrated urine is produced. This is the mechanism that prevents you from
producing litres and litres of dilute urine every day, and it is also the reason
why you don’t need to be continually drinking in order to stay hydrated. The
three segments of the loops of Henle have different characteristics that enable
countercurrent multiplication (Fig. 3.15).

• The thin descending limb is passively permeable to water but almost

impermeable to solutes. As the loop of Henle enters in the medulla
region the surrounding fluid is rich in salt concentration because of
reabsorption of solutes from the ascending limb of the loop of Henle. As
a result the concentration of solutes is increased within the interstitial
space (space between cells), which causes the water in the tubule
80 lumen to move down the concentration gradients into the interstitial
Unit 3 Osmoregulation and Excretion
space until the concentrations within the descending limb and the
interstitial space have equilibrated. In other words we can say, the
tubular fluid becomes steadily more concentrated or hypertonic
(compared to surrounding fluid) as it travels down the thin descending
limb of the tubule. Once the tubular fluid begins to flow up the ascending
limb it stops losing water.

• The thin ascending limb membrane is highly permeable to small solutes

( Na+ and Cl-), moderately permeable to urea, but is impermeable to
water, which means no more water is lost from this part of the loop.
However, Na+ and Cl- move out passively from the tubular fluid, and the
tubular fluid becomes hypotonic as it moves up the ascending limb of the
tubule.

• The thick ascending limb actively transports sodium, potassium and

chloride out into the interstitial fluid by a combination of the Na+/K+
ATPase pump, Cl- channels and Na+/2Cl-/K+ cotransporter molecules
leaving behind urea as the primary solute in the urine. The thick
ascending limb is also impermeable to water, which again means that
water cannot escape from the lumen of this part of the loop. This
segment is sometimes called the “diluting segment”.

Thus, the two sides of the loop of Henle perform opposing functions as
illustrated in Figure 3.15. Furthermore, since the loop of Henle uses energy to
generate an osmotic gradient that enables it to reabsorb water from the tubular
fluid and produce concentrated urine therefore, it acts as a countercurrent
multiplier.

Fig. 3.15: Urine concentration in the nephron. The Loop of Henle with its three
distinct regions. The descending limb is highly permeable to water; the
thin ascending limb is permeable to sodium and chloride ions while it
is impermeable to water; the ascending thick limb removes NaCl by
active transport against the concentration gradient. 81
Block 1 Fundamentals of Animal Physiology
The vasa recta (capillary network that supplies blood to the medulla of the
kidney), are highly permeable to solute and water. As with the loop of Henle,
the vasa recta which forms a parallel set of hairpin loops within the medulla)
around the loop of Henle acts as the countercurrent exchanger
(countercurrent exchanger refers to exchange of a substance between fluids
flowing in opposite directions) due to which water and salt that move out of the
lumen of this loop of Henle quickly diffuse into the (vasa recta).

As the blood flow through these capillaries is very slow, any solute that is
reabsorbed into the bloodstream has the time to diffuse back into the
interstitial fluid, which maintains the solute concentration gradient in the
medulla. This passive process is known as countercurrent exchange.

Urine Volume and Concentration in DCT

By the time the filtrate reaches the distal convoluted tubule of the ascending
limb (DCT) most of the solutes have been reabsorbed. The filtrate entering
DCT thus, has a high concentration of urea and other wastes though it is
hypo-osmotic to the interstitial fluid as NaCl has been transported out of the
tubular lumen. This region is important for transportation of K+, H+ and NH3
into the tubule from the interstitial fluid and Na+, Cl- and HCO3- out of the
tubule into the interstitial fluid. As the salts are pumped out of the lumen, water
also moves out passively. If the body requires additional water it is
reabsorbed here. The activity of the cells of DCT depends on the conditions or
osmotic stress faced by the body and is under hormonal control. Under
conditions of diuresis that is if no water has to be conserved in the body then
the fluid passes unchanged from the DCT in to the collecting duct. When water
needs to be conserved then the hypothalamus secretes the antidiuretic
hormone (ADH) which changes the permeability of the DCT walls to allow
water to move out of the tubule into the interstitial fluid of the kidney in order
to be retained by the body. When ADH titers are high in the body more
aquaporins are recruited in the epithelial cells of the tubule, water diffuses out
into the body and the urine fluid becomes iso-osmotic to blood when it enters
the collecting duct.

In amphibians and reptiles there is no loop of Henle so the fluid from the
proximal tube passes into the distal tubule directly and the final urine may be
hypoosmotic or at best isoosmotic to blood. Urine in these animals is made
hypoosmotic by resorption of salts and under dry conditions the volume is also
reduced by resorption of water in the distal tubule under the influence of ADH.
In lower vertebrates ADH is referred to as vasotocin.

Collecting Duct
The fluid that reaches the collecting duct may be hypoosmotic or isoosmotic
depending on the water status of the animal. Final adjustments in urine
composition are made in the collecting ducts whose walls become permeable
only under the influence of ADH. In the absence of ADH secretion the hypo-
osmotic urine that enters the collecting ducts is excreted unchanged and so a
relatively large volume of urine is excreted. Under situations demanding water
conservation, ADH is secreted and the walls of the collecting ducts become
permeable to water. The urine fluid passing through the collecting ducts,
becomes progressively concentrated and a urine hyperosmotic to blood and
82 isoosmotic to interstitial fluid is excreted.
Unit 3 Osmoregulation and Excretion
Tubular Secretion

Apart from K+, H+, NH3, some organic acids and organic bases that are not
bound to the plasma proteins are secreted from the plasma into the tubular
lumen for excretion. Organic anions are secreted along the sodium gradient
that is established by the N+/K+ pump. These secretory mechanisms are non
specific and as a result many new substances including drug molecules and
toxins can be secreted coupled to the transporting molecules. Most of the
tubular secretion happens in the DCT, but some occurs in the early part of the
collecting duct. Kidneys also maintain an acid-base balance by secreting
excess H+ ions.

SAQ 6
a) Tick the structures impermeable to water:

i) Thin ascending limb of Henle

ii) Thick ascending limb of Henle

iii) Thin descending limb of Henle

b) Why is the loop of Henle a counter current multiplier?

3.7 REGULATION OF KIDNEY FUNCTION

After studying about the structure of kidney and formation of urine in it, we
shall now learn how the function of the kidney is regulated so that water and
electrolyte balance is maintained. You have studied earlier in this unit that
urine production is dependent upon the filtration process. Reduction in blood
supply to the kidney results in a drop in the net filtration pressure and so, urine
production is greatly reduced. Under these conditions, a series of events are
initiated in the kidney itself to develop a hypertensive (increased pressure)
state so that there is better flow of blood to the kidney and resumption of urine
production.

3.7.1 Juxtaglomerular Apparatus

The autoregulation of renal blood flow involves a neuroendocrine
mechanism related to the juxtaglomerular apparatus in the kidney that is
responsible for the rennin- angiotensin - aldosterone regulatory system.

You can note in Fig. 3.16 that the juxtaglomerular apparatus (JGA) is located
between the afferent arteriole and the returning distal convoluted tubule of the
same nephron. It is responsible for regulating both the renal and the entire
body fluid volume status.

As you must have noticed in Fig. 3.16 the juxtaglomerular apparatus has three
components which are follows:

1. The juxtaglomerular cells of the afferent arteriole. Juxtaglomerular

cells synthesize and store renin. Renin acts as an enzyme and hormone 83
Block 1 Fundamentals of Animal Physiology
and is part of a physiological system that regulates blood pressure.
Renin is secreted when there is decrease in arterial blood pressure or a
decrease in sodium load delivered to the distal tubule or to the
sympathetic nervous system. Renin participates in the body's renin-
angiotensin-aldosterone system (RAAS) - also known as the renin-
angiotensin-aldosterone axis,

2. The macula densa, a region of the distal convoluted tubule which is

characterized by densely-packed tubular epithelial cells in comparison to
other regions of the nephron (and thereby leading to its characteristic
appearance under light microscopy). Macula densa detects sodium
concentration of the fluid in the tubule. In response to elevated sodium,
the macula densa cells trigger contraction of the afferent arteriole, thus,
reducing flow of blood to the glomerulus and as a result reducing the
glomerular filtration rate,

3. Mesangial cells, have smooth muscle cell activity. The mesangial cell
contracts or relaxes in response to a number of vasoactive agents. This
ability allows the cells to modify the glomerular filtration locally.

Fig. 3.16: Enlarged Structure of the Juxtaglomerular apparatus in mammals (human).

4.7.2 Renin-Angiotensin-Aldosterone System (RAAS)

Figure 3.17 will help you to understand RAAS. When the afferent arterial blood
pressure is low, the stretch sensitive receptors of the arterioles initiate nerve
impulses which induce the secretion of the proteolytic enzyme called renin by
the juxtaglomerular cells into the blood. This enzyme causes the release of a
decapeptide called angiotensin-I from a large globular plasma protein known
84 as angiotensinogen which is secreted by the liver. Another proteolytic enzyme
Unit 3 Osmoregulation and Excretion
the angiotensin converting enzyme (ACE), removes two amino acids from
angiotensin-I to form an octapeptide called angiotensin-II. ACE is found
mainly in lung capillaries and within the renal endothelium. Angiotensin II is
about 200 times more powerful than norepinephrine in its vasopressor activity
as it exerts a vasoconstricting effect that causes a rise in blood pressure. It
increases the blood pressure by two mechanisms. Firstly, it acts on the
smooth muscle of the arterioles and causes strong vasoconstriction. Secondly,
it stimulates the secretion of aldosterone by the adrenal cortex. Aldosterone
acts on the aldosterone receptors on the distal tubules and collecting tubules
and enhances the reabsorption of Na+ from the kidney tubules and hence
causes a rise in the plasma Na+ level. This results in an increase in the
extracellular fluid volume and consequently an elevation in blood pressure.
Angiotensin-II is degraded by an enzyme called angiotensinase, present in
plasma.

Anti-diuretuc Hormone

ADH is also known as vasopressin and helps in retention of water. It is

synthesized in the neurosecretory cells of the hypothalamus and stored in the
posterior pituitary. ADH acts in the kidney to regulate the volume and
osmolarity of the urine. Specifically, it acts in the cells of distal convoluted
tubule (DCT) and collecting duct (CT) cells.

Let’s consider an example where there is an increase in osmolarity of the

plasma. The increase in ADH release will mean that the permeability of the
DCT and CT cells to water increases, (water leaves the tubules) and volume
urine output decreases. This happens because ADH increases the aquaporin
channels in the apical membrane of the DCT and CT cells. This allows water
to move down its concentration gradient, out of the nephron and back into the
blood stream. This thus, causes an increase in the water reabsorbed from the
urine, a decrease in plasma osmolarity and an increase in total blood volume.
Water reabsorption in this manner is a passive process.

Fig. 3.17: The rennin - angiotensin - aldosterone system that regulates the ionic
and water balance in the system (adapted from Wikipedia). 85
Block 1 Fundamentals of Animal Physiology
Alternatively, when there is a decrease in osmolarity of the plasma then ADH
release is inhibited. This results in a decrease in the number of aquaporin
channels on the apical membrane of the DCT and CT cells. Therefore, there is
a reduction in the amount of water reabsorbed from the nephron back into the
blood stream.

SAQ 7
Fill in the blank spaces in the table given below:

Hormone/Enzyme Action

Renin

Antidiuretic enzyme

Angiotensin converting enzyme (ACE)

Angiotensinase

3.8 PATTERNS OF NITROGEN EXCRETION

Excretion is the removal of toxic waste products of metabolism from the
body. The end products of metabolism are either eliminated or conserved by
an animal depending on its physiology. We know that carbon, hydrogen,
oxygen and nitrogen are the end products of catabolism. Of these carbon
atoms are eliminated in the form of carbon dioxide, hydrogen in the form of
water and oxygen in the form of carbon dioxide and water. When amino acids
are catabolised, the amino group NH2 is released and is either transferred to
another group for reuse or removed. Any amino group that is not reused
needs to be dissolved in water and excreted as soon as possible in order to
avoid toxic rise of waste products in the plasma or must be converted into a
less toxic form before being eventually excreted.

You are aware that nitrogen is a characteristic constituent of amino acids and
proteins. Animals receive amino acids in diet and use them to synthesise a
variety of functional nitrogenous compounds such as nucleic acids, proteins
both enzymatic and non-enzymatic, some hormones, and neurotransmitters.
The amount of amino acids obtained in diet is usually in excess of the need for
the synthesis of the functional nitrogenous compounds. The excess of amino
acids is either catabolised for the release of energy or is used for the
synthesis of glycogen and fat. When amino acids, proteins or nucleic acids are
catabolised, nitrogen-containing excretory end-product such as ammonia
(NH3), or urea or uric acid which are formed are excreted. Excretion of
nitrogen, or urea or uric acid is closely related to the normal habitat of the
animal and to the availability of water.

Animals are often grouped according to their main excretory products as

follows:

i) Those that excrete mainly ammonia (NH) as the end-product of protein

86 metabolism are called ammonotelic.
Unit 3 Osmoregulation and Excretion
ii) Those that excrete mostly urea are ureotelic.

iii) Those that form mainly uric acid are uricotelic.

iv) Those that secrete guanine are guanotelic.

Ammonia is a strong base and it gains a proton to form the ammonium ion
(NH4+). Ammonia is toxic to cells and high concentration of ammonia raises
the pH of intracellular and extracellular fluids to an extent which is enough to
poison the enzymes in the cells. Different animal species have evolved various
strategies to prevent this from happening depending on their habitat. Those
that are aquatic can get rid of the ammonia because it diffuses easily in water
according to a concentration gradient. However, in terrestrial species ammonia
is converted into less toxic compounds like urea that can be excreted in urine
or uric acid which is a white paste like substance that you must have seen in
bird faeces. You already studied formation of these catabolic products of
nitrogenous compounds in the body in your graduation biochemistry course
so we are not discussing their formation here.

Excretion of nitrogen of ammonia or urea or uric acid as mentioned before is

closely related to the normal habitat of the animal and to the availability of
water. Table 3.2 shows that the type of nitrogenous waste produced by an
animal correlates with the type of osmotic stress it has to face in its natural
habitat. In all animals nitrogen excretion is not restricted to a single product.
Animals are designated as ammonotelic, ureotelic, uricotelic or guanotelic only
to indicate the predominant form in which nitrogen is excreted by them. The
term uricotelism, for example, does not preclude the excretion of ammonia and
urea in minor quantities.

The mode of nitrogen excretion may vary within a given group of animals
depending on the habitat of the species. For example, South African frog,
Xenopus laevis, which is aquatic in adult life is ammonotelic whereas, many of
the truly amphibious frogs and toads (e.g., Bufo species) excrete urea as
adults. Similarly, the aquatic chelonians (tortoise and turtles) excrete more or
less equal proportions of urea and ammonia. The semiaquatic forms are
ureotelic and the desert living forms are uricotelic.

Table 3.2: Characteristics of nitrogenous wastes produced by animals.

Characteristic Ammonia Urea Uric acid

Solubility in water High Medium Very low

Amount of water High Medium Very low

required for
excretion

Amount of energy Low High High

(ATP) required for
its excretion

Toxicity High Low Medium

87
Block 1 Fundamentals of Animal Physiology
Animal group Fish, aquatic Mammals, Birds, reptiles,
where it is the invertebrates sharks most terrestrial
primary waste insects, spiders

Method of Breakdown In liver starts Synthesis starts

synthesis of amino with amino with amino acids
acids and groups, and nucleic acids
nucleic acids generated from catabolic
from catabolic process of amino
process of acids and
amino acids nucleotides.

Method of In urine and In urine In faeces (in birds

excretion diffuses (mammals); uric acid is
through gills diffuses across derived from
gills (sharks) urine but is
excreted along
with the faeces)

SAQ 4
Fill in the blanks.

a) The mode of …………may vary within a given group of animals

depending on the habitat of the species.

b) The animals which secrete urea are called……………

c) Excretion of nitrogen, or urea or uric acid is closely related to the normal

habitat of the animal and ……………………

3.9 SUMMARY
Let us sum up what we have learnt in this unit:

• Osmoregulation is a process for the maintenance of osmotic

concentration of the body fluids. Animals have adapted various
physiological and behavioural mechanisms to cope up with the rigors of
the osmotic environments.

• Osmoregulation requires the exchange of salts and water between the

extra- cellular environment and the external environment to com-
pensate for obligatory, or uncontrolled, losses and gains. The transport
of solutes and water across epithelial layers is fundamental to all
osmoregulatory activity. The obligatory exchange of water depends on
(1) the osmotic gradient that exists between the internal and external
environments, (2) the surface-to-volume ratio of the animal, (3)the
permeability of the integument, (4) the intake of food and water, (5) the
evaporative losses required for thermoregulation, and (6) the disposal of
88 digestive and metabolic wastes in urine and feces.
Unit 3 Osmoregulation and Excretion
• Marine and terrestrial animals are faced with dehydration, whereas
freshwater animals must prevent hydration by uncontrolled osmotic
uptake of water. Marine birds, reptiles, and teleosts replace lost water by
drinking seawater and actively secreting salt through secretory epithelia.
Freshwater fishes do not drink water; they replace lost salts by active
uptake. Birds and mammals are the only vertebrates that secrete a
hypertonic urine. Many desert species in addition utilize mechanisms for
minimizing respiratory water loss.

• A variety of tubular organs are present in animals for the elimination of

unwanted materials and for osmotic and ionic regulation. Despite their
morphological diversity, their functional principles are remarkably similar,

• Osmoregulatory and excretory organs in invertebrate animals are the

contractile vacuoles of protists (protozoa), nephridia of worms,
molluscan kidneys, green glands of crustaceans, Malpighian tubules of
insects. The osmoregulatory and excretory organs of the vertebrates is
the paired kidney.

• Catabolism of proteins gives rise to nitrogen which is highly toxic to the

animal. Therefore, animals get rid of the toxic nitrogen in the form of
ammonia or convert it into less toxic forms such as urea and uric acid
before it is being excreted,

• Animals which predominantly excrete ammonia are called ammonotelic,

those that excrete urea are known as ureotelic ,uric acid excreting
animals are known as uricotelic and guanine excreting animals are
called guanotelic.

• The nephron is the functional unit of the vertebrate kidney, which actively
filters blood and forms urine. The nephron filters and exchanges water
and solutes with two sets of blood vessels and the interstitial fluid in the
kidneys,

• The glomerulus is a capillary bed that filters blood principally based on

particle size. The filtrate is captured by Bowman’s capsule and directed
to the PCT. A filtration membrane is formed by the epithelium, the fused
basement membranes of the podocytes and the capillary endothelial
cells.

• In vertebrates preurine filtrate enters the PCT of the nephron (uniforms

tubule). Absorption and secretion of solutes, several substances and
water occurs in the PCT, descending and ascending limbs and the loop
of Henle. Absorption and secretion continues in the DCT but to a lesser
extent than in the PCT. Each collecting duct collects formed urine from
several nephrons and responds to ADH hormone secreted by the
hypothalamus.Secretion of ADH causes the thick ascending loop of
Henle to recruit aquaporins water channels in cell membrane so that
water is rebsorbed from tubule back into the kidney tissues.

• Osmoregulation and excretion from kidneys is regulated by hormones.

• Regulation of urine output is regulated in the kidney by the Renin-

angiotensin - aldosterone system (RAAS). 89
Block 1 Fundamentals of Animal Physiology
• Contractile mesangial cells of the juxtaglomerular apparatus perform a
role in regulating the rate at which the blood is filtered in the glomerulus.
Specialized cells in the JGA produce signals to regulate blood flow and
filtration rates of the glomerulus by triggering the rennin - agiotensin-
aldosterone system, which plays a central role in homeostasis and blood
pressure regulation.

3.10 TERMINAL QUESTIONS

1. Write short notes on :

a) Green gland of crustaceans

b) Molluscan kidney

2. Differentiate between osmoregulators and osmo conformers.

3. Discuss the pattern of nitrogen excretion.

4. Draw a diagram showing the urine concentration in the nephron.

5. Summarize with the help of a diagram the movement and process of

urine formation in the mammalian nephron.

3.11 ANSWERS
Self-Assessment Questions
1) a) i) ions/solutes

ii) isotonic

iii) hypotonic, isoosmotic

iv) water, solutes

b) Na+/K+ ATPase pump functions to maintain osmotic balance and

electric gradient across biology/cell/plasma membranes by taking
3 Na+ out of the cell and at the same time bringing back 2 K+ into
the cell. The energy required for this comes from the
phosphorylation of this protein pump by ATP. Na+/K+ ATPase
pump is essential as it helps maintain the basal resting potential of
the cell/plasma membrane, and also helps maintain the osmolarity.
The pump also affects transport of electrolytes and regulates the
cellular volume.

2. a) apical, basal, basal basement membrane

b) excretion

c) ammonotelic, ureotelic

3. a) i) Malphigian tubule - insects

90 ii) Contractile vacuole - protists (amoeba)

Unit 3 Osmoregulation and Excretion
iii) Nephridia – worm (annelids)

iv) Green gland - crustacean

b) Draw Figure 3.6 and explain it with the help of Subsection 3.5.1.

4. a) Cortical nephrons - that have their glomeruli located nearer to the

enter part of the cortex and their short loops of Henle do not
extend into the medulla.

b) Juxtamedullary nephrons- that have their glomeruli located deep

within the cortex near the cortica medullary border and their long
loops of Henle extend deep into the medulla.

c) The blood enters via the afferent arteriole in the glomerular

capillaries.

d) The blood leaves via the efferent arteriole rather than via a vein as
is the case with other capillary networks.

5. a) The three layers of the capillary walls which make the filtration
membrane.

i) Epithelium (outer most layer)

ii) Glomerular Basement membrane (middle layer)

iii) Endothelium (innermost layer)

b) Name the three main processes which take place in the formation
of final urine that is excreted

i) Filtration

ii) Tubular reabsorption

iii) Tubular secretion

6. a) ii) Thick ascending limb of Henle,

iii) Thin descending limb of Henle

b) Refer to Section 4.7.3.

7.

Hormone/Enzyme Action

Renin regulates blood pressure

Anti diuretic enzyme helps in retention of water

Angiotensin converting removes two amino acids from

enzyme (ACE) angiotensin-I to form an octapeptide
called angiotensin-II

Angiotensinase degrades angiotensin-II.

91
Block 1 Fundamentals of Animal Physiology
Terminal Questions
1. Refer to Section 3.4.

2. Refer to Section 3.3.

3. Refer to Section 3.5.

4. Refer to Fig. 3.16.

5. Refer to Section 3.7, Fig. 3.13.

92
 UNIT 4
MUSCLE PHYSIOLOGY

Structure
4.1 Introduction Role of Calcium and Regulatory
Proteins
Objectives
Sequence of Molecular Muscle
4.2 Types of Muscles
Movement Cross Bridge Cycle
Smooth Muscles
Muscle Tone
Cardiac Muscles
4.4 Energetics of Muscle
Skeletal Muscles Contraction
4.3 Structure and Movement of 4.5 Adaptations of Muscles to
Vertebrate Skeletal Muscles Perform Different Activities in
Vertebrates
Structure of Vertebrate Skeletal
Muscles 4.6 Summary
Microscopic Anatomy of a 4.7 Terminal Questions
Skeletal Muscle Fiber
4.8 Answers
Molecular Basis of Muscle
Movement

Sliding Filament Mechanism

Nerve and Blood Supply

Neural Control of Muscle

Contraction

4.1 INTRODUCTION
In this block you have already read about the physiology of circulatory system,
respiratory system, osmoregulation, and excretion in animals. In this unit of
Block 1, you will learn about the physiology of movement in animals. Animals
sense the changes in the environment and respond accordingly and
appropriately. These responses usually are in the form of movement of some
sort. From one celled organism to humans, all respond to external and
internal stimuli and the response is usually in the form of movement.
Movement is a characteristic and fundamental property of animals. Often, we
Block 1 Fundamentals of Animal Physiology
consider movement in connection with locomotion, i.e., an organism moving
from place to place, or a cell moving away from or towards a source of
stimulus. However, even animals that remain attached and never show
locomotion, such as corals and sponges; barnacles and sea anemones exhibit
a variety of movements. They rely on the movement of their individual cells to
accomplish circulation, respiration, reproduction etc.

There are three basic mechanisms used by animals and individual cells to
achieve motion. These are: i) muscular movement, ii) amoeboid movement, iii)
ciliary and flagellar movement.

Muscular movement is the fundamental mechanism used in most animals for a

variety of movements. Muscle can exert force by shortening which is called
‘muscle contraction’. This force is the basis of all kinds of movements both
internal as well as external in animals. Individual cells also move. The internal
movement of individual cells is due to cytoplasmic streaming and their external
locomotion is brought about by amoeboid movement (which derives its name
from locomotion in Amoeba), or by cilia (sing: cilium) and flagella. Cilia are
found in all animal phyla and serve a variety of functions. They enable
movement of entire cells such as seen in the movement of single celled
Paramecium in water, set up currents that move water through the water
vascular systems of various invertebrates of which echinoderms are a good
example. Ciliated cells also line the lungs and respiratory tract of vertebrates
and slowly remove the foreign particles that lodge on cell surfaces. The single
celled sperms as you will recall also move with the aid of a tail that is a
flagellum.

In this unit you will learn about the structure of different types of muscles and
the mechanism of muscle contraction involved in the movement of skeletal
muscles. You will also learn about the various adaptations in muscle
physiology of animals to perform different functions.

Objectives
After studying this unit you should be able to:

 differentiate between the structure and function of skeletal, cardiac and

smooth muscles,

 explain the molecular basis of muscle contraction of skeletal muscles,

 describe the mechanisms that regulate skeletal muscle contraction,

 identify the components of the neuromuscular junction, and summarize

the events involved in the neural control of skeletal muscle contraction
and relaxation,

 describe the energetics of muscle contraction, and

 appreciate the adaptation in muscle physiology of animals to perform

94 different functions.
Unit 4 Muscle Physiology

4.2 TYPES OF MUSCLES

Before we start our study of muscle movement we must learn about the
muscle tissue and their types present in animals. Muscle cells or myocytes
have the special ability to contract in order to produce movement in the body
in organs and in the body as a whole. The contractile proteins actin and
myosin present in the muscle cells grant the muscle cells contractibility.
Vertebrate muscles can be categorized into three types: skeletal muscles,
cardiac muscles and smooth muscles (Fig 4.1 and 4.2). Each type of
muscle has a unique structure and specific role. However, all three muscle
tissues have the property of excitability- as their plasma membranes can
change their electrical state (from polarized to depolarized) and send an
electrical wave called an action potential along the entire length of the
membrane that results in the contraction of the muscle. Differences among
the three muscle types include the microscopic organization of their contractile
proteins, their location in the body, and their control mechanisms.
Morphologically vertebrate muscle tissues can be distinguished as
smooth and striated.

Smooth muscle cells do not show striations under the microscope.

Striated muscles on the other hand when seen under a microscope appear to
have alternating transverse dark and light bands, and hence they are
named as striated muscles. Striated muscles can be divided into two types
of muscles: skeletal and cardiac.

4.2.1 Smooth Muscles

Smooth muscle cells are fusiform and uninucleated (Fig. 4.1). Smooth
muscle cells contain contractile actin and myosin protein filaments called
myofibrils. These myofibrils are not arranged in a pattern in the cell, instead
they are scattered in the cell.

The mechanism of contraction of smooth muscles and the striated skeletal

muscles is similar. However, smooth muscle cells in comparison show greater
elasticity and can maintain their contractibility even when stretched. A
stretched smooth muscle adapts to its new length and retains its ability
to contract on demand. This ability of the smooth muscle cells is known as
plasticity and is especially important in digestive organs which are subject to
changes in volume (like the stomach). Smooth muscle contractions can be just
as powerful as those of skeletal muscle which are the striated muscle cells
and about which you will also learn in this section.

Smooth muscles in addition to being found in the stomach are also present in
the walls of internal organs such as large and small intestines, gall bladder
and large blood vessels. Contraction of smooth muscles causes the diameter
of blood vessels, gall bladder etc. to change by increasing or decreasing their
diameter. Smooth muscles also help propel the food down through the
digestive tract. Smooth muscles are known as involuntary muscles as they
are not under conscious control but are controlled by the autonomic nervous
system. 95
Block 1 Fundamentals of Animal Physiology

Fig. 4.1: a) Smooth muscle cells are uninucleated and regulated by the
autonomic nerves, therefore, not usually under voluntary control of the
animal; b) photomicrograph of a longitudinal section of a vertebrate
smooth tissue showing smooth muscle cells. (acknowledgement -Hill,
M.A. (2020, April 27) Embryology ANAT2241 Muscle Tissue :
from https://embryology.med.unsw.edu.au/embryology/index.php/ANA
T2241_Muscle_Tissue)

4.2.2 Cardiac Muscles

Cardiac muscles, as the name suggests are the muscles of the heart (Fig
A4.2 a). Cardiac muscles appear similar to skeletal muscle cells as similar to
skeletal muscles they are also striated (Fig A.4.2 b & c). However, the
striations in cardiac muscles are not aligned as seen in skeletal muscles.
Cardiac muscle cells are not as long as the skeletal muscle cells. Cardiac
muscles contain a large number of mitochondria and abundant reserves of
myoglobin as they solely depend on aerobic metabolism for their energy which
is required for heart muscle contraction.

Cardiac muscle cells are branched and mononucleated although a few may
have 2 or more nuclei (Fig. A 4.2b). One end of a cardiac muscle fibre (cell) is
connected to another cardiac muscle fibre by means of intercalated discs. The
branching of the cardiac muscle ensures that there is swift communication
between the cells, allowing coordinated wave like contractions. Thus, cardiac
muscle cells are specialized for continuous rhythmic contractions of the heart
which is needed for pumping blood. Cardiac muscles are not under
voluntary control and so are involuntary muscles. Their rhythmic
contractions (as you will recall from earlier “Unit 1-Circulatory system” of this
course) are regulated by the sinoatrial node of the heart, which serves as
96 the heart's pacemaker.
Unit 4 Muscle Physiology
4.2.3 Skeletal Muscles
Skeletal muscles are attached to the bones in the arms, legs, vertebral
column, and other parts of the skeleton (Fig. B 4.2 a) (a few skeletal muscles
attach to and move the skin or other skeletal muscles). The skeletal muscles
cells are long and multinucleated showing regular stripes or striations under
the microscope (Fig. B 4.2 b & c).These striations are due to the regular
arrangement and alignment of contractile protein myofibrils the actin and
myosin present in them.

Fig. 4.2: Striated muscles: A) Cardiac muscles (a) are present only in the heart.
(b) external appearance of cardiac muscles. (c) photomicrograph of a
longitudinal sections of the vertebrate cardiac muscle tissue showing,
cardiac cells which usually have one nucleus per cell. Cardiac cells are
branched and have intercalated discs that connect the cells; B) Skeletal
muscle fibers (a) are attached to bones; (b) external appearance of
skeletal muscles;(c) photomicrograph of a longitudinal section of a
vertebrate skeletal tissue showing skeletal muscle cells or fibers that
are multinucleated with definite striations (Acknowledgements-
photomicrograph of muscle cells or fibers (Retrieved from Hill, M.A.
(2020, April 27) Embryology ANAT2241 Muscle Tissue: from https://
embryology.med.unsw.edu.au/embryology/index.php/ANAT2241_Muscl
e_Tissue. 97
Block 1 Fundamentals of Animal Physiology
Skeletal muscles are the major muscles in the body and are responsible for all
the activities like walking, movements of the head, hand, etc. These muscles
are usually under voluntary control of the individual and exclusively get their
signals to contract through the spinal or cranial nerves. We will learn about the
structure and working in greater detail in the next section.

SAQ 1
a) List the three types of muscle tissue and give their major functions.

b) Fill in the blanks and compare your answers with those given at the end
of this unit:

i) ……………………… and ……………………. are called striated

muscles because under the microscope they exhibit transverse

light and dark bands.

ii) ……………….. muscles are under the control of will therefore are
usually called ……………….. while ……………. and -------------
muscles are not under the control of the conscious mind and are
therefore called ………..... .

4.3 STRUCTURE AND MOVEMENT OF

VERTEBRATE SKELETAL MUSCLES
Let us now study the structure and functioning of skeletal muscles. Recall from
the previous section that the most important property of skeletal muscle is
contraction and the resulting movement. Skeletal muscles not only cause
movement but also stop movement and resist gravity to maintain the body’s
posture. Each skeletal muscle is made up of integrated tissue. Let us learn
about the organization and ultra-structure of the skeletal muscle before we
study the mechanisms that cause it to contract.

4.3.1 Structure of Vertebrate Skeletal Muscles

Look at Figures 4.3 and 4.4 while you read this description of the structure of
the skeletal muscle. A skeletal muscle contains connective tissues, blood
vessels, nerves and skeletal muscle tissue. The entire skeletal muscle (Fig.
4.3 a & b, and Fig. 4.4 a & b) is enclosed by a connective tissue which is in the
form of a dense layer of collagen known as epimysium (Fig. 4.3 a & b, and
Fig. 4.4 a b& c) which contains blood vessels and nerves. The epimysium
separates the entire muscle from the surrounding tissues and allows it to
contract and move powerfully while still maintaining its structure.

A cross section (CS) of a skeletal muscle shows it to be further divided into

compartments by another layer of connective tissue known as perimysium.
Perimysium lies internal to epimysium (Fig. 4.3 a & b, Fig. 4.4 a, b & c). The
perimysium contains collagen, elastin. It encloses the blood vessels and
nerves that supply the muscle. Each compartment formed by the perimysium
98 contains a bundle of muscle fibers known as fascicle. This fascicular
Unit 4 Muscle Physiology
organization is common in muscles of the limbs. Each fascicle is made up of
several long, cylindrical, multinucleate muscle cells or muscle fibers,
arranged parallel to each other. The multinucleate structure of the muscle fiber
is due to the fact that during the development of the muscle tissues, the
individual precursor cells fuse together to form a new, combined mature
muscle fiber structure with many nuclei. That is why muscle fibers do not
follow the general rule of one nucleus per cell and each fiber is made up of
several cells that are combined into one. Some adult muscle fibers still have
one of these tiny precursor or stem cells called satellite cells attached to their
outer boundary (Fig. 4.4 d). During strength training these satellite cells being
stem cells fuse with myocytes in order to make bigger muscle fibers. The
satellite cells can also become active after a muscle injury to produce more
muscle fibers. Scientists are still trying to understand this mechanism better in
the hope of developing new therapies for replacing lost muscle tissues.

Each individual muscle fiber in the fascicle forms the individual contractile
units within a muscle. Each muscle fiber is enclosed in a delicate, flexible and Aponeurosis is a flat
elastic connective tissue layer – the endomysium that interconnects adjacent sheet or ribbon of
tendonlike material
muscle fibers. This endomysium layer also contains blood capillaries, and
composed of dense,
nerve fibers that supply the muscle fibers and also the embryonic muscle cells fibrous connective
(which function in muscle repair). tissue containing
fibroblasts (collagen-
In a skeletal muscle every muscle fiber is supplied by the axon branch of a secreting spindle-
somatic motor neuron, which signals the fiber to contract. Unlike cardiac and shaped cells) and
smooth muscles, the only way a skeletal muscle contracts functionally is bundles of
through signaling from the nervous system. collagenous fibres in
ordered arrays.
All the three connective tissue layers namely, epimysium, perimysium and
endomysium, are interwoven together and form the tendon at the end of the
muscle or a broad sheet called aponeurosis. Tendons and aponeurosis
connect the skeletal muscle to the bone (Fig 4.3 a and 4.4 a).

Fig: 4.3: a) Organization of the skeletal muscle showing the three connective
tissue coverings- epimysium, perimysium and endomysium, which are
interwoven together to form the tendon at the end of the muscle, which
connects to the bone tendon: b) diagrammatic representation of the
arrangement of the three connective tissue coverings of the muscle in
cross section. 99
Block 1 Fundamentals of Animal Physiology
4.3.2 Microscopic Anatomy of a Skeletal Muscle Fiber
Let us now examine the microscopic structure of the skeletal muscle cell or
fiber as it will help you to understand the physiology of muscle contraction in
skeletal muscles.

A skeletal muscle cell or fiber is different from a typical cell that you have read
about earlier. It is much larger and longer than most cells in the body and is
multinucleate. Furthermore, as mentioned earlier the skeletal muscle fibers are
arranged parallel to each other.

Fig: 4.4: a) Organization of the skeletal muscle and its angle of cross section
showing : b) cross section of the skeletal muscle; c) cross sections of a
muscle fascicle composed of a bundle of muscle fibres; and d) cross
section of an individual muscle fibre (cell) showing myofibrils.

Sarcoplasma

Each skeletal muscle fiber or muscle cell is enclosed by the plasma or cell
membrane known as sarcolemma. The sarcolemma encloses the
cytoplasm of the muscle fiber known as sarcoplasm. The sarcoplasm of
each muscle fiber contains hundreds of nuclei located, scattered in it (Fig. 4.4
a). The advantage of the skeletal muscle fiber being multinucleate is that the
multiple copies of genes in these nuclei can direct the production of proteins
and enzymes needed for muscle function at a much faster rate than it could
have done if there was only one nucleus per cell. The sarcoplasm of each
skeletal muscle fiber cell when seen under the microscope appears to be
stuffed with little thread like contractile organelles called the myofibrils which
100 are unique to them (Fig 4.4 d).
Unit 4 Muscle Physiology
Sarcolemma and Transverse Tubules

The sarcolemma of each muscle fiber cell at various points, extends inwards
at a right angle to the long axis of the muscle fiber cell into the sarcoplasm
towards the center of the muscle fiber to form Transverse (T) tubules. These
T tubules thus, extend transversely across the sarcoplasm of each muscle
fiber, (Fig 4.5). The T tubules DO NOT open into the interior of the muscle
cell instead, after returning from the muscle they open somewhere else
on the sarcolemma.T tubules are filled with interstitial fluid. Several
Transverse (T) tubules of the sarcolemma of a muscle fiber, pass down into
the muscle fiber cell and go around the little thread-like contractile myofibrils
present in large numbers in each skeletal muscle cell. Thus, the myofibrils
are encircled by the T tubules.

Fig.4.5: a) Structure of a single skeletal muscle fiber cell showing its striated
appearance and nuclei; b) Longitudinal view and cross section of a
single muscle fiber cell (enlarged) showing the organization of
myofibrils; c) Cutaway of a muscle fiber showing position of the
sarcoplasmic reticulum surrounding the myofibrils; d)The triad
structure formed by the Transverse(T) tubules and sarcoplasmic
reticulum in skeletal muscle fiber cell; Longitudinal section of a muscle
fiber cell showing the location of myofibrils and triads in it. 101
Block 1 Fundamentals of Animal Physiology
Sarcolemma like other plasma membranes has a transmembrane potential
difference. A sudden change in its membrane potential causes the muscle
fiber to contract. Even though the muscle fiber is quite long it can contract
quickly and simultaneously and this can happen only if the signal is
transmitted quickly throughout the muscle fiber. This quick and simultaneous
contraction of the muscle is due to the presence of the unique system of the
interconnecting transverse tubules (T tubules) in the muscle fiber which
enables for the quick transmission of signal. Muscle action potentials travel
along the sarcolemma and throughout the T tubules, quickly spreading
throughout the muscle fiber. This arrangement ensures that an action potential
excites all parts of the muscle fiber at essentially the same instant.

Inside the sarcoplasma of each muscle fiber the T tubules attach to the
sarcoplasmic reticulum(SR), at the point at which they encircle the myofibril.
The sarcoplasmic reticulum(SR) is similar to the endoplasmic reticulum of
cells and flanks the T tubules on either side. The sarcoplasmic reticulum
expands on either side of the T tubule and forms chambers called terminal
cisternae. The combination of a pair of terminal cisternae plus T tubuleis
known as a triad (Fig 4.5 c, d & e). The function of the triad is to conduct
electrical signals from the surface of the cell sarcolemma down and deeper
into the muscle cell, and then specifically, into the sarcoplasmic reticulum.

The terminal cisternae of the sarcoplasmic reticulum store Ca2+ which are
actively transported from the sarcoplasm into the terminal cisternae. The
concentration of bound and free Ca2+ in these cisternae can be up to 40,000
times more than that in the surrounding sarcoplasm. These storehouses of
calcium release calcium ions in the region where the thick and thin
contractile filaments contained in each myofibrils interact.

Myofibrils of Sarcomere

The sarcoplasm of each skeletal muscle fiber at high magnification, as

mentioned earlier is packed with a thousand or more of tightly packed
contractile myofibrils (Figure 4.4 b &4.5 b, c &4.6). These myofibrils are 1-2
μm in diameter and are as long as the cell. Myofibrils, in turn, are made up of
bundles of still finer fibers called myofilaments. You can see in Figures 4.5 c
and 4.6 at a glance that some of the filaments are much thicker than others.
The thicker fibers are myosin filaments while the thinner are actin filaments.
Although in the diagrams they are shown in contrasting colors however, they
are colorless.

The myofilaments of the myofibrils are made up of mainly three kinds of

protein filaments: 1) One type of protein filaments consist of the contractile
filaments which are of two types: i) thin actin filaments known as thin filaments
or simply actin and ii) the thick motor protein filaments called myosin filaments
or thick filaments or simply myosin .2) The second types of protein filaments
are those that stabilize the position of the actin and myosin filaments. While 3)
the third type of proteins are those that regulate the interactions between the
thick and thin filaments.

The thick and thin actin and myosin filaments respectively of the myofibril (Fig
102 4.6 a, b, c & d) do not extend the entire length of the muscle fiber, instead they
Unit 4 Muscle Physiology
are organized into compartments or repeated units known as sarcomeres (Fig
4.6a, b, c& d) that are arranged end to end. Sarcomeres are the functional
units of the muscle fibers. Narrow, plate-shaped regions of dense protein
material called Z discs or Z lines separate one sarcomere from the next. A
sarcomere is thus “the distance between two consecutive Z discs or Z
lines”. Therefore, a sarcomere extends from one Z disc to the next Z disc.

The distribution, arrangement and size of the thick and thin filaments give the
myofibril its typical banded or striated appearance. Figure 4.6 b,c and d,
shows the composition of a resting sarcomere. In a resting sarcomere you can
see the alternating dark and light bands present in it. The dark bands are
known as the A bands and the light bands are known as the I bands. The
structures and their location in the sarcomere are as follows (Fig 4.6 b, c & d):

Fig. 4.6: a) Cutaway section of a muscle fiber showing myofibrils; b) Longitudinal

view of a single myofibril showing asarcomere; c) diagrammatic
representation of a magnified view of a relaxed sarcomere showing the
position of the actin and myosin filaments and the various bands or areas
formed as a result and d) electron photomicrograph of a sarcomere
showing the position of the actin and myosin filaments and the various
bands or areas formed as a result. 103
Block 1 Fundamentals of Animal Physiology
1. A-BAND - The A band is the darker middle part of the sarcomere which
extends the entire length of the thick myofilaments. The length of the A
band is thus equal to the length of the myosin filaments and can be
further divided into three regions (see Fig. 4.6 to understand the
arrangement):

a) The central M (middle) line of the myofibril - is made up of

proteins that connect the central portion of each thick filament to
the neighboring thick filaments and so help stabilize the position of
all the thick filaments.

b) The H Zone - in a resting sarcomere is a slightly lighter band on

each side of the M line. This zone contains only myosin but no
actin.

c) The zone of overlap - consists of overlapping thick and thin

filaments that gives the striated appearance to the muscle. The
extent of overlap of the thick and thin filaments depends on
whether the muscle is contracted or relaxed. The pattern or the
length of distance of their overlap, gives rise to a variety of zones
and bands (Fig. 4.6 c & d).

2. The I. BAND-contains only the actin filaments and extends between

the A band of one sarcomere and the A band of the next sarcomere (Fig.
4.6 c).

3. The Z lines- form the boundaries of each sarcomere as mentioned

earlier. The Z lines are made up of a protein called actin (Fig 4.8 b).
The actin filaments of adjacent sarcomeres are attached to each other
by actinin protein. Actin fibers extend on each side of the Z lines towards
the center of the sarcomere making up the zone of overlap. Strands of
another protein titin extend from the tips of the myosin fibers and attach
to the Z lines, keeping the thick and thin filaments aligned and
preventing the sarcomere from stretching too far as this would disrupt
the contraction mechanism (Refer also to Fig 4.8 b).

At each end of the muscle fiber the myofibrils are attached to the inner surface
of the sarcolemma.The outer surface of the sarcolemma is attached to the
collagen fibers of the tendon by which the muscle is attached to the bone
(Refer again to Figs 4.3 and 4.4). Thus, when the myofilaments contract the
entire muscle cell shortens and the muscle is pulled. Scattered among the
myofilaments in the muscle cell are mitochondria and glycogen granules (the
storage form of glucose in animals) that provide the energy for the muscle
activity. Let us now look at the molecular structure and mechanism
responsible for skeletal muscle contraction.

4.3.3 Molecular Basis of Muscle Movement

You have studied in the earlier section that the thick filament is made up of
myosin and the thin filaments are mainly made up of actin. Each thick
filament is 10-12 nm in diameter and 1.6μm in length. It is roughly
composed of about 500 myosin molecules. Each myosin molecule has six
104 polypeptide chains and contains two heavy chains.
Unit 4 Muscle Physiology
Each heavy (Fig 4.7a) chain is organized into three structurally and
functionally different domains: i) the globular head domain that contains actin
and the ATP-binding sites and is responsible for generating force. This
globular head domain is the most conserved region among the various
myosins. Adjacent to the head domain lies ii) the α-helical neck region,
which is associated with two light chains and regulates the activity of the head
domain. After the neck domain and distal to it is the iii) tail domain
which contains the binding sites for the actin filaments.

Each myosin molecule appears as a double stranded globular protein twisted

around each other with two heads, two necks and a tail (Fig 4.7a). Each thick
filament as mentioned earlier contains about 500 myosin molecules which are
bundled together. The tails of these myosin molecules are bound together and
extend towards the M line in the center of the sarcomere. Whereas the
globular myosin heads of these myosin molecules are arranged in a spiral,
projecting towards the nearest thin filament and placed away from the M line.
(Figs 4.7 b and 4.8 e ). As a result, the central portion of a thick filament (in the
H zone) is smooth, but its ends are studded with a staggered array of myosin
heads.Thus the central region in the H zone appears lighter in colour as there
are no heads in that area (Figs 4.8 a & e.Also refer again to Figs 4.6 c, d).

Fig. 4.7: a) A single myosin molecule consists of two heavy chains and four light
chains and has two heads with the actin binding and ATP binding sites,
two neck and a tail containing the binding sites for the actin filaments;
b) The tail of the myosin molecule combines with the tails of other
myosin molecules to form the thick filament.

The heads of the myosin molecules interact with the thin filaments forming
cross bridges during the contraction mechanism as we will study in greater
detail in the next section. In the center of each thick filament and extending
from it on either side of the M line to the Z line, are present the thin strands of
titin proteins (Fig. 4.8 b & f). The exposed part of these titin protein in the I
band are highly elastic and coiled (Fig. 4.8 b & f). In the resting sarcomere this
protein remains relaxed and becomes tense only when the sarcomere
stretches.

The thin actin filaments are 5-6 nm thick and 1µm long. A single thin actin
filament is actually made up of 4 different proteins namely: i) Filamentous (F)
actin; ii) nebulin, and the two regulatory proteins; iii) tropomyosin; and iv)
troponin (Fig 4.8c). 105
Block 1 Fundamentals of Animal Physiology
F actin is a linear polymer made up of two rows of 300-400 molecules of a
globular protein G- actin which are twisted around each other in a helix
(Fig. 4.8 c). Each G-actin molecule contains an active site, where myosin
heads of the thick filaments can bind. Nebulin protein extends in the cleft
between the two strands of G actin molecules. The nebulin molecule holds the
two strands of G-actin molecules together and the length of the F actin strand
is determined by the length of the nebulin molecule. The two strands of G-
actin molecules along with the tubulin molecule collectively make up the F-
actin which is often simply referred to as actin.

When the muscle is at rest, strands of tropomyosin cover the active sites and
prevent the actin –myosin interaction. Each tropomyosin molecule is a double
stranded protein and covers 7 active sites and is bound to one molecule of
troponin in the middle of its length.

Fig. 4.8: a) Structure and arrangement of thick and thin myofilaments in a

relaxed sarcomere of the muscle cell fiber; b) diagram of a relaxed
sarcomere showing enlarged diagram of a part of the sarcomere in
which, position of the titin and actinin proteins which hold the actin and
myosin filaments in postion can be seen; c) structure of the two
regulatory proteins tropomyosin and troponin of the actin filament; d)
enlarged diagram of a part of the tropomyosin and troponin protein
showing the troponin molecule complex consisting of three subunits
namely, troponin I, troponin C, and troponin T, referred to ICT in short;
e) diagram of myosin filaments being held in place by the M Line,
showing the tails forming the shaft of the thick filaments and the
myosin heads projecting outwards toward the surrounding thin
filaments; f) An enlarged single myosin molecule showing the
106 movement of its head by its hinge-like neck.
Unit 4 Muscle Physiology
A troponin molecule is a complex of three subunits (ICT) (Fig. 4.8 d) which are
globular regulatory proteins and are referred to as troponin I, troponin C, and
troponin T. Troponin I subunit of the troponin molecule binds to the
tropomyosin forming the tropomyosin- troponin complex: Troponin I subunit
binds to one G actin molecule for keeping the tropomyosin - troponin complex
in position and troponin C subunit has a receptor that binds to a calcium ion
(Ca2+).Since the Ca ion concentration in the resting sarcomere is very low, this
site remains empty before contraction starts.

SAQ 2
a) Match the alphabet designating zones in the myofibril given in Column A
with their description given in Column B :

Column A Column B

i) H- zone a. central line of the sarcomere where

myosin filaments are anchored

ii) A- band b. the area where only actin filaments are

present

iii) M- line c. a line that separates one sarcomere

from another

iv) Z- disc d. the area where only myosin filaments

are present

v) I- band e. includes overlapping myosin and actin

filament

b) Fill in the blanks and compare your answers with those given at the end
of this unit:

i) The entire muscle is covered by an …………………… .

ii) Bundles of muscle fibers (cells) are sheathed by a ……………...

and each muscle fiber is surrounded by an ……………… .

iii) Myofibrils contain myofilaments called ……..…………. and

……………. filaments.

iv) Thin myofilaments consist of ………..…………, ………………,

……………….. and …………………… .

4.3.4 Sliding Filament Mechanism

Now that you know the molecular structure of individual myofibrils let us see
how they act to contract the muscle cells. Muscle contraction is a physical
activity that generates force and causes the shortening of the muscle. In order
for a muscle cell to contract, the sarcomere must shorten. When a sarcomere
contracts the distance between the Z discs is reduced. In addition, some
regions of the sarcomere shorten whereas others stay the same length. 107
Block 1 Fundamentals of Animal Physiology
Appearance of Bands in a Contracted Sarcomere

In this process of contraction of the sarcomere (See Fig. 4.9) the bands of the
sarcomere appear as follows:

 The A band that contains thick filaments of myosin, remains

constant in width while the width of other regions of the sarcomere
shorten.

 The width of both H zone that contains only myosin filaments and the I
band that contains only filaments of actin, decrease in length.

 Furthermore, the zone of overlap, in which thin filaments and thick

filaments occupy the same area, increases.

 The contraction weakens with the disappearance of the I bands, at

which point the Z lines come in contact with the end of the thick
filaments. The sarcomeres in the entire length of the myofibrils come
closer to one another thus, shortening the myofibril. All these changes
during the contraction of sarcomere can only be explained if we
consider that the thin filaments are sliding past the thick filaments in
order to move towards the center of the sarcomere.

Fig. 4.9: Steps in contraction of skeletal muscles showing the position of the
bands of the sarcomere in a) relaxed state; and b) in contracted state.
During contraction the Z lines move closer together, while the H zone
and I zone decrease in width. The width of A band remains the same.
This happens because the thin filaments move toward the M line of
108 each sarcomere and so the thin filaments slide past the thick filaments.
Unit 4 Muscle Physiology
These observations were first made in 1954 by two independent teams, Hugh
Huxley and Jean Hanson and Andrew. F. Huxley and Rolf Niedergerke who
proposed the theory of the ‘sliding filament model’ which has now been
experimentally confirmed. Thus, the theory of the sliding filament model is
no longer a theory, instead now it is a scientific become fact.

The next step in the study of muscle contraction by scientists was to find out
the molecular mechanism of how muscle contraction takes place. In the
forthcoming sections we will study this molecular mechanism of muscle
contraction as a result of the actin molecules sliding past the myosin
molecules. However, before we do so we will first examine in brief the blood
and the nerve supply of the skeletal muscles which are essential for these
muscle contraction. We will also study in subsection 4.4.7 the role of calcium
and regulatory proteins and various other molecular factors that cause skeletal
muscle contraction to take place

4.3.5 Nerve and Blood Supply

Skeletal muscles are well supplied with blood vessels and nerves. Generally, a
skeletal muscle fiber cell is supplied by a nerve, an artery and by one or two
veins.

Fig 4.10: Blood supply to muscle the muscle tissue, showing the capillaries
surrounding the muscle fiber cells.

Each muscle fiber cell is in close contact with one or more capillaries.
(Fig.4.10).The blood capillaries bring in oxygen and nutrients and remove heat
and the waste products of muscle metabolism. Especially during contraction, a
muscle fiber synthesizes and uses considerable ATP (adenosine
triphosphate). You will learn more about these reactions later, in the unit.
These reactions require oxygen, glucose, fatty acids, and other substances
that are delivered to the muscle fiber via blood. 109
Block 1 Fundamentals of Animal Physiology
4.4.6 Neural Control of Muscle Contraction
A normal skeletal muscle fiber contracts only under the control of the
nervous system. Each muscle fiber receives midway along its length, an
electrical signal by the axon of a motor neuron (Fig 4.11 a) originating in the
spinal cord or in the brain stem. The point where the muscle fiber receives the
axon terminal (Figs 4.11 a) is known as the neuromuscular junction (NMJ).
The axon of the neuron, branches at this junction to form a fine network. Each
neuron branch expands at its terminal end to form the synaptic terminal knob
which makes contact with the surface of the sarcolemma (Figs 4.11 a & b).The
junction between the synaptic terminal knob of the axon and the muscle fiber
with which it communicates is called the synapse. The part of the synapse
that is on the side of the axon is called the presynaptic terminal and the
part which is on the side of the skeletal muscle cell is called
the postsynaptic terminal. Between these terminals a gap, called the
synaptic cleft is present (Figs 4.11 a, b & c).

Fig 4.11: a) A diagrammatic representation of a muscle fiber with its

neuromuscular junction; b) enlarged view of a neuromuscular junction;
110 and c) detailed view of a terminal synaptic cleft and motor end plate.
Unit 4 Muscle Physiology
The neuromuscular junction is a microstructure through which the process of
contraction is initiated or halted in the muscles by action of the neurons.At the
junction of the presynaptic terminal of the axon and the sarcolemma
(post synaptic terminal), a neurotransmitter called acetylcholineis released
by the presynaptic terminal. Acetylcholine changes the permeability of the
muscle membrane. Both neurons and skeletal muscle cells are electrically
excitable, as they are able to generate action potentials across their
membranes. An action potential is a special type of electrical signal that can
travel along a cell membrane as a wave. This allows a signal to be transmitted
quickly and faithfully over long distances. You will learn in detail about this
property of the neuron in Unit 6 of Block 2 of this course.

Each pre-synaptic knob present on the terminal end of each branch of the
axon of the neuron contains many mitochondria. It also has several secretory
vesicles filled with neurotransmitter acetylcholine (ACh).Presynaptic terminal
also contains enzymes for the degradation of excess or extra
neurotransmitters (Fig 4.11 b). The synaptic cleft allows the neurotransmitters
to diffuse and reach the postsynaptic terminal present as mentioned earlier on
other side of the neuromuscular junction or the synapse.

The postsynaptic membrane (sarcolemma) of the skeletal muscle fiber lies

directly under the nerve terminal and is characterized by extensive
invaginations known as postjunctional folds which provide additional
surface area for ACh receptors. These postjunctional folds are often called
the motor end plate. The acetylcholine neurotransmitter released from the
pre-synaptic knob diffuses across the space of the synaptic cleft and binds to
its receptor in the sarcolemma (Fig.4.11 c) and changes the permeability of
the muscle membrane. This process takes place in the following steps:

1. An action potential or electrical impulse (which is a sudden change in the

membrane potential that travels along the length of the axon) arrives at
the presynaptic axon terminal of the synaptic junction. The action
potential changes the permeability of the presynaptic axon terminal
triggering the exocytosis of acetylcholine into the synaptic cleft.

2. The acetylcholine molecules diffuse across the synaptic cleft and bind to
their ACh receptor present on the surface of the sarcolemma on the
motor end plate on the postsynaptic terminal. The binding of an
acetylcholine molecule to its receptor opens a membrane channel in
the receptor for entry of positive ions Since the extracellular fluid
contains a high concentration of sodium ions and the sodium ion
concentration inside the muscle cell is very low, therefore, sodium ions
rush in, into the cytosol of the muscle

3. The sudden inrush of sodium ions results in the generation of an action

potential in the sarcolemma which begins to depolarize (becoming less
negative). The action potential thus generated travels along the surface
of sarcolemma and rapidly enters the T tubules to initiate the
“excitation-contraction coupling”.

4. The action potential on reaching the cisternae of the sarcoplasmic

reticulum causes the release of calcium ion (Ca2+). Within a millisecond
of the release of Ca2+ the level of Ca2+ in the sarcoplasma increases to 111
Block 1 Fundamentals of Animal Physiology
100 times the resting concentration. As the terminal cisternae are
situated at the zone of overlap of the thick and thin filaments, the effect
is instantaneous.

5. Even before the action potential has traveled through the sarcolemma
surface the acetylcholine molecules are degraded (broken down) by
enzyme acetylcholinesterase. Some of these molecules are reabsorbed
by the synaptic membrane to be reused and the sequence of steps is
ready to be repeated again.

The propagation of action potential along the sarcolemma as

described above is the excitation part of the excitation- contraction -
coupling.

4.3.7 Role of Calcium and Regulatory Proteins

Recall from the earlier subsection 4.4.3 that troponin has an active site for
binding calcium and a site that binds to tropomyosin. In a muscle at resting
state the tropomyosin prevents the interaction between the thick and thin
filaments by blocking the cross bridge binding sites of F-actin protein (actin
filament) (Fig. 4.12). The troponin which acts as a regulatory protein has a
high affinity for calcium ion. Thus calcium acts as a key to unlocking the sites
of F-actin protein. When Ca2+ ion binds to the active binding site of the
troponin protein it brings about conformational changes in both troponin and
tropomysin molecules. This effect induces the tropomyosin to move away from
the active sites present on the F-actin (actin filaments). This allows the myosin
heads to attach to the actin filament due to which the thin filaments slide past
the thick filament towards the center of the sarcomeres.

2+
Fig.4.12: Action of Ca in unlocking the active sites on actin filaments. In the
relaxed muscle the tropomyosin lies on the outside blocking the
myosin cross-bridge formation sites on the actin. In the contracting
muscle the tropomyosin is pulled away thus exposing the myosin
binding sites of the actin filaments.

4.3.8 Sequence of Molecular Muscle Movement Cross

Bridge Cycle
As you will recall from the previous subsection 4.3.7 the myosin heads attach
to the actin filaments due to which the actin filaments slide past the thick
filaments. However, each head that attaches to actin can pull it a very short
distance before it reaches its limit. So it needs to detach and ‘re-cock’ or
recharge itself to attach at the next site. This repeated attaching-detaching
cycle is known as cross bridge cycle and needs energy which is
112 supplied by ATP.
Unit 4 Muscle Physiology
In the resting sarcomere the myosin molecules are already charged with
energy required for binding to the actin molecules. This energy is obtained
from the breakdown of ATP. At the start of the contraction cycle the myosin
head splits a molecule of ATP into ADP and a molecule of phosphate (Pi) all of
which remain attached to the myosin head (Fig 4.13 a).

When the muscle is stimulated to contract, calcium ions enter the sarcoplasm
and bind to the troponin molecules. The tropomyosin-troponin complex moves
to expose the binding sites from the F actin molecule.

Myosin heads bind to the active sites on the actin molecules (F-actin
molecule), forming cross bridges (Fig. 4.13 b). In the resting sarcomere the
myosin heads face away from the M line like a spring and are packed with
energy obtained by the breakdown of ATP. In this position the ADP and Pi as
you are aware are still attached to the head. When the myosin heads forms
the cross bridges this locked energy is released as the head pulls the F- actin
molecules towards the M line. This action is known as the power stroke. After
the power stroke the ADP and Pi are released (Fig. 4.13 c).

Another ATP molecule now binds to the myosin head and the link between the
myosin head and the actin site is broken and this active site becomes free to
bind with another myosin head (Fig. 4.13 d).The myosin head that became
free now gets recharged by splitting another bound ATP molecule into ADP
and Pi and becomes ready to attach to a new site on the actin filament (Fig.
4.13 e).

Fig. 4.13: Steps of Skeletal Muscle Contraction: A) The active site (a) on actin is
exposed as calcium binds to troponin. The myosin head is attracted to
actin, and myosin binds actin at its actin-binding site, forming the
cross-bridge (b); B) During the power stroke(c), the phosphate
generated in the previous contraction cycle is released; C) This
results in the myosin head pivoting toward the center of the
sarcomere, after which the attached ADP and phosphate group are
released; D) A new molecule of ATP attaches to the myosin head,
causing the cross-bridge to detach. The myosin head hydrolyzes ATP
to ADP and phosphate, which returns the myosin to the cocked
position. 113
Block 1 Fundamentals of Animal Physiology
If calcium is sufficient and the supply of ATP is constant, then myosin heads
will repeat the cross-bridge cycle about 5 times per second. Each power
stroke shortens the sarcomere by about 1 percent which means that per
second the sarcomere can shorten by about 5 percent. Since all the
sarcomeres shorten at the same time, the entire muscle shortens at the same
rate.

The time taken by a skeletal muscle fiber to sustain a contraction depends on:
1) the duration of nervous stimulation; 2) the amount of Ca2+ released and
The greater the
3) the supply of ATP. Furthermore, the contraction can be sustained only if
muscle tone, the there are repeated action potentials arriving at the neuromuscular junction. If
higher the resting rate only one action potential arrives at the neuromuscular junction then the
of metabolism. In calcium ions (Ca2+) return to the resting concentration by moving back into the
weight loss programs extracellular fluid through active transport. In addition, the sarcoplasm
the aim is to reduce reticulum also pumps back the Ca ions using Ca2+ATPase. As the
your energy intake as concentration of the calcium ions fall the Ca ions detach from the troponin
well as increase the molecule and the troponin-tropomyosin complex is reformed so that the active
resting rate of
sites on the actin molecule are again covered by tropomyosin. Thus, as a
metabolism so that
more calories are
result the skeletal muscle contraction ends. When the contraction ends some
burnt even while you of the energy spent in stretching the elastic elements of the muscle is
are not exercising. recovered as the stretching elements recoil. This helps the muscle to relax
gradually and come to the resting state.

4.3.9 Muscle Tone

All muscle fibers controlled by a single motor neuron form a motor unit. The
size of the motor unit is indicative of the fineness of the motor control. For
example, in the eyelid which requires fine muscle control, the motor unit
controls 4-6 muscle fibers; in leg muscles a motor unit may control 1000-2000
muscle fibers as precise muscle control is not required. In a stimulated
muscle, the contraction starts with the activation of the smallest motor unit and
gradually the larger motor units are stimulated and tension rises steeply. This
smooth and steady rise in muscular tension by the increasing number of
activated motor units is known as recruitment or multiple motor unit
summation. During a sustained contraction the motor units are activated on a
rotating basis, so that some of them are resting while some are active. This
gives each motor unit some time to recover before it is stimulated again.
When muscle cells
However in any skeletal muscle some motor units are always active even if the
are damaged creatine
entire muscle is not contracting. Thus, at any given time some of the motor
phosphokinase leaks
out of the cells into units are stimulated/activated to produce tension to maintain the body posture.
the blood stream. This activation may not cause movement but gives the muscle a firmness or
Thus any increase in resting tension. This resting tension is also known as muscle tone. The
creatine position of bones and joints in the body is maintained because of the resting
phosphokinase in the muscle tone. Muscles with little tone appear limp and loose while those with
blood is indicative of moderate tone appear firm and solid while skeletal muscles with strong tone
serious muscle makes these skeletal muscles appear well defined as we see in the muscle
damage.
builders, and weight lifters.

4.4 ENERGETICS OF MUSCLE CONTARCTION

You know now that ATP is the immediate source of energy for muscle
contraction. A single muscle fiber may contain 15 billion thick filaments and
114 when that fiber is actively contracting then it uses upto 2500 ATP molecules
Unit 4 Muscle Physiology
per second. Even a small skeletal muscle tissue contains thousands of muscle
fibers and therefore imagine the number of ATP molecules required even if
the muscle was to contract for a short duration.

Skeletal muscle fibers unlike most cells of the body, often switch between a
low level of activity when they are relaxed and using only a modest amount of
ATP, and a high level of activity, when they are contracting and using ATP at a
rapid pace. A huge amount of ATP is needed to power the contraction cycle,
to pump Ca2+ into the sarcoplasmic reticulum, and for other metabolic
reactions involved in muscle contraction. However, the ATP present inside the
muscle fibers is only enough to power contraction for just a few seconds. If
muscle contractions continue past that time, the muscle fibers must make
more ATP. Skeletal Muscle fibers have three ways to produce ATP: 1) from
creatine phosphate, 2) by anaerobic glycolysis, and 3) by aerobic respiration
(Figure 4.14). The use of creatine phosphate for ATP production is unique to
muscle fibers, but all body cells can make ATP by the reactions of anaerobic
glycolysis and aerobic respiration. In this subsection we will consider how
generation of ATP occurs for sketelal muscle cell contraction in vertebrates.

1. Through Creatine phosphate: At rest a skeletal muscle produces more

ATP than it can use so where is this ATP stored in the muscles? In a
resting muscle fiber ATP transfers energy into creatine to form another
high energy compound molecule creatine phosphate in vertebrates and
into arginine to form arginine phosphate in invertebrates.

When the skeletal muscles in vertebrates starts to contract it needs

energy. During contraction of the skeletal muscles the ATP as you will
recall is broken down into ADP +P in order to liberate the energy
required. The recharge of the ADP produced then comes from creatine
phosphate which rephosphorylates ADP in a reversible reaction:

ATP + creatine → ADP + creatine phosphate → ATP + creatine (see

Fig 4.14 a)

The enzyme that is required for this transformation is creatine

phosphokinase. The pool of creatine phosphate in the muscle cell is
about 10 times larger than the ATP present. However as mentioned
earlier the, creatine phosphate-derived ATP powers only the first few
seconds of muscle contraction (Fig. 4.14 a) after which another energy
source has to be used.

2. Through glycogen: As the source of ATP from creatine phosphate gets

depleted, and the muscles continue to contract they turn to glycolysis as
an ATP source. They obtain their ATP from the breakdown of pyruvic
acid in the glycolysis pathway. Glycolysis is an anaerobic (non-oxygen-
dependent) process that breaks down glucose to produce ATP.
However, glycolysis cannot generate ATP as quickly as creatine
phosphate. The glucose used in glycolysis can be provided by blood
glucose or by metabolizing glycogen that is stored in the muscle. The
breakdown of one glucose molecule produces two ATP and two
molecules of pyruvic acid (Fig. 4.14 b). The two molecules of pyruvic
acid can be used in aerobic respiration or when oxygen levels are low 115
Block 1 Fundamentals of Animal Physiology
may be converted to lactic acid which may contribute to muscle fatigue.
Glycolysis itself cannot be sustained for very long (approximately 1
minute of muscle activity), but it is useful in facilitating short bursts of
high-intensity output. This is because glycolysis does not utilize glucose
very efficiently, producing a net gain of two ATPs per molecule of
glucose, and the end product of lactic acid.

Fig. 4.14: Biochemical pathways producing ATP utilized during vertebrate

skeletal muscle contraction : (a) Some ATP is stored in a resting
muscle. As contraction starts, it is used up in seconds. More ATP is
generated from creatine phosphate for about 15 seconds. Creatine
phosphate, which is formed from ATP while the muscle is relaxed,
transfers a high-energy phosphate group to ADP, forming ATP during
muscle contraction. In this process thus, oxygen is not needed for ATP
generation and 1 ATP is generated per creatine b) Breakdown of
muscle glycogen produces glucose. Each glucose molecule via
glycolysis produces two ATP and two molecules of pyruvic acid. In
glycolysis no oxygen is needed, and so this is an anaerobic pathway.
Energy provision by glycolysis lasts from 40 seconds to 2 minutes.
Pyruvic acid produced as a result of glycolysis can be used in aerobic
respiration or converted to lactic acid if oxygen is not available. When
oxygen is unavailable pyruvic acid is converted to lactic acid, which
may contribute to muscle fatigue. This occurs during strenuous
exercise when high amounts of energy are needed but oxygen cannot
be sufficiently delivered to muscle. (c) Aerobic respiration is the
breakdown of pyruvic acid obtained from glycolysis. This occurs within
the mitochondria, where pyruvic acid, fatty acids, and amino acids are
used via aerobic respiration to produce ATP, carbon dioxide, and water.
Approximately 95 percent of the ATP required for resting or moderately
active muscles is provided by aerobic respiration, which takes place in
mitochondria. In this process oxygen is needed for ATP generation and
30/32 ATPs are produced per glucose. Energy provision by aerobic
116 respiration of pyruvic acid lasts for several hours.
Unit 4 Muscle Physiology
3. Through aerobic respiration: Normally 95% ATP demand of the
contracting muscle fiber is met by aerobic metabolism that takes place in
the abundant mitochondria present in the muscle cells. For aerobic
respiration glucose, pyruvic acid and fatty acids are obtained from blood
circulation. Aerobic respiration is much more efficient than anaerobic
glycolysis. Aerobic respiration generates a net gain of approximately 32
ATPs per molecule of glucose versus a net gain of two generated from
glycolysis (Fig. 4.14 c).

Let us now see with the help of Fig 4.14 again how the production and usage
of energy for contraction changes as muscular activity increases:

• Recall that the demand for energy is low in a resting muscle and enough
oxygen is available for aerobic metabolism of glucose. The cell absorbs
glucose and fatty acids from the blood and is able to make enough ATP
for its use and stores the surplus in the form of creatine phosphate and
excess glucose is stored as glycogen.

• At moderate levels of muscle activity ATP demand increases. As more

mitochondrial ATP is formed oxygen consumption also increases which
is easily met by more oxygen diffusing into the cell. However, as the
contraction continues, no surplus ATP is available and the muscle now
relies on the breakdown of glycogen and other substrates such as lipids
and amino acids.

• When muscular activity is at its peak the energy demand also reaches its
maximum. The rate of ATP production is now limited by availability of
oxygen and so now only a third of the ATP is obtained from aerobic
mitochondrial activity and the rest is obtained from anaerobic conversion
of pyruvic acid (pyruvate) to lactate(lactic acid) to glucose.

• As more lactic acid is formed than is consumed, it accumulates in the

muscle fiber and changes its pH. The changed pH ultimately alters the
activity of the key enzymes in the muscle fiber. As a result the muscle
fiber cannot continue to contract further. Once the muscle slows down it
gets time for oxygen levels to be restored and for the lactates to revert to
pyruvate. This allows the aerobic metabolism to resume and supply
energy to the muscle cell. The build up of lactic acid in the muscle cells
gives the body the sensation felt after vigorous exercise. The soreness
felt after 24 hours of exercise is actually due to muscle damage and the
inflammatory repair response which leads to swelling and soreness that
lasts for a few days.

SAQ 3
Choose the correct options:

a) In a relaxed muscle fiber, the myosin-binding site on actin is blocked by

……………… .

i) titin ii) troponin

iii) myoglobin iv) tropomyosin 117

Block 1 Fundamentals of Animal Physiology
b) According to the sliding filament model, binding sites on actin open when
………………. .

i) creatine phosphate levels rise ii) ATP levels rise

iii) acetylcholine levels rise iv) calcium ion levels rise

c) The plasma membrane of a muscle fiber is called …………………. .

i) myofibril ii) sarcolemma

iii) sarcoplasm iv) myofilament

d) Muscle relaxation occurs when …………………. .

i) calcium ions are actively transported out of the sarcoplasmic

reticulum

ii) calcium ions diffuse out of the sarcoplasmic reticulum

iii) calcium ions are actively transported into the sarcoplasmic

reticulum

iv) calcium ions diffuse into the sarcoplasmic reticulum

e) During muscle contraction, the cross-bridge detaches when …………… .

i) the myosin head binds to an ADP molecule

ii) the myosin head binds to an ATP molecule

iii) calcium ions bind to troponin

iv) calcium ions bind to actin

4.5 ADAPTAIONS OF MUSCLES TO PERFORM

DIFFERENT ACTIVITIES IN VERTEBRATES
The principles that determine the mechanical properties of muscles are
illustrated in three very different kinds of motor activity: jumping of frogs,
swimming of fish, and sound production by toadfish and rattlesnakes. We will
consider each of these activities and the muscles that are used to produce
them.

Adaptation for Power: Jumping Frogs

When they jump, frogs move rapidly (in 50 to 100 milliseconds) from a
crouched position, in which their potential and kinetic energy is zero, to an
extended position, in which their potential and kinetic energy is high.
Mechanical work must be done if the potential and kinetic energy of a body is
to increase, and because that work must be performed in such a short time,
the muscles that produce the jump must generate high power (i.e., work per
unit time). In fact, the distance a frog jumps depends directly on how much
power its muscles produce. Thus we might expect that a frog's jumping
118 muscles would have properties that allow them to generate high power.
Unit 4 Muscle Physiology
A muscle generating high power exhibits three properties:

1) it operates within the plateau of the sarcomere length-tension curve

where maximal force is generated

2) it shortens at a rate at which maximal power is generated;and

3) it becomes maximally activated before shortening begins.

Diversity of Function: Swimming Fish

The study of muscles in fish has, for two reasons, been particularly useful in
helping to elucidate how muscular systems are organized. First, fish display a
wide diversity of movements that can be elicited readily and analyzed
quantitatively. Second, different kinds of movements are powered by different
muscle fibre types, which in fish are anatomically separated.

During the many movements of which fish are capable, the change in
sarcomere length is roughly proportional to the curvature of the spine.

The muscles of a fish is able to generate both slow, low-amplitude movements

and fast, high- amplitude movements. This is because fish has different
muscles, containing different fiber types. When a fish is swimming steadily,
only red muscles are active; these are composed of slow oxidative (type I)
fibers. In contrast, white muscles, composed of fast glycolytic (type IIa) fibers,
are recruited to produce fast swimming or very rapid movements such as the
escape response. A fish can produce very different kinds of movements well,
because for each movement it uses muscles that are specialized to match the
demands of the particular task.

Adaptation for Speed: Sound Production

Some animals produce sound through mechanisms-for example, the

movement of a column of air past a vibrating membrane or past the vocal
cords-that are not directly coupled to muscle contraction. In other animals,
however; sound is produced when muscles generate forces that directly cause
structures, such as the swimbladder of a toad-fish or the rattle on the tail of a
rattlesnake, to vibrate. In these animals, the sound-producing (or sonic)
muscles must undergo contraction-relaxation cycles at the frequency at which
the sound is made, which is 10 to 100 times faster than most locomotory
muscles operate.

Sonic muscles have unique properties that allow them to operate at the high
frequencies which are discussed in the following section.

Toadfish swim-bladder

The male toadfish, Opsanus tau, produces a "boat whistle" mating call ten to
twelve times per minute for many hours to attract females to its nest. This tone
is generated by oscillatory contractions of the muscles encircling the fish's
gas-filled swim-bladder.

Experiments with toadfish indicate that sonic fibers have a number of

adaptations that permit them to operate at very high frequencies.
Ultrastructural and biochemical studies, for example, suggest that the short 119
Block 1 Fundamentals of Animal Physiology
Ca2+ transient is achieved by an unusually high density of calcium channels in
the SR membrane through which Ca2+is released; an unusually high density
of calcium pumps through which Ca2+ is re-sequestered; an increased
concentration of certain calcium-binding proteins (e.g., troponin); and a fiber
morphology such that the distance between the SR membrane and the
myofilaments is particularly short, reducing the time required for diffusion. The
rapid release of Ca2+from troponin likely reflects decreased affinity of troponin
for Ca2+. Finally, the rapid detachment of cross-bridges implies that myosin in
sonic fibres also has special molecular properties.

Rattlesnakes

Rattlesnakes in the genus Crotalus also use special noise-making muscles,

but as a warning to members of other species rather than to attract
conspecifics for mating. Rattling is a loud and effective warning that renders
these snakes, like many venomous animals, very conspicuous. Unlike the
periodic toadfish boat whistle, rattling can be sustained continuously for up to
3 hours. However, the rapidity with which the rattle-producing muscles
contract suggests that these muscles might have many features in common
with the toadfish swim bladder sonic muscles.

The properties of rattlesnake shaker muscle suggest that a rapid calcium

transient alone is not sufficient for the production of very rapid contractions.
The release of Ca2+ from troponin and the detachment of cross-bridges from
actin filaments must be unusually fast as well.

Energetic and spatial constraints on muscle operation at high

frequencies

The rapid calcium kinetics that are required in muscle fibers that contract and
relax rapidly could potentially demand a relative increase in the surface area
(and volume) of the sarcoplasmic reticulum and in the number of mitochondria
within each fiber. Any such increase must reduce the space that is available
within each fiber for myofilaments, the structures that generate force. In
toadfish sonic fibers, for example, the rate of Ca2+uptake by the sarcoplasmic
reticulum is about 50-fold greater than it is in red fibers, and about 30% of the
entire volume of a sonic fiber is occupied by the sarcoplasmic reticulum. In
addition, if a fast muscle is to operate continuously, as the rattlesnake shaker
muscle does, it must use aerobic metabolism to generate enough ATP to fuel
a high rate of calcium pumping; thus each fiber must contain many
mitochondria, displacing even more myofilaments.

High-Power, High-Frequency Muscles: Asynchronous Flight

Muscles
Most species in the Hymenoptera (bees and wasps), Diptera (flies),Coleoptera
(beetles),and Hemiptera (true bugs) contain flight muscles that are a notable
exception to the rule that no more than one contraction is evoked by a single
depolarization of the surface membrane. This unusual type of striated skeletal
muscle does not exhibit a one- to-one relation between the arrival of motor
impulses and the timing of individual contractions. These flight muscles are
called fibrillar muscles, or more commonly asynchronous muscles, to
120 distinguish them from muscles that contract in synchrony with each action
Unit 4 Muscle Physiology
potential from a motor neuron. Although the timing of contraction is unrelated
to the timing of neuronal input to an asynchronous muscle, a constant train of
motor impulses and muscle depolarizations is required to maintain an
asynchronous muscle in an active condition.

In some species of small insects, the wingbeat frequency (and the frequency
of wing-muscle contractions) far exceeds the maximal maintained discharge
rates of which axons are capable. Wingbeat frequency has been found to vary
inversely with wing size. A tiny midge, for example, beats its wings at a
frequency in excess of 1000 Hz, producing a high-pitched sound that can be
perceived by the human ear.

The mechanics of flight differ considerably in insects possessing synchronous

flight muscles from those possessing asynchronous muscles. In those insects
with synchronous flight muscles (e.g., damselfly),the wings are elevated and
depressed by simple lever mechanics. Insects with asynchronous flight
muscles have a more complex musculoskeletal arrangement in which the
thorax can exist in either of two stable configurations. In these insects,
contractions of the antagonistically arranged flight muscles change the shape
of the thorax to generate only two wing positions-up or down.

SAQ 4
State ‘True’ or ‘False’:

a) The mechanical and energetic properties of different muscle fiber types

exhibit remarkable diversity and adaptation to various activities.

b) Electromyograms are the recordings of muscle response or electrical

activity in response to a nerve's stimulation of the muscle.

c) Fibrillar muscles are also known as synchronous muscles.

d) In asynchronous flight muscles at one point of time some motor units are
active others are inactive.

e) Vertebrate sonic muscles and some insect sonic muscles can operate at
frequencies well above 100Hz.

4.6 SUMMARY
In this unit you have read about the physiology of movement in animals. You
have read that:

• The primary function of the muscle tissue is to produce movement.

There are three types of muscle cells in the vertebrate body namely,
skeletal, cardiac, and smooth. Their structures match their specific
functions in the body. Skeletal muscle and cardiac muscles are striated.
The striated skeletal muscle is multinucleated, voluntary and responds to
conscious stimuli. Cardiac muscle is branched, uni-nucleated and is
found only in the heart and contracts autonomously and involuntarily.
Smooth muscle is involuntary. Each smooth muscle cell is spindle-
shaped. Striations are absent in smooth muscle cells. 121
Block 1 Fundamentals of Animal Physiology
• In skeletal muscle the sarcomere is the smallest contractile portion of a
muscle. Myofibrils are the contractile units of the muscle and are
composed of thick and thin filaments. Thick filaments are composed of
the protein myosin; thin filaments the actin are composed of the proteins
Filamentous (F) actin, nebulin, and two regulatory proteins iii)
tropomyosin and iv) troponin.

• Contraction in skeletal muscles is brought about by the sliding of the thin

actin filaments past the thick myosin filaments. A motor neuron controls
the activity of contraction of a skeletal muscle fiber at a neuromuscular
junction present on the skeletal muscle. When an action potential arrives
at the neuron’s axon (pre-synaptic) terminal, acetylcholine (ACh) is
released into the synaptic cleft. The binding of ACh to its receptors on
the post synaptic motor end plate on the sarcolemma leads to the
generation of an action potential which travels along the sarcolemma to
trigger calcium release from sarcoplasmic reticulum (SR). The actin sites
are exposed after Ca+ enters the sarcoplasm from its SR storage. This is
because Ca ions activate the troponin-tropomyosin complex so that the
tropomyosin shifts away from the actin binding sites. The cross-bridging
of myosin head which dock into actin-binding sites occurs and is
followed by the “power stroke” - the sliding of the thin actin filaments
between the thick myosin filaments. The power strokes are powered by
ATP. Ultimately, the sarcomeres, myofibrils, and muscle fibers shorten to
produce movement.

• Contraction of skeletal muscles requires large amounts of energy.

Creatine phosphate can release stored energy to convert ADP to ATP.
At rest or at moderate levels of activity, aerobic metabolism can provide
most of the ATP needed to support muscle contractions.At peak levels of
activity, the muscle cell relies heavily on anaerobic metabolism
(glycolysis) to generate ATP.

• The mechanical and energetic properties of different muscle fiber types

exhibit remarkable diversity and adaptation to various activities.
Muscular systems have evolved so that muscles operate at the optimal
myofilament overlap to generate maximal force, and they shorten at the
appropriate velocity (V/Vmax) to generate maximal mechanical power with
near-optimal efficiency.

• The kinetics of activation and relaxation tend to be relatively slow in

locomotory muscles, minimizing the high energetic cost of calcium
pumping by the sarcoplasmic reticulum.

• In contrast, vertebrate sound-producing muscles must operate at very

high frequencies, and thus have very rapid calcium pumping. Although
sonic muscles are accordingly very expensive energetically, they only
make up a small fraction of any animal's muscle mass.

• Finally, insect asynchronous flight muscles are able to generate high

mechanical power at very high frequency while avoiding the high
energetic cost of calcium pumping, because they use stretch activation
and shortening deactivation to achieve high-frequency modulation of
122 force.
Unit 4 Muscle Physiology

4.7 TERMINAL QUESTIONS

1. What is the role of nexin and the radial spokes in the structure and
movement of cilia and flagella? What will happen if we were to remove
the nexin links and the radial spokes by means of enzymes protease?

2. Explain briefly the role of myosin in muscle contraction.

3. Describe the role of calcium in the regulating muscle contraction.

4. Diagrammatically explain the biochemical pathways that produce ATP

for vertebrate muscle contraction.

5. How is sound produced by muscle contraction in toad and rattlesnake?

6. What modifications you see in flight muscle physiology and function as

compared to regular skeletal muscle?

4.8 ANSWERS
Self-Assessment Questions
1. a) The three types of muscle tissue are skeletal muscle, cardiac
muscle, and smooth muscle. Skeletal muscle tissues move the
body by pulling on our bones. Cardiac muscle tissues pump blood
which flows throughout the cardiovascular system. Smooth muscle
tissues push fluids and solids along the digestive tract and other
internal organs, and regulate the diameters of small arteries and
sphincters.

b) i) skeletal muscles, cardiac muscles

ii) Skeletal muscles, voluntary, smooth, cardiac,

involuntary

2. a) i) the area where only myosin filaments are present

ii) includes overlapping myosin and actin filaments

ii) central line of the sarcomere where myosin filaments are

anchored

iv) a line that separates one sarcomere from another

v) the area where only actin filaments are present

b) i) epimysium

ii) perimysium, endomysium;

iii) thin, thick;

iv) F-actin, nebulin, tropomyosin, troponin.

3. a) Tropomysin

b) calcium ions rise 123

Block 1 Fundamentals of Animal Physiology
c) sarcolemma

d) calcium ions are actively transported into the sarcoplasmic reticulum

e) myosin head binds to an ATP molecule.

4. a) True

b) True

c) False

d) True

e) True

Terminal Questions
1. Radial spokes and nexin are scaffolding proteins in the axoneme of
flagellum/cilium. The outer microtubule doublets are connected to the
central pair of microtubules by radial spokes that extend from the A
tubule of the doublet to the inner sheath covering the two central tubule.
The radial spokes thus keep the outer doublets linked to the sheath of
the central tubules. Nexin proteins cross- link each microtubule doublet
to its adjacent microtubule doublet. The nexin links are highly elastic. If
these two scaffolding proteins are removed by proteases then the
structural integrity of the axoneme will be destroyed.The nexin cross-
links would be broken, allowing the sliding of adjacent microtubules of
the doublets unimpeded and so will not lead to bending formation of the
cilium/flagellum.

2. A myosin molecule contains two heavy chains; part of a heavy chain

makes up one head and the other part forms the tail. The ‘heads of the
myosin molecules form crossbridges with the actin molecules. These
cross bridges cyclically attach to actin molecules and swivel, acting as
oars that pull the actin and myosin filaments past each other resulting in
sliding movement.

3. The thin filaments of the myofibril consist of F-actin simply called actin
and two regulatory proteins, troponin and tropomyosin. In a resting state
of the muscle, tropomyosin blocks the cross bridges binding sites of the
actin molecules. When a muscle is stimulated the calcium ion
concentration of the muscle fiber increases. The calcium ions bind to the
troponin molecules. Binding of calcium ions to the troponin causes
conformational changes in tropomyosin which moves away, thereby
uncovering the cross bridges binding sites of the actin molecules,
causing muscle contraction.

4. Refer to Subsection 4.3.8 and Fig. 4.14.

5. Refer to Section 4.5.

6. Refer to Section 4.5.

124
 UNIT 5
FEEDING SYSTEMS AND
DIGESTION

Structure
5.1 Introduction 5.4 Mechanism of Absorption in
Objectives Vertebrate Animals
5.2 Feeding Patterns Absorption of
Monosaccharides
Food Absorption through
Exterior Body Surfaces Absorption of Amino Acids,
Dipeptides, and Tripeptides
Endocytosis
Absorption of Lipids
Filter Feeding
5.5 Physiology of Digestion in
Fluid Feeding
Human Mammals
Cutting and Licking
Digestion in Oral Cavity
Seizing of Prey
Digestion in Stomach
Toxin
Digestion and Absorption in
Herbivory and Grazing to
Small Intestine
Collect Food
Absorption in Large Intestine
5.3 Mechanism of Digestion
5.6 Energy Metabolism in
and its Regulation
Animals
Action of Proteolytic Enzymes
5.7 Summary
Action of Carbohydrase
5.8 Terminal Questions
Enzymes
5.9 Answers
Action of Lipase Enzymes
Maintenance of Gut Lining
Coordination of Digestion in
Animals

5.1 INTRODUCTION
All organisms require a fairly steady supply of nutrient materials from the
environment to obtain energy in order to stay alive. You have already read that
in organisms cellular metabolism provides energy for various processes, like
movement, locomotion, excretion, osmoregulation, synthesis of new materials
for growth and maintenance and reproduction. To provide energy for these
Block 1 Fundamentals of Animal Physiology
processes raw material or nutrients are required which are supplied by food.
In addition, animals require certain amino acids, vitamins and minerals which
their body is unable to synthesize. The study of nutrition involves both the
need for food to provide energy and the need for specific food components for
various other activities of the body.

The process by which animals acquire and ingest their food is referred to as
feeding. Diverse types of feeding mechanisms have been evolved by different
groups of animals. There exists a relationship between the nature of ingested
food and type of feeding mechanisms used in acquiring of food. Virtually all
food ingested by animals whether of plant or of animal origin must be broken
down into simple compounds by the process of digestion. Digestion and
absorption of food constitute the essential link between nutrition and
metabolism. In this unit we shall first discuss briefly the different feeding
pattern observed in animals and the specialized feeding mechanisms. Then
we will consider the digestion and absorption of nutrients that takes place
within the body of the animals. Towards the end of the unit, we shall discuss
energy metabolism in animals.

Objectives
After studying this unit you should be able to:

 describe the various feeding patterns seen in animals,

 list the various feeding strategies evolved by the animals in relation to

the available food,

 distinguish between the process of digestion of proteins, carbohydrates

and fats,

 explain the role of gastrointestinal hormones in digestion,

 summarise the process of absorption of food from the alimentary canal,

 describe the digestion process in humans as an example of

mammalian digestion, and

 explain energy metabolism in animals in relation to oxygen

consumption.

5.2 FEEDING PATTERNS

Obtaining food adequate in both quantity and quality occupies much of the
routine behavior of most animals. Certainly an animal's physiology and
morphology are the result of natural selection that favours effective acquisition
of energy from food while avoiding becoming someone else's meal. In nature
you will see the variety of methods by which animals feed. Sessile (non-
mobile) bottom-dwelling species commonly resort to surface absorption, filter
feeding, or trapping. Mobile animals follow a more active sequence, which in
the extreme of many carnivores (meat eaters) includes searching, stalking,
126 pouncing, capturing, and killing.
Unit 5 Feeding System and Digestion
5.2.1 Food Absorption through Exterior Body Surfaces
The feeding method that is least dependent on specialized capture and
digestive organs involves absorption of nutrients directly across the body wall.
Certain protozoans, endoparasites (animals that live within other animals), and
aquatic invertebrates are able to take up nutrient molecules from the
surrounding medium directly through their soft body wall. Endoparasites such
as parasitic protozoans, tapeworms, flukes, and certain molluscs and
crustaceans are surrounded by host tissues or by alimentary canal fluids, both
of which are high in nutrients. Tapeworms, which may be many meters long,
lack even a rudimentary digestive system. Tapeworms evolved from a
primitive flatworm that lacked a body cavity (i.e., was acoelomic). However,
some endoparasites appear to have secondarily lost the digestive apparatus
present in their ancestors. For example, parasitic crustaceans, which belong to
the cirripeds (barnacle group),lack an alimentary canal, but they appear to
have evolved from nonparasitic ancestors possessing a gut.

Some free-living protozoans and invertebrates derive part of their nutrients by

direct surface uptake from the surrounding medium. Small molecules such as
amino acids are taken up from dilute solution by transport mechanisms against
what can often be a huge concentration gradient. In some of these organisms,
larger molecules or particles are taken up by a bulk process such as
phagocytosis, which is described next.

5.2.2 Endocytosis
Endocytosis represents a more active form of "feeding" than passive
absorption directly across the body wall. Like direct nutrient absorption,
however, it occurs at the local cellular rather than tissue or organismal level.
Endocytosis includes two processes, phagocytosis ("cell eating") and
pinocytosis ("cell drinking"). In phagocytosis, pseudopod- like protuberances
extend out and envelope relatively large nutrient particles. Pinocytosis occurs
when a smaller particle binds to the cell surface and the plasma membrane
invaginates (folds inward) under it, forming an endocytic cavity. Whether
captured by phagocytosis or pinocytosis, the morsel is then engulfed in a
membrane-enclosed vesicle that pinches off from the bottom of the cavity.
Feeding by pinocytosis and phagocytosis is familiar in protozoans such as
Paramecium, but also occurs in the lining of the alimentary canals and other
tissues of many multicellular animals. The detailed process of phagocytosis
are described further in this unit.

5.2.3 Filter Feeding

Many aquatic animals use filter feeding, also called suspension feeding, to
capture food. Food items (usuallyphytoplankton or zooplankton) are carried to
specialized entrapment devices either on the body surface or within it.

Most marine filter feeders are small, sessile animals, such as sponges,
brachiopods, lamellibranchs, and tunicates. Food items are carried along on
water currents that either occur naturally or are generated by the movements
of body parts of the filter-feeding animal itself, such as cilia or flagella.
Brachiopods respond behaviourally to currents, rotating on their pedal stalks to 127
Block 1 Fundamentals of Animal Physiology
present the most efficient hydrodynamic orientation for capturing the water
current. A number of other sessile animals located in moving water make use
of Bernoulli's effect (i.e., a drop in fluid pressure as fluid velocity increases)to
increase the rate of water flowing through the entrapment sites, at no energy
cost to themselves. An example of such passively assisted filter feeding is
seen in sponges (Fig. 5.1).The flow of water across the large terminal opening
causes a drop in pressure (Bernoulli's effect) outside the osculum. As a result,
water is drawn out of the sponge through the osculum, and is drawn in through
the numerous ostia (mouth-like openings) in the body wall. The drop in
pressure is facilitated by the shape of the sponge's exterior, which causes the
water over the osculum to flow with greater velocity than the water flowing past
the ostia. Food particles, swept into the ostia of the sponge along with the
water, are engulfed by choanocytes, the flagellated cells lining the body cavity.
The flagella of the choanocytes also create internal water currents within
spongocoel, the hollow water-filled interior. Some sponges living in moving
water "pump" a volume of water equivalent to up to 20,000 times their body
volume per day.

Mucus, a sticky mixture of mucopolysaccharides, often plays an important role

in filter feeding. Waterborne microorganisms and food particles are trapped in
a layer of mucus that covers a ciliated epithelium. The mucus is then
transported to the oral parts by beating cilia. The cilia propel water through
sessile animals not only to capture suspended food but also to aid in
respiration. This is of greatest importance in still water. In molluscs such as the
mussel, Mytilus, the cilia on the surface of the ctenidium draw a stream of
water through the inhalant siphon, passing the water between the gill filaments
(Fig. 5.2). These cilia are also responsible for keeping mucus traveling down
along the filaments (i.e., 90" to the water flow) to the tip of the gill, where it
travels in a special groove under ciliary power toward the mouth in a rope-like
string of mucus. Sand and other indigestible particles are sorted out and
rejected (presumably on the basis of texture), passing out with the water
leaving the exhalant siphon.

128 Fig: 5.1 Water flow in syconoid sponges.

Unit 5 Feeding System and Digestion

Fig. 5.2: Lamellibranch molluscs employ ciliary feeding.

Non-sessile animals filter-feed by various mechanisms. A number of fishes are

planktivorous, using modified gill rakers to strain food out of the flow of water
passing through the mouth and over the gills. Juveniles of the paddlefish,
Polyodon spathula, swim rapidly and continuously both to ventilate their gills
and to filter out food items. Filter feeding is also very common in amphibian
larvae. In Xenopus laevis, the South African clawed toad, the branchial
chamber contains gills bearing branchial filter plates that entrap suspended
organic material. The material becomes entrapped in mucus, which is then
swept by cilia into the oesophagus to be swallowed. In Xenopus, branchial
respiration and food ingestion may present functional conflicts. As the gill filter
plates load up with suspended food items, the resistance to water flow through
the gills rises sharply. Indeed, in larval Xenopus, gill ventilation decreases in
proportion to food density in inspired water, presumably maintaining a
constant rate of food ingestion. Increased cutaneous and pulmonary
respiration apparently can compensate for the lack of branchial gas exchange
when optimal conditions for filter feeding result in reduced branchial water
flow.

The largest filter feeders are the baleen whales, such as the right whale.
Horny baleen plates bear a fringe of parallel filaments of hair-like keratin that
hang down from the upper and lower jaws and act as strainers analogous to
the gill rakers of fishes or larval amphibians (Fig. 5.3 A). These whales swim
with jaws open into schools of pelagic crustaceans such as krill, engulfing vast
numbers suspended in tons of water. As the jaws close, the water is squeezed
back out through the baleen strainers with the help of the large tongue, and
the crustaceans, left behind inside the mouth, are swallowed. Clearly, filter
feeding can be a very effective form of food capture and can support an
animal of huge dimensions.

Birds such as flamingos also use filter feeding to capture small animals and
other morsels they find in the muddy bottoms of their freshwater habitat (Fig.
5.3 B). The flamingo and the right whale exhibit remarkable convergent
evolution: they both have a deep-sided lower jaw, a re- curved rostrum,
fibrous-fringe filters suspended from the upper jaw, and a large, fleshy tongue. 129
Block 1 Fundamentals of Animal Physiology
Both feed by filling the mouth cavity with water, and then using the tongue as a
piston to force water out through the filters, trapping and retaining waterborne
food particles.

Fig. 5.3: Filter feeding mechanism in A) whale and B) flamingo. Please refer to
text for description.

5.2.4 Fluid Feeding

Fluid feeding involves a variety of structures and mechanisms, including
piercing and sucking, and cutting and licking.

Piercing and Sucking

Feeding by piercing a prey or food item and sucking fluids from it occurs
among the platyhelminths, nematodes, annelids, and arthropods. Leeches,
among the annelids, are true bloodsuckers, using an anticoagulant in their
saliva to prevent clotting in their prey's blood.

Large numbers of arthropods feed by piercing and sucking. Most familiar and
irksome of these to humans are mosquitoes, fleas, bedbugs, and lice, which
can be vectors of disease. The majority of sucking arthropods victimize animal
hosts. However, especially among the Hemiptera (true bugs) are species that
pierce and suck plants, from which they draw sap. Sucking insects generally
possess fine piercing mouthparts in the form of a proboscis (Fig. 5.4 A).Often,
130 the two maxillae are shaped so that they make up two canals that run to the tip
Unit 5 Feeding System and Digestion
of the proboscis (Fig. 5.4 B and C).One of these, the dorsal canal, is the
passage for blood or sap sucked from the host. The other, the ventral canal,
carries saliva, containing anticoagulants or enzymes, from the salivary glands
into the host. Sucking occurs by the action of a muscular pharynx. After
feeding, most insects are able to fold the proboscis back out of the way.

Fig. 5.4: Mouth parts of mosquito A) lateral view, B) cross-section and C) lateral
view of butterfly mouth parts.

5.2.5 Cutting and Licking

Numerous invertebrates and a few vertebrates feed by cutting the body wall of
a prey item and then licking, or sponging the body fluids that leak from the cut.
The blackfly and related biting flies have mouthparts with a sharpened
mandible for cutting, and a large, sponge-like labium for transferring the body
fluid (usually blood) to the esophagus. Among the chordates, a few phyletically
ancient fishes (lampreys, hagfishes) use rasp-like mouths to make large,
circular flesh wounds on the host. They feed on the blood created from these
wounds. Vampire bats use their teeth to make puncture wounds in cattle, from
which they lick oozing blood. The saliva of these bats contains an
anticoagulant, as well as an analgesic to prevent the host from feeling the
effects of the bite, at least until the bat has finished feeding. 131
Block 1 Fundamentals of Animal Physiology
5.2.6 Seizing of Prey
Predators use various types of mouthparts and other appendages to capture
and masticate animals and plants. Often toxins are used to further immobilize
items of prey.

Jaws, teeth, and beaks

Although no true teeth occur among the invertebrates, various invertebrates

have beak-like or tooth-like chitinous structures for biting or feeding.
Invertebrates such as the praying mantis and the lobster also have anterior
limbs modified for prey capture (Fig. 5.5). Spiders and their relatives have
needle-like mouthparts for injection of venom, while cephalopods like the
octopus has a sharp, tearing beak. Among the vertebrates, hagfishes, sharks,
bony fishes, amphibians, and reptiles have pointed teeth, mounted on the jaws
or palate, that aid in holding, tearing, and swallowing prey.

Fig. 5.5: Mouthparts modified for holding and tearing the prey in A) mantispid
insect B) lobster.

The teeth of non-mammalian vertebrates are usually non-differentiated, with a

single tooth type found through-out the mouth. One notable exception is found
among the poisonous snakes, such as vipers, cobras, and rattlesnakes, which
have modified teeth, called fangs, that they use to inject venom (Fig. 5.6).
These fangs either are equipped with a groove that guides the venom or are
132 hollow, very much like a syringe. In rattlesnakes, the fangs fold back against
Unit 5 Feeding System and Digestion
the roof of the mouth, but extend perpendicularly when the mouth is opened to
strike at prey. A snake's jaws are held together with an elastic ligament that
allows them to spread apart during swallowing. This enables the snake to
swallow animals larger than the diameter of its head (see Fig. 5.6).Swallowing
whole prey is relatively common, and very evident in prey capture and
consumption in snakes.

Fig. 5.6: Modified teeth of Rattlesnake, known as fangs A) striking position, B)

non-striking position.

Mammals use their teeth for seizing and masticating their prey. Their teeth
have developed very different shapes during evolution (Figure 5.7).Chisel-like
incisors are used for gnawing, especially by rodents and rabbits. In the
elephants (and before them, mammoths), the incisors are modified into a pair
of tusks. Pointed, dagger-like canines are used by the carnivores,
insectivores, and primates for piercing and tearing food. In some groups like
wild pigs and walruses, the canines are elongated as tusks, which are used for
preying and fighting. Most complex and interesting in their form are the molars
of some herbivorous groups such as cattle, including oxen, pigs,
hippopotamuses, and horses and zebras. These teeth, which are used in a
side-to- side grinding motion, are composed of folded layers of enamel,
cement, and dentine, all of which differ in hardness and in rate of wearing.
Because the softer dentine wears rather quickly, the harder enamel and
cement layers form ridges that enhance the effectiveness of the molars for
chewing grass and other tough vegetation. Many mammals, such as the cats
(the domestic cat and the great cats, such as the lion) use limbs equipped with
sharp claws to supplement the teeth as food-capturing structures. 133
Block 1 Fundamentals of Animal Physiology

Fig. 5.7: Variously modified dentition in placental mammals A) generalised

dentition B) Squirrel dentition with modified incisors C) Lion dentition
with modified canines D) Ox dentition with modified molars

Instead of teeth, birds have horny beaks, in a multitude of shapes and sizes,
evolved to adapt to each species' unique food sources and methods of
obtaining them. For instance, beaks may have finely serrated edges, sharp,
hook-like up- per bills, or sharp, wood-pecking points (Figure 5.8). Seed-eating
birds eat their food whole (perhaps after removing the outer hull), but may
grind the swallowed seed in a muscular crop or gizzard containing pebbles
that act like "millstones." Raptorial birds (hawks,eagles),endowed with
excellent vision and flight mobility, capture prey with their talons as well as
their beaks.

134 Fig. 5.8: Birds beaks adapted for herbivory, omnivory and carnivory.
Unit 5 Feeding System and Digestion
5.2.7 Toxins
A large number of animals from different phyla use toxins either to subdue
prey or to fend off predators. Most of these toxins act at synapses in the
nervous system. Surprisingly simple animals can use sophisticated arrays of
venom- producing cells. Among the coelenterates (hydras, jellyfish,
anemones, corals) for example, there is extensive use of nematocysts
(stinging cells). Concentrated in large numbers on the tentacles, the
nematocysts inject paralytic toxins into prey and immobilize it while the
tentacles carry it to the mouth. Many nemertine worms paralyze their prey by
injecting venom through a stiletto-like proboscis. Venoms are also used by
annelids, gastropod molluscs (includingone species of octopus), and a wide
variety of arthropods.

Among the last group, scorpions and spiders are most notorious for their
toxins, which are usually highly specific chemicals that bind to specific
receptor types. After grabbing its prey with its large chelae (pincer-like
organs), a scorpion will arch its tail and then plunge its sting into its prey
(Figure 5.9).The scorpion then injects the victim with a poison containing a
neurotoxin that interferes with the proper firing of nerve impulses. Spider
poisons also contain neurotoxins. The venom from the black widow spider
contains a substance that induces massive release of neurotransmitter at the
motor endplate in muscle. A neurotoxin, a-bungarotoxin, found in the venom of
the cobra-like krait, binds to nicotinic acetyl-choline (ACh) receptors, thereby
blocking neuromuscular transmission in vertebrates. The venoms of various
species of rattlesnake contain haemolytic (blood cell-destroying) substances.

Fig. 5.9: The Scorpion captures prey and then injects poison to subdue it. 135
Block 1 Fundamentals of Animal Physiology
Toxins, although highly effective, are generally expensive to produce. Usually
carefully measured doses of toxins are delivered during a bite or sting. Toxins
also must be specially stored before administration to avoid self- poisoning.
Toxins are generally proteins and, as such, are rendered harmless by the
proteolytic enzymes of the predator's digestive system when it ingests its
poisoned prey.

5.2.8 Herbivory and Grazing to Collect Food

Herbivores often have mouth parts specialized for feeding on plant material.
Microtines belong to
Many gastropods use a rasp-like structure termed as radula to scrape algae
the subfamily
Microtinae, that from rock surfaces or to rasp through vegetation (Fig. 5.10).Vertebrate
includes, voles, herbivores have bony plates (some fish and reptiles) or teeth primarily in the
lemmings and form of molars with wide flat surfaces modified for grinding plant material.
muskrats. Plants (especially some grasses) contain relatively large amounts of silicates,
and can be tremendously abrasive. Consequently, the molars of herbivores
often are coated in especially tough enamel to resist wear. Alternatively, some
herbivores such as small rodents (microtines) have continuously growing,
rootless teeth.

Fig. 5.10: Gastropod head saggital section displaying A) radula location with
respect to mouth; B) Protraction and C) retraction of radula while
136 grazing.
Unit 5 Feeding System and Digestion
By now you know the different feeding strategies adopted by various
invertebrate groups and by different vertebrates. Next we have listed the major
feeding methods adopted by various animal groups, based on the type of food
available to them (Table 5.1). You can note that taxonomically different animal
groups living in the same habitat obtain food in a similar manner. For example,
many marine animals (annelids, molluscs, crustaceans, herring fish, petrel
birds, flamingo birds, baleen whales etc) may be filter feeders though the
organs involved with the process of filtration may not be anatomically similar.
Table 5.1: Feeding methods classified according to type of food.
Sl. Type of Food Method of feeding Animals using the method
No.
I. Small Particles Digestive vacuoles Amoeba, Radiolarians
Use of cilia Sponges, Bivalves, Tadpoles
and Ciliates including
Paramecium
Mucous traps Gastropods, Tunicates
Tentacles Sea cucumbers
Filter feeding Annelids, Molluscs, small
Crustacean, Herring fishes,
Flamingoes, Petrels birds,
Baleen Whales
II. Large food Ingestion of inactive Detritus feeders, Earthworm
masses masses

Scraping, chewing, Sea urchins, Snails, Insects,

boring Vertebrates
Capture and swallowing Coelenterates, Fishes,
of prey Snakes, Birds, Bats,
Chewing and swallowing Mammals
III. Fluid or soft Sucking plant sap, nectar Aphids, Bees, Humming-
tissue birds,
Ingestion of blood Leeches, Ticks, Lice, Insects
Vampire bats
Sucking milk or similar Young Mammals and Birds
secretions

External digestion Spiders

Nutrient molecules Parasites, Tapeworm
uptake through the
integument of the host or
from the host medium in
which they live.
IV. Dissolved organic Uptake from dilute Aquatic invertebrates
solution solution
V. Symbiotic supply Intracellular symbiotic Sponges, Corals, Hydras,
of nutrients algae Flatworms, Clams
137
Block 1 Fundamentals of Animal Physiology

SAQ 1
Name one animal in each category which follows the following feeding
methods.

a) Food absorption by external body surface

b) Endocytosis
c) Filter feeding
d) Fluid feeding
e) Prey capture using toxins

5.3 MECHANISM OF DIGESTION AND ITS

REGULATION
The process by which the food is broken down into simpler molecules with the
aid of enzymes is known as digestion Figure 5.11 summarizes the process of
digestion and the absorption of ingested food which is applicable to all
animals. As you can see in the figure, ingested food which is in the form of
complex macromolecules like proteins, carbohydrates and lipids and nucleic
acids is digested or broken down by means of enzymes into simpler food
molecules, amino acids, simple sugars and glycerols, fatty acids and
nucleotides respectively, so that they can be easily absorbed by the body.
Important minerals, vitamins, and water present in the ingested foods are also
extracted and absorbed from the ingested food by the body during the process
of digestion. We will study the mechanism of digestion and its regulation in this
section.

Fig. 5.11: Digestion is the breakdown of complex, ingested food into simpler
food molecules with the help of digestive enzymes for absorption by
the body. The digested simpler food molecules help in growth, tissue
138 building and generating energy for various functions of the body.
Unit 5 Feeding System and Digestion
The process of digestion is mediated through various digestive enzymes.
Digestive enzymes as you know chemically breakdown complex, large,
ingested food molecules by hydrolysis into simpler, smaller, soluble units.
The enzymes to digest food rupture the chemical bonds of the food to be
digested by addition of one or more than one of their water molecules to the
ingested food, so that the food is broken down into simpler molecules. The
properties and nature of digestive enzymes show specific differences from
enzymes in general in the following ways:

1. Digestive enzymes are not as narrowly specific as other enzymes and

instead show group specificity. For example, enzymes that digest
carbohydrates can digest polysaccharides of both animal and plant origin.

2. Even though enzymes performing similar functions in different animals are

given the same names, they are not however, identical chemically. For
example, trypsin (an enzyme, that hydrolyses proteins) in humans is not
identical to that found in fish. Temperature and pH for optimum activity
also differs. For example, trypsin from vertebrate pancreas acts best in
the pH range of 7-9 while in silkworm the ideal pH range is 6.2 to 9.

3. Digestive enzymes secreted by pancreas particularly those that digest

proteins are secreted in an inactive form.

The four major classes of digestive enzymes in animals are as follows:

i) Proteases that hydrolyse peptide bonds in proteins,

ii) Carbohydrases that hydrolyse glycosidic bonds in carbohydrate,

iii) Lipases that hydrolyse ester bonds in fats

iv) Nucleases that hydrolyse DNA and RNA into nucleotides

In this unit we will study the role of only three classes of digestive enzymes
namely proteases, carbohydrases and lipases involved in digestion of food.

5.3.1 Action of Proteolytic Enzymes

Protein digestion begins in the stomach and is completed in the small
intestine. It is brought about by proteolytic enzymes known as proteases that
are synthesized as larger, inactive forms (zymogens) which after secretion,
are cleaved to produce active proteases. Enzymes that digest proteins are
categorized, based on the site of their action in the protein molecules into two
groups : i) endopeptidases and ii) exopeptidases :

i) Endopeptidases confine their attack to the interior part of the protein

molecule so that the large peptide chains are broken into smaller
fragments. These smaller fragments provide many more sites for
exopeptidases to act upon as endopeptidases cannot breakdown
proteins into their monomers (simpler forms).

ii) Exopeptidases digest the smaller protein fragments produced by the

action of the endopeptidases. They attack only peptide bonds present at
the end of a peptide chain and cause the release of amino acids,
dipeptides and tripeptides. 139
Block 1 Fundamentals of Animal Physiology
There are several types of endopeptidases and exopeptidases of which the
main ones are listed in Table 5.2.

As you can see, in the table, the exopeptidases and endopeptidases, attack
specific peptide bonds depending on the chemical group near them. The
inactive forms of the proteolytic enzymes need activators and autocatalysts for
converting them into active forms. For example pepsinogen, secreted by chief
cells of the vertebrate stomach is only activated when hydrochloric acid (HCl)
which is secreted by the parietal cells of the stomach makes the medium of
the stomach acidic (pH 2).The acidic environment of the stomach activates the
inactive pepsinogen into the active pepsin by altering the conformation of
pepsinogen so that it can cleave itself to yield active pepsin.

Pepsin acts as endopeptidase and cleaves dietary proteins, composed of

large polypeptide chains into a number of short fragments of polypeptides.
Pepsin can hydrolyze virtually any peptide bond except for those present
in keratin, mucoproteins, and protamines (protamine sulfate is a small Salmon
positively charged protein derived from Salmon sperm which is widely used to
neutralize the anticoagulant activity of heparin following cardiac procedures).
Pepsin has a broad spectrum of specificity though it prefers to cleave
specifically the peptide bond on the amino group of an aromatic amino acid
(phenylalanine, tryptophan and tyrosine) (Table. 5.2 and Fig.5.14 a).In this
way short fragments of polypeptide chains are formed. Invertebrates seem to
lack pepsin and their main endopeptidases is more like trypsin. In Table 5.2
you can further note that endopeptidase chymotrypsin also attacks a peptide
bond involving an aromatic amino acid but it attacks on the carboxyl terminal
end of the molecule while pepsin attacks on the amino terminal end.

Table 5.2: Proteases of animals. Some enzymes are secreted as

zymogens that need other enzymes to convert them into their
active forms while some proteases are secreted in their active
forms but require the presence of ions to be active.

Extracellular Enzyme

Preferred Peptide Link

Activator for Attack
Inactive form Active form (Origin)
autocatylst
I. ENDOPEPTIDASES

HCL Link to amino group of

pepsin aromatic amino acid
Pepsinogen Pepsin (tyrosine, tryptophan,
(Stomach) and phenylalanine)

Trypsinogen Enterokinase Link to carboxyl end

Trypsin
Trypsin of arginine or lysine
(Pancreas)

Chymotrypsinogen Trypsin Chymotrypsin Link to carboxyl group

of aromatic amino
(Pancreas)
acid (phenylalanine,
140 tyrosine, tryptophan)
Unit 5 Feeding System and Digestion
and also bonds
adjacent to
methionine and
leucine when they are
present.

Proelastase Trypsin Elastase Link to carboxyl group

of alanine, glycine,
(Pancreas)
and serine

II. EXOPEPTIDASES

Aminopeptidases (Mn2+, Mg2+, Zn2+) Link to terminal

amino acid with free
(Lining of intestine)
amino group

Trypsin Link to terminal

amino acid with free
Procarboxypeptidases Carboxypeptidases
(Zn2+) carboxyl group

Dipeptidases (Mn2+, Mg2+, Zn2+) Bonds between pairs

of amino acids
(Lining of intestine)

The broken down polypeptide fragments are further digested in the small
intestine by the endopeptidases like trypsin (active form of inactive
trypsinogen) and several other inactive digestive proenzymes (zymogens) that
are secreted by the pancreas into the duodenum of the small intestine and
which, when activated, collectively digest the peptides into single amino acids
under alkaline conditions needed for digestion.

Fig. 5.12: Protein digesting enzymes: a) attacking specific peptide bonds in a

protein fragment; and b) breakdown of a tripeptide. However, both
exopeptidases cannot attack the same tripeptide.

Trypsinogen secreted by the pancreas into the duodenum of the small

intestine is activated into trypsin by another enteropeptidase enzyme (also 141
Block 1 Fundamentals of Animal Physiology
called enterokinase), secreted by the brush border cells of the small intestine.
As trypsin is formed, it autocatalytically activates more trypsinogen to be
converted into trypsin. Trypsin acts in an alkaline medium between pH 7-9.It
breaks a peptide bond next to a basic amino acid like arginine or lysine.

Trypsin cleaves the following proteolytic enzymes at the carboxyl side of

arginine and lysine: 1) inactive chymotrypsinogen to yield active
chymotrypsin. Chymotrypsin enzyme hydrolyses proteins that enter the
duodenum, into smaller peptides and amino acids. 2) inactive proelastase, to
yield active enzyme elastase. Elastase breaks down elastin (an elastic fibre
protein that, together with collagen, determines the mechanical properties
of connective tissue) and 3) inactive procarboxypeptidases, to yield active
carboxypeptidases, an exopeptidase. Enzyme carboxypeptidase splits one
amino acid at a time. Thus, trypsin plays a central role as the master activator
enzyme because it cleaves dietary proteins and activates other proteases that
cleave dietary protein. Apart from carboxypeptidase the other exopeptidases
are secreted in an active form but all need metal ions as cofactors to increase
their activity.

Each of these activated protein digesting enzyme exhibits cleavage specificity

as you can see in Table 1.6: i) chymotrypsin cleaves at the carboxyl side of
phenylalanine, tyrosine, tryptophan and leucine; ii) elastase cleaves at the
carboxyl side of alanine, glycine and serine. Carboxypeptidase in the presence
of zinc ion and trypsin cleaves single amino acids from the carboxyl terminus.

The polypeptide fragments produced due to endopeptidase proteolytic

enzymes are further digested as you are aware by the exopeptidases.

The action of the endopeptidase is shown in Fig. 5.12 (a &b) which illustrates
the action of i) aminopeptidase which removes terminal amino acids having
free amino groups and ii) carboxypeptidase which removes terminal amino
acids possessing a free carboxyl group. In this way aminopeptidase and
carboxypeptidase enzymes progressively remove peptides from each end
until a dipeptide fragment consisting of only two amino acids remains. Bonds
between these pairs of amino acids are split further by dipeptidases which
results in release of free amino acids, which are the simplest components of
protein that can be absorbed through the cells lining the wall of the small
intestine.

5.3.2 Action of Carbohydrase Enzymes

Enzymes that digest carbohydrates are called carbohydrases. Simple sugars
like glucose and fructose can be absorbed and metabolized directly by the
lining of the small intestine. However, disaccharides such as sucrose or
lactose and polysaccharides such as starch and glycogen have to be broken
down to monosaccharides before they can be absorbed and used in metabolic
pathways. Carbohydrases can be grouped into the following two categories:

i) Polysaccharases that split polysaccharides into disaccharides or

trisaccharides/oligosaccharides (short chains of three or more
monosaccharides).

ii) Glycosidases that break up the disaccharides or trisaccharides to

142 monosaccharides.
Unit 5 Feeding System and Digestion
The digestion of carbohydrates, similar to proteins occurs in a stepwise
manner. The polysaccharase enzyme first splits the polysaccharides into
disaccharides or oligosaccharides. After this, the disaccharides or
trisacchrides or oligosaccharides (the products resulting, due to the action of
the polysaccharase enzyme) are further split by the action of glycosidase (the
second enzyme) into monosaccharides. The enzymes responsible for
digestion of carbohydrates and the end products of carbohydrate digestion are
given in Table 5.3.

Table 5.3: Digestion of carbohydrates.

Polysaccharases
Polysaccha  Glycosida ses
   → Disaccharides      → Monosaccharide
rides
(C12H22O11) (CnH2nOn)
(C6H10O5)X

Glycogen Amylase Maltose Maltase Glucose

→   →
(animals)

Starch Amylase Maltose Maltase Glucose

→   →
(Plants)

Cellulose Cellulase Cellobiose Cellobiose s Glucose

  →    →
(Plants &
animals)

Trehalose Trehalase Glucose

  →
(insects and
some plants)

Lactose Lactase Galactose +

  →
Glucose

Sucrose Invertase Fructose +

 →
Glucose

Carbohydrate digestion in vertebrates and invertebrates is very similar. All the

enzymes shown in Table 5.3 are not required by all animals. However,
amylase and maltase are of universal occurrence. The presence of enzymes
in the digestive tract of an animal group is related to the food habits. For e.g.,
carbohydrate digestion in the human alimentary tract occurs in the following
two steps :i) step one, involves digestion of carbohydrates by enzyme amylase
of the salivary glands followed by digestion by means of alpha (α)-amylase
enzyme of pancreas which is secreted into the duodenum in larger amounts in
comparison to salivary amylase. The activity of amylase secreted by the
salivary glands and pancreas of humans works best at pH of 6.7- 7.5. ii) Step
two involves digestion of carbohydrates by the enzymes of the brush border
cells present in the epithelial lining of the duodenum.

Enzyme production in some animals is also influenced by genetic

characteristics and enzyme induction. For example, production of maltase and
sucrase by the intestinal villi depends on the amount of ingested sugar. If a
143
Block 1 Fundamentals of Animal Physiology
high maltose or sucrose diet is taken, it induces the villi to produce more
maltase and sucrose within 2-5 days. Lactase production declines in humans
Between 2-18 per
cent of Caucasians as gut develops after infancy. It ceases in some individuals so that lactose
lose the capacity to sugar can no longer be digested by them.
produce lactase and
between 95-100 per Now let us consider the digestion of cellulose, the most important structural
cent of Oriental and material of plants and a major component of the diet of herbivores. Very few
native African races animals possess the enzyme cellulase. Then how do animals that feed on
lose the ability to plants break down this carbohydrate? Cellulase enzymes are synthesized by
produce lactase as
several microorganisms such as bacteria and protists (protozoans) that live
they grow older. As a
result they can no symbiotically in the gut of many herbivores and insects. In herbivores such as
longer digest milk ruminants (i.e., cow, sheep, etc.) cellulose digestion is carried on with the help
which as a of these symbiotic microorganisms which live in the stomach. The
consequence breakdown products of cellulose are then utilized by the host. In some
ferments in their gut
invertebrates like the arthropod silver fish (Ctenolepisma lineate) true cellulase
and produces
diarrhoea and related
enzymes have been reported but the insect cannot survive exclusively on a
problems. cellulose diet. Some other invertebrates also have some cellulases that partly
Interestingly yoghurt digest cellulose. So far however, no conclusive evidence of a complete
and cheese do not breakdown of cellulose into glucose without the help of symbionts has been
create any problems reported.
as these contain less
than 2 per cent 5.3.3 Action of Lipase Enzymes
lactose due to action
of bacteria. Digestion of fats or lipids is similar in both invertebrates and vertebrates.
Lipases are the enzymes that hydrolyse fats. A single lipase can catalyse
many steps in the breakdown of fat. In vertebrates, lingual lipase secreted by
the tongue, gastric lipase (these two lipases are acid stable) in the stomach
Bile acids are steroid and pancreatic lipase (works in alkaline medium) secreted into the duodenum
acids found are responsible for digesting lipids. Lipids/triglycerides are hydrophobic, and
predominantly in
poorly soluble in the aqueous environment of the digestive tract.Thus, they
the bile of mammals
and other vertebrates.
have to be broken down into fat droplets by the action of lingual lipase and
Diverse bile acids are gastric lipase before pancreatic lipase can act on them in an alkaline medium.
synthesized in However the key players in the digestion of fats to yield monoglycerides and
the liver. Bile acids fatty acids are bile and pancreatic lipase as very little digestion of fat occurs
are conjugated with in the mouth and the stomach. Let us now study the digestion of
residues of amino
fats/triglycerides in detail.
acids taurine or
glycine to give anions
In vertebrates, the bulk of dietary lipid are neutral fats/lipids or triglycerides,
called bile salts.
Primary bile
composed of a glycerol backbone in which each carbon is linked to a fatty
acids are those acid. Foodstuffs typically also contain phospholipids, sterols like cholesterol is
synthesized by the and many minor lipids, and fat-soluble vitamins. Finally, small intestinal
liver while secondary contents contain lipids from sloughed epithelial cells and considerable
bile acids result from cholesterol is delivered in bile.
bacterial actions in
the colon. In order for the triglyceride to be digested and absorbed into the blood stream
the following two processes must occur:

i) Large aggregates of dietary triglyceride, which are hydrophobic and

virtually insoluble in an aqueous environment, must be broken down
physically into micelles which are held in suspension by a process called
144 emulsification.
 Unit 5 Feeding System and Digestion
ii) Triglyceride molecules must be enzymatically digested to yield
monoglyceride and fatty acids, both of which can efficiently diffuse or be
transported into the enterocyte (epithelial absorptive cells of the gut).

The key players in the digestion of fats to yield monoglycerides and fatty acids
as mentioned earlier are bile and pancreatic lipase, both of which are mixed
with chyme and act in the lumen of the small intestine. Bile acids from bile
juice are also necessary to solubilize other lipids, including cholesterol.

Emulsification, Hydrolysis and Micelle Formation

Bile juice simply referred to as bile plays a vital role in lipid assimilation by
promoting emulsification. Bile juice is produced in the liver, flows through the
biliary tract and gets stored in the gallbladder from where it is secreted into the
duodenum of the small intestine. It is a complex fluid containing water,
electrolytes and a large number of organic molecules including bile acids,
cholesterol, phospholipids and bilirubin. The bile juice is largely responsible for
making the environment in the duodenum alkaline which enables the
pancreatic digestive enzymes to function. Bile acids that form bile salts play
the first critical role in lipid assimilation by promoting emulsification (Fig. 5.15 a
and b). As derivatives of cholesterol, bile acids have both hydrophilic and
hydrophobic domains (i.e., they are amphipathic). On exposure to a large
aggregate of triglycerides, the hydrophobic portions of the bile acids get
intercalated (inserted) into the lipid, while the hydrophilic domains remain at
the surface which results in the formation of micelles. Such a coating formed
by the bile acids aids in the breakdown of large aggregates or droplets into
smaller and smaller droplets (Fig. 5.13 a and b).

Fig. 5.13: Digestion of fats. a) bile juice emulsifies the fat globules into smaller
droplets. b) pancreatic lipase cleaves the triglycerides and
diglycerides in the emulsified fat droplets into free fatty acids and 2-
monoglyceride.

Break down or hydrolysis of triglyceride is accomplished predominantly by

pancreatic lipase which is a water-soluble enzyme. Shortly after a meal,
pancreatic lipase is present within the small intestine in rather huge quantities,
but can act only on the surface of the triglyeride droplets. The pancreatic
lipase enzyme breaks down triglycerides into fatty acids, monoacylglycerides
(glycerol backbone with one fatty acid still attached), monogylecerides and
some free glycerol. Cholesterol and fat-soluble vitamins do not need to be
enzymatically digested. After this the next process requires that the products
of fat digestion (fatty acids, monoglycerides, monoacylglycerides glycerol,
cholesterol, and fat-soluble vitamins) enter into the circulation so that they can
be used by the body. 145
Block 1 Fundamentals of Animal Physiology
About 95 percent of lipids are absorbed in the small intestine. Bile salts not
only speed up lipid digestion, they are also essential for the absorption of the
end products of lipid digestion. Short-chain fatty acids are relatively water
soluble and can enter the absorptive cells (enterocytes) of the small intestine
directly. Despite being hydrophobic, the small size of short-chain fatty acids
enables them to be absorbed by enterocytes via simple diffusion, and then
take the same path as monosaccharides and amino acids into the blood
capillary of a villus. However, the large hydrophobic fatty chains (with more
than 10-12 carbon atoms), long chain fatty acids and monoglycerides that are
present in the duodenal chyme are too large in size due to which they have
difficulty being suspended in the watery environment of the intestinal chyme.
Bile salts formed from bile acids of the bile juice and lecithin present in the
chyme resolve this by enclosing these insoluble fat molecules into micelles. A
micelle is a tiny sphere with polar (hydrophilic) ends facing the watery
environment and hydrophobic tails turned to the interior (Fig. 5.14). The core
of the micelle also includes cholesterol and fat-soluble vitamins. Micelles
reduce surface tension at the fat-water interphase. Micelles are essentially
small aggregates (4-8 nm in diameter) of a mixture of lipids and bile acids,
suspended within the aqueous medium of the digested food in the intestine
(ingesta). As the ingesta is mixed, the micellefs bump into the brush border of
the epithelium of the small intestine , with the result that the lipids, including
monoglycerides monoacyl glycerol and fatty acids, are taken up into the
epithelial cells enterocytes.

Fig. 5.14: Scheme of a micelle formed by phospholipid in an aqueous solution.

SAQ 2
a) Choose the correct answer from the following:

i) Enzyme lipase digests lipids while trypsin digests proteins. Both

these digestive processes:

1. occur inside cells.

146 2. add a water molecule to break bonds

Unit 5 Feeding System and Digestion
3. require zinc to activate the enzymes

4. consume ATP.

ii) What role do bile acids and salts play in digestion of complex fats?

1. They catalyze the cleavage of bonds that release fatty acids

2. They emulsify the large fat droplets

3. They digest the complex fats.

4. They activate the lipases

iii) Carbohydrates are digested by:

1. Chymotrypsin in the duodenum

2. Lingual lipase in the oral cavity

3. Salivary and pancreatic amylase

4. HCl and pepsin in stomach

b) Which of the following organ is not correctly paired with its function?

i) Stomach - protein digestion

ii) Gall bladder - bile juice synthesis

iii) Small intestine - nutrient absorption

iv) Pancreas - enzyme production

c) Which of the following is not a major activity of the stomach?

i) Storage

ii) HCl production

iii) Nutrient absorption

iv) Churning of large food particles

5.3.4 Maintenance of Gut Lining

After studying the digestive enzymes you must be wondering as to why the gut
linings are not digested themselves. This is because animals have several
mechanisms that protect their gut lining from auto digestion that is self
digestion. Mucous cells of the vertebrate gut secrete a slightly alkaline mucous
that lubricates the food and protects the cells lining the gut from corrosive
secretions. In addition, the lateral surfaces of exposed epithelial cells of the
digestive tract are joined by tight junctions that prevent the digestive
secretions from penetrating between them. Careful studies have also revealed
that the entire lining of the gut is renewed every third day in rats and every 5-6 147
Block 1 Fundamentals of Animal Physiology
days in humans. Similar mechanisms are present in invertebrates also. In
insects, the fore-gut and hind gut are lined by cuticle. This lining is known as
intima. The epithelial cells are exposed only in the midgut, where most of the
digestion occurs. The midgut is lined by a delicate lining the peritrophic
membrane in some insects.

5.3.5 Coordination of Digestion in Animals

You have learnt that digestion is a process in which large food molecules are
broken down step by step into their constituents. In primitive metazoans that
are continuous feeders, the enzyme producing cells, secrete continuously. In
higher animals more precise controls are needed to regulate the release of
food from the stomach into intestine and also for the release of digestive
enzymes at the proper time. The interplay of nervous and hormonal control is
beautifully illustrated when we study the coordination of digestive activity.

In the mammalian mouth, control of salivary gland secretion is entirely

controlled by the nervous system. In the stomach the gastric secretions are
under hormonal and neural control while intestinal also called gastro intestinal
secretions are slower and are primarily under hormonal control.

Gastrointestinal secretion is largely under the control of gastrointestinal

hormones secreted by the endocrine glands present in the gastric and
intestinal mucosa of the gut.

Vertebrate Gastrointestinal Hormones: Example mammals

The three main mammalian gastrointestinal hormones are secretin, gastrin

and cholecystokinin (CCK). There are several other hormones, all peptides.
The physiology of only three major hormones is listed in Table 5.4. while Fig.
5.17 summarizes the action of these three GI hormones.

Gastrin is secreted by the stomach and is responsible for control of HCl

volume; the presence of HCl in turn inhibits further gastrin secretion.

Secretin regulates water homeostasis throughout the body and influences the
environment of the duodenum by regulating secretions in the stomach,
pancreas, and liver. It is a peptide hormone which is secreted in the
duodenum in response to acidic chyme and stimulates the pancreas and gall
bladder to release a large amount of water and bicarbonates causing the
environment of the duodenum to become alkaline and thus neutralize the
acidic chyme. Secretin is also of some historical interest, as it was the first
hormone to be discovered in 1902 by Sir William Maddock Bayliss and
Ernest Starling.

Cholecystokinin (CCK) hormone is produced by the cells lining the

duodenum. The arrival of partially digested fats and proteins from the stomach
into the duodenum initiates the release of CCK by the intestinal mucosa of the
duodenum.CCK induces the secretion and flow of bile and pancreatic juice
which are alkaline in nature into the duodenum. Bile juice aids in fat digestion
148 in the duodenum by emulsifying fats while pancreatic juice digest the fats.
Unit 5 Feeding System and Digestion
Table 5.18: Mammalian gastrointestinal hormones; − denotes inhibition;
+ stimulation; +++ hormone more important than other two

Features of Gastrin Secretin Cholecystokinin

gastrointestinal
hormones-Gastrin,
Secretin and
Cholecystokinin

Secreted by Stomach Duodenum Duodenum

Stimulus for release Peptides Acid (HCI) Amino acids,

fatty acids

Parasympathetic

Effect on : Nerves (Vagus

nerve)

Gastric Motility + _ _

Gastric HCl Secretion +++ _ _

Pancreatic secretion

bicarbonates + +++ +

enzymes + + +++
Sometimes, some
areas of the stomach
The two hormones Secretin and CCK inhibit stomach motility. Fig. 5.15 lining gets damaged
summaries the action of the GI hormones. and gastric ulcers
develop. Earlier it was
thought that ulcers
were caused by
psychological stress
which resulted in
excess acid
secretion. However,
Australian
researchers Barry
Marshall and Robin
Warren discovered
that ulcers are due to
infection by the acid-
tolerant bacterium
Helicobacter pylori.
They also proved that
an antibiotic could
cure most gastric
Fig. 5.15: Action of the three gastrointestinal hormones: Gastrin is secreted in ulcers. They were
response to intragastric proteins, stomach distention and stimulation awarded the Nobel
by the vagus nerve. Gastrin from the lower stomach stimulates HCl Prize in 2005 for
and pepsin secretion from the secretory cells. Cholecystokinin (CCK) these findings.
stimulates pancreas to secrete digestive enzymes and alkaline
secretions in order to neutralize and digest chyme. It also induces
contraction of gall bladder to secrete bile salts. CCK is secreted in
response to the arrival of amino acids and fatty acids in duodenum
from stomach. 149
Block 1 Fundamentals of Animal Physiology

SAQ 3
Fill in the blanks:

Three gastrointestinal hormones stimulate the release of digestive materials of

which, …………………. stimulates the release of gastric juices, ………………
stimulates the release of bicarbonate ions and ……………….. stimulates the
release of bile and pancreatic enzymes.

5.4 MECHANISM OF ABSORPTION IN

VERTEBRATE ANIMALS
In Section 5.3 you have read that once the food has been digested it is
absorbed in the small intestine. In animal physiology, absorption in higher
multicellular organisms is defined as, “the transfer of products of extracellular
digestion from the lumen of the gastrointestinal tract into the blood or lymph”.
In extracellular digestion, absorption occurs in sequence: digestion first,
absorption second. These organizing concepts, however, are inadequate for
the animal kingdom as a whole. When digestion is only intracellular, the same
cells are involved both in digestion and absorption. In those multicellular
animals where some of the digestion is extracellular and most is intracellular,
absorption in them is takes place first by the transfer of food particles from
the gut lumen into the get cells that digest the food particles intracellularly and
only later are the digestive products passed into the blood. In higher
multicellular animals however, in which digestion is totally extracellular there
are usually separate tissues and areas of the gut for enzyme production,
digestion and absorption. Thus a general definition of absorption, also called
assimilation, common to all animal groups would be, “the entry of
digested food molecules into tissues of the animal”.

Fig. 5.16: Epithelial lining of the mammalian small intestine : a) Villus covered with
digestive epithelium which consists of absorptive cell and occasional
goblet cells; b) An absorptive cell under higher magnification showing its
150 apical surface bearing a brush border of microvilli.
Unit 5 Feeding System and Digestion
In this section we will mainly be concerned with absorption of amino acids,
sugars and fats released during extracellular digestion in vertebrates. All of the
chemical and mechanical phases of digestion in the alimentary tract in the
vertebrates are directed toward changing food into forms that can pass
through the absorptive epithelial cells that line the mucosa of the small
intestine and from there into blood and lymph. As you already know from
earlier courses, the wall of the vertebrate intestine is folded and ridged to
increase the absorptive surface (Fig. 5.16 a). These ridges or folds are
covered by minute absorptive villi (Fig. 5.16 b) which further bear microvilli.
The villi are highly specialized absorptive organs with a core containing a
network of capillaries derived from blood vessels in the gut wall. Each villus
also contains a central lymph vessel known as a lacteal which begins blindly
at the tip of the villus and drains into the main lymph channels of the gut wall.

The end products of digestion, namely, amino acids, monosaccharides and The transport of
some of the lipids pass from the epithelial cells of the small intestine into the sugars and of amino
underlying blood capillaries. The rest of the lipids due to their large size cannot acids takes place
enter the capillaries and so enter the underlying comparatively larger lacteal through protein
transport molecules
(lymphatic vessel). In the present section we will study the movement of the
that depend on the
end products of digestion into the blood circulation which is accomplished via
action of sodium
any of the following processes- 1) active transport, 2) passive diffusion, 3) pumps and can be
facilitated diffusion, 4) co-transport (or secondary active transport), and 5) blocked by metabolic
endocytosis. The types of movements are briefly defined below: inhibitors such as
cyanide.
1. Active transport refers to the movement of a substances or solutes
across a cell or biological membrane from an area of lower concentration
to an area of higher concentration (against the concentration gradient).
In this type of transport, proteins within the cell membrane act as
“pumps,” using cellular energy (ATP) to move the substance.
+
Electrolytes like Na ions (Na ) are absorbed by active transport into the
blood.

2. Passive diffusion refers to the movement of substances or solutes from

an area of higher concentration to an area of lower concentration across
a biological or cell membrane. The solute diffuses down its
electrochemical gradient and at the end of the process the concentration
is equal in both areas.

3. Facilitated diffusion refers to the movement of solutes or substances

across the biological or cell membrane along the concentration gradient.
In facilitated diffusion, solutes from an area of higher concentration are
transported to an area of lower concentration by means of a carrier
protein molecule present in the cell membrane. The concentration
gradient of the solute in the area into which it diffuses never exceeds
that of the area from where the solute diffuses.

4. Co-transport (or secondary active transport) uses the movement of

one molecule through the membrane from a higher to lower
concentration to power the movement of another molecule from lower to
higher concentration.

5. Endocytosis is a transportation process in which the cell membrane

engulfs material. It requires energy, generally in the form of ATP. 151
Block 1 Fundamentals of Animal Physiology
All animal groups make use of some of the five processes in the transport of
digested nutrients. The absorption of most nutrients is through the mucosa of
the intestinal villi via the epithelial cells lining the mucosa. This absorption
usually requires active transport, and so is fueled by ATP. Sugars and amino
acids are absorbed directly by the blood capillaries while most but not all of the
Glu T2 is one of the 5 digested lipids pass into the lacteals and then into the blood circulation. Now
glucose transport let us examine the actual processes involved in absorption of the digested
molecules known to food.
carry glucose across
cell membranes. 5.4.1 Absorption of Monosaccharides
These transporters
are broadly similar in All carbohydrates are absorbed as monosaccharides. All dietary
structure and function carbohydrates that are normally digested are absorbed, leaving only
and are numbered in
indigestible cellulose and fibers in the faeces. Monosaccharides pass from the
the order of their
discovery. These lumen of the small intestine through the apical cells of the epithelial membrane
transporters change via facilitated diffusion or active transport into the blood.
their configuration, so
that one shape of the Fructose, a monosaccharide found in fruits, is carried down its concentration
transporter molecule gradient by facilitated diffusion. Glucose and galactose are transported into
binds glucose on its absorptive cells of the villi via secondary active transport process, by being
extracellular side and coupled to the active transport of Na+(Fig 5.17). The best-known transporter
the other shape binds molecule of glucose and galactose in the apical (epithelial) cells is a co-
glucose in the
transporter protein molecule termed as sodium-glucose transporter 1
intracellular side. This
(SGLT1).The co-ransporter protein molecule is so called because it couples
binding and
transportation in a the transport of a glucose or galactose molecule with that of a sodium ion. The
flip-flop manner is energy needed in the transport of glucose or galactose is provided by the
very rapid. movement of sodium ion along its gradient. The co-transporter protein
molecule enables the epithelial cells lining the lumen of the intestine to absorb
even very small traces of glucose or galactose from the digested food despite,
the fact that the epithelial cells may already have high concentrations of
glucose or galactose inside them.

+
Fig. 5.17: Mechanism for absorption of glucose: Na and glucose/galactose are
transported together from the gut lumen into the intestinal epithelial
cells by the carrier protein molecule also called cotransporter or
sodium glucose transporter 1 (SGLT 1) molecule located on the
+
epithelial cell membrane. Inside the epithelial the cell, sodium (Na ) is
moved out by the ATP pump while the glucose is taken up by another
transporter protein molecule termed as Glu T2 and transported into
152 the blood capillaries via facilitated diffusion.
Unit 5 Feeding System and Digestion
Once the glucose or galactose is inside the epithelial cell, the sodium ion is
pumped out by ATP energized active transport and the glucose or galactose
molecule is transferred to the blood stream through another transporter
molecule, called Glucose transporter 2 (Glu T2), along its concentration
gradient (Fig. 5.17). Glu T2 transports glucose or galactose via facilitated
diffusion into the blood capillaries of the villi. Glu T2 transports glucose into the
blood capillaries in proportion to the sugar concentration present in the blood.
If more glucose or galactose is present in the blood, then the transport of
glucose slows down and if glucose content of blood is low then transport of
glucose is accelerated. Galactose competes with glucose to ride the same
transporter molecule which is also termed as a symporter (defined as an The ability of the
integral membrane protein involved in the movement of different molecules or absorptive epithelial
ions in the same direction across a biological or phospholipid membrane) cells of the gut to
absorb intact proteins
since Na+ and both glucose or galactose move in the same direction, with the
even for a few days
help of the same transporter molecule.
after birth is of
immense importance
5.4.2 Absorption of Amino Acids, Dipeptides, because it allows the
and Tripeptides newborn animal to
acquire passive
Most proteins are absorbed as amino acids via active transport processes that immunity by
occur mainly in the duodenum and jejunum of the small intestine during absorbing
digestion. Normally, 95–98% of the proteins present in the small intestine are immunoglobulins
present in the
digested and absorbed. Absorption of most amino acids into the absorptive
colostral milk.
cells of the villi of the small intestine occurs via Na+ dependent secondary
active transport processes that are similar to the glucose transporter of
monosaccharides. The plasma membrane of the absorptive cell of the small
intestine bears at least four sodium-dependent amino acid transporter
molecules of which two are for transport of neutral amino acids, one for basic
and one for acidic amino acid. These transporter molecules are termed as
sodium-dependent amino acid transporters since they bind to amino acids
only after binding to a sodium ion. The fully loaded transporter molecule with
the sodium ion and amino acid at the site of release undergoes a
conformational change that releases the sodium ion and the amino acid into
the cytoplasm of the epithelial cells of the duodenum or jejunum of the small
intestine. This is followed by the absorption of the amino acid by the blood
capillaries surrounding the villi and reorientation of the transporter molecules
back into their original form.

Another separate transport system exists for the absorption of dipeptides and
tripeptides into the absorptive epithelial cells of the small intestine. A
symporter molecule brings in dipeptides and tripeptides together with a
hydrogen ion (H+) inside the epithelial absorptive cells of the small intestine
where they are hydrolyzed into single amino acids by intracellular peptidases.

There is virtually no absorption of peptides that are longer than four amino
acids, except in neonates of humans and rodents whose absorptive gut
epithelium for a very few days after birth has the ability to absorb intact
proteins after which this ability is rapidly lost.

In the adult mammals, all the amino acids, which move into the absorptive
epithelial cells of the small intestine after protein digestion, exit the absorptive 153
Block 1 Fundamentals of Animal Physiology
cells via diffusion and enter the blood capillaries. The absorbed amino acids,
similar to monosaccharides are ultimately transported via the blood stream
into the liver by the hepatic portal system. If not removed by hepatocytes, they
enter the general blood circulation.

5.4.3 Absorption of Lipids

The products of digestion of fats and oils (triacylglycerols) by the digestive
activities of enzyme lipases are free fatty acids, glycerol, and 2-
monoacylglycerols (glycerol with a single fatty acid). About 95 percent of lipids
Chyme is a thick are absorbed by diffusion into the epithelial cells lining in the small intestine.
acidic semifluid mass
Small short-chain fatty acids are more water-soluble and can dissolve in the
of partially digested
food and digestive watery intestinal chyme and pass through the absorptive epithelial cells via
secretions that is simple diffusion. They follow the same route taken by monosaccharides and
formed in the amino acids to enter the blood capillary of a villus (Fig. 5.20). However, as
stomach and intestine mentioned earlier in subsection 5.4.3 the hydrophobic large, short-chain fatty
during digestion. acids (with more than 10-12 carbon atoms), long-chain fatty acids, and
monoacylglycerides present in the chyme that enters the duodenum from the
stomach being large and not being water-soluble, are present in the form of
micelles. These micelles move from the lumen of the small intestine and bump
into the brush border of the epithelial absorptive cells. At this point the
hydrophobic but digested products of lipids fats (large chain fatty acids, long
chain fatty acids and monoacylgylycerides diffuse out of the micelles into the
absorptive epithelial cells of the small intestine, leaving the micelles behind in
the chyme. The micelles continually repeat this ferrying function as they move
from the brush border back through the chyme to the lumen of the small
intestine to pick up more of insoluble fatty acids, monoglycerides cholesterol
and fat-soluble vitamins.

The free fatty acids and monoacylglycerides that enter the absorptive
epithelial cells of the small intestine recombine together to reform into
triglycerides within the absorptive epithelial cells. The recombined triglycerides
mix with phospholipids and cholesterol present in the absorptive cells, and get
enclosed in a protein coat, forming a new water-soluble lipoprotein complex
called chylomicron (Fig.5.18). The large sized chylomicrons are processed
by the Golgi apparatus within the absorptive, epithelial cells after which they
exit these epithelial cells of the small intestine. The chylomicrons are unable to
directly enter the blood capillaries due to their large size and so instead enter
into the central lacteal vessel of the small intestine. The central lacteal vessel
is the lymph vessel located at the core of each villus. The lacteals contain
patches of endothelial cells that are held together by specialized "button
junctions" which are much more permeable to chylomicrons than the normal
cellular junctions. The lymph containing the chylomicrons passes through the
button junctions into the lacteals and from there into larger vessels of the
lymphatic system and eventually reaches the circulatory system. Once in the
circulatory system, the enzyme lipoprotein lipase breaks down the
triglycerides of the chylomicrons into free fatty acids and glycerol which can
then pass through capillary walls either to be used for energy by cells or for
getting stored in adipose tissue as fat. Liver cells combine the chylomicrons
remnants with proteins to form lipoproteins which transport cholesterol in the
154 blood. Two or three hours after a meal, few chylomicrons remain in the blood.
Unit 5 Feeding System and Digestion
After participating in the emulsification and absorption of lipids, most of the bile
salts are reabsorbed by active transport in the ileum of the small intestine and
are returned by the blood to the liver through the hepatic portal system for
recycling.

Fig. 5.18: Fatty acids and monoglycerides from the micelles are absorbed by the
epithelial cells of the small intestine and resynthesised into
triglycerides. These are then coated by protein to form chylomicrons
which enter the lacteals of the small intestine.

SAQ 4
a) Which chemical among the following is absorbed mainly in the large
intestine?

i) glucose

ii) amino acid

iii) sodium

iv) water

b) Why are villi present in the small intestine and not in the stomach?

5.5 PHYSIOLOGY OF DIGESTION IN HUMAN

MAMMALS
In the earlier sections you have read that the process of digestion is similar in
vertebrates, including mammals. Now that you know the general mechanisms
of digestion and absorption let us see how those can be applied in the
physiology of digestion in humans which we take as an example of a mammal.

5.5.1 Digestion in Oral Cavity

A pair of lips protects the oral cavity (mouth) and helps retain the food as it is
being chewed. The oral cavity contains the tongue and teeth (Fig. 5.19) .The 155
Block 1 Fundamentals of Animal Physiology
process of food digestion begins in the oral cavity as soon as a bite of food
enters into it. The mechanical action of the teeth and tongue and the chemical
action of saliva begin to break down the ingested food. Three major pairs of
salivary glands constantly secrete watery fluid called saliva.

Fig.5.19: Human Digestive system and its associated glands. Digestion begins
in the mouth and continues in the stomach and small intestine as the
food travels through the alimentary canal. Most absorption occurs in
the small intestine while some absorption especially water takes place
in the large intestine.

Among the three pairs of salivary glands: i) parotid glands produce a thin,
watery, serum like secretion that contains a large amount of salivary amylase
enzyme which breaks down starches (complex carbohydrates); ii) sublingual
glands produce mucous that acts as a buffer and lubricant; and iii)
submandibular glands secrete a mixture of buffers, salivary amylase and
glycoproteins called mucins which in contact with water produce mucous.
Lingual glands are present in the tongue and secrete both mucous and a
watery lingual lipase which acts on as much as 30% of dietary triglycerides
converting them into simpler fatty acids and diglycerides.

The salivary mucous serves to moisten and lubricate the food for easier
swallowing, protects the gums against abrasions, and facilitates taste and
smell. The saliva contains bicarbonate ions (HCO3-), which buffers chemicals
in the mouth, and help prevent tooth decay. The saliva also contains
lysozymes which kill microorganisms that enter the mouth with food.

The function of the salivary amylase is to begin starch (a polysaccharide)

digestion by cleaving the bonds that link glucose monomers to produce
disaccharide such as maltose, trisaccharide such as maltotriose, and short-
156 chain glucose polymers called α-dextrins.
Unit 5 Feeding System and Digestion
The tongue movements manipulate the mixture of saliva and food, into a ball
shaped structure called a food bolus. The food bolus, once formed is pushed
with the help of the tongue to the back of the oral cavity (Fig. 5.20) and
triggers swallowing or deglutition of the bolus into pharynx/throat. Swallowing
can be voluntary, but most of the time it is involuntary. At the start of
swallowing, food is prevented from entering into the respiratory system by the
larynx which moves upwards, closing the epiglottis and so prevents the food
from entering into the respiratory tract (Fig. 1.12). Once swallowing of food is
begun the nervous system completes it due to a nerve reflex. Prior to the entry
of the food bolus into the oesophagus, the oesphageal sphincter relaxes,
allowing the bolus to enter from the pharynx. Unlike the pharynx the hollow,
muscular tube like, oesophagus actively moves the food bolus into the
stomach by peristalsis which is automatic and is caused by alternate
contraction and relaxation of the circular and longitudinal muscles which line
the alimentary canal from the esophagus to the anus. Mucous secreted by the
esophageal glands lubricates the bolus and reduces friction. The passage of
solid or semisolid food from the mouth to the stomach takes 4 to 8 seconds;
very soft foods and liquids pass through in about 1 second. Neither the
pharynx nor the esophagus contributes to digestion.

Fig. 5.20: Intersection of the human airway and digestive tract: In humans, the
pharynx is the common passage for the oesophagus of the digestive
system and the trachea of the respiratory system and at most times :
a) a contracted sphincter seals off the oesophagus while the trachea
remains open; b) When a food bolus arrives at the pharynx, the
swallowing reflex is triggered due to which the upper part of the
airway, the larynx moves upwards which tips a flap of tissue called
the epiglottis downwards, thus preventing the food from entering the
trachea. At the same time, the esophageal sphincter relaxes, allowing
the bolus to pass into the oesophagus. Once the bolus passes into the
oesophagus the trachea then reopens, and peristaltic contractions of
the esophagus move the bolus to the stomach.

5.5.2 Digestion in Stomach

The stomach, located just below the diaphragm is a muscular, distensible sac
with accordion-like folds to (Fig. 5.21 a) accommodate about 2 liters of food
and fluid and is guarded by valves called sphincters on both its ends - the 157
Block 1 Fundamentals of Animal Physiology
gastro-oesphageal sphincter at the oesophageal end of the stomach and the
pyloric sphincter located at the lower end of the stomach. The stomach
mechanically and chemically digests the food bolus by its muscular
contractions and by the action of hydrochloric acid and gastric enzymes. The
wall of the stomach as you have learnt earlier in the second semester is
composed of the same four layers that make up most of the
gastrointestinal(GI) tract : innermost mucosa, which is enclosed by
submucosa. Submucosa is enclosed by muscularis mucosa which is covered
by serosa (Fig. 5.21 a). The epithelial layer of the stomach mucosa is a
secretory sheet consisting of simple columnar epithelium that contains many
types of cells (Fig. 5.21 b).

Fig. 5.21: a) The human stomach with; b) its absorptive cells and secretions;
c) production of HCl by the parietal cells.

Glands in the pyloric part of the stomach secrete alkaline mucous rather than
enzymes or acids. Mucous covers the entire surface of the stomach protecting
its inner epithelial lining against the action of acid and enzymes secreted by
the stomach cells. The fundus region of the stomach contains thousands of
gastric glands that secrete gastric juices .Gastric glands are dominated by two
types of secretory cells namely, parietal cells and chief cells. Parietal cells
secrete hydrogen ions that form the hydrochloric acid in the lumen of the
stomach (Fig. 5. 21 c). Parietal cells also secrete an intrinsic factor, a
glycoprotein that helps in the absorption of vitamin B12 across the lining of the
small intestine. The chief cells secrete pepsinogen, the precursor of the
protease enzyme pepsin. Both the cells are present in the pits of the gastric
158 glands. The secretions of the mucous, parietal, and chief cells constitute the
Unit 5 Feeding System and Digestion
gastric juice, which totals 2000–3000 milli liter (ml)/day and which is a
nearly colorless, strongly acidic, liquid consisting of mucous, hydrochloric acid
(HCl), and the proteolytic digestive enzymes pepsin and rennin (found
primarily in the gastric juice of infants which acts on milk proteins converting
casein into paracasein).

In addition, several types of enteroendocrine cells are scattered among the

mucous secreting cells lining the stomach. These enteroendocrine cells
produce at least seven hormones, of which the most notable Gastrin, is
produced by G cells, which are most abundant in the gastric pits of the pyloric
region. Gastrin (Refer again to subsection 15.5 of this unit) stimulates the
secretion of both parietal and chief cells, as well as contractions of the gastric
wall that mix and stir the gastric contents. The pyloric region also contains
enteroendocrine D cells which secrete somatostatin, a hormone that inhibits
the release of gastrin. D cells continuously release their secretions into the
interstitial fluid adjacent to the G cells. Neural and hormonal stimuli can
override this inhibition of gastrin production when the stomach is preparing for
digestion or is already engaged in digestion. Several other hormones play a
role in hunger and satiety (feeling of not being hungry). Another hormone from
the stomach and small intestine, the obestatin, is thought to decrease
appetite and inhibit thirst.

The food bolus may remain in the stomach for about an hour without
becoming mixed with gastric juice. During this time, digestion by salivary
amylase from the salivary glands continues. Soon, however distention of the
stomach due to the arrival of food as well as the act of eating causes the
gastric pits to secrete HCl and pepsinogen. The HCl inactivates the salivary
amylase and activates the lingual lipase that acts in an acidic medium.

Parietal cells use an ATP driven pump to expel hydrogen ions produced by
The gastro-
them into the stomach lumen. At the same time, chloride ions present in the oesphageal sphincter
parietal cells diffuse into the stomach lumen through specific membrane at the promimal end
channels of the parietal cells. It is therefore only within the stomach lumen that of the stomach
hydrogen and chloride ions combine to form HCl (see Fig. 5.21 b).The occasionally, allows a
concentration of HCl is high in the stomach, causing the pH of gastric juice to back movement, or
flux, of the chyme
be about 2, which is acidic enough to dissolve iron nails (and to kill most
from the stomach
bacteria) back into the lower
end of the esophagus
Hydrochloric acid (HCl) in the stomach lumen disrupts the extracellular matrix which causes painful
that binds cells together in meat and plant material. It further increases the irritation of the
exposure of the peptide bonds of the food present in the stomach for the esophagus. This is
digestive enzymes to act upon. Protein digestion in humans starts only from referred to as acid
the stomach. HCl converts pepsinogen to active enzyme pepsin (a protease, reflux but is more
commonly known as
or protein-digesting enzyme), by clipping off a small portion of it and exposing
“heartburn”.
its active site. As you must have noted, both HCl and the active pepsin form in
the lumen (cavity) of the stomach and not within the cells of the gastric
epithelium. This ensures that parietal and chief cells are not digested from
within by their secretions. Furthermore, the lining of the stomach is replaced
every 5-6 days to prevent damage to the stomach. The hydrochloric acid
converts a small amount of inactive pepsinogen into active pepsin by clipping
off a small portion of it and exposing its active site. The pepsin, then itself 159
Block 1 Fundamentals of Animal Physiology
autoactivates pepsinogen into pepsin. Pepsin, similar to HCl, can clip
pepsinogen to expose the enzyme’s active site (see Section 1.4 again).
Digestion of protein is also aided by the mechanical action of the mucosal
muscles of the stomach which contract and vigorously churn and mix the
stomach contents about every 20 seconds. About three to four hours after a
meal, the stomach contents are sufficiently mixed to the form an acidic, semi-
liquid chyme which contains the hydolysed proteins that have been converted
into smaller and simpler polypeptides by the action of enzyme pepsin.

The pyloric sphincter of the stomach regulates the passage of chyme into the
duodenerm of the small intestine, allowing only one squirt of chyme to enter at
a time.” When the stomach is empty, peristaltic waves cease. However, after
about 10 hours of fasting, new waves may occur in the upper region of the
stomach. These waves can cause “hunger pangs’’ as sensory nerve fibers
carry impulses to the brain.

5.5.3 Digestion and Absorption in Small Intestine

The stomach leads into the small intestine which is the longest section of the
digestive tube and begins at the pyloric sphincter of the stomach. The small
intestine coils through the central and inferior part of the abdominal cavity, and
eventually opens into the large intestine. In humans the small intestine
consists of three segments that form a passage from the beginning of the
small intestine upto the beginning of the large intestine. The three segments of
the small intestine are as follows:

• Duodenum: is the first segment of the small intestine. It is a short

section that receives secretions from the pancreas and liver via
the common bile ducts respectively.

• Jejunum: is the mid segment part of the small intestine after the
duodenum and roughly forms 40% of the small intestine in humans,

• Ileum: is the posterior most part of the small intestine after the
jejunum and forms about 60% of the small intestine in humans. Ileum
empties into the large intestine.

The wall of the small intestine similar to the stomach is composed of: mucosa,
submucosa, muscularis, and serosa and similarly the mucosa is lined by a
layer of epithelium consisting of simple columnar epithelial cells. This
epithelium also contains many types of secretory cells. As you will recall from
the previous course BZYCT-133, the total surface area of the small intestine is
greatly increased by large circular folds present in them and by the finger
shaped projections called villi that project from the epithelial lining of the
circular folds (Fig. 5.22 also see Fig. 5.23). This total surface area of the small
intestine is further increased by the fact that the villi also have microvilli which
are projections of the apical (free) membrane of the villi (Fig.5.23 b & c).

Absorptive cells also called enterocytes (Fig. 5.23) present in the intestinal
epithelium of the muscosa of the small intestine by means of their microvilli
take up and deliver into the blood virtually all nutrients from the ingested food.
Some other cells are also present in the epithetium lining the lumen of the
160 small intestine and release enzymes that digest the food in the chyme. Also
Unit 5 Feeding System and Digestion
present in is epithelium are goblet cells, which secrete mucous (Fig. 5.23).
The mucosa of the small intestine contains many deep crevices lined with
glandular epithelium. Cells lining the crevices between the villi form crypts of
Lieberkühn (Fig. 5.23) which are involved primarily in the secretion of
electrolytes.

Fig. 5.22: Micrographs of Intestinal folds or villi enlarged to show their

composition in the human intestine. (Sourse: http://www.vivo.colostate
.edu/hbooks/pathphys/digestion/smallgut/anatomy.html).

Fig. 5.23: Diagrammatic representation of cross section of small intestine,

showing the major constituents of the epithelium. The intestinal
epithelium in lined with a single layer of epithelial cells among which
the major types include enterocytes, globlet cells, Paneth cells, stem
cells etc.

The absorptive cells or enterocytes present in the epithelial lining of the small
intestine are major cell types and play an important role in nutrient absorption
(water, sugar, peptides, and lipids and secreting immunoglobulins. Besides
absorptive cells and goblet cells, the intestinal epithelial lining of the small
intestine also contains paneth cells and three types of enteroendocrine (earlier
called neuro endrocrine) cells (Fig. 5.24). Paneth cells may have a role in
regulating the microbial population in the small intestine as they are capable of 161
Block 1 Fundamentals of Animal Physiology
phagocytosis and secrete the bactericidal enzyme lysozyme. The three types
of enteroendocrine cells found in the intestinal epithelium of the small intestine
are S cells, CCK cells, and K cells, which secrete the hormones secretin,
cholecystokinin (CCK) and glucose-dependent insulinotropic peptide
(GIP) respectively. Importantly, towards the base of the crypts of Lieberkühn
are stem cells (Fig. 1.15), which continually divide and provide the source of
all the epithelial cells. Stem cells in the crypts divide to form daughter cells.
One daughter cell from each stem cell division is retained as a stem cell. The
other becomes committed to differentiate along one of four pathways to
become absorptive cells, or goblet cells, or paneth cells, or enteroendocrine
The fat-soluble cells.
vitamins A, D, and E
are absorbed in the The chyme containing, partially digested carbohydrates, proteins, and lipids,
upper small intestine. enters the duodenum which is the first section of the small intestine, and is
The factors that primarily involved in further digestion of the food. The submucosa of the
cause the duodenum contains duodenal glands, called Brunner’s glands, which secrete
malabsorption of fat an alkaline mucous that helps neutralize the gastric acid of the chyme entering
can also affect the
the duodenum from the stomach. The complete digestion of carbohydrates,
absorption of these
proteins, and partial digestion of lipids, is accomplished in the duodenum by
vitamins. Vitamin B12
is absorbed in the the collective action of: 1) intestinal gland cells, located in the duodenum wall,
ileum and must be which secrete daily about 1-2 liters of intestinal juice which is a clear, slightly
bound to an intrinsic alkaline (pH 7.6), yellow fluid, containing water and mucous. The alkaline pH
factor, a protein of intestinal juice is due to its high concentration of bicarbonate ions (HCO3-).
secreted in the 2) alkaline bile juice containing bile salts, produced by liver, stored in the gall
stomach, in order to bladder and released from it into the duodenum and (3) the alkaline digestive
be absorbed. If
juices from the pancreas. Thus an alkaline liquid medium is provided in the
intrinsic factor is
missing, then Vitamin duodenum that aids the digestion and absorption of nutrients from chyme.
B12 is not absorbed
The presence of partially digested proteins and the entry of the acidic chyme
and pernicious
into the duodenum, triggers the CCK cells in the small intestine to secrete
anaemia results.
hormone cholecystokinin. Pancreatic enzymes enter the duodenum in
response to secretion of cholecystokinin. Most of the digestive enzymes that
act on the chyme in the small intestine are secreted by the pancreas and enter
the small intestine via the common bile duct. You already know from Section
1.5 that the pancreatic enzymes include the following: 1) proteolytic enzymes
which include - i) proteases which break apart large protein complexes, and ii)
peptidases, which break small peptide chains into individual amino acids. 2)
alpha (α) amylase, a carbohydrate digesting enzyme that breaks down certain
starches (pancreatic alpha (α) -amylase is almost identical to salivary
amylase). 3) pancreatic lipase, which breaks down certain complex lipids/fats,
releasing products (such as fatty acids) that can be easily absorbed. 4)
Nucleases, which break down RNA or DNA. Nucleotides that result from the
action of the nucleases are further digested by brush-border enzymes called
nucleosidases and phosphatases into pentoses, phosphates, and
nitrogenous bases. These products are absorbed via active transport by the
small intestine.

Only a few starches are digested in the chyme leaving the stomach. Most of
the carbohydrate digestion also takes place in the duodenum of the small
intestine. The pancreatic enzyme amylase digests those starches that have
162 not already been broken down in the stomach. Although pancreatic amylase
Unit 5 Feeding System and Digestion
acts on both glycogen and starches, it has no effect on another polysaccharide
called cellulose, an indigestible plant fiber that is commonly referred to as
“roughage” as it moves through the digestive system.

The brush-border cells of the duodenum of the small intestine secrete four
carbohydrate-digesting enzymes called alpha (α) -dextrinase, maltase,
sucrase, and lactase which digest ingested molecules of carbohydrates into
disaccharides and monosaccharides. Alpha (α)-dextrinase acts on α-dextrins,
clipping off one glucose unit at a time. Maltase splits maltose and maltotriose
into two or three molecules of glucose. Sucrase breaks sucrose into a
molecule of glucose and a molecule of fructose. Lactase digests lactose into a
molecule of glucose and a molecule of galactose. Digestion of carbohydrates
ends with the production of monosaccharides, which the small intestine is able
to absorb.

Fats in the form of triglycerides and phospholipids are also digested primarily
in the duodenum of the small intestine. We have already explained in
subsection 1.6.3 how the end products of lipid digestion are absporbed by the
absorptive cells of the small intestine. The absorption process of lipids in
humans occurs in the same manner.

The chyme of the duodenum next moves by peristalsis into the jejunum which
similar to duodenum is involved in both digestion and absorption of nutrients
and so also secretes fluids. From jejunum, the contents of digested food move
by peristalsis into the ileum, which is the posterior most part of the small
intestine and is primarily involved in absorption of nutrients that have been
digested in the duodenum. Thus the digestion and absorption of
carbohydrates and proteins is completed in the small intestine. Most of the
lipids are also digested and absorbed in the small intestine.

Once digestion is largely complete in the duodenum the absorption of the end
products of digestion namely amino acids, simple sugars, fatty acids, glycerol,
and nucleotides takes place across the lining of the jejunum and ileum of the
small intestine. In addition to absorbing nutrients, the jejunum and ileum also
absorb most of the water, vitamins and dissolved mineral ions. Depending on
the nutrients, absorption of the digested nutrients across the epithelial cells of
the small intestine can be by the passive or active process and is the same as
described in Section 1.6. Thus absorption in mammals (humans) involves
active transport and sodium dependent ATPase pump about which you have
already learnt in Section 1.6 in greater detail.

5.5.4 Absorption in Large Intestine

The digested food from the small intestine reaches the large intestine. The
large intestine includes as you know the colon, caecum and rectum. The
caecum is important in mammals that eat large amounts of plant material as
fermenting of ingested material, occurs in it. Compared with many other
mammals, humans have a small caecum.

The wall of the large intestine contains the typical four layers found in the rest
of the GI tract: mucosa, submucosa, muscularis, and serosa. The mucosa is
similarly covered by simple epithelial layer of columnar cells, which contains
mostly goblet and absorptive cells. The goblet cells secrete mucous that 163
Block 1 Fundamentals of Animal Physiology
lubricates the passage of the colonic contents while the absorptive cells are
primarily involved in water re-absorption. The major function of the large
intestine is reabsorption of water and minerals, and the formation and storage
of faeces. It takes 12-24 hours for food material to travel the length of the
colon, which similar to the small intestine also osmotically recovers some of
the remaining water which then returned to the lymphatic system and
bloodstream. Typically most of the water consumed by a person in a day is
reabsorbed by the small and large intestines per day and only about 0.1 litre of
the water is not reabsorbed. By the time the chyme has remained in the large
intestine for 3-10 hours, it becomes increasingly solid or semisolid because of
water absorption. Insufficient, water reabsorption by the small and large
intestine causes diarrhea while excessive reabsorption of water by the small
and large intestine causes the faecal matter to become too thick, resulting in
constipation.
Bile secretion besides Minerals diffuse or are actively transported from the food residue across the
its digestive function epithelial surface of the large intestine into the bloodstream as peristaltic
also serves as the
waves move food residues along, in the large intestine.
route of excretion of
bilirubin, pigments The final stage of digestion occurs in the colon through the activity of bacteria
generated due to the that inhabit the lumen. Bacterial products that are absorbed in the colon
destruction of red
include several vitamins needed for normal metabolism. Chyme for elimination
blood cells that are no
is prepared, by the action of bacteria, which ferment any remaining
longer fully functional.
Bilirubin pigments carbohydrates and release hydrogen, carbon dioxide, and methane gases.
released as a result These gases contribute to flatulence in the colon. Bacteria also convert any
of red blood cell remaining proteins to amino acids and breakdown the amino acids into simpler
destruction are substances, some of which get eliminated in the faeces and contribute to the
incorporated into bile odour of the faeces while the rest are absorbed and transported to the liver. In
pigments, and the liver these simipler compounds are converted to less toxic compounds and
eliminated from the
excreted in the urine. Bacteria also decompose bilirubin of the bile juice into
body along with
faeces. In some liver
simpler pigments, including stercobilin, which gives faeces their brown color.
and blood disorders, The faeces is a mixture of water, inorganic salts, bacteria, products of bacterial
bile pigments decomposition, undigested plant fiber, sloughed-off intestinal cells, and other
accumulate in the waste products. Faeces are stored in the rectum, before being discharged
skin, resulting in its outside by the anus. This process of the faeces being discharged outside
yellow appearance. through the anus is called egestion (take care not to confuse egestion
The disease is called with excretion).
jaundice.

SAQ 5
a) The mammalian trachea and esophagus both connect at their proximal
end to the:

i) pharynx

ii) stomach

iii) large intestine.

iv) Rectum

b) Where does the chemical digestion of starch begin?

164 i) mouth
Unit 5 Feeding System and Digestion
ii) esophagus

iii) stomach

iv) small intestine

c) Which among these enzymes is involved in the chemical digestion of

protein?

i) pancreatic amylase

ii) trypsin

iii) sucrase

iv) pancreatic nuclease

d) Digestion does not take place in the:

i) duodenum

ii) oesophagus

iii) mouth

iv) stomach

e) Which group of nutrients do not require to be digested?

i) Proteins, and vitamins, water

ii) Fats, minerals and proteins

iii) Minerals, vitamins and water

iv) Water, carbohydrates and minerals

f) What would happen if the stomach is unable to secrete hydrochloric acid?

g) Why is bile juice important in digestion?

5.6 ENERGY METABOLISM IN ANIMALS

In the preceding sections we have studied how the products of digestion are
absorbed and transported to the tissues via the blood. You have also learnt
that generally carbohydrates and fats are the fuel that provide energy. The
oxidation of these compounds yields virtually all the chemical energy required
by the body of the animal. This use of chemical energy is referred to as the
animal’s energy metabolism. We can also define energy metabolism as the
process of generation of ATP from the nutrients by a series of interconnected
pathways that function in the presence or absence of oxygen. You will learn
more about the chemical pathways in Volume 2 of this course. Aerobic
metabolism converts one molecule of glucose into 30-32 ATP molecules.

The rate at which energy is used by the body during both external and internal
activities is known as the metabolic rate. Most of the body’s energy
expenditure or usage eventually appears in the form of heat, and so metabolic 165
Block 1 Fundamentals of Animal Physiology
rate is normally expressed in terms of the rate of heat production in
kilocalories per hour. The basic unit of heat energy is the calorie, which if you
can recall “is the amount of heat required to raise the temperature of 1 g of
water (H2O) by 1°C”. This unit is however, considered too small to be
convenient when discussing the enormous amount of heat involved in the
human body and so as a result the energy in the human or mammalian body is
calculated in kilocalorie or Calorie, which is equivalent to 1000 calories. Thus,
when nutritionists speak of “calories” in quantifying the energy content of
various foods, they are actually referring to kilocalories or Calories.

One of the ways used to find out the rate of metabolism or the actual amount
of energy liberated in oxidative metabolism is to measure the total heat
produced by the organism per unit of time that is

Metabolic rate = energy expenditure/unit of time or

Metabolic rate = Rate of energy intake – rate of energy loss per unit
time

Energy intake is the chemical energy content of the ingested food over a given
time period and energy loss is the chemical energy that remains in the faeces
and urine produced by the animal over the same time period. Through
experiments it has been found out that carbohydrates and lipids are fully
oxidized to carbon dioxide and water during metabolism but proteins are not
fully oxidized since the end product of protein metabolism is i.e. urea
possesses some energy. The energy contained in energy-yielding nutrients
such as carbohydrates and lipids differ because the energy-yielding nutrients
are composed of different types of chemical bonds. Carbohydrate or protein in
food yields approximately 4 kilocalories per gram of energy, whereas the
triglycerides that compose fat in food yield 9 kilocalories per gram of energy.

Although carbohydrates, proteins, and fats require different amounts of O2 for

their oxidation and yield different amounts of kilocalories when oxidized, an
average estimate referred to as energy equivalent of O2, can be made on the
basis of the quantity of heat produced per liter of O2 consumed. The most
convenient and commonly used measure to determine metabolic rate of an
animal is to find the oxygen consumption per unit of time since the heat
produced for each liter of oxygen used is the same for all nutrients (proteins,
lipid and carbohydrates) irrespective of the type of food oxidized. Therefore, it
has been found that on an average the value of energy (in the form of heat)
liberated per liter of O2 consumed, is 4.8 kcal. It has also been determined that
oxygen consumption per unit weight per unit time decreases with higher body
weight of animals. In other words a mouse or shrew has a much higher
metabolic rate than an elephant; consequently smaller animals need to feed
constantly while larger animals can survive without food for a longer time.

Energy storage

Food intake and energy expenditure for animals is approximately equal. If

energy expenditure exceeds food intake, then the excess energy is obtained
by utilization of body fat. However, if food intake is excess, then the surplus is
166 stored as fat irrespective of the kind of food eaten.
Unit 5 Feeding System and Digestion
Excess carbohydrates are changed to fats. This is because fats contain
relatively less oxygen and the excess oxygen of carbohydrates is used in the
metabolism. Conversion of carbohydrate to fat thus reduces the oxygen
uptake by the animal and as a result the respiratory carbon dioxide to oxygen
ratio is increased. For this reason fat is an ideal storage material for energy. It
is much lighter and yields twice as much energy as the carbohydrates.
Migratory birds that may have to fly more than 1000 km non-stop, carry fat as
40% to 50% of their body weight.

Nonetheless, some carbohydrates are important in energy storage. Glycogen,

a starch-like carbohydrate polymer is stored as granules in the skeletal
muscles and liver of vertebrates. During heavy muscular exercise when blood
does not deliver sufficient oxygen to meet demands, glycogen provides the
energy. Glycogen is broken down directly into glucose-6-phosphate, providing
fuel for carbohydrate metabolism more directly than fat does.

On the other hand, many animals that do not move about, also store glycogen
as an excess energy source. For example, clams, oysters and many intestinal
parasites like Ascaris use glycogen as the energy storage material. These
animals have to face anaerobic conditions and in such situations glycogen
breaks down to acetic acid to yield energy.

SAQ 6
A person who is on a diet and does not consume any fat but only rice and
sweets still keeps gaining weight. Why?

5.7 SUMMARY
In this unit you have studied that:

• Food is obtained by animals in different ways and feeding strategies

have evolved according to the nature of food required.

• Digestion is the breakdown of complex food molecules into simpler

constituents. Two broad categories of chemical digestion are seen -
intracellular and extracellular. Intracellular digestion is characteristic of
primitive animals and extracellular digestion of the higher forms.
Extracellular digestion of proteins, carbohydrates and lipids is a step by
step process in which large food molecules are broken down by the
action of specific enzymes. Enzyme lipases are not as specific as
carbohydrases and proteases.

• Secretion of digestive juices is under neural and hormonal control. All

gastrointestinal hormones are peptides. Both direct activation by the
arrival of food chyme in the gut and neural activation stimulate the
endocrine cells of the alimentary canal to secrete hormones.

• The products of digestion are taken up by the absorptive cells of the

intestine and transferred to the blood capillaries and lymphatic system.
Transport of some sugars occurs by facilitated diffusion which does not 167
Block 1 Fundamentals of Animal Physiology
require metabolic energy. Most sugars and amino acids are transported
with Na+ and a carrier molecule requiring energy. The products of fat
digestion are absorbed by diffusion across cell membranes.

• The absorbed nutrients provide fuel for body metabolism. The chemical
energy used in metabolism can be measured as heat energy. A given
class of food will liberate the same amount of heat and require the same
amount of oxygen when oxidized to its end products of carbon dioxide
and water. The heat production and oxygen consumption helps provide
information about the metabolic rate of an animal.

5.8 TERMINAL QUESTIONS

1. What prevents the epithelial lining of the walls of the stomach of animals
from being digested by the HCl secreted by it?

2. What are the end-products of food that can be absorbed by the body?
Explain how absorption of fats differs from absorption of proteins and
sugars.

3. Fill in the names of the enzymes associated with carbohydrate digestion

in humans in the chart given below:

5.9 ANSWERS
Self-Assessment Questions
1. a) Tapeworm

b) Amoeba

c) Hydra

d) Aphids

168 e) Spiders
Unit 5 Feeding System and Digestion
2. a) i) Add a water molecule to break bonds
ii) They emulsify the large fat droplets
iii) Salivary and pancreatic amylase

b) Gall bladder - bile juice synthesis

c) Nutrient absorption
3. Gastrin, Secretin, Cholecystokinin.

4. a) Water
b) The small intestine is mainly responsible for the absorption of the end
products of digestion. The villi and their microvilli in the small
intestine increase the absorptive surface area of the small intestine
so that a considerably larger amount of the end products of digestion
can be absorbed. The stomach, on the other hand, secretes HCl and
digestive enzymes which aid in food digestion and so the stomach
does not require villi like structures.
5. a) pharynx
b) mouth
c) trypsin
d) oesophagus
e) Minerals, vitamins and water
f) Hydrochloric acid disrupts the extracellular matrix that binds cells
together in meat and plant material ingested by the mammal. HCl
also activates pepsinogen into pepsin which requires an acidic
environment for activation. The activated pepsin in the stomach
digests proteins. Lingual lipase also acts on lipids in an acidic
medium. Digestion will not take place in stomach if HCl is not
secreted by it as in the absence of HCl the extracellular matrix that
binds the cells together in meat and plant material will not be
disturbed and the partial lipid and protein digestion by lingual and
gastric lipase and pepsin respectively will not take place as both
lipase and pepsin act in an acidic medium. Furthermore, activation of
pepsinogen into pepsin will not take place so protein will not be
digested in the stomach. HCl also kills the harmful bacteria in the
stomach and so in the absence of its production the growth of these
harmful bacteria may occur in the stomach which may adversely
affect the health of the individual.
g) Bile is secreted by the liver. It plays an important role in digestion.
First being alkaline in nature the bile juice helps in neutralizing the
acidic chyme and making the environment of the duodenum of the
small intestine alkaline. The most important role the bile juice plays is
that its causes the emulsification of fats. Bile juice also helps in the
absorption of fats. The bile salts reduce the surface tension of the
large fat droplets and convert them into smaller droplets. Lipase
which acts on these smaller fat droplets is activated by bile salts.
6. Excess carbohydrates that are not used for metabolism are converted to
fats and stored in the body. This increases the body weight. 169
Block 1 Fundamentals of Animal Physiology
Terminal Questions
1. Mucous is secreted by the goblet cells of the epithelial lining of the
stomach which coats the entire digestive tract and prevents the action of
HCl and enzymes. Furthermore, most of the digestive enzyme especially
the proteases are present in inactive form and HCl does not form in the
cells lining the stomach, instead it forms in the lumen of the stomach and
so the cells lining the stomach are not corroded by the action of HCl. In
addition the stomach lining is replaced within a few days so in case the
lining gets damaged it would soon be replaced.

2. Carbohydrates are hydrolysed to monosaccharides. Proteins are

hydrolysed to amino-acids and fats are hydrolysed to the free fatty acids
and glycerol.

i) Amino acids and glucose enter the intestinal epithelial cells by a

carrier or cotransporter protein molecules that depend on the
action of a sodium ion pump. Small short chain fatty acids are
water soluble and diffuse into the epithelial cells of the duodenum.
However, the hydrophobic large, chain fatty acids, long, chain fatty
acids and monoacylglycerides form micelles with bile salts and
lecithin before entering the intestinal epithelium through diffusion.

ii) Amino acids and glucose are passed on to the blood stream
through another transporter molecule. Small, short, chain acids
diffuse into the blood stream while fatty acids and
monoacylglycerides form chylomicrons within the epithelial cells of
the mucosa of the small intestine. Chylomicrons then move from
the epithelial cells of the mucosa of the small intestine and enter
the lymphatic lacteals from where they enter the blood stream.

170