COURSE

GUIDE

BIO 314
ANIMAL ECOLOGY

Course Team Miss. Fisayo Chistie Olakolu (Course Writer )

Nigerian Institute for Oceanography and Marine
Research, Victoria Island, Lagos, Nigeria
Dr. Adefunke M. Adesina (Course Editor)-
Ministry of Health Ala usa, Lagos, Nigeria
Dr. Maureen N. Chukwu (Course Coordinator)-
Noun

NATIONAL OPEN UNIVERSITY OF NIGERIA

BIO 314 COURSE GUIDE

© 2023 by NOUN Press

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Published by
National Open University of Nigeria

Printed 2015, Reprinted 2023

ISBN: 978-058-535-7

Reviewed 2023

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BIO 314 COURSE GUIDE

INTRODUCTION

Animal Behaviour (BIO314) is a fundamental course for undergraduate

students of Biology that deals with the theories and principles of adaptive
behaviour and evolution of animals. The course contents are history
of ethology. Reflex and complex behaviour. Orientation and taxes.
Fixed action patterns, releasers, motivation and driver. Displays,
displacement activities and conflict behaviour. Learning communication
and social behaviour. The social behaviour of primates. Hierarchical
organization. The physiology of behaviour. Habitat selection, homing
and navigation. Courtship and parenthood. Biological clocks.

COURSE COMPETENCIES

This course aims to enable you to understand the theories and principles
of adaptive behaviour and evolution of animals, Behavioural Patterns
and Mechanism of Adaptation

COURSE OBJECTIVES

The Comprehensive Objectives of the Course are to;

1. Explain emotion in animals with examples

2. Explain the different forms of communication
3. Explain the different taxes according to their response to
stimulus
4. The phenomenal behind biological clocks.

WORKING THROUGH THIS COURSE

To successfully complete this course, you are required to read each study
unit, read the textbooks and other materials provided by the National
Open University.

Reading the reference materials can also be of great assistance. Each unit
has self –assessment exercise which you are advised to do.

There will be a final examination at the end of the course. The course
should take you about 8 weeks to complete.

This course guide provides you with all the components of the course,
how to go about studying and how you should allocate your time to each
unit so as to finish on time and successfully.

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BIO 314 COURSE GUIDE

STUDY UNITS

The course is divided into 3 modules and study units in this course are
given below::

BIO 314 ANIMAL Behaviour (2 UNITS)

Module 1 History of Ethology

Unit 1 Historical Background of Ethology

Unit 2 Reflex and Complex bahaviour
Unit 3 Orientation in Animals
Unit 4 Taxes in Animals
Unit 5 Fixed action Pattern, Motivation and Drive

Module 2 Learning, Communication and Social Bahvaiour

Unit 1 Display, Displacement Behaviour and Conflict

Behaviour
Unit 2 Learning and Communication in Animal
Unit 3 Social Behaviour
Unit 4 Social Behaviour of Primates
Unit 5 Hierarchical Organisation

Module 3 Habitat Selection, Homing and Navigation and

Courtship Behaviour

Unit 1 Physiology of Behaviour

Unit 2 Habitat Selection
Unit 3 Homing and Navigation in Birds
Unit 4 Courtship bahviour and Parenthood
Unit 5 Biological Clocks

REFERENCES AND FURTHER READINGS

You would be required to read the recommended references and

textbooks in each unit of the course materials.

PRESENTATION SCHEDULE

There is a time-table prepared for the early and timely completion and
submissions of your TMAs as well as attending the tutorial classes. You

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BIO 314 COURSE GUIDE

are required to submit all your assignments by the stipulated date and
time. Avoid falling behind the schedule time.

ASSESSMENT

There are three aspects to the assessment of this course.

The first one is the in-text questions and the second is self-assessment
exercises, while the third is the written examination or the examination to
be taken at the end of the course.

Review the exercises or activities in the unit by applying the information

and knowledge you acquired during the course.

The work submitted to your tutor for assessment will account for 30% of
your total work.

At the end of this course you will have to sit for a final or end of course
examination of about a three hour duration and this will account for 70%
of your total course mark.

HOW TO GET THE MOST FROM THE COURSE

In this course, you have the course units and a course guide. The course
guide will tell you briefly what the course is all about. It is a general
overview of the course materials you will be using and how to use those
materials. It also helps you to allocate the appropriate time to each unit so
that you can successfully complete the course within the stipulated time
limit.

The course guide also helps you to know how to go about your in-text
questions and Self-assessment questions which will form part of your
overall assessment at the end of the course. Also, there will be tutorial
classes that are related to this course, where you can interact with your
facilitators and other students. Please I encourage you to attend these
tutorial classes.

This course exposes you to Animal Behaviour, a sub-discipline and very

interesting field of Biology.

ONLINE FACILITATION

Eight weeks are provided for tutorials for this course. You will be notified
of the dates, times and location for these tutorial classes.

v
BIO 314 COURSE GUIDE

As soon as you are allocated a tutorial group, the name and phone number
of your facilitator will be given to you.

The duties of your facilitator are to monitor your progress and provide
any necessary assistance you need.

Do not delay to contact your facilitator by telephone or e-mail for

necessary assistance if

• You do not understand any part of the study in the course material.
• You have difficulty with the self-assessment activities.
• You have a problem or question with an assignment or with the
grading of the assignment.

It is important and necessary you attend the tutorial classes because this
is the only chance to have face to face contact with your facilitator and to
ask questions which will be answered instantly. It is also a period where
you can point out any problem encountered in the course of your study.

COURSE INFORMATION
Course Code:BIO 314
Course Title: ANIMAL BEHAVIOUR
Credit Unit: 2
Course Status: ELECTIVE
Course Blub:
Semester: FIRST SEMESTER
Course Duration: EIGHT WEEKS
Required Hours for Study

ICE BREAKER

Dr. Esenowo, Imeh Kokoete is a Senior Lecturer of Ecology and

Environmental Biology in the Department of Animal and Environmental
Biology, University of Uyo. Dr. Esenowo moderate and facilitate courses
in the National Open University. He has supervised student projects and
seminar review in the Department of Biology, Faculty of Science.

Dr. Esenowo research interests are; physico-chemical aspects of

freshwater and terrestrial ecosystem, fish biology and environmental
toxicology.

vi
 MAIN
COURSE

CONTENTS

Module 1 History of Ethology ……………………………. 1

Unit 1 Historical Background of Ethology……………… 1

Unit 2 Reflex and Complex bahaviour ………………… 13
Unit 3 Orientation in Animals…………………………. 22
Unit 4 Taxes in Animals…………………………….. 31
Unit 5 Fixed action Pattern, Motivation and Drive……. 38

Module 2 Learning, Communication and Social Bahvaiour 50

Unit 1 Display, Displacement Behaviour and Conflict

Behaviour………………………………………….. 50
Unit 2 Learning and Communication in Animal………… 60
Unit 3 Social Behaviour ………………………………….. 72
Unit 4 Social Behaviour of Primates……………………… 80
Unit 5 Hierarchical Organisation……………………….. 88

Module 3 Habitat Selection, Homing and Navigation

and Courtship Behaviour …………………… 98

Unit 1 Physiology of Behaviour……………………… 98

Unit 2 Habitat Selection……………………………… 106
Unit 3 Homing and Navigation in Birds…………….. 114
Unit 4 Courtship bahviour and Parenthood…………. 119
Unit 5 Biological Clocks…………………………… 129
BIO 314 ANIMAL ECOLOGY

Module 1 History of Ethology

Module Introduction

In Module One, deals with the historical aspects of ethology, emotions

in animals, the different forms of communication in animals and how
animal respond to emotions through communication.

Unit 1 Historical Background of Ethology

Unit 2 Reflex and Complex bahaviour
Unit 3 Orientation in Animals
Unit 4 Taxes in Animals
Unit 5 Fixed action Pattern, Motivation and Drive

Glossary

Unit 1 Historical Background of Ethology

Contents
1.1 Introduction
1.2 Intended Learning Outcomes (ILOs)
1.3 Historical Background of Ethology
1.4 Emotion in Animal
1.4.1 Examples of Emotion in Animals
1.5 Animal Communication
1.5.1 Functions of Communication
1.6 Summary
1.7 References/Further Readings/Web Sources
1.8 Possible Answers to Self-Assessment Exercises

1.1 Introduction

Ethology (from Greek: ἦθος, ethos, "character"; and -λογία, -logia, "the
study of") is the scientific study of animal behavior, and a sub-topic
of zoology. Although many naturalists have studied aspects of animal
behavior throughout history, the modern discipline of ethology is
generally considered to have begun during the 1930s with the work of
Dutch biologist Nikolaas Tinbergen and Austrian biologists Konrad
Lorenz and Karl von Frisch, joint winners of the 1973 Nobel Prize in
Physiology or Medicine. Ethology is a combination of laboratory
and field science, with a strong relation to certain other disciplines e.g.,
neuroanatomy, ecology, evolution.

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BIO 314 ANIMAL ECOLOGY

Ethologists are typically interested in a behavioral process rather than

in a particular animal group and often study one type of behavior (e.g.
aggression) in a number of unrelated animals. The desire to understand
animals has made ethology a rapidly growing topic, and since the turn
of the 21st century, many prior understandings related to diverse fields
such as animal communication, personal symbolic name use, animal
emotions, animal culture, learning, and even sexual conduct long thought
to be well understood, have been modified, as have new fields such as
neuroethology

1.2 Intended Learning Outcomes (ILOs)

At the end of unit, students

should be able to:

 Define ethology
 State the Tinbergen’s four questions for ethology.
 Explain emotion in animals under ethology.

1.3 Historical Background of Ethology

The first modern ethologist was Charles Darwin, whose book, The
Expression of the Emotions in Man and Animals, influenced many
ethologists. He pursued his interest in behaviour by encouraging his
protégé George Romanes, who investigated animal learning and
intelligence using an anthropomorphic method, anecdotal cognitivism,
that did not gain scientific support.

The natural history approach of Darwin and his predecessors gradually

evolved into the twin sciences of animal ecology, the study of the
interactions between an animal and its environments and ethology the
biological study of animal behaviour. The roots of ethology can be traced
to the late 19th and early 20th centuries, when scientists from
severalcountries began exploring the behaviours of
selected vertebrate species: dogs by the Russian physiologist Ivan
Pavlov; rodents by American psychologists John B. Watson, Edward
Tolman, and Karl Lashley; birds by American psychologist B. F. Skinner;
and primates by German American psychologist Wolfgang Köhler and
American psychologist Robert Yerkes. The studies were carried out in
laboratories, in the case of dogs, rodents and pigeons, or in artificial
colonies and laboratories, in the case of primates. These studies were
oriented toward psychological and physiological questions rather than
ecological or evolutionary ones.

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BIO 314 ANIMAL ECOLOGY

It was not until the 1930s that field naturalists—such as English

biologist Julian Huxley, Austrian zoologist Konard Lorenz, and Dutch-
born British zoologist and ethologist Nikolaas Tinbergen studying birds
and Austrian zoologist Karl von Frisch and American entomologist
William Morton Wheeler examining insects—gained prominence and
returned to broadly biological studies of animal behaviour. These
individuals, the founders of ethology had direct experience with the
richness of the behavioural repertoires of animals living in their natural
surroundings. Their “return to nature” approach was, to a large extent, a
reaction against the tendency prevalent among psychologists to study just
a few behavioral phenomena observed in a handful of species that were
kept in impoverished laboratory environments.

One of Tinbergen’s most important contributions to the study of animal

behaviour was to stress that ethology is like any other branch of biology,
in that a comprehensive study of any behaviour must address four
categories of questions, which today are called “levels of analysis,”
including causation, ontogeny, function and evolutionary history.
Although each of these four approaches requires a different kind of
scientific investigation, all contribute to solving the enduring puzzle of
how and why animals, including humans, behave as they do. A familiar
example of animal behaviour—a dog wagging its tail—serves to illustrate
the levels of analysis framework. When a dog senses the approach of a
companion (dog or human), it stands still, fixates on the approaching
individual, raises its tail, and begins swishing it from side to side. Why
does this dog wag its tail? To answer this general question, four specific
questions must be addressed.

Both the biological and the physical sciences seek explanations of natural
phenomena in physicochemical terms. The biological sciences (which
include the study of behaviour), however, have an extra dimension
relative to the physical sciences. In biology, physicochemical
explanations are addressed by Tinbergen’s questions on causation and
ontogeny, which taken together are known as “proximate” causes. The
extra dimension of biology seeks explanations of biological phenomena
in terms of function and evolutionary history, which together are known
as “ultimate” causes. In biology, it is legitimate to ask questions
concerning the use of this life process today (its function) and how it came
to be over geologic time (its evolutionary history). More specifically, the
words use and came to be are applied in special ways, namely “promoting
genetic success” and “evolved by means of natural selection.” In physics
and chemistry, these types of questions are out of bounds. For example,
questions concerning the use of the movements of a dog’s tail are
reasonable, whereas questions regarding the use of the movements of an
ocean’s tides are more metaphysical.
What is Ethology?

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BIO 314 ANIMAL ECOLOGY

1.4 Emotion in Animals

There is no scientific consensus on emotion in animals, that is, what

emotions certain species of animals, including humans, feel. Animal
expressions of apparent pleasure are ambiguous as to whether this is
emotion, or simply innate responses, perhaps for approval or other hard-
wired cues. The ambiguity is a source of controversy as there is no
certainty which views, if any, reflect reality. In recent years, research has
become available which expands prior understandings of animal
language, cognition and tool use, and even sexuality. Emotions arise
in the mammalian brain, or the limbic system, which human beings share
in common with other mammals as well as many other species.

1.4.1 Examples of emotion in animal

i. Primates

Primates and in particular great apes are candidates for highly

developed capabilities for empathy and theories of mind. Great apes
have highly complex social systems. Young apes and their mothers
have very strong bonds of attachment. Often when a baby chimpanzee
or gorilla dies, the mother will carry the body around for several days.
Jane Goodall has described chimpanzees as exhibiting mournful
behavior. See notably the example of the gorilla Koko, who expressed
sadness over the death of her pet cat, All Ball.

ii. Fish

A 2007 study by the University of Guelph Scientists in Canada suggests

that fish may have their own separate personalities. The study examined
a group of trout that were visually identical. The study concluded that
different fish within the same group exhibited different personality traits.
Some fish were more willing to take risks in unknown waters than others
when taken from their environment and introduced to a dark tube. Some
fish were more social than others while some fish preferred being alone.
Fish were also shown to have different preferences as far as eating habits.

iii. Felines

The emotions of cats have also been studied scientifically. It has been
shown that cats can learn to manipulate their owners through
vocalizations that are similar to the cries of human babies. Some cats
learn to add a purr to the cry, which makes it less harmonious to humans
and therefore harder to ignore. Individual cats learn to make these cries

4
BIO 314 ANIMAL ECOLOGY

through operant conditioning; when a particular cry elicits a positive

response from a human, the cat is more likely to use that cry in the future.

iv. Canines

Research suggests that canines can experience negative emotions in

a similar manner to people, including the equivalent of certain chronic
and acute psychological conditions. The classic experiment for this was
Martin Seligman's foundational experiments and theory of learned
helplessness at the University of Pennsylvania in 1965, as an extension of
his interest in depression:

A further series of experiments showed that under conditions of long

term intense psychological stress, around 1/3 of dogs do not develop
learned helplessness or long term depression. Instead these animals
somehow managed to find a way to handle the unpleasant situation in
spite of their past experience. The corresponding characteristic in
humans has been found to correlate highly with an explanatory style
and optimistic attitude and lower levels of emotion dog that had
earlier been repeatedly conditioned to associate a sound with electric
shocks did not try to escape the electric shocks after the warning was
presented, even though all the dog would have had to do is jump over a
low divider within ten seconds, more than enough time to respond. The
dog didn't even try to avoid the "aversive stimulus"; it had previously
"learned" that nothing it did mattered. A follow-up experiment involved
three dogs affixed in harnesses, including one that received shocks of
identical intensity and duration to the others, but the lever which would
otherwise have allowed the dog a degree of control was left disconnected
and didn't do anything. The first two dogs quickly recovered from the
experience, but the third dog suffered chronic symptoms of clinical
depression as a result of this perceived helplessness. Such studies
highlighted similar distinctions between people who adapt and those who
break down, under long term psychological pressure, which were
conducted in the 1950s in the realm of brainwashing.

1.5 Animal Communication

Animal communication is any behavior on the part of one animal that

has an effect on the current or future behaviour of another animal. The
study of animal communication, sometimes called Zoosemiotics
(defined as the study of sign communication or semiosis in animals;
distinguishable from anthroposemiotics, the study of human
communication) has played an important part in the methodology of
ethology, sociobiology, and the study of animal cognition. The best

5
BIO 314 ANIMAL ECOLOGY

known form of communication involves the display of distinctive body

parts, or distinctive bodily movements; often these occur in
combination, so a distinctive movement acts to reveal or emphasize
a distinctive body part.

Many animals communicate through vocalization. This is essential for

many tasks including mating rituals, warning calls, conveying location
of food sources, and social learning. Male mating calls are used to signal
the female and to beat competitors in species such as hammer-headed
bats, red deers, humpback whales and elephant seals. In whale species,
whale song has been found to have different dialects based on location.
Other instances of communication include the warning cries of the
Campbell monkey, the territorial calls of gibbons, the use of frequency
in Greater Spear-nosed bats to to distinguish between groups.

Less obvious (except in a few cases) is olfactory communication. Many

mammals, in particular, have glands that generate distinctive and
long-lasting smells, and have corresponding behaviours that leave these
smells in places where they have been. Often the scented substance is
introduced into urine or feces. Sometimes it is distributed through sweat,
though this does not leave a semi-permanent mark as scents deposited
on the ground do. Some animals have glands on their bodies whose
sole function appears to be to deposit scent marks: for example
Mongolian gerbils have a scent gland on their stomachs, and a
characteristic ventral rubbing action that deposits scent from it. Golden
hamsters and cats have scent glands on their flanks, and deposit scent by
rubbing their sides against objects; cats also have scent glands on their
foreheads. Bees carry with them a pouch of material from the hive which
they release as they reenter, the smell of which indicates that they are a
part of the hive and grants their safe entry. Ants use pheromones to
create scent trails to food as well as for alarm calls, mate attraction and
to distinguish between colonies. Additionally, they have pheromones
that are used to confuse an enemy and manipulate them into fighting
with themselves.

A rarer form of animal communication is electrocommunication. It is

seen primarily in aquatic life, though some mammals, notably the
platypus and echidnas are capable of electroreception and thus
theoretically of electrocommunication.

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BIO 314 ANIMAL ECOLOGY

1.5.1 Function of communication

i. Agonistic interaction: everything to do with contests and

aggression between individuals. Many species have distinctive
threat displays that are made during competition over food, mates
or territory; much bird song functions in this way. Often there is
a matched submission display, which the threatened individual
will make if it is acknowledging the social dominance of the
threatener; this has the effect of terminating the aggressive
episode and allowing the dominant animal unrestricted access
to the resource in dispute. Some species also have affiliative
displays which are made to indicate that a dominant animal
accepts the presence of another.

ii. Courtship rituals: signals made by members of one sex to attract

or maintain the attention of potential mate, or to cement a pair
bond. These frequently involve the display of body parts, body
postures (gazelles assume characteristic poses as a signal to
initiate mating), or the emission of scents or calls, that are unique
to the species, thus allowing the individuals to avoid mating with
members of another species which would be infertile. Animals
that form lasting pair bonds often have symmetrical displays that
they make to each other:

iii. Food-related signals: many animals make "food calls" that

attract a mate, or offspring, or members of a social group
generally to a food source. When parents are feeding offspring,
the offspring often have begging responses (particularly when
there are many offspring in a clutch or litter - this is well known
in altricial songbirds, for example). Perhaps the most elaborate
food-related signal is the dance language of honeybees studied.

iv. Alarm calls: signals made in the presence of a threat from a

predator, allowing all members of a social group (and often
members of other species) to run for cover, become immobile, or
gather into a group to reduce the risk of attack.

v. Ownership / territorial: signals used to claim or defend a territory,

food, or a mate.

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BIO 314 ANIMAL ECOLOGY

vi. Metacommunications: signals that modify the meaning of

subsequent signals. The best known example is the play face
in dogs, which signals that a subsequent aggressive signal is
part of a play fight rather than a serious aggressive episode
vii. What is animal communication?
viii. What is Zoosemiotics?

Self-Assessment Exercises 1

Attempt these exercises to measure what you have learnt so far.

This should not take you more than 5 minutes.
1. Briefly explain emotions in cat.
2. Write on courtship rituals

1.6 Summary

One of Tinbergen’s most important contributions to the study of animal

behaviour was to stress that ethology is like any other branch of biology,
in that a comprehensive study of any behaviour must address four
categories of questions, which today are called “levels of analysis,”
including causation, ontogeny, function and evolutionary history.
Although each of these four approaches requires a different kind of
scientific investigation, all contribute to solving the enduring puzzle of
how and why animals, including humans, behave as they do.

The desire to understand animals has made ethology a rapidly growing

topic, and since the turn of the 21st century, many prior understandings
related to diverse fields such as animal communication, personal
symbolic name use, animal emotions, animal culture, learning, and even
sexual conduct long thought to be well understood, have been
modified, as have new fields such as neuroethology

1.7 References/Further Readings/Web Sources

Balcombe, J.P. (1990). Vocal recognition of pups by mother Mexican

free-tailed bats, Tadarida brasiliensis mexicana. Animal
Behaviour 39, 960-966

Berglund, J., Rosenqvist G. and Bernet P. (1997). Ornamentation

8
BIO 314 ANIMAL ECOLOGY

predicts reproductive success in female pipefish. Behavioral

Ecology and Sociobiology 40, 145-150

Clutton-Brock, T., Albon S., Gibson S. & Guinness F. (1979). The logical
stag: Adaptive aspects of fighing in the red deer. Animal
Behaviour 27, 211-225

de Waal F.B.M. (1989). Food sharing and reciprocal obligations among

chimpanzees. Journal of Human Evolution 18, 433–459

Hauser, M. (1997). The Evolution of Communication. Cambridge, MA:

MIT Press.

Lloyd, J.E. (1975). Aggressive mimicry in Photuris: signal repertoires by

femmes fatales. Science 197, 452-453.

Marler, P. (1955). Characteristics of some animal calls. Nature 176, 6-8

Martinez Well, M. and Henry C. S. (1992).The role of courtship songs in

reproductive isolation among populations of green lacewings of
the genus Chrysoperla. Evolution 46, 31-43.

McCracken, G.F. (1984). Communal nursing in Mexican free-tailed bat

maternity colonies. Science 223, 1090-1091.

Rabin, L.A., McCowan B., Hooper S.L and Owings D.H. (2003).
Anthropogenic noise and its effect on animal communication: an
interface between comparative psychology and conservation
biology. International Journal of Comparative Psychology 16,
172-192.

Ryan M.J., Tuttle M.D., and Rand A.S. (1982). Sexual advertisement and
bat predation in a neotropical frog. American Naturalist 119, 136–
139.

Schneider, D. (1974). The sex attractant receptors of moths. Scientific

American 231, 28-35.

Seyfarth, R.M., Cheney D.L. and Marler P. (1980). Monkey responses to

three different alarm calls: Evidence for predator classification and
semantic communication. Science 210, 801-803.

Smith, D. (1972). The role of the epaulets in the red-winged blackbird,

(Agelaius phoeniceus) social system. Behaviour 41, 251-268.

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BIO 314 ANIMAL ECOLOGY

Vehrencamp, S.L., Bradbury J.W., and Gibson R.M. (1989). The

energetic cost of display in male sage grouse. Animal
Behaviour 38, 885-896.

Verbeek, P. (2008) "Peace Ethology." Behaviour 145, 1497-1524

von der Emde, G. (1998). Electroreception. In D. H. Evans (ed.). The

Physiology of Fishes, pp. 313-343. Boca Raton, FL: CRC Press.

Wilson, E.O. (1975). Sociobiology: The New Synthesis.

Cambridge, MA: Harvard University Press.

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BIO 314 ANIMAL ECOLOGY

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BIO 314 ANIMAL ECOLOGY

1.8 Possible Answers to SAEs

Answers to SAEs 1

1. The emotions of cats have also been studied scientifically. It has

been shown that cats can learn to manipulate their owners through
vocalizations that are similar to the cries of human babies. Some
cats learn to add a purr to the cry, which makes it less harmonious
to humans and therefore harder to ignore. Individual cats learn
to make these cries through operant conditioning; when a
particular cry elicits a positive response from a human, the cat is
more likely to use that cry in the future.

2. Courtship rituals are signals made by members of one sex to attract

or maintain the attention of potential mate, or to cement a pair
bond. These frequently involve the display of body parts, body
postures (gazelles assume characteristic poses as a signal to
initiate mating), or the emission of scents or calls, that are unique
to the species, thus allowing the individuals to avoid mating with
members of another species which would be infertile. Animals that
form lasting pair bonds often have symmetrical displays that they
make to each other

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BIO 314 ANIMAL ECOLOGY

Unit 2 Reflex and Complex behaviour

Contents
2.1 Introduction
2.2 Intended Learning Outcomes (ILOs)
2.3 Reflex
2.4 Identified Neuron
2.5 Summary
2.6 References/Further Readings/Web sources
2.7 Possible Answers to Self-Assessment Exercises

2.1 Introduction

The nervous system of vertebrate animals (including humans) is divided

into the central nervous system (CNS) and peripheral nervous system
(PNS); aneurone nervous system.

The central nervous system (CNS) is the largest part, and includes the
brain and spinal cord. The spinal cavity contains the spinal cord, while
the head contains the brain. The CNS is enclosed and protected by
meninges, a three-layered system of membranes, including a tough,
leathery outer layer called the dura mater. The brain is also protected by
the skull, and the spinal cord by the vertebrae.

The peripheral nervous system (PNS) is a collective term for the nervous
system structures that do not lie within the CNS. The large majority of
the axon bundles called nerves are considered to belong to the PNS, even
when the cell bodies of the neurons to which they belong reside within
the brain or spinal cord. The PNS is divided into somatic and visceral
parts. The somatic part consists of the nerves that innervate the skin,
joints, and muscles. The cell bodies of somatic sensory neurons lie in
dorsal root ganglia of the spinal cord. The visceral part, also known as
the autonomic nervous system, contains neurons that innervate the
internal organs, blood vessels, and glands. The autonomic nervous
system itself consists of two parts: the sympathetic nervous system and
the parasympathetic nervous system. Some authors also include sensory
neurons whose cell bodies lie in the periphery (for senses such as
hearing) as part of the PNS; others, however, omit them.

The vertebrate nervous system can also be divided into areas called grey
matter ("gray matter" in American spelling) and white matter. Grey

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BIO 314 ANIMAL ECOLOGY

matter (which is only grey in preserved tissue, and is better described as

pink or light brown in living tissue) contains a high proportion of cell
bodies of neurons. White matter is composed mainly of myelinated
axons, and takes its color from the myelin. White matter includes all of
the peripheral nerves, and much of the interior of the brain and spinal
cord. Grey matter is found in clusters of neurons in the brain and spinal
cord, and in cortical layers that line their surfaces. There is an
anatomical convention that a cluster of neurons in the brain or spinal
cord is called a nucleus, whereas a cluster of neurons in the periphery
is called a ganglion. There are, however, a few exceptions to this rule,
notably including the part of the forebrain called the basal ganglia.

2.2 Intended Learning Outcomes (ILOs)

At the end of this unit, students should be able to;

 Explain the action of reflex in animals
 Explin the term identified neuron

2.3 Reflex

Reflex, in biology is an action consisting of comparatively segments of

behaviour that usually occur as direct and immediate responses to
particular stimuli uniquely correlated wit them.

Many reflexes of placental mammals appear to be innate. They are

hereditary and are a common feature of the species and often of the genus.
Reflexes include not only such simple acts as chewing, swallowing,
blinking, the knee jerk, and the scratch reflex, but also stepping, standing,
and mating. Built up into complex patterns of many coordinated muscular
actions, reflexes form the basis of much instinctive behaviour in animals.

Humans also exhibit a variety of innate reflexes, which are involved with
the adjustment of the musculature for optimum performance of the
distance receptors (i.e., eye and ear), with the orientation of parts of the
body in spatial relation to the head, and with the management of the
complicated acts involved in ingesting food. Among the innate reflexes
involving just the eyes, for example, are:

i) paired shifting of the eyeballs, often combined with turning of the

head, to perceive an object in the field of vision;

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BIO 314 ANIMAL ECOLOGY

ii) contraction of the intraocular muscles to adjust the focus of the

retina for the viewing of near or far objects;
iii) constriction of the pupil to reduce excessive illumination of the
retina; and
iv) blinking due to intense light or touching of the cornea.
In its simplest form, a reflex is viewed as a function of an idealized
mechanism called the reflex arc. The primary components of the reflex
arc are the sensory-nerve cells (or receptors) that receive stimulation, in
turn connecting to other nerve cells that activate muscle cells (or
effectors), which perform the reflex action. In most cases, however, the
basic physiological mechanism behind a reflex is more complicated than
the reflex arc theory would suggest. Additional nerve cells capable of
communicating with other parts of the body (beyond the receptor and
effector) are present in reflex circuits. As a result of the integrative action
of the nervous system in higher organisms, behaviour is more than the
simple sum of their reflexes; it is a unitary whole that exhibits
coordination between many individual reflexes and is characterized by
flexibility and adaptability to circumstances. Many
automatic, unconditioned reflexes can thus be modified by or adapted to
new stimuli, making possible the conditioning of reflex responses. The
experiments of the Russian physiologist Ivan Petrovich Pavlov, for
example, showed that if an animal salivates at the sight of food while
another stimulus, such as the sound of a bell, occurs simultaneously, the
sound alone can induce salivation after several trials. The animal’s
behaviour is no longer limited by fixed, inherited reflex arcs but can be
modified by experience and exposure to an unlimited number of stimuli.

2.4 Identified Neuron

A neuron is called identified if it has properties that distinguish it from

every other neuron in the same animal—properties such as location,
neurotransmitter, gene expression pattern, and connectivity—and if
every individual organism belonging to the same species has one and
only one neuron with the same set of properties. In vertebrate nervous
systems very few neurons are "identified" in this sense—in humans,
there are believed to be none—but in simpler nervous systems, some
or all neurons may be thus unique. In the roundworm C. elegans,
whose nervous system is the most thoroughly described of any animal's,
every neuron in the body is uniquely identifiable, with the same location
and the same connections in every individual worm. One notable
consequence of this fact is that the form of the C. elegans nervous

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BIO 314 ANIMAL ECOLOGY

system is completely specified by the genome, with no experience-

dependent plasticity.

The brains of many molluscs and insects also contain substantial

numbers of identified neurons. In vertebrates, the best known identified
neurons are the gigantic Mauthner cells of fish. Every fish has two
Mauthner cells, located in the bottom part of the brainstem, one on the
left side and one on the right. Each Mauthner cell has an axon that
crosses over, innervating neurons at the same brain level and then
travelling down through the spinal cord, making numerous connections
as it goes. The synapses generated by a Mauthner cell are so powerful
that a single action potential gives rise to a major behavioral
response: within milliseconds the fish curves its body into a C-
shape, then straightens, thereby propelling itself rapidly forward.
Functionally this is a fast escape response, triggered most easily by a
strong sound wave or pressure wave impinging on the lateral line organ
of the fish. Mauther cells are not the only identified neurons in fish—
there are about 20 more types, including pairs of "Mauthner cell analogs"
in each spinal segmental nucleus. Although a Mauthner cell is capable
of bringing about an escape response all by itself, in the context of
ordinary behavior other types of cells usually contribute to shaping
the amplitude and direction of the response. Mauthner cells have been
described as command neurons. A command neuron is a special type of
identified neuron, defined as a neuron that is capable of driving a specific
behavior all by itself. such neurons appear most commonly in the fast
escape systems of various species—the squid giant axon and squid
giant synapse, used for pioneering experiments in neurophysiology
because of their enormous size, both participate in the fast escape circuit
of the squid. The concept of a command neuron has however, become
controversial, because of studies showing that some neurons that
initially appeared to fit the description were really only capable of
evoking a response in a limited set of circumstances

i. Neural precursors in sponges

Sponges have no cells connected to each other by synaptic junctions,

that is, no neurons, and therefore no nervous system. They do, however,
have homologs of many genes that play key roles in synaptic function.
Recent studies have shown that sponge cells express a group of
proteins that cluster together to form a structure resembling a
postsynaptic density (the signal-receiving part of a synapse). However,

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BIO 314 ANIMAL ECOLOGY

the function of this structure is currently unclear. Although sponge cells

do not show synaptic transmission, they do communicate with each
other via calcium waves and other impulses, which mediate some simple
actions such as whole-body contraction.

ii. Radiate

Jellyfish, comb jellies, and related animals have diffuse nerve nets rather
than a central nervous system. In most jellyfish the nerve net is
spread more or less evenly across the body; in comb jellies it is
concentrated near the mouth. The nerve nets consist of sensory neurons
that pick up chemical, tactile, and visual signals, motor neurons that can
activate contractions of the body wall, and intermediate neurons that
detect patterns of activity in the sensory neurons and send signals to
groups of motor neurons as a result. In some cases groups of intermediate
neurons are clustered into discrete ganglia.

iii. Bilateria

The vast majority of existing animals are bilaterians, meaning animals

with left and right sides that are approximate mirror images of each
other. All bilateria are thought to have descended from a common
wormlike ancestor that appeared in the Cambrian period, 550–600
million years ago. The fundamental bilaterian body form is a tube with
a hollow gut cavity running from mouth to anus, and a nerve cord with
an enlargement (a "ganglion") for each body segment, with an especially
large ganglion at the front, called the "brain". Even mammals, including
humans, show the segmented bilaterian body plan at the level of the
nervous system. The spinal cord contains a series of segmental ganglia,
each giving rise to motor and sensory nerves that innervate a portion of
the body surface and underlying musculature. On the limbs, the layout
of the innervation pattern is complex, but on the trunk it gives rise to a
series of narrow bands. The top three segments belong to the brain,
giving rise to the forebrain, midbrain, and hindbrain.

Bilaterians can be divided, based on events that occur very early in

embryonic development, into two groups (superphyla) called
protostomes and deuterostomes. Deuterostomes include vertebrates as
well as echinoderms, hemichordates (mainly acorn worms), and
Xenoturbellidans. Protostomes, the more diverse group, include
arthropods, molluscs, and numerous types of worms. There is a

17
BIO 314 ANIMAL ECOLOGY

basic difference between the two groups in the placement of the

nervous system within the body: protostomes possess a nerve cord on
the ventral (usually bottom) side of the body, whereas in deuterostomes
the nerve cord is on the dorsal (usually top) side. In fact, numerous
aspects of the body are inverted between the two groups, including the
expression patterns of several genes that show dorsal-to-ventral
gradients. Most anatomists now consider that the bodies of protostomes
and deuterostomes are "flipped over" with respect to each other, a
hypothesis that was first proposed by Geoffroy Saint-Hilaire for insects
in comparison to vertebrates. Thus insects, for example, have nerve
cords that run along the ventral midline of the body, while all
vertebrates have spinal cords that run along the dorsal midline.
iv. Worms

Worms are the simplest bilaterian animals, and reveal the basic structure
of the bilaterian nervous system in the most straightforward way. As an
example, earthworms have dual nerve cords running along the length
of the body and merging at the tail and the mouth. These nerve cords
are connected by transverse nerves like the rungs of a ladder. These
transverse nerves help coordinate the two sides of the animal. Two
ganglia at the head end function similar to a simple brain.

Photoreceptors on the animal's eyespots provide sensory information on

light and dark. The nervous system of one very small worm, the
roundworm Caenorhabditis elegans, has been mapped out down to the
synaptic level. Every neuron and its cellular lineage has been recorded
and most, if not all, of the neural connections are known. In this species,
the nervous system is sexually dimorphic; the nervous systems of the two
sexes, males and hermaphrodites, have different numbers of neurons and
groups of neurons that perform sex-specific functions. In C. elegans,
males have exactly 383 neurons, while hermaphrodites have exactly 302
neurons.
v. Arthropods

Arthropods, such as insects and crustaceans, have a nervous system

made up of a series of ganglia, connected by a ventral nerve cord
made up of two parallel connectives running along the length of the
belly. Typically, each body segment has one ganglion on each side,
though some ganglia are fused to form the brain and other large ganglia.
The head segment contains the brain, also known as the supraesophageal
ganglion. In the insect nervous system, the brain is anatomically divided
into the protocerebrum, deutocerebrum, and tritocerebrum.
Immediately behind the brain is the subesophageal ganglion, which is

18
BIO 314 ANIMAL ECOLOGY

composed of three pairs of fused ganglia. It controls the mouthparts,

the salivary glands and certain muscles. Many arthropods have
well-developed sensory organs, including compound eyes for vision
and antennae for olfaction and pheromone sensation. The sensory
information from these organs is processed by the brain. What is Reflex?
The best known identified neurons are found inwhere?

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is identified neurons?
2. Write on neurons in radiate

2.5 Summary

A reflex is viewed as a function of an idealized mechanism called

the reflex arc. The primary components of the reflex arc are the sensory-
nerve cells (or receptors) that receive stimulation, in turn connecting to
other nerve cells that activate muscle cells (or effectors), which perform
the reflex action. Animal’s behaviour is no longer limited by fixed,
inherited reflex arcs but can be modified by experience and exposure to
an unlimited number of stimuli.

2.6 References/Further Readings/Web Sources

Dawkins, R. & Krebs, J. R. (1978). Animal Signals: Information or

Manipulation. In Krebs, J. R. & Davies, N. B. (Eds.), Behavioural
Ecology: An Evolutionary Approach (pp. 282–
309). Oxford: [Link] Scholar

Kandel E.R, Schwartz J.H, Jessel T.M, ed (2000). "Ch. 2: Nerve

cells and behavior". Principles of Neural Science.
McGraw-Hill Professional.

Ruppert EE, Fox RS, Barnes RD (2004). Invertebrate Zoology (7 ed.).

Brooks / Cole. pp.

[Link]
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BIO 314 ANIMAL ECOLOGY

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[Link]

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2.7 Possible Answers to SAEs

Answers to SAEs 1

1. A neuron is called identified if it has properties that distinguish it

from every other neuron in the same animal—properties such as
location, neurotransmitter, gene expression pattern, and
connectivity—and if every individual organism belonging to the
same species has one and only one neuron with the same set of
properties

2. Jellyfish, comb jellies, and related animals have diffuse nerve nets
rather than a central nervous system. In most jellyfish the nerve
net is spread more or less evenly across the body; in comb jellies
it is concentrated near the mouth. The nerve nets consist of sensory
neurons that pick up chemical, tactile, and visual signals, motor
neurons that can activate contractions of the body wall, and
intermediate neurons that detect patterns of activity in the sensory
neurons and send signals to groups of motor neurons as a result.
In some cases groups of intermediate neurons are clustered into
discrete ganglia.

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BIO 314 ANIMAL ECOLOGY

Unit 3 Orientation in Animals

Contents

3.1 Introduction
3.2 Intended Learning Outcomes (ILOs)
3.3 Orientation
3.3.1 Steering
3.4 Summary
3.5 References/Further Readings/Web sources
3.6 Possible Answers to Self-Assessment Exercises

3.1 Introduction

Orientation is the ability of an animal to determine its position in space

and among individuals of the same or other species. Animal orientation is
complex process that includes receiving information about the external
world through various channels of communication, processing it,
correlating it in the central nervous system, and forming a response. The
reception and processing of signals consist of recognizing and locating an
imagine (information content of the signal) that is determining by various
receptor system the source of the signal in relation to the body
(biolocation).

There are diiferent forms of orientation. Object orientation takes place

when the animal tries to approach an object which may be food or water.
Aquatic animals move vertically in pond or lake which is called strato-
orientation. When the animals try to move from grassland to forests,
deserts or mountains it is called zonal orientation. Animals which
migrate long distances generally possess topographical or geographical
orientation.

3.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Define orientation in animal
 Explain the different forms of orientation

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BIO 314 ANIMAL ECOLOGY

3.3 Orientation in Animal

Orientation of locomotor behaviour is usually categorized as

either kinesis or taxis. In kinesis, an animal’s body is not oriented in
relation to a sensory stimulus; rather, the stimulus causes an alteration in
speed or direction of movement. In wood lice, for example, the kinetic
response alters only the rate of movement. Because wood lice tend
to aggregate in moist areas, their ambulatory activity increases or
decreases as the relative humidity decreases or increases, respectively. In
the planarian (an aquatic, ciliated flatworm), on the other hand, the kinetic
response affects only the rate at which the planarian changes its direction.
Because planaria tend to stay in or return to darker areas, an increase
in light intensity causes an increase in their turning responses. Generally,
however, animals tend to alter both direction and speed as a single kinetic
response.

In taxis, an animal orients itself in a specific spatial relationship to a

stimulus. The orientation may be simply an alteration of body position or
it may be an alteration of locomotor direction so that the animal moves
toward, away from, or at a fixed angle to the source of the stimulus.
Sources that elicit a taxis response, which may cause a modification of
speed, direction, or both, seem to encompass the entire range of
environmental stimuli, such as gravity (geotaxis), temperature
(thermotaxis), light (phototaxis), or chemicals (chemotaxis). If the
response is negative, the animal moves away from the source; if it is
positive, the animal moves toward the source.

The control of the response to a taxis is of two types. In open-system

control, the initial response to a stimulus has no effect on subsequent
responses to the same stimulus. A male firefly, for example, locates a
female by the latter’s brief flashes of light. When a male sees a female’s
flash, the male turns in the direction of the female, even though the source
is no longer visible. If another female flashes, however, the male responds
to the second flash in exactly the same manner as it did to the first. In
close-system control, on the other hand, the response is progressively
altered by feedback so that all subsequent responses are adjusted to the
initial response. A bat chasing a flying insect will alter its flight path to
intercept that of the insect. The bat’s initial change in direction is only a
general alteration of its course, but, as it approaches the insect, the bat
constantly modifies its course to obtain an accurate interception.

Experiments have shown that the orientation of birds is based on celestial

bearings. The Sun is the point of orientation during the day, and birds are
able to compensate for the movement of the Sun throughout the day. A

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BIO 314 ANIMAL ECOLOGY

so-called internal clock mechanism in birds involves the ability to gauge

the angle of the Sun above the horizon. Similar mechanisms are known in
many animals and are closely related to the rhythm of daylight,
or photoperiodism. When the internal rhythm of birds is disturbed by
subjecting them first to several days of irregular light–dark sequences,
then to an artificial rhythm that is delayed or advanced in relation to the
normal rhythm, corresponding anomalies occur in the homing behaviour.

Two theories have been formulated to explain how birds use the Sun for
orientation. Neither, however, has so far been substantiated with proof.
One theory holds that birds find the right direction by determining the
horizontal angle measured on the horizon from the Sun’s projection. They
correct for the Sun’s movement by compensating for the changing angle
and thus are able to maintain the same direction. According to this theory,
the Sun is a compass that enables the birds to find and maintain their
direction. This theory does not explain, however, the manner in which a
bird, transported and released in an experimental situation, determines the
relationship between the point at which it is released and its goal.

The second theory, proposed by British ornithologist G.V.T. Matthews,

is based on other aspects of the Sun’s position, the most important of
which is the arc of the Sun—i.e., the angle made by the plane through
which the Sun is moving in relation to the horizontal. Each day in the
Northern Hemisphere, the highest point reached by the Sun lies in the
south, thus indicating direction; the highest point is reached at noon, thus
indicating time. In its native area a bird is familiar with
the characteristics of the Sun’s movement. Placed in different
surroundings, the bird can project the curve of the Sun’s movement after
watching only a small segment of its course. By measuring maximum
altitude (the Sun’s angle in relation to the horizontal) and comparing it
with circumstances in the usual habitat, the bird obtains a sense of latitude.
Details of longitude are provided by the Sun’s position in relation to both
the highest point and position it will reach—as revealed by a precise
internal clock.

Migrant birds that travel at night are also capable of directional

orientation. Studies have shown that these birds use the stars to determine
their bearings. In clear weather, captive migrants head immediately in the
right direction using only the stars. They are even able to orient
themselves correctly to the arrangement of night skies projected on the
dome of a planetarium; true celestial navigation is involved because the
birds determine their latitude and longitude by the position of the stars. In
a planetarium in Germany, blackcaps (Sylvia atricapilla) and garden
warblers (S. borin), under an artificial autumn sky, headed “southwest,”
their normal direction; lesser whitethroats (S. curruca) headed
“southeast,” their normal direction of migration in that season.

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BIO 314 ANIMAL ECOLOGY

It is known, then, that birds are able to navigate by two types of

orientation. One, simple and directional, is compass orientation; the
second, complex and directed to a point, is true navigation, or goal
orientation. Both types apparently are based on celestial bearings, which
provide a navigational “grid.”

The methods of directional orientation used by birds are similar to those

used by other animals. Orientation to the Sun has been demonstrated in
various crustaceans, particularly in the sand flea (Talitrus saltator).
Various insects, particularly bees and certain beetles
(families Scarabaeidae, Tenebrionidae, and Carabidae), use the Sun to
plot their course with remarkable accuracy.

Fishes also are able to use celestial bearings; salmon presumably use the
Sun. Experiments with the parrot fish (Scarus) have demonstrated a Sun
compass reaction that may also occur in other fishes. Localization of the
Sun is, however, much more difficult in water than in the air, because of
the characteristics of light rays passing through water. Experiments
suggest that topographical clues are also used by fishes to recognize their
range, particularly their spawning grounds. Visual bearings in this respect
have great importance. It is possible that chemical substances also provide
clues.

Visible landmarks are used by mammals, at least for orientation within

short distances. Scented trails are probably helpful within a limited area,
proportionate to the size of the animal; olfaction plays an important role
in the life of mammals. Some mammals, however, migrate over enormous
distances and are able to return after being taken far away from their home
territory; bats, for example, have returned 265 kilometres (165 miles) to
their caves. Random exploration plays a part in such movements, but it is
possible that some type of true navigation is involved in certain of these
movements.

3.3.1 Steering

Animals obtain accurate directional response (steering) by changing their

propulsive response. Because steering relies heavily on continuous
feedback (the communication cycle in which the motor output, or
behaviour, is constantly being modified by the sensory input, or stimulus),
it requires a precise integration of the central and peripheral nervous
systems. (The central nervous system—in vertebrates, the brain and spinal
cord—is that part of the nervous system that receives sensory impulses
and sends out motor impulses; the peripheral nervous system consists of
all the nerves that carry impulses between the central nervous system and
other parts of the body.) Exteroceptive stimuli (those that originate

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BIO 314 ANIMAL ECOLOGY

outside the body) received by the peripheral nervous system establish the
animal’s spatial position in the environment; proprioceptive stimuli (those
that originate inside the body), also received by the peripheral nervous
system, establish the relative position of the body units to each other.
Through integration of these two sets of stimuli, the central nervous
system continuously adjusts the contraction of the motor units (e.g.,
muscles) to obtain the desired orientation.

During locomotion, steering is a continual process. The direction of

movements must be constantly adjusted to counteract environmentally
produced deviations of direction. The apparently simple act of
a bird flying from a tree to the ground illustrates the complexity of
directional control. As the bird flies to the ground, it must be constantly
aware of its height above the ground, the orientation of its body axis
relative to the ground, deviations in flight direction resulting from air
currents, and its speed of fall. All these parameters are determined
primarily by exteroceptive stimuli received through the eyes and inner
ears. The downward flight is constantly adjusted in response to these
exteroceptive stimuli, and the fine control necessary for these adjustments
is obtained by proprioceptive feedback. What is Orientation?
Birds traveling of flying at night are capable of what?

Self-Assessment Exercises 1

Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. Birds are able to navigate with how many types of
orientation?

3.4 Summary

Orientation is the position of the animal with reference to gravity or

resource. This is the position the animal maintains in order to reach the
resource. The ability of invertebrate and especially vertebrate to see
individual objects makes orientation in an enviroenmnet much easier.
This is particularly important for every mobile animals. Invertebrates
and lower vertebrate are not capable of a detailed and coplex analysis
of the visble world. They discern only a few biologically important
signals against an undifferentiated background.

26
BIO 314 ANIMAL ECOLOGY

3.5 References/Further Readings/ Web Sources

Blass, E. M (1987). "Opioids, sweets and a mechanism for positive

affect: Broad motivational implications". In Dobbing, J.
Sweetness. London: Springer-Verlag. pp. 115–124. ISBN 0-387-
17045-6

Dusenbery, D. B. (1988) Behavioral Ecology and Sociobiology,

22:219-223 (1988). Avoided temperature leads to the surface

Dusenbery, D. B. (1992). Sensory Ecology, p.114. W.H. Freeman, New

York. ISBN 0-7167-2333-6.

Dusenbery, D.B. ( 1 9 8 9 ) Biological Cybernetics, 60:431-437

(1989). A simple animal can use a complex stimulus patter to
find a location:

Dusenbery, D. B. (2009). Living at Micro Scale, pp.164-167.

Harvard
University Press, Cambridge, Mass. ISBN 978-0-674-03116-6

Kendeigh, S. C. (1961). Animal Ecology. Prentice-Hall, Inc., Englewood

Cliffs, N.J.. pp. 468 pp.

Martin, E. A., ed (1983). Macmillan Dictionary of Life Sciences

(2nd ed.). London: Macmillan Press. p. 362. ISBN 0-333-34867-
2

Menzel, Randolf (1979). "Spectral Sensitivity and Color Vision in

Invertebrates". In H. Autrum (editor). Comparative Physiology
and Evolution of Vision in Invertebrates- A: Invertebrate
Photoreceptors. Handbook of Sensory Physiology VII/6A New
York: Springer-Verlag. pp. 503–580. See section D:
Wavelength- Specific Behavior and Color Vision. ISBN
3540088377.
[Link]
TY5NDMwNDAwMCZpZ3VpZD0xM2EwYmJmYS05Njk3L
TY0MDUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9
NTIwNA&ptn=3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
a992978a655d&psq=list+of+related+materials++in+Orientati
on+and+Taxes+in+animal&u=a1aHR0cHM6Ly9hY2FkZW1

27
BIO 314 ANIMAL ECOLOGY

pYy5vdXAuY29tL2ljYi9hcnRpY2xlLzMxLzEvMTU3LzE1M
Tg1Mg&ntb=1

[Link]
Y5NDMwNDAwMCZpZ3VpZD0xM2EwYmJmYS05Njk3LTY0MD
UtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTM1OA&ptn
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oYXZpb3Jz&ntb=1

[Link]
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3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
a992978a655d&psq=list+of+related+materials++in+Orientation+an
d+Taxes+in+animal&u=a1aHR0cHM6Ly93d3cuaWFzem9vbG9neS
5jb20vb3JpZW50YXRpb24v&ntb=1

[Link]
on+and+Taxes+in+animals&&view=detail&mid=E79A14CF7E755F
1E32BBE79A14CF7E755F1E32BB&&FORM=VRDGAR&ru=%2F
videos%2Fsearch%3Fq%3Dlectures%2520video%2520of%2520Or
ientation%2520and%2520Taxes%2520in%2520animals%26qs%3D
n%26form%3DQBVR%26%3D%2525eManage%2520Your%2520
Search%2520History%2525E%26sp%3D-
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50%26sk%3D%26cvid%3DE0F48087BC294CE7B627EA8AB3E7E
693%26ghsh%3D0%26ghacc%3D0%26ghpl%3D

[Link]
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UtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTIwNg&ptn=
3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
a992978a655d&psq=lectures+video+of+orientation+and+taxes+in+a
nimals&u=a1aHR0cHM6Ly93d3cueW91dHViZS5jb20vd2F0Y2g_dj
1ldUl3TVBtblk2QQ&ntb=1

[Link]
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DUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTMwMw&
ptn=3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-

28
BIO 314 ANIMAL ECOLOGY

a992978a655d&psq=lectures+video+of+orientation+and+taxes+in+a
nimals&u=a1aHR0cHM6Ly93d3cueW91dHViZS5jb20vd2F0Y2g_dj
1ZRVhpR1ZKLTBkSQ&ntb=1

29
BIO 314 ANIMAL ECOLOGY

3.6 Possible Answers to SAEs

Answers to SAEs 1

1. Birds are able to navigate by two types of orientation. One, simple

and directional, is compass orientation; the second, complex and
directed to a point, is true navigation, or goal orientation

30
BIO 314 ANIMAL ECOLOGY

Unit 4 Taxes in Animals

Contents
4.1 Introduction
4.2 Intended Learning Outcomes (ILOs)
4.3 Taxes
4 .3.1 Examples of Taxes in Animals
4.4 Summary
4.5 References/Further Readings/Web sources
4.6 Possible Answers to Self-Assessment Exercises

4.1 Introduction

A taxis (plural taxes, pronounced / tæksi z/) is an innate behavioral

response by an organism to a directional stimulus or gradient of
stimulus intensity. A taxis differs from a tropism (turning response,
often growth towards or away from a stimulus) in that the organism has
motility and demonstrates guided movement towards or away from the
stimulus source. It is sometimes distinguished from a kinesis, a non-
directional change in activity in response to a stimulus.

4.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Define taxes in animal

 Explain the different forms of taxes in animals

4.3 Taxes

4.3.1 Examples of Taxes in Animals

i. Aerotaxis

Aerotaxis is the response of an organism to variation in oxygen

concentration, and is mainly found in aerobic bacteria.

ii. Chemotaxis

Chemotaxis is a migratory response elicited by chemicals: that is, a

response to a chemical concentration gradient. For example, chemotaxis
in response to a sugar gradient has been observed in motile bacteria such
31
BIO 314 ANIMAL ECOLOGY

as E. Coli. Chemotaxis also occurs in the antherozoids of liverworts,

ferns, and mosses in response to chemicals secreted by the archegonia,
also in higher animals e.g Dogs for sexual attraction. Unicellular (e.g.
protozoa) or multicellular (e.g. worms) organisms are targets of
chemotactic substances. A concentration gradient of chemicals
developed in a fluid phase guides the vectorial movement of responder
cells or organisms. Inducers of locomotion towards increasing steps of
concentrations are considered as chemoattractants, while
chemorepellents result moving off the chemical. Chemotaxis is
described in prokaryotic and eukaryotic cells, but signalling mechanisms
(receptors, intracellular signaling) and effectors are significantly
different.

iii. Energy taxis

Energy taxis is the orientation of bacteria towards conditions of optimal

metabolic activity by sensing the internal energetic conditions of cell.
Therefore in contrast to chemotaxis (taxis towards or away from a
specific extracellular compound), energy taxis responds on an
intracellular stimulus (e.g. proton motive force, activity of NDPH- 1)
and requires metabolic activity.

iv. Phototaxis

Phototaxis is the movement of an organism in response to light:

that is, the response to variation in light intensity and direction.
Negative phototaxis, or movement away from a light source, is
demonstrated in some insects, such as cockroaches.

Positive phototaxis, or movement towards a light source, is

advantageous for phototrophic organisms as they can orient themselves
most efficiently to receive light for photosynthesis. Many
phytoflagellates, e.g. Euglena, and the chloroplasts of higher plants
positively phototactic, moving towards a light source. Two types of
positive phototaxis are observed in prokaryotes.

Scotophototaxis is observable as the movement of a bacterium out

of the area illuminated by a microscope. Entering darkness signals the
cell to reverse direction and reenter the light. A second type of positive
phototaxis is true phototaxis, which is a directed movement up a
gradient to an increasing amount of light.

v. Thermotaxis

Thermotaxis is a migration along a gradient of temperature. Some

slime molds and small nematodes can migrate along amazingly small

32
BIO 314 ANIMAL ECOLOGY

temperature gradients of less than 0.1C/cm. They apparently use this

behavior to move to an optimal level in soil.

vi. Geotaxis

Geotaxis is a response to the attraction due to gravity. The planktonic

larvae of the king crab Lithodes aequispinus use a combination
of positive phototaxis (movement towards the light) and negative
geotaxis (upward movement). Both positive and negative geotaxes are
found in a variety of protozoans.

vii. Rheotaxis

Rheotaxis is a response to a current in a fluid. Positive rheotaxis is shown

by fish turning to face against the current. In a flowing stream, this
behavior leads them to hold their position in a stream rather than being
swept downstream. Some fish will exhibit negative rheotaxis where they
will avoid currents.

viii. Magnetotaxis

Magnetotaxis is the ability to sense a magnetic field and coordinate

movement in response. However, the term is commonly applied to
bacteria that contain magnets and are physically rotated by the force
of the Earth's magnetic field. In this case, the "behavior" has nothing
to do with sensation, and the bacteria are more accurately described as
"magnetic bacteria".

ix. Phonotaxis

Phonotaxis is the movement of an organism in response to sound

x. Galvanotaxis / electrotaxis

Galvanotaxis or electrotaxis is directional movement of motile cells in

response to an electric field. It has been suggested that by detecting
and orientating themselves toward the electric fields, cells are able
to direct their movement towards the damages or wounds to repair the
defect. It also is suggested that such a movement may contribute to
directional growth of cells and tissues during development and
regeneration. This notion is based on

a. the existence of measurable electric fields that naturally occur

during wound healing, development and regeneration; and
b. cells in cultures respond to applied electric fields by
directional cell migration – electrotaxis / galvanotaxis.
33
BIO 314 ANIMAL ECOLOGY

What is Taxis? What is Aerotaxis

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1 Write on Energy taxis
2. What is Scotophototaxis?

4.4 Summary

A taxis differs from a tropism (turning response, often growth towards

or away from a stimulus) in that the organism has motility and
demonstrates guided movement towards or away from the stimulus
source. It is sometimes distinguished from a kinesis, a non-directional
change in activity in response to a stimulus.

4.5 References/Further Readings/ Web Sources

Adams, C. F. and Paul, A. J. (1999). "Phototaxis and geotaxis of light-

adapted zoeae of the golden king crab Lithodes aequispinus
(Anomura: Lithodidae) in the laboratory". Journal of Crustacean
Biology 19 (1): 106–110. doi:10.2307/1549552.
[Link]

Blass, E. M (1987). "Opioids, sweets and a mechanism for positive

affect: Broad motivational implications". In Dobbing, J.
Sweetness. London: Springer-Verlag. pp. 115–124. ISBN 0-387-
17045-6

Dusenbery, D. B. (1988) Behavioral Ecology and Sociobiology,

22:219-223 (1988). Avoided temperature leads to the surface

Dusenbery, D. B. (1992). Sensory Ecology, p.114. W.H. Freeman, New

York. ISBN 0-7167-2333-6.

Dusenbery, D. B. (2009). Living at Micro Scale, pp.164-167.

Harvard
University Press, Cambridge, Mass. ISBN 978-0-674-03116-6

34
BIO 314 ANIMAL ECOLOGY

Fenchel, T. and Finlay, B. J. (1 May 1984). "Geotaxis in the ciliated

protozoon Loxodes". Journal of Experimental Biology 110 (1):
110–133. [Link]

Kendeigh, S. C. (1961). Animal Ecology. Prentice-Hall, Inc., Englewood

Cliffs, N.J.. pp. 468 pp.

Martin, E. A., ed (1983). Macmillan Dictionary of Life Sciences

(2nd ed.). London: Macmillan Press. p. 362. ISBN 0-333-34867-
2

Schweinitzer T, Josenhans C. ( 2 0 1 0 ) Bacterial energy taxis: a

global strategy? Arch Microbiol. 2010 Jul;192(7):507-20.
[Link]
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TY0MDUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9
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pYy5vdXAuY29tL2ljYi9hcnRpY2xlLzMxLzEvMTU3LzE1M
Tg1Mg&ntb=1

[Link]
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UtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTIyOA&ptn=
3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
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d+Taxes+in+animal&u=a1aHR0cHM6Ly93d3cuaWFzem9vbG9neS
5jb20vb3JpZW50YXRpb24v&ntb=1

[Link]
on+and+Taxes+in+animals&&view=detail&mid=E79A14CF7E755F
1E32BBE79A14CF7E755F1E32BB&&FORM=VRDGAR&ru=%2F
videos%2Fsearch%3Fq%3Dlectures%2520video%2520of%2520Or
ientation%2520and%2520Taxes%2520in%2520animals%26qs%3D

35
BIO 314 ANIMAL ECOLOGY

n%26form%3DQBVR%26%3D%2525eManage%2520Your%2520
Search%2520History%2525E%26sp%3D-
1%26lq%3D0%26pq%3Dlectures%2520video%2520of%2520orien
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693%26ghsh%3D0%26ghacc%3D0%26ghpl%3D

[Link]
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UtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTIwNg&ptn=
3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
a992978a655d&psq=lectures+video+of+orientation+and+taxes+in+a
nimals&u=a1aHR0cHM6Ly93d3cueW91dHViZS5jb20vd2F0Y2g_dj
1ldUl3TVBtblk2QQ&ntb=1

[Link]
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DUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTMwMw&
ptn=3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
a992978a655d&psq=lectures+video+of+orientation+and+taxes+in+a
nimals&u=a1aHR0cHM6Ly93d3cueW91dHViZS5jb20vd2F0Y2g_dj
1ZRVhpR1ZKLTBkSQ&ntb=1

36
BIO 314 ANIMAL ECOLOGY

4.6 Possible Answers to SAEs

Answers to SAEs 1

2. Energy taxis is the orientation of bacteria towards conditions of

optimal metabolic activity by sensing the internal energetic
conditions of cell. Therefore in contrast to chemotaxis (taxis
towards or away from a specific extracellular compound), energy
taxis responds on an intracellular stimulus (e.g. proton motive
force, activity of NDPH- 1) and requires metabolic activity.

3. Scotophototaxis is observable as the movement of a bacterium

out of the area illuminated by a microscope

37
BIO 314 ANIMAL ECOLOGY

Unit 5 Fixed Action Pattern, Motivation and Drive

Contents
5.1 Introduction
5.2 Intended Learning Outcomes (ILOs)
5.3 Fixed Action Pattern
5.4 Motivation and Drive
5.4.1 Drive or Urge in Animals
5.5 Summary
5.6 References/Further Readings/Web sources
5.7 Possible Answers to Self-Assessment Exercises

Glossary

5.1 Introduction

In ethology, a Fixed Action Pattern (FAP) is an instinctive behavioral

sequence that is indivisible and runs to completion. Fixed action patterns
are invariant and are produced by a neural network known as the
innate releasing mechanism in response to an external sensory
stimulus known as a sign stimulus or releaser (a signal from one
individual to another). A fixed action pattern is one of the few types of
behaviors which can be said to be hard-wired and instinctive

5.2 Intended Learning Outcomes (ILOs)

At the end of this topic students should be able to;

 Explain Fxed Action Pattern

 Define motivation
 Explain theories types of motivation and emotion.

5.3 Fixed Action Pattern

Many mating dances, commonly carried out by birds, are examples of

fixed action patterns. In these cases, the sign stimulus is typically the
presence of the female. Another example of fixed action patterns is
aggression towards other males during mating season in the red-bellied
stickleback. A series of experiments carried out by Niko Tinbergen
showed that the aggressive behavior of the males is a FAP triggered

38
BIO 314 ANIMAL ECOLOGY

by anything red, the sign stimulus. The threat display of male stickleback
is also a fixed action pattern triggered by a stimulus.

Another well-known case are the classic experiments by Tinbergen and

Lorenz on the Graylag Goose. Like similar waterfowl, it will roll a
displaced egg near its nest back to the others with its beak. The sight of
the displaced egg triggers this mechanism. If the egg is taken away,
the animal continues with the behavior, pulling its head back as if
an imaginary egg is still being maneuvered by the underside of its
beak. However, it will also attempt to move other egg shaped objects,
such as a golf ball, door knob, or even an egg too large to have possibly
been laid by the goose itself (a supernormal stimulus). Kelp Gull chicks
are stimulated by a red spot on the mother's beak to peck at spot, which
induces regurgitation. Some moths instantly fold their wings and drop to
the ground if they encounters ultrasonic signals such as those produced
by bats; see ultrasound avoidance. Mayflies drop their eggs when they
encounter a certain pattern of light polarization which indicates they
are over water.

i. Important/Significant of Fixed Action Pattern

A Fixed Action Pattern (FAP) is significant in animal behavior because

it represents the simplest type of behavior, in which a readily defined
stimulus nearly always results in an invariable behavioral response. A
FAP can truly be said to be "hard-wired." FAPs are also unusual in that
almost behaviors are modulated by the environment; a fixed response
can lead to maladaptive results, whereas flexible behaviors are generally
more useful. Because of this, most behaviors which are both FAPs and
occur in more complex animals are usually essential to the animal's
fitness or in which speed is a factor. For instance, the Greylag Goose's
egg rolling behavior is so essential to the survival of its chicks that its
fitness is increased by this behavior being hard-wired. A chick which
cannot consistently feed will die. A moth's response to encountering
echolocation needs to be immediate in order to avoid predation. An
attacking stickleback is placed at an advantage if it reacts quickly to a
threat. However, because these behaviors are hard-wired, they are also
predictable. This can lead to their exploitation by humans or other
animals.

ii. Exploitation of Fixed Action Pattern

Some species have evolved to exploit the fixed action patterns of other
species by mimicry of their sign stimulus. Replicating the releasing
mechanism required to trigger a FAP is known as code-breaking. A well-
known example of this is brood parasitism, where one species will lay its
eggs in the nest of another species, which will then parent its young. A
young North American cowbird, for example, provides a supernormal

39
BIO 314 ANIMAL ECOLOGY

stimulus to its foster parent, which will cause it to forage rapidly in

order to satisfy the larger bird's demands. In a natural situation a
nestling will provide higher levels of stimulus with noisier, more
energetic behavior, communicating its urgent need for food. Parents in
this situation should work extra hard to provide food, otherwise their own
offspring are likely to die of starvation.
What is Fixed Action Pattern?

Self-Assessment Exercises 1

Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is the important of fixed action pattern?

5.4 Motivation and Drives

Motivation is the various internal factors that contribute to an animal

behaving in a particlur way. Motivation is an intervening variable used
to account for factors with in the organism that arouse, maintain and
channel behaviour towards a goal. Motivated behaviour is a drive that
leads to goal oriented behaviour and satiation.

The term motivation defines some kind of internal variable which

influences the relationship between stimulus and response. Motivation
was previously described as drives, which build up, the animals threshold
of response to functionally related groups of environmental stimuli.
(Thus the buildup of feeding drive resulted in an increased
responsiveness to stimuli connected with food, like the sight or smell of
prey). Motivation or drive was resulted in rise or fall in threshold levels
of responsiveness to functionally related stimuli. Behaviours which
depend on an internal state are said to be motivated, and the study of
animal motivation is an important part of ethology.
Motivated behaviour has 3 distinct phases,

(1) searching phase or phase of appetitive behaviour,

(2) orientation phase or phase of consummatory behaviour and
(3) quiescent phase.

i. Lorenz' psychohydraulic model.

Lorenz’s hydraulic model, one of the earliest attempts to model

motivation used the analogy of a hydraulic flow system. The fluid in its
resting state, the valve blocks the next centre in the hierarchy and inhibits
the performance of behaviours beyond that point. When it is excited, the

40
BIO 314 ANIMAL ECOLOGY

block is removed and the centre is freed for propagation. Impulses can
now travel down to lower centres controlling for e g; brood care, resting
and fighting but each of these centers is blocked until their appropriate or
key stimulus appears. In the case of fighting the key stimulus would be a
rival. The rival must be then providing a range of further key stimuli to
elicit particular fighting behaviours like biting or chasing, etc.

ii. Deutsch’s model.

An early attempt at the kind of motivational model which has proved very
powerful in recent years is the loop system proposed by deutsch.
Although deutsch’s model was designated to cover much wider aspects
of behaviour, only the part representing motivation is reproduced here.
The model operates in the following way. A deficit or imbalance in the
animals physiology is detected by and excites ‘central structure ‘or ‘link’.
The persistence and strength of this excitation depends on the magnitude
of the imbalance. The excited link then activated an appropriate motor
system which makes corrective behaviour in motion. As a result of the
animal’s behaviour some aspect of its internal and external environment
changes. (For example it has eaten and its stomach is now full of food),
the change in the environment is registered by the analyzer which inhibits
the link so that it no longer responds to inputs from the internal/external
environment. This inhibition slowly decays until the link is once again
sensitive to excitation.

5.4.1 Drive or Urge in Animals

The term Drive was introduced by Woodworth (1918) as motivational

concept. Animals experienced drive as biological needs such as eating
and drinking and alteration in their behaviour. Drive theories were later
given by Sigmund Freud (1915) and Clark Hull (1943). Freud, who was
physiologist by training, believed that drives and urges such as hunger
were recurring conditions in the body of animal that produced energy
build up in the nervous system.

This energy build up caused psychological discomfort and restlessness

that kept on increasing unless the urge was satisfied. Drive arose from
a range of bodily disturbances, such as deprivation of food, water, air,
sleep or temperature regulation, injury or activities like nest building.
Freudian drive theory was based on the following three principles:

a.) Drive emerged from bodily needs of the animal.

b.) Drive energized the behaviour of animal due to restlessness.
c.) Reduction of drive by satisfying needs produced learning.

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BIO 314 ANIMAL ECOLOGY

Konrad Lorenz (1950) proposed the Psycho-hydraulic model or Flush

toilet model to explain the drive and consummatory behaviour, which
has three steps:

1.) Drive causes action specific energy to accumulate with time and
causes increased restlessness in the animal, which results in
searching behaviour for food, water, mate etc.
2.) Consummatory behaviour starts after achieving the goal such as
food or any other sign stimuli. The innate release mechanism
releases the accumulated energy in the animal.
3.) After consummatory behaviour there is a quiescent period in the
animal as the accumulated energy has been released and the action
stoped. This is called refractory behaviour of the animal.

The following are the different forms of Drive in Animal.

i. Hunger & Thirst Drive

Hunger drive is controlled by lateral hypothalamus and ventro-median

nucleus, the former is stimulatory in function while the latter is
inhibitory. Glucocorticoids inhibit the hunger drive. Lateral
hypothalamus can be stimulated by epinephrine. The hunger and thirst
drives depend on hours of deprivation of feeding on dry food.

ii. Hoarding Drive

Many mammals such as male gerbil and squirrels possess hoarding

drive as the lean season approaches. Low estrogens and testosterone
levels stimulate hoarding drive in mammals. Castrated individuals show
increased hoarding drive, which can be reduced by giving testosterone
treatment.

iii. Migratory Drive

Migratory drive occurs in fishes and birds and may be seasonal or

related to reproduction. Pineal glands, which is affected by day-light
hours, affects migration in birds. In warblers pituitary gland influences
migratory urge as well as excessive eating to deposit fat energy in the
body. In stickleback fish thyroxin injection caused them to migrate.
In Salmon and Anguilla, maturation of gonads produces migratory
drive, so much so that they stop eating and set out to the course of
migration crossing all obstacles on the way.

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BIO 314 ANIMAL ECOLOGY

iv. Aggression Drive

Aggression is controlled by amygdala of the limbic system of brain and

posterior hypothalamus is also involved to some extent. In most of the
male mammals testosterone causes aggression while in females high
oestrogen levels reduce aggression and make the female peaceful.
Hydrocortisone also increases aggression while hydroxydione
decreases it. In ringed dove implants of testosterone propionate at
specific sites of hypothalamus causes aggressiveness.

v. Territorial Drive

Many vertebrates mark and defend their territory. Dogs, hyenas and
some prosimians mark their territory by their own urine. Monotremes
and marsupials have anal glands which they rub on the ground to mark
territory. In tigers and cheetahs also there are anal glands which spray
the secretion on the trees to mark their territory. Gazelles possess orbital
glands below the eyes which secrete a tar-like substance that they apply
on grasses and bushes. Territorial behaviour is also hormone dependent.
Yahr & Thiessen (1972) isolated 11 different hormones that influence
territorial behaviour in vertebrates.

vi. Sexual Drive

Sexual drive involves courtship behaviour such as singing and dancing

in birds, croaking in frogs and fighting in males of many vertebrates. In
insects courtship behaviour is stopped if corpora allata are removed.
Hormonal levels increase in breeding season. Castrated males and
females do not show sexual behaviour in vertebrates while testosterone
injections elicit sexual behaviour. According to Johnson (1976)
oestrogen enhances female attractiveness and receptivity and causes
oestrous in females. Hypothalamic Releasing Factor (LH-RF) and
ACTH are known to affect copulatory behaviour in many animals.

vii. Parental Care Drive

Gonadotropin secretion by pituitary gland cause not only courtship

display but also parental care in birds. Progesteron injections made the
birds sit on the eggs to incubate within 20 minutes. In pigeons, secretion
of prolactin from pituitary causes enlargement of crop to produce
pigeon-milk which is fed to the chicks. Prolactin also acts directly on
brain and makes the preoptic nucleus of hypothalamus in birds to
respond to chicks calls. What is Motivation?

Answer: Motivation is the various internal factors that contribute to

an animal behaving in a particlur way. Motivation is an intervening

43
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variable used to account for factors with in the organism that arouse,
maintain and channel behaviour towards a goal. What is Drive?

Self-Assessment Exercises 2
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. List the three step involves in expliaing the drive and
consummatory bahaviour?
2. Write on hunger and thrist drive.
5.5 Summary

A fixed action pattern is one of the few types of behaviors which can be
said to be hard-wired and instinctive. They are modulated by the
environment; a fixed response can lead to maladaptive results, whereas
flexible behaviors are generally more useful.

5.6 References/Further Readings

Alcock, J. (1998) Animal Behavior: An Evolutionary Approach (6th

edition), Chapter 5. Sinauer Associates, Inc. Sunderland,
Massachusetts. ISBN 0-87893-009-4

Breland, K. & Breland, M. (1961) The misbehavior of

organisms. American Psychologist 16: 681–84.
[Link]

Catania, A. C. (1984) Learning, 2d [Link]-Hall. ISBN-10:

0135276977.

Colgan, P.W. (1989) Animal Motivation. Springer Netherlands.

DOI:10.1007/978-94-009-0831-4.

Cooper JO (2007) Applied Behavior Analysis. Upper Saddle River, NJ,

USA: Pearson Education. ISBN: 978-0-13-129327-4.

Dawkins, M. (1990) From an animal’s point of view: Motivation, fitness,

and animal welfare. Behavioral and Brain Sciences, 13(1), 1-9.
DOI:10.1017/S0140525X00077104.

Dawkins, R. (1982) The extended phenotype. W. H. Freeman. ISBN-10:

0716713586.

Freud, S. (2012). A General Introduction to Psychoanalysis. Renaissance

Classics. ISBN: 9781484156803.

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BIO 314 ANIMAL ECOLOGY

Hughes, B.O. and Duncan, I.J.H. (1988) The notion of ethological ‘need’,
models of motivation and animal welfare. Animal Behaviour,
Volume 36, Issue 6, 1988, Pages 1696-1707. ISSN: 0003-3472.

James E. M. (2005) Learning and Behavior, (6th Edition]. New

Jersey: Prentice Hall. ISBN 978-013-193163-3

Lorenz, K. (1950) The comparative method in studying innate behaviour

patterns. Symposium of the Society for Experimental Biology 4:
221–68. [Link]

Lorenz, K. (1981) The Foundations of Ethology. Springer. ISBN: 978-3-

211-81623-3.

McFarland, D. J. (1989) Problems of Animal Behaviour. Longman

Scientific & Technical. ISBN-10: 0582468205.

Salamone, J. D. and Correa, M. (2012) The mysterious motivational

functions of mesolimbic dopamine. Neuron. 76 (3): 470–85. DOI:
10.1016/[Link].2012.10.021

Tinbergen, Nikolaas (1951). The Study of Instinct. Oxford University

Press. ISBN: 9780198577225.

Tinbergen, N. (1963). On aims and methods of Ethology. Zeitschrift für

Tierpsychologie. 20 (4): 410–433. DOI:10.1111/j.1439-
0310.1963.tb01161.x.

Wickler, W. (1968) Mimicry in Plants and Animals. World University

Library, London
Wilson, E.O. (1975) Sociobiology: The New Synthesis. Harvard
University Press. ISBN: 0-674-00089-7.

[Link]
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DUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTE4MA
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a992978a655d&psq=list+of+related+materials++in+Fixed+Actio
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WFsLWJlaGF2aW9yLWZhbGwtMjAxMy9hZGFlMTM0NjVm
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zX0xlYzgucGRm&ntb=1

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BIO 314 ANIMAL ECOLOGY

[Link]
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DUtMWYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTMyM
A&ptn=3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-
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GUtZW52aXJvbm1lbnQvYS9pbm5hdGUtYmVoYXZpb3Jz&nt
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WYxNS1hOTkyOTc4YTY1NWQmaW5zaWQ9NTMxMw&ptn=3&hs
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3d3cuY291cnNlaGVyby5jb20vZmlsZS8xNzgxMTU0OS9DaWFsZGlu
aS1DaGFwdGVyLTEtMy8&ntb=1

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FpbCZGT1JNPVZJUkU&ntb=1

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[Link]
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n=3&hsh=3&fclid=13a0bbfa-9697-6405-1f15-

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BIO 314 ANIMAL ECOLOGY

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tivation%2Band%2BDrive%2Bin%2Banimals%26FORM%3DVD
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5.7 Possible Answers to SAEs

Answers to SAEs 1

1. A FAP is significant in animal behavior because it represents the

simplest type of behavior, in which a readily defined stimulus
nearly always results in an invariable behavioral response. A FAP
can truly be said to be "hard-wired." FAPs are also unusual in
that almost behaviors are modulated by the environment; a
fixed response can lead to maladaptive results, whereas flexible
behaviors are generally more useful.

Answers to SAEs 2

1. Drive causes action specific energy to accumulate with time and

causes increased restlessness in the animal, which results in
searching behaviour for food, water, mate etc.

Consummatory behaviour starts after achieving the goal such as

food or any other sign stimuli. The innate release mechanism
releases the accumulated energy in the animal.

After consummatory behaviour there is a quiescent period in the animal

as the accumulated energy has been released and the action stoped.
This is called refractory behaviour of the animal.

2. Hunger drive is controlled by lateral hypothalamus and ventro-

median nucleus, the former is stimulatory in function while the
latter is inhibitory. Glucocorticoids inhibit the hunger drive.
Lateral hypothalamus can be stimulated by epinephrine. The
hunger and thirst drives depend on hours of deprivation of
feeding on dry food

Glossary

°C = degrees Celsius
cm = centimeters
CNS = Central nervous system
CH4 = Methane
CO = Carbon
CH4 = methane (CH4)
DNA = Deoxyribonucleic acid

48
BIO 314 ANIMAL ECOLOGY

E. coli = Escherichia coli

FAP = Fixed Action Pattern
°F = degree Fahrenheit
Ft = feet
h = hour
in = inches
Kgs = kilograms.
km = kilometers
kph = kilometer per hour
m = meters
mm = millimeters
mph = meter per hour
NH2OH = Hdroxylamine
NH3 = Ammonia
O2 = Oxygen
PNS = Peripheral Nervous System
PRC = Phase Response Curve
RNA = Ribonucleic acid
L = length
HIV = Human immunodeficiency virus
% = percentage
g = grams
spp = species
UV = Ultraviolet

End of the Module Questions

1. What is animal communication?

2. What is courtship rituals?
3. What is reflex?
4. What is object orientation?
5. What is Phototaxis?

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BIO 314 ANIMAL ECOLOGY

Module 2 Learning, Communication and Social Behaviour

Module Introduction

In Module two, unit one deals with Display and displacement of animal
behavior, types of conflict behavior. While unit two deal with learning
and communication in Animal and the different types of learning behavior
exhibited by different organism.

Unit 1 Display and displacement behavior and conflct behavior

Unit 2 Learning and Communication in Animal
Unit 3 Social Behaviour
Unit 4 Social Behaviour of Primates
Unit 5 Hierarchical Organisation

Glossary

Unit 1 Displays, Displacemnet Behaviour and Conflict

Behaviour

Contents
1.1 Introduction
1.2 Intended Learning Outcomes (ILOs)
1.3 Display and Displacement behvaiour in Animal
1.3.1 Displacement activities
1.3.2 Examples of Displacement behavior in animal
1.4 Conflict Behaviour
1.4.1 Types of Conflict behavior in animal
1.5 Summary
1.6 References/Further Readings/Web sources
1.7 Possible Answers to Self-Assessment Exercises

1.1 Introduction

Display behaviour is a ritualized behaviour by which an animal provides

specific information to others, usually members of its own species while
displacement behavior is usually thought of as self-grooming, touching,
or scratching, which is displayed when an animal has a conflict between
two drives, such as the desire to approach an object, while at the same
time being fearful of that object.
.

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BIO 314 ANIMAL ECOLOGY

1.2 Intended Learning Outcomes (ILOs)

At the end of this unit, students should be able to:

 Explain displacement behavior in animals
 List examples of displacement behvaiour in animal
 Explain conflict behaviour

1.3 Display and Displacement behvaiour in Animal

Virtually all higher animals use displays to some extent. The best-known
displays are visual ones—and some biologists restrict the term display to
visual signals or gestures—but many also incorporate sound, smell, or
even touch. Displays evolve through the ritualization of specific
behaviour patterns. Some mating displays evolve from food-giving
behaviours; the male bobwhite quail gives a food call and offers a tidbit
to his potential mate. In many birds the food-giving behaviour is
completely ritualized and proceeds without any exchange of food;
domestic cocks, for example, call and peck at bare ground to attract a hen.

Agonistic (aggressive) displays usually occur near the borders of a

territory. When a strange howler monkey approaches the territory of
others, resident males set up a tremendous din, warning the intruder off.
Many songbirds sit on highly visible perches while singing, providing
both auditory and visual displays. Agonistic display is adaptive in
conserving energy, making it unnecessary for the resident animal to chase
others away. Furthermore, where display occurs, injury is rare, as physical
contact is rarely required. An impending threat to the group may provoke
display behaviour that is protective, signaling danger at the approach of
a predator.

Another type of display behaviour is that designed to deceive a predator

or lure it away from vulnerable young. An example is the broken-wing
display-where the parent flutters along the ground as if injured-used by
many birds to lure predators away from the nests. See also alarm
signal; courtship; territorial behaviour.

1.3.1 Displacement activities

Displacement activities as described by Lorenz are motor programs that

seem to discharge tension or anxiety. For example, if one is trying to
entice a squirrel to come up and take a peanut, the squirrel becomes
conflicted-caught between two incompatible drives. It wants the nut, but
it fears humans. The squirrel is caught between approach and avoidance

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BIO 314 ANIMAL ECOLOGY

tendencies, but it cannot do both at once. It becomes visibly edgy.

It may take a few hops toward the human holding the peanut, then
scratch itself suddenly or make a few digging movements. This does not
mean the squirrel itches or needs to dig a hole. Lorenz suggested it
was "breaking the tension" caused by competing urges

In 1940 Tinbergen and Kortlandt independently drew attention to a

behavioural phenomenon which has since been called displacement
activity and has received a good deal of attention. Although no binding
rules exist by which displacement behaviour can be recognized, the
term is applied to behaviour patterns which appear to be out of context
with the behaviour which closely precedes or follows them either in the
sense that they do not seem functionally integrated with the preceding or
following behaviour or that they occur in situations in which causal
factors usually responsible for them appear to be absent or at least weak
compared with those determining the behavioural envelope.

Displacement activities occur in three

situations:
i. Conflict
ii. frustration of consummator acts
iii. physical thwarting of performance

.
1.3.2 Examples of displacement behaviour in animals

i) Displacement behaviour in dog

There are basically two types of displacement behaviours: those that are
self- directed to something the dog does to himself, and those that are
re-directed to something external. A common example of a self-directed
displacement behaviour in dogs is self-grooming, most often licking
the genital area. Another common self-directed behaviour is yawning.

Common examples of re-directed displacement behaviours are

finding, picking up and carrying a toy, barking, circling, grazing grass
and gulping water as the reader describes. In a multi-dog household, re-
directed behaviour often takes the form of one dog jumping onto and
engaging in play with another dog, grabbing, wrestling and the like. This
is what displacement behaviours are; now to the bigger question of why
dogs and other animals (including us) engage in them.

Displacement behaviour occurs at times of emotional conflict, serving as

an outlet to dissipate energy. Using the reader’s question for example,
the behaviours that are in conflict have to do with excitement and
expression of greeting behaviour. Let’s explore what normal greeting

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BIO 314 ANIMAL ECOLOGY

behaviour is, and why a dog might have conflicting emotions about it:
For a dog, greeting involves two major areas and behaviours: licking
the mouth of the returning pack member (or visitor), and sniffing the
genital area. While both these behaviours are normal for dogs, most of
us humans discourage such expressions of friendship.

Jumping up on us in greeting is because the dog is trying to lick us

around the mouth. Since we are upright rather than on all fours, dogs
can’t reach our mouths without jumping up. Most of us don’t want our
dogs to jump on us, so we discourage this normal dog behaviour. In most
cases, such discouragement is a verbal reprimand or scolding, and
sometimes involves some form of physical punishment such as
applying a knee to the dog’s chest. Embarrassing sniffing behaviour,
as well, is most often strongly reproached.

Reprimanding or punishing what is normal behaviour for a dog

(inappropriate though we may consider it) makes the dog feel anxious
and stressed. Over time, these feelings become intrinsically associated
with the situation that triggers them, so even once the dog has learned
to not jump up, he is conditioned to feel anxious in this situation.

Just as importantly, chastising the dog for what is normal does not
provide an alternative outlet for the energy of this behaviour. For
example, teaching the dog to sit or get a toy and carry it around when
someone comes to the door creates an alternative behaviour outlet for
his energy. Displacement behaviour occurs in the absence of learning a
positively reinforced alternative behavior to replace his normal greeting
behaviour.

It isn’t just dogs that operate this way. Consider how you would feel if
you suddenly find yourself in an unfamiliar culture, with different,
unknown greeting rituals. You offer your hand to shake hands and the
person looks at you with disgust and turns away. Standing there foolishly
with your hand outstretched you might laugh uncomfortably, cough and
cover your mouth with your outstretched hand, or reach to pick up
something, as if that’s what you intended all along all displacement
behaviour.

Now think of how much better you would feel when you have been
forewarned as to proper greeting in this unfamiliar culture. Feeling no
anxiety you would offer the appropriate behaviour. This is just how we
should approach greeting behaviour, or any other “normal” dog
behaviour that we consider inappropriate or unacceptable in our society
and culture. Rather than simply expressing our dismay or disgust, the
best approach is to teach your dog an alternative, acceptable behaviour
so his emotions will no longer be in conflict.

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BIO 314 ANIMAL ECOLOGY

While the reader’s dog doesn’t seem too terribly upset, and has found an
acceptable displacement behaviour, the reader could ask the dog to
sit, lie down, or offer another learned behaviour during greeting, and see
if it doesn’t change her need to drink water.

And finally, I want to thank the reader for noticing that I invite
questions. I try to answer all emails I receive, and am most appreciative
of getting them. Readers’ questions often trigger column topics I hadn’t
thought of, and I’m always grateful for new ideas!

ii) Displacement behaviou in cat

In the cat Parmeggiani stimulated several different and separate areas

of the forebrain and brainstem and obtained a behavioural complex
consisting of sniffing, grooming, yawning, lying down, curling up,
dozing and sleeping. He emphasizes that this behaviour is normal in
unstimulated cats. Rowland and Gluck present some evidence that in a
certain conditioning procedure grooming replaced the synchronization of
the electroencephalogram shown by sleeping cats when those were
tested awake. Again, Leyhausen lists grooming, sniffing and lying
down as displacement behaviour for the cat.

For the rat, Caspers has shown that grooming and some other
unfortunately unspecified "motorautomatisms" are associated with shifts
in the cortical d.c. potential making the surface positive, which otherwise
are typical of sloop, while shifts towards a negative surface are
characteristic of the waking animal. Grant mentions grooming, digging
and sniffing as typical displacement activities of rats.

These facts suggest that grooming or preening and certain other

movements are largely controlled by neurophysiological mechanisms
which are also responsible for de-arousal and sleep. On the other
hand they are often involved in displacement behaviour. There is little
doubt, however, that the behavioural situations loading to displacement,
that is, conflict, frustration and thwarting, are effective in increasing
arousal. I suggest that the occurrence of at least some displacement
activities is the reflexion of a homeostatic process operating towards
cancelling the arousal increment so generated, through the activation of
an arousal inhibiting system. The existence of arousal homeotasis has
been suggested by Borlyne, who also marshals supporting empirical
evidence. Such regulation appears logically necessary if arousal is
correlated with the rate at which information is handled and if the
nervous system is considered as a communication channel of restricted
and specific capacity whore for maximum efficiency the information
handling rate must be hold within certain
What is Displacement activities?

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BIO 314 ANIMAL ECOLOGY

What are the common examples of re-directed displacement behavior

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is Display behavior?
1.4 2. Conflict
MentionBehavior
the three situation in which displacement activities
occur.

Conflict behavior is a state of motivation in which tendencies to perform

more than one activity are expressed simultaneously. At any particular
moment, an animal has many different incipient tendencies, but by a
process of decision-making, one of these becomes dominant. Generally,
only one tendency becomes dominant, but in certain circumstances
more than one competes for dominance, and conflict arises.

1.4.1 Types of conflict behavior in animals

Conflict has traditionally been divided into three main types:

i) Approach–approach: conflict occurs when two tendencies in

conflict are directed towards different goals. In such a case the
animal may reach a point where the two tendencies are in balance.
However, the tendency to approach a goal generally increases
with proximity to the goal. This makes approach– approach
conflict unstable, because any slight departure from the point
of balance, towards one goal, will result in an increased
tendency towards the other, thus resolving the conflict.

ii) Avoidance–avoidance: conflict occurs when the two tendencies in

conflict are directed away from different points. Since the
tendency to avoid objects generally increases with proximity to the
object, movement toward either object is likely to result in a return
to the point of balance. Such situations are not normally stable,
because the animal can escape in a direction at right angles to the
line between the two objects.

iii) Approach–avoidance: conflict occurs when one activity is

directed towards a goal, and another away from it. For example,
an animal may have a tendency to approach food, but be
frightened of the strange food dish. The nearer it approaches the
food, the stronger the approach tendency, but the nearer it gets
to the dish the stronger the avoidance tendency. The animal can
reach the food, only when the approach tendency is larger than
the avoidance tendency. Often there is an equilibrium point,
some distance from the goal. When the animal approaches beyond

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BIO 314 ANIMAL ECOLOGY

this point, avoidance is greater than approach. When the animal

retreats, approach is larger than avoidance. Such situations tend
to be stable, because the animal is always pulled towards the
equilibrium point.

Approach–avoidance conflict is by far the most important and most

common form of conflict in animal behaviour. Typically such conflict is
characterized by compromise and ambivalence, especially near the point
of equilibrium. Irrelevant behaviour, such as displacement activity, is also
common, as are various forms of display and ritualization. Ritualized
conflict behaviour often occurs in territorial disputes, and often forms the
basis of threat display.
What is conflict behavior?

Self-Assessment Exercises 2
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. Write on Avoidance – avoidance as a form of conflict
behavior.

1.4 Summary

Display behaviour is a ritualized behaviour by which an animal provides

specific information to others, usually members of its own species while
displacement behavior is usually thought of as self-grooming, touching,
or scratching, which is displayed when an animal has a conflict between
two drives.

1.5 References/Further Readings/Web Sources

Baker K. C., Aureli F. 1997. Behavioural indicators of anxiety: An

empirical test in chimpanzees. Behaviour 134: 1031-1050.

Castles D. L., Whiten A., Aureli F. 1999. Social anxiety, relationships

and self-directed behaviour among wild female olive baboons
Animal Behaviour 58: 1207-1215.

Hughes B. O., Duncan I. J. H. 1988 The notion of ethological need,

models of motivation and animal-welfare. Animal Behaviour 36:
1696-1707.

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Leavens DA, Aureli F, Hopkins WD, Hyatt CW 2001 Effects of

cognitive challenge on self-directed behaviors by chimpanzees
(Pan troglodytes) American Journal of Primatology 55 (1): 1-14

Maestripieri D., Schino G., Aureli F., Troisi A. 1992. A modest proposal
- displacement activities as an indicator of emotions in primates.
Animal Behaviour 44: (5) 967-979

Maestripieri, D., Schino, G., Aureli, F. & Troisi, A. A modest proposal:

displacement activities as an indicator of emotions in
primates. Anim. Behav. 1992; 44; 967-979

Manson JH, Perry S 2000 Correlates of self-directed behaviour in

wild white faced capuchins Ethology 106 (4): 301-317

Troisi A., Belsanti S., Bucci A. R., Mosco C., Sinti F., Verucci M.
2000 Affect regulation in alexithymia - An ethological study
of displacement behavior during psychiatric interviews. Journal
Of Nervous And Mental Disease 188: (1) 13-18

[Link]
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1.6 Possible Answers to SAEs

Answers to SAEs 1

1. Display behaviour is a ritualized behaviour by which an animal

provides specific information to others, usually members of its
own species.
2. Displacement activities occur in three situations:
i. Conflict
ii. frustration of consummator acts
iii. physical thwarting of performance

Answers to SAEs 2

1. Drive causes action specific energy to accumulate with time and

causes increased restlessness in the animal, which results in
searching behaviour for food, water, mate [Link] Avoidance–
avoidance: conflict occurs when the two tendencies in conflict are
directed away from different points. Since the tendency to avoid
objects generally increases with proximity to the object,
movement toward either object is likely to result in a return to the
point of balance. Such situations are not normally stable, because
the animal can escape in a direction at right angles to the line
between the two objects

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Unit 2 Learning and Communication in Animal

Contents

2.1 Introduction
2.2 Intended Learning Outcomes (ILOs)
2.3 Learning and Communication in Animal
2.3.1 Types of Learning behavior in Animal
2.4 Communication in Animals
2.5 Summary
2.6 References/Further Readings/Web Sources
2.7 Possible Answers to Self-Assessment Exercises

2.1 Introduction

Through learning, animals can adjust quickly to changes in their

environment. Learning is adaptive for animals in an environment where
changes are not predictable. Learning produces changes in the behavior
of an individual that are due to experience. Once an animal learns
something, its behavioral choices increase.

An animal’s ability to learn may correlate with the predictability of certain

characteristics of its environment. Where certain changes in the habitat
occur regularly and are predictable, the animal may rapidly respond to a
stimulus with an unmodified instinctive behavior. An animal would not
necessarily benefit from learning in this situation. However, where certain
environmental changes are unpredictable and cannot be anticipated, an
animal may modify its behavioral responses through learning or
experience. This modification is adaptive because it allows an animal to
not only change its response to fit a given situation, but also to improve
its response to subsequent, similar environmental changes.

2.2 Intended Learning Outcomes (ILOs)

At the end of lecture, the students should be to:

 Understand the biological meaning of learning and the categories

of learning.
 Define communication and the different types of communication.

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 Modalities for communication in animals

2.3 Learning and Communication in Animal

2.3.1 Types of Learning Behaviour in Animals

i. Habituation

Habituation is the simplest and perhaps most common type of behavior

in many different animals. It involves a waning or decrease in response
to repeated or continuous stimulation. Simply an animal learns not to
respond to stimuli in its environment that are constant and probably
relatively unimportant thereby saving time as well as conserving energy.
For example, after time, birds learn to ignore scarecrows that previously
cause them to flee. Squirrels in a city park adjust to the movements of
humans and automobiles. Habituation is believed to be controlled
through central nervous system and should be distinguished from
sensory adaptation. Sensory adaptation involves repeated stimulation of
receptors until they stop responding. For example, if you enter a room
with an odor, your olfactory sense organs soon stop responding to these
odors.

ii. Classical conditioning

Classical conditioning is a type of learning documented by Russian

physiologist, Ivan Pavlov (1849-1936). In his classic experiment on the
salivary reflex in dogs, Pavlon presented food right after the sound of
bell. After a number of such presentations, the dogs were conditioned-
-- they associated the sound of the bell with food. It was then possible
to elicit the dog’s usual response to food – salivation—with just the
sound of the bell. The food was a positive reinforcement for salivating
behavior, but responses could also be conditioned using negative
reinforcement. Classical conditioning is very common in the animal
kingdom. For example, birds learn to avoid certain brightly colored
caterpillars that have a noxious taste. Because birds associate the color
pattern with the bad taste, they may also avoid animals with a similar
color pattern.

iii. Instrumental conditioning

In instrumental conditioning (also known as trial-and-error learning), the

animal learns while carrying out certain searching actions, such as
walking and moving about. For example, if the animal finds food during
these activities, the food reinforces the behavior and the animal

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associates the reward the reward with the behavior. If this association is
repeated several times, the animal learns that the behavior leads to
reinforcement; the animal then tends to repeat or avoid that behavior,
depending on whether the reinforcement is positive or negative. For
example, an American psychologist B.F. Skinner placed a rat in a
“Skinner box’’ which have a choice of various levers it might push, some
of which reward the animal by releasing food. The animal’s choices may
be random at first, but quickly learns to choose those levers that provide
food. Such learning is the basis for most of the animal training done by
humans, in which the trainer typically induces a particular behavior at
first by rewarding the animal.

Instrumental conditioning is undoubtedly very common in nature. For

example, animals quickly learn to associate eating particular food items
with good or bad tastes and modify their behavior accordingly. In some
cases animals may be able to skip some of their own trial and error and
learn simply by watching the behavior of others. A good example is that
of tits (chickadee-like birds) in England. Sometime in the early 1950s,
one of these birds apparently learned to peck through the paper tops of
milk bottles left on doorsteps and drink the cream on top. This
was probably a case of instrumental conditioning with the bird learning
that its general pecking, probing behavior was rewarded if directed at the
bottles. But the behavior quickly spread through the population and was
“handed down” to the succeeding generation that that learned the
behavior by watching adults.

iv. Latent learning

Latent learning sometimes called exploratory learning, involves

making associations without immediate reward. The reward is not
obvious. An animal is apparently motivated, however, to learn about its
surroundings. For examples, if a rat is placed in a maze that has no food
or reward, it explores the maze, although rather slowly. If food or another
reward is provided, the rat quickly runs the maze. Apparently previous
learning of the maze had occurred but remained latent, or hidden, until an
obvious reward was provided. Latent learning allows an animal to learn
about its environment as it explores. Knowledge about an animal’s home
area may be important for its survival, perhaps enabling it to escape from
a predator or capture pray.

v. Insight learning

Insight learning is the ability to perform a correct or appropriate

behavior on the first attempt in a situation with which the animal
have no previous experience (some prefer to call it reasoning, rather
than learning). For example, if a chimpanzee is placed in an area with a

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banana hung too high above its head to be reached and several boxes on
the floor, the chimp can size up the situation and then stack the boxes to
allow it reach the food. In general insight is best developed in primate
and other mammals but even in these groups the level of insight
often varies from one situation or species to another. The great
majority of animals display little or no ability to use insight.

Very broadly the capacity to learn can be thought of as another adaptation

that enhances survival and reproductive success and must have some
genetic bases. However, the internal mechanisms of learning are very
poorly known and it is only recently that progress has begun on linking
some simple kinds of learning to internal biochemical or physiological
changes. Although animals frequently appear to do complex things, most
behaviors can be understood as relatively fixed patterns that are often
modified in their frequency and orientation by simple kinds of learning.
Put in another way, animals in general are not all that “smart” rather they
have been fine-tuned by natural selection and their limited repertoire
of abilities work very well in normal circumstances.
What is habituation?

What is Insight learning?

2.4 Communication in Animals

Communication is the transfer of information from one animal to

another. It requires a sender and receiver that are mutually adapted to
each other. The sender must send a clear signal to the receiver.
Communication can occur within species (intraspecific) or between
species (interspecific). Intraspecific communication in animal is very
important for reproductive success. Examples of intraspecific
communication include warning signals, such as the rattle of a rattle
snake’s tail and the skunk’s presentation of its hindquarters and tail.

Animals use a variety of modalities for communication, including visual,

auditory, tactile, and chemical signals. Natural selection has influenced
the characteristics of a signal system. Animals have evolved
combinations of signals that may be more effective than any single
signal.

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2.4.1 Types of Communication in Animals

i. Visual communication

Visual communication is important to many animals because a large

amount of information can be conveyed in a short time. Most animals
(e.g., cephalopod mollusk, arthropods and most vertebrates other than
mammals) with well- developed eyes have color vision. Many fishes,
reptiles and birds exhibit brilliant color patterns that usually have a
signaling function. Most mammals have plain, darker colors and lack
color vision because they are nocturnal, as were their probable
ancestors- nocturnal insectivores. Primates are a notable exception in that
they have both color vision and colorful displays.

A visual signal may be present at all times, as are the bright facial
markings of a male mandrill. The signal may be hidden or located on a
less exposed part of an animal’s body and then suddenly presented. Some
lizards, such as green anoles can actually change their color through
activities of pigment cells in the skin.

Visual signals have some disadvantages in that various objects in the

environment may block the line of sight and / or the signals may be
difficult to see over a long distance. Also, the signals are usually not
effective at night and may be detected by predators.

ii. Acoustic communication

Arthropods and vertebrates commonly use acoustic or sound

communication. These animals must expend energy to produce
sounds but sounds can be used during the night or day. Sound waves
also have the advantage of traveling around objects and may be
produced or received while an animal is in the open or concealed.
Sounds can carry a large amount of information because of the many
possible variations in frequency, duration, volume and tone.

Acoustic communication systems are closely adapted to the

environmental conditions in which they are used and the function of the
signal. For example, tropical forest birds produce low frequency calls
that pass easily through dense vegetation. Many primates in tropical
forests produce sounds that travel over long distances. Other examples
include the calls of territorial birds that sit on a high perch to deliver
the signal more effectively and the alarm calls of many small species
of birds. Some of the more complex acoustic signals that have been
studied are birdsong and human speech.

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iii. Tactile Communication

Tactile communication refers to the communication between animals in

physical contact with each other. The antennae of many invertebrates
and the touch receptors in the skin of vertebrates function in tactile
communication. Some examples of tactile communication are birds
preening the feathers of other birds and primates grooming each other.

iv. Chemical communication

This is another common mode of communication. Unicellular organisms

with chemoreceptor can recognize members of their own species.
Chemical signals are well developed in insects, fishes, salamanders and
mammals. Advantages of chemical signals are that they (1) usually
provide a simple message that ca last for hours or days; (2) are effective
night or day (3) can pass around objects; (4) may be transported over
long distances; (5) take relatively little energy to p r o d u c e . T h e
disadvantages of chemical signals are that they cannot be changed
quickly and are slow to act.

Chemicals that are synthesized by one organism and that affect the
behavior of another member of the same species are called pheromones.
Olfactory receptors in the receiving animal usually detect chemical
signals. Many animals mark their territories by depositing odors that
act as chemical signals to other animals of the same species. For
examples, many male mammals mark specific points in their territories
with pheromones that warn other males of their presence in the area. The
same pheromones may also attract females that are in the breeding
condition.

Differences in the chemical structure of pheromones may be directly

related to their function. Pheromones used for making territories and
attracting mates usually last longer because of their higher molecular
weights. Airborne signals have lower molecular weight and disperse
easily. For example, the sex attractant pheromones of female moths that
are ready to mate are airborne and males several kilometers away can
detect them.

The following are communication systems associated with the different

animals:

1. Auditory - birds and humans, some insects (crickets)

2. Olfactory - most mammals, moths
3. Visual - bees and dancing, fireflies at night
4. Altruism - personal sacrifice for the good of the group
5. Alarm call of mammals - Belding's Ground Squirrels

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6. Cooperative breeding - African bee-eaters, Scrub Jays Bees and

other hymenopterous insects
What is Communication?
What is another name for Latent Communication?#

Self-Assessment Exercises 1
Attempt these exercises to measure what you have learnt so far.
This should not take you more than 5 minutes.
1. What is tactile communication?
2. Which group of animals uses acoustic communication?

2.5 Summary

Through learning an animal can adjust quickly to changes in its

environment. Learning is adaptive for animals in an environment where
changes are not predictable. The types of learning known to occur in
animals include habituation, classical conditioning, instrumental
conditioning, latent learning and insight learning. Communication is
an essential aspect of animal behavior which enhances continuity in
existence. The different types of communication showed how animals
relate to one another. Communication in animals requires the use of clear
signals by one animal and their reception by another. Visual, acoustic,
tactile, and chemical signals are important channels in communication
systems. Communication can occur within species (intraspecific) or
between species (interspecific). Intraspecific communication in animal
is very important for reproductive success.

2.6 References/Further Readings/Web Sources

Bradbury, J. W. & Vehrencamp, S. L. (1998). Principles of Animal

Communication. Sunderland, MA: Sinauer Associates.

Dawkins, R. & Krebs, J. R. (1978). Animal Signals: Information or

Manipulation. In Krebs, J. R. & Davies, N. B. (Eds.), Behavioural
Ecology: An Evolutionary Approach (pp. 282–
309). Oxford: Blackwell

Hockett, C. F. (1960). The origin of speech. Scientific

American, 203(3), 88–97.

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Laidre, M. E. & Johnstone, R. A. (2013). Animal signals. Current

Biology, 23(18):829–833

Lattenkamp, E. Z. & Vernes, S. C. (2018). Vocal learning: A language-

relevant trait in need of a broad cross-species approach. Current
Opinion in Behavioral Sciences, 21, 209–215.

Maynard Smith, W. J. (1977). The Behavior of

Communicating. Harvard, MA: Harvard University Press

Owings, D. H. & Morton, E. S. 1998. Animal Vocal Communication: A

New Approach. Cambridge: Cambridge University Press

Owren, M. J., Rendall, D., & Ryan, M. J. (2010). Redefining animal

signaling: Influence versus information in
communication. Biology & Philosophy, 25(5), 755–780.

Partan, S. R. & Marler, P. (2005). Issues in the classification of

multimodal communication signals. American
Naturalist, 166(2), 231–245.

Rendall, D., Owren, M. J., & Ryan, M. J. (2009). What do animal signals
mean? Animal Behaviour, 78(2), 233–240.

Searcy, W. A. & Nowicki, S. (2005). The Evolution of Animal

Communication: Reliability and Deception in Signaling
Systems. Princeton, NJ: Princeton University Press.

Slocombe, K. E. & Zuberbühler, K. (2005). Functionally referential

communication in a chimpanzee. Current Biology, 15(19), 1779–
1784.

Smith, M. J. & Harper, D. G. (1995). Animal signals: Models and

terminology. Journal of Theoretical Biology, 177(3), 305–311.

Tanner, J. E., Patterson, F. G., & Byrne, R. W. (2006) The development

of spontaneous gestures in zoo-living gorillas and sign-taught
gorillas: From action and location to object representation. Journal
of Development Processes, 1, 69–102

Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift für

tierpsychologie, 20(4), 410–433

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Uetz, G. W., Roberts, J. A., & Taylor, P. W. (2009). Multimodal

communication and mate choice in wolf spiders: Female response
to multimodal versus unimodal signals. Animal
Behaviour, 78(B2), 299–305

Watson, S. K., Townsend, S. W., Schel, A. M., Wilke, C., Wallace, E.

K., Cheng, L., West, V., & Slocombe, K. E. (2015). Vocal
learning in the functionally referential food grunts of
chimpanzees. Current Biology, 25(4), 495–499.

Wiener, N. (1948). Cybernetics or Control and Communication in the

Animal and the Machine. Technology Press. New York: Wiley.

Zuberbühler, K. (2000). Interspecies semantic communication in two

forest primates. Proceedings of the Royal Society of London.
Series B: Biological Sciences, 267(1444), 713–718.
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2.7 Possible Answers to SAEs

Answers to SAEs 1

1. Tactile communication refers to the communication between

animals in physical contact with each other. The antennae of
many invertebrates and the touch receptors in the skin of
vertebrates function in tactile communication. Some examples of
tactile communication are birds preening the feathers of other
birds and primates grooming each other.

2. Arthropods and vertebrates commonly use acoustic or sound

communication

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Unit 3 Social Behaviour

Contents
3.1 Introduction
3.2 Intended Learning Outcomes (ILOs)
3.3 Social Behaviour
3.4 Summary
3.5 References/Further Readings/Web sources
3.6 Possible Answers to Self-Assessment Exercises

3.1 Introduction

Social behaviour typically refers to any interactions among members of

the same species, but it also applies to animals of different species,
excluding predator- prey interactions. A group of animals may form
an aggregation for some simple purpose, such as feeding, drinking or
mating. A true animal society is a stable group of individuals of the
same species that maintains a cooperative social relationship. This
association typically extends beyond the level of mating and taking
care of young. Social behaviour has evolved independently in many
species of animals; invertebrates have complex social organizations.

3.2 Intended Learning Outcomes (ILOs)

At the end of lecture the students should be able to;

 Understand the meaning of social behavior and

 Identify different types of social behavior among animals.

3.3 Social Behaviour

In the animal kingdom, social species group together with the goal of
increasing the overall survival of the species. A solitary antelope is more
likely to get picked off by a lion than if it had the protection of the group
around it. Pack animals, like wolves and dolphins, work together to obtain
food, raise young and protect the group. While they may have to share the
food with others in the group, their overall survival is increased by the
presence of the group.

Animal populations are often organized into groups. A group of animals

may form an aggregation for some simple purpose, such as feeding,

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drinking or mating. Several Drosophila flies on a piece of rotting

fruit are an example of an aggregation. A true animal society is a
stable group of individuals of the same species that maintains a
cooperative social relationship. This association typically extends
beyond the level of mating and taking care of young. Social behavior has
evolved independently in many species of animals; invertebrates have
complex social organizations.

One major benefit of belonging to a group may be that it offers protection

against predators. There is safety in numbers and predator detection may
be enhanced by having several group members on alert to warn
against an intruder. Also, cooperative hunting and capture of prey
increase the feeding efficiency of predators. Living in social groups
is also advantageous in some instances due to the ability to gain
protection from the elements (e.g. huddling together in cold weather) and
during the processes of mate finding and rearing of young. In many
species, most notably the social insects, living groups has resulted
in the evolutionary division of labor, with specific individuals
performing specialized tasks (defense, food procurement, feeding of
young.

A disadvantage of group living may be competition for resources. Other

disadvantages include the diseases and parasites that may spread more
rapidly in group-living animals and interference between individuals
with regard to reproduction and rearing of young. The value of group
living depends on the species and behaviors involved. Below is an
example of social behavior in animals:

i. Altruism and Kin Selection

Sometimes one animal in the group will do something to increase the

survival of another at the potential cost to its own fitness or survival. This
behavior is known as altruism. For example, if we are stranded on a
deserted island and I find food and share it with you, I am performing an
altruistic act.

Altruism is performed to better the survival potential of the group rather

than the individual. Many animals, like prairie dogs, will risk the notice
of a predator by sounding an audible alarm call. Others, like deer, use a
visual alarm call to warn other members of the social group that a predator
is near. But, in doing so, the individual sounding the alarm attracts the
notice of the predator, potentially risking one's own life to save the group.

When an altruistic act is performed for a member of one's own family it

is called kin selection. This selection of related animals can affect the
fitness of an individual. This increases the reproductive fitness (survival)

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of future generations. A gene that a particular individual carries passes to

the next generation through a related animal. Therefore, the fitness of an
individual is based on the genes it passes on. It is -also based on those
common genes that its relatives pass on. Therefore, altruism is a
genetically based tendency. It is passed on by the individual carrying it or
by a relative who also carries it. But the individuals of a group must
identify its relatives for kin selection to work. It is done by small groups
of primates and social insects. Lets consider few examples of altruism in
some animals;

a. Altruism in crows: One individual of a group of crows gives an

alarm call. It
warns the other individuals of the group of a predator. This call
may attract he predator to the sender of the signal.

b. Altruism in Honey bee: Altruism is also present in societies of

hymenopteran insects like honeybees. The male drones are
haploid. The female workers and queen are diploid. It develops
genetic asymmetry. Diploid workers share three fourths of their
genes with their full sisters. If they reproduced, they will share only
half of their genes with hypothetical offspring. Thus, female
honeybees have more genes common with their sisters than their
offspring. The workers help their mother to produce more sisters.
Thus the worker passes more genes to the next generation by their
mother. Some of their sisters become reproductive queens.

ii. Dominance and Hierarchy

The organization of group of animals in such a way that some members

of the group have greater access to resources like food or mates than others
is called dominance hierarchies. Some animals are present near the top of
the order. They have first choice of resources. The animals present near
the bottom do not get sufficient resources.

Many primate, canine and other social groups are organized through
a dominance hierarchy. The male or female in charge is the most
dominant, the alpha, the leader of the group, like the president. While
many groups are male dominant, some groups, like the spotted hyena and
bonobo, are female dominant.

Example is the Pecking (fight with beaks) order is an example of a

dominance hierarchy. Peck order is present in chickens in a pen. The
chickens are placed together. They fight among themselves. Finally, a
linear hierarchy of dominance is formed. Higher-ranked chickens are first
to eat. They peck lower-ranked chickens. Peaceful coexistence is possible

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after the setting of the hierarchy. Sometimes, a bird tries to move up in the
order. Therefore, occasional fights o Cur.

Dominance hierarchies also exist in many vertebrate groups. The most

common dominance hierarchy is present in the form of linear
relationships. Sometime, triangular relationships may be formed. The
strongest male is highest in the rank order in baboons. But sometimes,
older males may form coalitions. They subdue a stronger male and lead
the troop.

iii. Agnostic behaviour

The behavior in which one animal is aggressive or attacks another animal,

the other responds by returning the aggression or submitting is called
agnostic behavior. These behaviors are used to display one's fitness to a
challenger in an attempt to intimidate him or her into backing down, like
ritualistic fighting displays. A society of animals maintains social
structure. Agnostic behavior is lethal in rare cases. Usually the animals
are not killed or severely injured. Think of a dog growling and bearing its
teeth, or an elephant stomping its feet and flapping its ears.

The males show their aggression in the form of threat displays. The
aggression displays involve signals. It warns other males of an intention
to defend an area or territory. Agnostic behaviour seems antisocial. But it
maintains the social order. It is important in the maintenance of territories
and dominance hierarchies.

iv. Territory

The site defend by territorial animal by agnostic behaviour is called

territory of the animal. They excluded the competing individuals from this
site. Many male birds and mammals occupy a breeding territory. A male
actively defends his area against other males. He attracts a female in. his
territory and courts (mating) her without interference. Some
territories contain a food supply. Some territories provide shelter. It
protects the animal from predators and unfavorable climate. What is
Social behvaiour?
What is Altruism?

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. Write on agnostic behavior.
2. What is dominance hierarchies?

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3.4 Summary

Social behaviors in animals have both advantages and disadvantages.

Many animals’ species live in groups that provide various benefits.
Groups range from simple aggregations to more complex social
organization or societies.

3.5 References/Further Readings/Web Sources

Laidre, M. E. and Johnstone, R. A. (2013). Animal signals. Current

Biology, 23(18):829–833

Maynard Smith, W. J. (1977). The Behavior of

Communicating. Harvard, MA: Harvard University Press

Miller, S. A. and Harley, J. B. (1999) Zoology (fourth edition).

WCB McGraw- Hill Boston 750 pgs

Owings, D. H. and Morton, E. S. 1998. Animal Vocal Communication:

A New Approach. Cambridge: Cambridge University Press

Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift für

tierpsychologie, 20 (4): 410–433

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3.6 Possible Answers to SAEs

Answers to SAEs 1

1. The behavior in which one animal is aggressive or attacks another

animal, the other responds by returning the aggression or
submitting is called agnostic behavior. These behaviors are used to
display one's fitness to a challenger in an attempt to intimidate him
or her into backing down, like ritualistic fighting displays. A
society of animals maintains social structure. Agnostic behavior is
lethal in rare cases. Usually the animals are not killed or severely
injured. Think of a dog growling and bearing its teeth, or an
elephant stomping its feet and flapping its ears.

2. The organization of group of animals in such a way that some

members of the group have greater access to resources like food
or mates than others is called dominance hierarchies.

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Unit 4 Social Behaviour of Primates

Contents
4.1 Introduction
4.2 Intended Learning Outcomes (ILOs)
4.3 Social structure compositions among primates
4.4 Summary
4.5 References/Further Readings/Web Sources
4.6 Possible Answers to Self-Assessment Exercises

4.1 Introduction

Most primates, including humans, spend their lives in large social groups.
In the case of semi-terrestrial species, such as baboons, being in a large
community helps provide protection against predatory cats, dogs, and
hyenas. It also helps protect scarce food resources. This is especially
true for non-human primates when the food is fruit. Leaf-eaters, such
as colobus monkeys and langurs, tend to form smaller social groupings
since there is little competition for their food. The very few nocturnal
species of primates are mostly small, relatively solitary hunters.

As noted earlier, all primates (including us) form groups. Groups

involve such cohesive (bonding) activities such as grooming and
mother-infant bonding. It is a way to defend resources against intruders
and to fend off predators, such as leopards that inhabit the forests. Group
behavior among primates is the most complex among all gregarious
animals (animals that form groups, which also include gazelles, cattle,
bison, zebras, and others).

Most non-human primate communities are more or less closed to

contact with members of other communities. Most often, they are tied
to a particular locale and rarely migrate outside of their home range. This
aloofness from other troops prevents high concentrations of individuals
which could result in rapid depletion of local resources. Communities
usually avoid each other and are aggressive towards outsiders. As a
result, social interactions between members of different troops are usually
very rare, especially for females. Chimpanzees are a notable exception.
When chimpanzees from different troops come together, there is often an
exciting, friendly encounter lasting several hours, following which,
some of the adult females switch groups. Apparently, they are seeking
new mates.

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4.2 Intended Learning Outcomes (ILOs)

At the end of lecture the students should be able to;

 Understand the social structure among primate

 Identify different types of social behavior among primate

4.3 Social structure compositions among primates

Interactions within non-human primate communities are usually

unlimited. Subgroups are rarely closed from group interaction. All
members of a community have daily face to face, casual communication.
The most common type of subgroup are as follow:

i. Single female and her offspring

The single female and her offspring group pattern is rare for primates but
common for other mammals. It is found among the orangutans and
some of the small nocturnal prosimians (e.g., mouse lemurs and
galagos). The adult males lead their lives mostly alone. However, they
come together with females occasionally for mating. The males of
these species generally have large territories that overlap those of
several females. Both male and female children usually leave their
mother when they reach sexual maturity.

ii. Monogamous Family Group

Monogamous groups consist of an adult male and female with their

children. When they are grown, the children leave to create their
own nuclear families. While this group pattern is the most common one
for humans, it is rare for non- human primates. It is found among the
small Asian apes as well as some of the New World monkeys and
prosimians. Specifically, monogamous family groups are the common
pattern for gibbons, siamangs, titi monkeys, indris, tarsiers, and
apparently some pottos.

iii. Polyandrous Family Group

The smallest New World monkeys, the marmosets and tamarins, form
both monogamous and polyandrous family units. They generally start
with a monogamous mating pair. Later, a second adult male may
join the family and assist in child rearing. When this occurs, both

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adult males will potentially mate with the adult female. This
polyandrous mating pattern is extremely rare among non-human
primates but does occur in some human societies in isolated rural
regions of India, Sri Lanka, and especially Nepal, and Tibet.

iv. One-Male-Several Female group

One-male-several-female groups have polygynous mating patterns.

That is to say, one male regularly mates with more than one female.
Polygyny is generally not a promiscuous mating pattern. Rather, the male
and his female mates form a distinct mating and child rearing group. This
pattern is found among hamadryas baboons, geladas, langurs, howler
monkeys, gorillas and many human societies.

It would be a mistake to automatically assume that non-human primate

one-male- several-female groups are dominated by males. Among
geladas, females largely control the social group. This is despite the fact
that the males are larger, stronger, and more aggressive. Mothers, sisters,
and aunts act as a team in chasing off other unrelated females. They also
collectively select their mutual mate among a number of potential suitors
roaming in and out of their territory. The male that is chosen usually is
one that does not act abusively towards them and is willing to cooperate
with them in defending their territory. The relationship with any
particular male may be short-term. The stable core of the community
is the group of related females. This is a long way from stereotypical
male domination.

One-male-several female groups may take a different form when

predator pressure is a problem. In open grasslands, hamadryas baboon
communities are much larger, often consisting of a number of
polygynous families. In such multiple one-male- several-female group
societies, males are the dominant, controlling members. The adult males
not only "herd" their own sexually mature females, but also maintain
order and protect the community from predators.

In contrast, gorillas rarely have to be concerned about predator dangers.

Subsequently, their communities usually consist of a single dominant
adult male, his mates, and their children. When males reach maturity,
they usually are driven off by the dominant silverback male. These
exiled males ultimately form their own one-male-several-female groups.
As females reach sexual maturity, they also leave their natal families and
disperse. They later join with single males to form new families or they
join the families of males who already have mates. When the silver back
males have usually peaceful personalities, the gorilla community may
have several of them.

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v. Multimale- Multifemale Group

The most common social group pattern among semi-terrestrial primates

is the multimale-multifemale group. With this pattern, there are no
stable heterosexual bonds to both males and females have a number of
different mates. This is characteristic of savanna baboons, macaques, as
well as some colobus and New World monkey species.

Multimale-multifemale groups commonly have a dominance hierarchy

among both males and females. Each individual is ranked relative to all
other community members of the same gender. This tends to reduce
serious violence within the community since everyone knows in
advance who they must defer to and who must be submissive to them.
Among rhesus macaques, one's position in the dominance hierarchy is
determined by the rank of his or her mother. The top ranking
individuals are referred to by primatologists as the alpha male and
the alpha female. All other community members defer to them. A
female's rank in the hierarchy stays with her throughout life. However,
most young adult male rhesus macaques leave their natal community
and ultimately join others to find mates. When they do so, they start at
the bottom of the male dominance hierarchy again. Alpha males usually
mate more often than others. This makes the social organization
superficially look like one-male-several-female group. However,
younger females often sneak off to mate with males lower down on the
dominance hierarchy. The stable core of rhesus macaque communities is
the group of female relatives. They stay within their natal community
throughout life and work as a team to defend it against other females

vi. Fission-Fusion Society

A fission-fusion society is one in which the social group size and

composition change throughout the year with different activities and
situations. This is the social pattern typical of chimpanzees.
Individuals enter and leave communities from time to time. Adult males
occasionally wander off and forage alone or join a few other males in a
hunting party. Females casually change membership from one group
to the other. This occurs especially when females are in estrus and
seeking mates. As a result, foraging and sleeping groups reform
frequently. Male chimps are the relatively stable core of the community
since they rarely join other troops.

What allows for the generally loose relationship between chimpanzee

communities is that they apparently recognize a wider range of social
bonds than do monkeys. They often have relatives and friends in
several different neighboring troops. When chimpanzee communities
come together, they usually exchange friendly greetings rather than show

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aggression. However, it would be a mistake to assume from this that

chimpanzee society is always peaceful. The adult males within each
community are frequently engaged in complex political activities
involving scheming and physical intimidation in order to move up the
dominance hierarchy. They develop short-term alliances with other
males by mutual support, sharing meat, and allogrooming (grooming
others). It isn't always the largest and strongest males who make it to the
top of the hierarchy. Often teamwork used to frighten and impress is
more effective than any one individual's muscles in achieving
chimpanzee goals. This is an indication of their intelligence.

Chimpanzees are not the only primates that change group membership
from time to time. For instance, adult rhesus macaque males usually must
permanently leave the community of their birth and try to join others in
order to find mates. This is not easy since they are not warmly
welcomed in their adoptive troop. Group composition of some langur
and baboon species also change as a result of the availability of food and
mates. Evidently, none of these monkey species change group
composition with the ease and frequency of chimpanzees. As a result,
their societies are not usually referred to as fission-fusion types.

What is Fission – Fussion Society?

Self-Assemeement Exercises 1

Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. Write on Monogamous groups
2. What do you understand by Polyandrous family group?

4.4 Summary

Group behavior among primates is the most complex among all

gregarious animals (animals that form groups, which also include
gazelles, cattle, bison, zebras, and others). Primates live in social groups
that provide various benefits like food, protection, mates, learning of
skills, grooming and parental care.

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4.5 References/Further Readings/Web Sources

Laidre, M. E. and Johnstone, R. A. (2013). Animal signals. Current

Biology, 23(18):829–833

Maynard Smith, W. J. (1977). The Behavior of

Communicating. Harvard, MA: Harvard University Press

Miller, S. A. and Harley, J. B. (1999) Zoology (fourth edition).

WCB McGraw- Hill Boston 750 pgs

Owings, D. H. and Morton, E. S. 1998. Animal Vocal Communication:

A New Approach. Cambridge: Cambridge University Press

Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift für

tierpsychologie, 20 (4): 410–433
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4.6 Possible Answers to SAEs

Answers to SAEs 1

1. Monogamous groups consist of an adult male and female with

their children. When they are grown, the children leave to
create their own nuclear families. While this group pattern is
the most common one for humans, it is rare for non- human
primates. It is found among the small Asian apes as well as
some of the New World monkeys and prosimians. Specifically,
monogamous family groups are the common pattern for gibbons,
siamangs, titi monkeys, indris, tarsiers, and apparently some
pottos.

2. The smallest New World monkeys, the marmosets and tamarins,

form both monogamous and polyandrous family units. They
generally start with a monogamous mating pair. Later, a second
adult male may join the family and assist in child rearing.
When this occurs, both adult males will potentially mate with
the adult female. This polyandrous mating pattern is extremely
rare among non-human primates but does occur in some human
societies in isolated rural regions of India, Sri Lanka, and
especially Nepal, and Tibet.

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Unit 5 Herachical Organisation

Contents

5.1 Introduction
5.2 Intended Learning Outcomes (ILOs)
5.3 Dominance Hierarchies
5.4 Summary
5.5 References/Further Readings/Web sources
5.6 Possible Answers to Self-Assessment Exercises

5.1 Introduction

A dominance hierarchy is the organization of individuals in a group

according to their dominance. These hierarchies are often linear, each
individual dominating all individuals below him and not those above him.
The most dominant individual is often referred to as the alpha male (or
female), followed by the beta male and so on. These social structures were
first observed in chickens and are thus sometimes referred to as a “pecking
order”. The alpha male always eats first but also ensures that all members
of the group get something, even the least dominant. He has a
responsibility to ‘look after’ and make decisions for the group

The evolution of dominance hierarchies in a species is indicative that

there is competition for resources. Members of a dominance hierarchy are
aware of how they are positioned within that hierarchy and they behave
appropriately. Of particular importance, the establishment of dominance
hierarchies allows for the resolution of conflict between individuals
without costly fighting that can result in serious injury or even death. In
species where organized group living is essential to survival, it also serves
to maintain order among pack members

5.2 Intended Learning Outcomes (ILOs)

At the end of lecture the students should be able to;

 Explain dominance hierarchy

 Identify different ty

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5.3 Dominance Hierarchies

Dominance hierarchies characterize many species in which individuals

live in close proximity to one another. The dominance hierarchy is a social
structure within a group of animals in which certain individuals are
dominant over others, and are therefore able to claim access to better
resources in the form of food, mates, shelter, and other desirable
commodities.

Those near the top of the order have first choice of recourses, whereas
those near the bottom go last and may do without if resources are in short
supply. An example of dominance hierarchy is the pecking order of
chickens in pen. When chickens are placed together, they fight among
themselves until a linear hierarchy of dominance is established. Higher-
ranked chickens are among the first to eat and may peck lower–ranked
chickens. Once the hierarchy is set, peaceful coexistence is possible.
Occasional fights will occur if bird tries to move up in the order.

i. Establishment of Dominance Hierarchies

Dominance hierarchies are often established through ritualized displays

or mild fighting, rather than all-out battle. The loser in a battle for
dominance typically moves away from a choice habitat or a disputed
mate. Among primates, dominance conflicts frequently involve no more
than the display of enlarged canines, sometimes through yawning. Bears,
also, will roar or wave their open mouths at social inferiors. Behaviors
like these do not require fighting, but do result in the prominent exhibition
of potentially formidable fighting weapons. In other cases, as in elephant
seals, there actually can be prolonged, often bloody fighting. However,
once the hierarchy is established, subsequent fighting is less frequent. In
many cases, there is a strong correlation between dominance and large
size.

Dominance hierarchies exist in many vertebrate groups, the most

common being in the form of linear relationships, although triangular
relationships may form. In baboons, the strongest male is usually highest
in the rank order. But sometimes, older males may form coalitions to
subdue a stronger male and lead the troop.

All primates have dominance hierarchies of some short. One chimp

asserts its lordship over another individual. Chimps do shift in who
is the top chimp. In Gombe, Freud was the alpha male, then his brother
Frodo use bullying to displace Freud, who was getting ill. Then, having
had enough bullying, the other chimps kicked out Frodo. Baboons have

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more rigid hierarchies. Bonobos (below) have another system. The

females form a hierarchy, and they pass it down to their sons. Yet,
ironically, sisterhood is not possible. For one thing, there is no
sisterhood. Females leave their troops to join another one, and the
hierarchy is formed among total strangers.

Dominance hierarchies have to be reestablished when certain individuals

feel prepared to move up within the hierarchy, or when new individuals
are introduced into an area. During such time a series of challenges may
occur. This can be a stressful period for all individuals involved.

ii. Dominance and Mate Competition

Mate competition is extremely common in the animal kingdom, and many

dominance hierarchies relate to competition either for mates, or for those
resources such as admirable territories that will attract them. In most cases
males compete for females, although there are also a few instances of
females fighting for males.

There are clearly advantages to dominance. Dominant males have been

shown in many species to copulate more frequently or to produce more
off-spring. In cowbirds, for example, only the dominant male is allowed
to sing the songs that are most effective in attracting females. If
subordinate males attempt to sing these highly charged songs, they are
attacked, often brutally, by more dominant individuals.

Elephant seals are another group in which reproductive success is linked

to dominance. Dominance battles in this species involve two males
posturing chest to chest and attempting to bite each other, with the loser
ultimately retreating. In a few species, such as wolves, the dominant
members of a group are the only ones that reproduce.

One tell-tale sign of competition for mates is sexual size dimorphism,

which describes a situation where one sex of a given species has much
greater body size than the other. In the case of mate competition, it is the
males that are larger than females. (There are other species where the
females are larger, including, the large majority of frogs. However, in
these species the large size of females appears to be associated with
increased fertility rather than with the establishment of dominance.)

Sexual size dimorphism is often particularly pronounced in species where

it is possible for a single male to monopolize many females, as in elephant
seals. In fact, a study across various pinniped species (seals, sea lions,
etc.) suggests that the degree of sexual size dimorphism is positively
correlated with the size of the harem.

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iii. Dominance and Submissive Displays

Even simpler behavioural displays are often use to maintain a dominance
hierarchy. Agonistic displays are used to assert dominance. These often
include a more upright, aggressive stance. For instance, this first pukeko's
upright stance and raised wing is a dominance display. Subordinate
individuals (lower in the hierarchy) will respond with submissive displays
(appeasement gestures) such as making themselves look small, bowing
the head and exposing vulnerable parts. The second pukeko's bowed head
is a submissive display. When young dogs roll onto their back, this is also
a submissive /appeasement gesture. The young dog is exposing its
vulnerable areas to communicate that it is no threat to your dominance.

iv. Dominant hierarchical behaviour in spotted hyena

A particularly interesting example of the dominance hierarchy is that of

the spotted hyena. It is the largest species of hyena and has also been
called the laughing hyena because of the calls that individuals make when
they are in danger. Spotted hyenas live in social groups that vary greatly
in size, with the largest having as many as eighty members. Each group
defends a territory and hunting occurs in packs.

What is unusual about social organization in this species is that females

are dominant within the group and at the same time possess reproductive
organs that very much resemble those of males. In fact, female genitalia
resemble the scrotum and testes of males so closely that it is almost
impossible to determine the sex of individuals in the field.

One early hypothesis to explain this male-mimicking anatomy was that

females evolved it in order to participate in the hyena greeting ritual, in
which members of the same social group sniff each other’s' erect penises
when they meet again after an absence. Because greeting behaviour is
important to group solidarity, it was argued that females evolved male-
like anatomy so they could participate as well.

However, the greeting ritual theory has since been abandoned in favour
of an argument based on fighting for dominance within the hierarchy.
There are numerous benefits to being the dominant female within a
spotted hyena clan. Females who are high in the hierarchy have priority
at kills, and obtain more food than subordinate females or males.
Dominant females tend to be the largest hyenas of a pack. They also tend
to produce dominant off-spring. The production of a dominant male is
particularly advantageous because only the dominant male within a pack
mates.

Many scientists believe that because aggressive behaviour is

advantageous in competitions for dominance, female hyenas have
evolved high circulating levels of androgens (male sex hormones) such as

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testosterone, which promote aggression. The curious male-mimicking

genitalia are now believed to be a mere side effect of the unusually high
testosterone levels. The testosterone circulating in the female's
bloodstream while she is pregnant results in the masculinization of the
anatomy of both her male and female offspring. It was indeed confirmed
that female spotted hyenas do in fact have unusually high testosterone
concentrations in their blood.

v. Dominant hierarchical behaviour in wolves

Most social mammals have some form of dominance hierarchies. Perhaps

the most famous example of dominant hierarchical behavior is wolves.
White wolves are displayed here, exhibiting the effects of their dominance
hierarchy. One can see that a single wolf is lying down on the highest
ground available. The alpha wolf and his chosen alpha female mate for
life and are the only group members to procreate. All pack members are
responsible for looking after the pups and providing them with food,
water, and shelter. While many species, like lions, will kill the young
offspring of dethroned alphas, wolves always adopt the pups as their own
and raise them to maturity. This altruism often creates better pack unity
and shows wolves' strong social bonds within these dominance
hierarchies.

Domesticated chickens are another example of dominant behavior. Even

in the absence of a rooster, hens will create a pecking order based on their
physical and personality characteristics. The more dominant hens receive
preference on roosting positions and access to food and water, which
prevents hens from having constant disputes over resources.

In-Text Question(s)

What is dominance hierarchy?

Answer: A dominance hierarchy is the organization of individuals in a
group according to their dominance. These hierarchies are often linear,
each individual dominating all individuals below him and not those above
him. The most dominant individual is often referred to as the alpha
male (or female), followed by the beta male and so on.

Self-Assessment Exercises 1

Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. How is Dominance Hierarchies is established?

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5.4 Summary

Many animal species live in groups that provide various benefits.

Groups range from simple aggregations to more complex social
organizations or societies. Some animals organize themselves in
hierarchies which rank members in order from the most dominant
individual to the most subordinate individual. Once the hierarchy is
established, agonistic behavior is reduced in the group.

5.5 References/Further Readings

Alcock, John. (1989). Animal Behavior, 4th ed. Sunderland, MA: Sinauer
Associates.

Gould, James L., and William T. Keeton. (1996). Biological Science, 6th
ed. New York: W. W. Norton.

Halliday, Time. (1994). Animal Behavior. Norman: University of

Oklahoma Press.

Krebs, John R., and Nicholas B. Davies.(1997). Behavioural Ecology:

An Evolutionary Approach,4th ed. Cambridge, MA: Blackwell
Science.

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5.6 Possible Answers to SAEs

Answers to SAEs 1

1. Dominance hierarchies are often established through ritualized

displays or mild fighting, rather than all-out battle. The loser in a
battle for dominance typically moves away from a choice habitat
or a disputed mate.

Glossary

°C = degrees Celsius
cm = centimeters
CNS = Central nervous system
CH4 = Methane
CO = Carbon
CH4 = methane (CH4)
DNA = Deoxyribonucleic acid
E. coli = Escherichia coli
FAP = Fixed Action Pattern
°F = degree Fahrenheit
Ft = feet
h = hour
in = inches
Kgs = kilograms.
km = kilometers
kph = kilometer per hour
m = meters
mm = millimeters
mph = meter per hour
NH2OH = Hdroxylamine
NH3 = Ammonia
O2 = Oxygen
PNS = Peripheral Nervous System
PRC = Phase Response Curve
RNA = Ribonucleic acid
L = length
HIV = Human immunodeficiency virus
% = percentage
g = grams
spp = species
UV = Ultraviolet

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End of the Module Questions

1. What is conflict behavior?

2. What is habituation?
3. List the different types of communication in animal?
4. What is pheromones?
5. What is a fission – fusion society?

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Module 3 Habitat Selection, Homing and Navigation

Module Introduction

In Module three, unit one deals with the physiology of animal behavior,
unit two deals with the different types of habitats, history and current
understanding of Animal Ecology and how organisms and its
environment relate and influence one another in their various ecosystems.
You are taught about the fundamentals of ecology; interaction in animals
and energy flow within the environment.

Unit 1 The Physiology of Bahviour

Unit 2 Habitat Selection
Unit 3 Homing and Navigation in Birds
Unit 4 Courtship bahviour and Parenthood
Unit 5 Biological Clocks

Glossary

Unit 1 The Physiology of Bahviour

Contents

1.1 Introduction
1.2 Intended Learning Outcomes (ILOs)
1.3 The Physiology of behaviour
1.4 Summary
1.5 References/Further Readings/Web sources
1.6 Possible Answers to Self-Assessment Exercises

1.1 Introduction

Behavior is the varied activities that an animal perform during its

lifetime. Internal physiological conditions, environmental stimuli and
social situations influence specific behavioral responses. Animals are
faced with two key problems finding food and a place to live. The
evolution of various social systems, in which animals live in groups,
affects many aspects of their behavior.

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Animal behavior refers to the activities animals perform during their

lifetime, including locomotion, feeding, breeding, capture of prey,
avoidance of predators and social behavior. Animals send signals,
respond to signals or stimuli, carry out maintenance behavior, make
choices and interact with one another.

1.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Explain physiology of animal behaviour

 Expalin the four approaches to animal behaviour
 Defne anthropomorphism
.

1.3 The Four Approaches to Animal Behavior

Naturalists and philosophers have observed animal behavior for

centuries. Only in the last century, however has there been significant
progress in understanding this behavior.

One approach to the study of animal behavior is comparative

psychology. Comparative psychologists emphasize studies of the
genetic, neural and hormonal bases of animal behavior. Psychologists
conduct experimental studies in both laboratory and field settings, that
relate to animal learning and to development behavior. They explore how
animals receive information and the processes and nature of the behavior
patterns constituting the animals’ responses to the surroundings.

Ethology is the study of animal behavior that focuses on evolution and

natural environment. The leaders of this approach have been Konrad
Lorenz, Niko Tinbergen and Karl von Frisch, who were awarded the
Noble price in physiology or Medicine in 1973. Ethologists observe
the behavior of a variety of animals in their natural environment and
study the behavior of closely related species to consider the evolution
and origin of certain behavior patterns. Ethologists rarely deal with
learning and are intereste instead in animal communication, mating
behavior and social behavior.

Behavioral ecology emphasizes the ecological aspects of animal

behavior. Predator-prey interaction, foraging strategies, reproductive
strategies, habitat selection intraspecific and interspecific competition
and social behavior are topics of interest to behavioral ecologists.

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Sociobiology is the study of the evolution of social behavior. It combines

many aspects of ethology and behavioral ecology. Sociobiologists
emphasize the importance of natural selection on individuals living in
group.

There are two major causes of behavior namely PROXIMATE

AND ULTIMATE CAUSES

a. Proximate causation - immediate causes. This explains how

behavior works and what stimulates behavior to occur. It could
be studied by measuring or describing the stimuli that elicit
behavior. It involves Internal - physiological events (hormones,
nervous system) and External - environmental stimuli like
changes in daylength.

b. Ultimate causation - historical explanations: this explains why a

behavior evolved. It is studied by measuring influence on
survival or reproduction Example; bird migration - birds that
migrate have a selective advantage over birds that don't/didn't,
selected for over time, could be due to long term climate changes,
glaciation, disease, taking advantage of food sources, etc.

Behavioral scientists frequently ask, “Why do animals do what they do?”

more immediate ecological and physiological causes of behavior,
such as eating to satisfy hunger, are called proximate causes. Another
level of causation in behavior occurs on the evolutionary time scale and
that is of ultimate causes. For example, a display not only attracts a male,
but also increases the likelihood of passing genetic information to the
next generation.

i. Timing of behavior

Circaannual - behavior occurs on a seasonal/annual basis. Examples is

the hibernation in bears, frogs, toads, salamanders bury themselves in
mud during the winter, while Circadian - behavior occurs on a daily basis.

i. Components of Behavior

There are two Components of behavior and they are;

1) Innate behavior: instinct and genes determine behavior

2) Learned behavior: experience and learning influence behavior

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The two components of behaviour are not mutually exclusive, but work
together to influence behavior. Examples of Innate behavior are;

a. The Nest building in Lovebirds by Dilger such as;

- Fischer's Lovebird - uses long strips of nest materials, carries in
beak, one at a time
- Peach-faced Lovebird - tucks several short strips in feathers
- Hybrids - intermediate lengths of nest materials, clumsy behavior
trying to tuck strips into feathers, later will carry strips in bill
but will still try and tuck into feathers

b. Egg ejection by cuckoos (brood parasites)

c. Freezing behavior of nestling birds when exposed to silhouettes
(raptors versus waterfowl)
d. Parental feeding - brood parasites take advantage of parents
e. Freezing behavior of nestlings
f. Incubation behavior of some birds (Oystercatchers)
g. Drosophila - 2 alleles of the Dg2 gene sitter allele (sedentary
behavior) rover allele (hyperactive, mobile)

Components of Innate behavior is the fixed action pattern, all or none

response and

Sign stimulus which causes release of FAP. Examples are colors of

stickleback males during mating, oyster catchers and eggs during
incubation (super-normal releaser). The Nature of sign stimulus are
usually an obvious aspect of the morphology such as red mark on beak
of Herring Gulls, red belly of Sticklebacks, detection of ultrasounds
from bats by prey species of moths.

2. Learned Behavior

a. Simple learning - habituation, species of prey and the presence of

predators. Lehrman's study of gull chick feeding behavior - how
an instinct is learned

b. Learning and development - imprinting and Lorenz's classic

experiments with Greylag Goose (critical period for learning)
- geese forms social attachments shortly after birth , salmon
and home stream, birds and breeding range, nesting materials,
etc.

c. Sexual imprinting - Direct sexual behavior at member of one’s

own species - cross-fostering studies, individuals raised by
another species, recognizes foster species as its own when
sexually mature, will attempt to mate with foster species.

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d. Classical/Pavlovian conditioning;- Animals make associations

such as Pavlov's dog associates bell with food, begins to
salivate, can be extinguished and later followed by recovery
(unconditioned stimulus - meat, unconditioned response -
salivation, conditioned stimulus - bell, conditioned response –
salivation).

e. Operant conditioning- Reward/punishment for behavioral

response, rats bar press for food.

f. Observational learning - social imitation

g. Insight Learning – the ability to respond correctly to a situation

that is experienced for the first time and that is different from experience
encountered previously. Examples are Chickadees/tits and opening milk
bottles, Egyptian Vulture - uses rocks, Cocos Finch - uses splinters of
wood, North American Gulls, Northwestern Crow - smash clams on
sandy beaches

1.4 Anthropomorphism

Anthropomorphism is the application of human characteristics to

anything not human. In observing animals, assigning human feelings
to animal behavior is not likely to be accurate, especially with
invertebrate animals. Consider the example of placing an earthworm, on
a fishhook. Does the fishhook hurt the earthworm, causing it to writhe
in pain? Both of the descriptive words hurt and pain, are based on
human experience and conscious awareness. A better explanation that
reduces the anthropomorphic interpretation is that placing the
earthworm on the hook stimulates certain receptors which generate nerve
impulses that travel along reflex neural circuits. The impulses stimulate
muscles that allow the worm to wriggle in an attempt to escape from
the hook. This explanation more closely describes what has been
observed and does not attempt to suggest what earthworm “feels.”
What is Behaviour? What is Circaannual?

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is Insight Learning?
2. What is Anthropomorphism?

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1.4 Summary

Many behavior patterns require instinctive and learned components for

efficient performance. In some instances, an animal may inherit a
disposition to learn a specific behavior. Also, an animal may learn certain
behavior patterns only during a specific sensitive period early in life.

1.5 References/Further Readings/Web Sources

Campbell, N.A (1987): Biology. The Benjamin/Cummings publishing

company, Inc, California 1101pp

Miller, S.A. and Harley, J.B. (1999): Zoology (Fourth Edition). WCB
McGraw- Hill, Boston
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1.6 Possible Answers to SAEs

Answers to SAEs 1

1. Insight Learning is the ability to respond correctly to a situation

that is experienced for the first time and that is different from
experience encountered previously.

2. Anthropomorphism is the application of human characteristics

to anything not human.

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Unit 2 Habitat Selection

Contents

2.1 Introduction
2.2 Intended Learning Outcomes (ILOs)
2.3 Habitat selection
2.4 Summary
2.5 References/Further Readings/Web Sources
2.6 Possible Answers to Self-Assessment Exercises

2.1 Introduction

Habitat selection is the choice by an organism of a particular habitat in

preference to others. e.g may fly nymphs inhabit the underside of
stones in fast-flowing streams and burrow in sediment in still water
while habitat is a place where species get what they need to survive:
food, water, cover, and a place to raise young. In other words, a habitat
is a plant or animal's home. For people, habitat might stretch from their
home (where they have water, cover and a place to raise young), to the
supermarket (where they buy food). All the places people go to get what
they need to survive can be considered part of their habitat.

2.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Define habitat and habitat selection

 Explain the different types of habitat.

2.3 Types of Habitat

The following are different types of habitats: Polar/arctic areas,

Mountains, Oceans, Deserts, Savannah/grasslands/prairies, Tropical
rainforest, Woodland/forest, Tundra, Taiga, Wetland areas/marshes,
Pond Rivers/lakes, Coral reef, Deciduous forest, Tide pool Cave

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i.) Forest habitat

Forests are fascinating ecosystems. The defining feature of a forest is its

dense growth of trees. Generally speaking, two key variables dictate the
geographical distribution of Earth's different habitat types: Humidity and
temperature. Forests grow where there is enough water available to
fulfill trees' needs. The extent of forest growth also depends on
temperature ranges, soil nutrients, adequate growing season and altitude.

All of the forests in the continental United States are temperate forests
(located between the boreal and sub-tropical zone). Eastern temperate
forests tend to have cold winters and wet, hot summers. Deciduous trees
(those that lose their leaves in the fall) like oak and maple thrive in these
conditions. In fact, most eastern forests are defined by the mix of oak,
maple, birch and other trees that grow there. These trees create a canopy
that shades the forest floor and provides a variety of habitats.

For many creatures, such as gray squirrels, white-footed mice, white-

tailed deer, blue jays. Deciduous trees are also found in continental
Africa and Nigeria Southwest.
Generally speaking, deciduous trees dominate the forests of the Eastern
United States, while coniferous trees (those that keep their leaves year-
round) predominate in western forests. What kind of wildlife would you
expect to live in the forests Western United States?

ii.) Grasslands habitat

Grasslands are characterized as areas where grasses are the predominant

vegetation and the subsoil is dry with seasonal moisture in the upper
soil layers. Their evolution was shaped by periodic fires and the
presence of grazing animals. These conditions resulted in the
establishment of vast areas of grassland on all of the continents except
Antarctica. Today, a quarter of the earth's land surface remains covered
by this rapidly vanishing ecosystem. Example in Nigeria is Osun State
extending to part of kwara / kogi.

All grasslands share several common traits. In general, the term

grassland refers to land which: is dominated by grasses; occurs on flat
or rolling terrain; has similar soils (alkaline, lots of organic matter, very
fertile, and fine- grained); has soil that is almost completely covered by
vegetation, commonly has fires and high winds (which lead to high
evaporation rates and the spread of fires); is characterized by periods of
rain followed by periods of drought.

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iii.) Desert habitat

As different from one another as deserts of the world are, they all
share one characteristic: they are very dry. Scientists define deserts as
areas that get less than 10 inches of rainfall a year and have a very high
rate of evaporation. If the annual evaporation rate of an area is higher
than the annual amount of rainfall, the area is considered a desert.
Evaporation rates are high because deserts tend to have very little cloud
cover and strong winds.

Another characteristic of deserts is sporadic rainfall. If the limited

rainfall in deserts fell a little at a time throughout the year, many deserts
probably would not look much like deserts. Instead, they'd have a lot
more vegetation. Rain doesn't fall evenly throughout the year in a desert,
though. It usually comes in big bursts. In some deserts, none at all may
fall for more than a year. And then a huge thunderstorm may dump over
5 inches all at once!

Deserts have some of the most variable temperatures of any places

on earth. Because the desert skies are nearly cloudless, the temperatures
during the day may sizzle. But without cloud cover to hold in the heat, it
radiates into the atmosphere very quickly once the sun goes down. In
some deserts, the temperature may drop as much as 77 degrees
Fahrenheit in 12 hours. For example Bauchi and Maiduguri.

iv.) Wetlands habitat

As the name implies, wetlands are areas where water is present at least
part of the year, generally for at least a portion of the plant-growing
season. In addition, wetland soils differ considerably from nearby or
surrounding uplands. Hydric soils, found in wetlands, are wet, low in
oxygen, and often black with muck. Finally, wetlands support plants —
called hydrophytes — that are adapted to living in wet, oxygen-poor
soils. Together, these water, soil and vegetation characteristics make
up a broad definition for wetlands.

Though all wetlands contain water at least periodically, the volume of

water and the amount of time a wetland is "wet" varies greatly. They
also vary in size, from wading-pool sized vernal pools to thousands of
acres along coastlines or rivers.

Wetlands are found all over North America, along coastlines, far inland,
in rural areas, and even in the middle of well-populated urban areas.

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There are generally five kinds of areas where we find wetlands:

a.) rivers;
b.) near coasts and inland lakes;
c.) in depressions where land is low compared to
surrounding landscapes;
d.) areas where groundwater seeps out of the ground, and;
e.) in broad, flat areas that receive significant rainfall (such as
the Everglades).

v.) Arctic Tundra habitat

The arctic tundra is circumpolar, meaning that it is an ecosystem

surrounding the polar region, above roughly 60 degrees north latitude.
The Arctic Circle occurs at 66 degrees north latitude.
In the tundra, short days for much of the year and a harsh cold climate
result in a brief growing season of 50-60 days. By contrast, the growing
season in temperate forests is about six months long and in tropical
forests lasts the entire year.

Strong winter winds challenge the stability of any plants that grow
more than an inch or two above ground surface. Below a thin layer of
soil that thaws every summer is ground that remains frozen year-
round, called permafrost. The permafrost may be very deep,
reaching more than 1000 feet thick in some locations. Although the
tundra receives less than ten inches of precipitation each year (which
is why it is sometimes referred to as an arctic desert), there can be
plenty of standing water when the upper layer of soil thaws each summer.

Due to its high latitude and the tilt of the earth, the arctic experiences
light and temperature extremes throughout the calendar year. The plants
and animals of the tundra must be adapted to face these challenges,
including not only extremes of day length and temperatures, but also
harsh winter winds, long periods of below- freezing temperatures, and
permanently frozen ground.

2.3.1 Habitat selection by some animals

African Elephant are Found in forests, grasslands, marshes, scrub, and

semi- desert areas. Elephants live in a highly organized social structure
referred to as a matriarchal herd. The herd is typically composed of
up to ten females and their young. All of the females in the herd are
directly related to the matriarch who is typically the oldest and largest
female. Males beyond the age of maturity are with the herd only during
mating.

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Penguin tend to inhabit islands and remote landmasses that are

relatively free from land predators. Some species spend nearly 75% of
their life at sea. All penguins live south of the equator, from the icy
waters of Antarctica to the tropical Galapagos Islands off the coast of
Ecuador, almost astride the equator. Penguins are specialized marine
birds adapted to living at sea. Some species spend as much as 75% of
their lives in the sea - only coming ashore for breeding and molting.
Penguin wings are paddle-like flippers used for swimming, not flying.

Sea Stars are found from the bearing sea, usually resting on broken or
solid rocks. They are usually found on gravel, rocks and sand in the low
intertidal zone.

What is habitat selection? What is Grassland Habitat?

Self-Assessment Exercises 1

Attempt these exercises to measure what you have learnt so far.

This should not take you more than 5 minutes.
1. What is Desert?
2. List the five kinds of areas where wetland is found?

2.4 Summary

Habitat selection is the choice by an organism of a particular habitat in

preference to others. e.g may fly nymphs inhabit the underside of
stones in fast-flowing streams and burrow in sediment in still water
while habitat is a place where species get what they need to survive:
food, water, cover, and a place to raise young.

2.5 References/Further Readings/Web Sources

Miller, S.A. and Harley, J.B. (1999): Zoology (Fourth Edition). WCB
McGraw- Hill, Boston

Chapman JL and Reiss MJ (1995): ECOLOGY, Principles and

[Link] University Press, UK

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[Link]
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saXBzLXZpZGVvL2tzMS1zY2llbmNlLWhhYml0YXRzLWVudmlyb2
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[Link]
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[Link]
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2.6 Possible Answers to SAEs

Answers to SAEs 1

1. Scientists define deserts as areas that get less than 10 inches of

rainfall a year and have a very high rate of evaporation.
2. There are generally five kinds of areas where we find wetlands:
a.) rivers;
b.) near coasts and inland lakes;
c.) in depressions where land is low compared to
surrounding landscapes;
d.) areas where groundwater seeps out of the ground, and;
e.) in broad, flat areas that receive significant rainfall (such as
the Everglades).

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Unit 3 Homing and Navigation

Contents

3.1 Introduction
3.2 Intended Learning Outcomes (ILOs)
3.3 Factors affecting initiation of Navigation / Migration
3.3.1 Orientation and navigation
3.3.2 Navigation Methods in Animals
3.4 Summary
3.5 References/Further Readings/Web sources
3.6 Possible Answers to Self-Assessment Exercises

3.1 Introduction

Homing is the ability of certain animals to return to a given place when

displaced from it, often over great distances. This may occur in any
compass direction and at any season. Navigation clues used by homing
animals are the sun angle, star patterns etc. very strong homing ability
are found among birds seabirds and swallows eg. A Manx Shearwater
(Puffinus puffinus) transported in a closed contain to a point about
5,500 km (3,400 miles) from its nest and returned to the nest in 12 ½
days.

Navigation means migration which is the movements of animals in large

numbers from one place to another. In modern usage the term is usually
restricted to regular, periodic movements of populations away from and
back to their place of origin. A single round trip may take the entire
lifetime of an individual, as with the Pacific salmon salmon, member of
the Salmonidae, a family of marine fish that spawn in freshwater,
including the salmons, the trouts, and the chars. Many authorities place
the whitefish and the grayling among the Salmonidae, so similar
are they in structure and habits. An individual may make the same
trip repeatedly, as with many of the migratory birds and mammals.
The animals may travel in groups along well-defined routes; or
individuals may travel separately, congregating for breeding and then
spreading out over a wide feeding area.

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3.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Define homing and navigation.

 Explain navigation mechanism in animals.

3.3 Factors affecting initiation of Navigation/Migration

Various factors determine the initiation of migration. In some cases

external pressures—temperature, drought, food shortage—alone may
cause the animals to seek better conditions. For example, most of the
mule deer of Yellowstone Park, Wyo., migrate between summer and
winter pastures, but those living near hot springs, where grazing is
available all year, do not. In many species migration is initiated by a
combination of physiological and external stimuli. In birds the migratory
instinct is related to the cycle of enlargement of the reproductive organs
in spring and their reduction in fall. Experiments have shown that
variation in day length is the chief external stimulus for this cycle: light
received by the eye affects production of a hormone by the anterior
pituitary gland, which stimulates growth of the reproductive organs.

3.3.1 Orientation and navigation

Much work has been done on orientation and navigation in migrating

animals, although the subject is still not well understood. Studies of
salmon indicate that they depend on the olfactory sense to locate and
return to their stream of origin. Herbivorous mammals often follow
well- established trails and probably also use their sense of smell. Bats,
whales, and seals use echolocation to navigate in the dark or underwater;
in addition, some whales appear to take visual bearings on objects on the
shore in their migrations.

Migratory birds are believed to use the stars, sun, and geographic
features as guides. The probability that stellar navigation is used has
been strengthened by experiments in planetariums indicating that birds
navigate at least in part by the stars. Night-migrating birds are
sometimes disoriented in prolonged heavy fog. Day-flying birds
navigate by the sun and also make some use of geographic features,
particularly of shorelines. It has long been proposed that birds perceive
the direction of the earth's magnetic field and use it for navigation,
but experimental evidence for that hypothesis is inconclusive. Most

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migratory birds travel within broad north-south air routes known as

flyways. There are four major flyways in North America, called the
Pacific, central, Mississippi, and Atlantic flyways. The space within the
flyway used by a particular group of birds is called a corridor. Bird
migration is not always in a north-south direction. Many European birds
migrate in an east-west direction, wintering in the more temperate
British Isles, and many mountain-dwelling birds descend to lower
altitudes in winter. The breeding grounds of a bird species are regarded
as its home territory. Some migratory birds winter only a few hundred
miles from their breeding grounds, while others migrate between the
cold or temperate zones of the two hemispheres. The longest journey is
made by the arctic tern, common name for a sea bird of the Old and New
Worlds, smaller than the related gull. Because of their graceful flight and
their long pointed wings and forked tails, some terns are called sea
swallows. They plunge headlong into the water to catch small fish.
The monarch butterfly has a north-south migration pattern that resembles
that of many birds. One monarch population that inhabits northeastern
and midwestern North America averages c.12 mph (19 kph) as it heads
for the winter to Mexico's Sierra Madre mountains. Monarchs start the
return trip in the spring, but they breed along the way and then die; the
new generation completes the journey.

There are various tools for studying migration. The movements of

migrating animals are often studied by tagging individuals. Bird banding
has been carried on extensively since the 1920s; more recently there has
been tagging of fishes, butterflies, and marine mammals. Use is now
made of radar, sonar, and radio for following migrations, particularly
those of marine animals. Radio transmitters attached to whales or seals
emit signals that can be picked up by weather satellites at regular
intervals.

3.3.2 Navigation Methods in Animals

i.) The sun : starlings and ants navigate this way. Some birds can
travel at night using the sun – theories suggested that they take a
‘reading’ from where the sun sets and use that to est their course.
ii.) Landmarks: fly towards those mountains head to the left a little
when you see the ocean, nest in the first nice, looking tree you can
find. E.g Whales traveling in the pacific ocean near the North
American West.
iii.) Moon and stars: Planetarium experiments’ have proved that many
birds rely on stars cues to figure out which way to migrate e.g
indigo.
iv.) Scent: scent can pin point specific location. E.g Salmon find their
exact spawning ground through scent.

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v.) Weather: wind condition are often used as supplementary

navigation aids by [Link]
vi.) Magnetic field: the earth has a magnetic field that is usually
undetectable to human who are not holding a compass. Bat and
sea turtles use magnetic information to find their way. Bacteria
even rely on the magnetic field to orient them.

What is Homing? What is Navigation?

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is the space within the flyway used bhy a particular
group of birds called?
2. 3.4
List the four major flyways in North America?
Summary

In animal behavior homing and navigation / migration are important

issues that cannot be overemphasis. This topic had being dealt with but
more information are still needed in the area of orientation and
navigation. There are various tools for studying migration. The
movements of migrating animals are often studied by tagging individuals.
Bird banding has been carried on extensively since the 1920s; more
recently there has been tagging of fishes, butterflies, and marine
mammals.

3.5 References/Further Reading/Web Sources

Miller, S.A. and Harley, J.B. (1999): Zoology (Fourth Edition). WCB
McGraw- Hill, Boston

Moore, F.R. (1987). Sunset and the orientation behavior of

migratory [Link] Reviews, 62, 65-86.

MacDonald, David. The Encyclopedia of Mammals: 2. London: George

Allen & Unwin Co., 1985.

Mouritsen, H., & Frost, B.J. (2002). Virtual migration in tethered flying
monarch butterflies reveals their orientation mechanisms.
Proceedings of the National Academy of Sciences of the United
States of America, 99, 10162-10166.

Wiltschko, R., & Wiltschko, W. (1981). The development of sun

compass orientation in young homing pigeons. Behavioral
Ecology and Sociobiology, 9, 135 - 141.
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Wiltschko, W., & Gwinner, E. (1974). Evidence for an innate magnetic

compass in garden warblers. Naturwissenschaften, 61, 406.

Wiltschko, W., & Wiltschko, R. (1972). Magnetic compass of

European robins. Science, 176, 62-64.

Wiltschko, W., & Wiltschko, R. (1975). The interaction of stars and

magnetic field in the orientation systems of night migrating
birds. II. Spring experiments with European robins (Erithacus
rubecula) Zeitschrift für Tierpsychologie, 39, 265-282
[Link]
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3.6 Possible Answers to SAEs

Answers to SAEs 1

1. The space within the flyway used by a particular group of birds

is called a corridor.
2. There are four major flyways in North America, called the Pacific,
central, Mississippi, and Atlantic flyways

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Unit 4 Courtship Behaviour and Parenthood

Contents
4.1 Introduction
4.2 Intended Learning Outcomes (ILOs)
4.3 Courtship
4.4 Summary
4.5 References/Further Readings/Web Sources
4.6 Possible Answers to Self-Assessment Exercises

4.1 Introduction

Courtship is the period in a couple’s relationship which precedes

their engagement and marriages or establishment of an agreed
relation of a more enduring kind. A courtship may be an informal
and private matter between two people or may be a public affair or a
formal arrangement with family approval. The average duration of
courtship varies considerably throughout the world, this depends on
individual.

Parenthood means parentage i.e. the state of being a parents. Parenthood

is the intersection of two distinct relationships, that between parents
and child, and that between the parents (or family) and the larges society
or other collective. This is called relationship. Parenting is the processs
of promoting and supporting the physical, emotional, social and
intellectual development of a child from infancy to adulthood.

4.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Expalin courtship behavior in animals

 Identify the different forms of courtship behavior

4.3 Courtship

Many non-human animal species have mate-selection rituals also

referred to as ‘courtship’ in an anthropomorphic (misleading) manner.
Animal courtship may involve complicated dances, or touching,
vocalization, displays of beauty or fighting prowess.

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From the scientific point of view, courtship in the animal kingdom is

the process in which the different species select their partners for
reproduction purpose. Generally speaking, the male initiates the
courtship and the female chooses to either mate or reject the male
based on performance. for example, the Selfish Gene model was
proposed by Richard Dawkins which state that an individual of a
particular species will mate with individuals from the same species
that display good genes.

In this case, courtship is a display of “genes” carried by a particular

organism looking forward to mix with the genes of another organism in
generation, thereby, ensuring the survival of the genes themselves.
Examples of courtship behavior in animal kingdom are as followed;

i.) Insects: female uses odorous substances called pheromones to

attract males from a distance eg. Gypsy moth (lymantria dispar).
ii.) Birds: boobies perform ritualized dances with many components,
including whistling and an elaborate gesture known to
Ornithologists as sky-pointing. The male peacock displays his
glouries plumage the female.
iii.) Amphibians: Courtship of songs in frogs (Rana species).

a. Humans

Human sexuality, besides ensuring biological reproduction, has

important social functions: it creates physical intimacy, bonds, and
hierarchies among individuals; and in a hedonistic sense to the
enjoyment of activity involving sexual gratification. Sexual desire,
or libido, is experienced as a bodily urge, often accompanied by strong
emotions such as love, ecstasy and jealousy. The extreme importance of
sexuality in the human species can be seen in a number of physical
features, among them hidden ovulation, the evolution of external
scrotum and penis suggesting sperm competition, the absence of an os
penis, permanent secondary sexual characteristics, the forming of
pair bonds based on sexualattraction as a common social structure and
sexual ability in females outside of ovulation. These adaptations indicate
that the importance of sexuality in humans is on a par with that found
in the Bonobo, and that the complex human sexual behaviour has a
long evolutionary history.

Human choices in acting on sexuality are commonly influenced by

cultural norms, which vary widely. Restrictions are often determined by
religious beliefs or social customs. The pioneering researcher Sigmund
Freud believed that humans are born polymorphously, perverse, which
means that any number of objects could be a source of pleasure.
According to Freud, humans then pass through five stages of

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psychosexual development (and can fixate on any stage because of

various traumas during the process). For Alfred Kinsey, another
influential sex researcher, people can fall anywhere along a continuous
scale of sexual orientation (with only small minorities fully heterosexual
or homosexual). Recent studies of neurology and genetics suggest
people may be born predisposed to various sexual tendencies.

b. Dog

In domestic dogs, sexual maturity begins to happen around age six to

twelve months for both males and females, although this can be delayed
until up to two years old for some large breeds. This is the time at
which female dogs will have their first estrous cycle. They will
experience subsequent estrous cycles biannually, during which the
body prepares for pregnancy. At the peak of the cycle, females will
come into estrus, being mentally and physically receptive to
copulation. Because the ova survive and are capable of being fertilized
for a week after ovulation, it is possible for a female to mate with more
than one male.

Dogs bear their litters roughly 56 to 72 days after fertilization, with an

average of 63 days, although the length of gestation can vary. An
average litter consists of about six puppies, though this number may
vary widely based on the breed of dog. Toy dogs generally produce from
one to four puppies in each litter, while much larger breeds may average
as many as twelve.

Some dog breeds have acquired traits through selective breeding that
interfere with reproduction. Male French Bulldogs, for instance, are
incapable of mounting the female. For many dogs of this breed, the
female must be artificially inseminated in order to reproduce.

c. Raccoon

Raccoons usually mate in a period triggered by increasing daylight

between late January and mid-March. However, there are large
regional differences which are not completely explicable by solar
conditions. For example, while raccoons in southern states typically
mate later than average, the mating season in Manitoba also peaks
later than usual in March and extends until June. During the mating
season, males roam their home ranges in search of females in an attempt
to court them during the three to four day period when conception is
possible. These encounters will often occur at central meeting places.
Copulation, including foreplay, can last over an hour and is repeated
over several nights. The weaker members of a male social group also

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are assumed to get the opportunity to mate, since the stronger ones
cannot mate with all available females.

d. Chicken

To initiate courting, some roosters may dance in a circle around or near

a hen ("a circle dance"), often lowering his wing which is closest to the
hen. The dance triggers a response in the hen's brain, and when the hen
responds to his "call", the rooster may mount the hen and proceed with
the fertilization.

e. Desert Tortoise

Tortoises mate in the spring and in the fall. The female will lay a
clutch of 3 - 5 hard-shelled-eggs (which are the size and shape of
ping-pong balls), usually in June or July, and they hatch in August or
September. Wild female tortoises can produce 2 or possibly 3 clutches
a year.

f. Wood Turtles

The wood turtle takes a long time to reach sexual maturity, has a low
fecundity (ability to reproduce), but has a high adult survival rate.
However, the high survival rates are not true of juveniles or
hatchlings. Although males establish hierarchies, they are not
territorial. The wood turtle becomes sexually mature between 14 and 18
years of age. Mating activity among wood turtles peaks in the spring
and again in the fall, although it is known to mate throughout the portion
of the year they are active. However, it has been observed mating in
December. In one rare instance, a female wood turtle hybridized with a
male Blanding's turtle.

The courtship ritual consists of several hours of 'dancing,' which usually

occurs on the edge of a small stream. Males often initiate this behavior:
starting by nudging the females shell, head, tail, and legs. Because of
this behavior, the female may flee from the area, in which case the male
will follow. After the chase (if it occurs), the male and female approach
and back away from each other as they continually raise and extend their
heads. After some time, they lower their heads and swing them from
left to right. Once it is certain that the two individuals will mate, the
male will gently bite the female's head and mount her. Intercourse lasts
between 22 and 33 minutes. Actual copulation takes place in the water,
between depths between 0.1 and 1.2 meters (0 and 4 ft). Although
unusual, copulation does occur on land. During the two prominent times
of mating (spring and fall), females are mounted anywhere from one to
eight times, with several of these causing impregnation. For this reason,

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a number of wood turtle clutches have been found to have hatchlings

from more than one male.

Nesting occurs from May until July. Nesting areas receive ample
sunlight, contain soft soil, are free from flooding, and are devoid of rocks
and disruptively large vegetation. These sites however, can be limited
among wood turtle colonies, forcing females to travel long distances in
search of a suitable site, sometimes a 250 meters (820 ft) trip. Before
laying her eggs, the female may prepare several false nests. After a
proper area is found, she will dig out a small cavity, lay about seven eggs
(but anywhere from three to 20 is common), and fill in the area with
earth. Oval and white, the eggs average 3.7 centimeters (1.5 in) in
length and 2.36 centimeters (0.93 in) in width, and weigh about 12.7
grams (0.45 oz). The nests themselves are 5 to 10 centimeters (2.0
to 3.9 in) deep, and digging and filling it may take a total of four
hours. Hatchlings emerge from the nest between August and October
with overwintering being rare although entirely possible. An average
length of 3.65 centimeters (1.44 in), the hatchlings lack the vibrant
coloration of the adults. Female wood turtles in general lay one clutch
per year and tend to congregate around optimum nesting areas.

The wood turtle, throughout the first years of its life, is a rapid grower.
Five years after hatching, it already measures 11.5 centimeters (4.5 in),
at age 16, it is a full 16.5 to 17 centimeters (6.5 to 6.7 in), depending
on gender. The wood turtle can be expected to live for 40 years in the
wild, with captives living up to 58 years.

In-Text Question(s)

What is Courtship?

Answer: Courtship is the period in a couple’s relationship

which precedes their engagement and marriages or establishment
of an agreed relation of a more enduring kind.
At what age do sexual maturity begins with domestic dogs?
What period do Raccoon usually mate?

Self-Assemeement Exercises 1

Attempt this exercises to measure what you have learnt so far.

This should not take you more than 5 minutes.
1. Write briefly on desert tortoise courtship.
2. Exlain courtship behavior in chicken.

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4.4 Summary

In summary, courtship in the animal kingdom is the process in which

the different species select their partners for reproduction purpose.
Generally speaking, the male initiates the courtship and the female
chooses to either mate or reject the male based on performance.

4.5 References/Further Readings/Web Sources

Bagemihl, Bruce (1999). Biological Exuberance: Animal

Homosexuality and Natural Diversity. St. Martin's Press ISBN
0-312-19239-8s

Ernst, Carl; Jeffrey Lovich (2009). Turtles of the United States and
Canada. Baltimore, Maryland: The Johns Hopkins
University Press. pp. 250–262. ISBN 978-0-8018-9121-2

Miller, S.A. and Harley, J.B. (1999): Zoology (Fourth Edition). WCB
McGraw- Hill, Boston
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1Raw&ntb=1

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4.6 Possible Answers to SAEs

Answers to SAEs 1

1. Tortoises mate in the spring and in the fall. The female will
lay a clutch of 3 - 5 hard-shelled-eggs (which are the size and
shape of ping-pong balls), usually in June or July, and they
hatch in August or September. Wild female tortoises can produce
2 or possibly 3 clutches a year.

2. To initiate courting, some roosters may dance in a circle around

or near a hen ("a circle dance"), often lowering his wing which is
closest to the hen. The dance triggers a response in the hen's
brain, and when the hen responds to his "call", the rooster may
mount the hen and proceed with the fertilization.

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Unit 5 Biological Clocks

Contents

5.1 Introduction
5.2 Intended Learning Outcomes (ILOs)
5.3 Circadian Clocks
5.3.1 Circadian Timing
System
5.4 Summary
5.5 References/Further Readings/Web sources
5.6 Possible Answers to Self-Assessment Exercises

5.1 Introduction

Biological clocks are mechanisms internal to the animal, that has

rhythmic influence upon its physiology and behavior, synchronizing
them to cyclic changes in the environment. Biological clocks are internal
timing mechanisms which can have a period of several hours, a day, or
a year.

The circadian clock runs with a period of about 24 hours. Circadian

clocks have two functional characteristics:

i.) they will persist (=free-run) with a period of about a 24 hours in

the absence of environmental cues; and
ii.) they will synchronize (=entrain) to a 24 hour environmental cue,
such as the light-dark cycle.

Entrainment is important because it permits animals to synchronize to

a changes in the seasonal photocycle. How they synchronize to the cue
is determined by the phase response curve (PRC).

Biological clocks are groups into the following;

i.) Exogenous: this is a direct response to various changes in

external (exogenous) geophysical stimuli
ii.) Endogenous: this is an internal (endogenous) rhythm that
programs the animals’ behavior in synchrony with the
exogenous temporal period, particularly a 24- hours or 365-
days period.

An animal may use many features of the external environment to gain

information about the passage of time. The most important of these is
the apparent movement of celestial bodies e.g sun, moon and stars, such
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influences have been much studies in birds and in bees (Apidae). In

addition, it is possible that animals can obtain time cues from changes
in environmental temperature, barometric pressure and magnetic
phenomena.

Endogenous daily rhythms are termed “circadian” and usually fall short
of a 24- hour periodicity. Endogenous annual rhythms are termed
“circannual” and usually less than 365 days.
Our biological clocks measure the day length and change our behavior
according to the seasons circadian rhythms and circannual rhythms
are internal calendars built into an animals’ nervous system, especially
in the brain.

5.2 Intended Learning Outcomes (ILOs)

At the end of this course, students should be able to:

 Explain biologica clock

 Identify the different types of biological clocks
 Differentiate between circadian and circannual rhythm

5.3 Circadian Clocks

Circadian clocks are important in photoperiod time measurement. There

are two models for how clocks might be important in measurement
of photoperiod:

i.) External coincidence model (external light occurs at a critical

phase in the circadian oscillation) and
ii.) Internal coincidence model (internal phase of multiple circadian
oscillators is set by dusk and dawn).

Circadian clocks are also important in animal orientation. Many species

of bird and fish use the sun for orientation. To be successful, however,
adjustment to the sun's daily movement is necessary. A circadian clock
makes this adjustment.

The concept of time has always perplexed and fascinated people.

Although the ancient Greek philosophers believed that time was infinite,
they perceived it as following endless cycles where the universe is born
and dies, with an exact recurrence of everything in each cycle. The
Newtonian view held that time is an immutable entity flowing on an

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infinite linear scale operating independent of nature’s forces. Einstein’s

theory of relativity radically changed this conventional wisdom by
revealing that time is actually embedded in the very fabric of the physical
universe giving rise to a reality that is more accurately expressed as
a four-dimensional space-time continuum. It follows from the big
bang theory that time and our physical universe were jointly created
at a singular event that, according to current estimates, occurred ~12–15
billion years ago, at least from our frame of reference. Interestingly,
Kabbalists had intimate knowledge of old oral traditions that
discussed the deep relationship between time, corporeal entities, and
their creation. For example, almost 800 years ago a famous rabbi known
as Nachmanidies, in his commentary to Genesis, wrote, "with this
primeval creation, which was like a very small point having no
substance, everything in the heavens and the earth was formed, and
when the heavens and the earth came forth from nothing into
existence, time came into being and from the moment some substance
came into existence time was already part of it”

Despite our changing understanding of the nature of time, one thing has
remained constant, the human obsession with harnessing this elusive
entity. From sundials to calendars to cesium clocks, the quest to capture
the essence of time and measure its passage has significantly influenced
human history.

But on what basis do we rationalize the units we use to measure time?

Essentially, this is a tale of spheres and cycles. The passage of time has
been recorded (at least historically) by observing the rhythms of
"heavenly bodies", most notably the daily rotation of the Earth on its
axis, the monthly cycle of the moon around the Earth, and the yearly
journey of our planet around the Sun. Considering the average
human lifespan and spatial distribution on our tilted planet, the broad
frequencies encompassed by these reliable celestial rhythms (day,
month, and year) provide quite relevant and useful units for recording
time. These predictably recurring physical events inspired humans to
design timing devices that could measure the passage of a known
amount of time (e.g., sand-filled hourglasses), identify a specific
phase in a cycle (e.g., sundials, Stonehenge?), or both (e.g., modern
clocks).

The most influential physical oscillation that reminds us of our

inescapable rhythmic relationship with time is the day-night cycle.
Although we intuitively know that time proceeds in a unidirectional
flight into the future, our lives are largely organized into a 24-h
schedule dominated by periods of wakefulness and sleep. Thus we
perceive time as being spiral in its geometry; there is ever forward
progression into the future (e.g., counting years) coupled with time

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coordinates that are revisited in a cyclical manner (e.g., measuring time

of day).

In 1729, a French astronomer named Jean Jacques d’Ortous de Marian

took plants that displayed daily leaf movements and put them in the dark
for several days. He noted that the leaves of the plants continued to
open during the day and close at night despite the absence of sunlight.
Based on this seminal experiment, he concluded that the observed
rhythm was not passively driven by a cyclic environment but was an
innate property of the plant. From these humble beginnings, the formal
study of circadian (endogenously driven biological rhythms with periods
of ≈24h) biology can be traced. It is now quite clear that living
organisms have been tracking the passage of daily time long before
we ever invented clocks.

However, the contention that life forms have internal time keeping
devices was only widely accepted about 60 years ago. For some it
seemed too incredulous that organic material had properties similar to
clocks fashioned by human hands. After all, it was claimed that
although daily pacemakers keep on ticking with a reasonably constant
period in the absence of environmental cues (zeitgebers), in natural
conditions they are accurately synchronized to local time. Adding to the
dismay was a peculiar attribute of circadian rhythms; the length of the
period is invariant over a wide range of temperatures. This still
mysterious property, known as temperature compensation, was viewed
largely enigmatic from a chemical or biochemical perspective because
most chemical reactions speed up about two- to threefold for every 10°C
increase in temperature (or Q10). Moreover, the prevailing thought at the
turn of the previous century was that physiology is governed by the
principles of homeostasis, effectively dismissing any observed
oscillatory behavior as nothing more than random fluctuations of little
or no significance.

With the eventual realization that endogenously driven daily rhythms

are "real" and widespread, occurring in virtually all organisms, much
interest was placed on elucidating the nature of the underlying
pacemaker or clock.

Then, almost overnight, there are now a bunch of clock genes identified
in humans, rodents, fish, frogs, insects, plants, and even cyanobateria.
This gold mine led to important paradigm shifts in how the organization
of the circadian timing system is viewed. From a molecular perspective,
the basic message from these recent studies is that circadian clocks use
the same design principles; namely, the period, amplitude, and phase of
a circadian clock are determined by a specialized set of interconnected
proteins, many of which undergo daily rhythms in one or more

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character traits, most notably abundance. From a theoretical point of

view, this framework for understanding the molecular underpinnings
governing circadian rhythms is very satisfying, because regular change
is the basis for timekeeping devices. However, although the RNA and
protein products from many genes display cyclical behavior, those
that define the clock operate within molecular.

5.3.1 Circadian Timing System

Circadian rhythms are operationally defined as biorhythms that exihibit the

following three properties:

i.) Persist (or free run ) with a period of ≈24 h in the absence of external
time cues (or zeitgebers)
ii.) Reset by changes in environmental conditions, most notably the
daily light –dark and temperature cycyles, and
iii.) Have an invariant period length over a wide range of physiological
relevant tempareture (temperature compensation, see above).

It is not clear why circadian clocks are designed with the capacity
to keep on ticking for long periods of time in the absence of a cyclical
environment (first property), a situation not normally faced in nature. A
free-running oscillator might enable animals to maintain synchrony
even during days when adverse weather conditions or other
unfavorable settings force them to seek shelter in places that receive little
or no light. Alternatively, this self-sustaining property might reflect some
peculiarity of the design principles required to build these oscillators and
not an adaptation with a particular advantage. The ability to reset a
circadian clock (second property) allows it to maintain temporal
alignment with local time. The last property (i.e., temperature
compensation) makes biological sense, because regardless of whether
it is a cold day or a warm day, it still lasts 24 h. A mechanism
that can offset the effects of temperature on the periodicity of an
oscillator is likely to be absolutely necessary in non-homeotherms, if
they are to maintain accurate timekeeping. Indeed, early versions of
man-made clocks were not very accurate, because increases in
temperature lengthened the pendulum causing the clock to slow down.
The circadian timing system is usually depicted as being composed of
three interconnected parts:

i.) input pathways that can receive and transmit environmental

cues, such as light and temperature, to a
ii.) clock or pacemaker, connected to
iii.) downstream effector pathways that manifest overt rhythms.

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Although the input to clock to output paradigm is usually depicted as

moving from left to right, there are examples where the flow of
information occurs in the opposite direction. These findings support
a recent model whereby photic input and circadian clock are viewed
in a more fluid relationship being composed of overlapping molecular
loops.

Whatever the early driving force(s), from our present vantage point it
appears that the most critical property of circadian clocks under natural
conditions is that they can be reset by external time cues. This property
was not merely selected so that we could avoid perpetual jet lag
following transmeridian flight. Rather, the ability to anticipate
environmental changes enables organisms to organize their physiology
and behavior such that they occur at biologically advantageous times
during the day. In addition, a second function that is widely regarded as
important is that these endogenous timekeeping devices also serve to
impose internal alignments between different biochemical and
physiological oscillations.

With this in mind we can appreciate why circadian rhythms are

observed at all levels of cellular organization. There are daily
oscillations in the levels of enzymes and hormones that affect the timing
of cell function, division, and growth. Physiological parameters such as
body temperature, immune responses, digestion, susceptibility to
anesthesia, and dental pain threshold. all undergo cyclic changes peaking
at fixed times during the day. Our visual and mental acuity fluctuate
during the day, affecting complex behaviors.

In addition to circadian rhythms that are manifested by and within

individuals, there are also group or population rhythms. Some of these
rhythms occur multiple times during the lifetime of the organism. For
example, in many Diptera, males and females have the same peak time
for activity during a daily cycle, increasing the chances of productive
encounters between the sexes. In this regard it is interesting to note that
because related species of insects have varying daily distributions of
activity , the circadian clock might have contributed to insect
speciation by establishing temporal barriers limiting the mating
opportunities of individuals sharing the same spatial constraints. Other
population rhythms involve events that occur once in a lifetime. A well-
studied example is the eclosion (emergence from pupal cases) rhythm in
Drosophila, which is only apparent in a group of individuals comprising
mixed developmental stages. The circadian clock gates the timing of
eclosion such that it happens in the early morning when the relative
humidity in the air is high. This is important because upon emerging
from its pupal case the fruit fly is susceptible to desiccation, and its wings
do not readily expand at low humidities. An interesting example of a

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population rhythm that is composed of many synchronized once-in-a-

lifetime events is the daily oscillation in luminescence displayed by the
cyanobacterium Synechocystis sp. It is interesting to note that this rhythm
is somehow transmitted from mother to daughter in mid- stride without
missing a beat, as the replication cycle is shorter than 24 h. Stable
population rhythms are not restricted to individuals of the same
species. The classic tango between bees and plants is a case in point.
Different flowering plants have characteristic times during the day when
they open and close their petals, making nectar available only at
restricted times. The presence of an endogenous and synchronizable
clock maximizes the feeding success of bees by enabling them to return
to the same plants at times in the day when their nectar is available.

These rhythms also highlight the fact that the "adaptive value" of a
circadian rhythm might only be understood within the framework
of the dynamic interactions occurring in particular habitats. On a more
global perspective, it is important to consider that organisms do not
adapt to a static environment but one that undergoes daily changes.
Oscillations in physical parameters (e.g., intensity of visible light, water
and air temperature, relative humidity) will pervade natural habitats
and their occupants, adding a strong daily component to the intricate
relationships that govern ecosystems. Whether physical or biotic factors
play primary or secondary roles in influencing the daily activity patterns
of animals is likely to be largely dependent on the species in question. It
has been suggested that the high rate of water loss in dry air might
be the main driving force for the nocturnal activities of some small
animals. On the other hand, biotic factors such as predation are
relatively more significant in determining the daily activity patterns
of larger animals. In any case, it is almost certain that many behaviors
involved in mating, reproduction, seeking shelter, hunting for food, and
avoiding predators evolved to take advantage of temporal niches. A
recent study showed that the ability of Drosophila to smell odors is
under circadian regulation (90), suggesting that many cyclical behaviors
are ultimately "hardwired" into clocks that regulate physiological
changes in the ability to sense, interpret, and respond to various cues
in the environment.

Circadian clocks are not limited to timing daily events, but also play a
role in adapting to seasonal changes in day length (photoperiod). By
distinguishing between the long days (or short nights) of
summer/spring and the short days (or long nights) of autumn/winter,
organisms that live in temperate latitudes can anticipate and respond to
seasonal changes in external conditions by controlling appropriate
developmental, physiological, and behavioural switches. For example,
certain species of insects enter diapause, a period of growth
arrestment that is induced by short photoperiods or cold temperatures.

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The Siberian hamster typically breeds only in the spring and summer
months, a seasonal adaptation that is partly regulated by regression of
the gonads induced by the expanded nocturnal release of melatonin.
Photoperiodism has been extensively studied in plants, where floral
initiation can be experimentally controlled by altering day length. Recent
genetic evidence in the flowering plant Arabidopsis clearly indicates that
common elements participate in circadian clock function and in eliciting
photoperiodic responses.

In addition to day length, circadian rhythms are modulated by seasonal

changes in average daily temperatures. Diurnal animals typically respond
to colder temperatures by displaying a greater proportion of their activity
during daytime hours, whereas nighttime activity predominates at
warmer temperatures. This directional response has a clear adaptive
value, ensuring that the activity of an organism is maximal at a time of
day when the temperature would be expected to be optimal for
activity. Direct evidence for circadian clock function in
temperature-induced alterations in the timing of the daily distribution
of activity has been shown in Drosophila melanogaster, where a
thermosensitive splicing event in per RNA contributes to preferential
daytime activity on cold days.

A less classic example in which circadian clocks are used is during long
distance navigation of birds, insects, and other animals to predetermined
target areas using the azimuth of the sun as a compass. By artificially
resetting the circadian clock, the animal misrepresents the position of
the sun leading to a predictable change in the direction of navigation.
What is Biological clocks? What is Exogenous?

Self-Assessment Exercises 1
Attempt this exercises to measure what you have learnt so far. This
should not take you more than 5 minutes.
1. What is Circadian Timing System?

5.4 Summary

Biological clocks are internal timing mechanisms which can have a

period of several hours, a day, or a year. The circadian clock runs with
a period of about 24 hours. Circadian clocks have two functional
characteristics where they will persist (=free-run) with a period of about

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a 24 hours in the absence of environmental cues; and will synchronize

(=entrain) to a 24 hour environmental cue, such as the light-dark cycle.

5.5 References/Further Readings/Web sources

Adams MD, Kerlavage AR, Fleischmann RD, Fuldner RA, Bult CJ,
Lee NH, Kirkness EF, Weinstock KG, Gocayne JD, White O,
(1995). Initial assessment of human gene diversity and
expression patterns based upon 83 million nucleotides of cDNA
sequence. Nature 377, Suppl: 3–174.

Akiyama M, Kouzu Y, Takahashi S, Wakamatsu H, Moriya T,

Maetani M, Watanabe S, Tei H, Sakaki Y, and Shibata S. (1999).
Inhibition of light- or glutamate-induced mPer1 expression
represses the phase shifts into the mouse circadian locomotor
and suprachiasmatic firing rhythms. J Neurosci 19: 1115–1121

Albrecht U, Sun ZS, Eichele G, and Lee CC. ( 1 9 9 7 ) . A differential

response of two putative mammalian circadian regulators,
mper1 and mper2, to light. Cell 91: 1055–1064.

Allada R, White NE, So WV, Hall JC, and Rosbash M. A (1998). mutant
Drosophila homolog of mammalian Clock disrupts circadian
rhythms and transcription of period and timeless. Cell 93: 791–
804

Bae K, Lee C, Hardin PE, and Edery I. (2000). dCLOCK is present in

limiting amounts and likely mediates daily interactions between
the dCLOCK-CYC transcription factor and the PER-TIM
complex. J Neurosci 20: 1746–1753

Bae K, Lee C, Sidote D, Chuang KY, and Edery I. (1998). Circadian

regulation of a Drosophila homolog of the mammalian Clock
gene: PER and TIM function as positive regulators. Mol
Cell Biol 18: 6142–6151.

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5.6 Possible Answers to SAEs

Answers to SAEs 1

1. Circadian rhythms are operationally defined as biorhythms that

exihibit the
following three properties:

i. Persist (or free run ) with a period of ≈24 h in the absence of external
time cues (or zeitgebers)
ii. Reset by changes in environmental conditions, most notably the
daily light –dark and temperature cycyles, and
iii. Have an invariant period length over a wide range of
physiological relevant tempareture (temperature compensation,
see above).

Glossary

°C = degrees Celsius
cm = centimeters
CNS = Central nervous system
CH4 = Methane
CO = Carbon
CH4 = methane (CH4)
DNA = Deoxyribonucleic acid
E. coli = Escherichia coli
FAP = Fixed Action Pattern
°F = degree Fahrenheit
Ft = feet
h = hour
in = inches
Kgs = kilograms.
km = kilometers
kph = kilometer per hour
m = meters
mm = millimeters
mph = meter per hour
NH2OH = Hdroxylamine
NH3 = Ammonia
O2 = Oxygen
PNS = Peripheral Nervous System
PRC = Phase Response Curve
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RNA = Ribonucleic acid

L = length
HIV = Human immunodeficiency virus
% = percentage
g = grams
spp = species
UV = Ultraviolet

End of the Module Questions

1. What is proximate causation?

2. List examples of innate behaviour?
3. What is Anthropomorphism?
4. What is habitat selection?
5. What is Parenthood?

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