Introduction

All animals have a variety of complex relationships

with members of their own species, with members of other
species, and with physical environment. The survival of a
species depends on its ability to obtain food and shelter, to
find mate and reproduce, and to protect themselves from
predators and parasites. In many ways adaptive behavior
ensures survival and survival ensures evolutionary success.

Why we study animal behavior

The study of animal behavior is enormously important,

both scientifically and economically. There are researchers
who study the behavior of animals in order to conserve and
protect endangered species, while others study animals in
captivity in order to preserve and exhibit them for the
education of humanity. Other behaviorists are interested
solely in the actions of economically important predators,
pests, and parasites. A few researcher study the behavior of
domestic animals that provide for our well being.

Behavior and animal welfare

Great concern with the welfare of captive animals was

taken recently. Captive animals include those species

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reared on farms, exhibited in zoos, kept as companions by
humans, wild animals that are hunted or trapped, and those
that are managed for conservation. Animals that enjoys
good welfare will not want for food, water, or shelter. It
will have space to move and opportunities for socialization
appropriate to its needs. A deficiency in any of these will
result in stress, a physical response to poor welfare. The
consequences of stress can be extreme. short-term stress
can be sufficient to have a negative impact upon the
immune system and a sufferer of prolonged stress will
often exhibit physiological, physical and psychological
damage. Animals that are confined for prolonged periods in
suboptimal conditions may develop characteristically
stereotyped patterns of behavior that they will repeat and
repeat and repeat. These stereotypies might be as
uncomplicated as repeatedly pacing a fixed route, or they
may be quite elaborate involving a set routine of paces,
head weaving, and other gestures. In some cases
stereotypical rubbing, digging, biting, and chewing, etc. can
even result in the performing animal damaging itself
physically.

E.g: If a wild male rat blunders into the territory of another

male the dominant resident will attack it. The intruder will

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flee, and once it is clear of the territory it will be safe from
further attack. But if we were to replicate this scenario in
the laboratory, in an enclosure from which the intruder
could not escape, we would see just how extreme the effect
of unmanageable stress can be. In this situation the intruder
rat’s physical health would deteriorate very quickly and it
may well die.

History of the study of animal behavior

1-Early humans

For many thousands of year humans were hunters and

meat eater, used fire, and made tools from animal bones.
Leaky, an anthropologist, proposed and tested a hunting
strategy that was based on animal behavior. He suggested
that upon sighting the prey at about fifteen meters distance,
the hunter should sprint directly toward the animal (small
animal often freeze in this situation. Within two or three
meters of the prey, the hunter should turn left or right
because the escape behavior is to make a sudden dash in
one direction or the other.

Prehistoric cave painting in France and Spain reveal

other aspects of humankind’s relationship to animals. Some

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of the drawing are symbolic representation of actual
hunting scenes.

2-Aristotle (384-322 B.C)

Aristotle wrote ten volumes on the natural history of

animals and used the observational method.

3-Theory of natural selection

Darwin (1809- 1882) clearly recognized the central role of

animal behavior in determining the outcome of competition
between animals.

4-Comparative method (George Roman, 1848-1892)

The comparative method involved studying animal to

gain insights into the behavior of humans. Roman proposed
that mental processes evolve from lower to higher forms.
This proposal relied on inferences rather than recorded
facts and observations.

5-Morgan (1852-1936)

Morgan used the observational method and experiments to

make theory.

6- Theories of genetics and inheritance

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 Theories of genetics and inheritance influenced research
in animal behavior. We now know that animal behavior
has a genetic components. Thus behavior may change as a
species evolve.

How to understand the animal behavior

Tinbergen’s four questions

In 1963 Niko Tinbergen, a recognized pioneer of the study

of animal behavior, suggested that there are four ways in
which we can ask the question “why?” and four different
ways of answering this question to understand certain
behavior.

Example: why common shrews (Sorex araneus) select

specific prey? Shrews choose the most nutritious prey.
Why do they do this?

Answers at Tinbergen’s four different levels might be as

follows:

1-The function of behavior (what is the behavior for?).

Answers at this level are concerned with its role in the life
of the organism. Our shrew may thus feed selectively to
make sure it takes only the most nutritious prey, so
maximizing its foraging efficiency.
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2-The mechanism of behavior (how is the behavior
achieved?). Answers here are concerned with how the
system operates in terms of underlying mechanism and
organization. So, for example, our shrew’s tactile and
visual senses may be most responsive to large, active prey
(which also turn out to be the most nutritious), thereby
biasing the animal’s intake towards these items.

3- Development of the behavior (how does the behavior

develop?). These kinds of answer are concerned with the
way the system reflects its embryological, or other
developmental influences. Our shrew’s early foraging
experience may therefore teach it which types of prey are
easy to locate and subdue, and so the most efficient to deal
with in terms of energy return.

4- The evolution of the behavior (where has the behavior

come from?). Here we are concerned with the ancestral
selection pressures and phylogenetic pathways that have
shaped and constrained the system. Thus shrews may be
selective foragers because less discriminating ancestors
foraged inefficiently so were less likely to survive and
reproduce. Selective foraging is thus the result of
generations of natural selection for increased foraging.

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 Approaches and methods

I-Ethology

Ethology is the biology of behavior: the exploration of

functional and evolutionary questions, and the mechanisms
underlying why an animal exhibits certain behavior.

Ethological study

The ethological study use observations from nature

and field experiments to formulate and test questions.
The behavior of black headed gull studied by Niko
Tinbergen is an example for the ethological method.
Tinbergen observed that after the young black headed
gull chick hatches, the parents carefully pick up the
remnant pieces of shell, fly off to some distant location
and drop them. Why do parent birds remove the egg
shell?

Tinbergen and his colleagues considered several

explanations to this behavior. The sharp edges of the
shells might injure the checks or the shell may clutter the
nest and hamper the parents. These explanations were
not correct because the egg shell is thin. They
hypothesized that, the parents remove the shells because

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 it may attract predators. They make two experiments to
test the hypothesis.

Black-headed gulls build their nests in close proximity to

one another

Experiment 1: They scattered eggs outside the gallery.

Some eggs have natural color and others were painted with
white. They recorded predation rate and found that white
eggs were more taken by predators.

Experiment 2: They laid eggs outside the dunes and

scattered half eggs at different distances from it. The result
indicated that, the greater the distance between the half
eggs and whole eggs, the lower the probability to detect the
position of the whole eggs.

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Conclusion: The parents remove egg shell to protect chick
from predators.

Advantages of the ethological method:

The function and adaptive significance of behavior can be

discerned best under field condition.

1- In natural setting, animal behavior is not be

restricted.

Disadvantages of the ethological method:

1- Ethologists can not control environment and lack of

knowledge of the observed animals.
2- Ethologists can not design suitable experiment in the
field conditions.

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II-Comparative psychology method:

Comparative psychology is a comparative study of

behavior in animals. Comparative psychology research
depends on two ways:

1- Identify and characterize classes of behavioral

patterns in two or more species.
2- Select species that is the most appropriate for
investigation of a particular problem.

Example of the study:

Test the ability of snake to respond differentially to

airborne odorants. In the first portion of experiment, snakes
were placed in glass tank divided into 3 compartments. The
snakes placed in the middle and chips with lemon extract
put in another compartment. Partitions then removed and
when the snakes head removed toward the lemon odor
scientists gave them reward. It was found that 70% of
snakes move toward the odor. To test snake ability to learn
the experiment repeated but the reward was given when
snakes move their heads far from the odor. After 50 trials
the majority of snakes move their heads far from the odor.

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Conclusions: The snakes had the ability to discriminate
airborne odorant and use this ability to detect prey, also
they had the ability to learn.

Advantages of Comparative psychology

1- The experience and the environment can be

controlled.
2- The ability to design suitable experiment.

Disadvantages of Comparative psychology

1- Animal behavior can be affected by captivity.

2- Can not study in their natural environment.

III-Behavioral ecology and sociobiology

Behavioral ecology and sociobiology involves

experimental manipulation under field condition.
Behavioral ecology deals with habitat selection, feeding,
and other aspects of species ecological niche, with
particular reference to behavior. Sociobiology is concerned
with social behavior and social organization.

Example of study: Scientists study whether energy level

affect mating behavior in Sand goby fish. They divided
Sand gobies into 2 groups in 2 artificial nests in natural

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setting. One group supplied by food and stay long time in
the nest but the other not supplied with food and leave the
nest to obtain food. The fed group spend more time in the
nest and more time for mating than do the unfed group. The
fed group had big egg mass than the unfed group.

Advantages of the Behavioral ecology and sociobiology

1- Animals studied in their natural habitat.

2- Provide specific data to test particular hypothesis.

Disadvantages of the Behavioral ecology and sociobiology

1- Scientists can not control environment.

2- Scientists can not design suitable experiment in the
field conditions.

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 Behavioral genetics

Many pattern of animal behavior are adaptations that

have resulted from the process of natural selection. In order
for natural selection to operate on behavior, behavioral
pattern must be at least partly genetically based.

Studying single gene effect

While no behaviour is encoded at a single locus in a

literal sense, there is a wealth of evidence that changes at
single loci can affect behaviour through the cascade of
developmental processes influenced by their expression.

Mendelian crosses

Classical genetic crosses can sometimes be used to

investigate simple pattern of inheritance for behavioural
trait. A famous example is the ‘hygienic’ behaviours in
honey bees. Honey bee larvae are susceptible to various
diseases, among them American foul brood caused by the
bacterium Bacillus larvae. Some strains of honey bee are
called hygienic because when larvae die inside their cells,
workers uncap the cells and remove them. Dead larvae in
unhygienic strains are left in their cells to decompose, thus
risking the spread of disease. If unhygienic and hygienic

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strains are crossed, the F1 progeny are all unhygienic.
Unhygienic behaviour is thus dominant and hygienic
behaviour recessive. F1 bees back crossed (mated them
with the homozygous recessive parental strain) with the
following outcomes:

-Nine of the resultant colonies uncapped cells in which

larvae had died but did not remove the corpses.
-Six colonies would remove dead larvae from uncapped
cells, but would not uncap the cells themselves.
- Eight colonies were unhygienic and neither uncapped
cells nor removed dead larvae if the cells were uncapped
for them.
-Six colonies were hygienic and did both.
The hygienic trait can thus be explained in terms of two
independently segregating genes each with two alleles. One
gene codes for uncapping behaviour as dominant
unhygienic (U) and recessive hygienic (u) alleles, the other
codes for removing the larva, again with unhygienic alleles
(R) being dominant and hygienic (r) alleles recessive. Since
honey bee workers are diploid, they possess two copies of
each of the uncapping and removal genes. Workers of the
pure breeding hygienic and unhygienic strains in therefore
had the genotypes uurr and UURR respectively.

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 Mendelian crosses of honey bees

Mutations
Until relatively recently, studies of single gene effects
have relied on mutations arising at particular loci and then
looking for differences in behaviour between mutant and
non-mutant (wild-type) individuals. Useful mutations can
arise spontaneously, or, more usually, artificially by
exposing organisms to chemical or other mutagens. A
useful property of many single gene mutations affecting
behaviour is that they also have other phenotypic effects
(i.e. they show pleiotropy) so that carriers are
easy to identify even when they are not performing the
behaviour in question.
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Transgenic technique
The advent of genetic manipulation techniques has
provided opportunities for probing single gene effects more
directly. Identified genes can be inserted into (so-called
‘knock in’ procedures) or removed (‘knock-out’
procedures) from the organism’s cells, a process called
transgenesis, or they can be activated and deactivated in
situ, to see what effect they have.
Multiple gene effect
While single gene mutations show that small differences
in genotype can have pronounced effects on behaviour,
many, if not most, behavioural differences are likely to be
underpinned by many different loci with complex epistatic
relationships between them. The evidence for multiple gene
effects is diverse, and much of it long established in the
behaviour genetics literature.
Cricket songs are also mechanical in origin and they are
produced when the animals open and close their specially
modified forewing cases. When the wings open no noise is
emitted, but as they close they rasp against one another and
a sound pulse results. The members of each species sing a
specific song characterized mainly by the duration of these
pulses and the silences between them. The sonogram
represent pictorially the songs of two cricket species and of

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hybrids between them. A comparison of these sonograms
reveals that the hybrid songs have features of the songs of
both of its parent species, but are distinct from them and
from one another. This observation, together with the fact
that song is a product of wing-case shape, musculature, and
nervous system, suggests strongly that song production in
these species is polygenic.

Sonogram

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Genes versus environmental influences

The ability of mice to find food in a maze is an example

to clarify the interaction between environment and the
genes to create certain behavior. Experiments were
designed to compare the speed with which mice of
different strains learned to negotiate a maze of ladders,
tightropes and other challenges to get to the food. When
mice were all reared in the same standard laboratory cage,
there was little difference in the speed of learning between
some of the strains so genetic differences appeared to
contribute little to differences in learning ability. However,
when mice were reared in cages enriched with toys and
climbing surfaces, differences between these previously
similar strains began to emerge. While learning speed was
enhanced in all mice reared in enriched environments, mice
of some strains were very much quicker than those of
others, a difference that was attributable to the genetic
difference between them. Thus, a particular kind of
environment was necessary for the genetic difference in
learning ability to be expressed. Of course, some traits,
such as blood group and eye colour, are fixed by the
individual’s genes, but many others –especially behavioural

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traits – are subject to considerable modification by the
environment.

Some extreme examples of how powerful environmental

influences can be come from the caste systems of eusocial
insects. For example, depending on how they are fed as
larvae, closely related sisters of the haplodiploid ant species
Pheidole kingi develop into very different kinds of adult,
queens or various forms of worker, not only performing
different suites of behaviour but also looking very different
morphologically.

Different castes of the ant Pheidole kingi (a) queen, (b) worker, (c)

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 Mechanisms of behavior

Mechanisms of behavior include the neural, hormonal,

and biological clock effects on behavior. The experiences
gained by animals (learning) are also one of the
mechanisms of behavior.

I -Nervous system

Every living organism is continually bombarded by

environmental stimuli. Animals have sensory receptors to
receive information from the environment, sensory neurons
to transmit stimuli, nervous system to sort out and interpret
the stimuli, interneuron and motor neuron to transmit
stimuli to effector (e.g., muscles and glands) system to
produce appropriate behavioral responses.

Examples of neurobiology and behavior

1-Mechanics of feeding:

All animals need to feed. This simple behavior generates

many questions. What prompts an animal to begin feeding?
What signals them to stop feeding?

E.g., blow fly: when a blowfly is hungry and lands on

potential food source, chemoreceptors on its legs are

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stimulated. It extends its proboscis, thereby stimulating its
oral taste receptors and begins to feed until the fullness of
its foregut. Ethologists hypothesized that hunger and satiety
might be controlled by the fullness of the foregut. There are
receptors on the wall of the fore gut that stretched when the
foregut is full and send message to the brain via specific
nerve to inhibit further eating.

2-Escaping danger

When an animal is suddenly confronted with danger, its

survival depends on fast action such as running, flying, or
pulling back into a shell.

E.g., Cockroach: cockroaches have cerci projecting

posteriorly off their abdomen. These are mechanoreceptors
covered by hair that are highly sensitive to the movement
of air currents. When cockroach senses the approaches of a
crushing foot they run quickly.

Escape response is that: receptors transmit stimuli to giant

interneuron that transmit the stimulus to the leg muscle in
the thorax.

E.g. (2) Moth escape from bats: bats detect prey by sending
ultrasound and listen to the echo. Moth auditory receptors

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detect the sound of the bat and the message send to their
wing muscles and fly.

3-Detecting prey

Predators must have the ability to distinguish preys that

are not dangerous to them. Many animals have feature
detectors, which are neurons respond only to stimuli with
specific characteristics.

Example 1: Dragonfly has 8 feature detectors. Some of

these neurons respond to target with specific size, whereas
others help to steer the dragonfly during prey tracking.

Example 2: Prey capture by the toad

When a prey item (usually a worm or an insect) moves

through the visual field of a toad a sequence of behaviors
will be performed:

1. The toad moves so that it faces the prey animal.

2. The toad approaches the prey animal to within striking

distance.

3. The toad uses its tongue to strike at the prey animal and
captures it.

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4. The toad swallows the animal.

5. The toad wipes its mouth.

Mechanism of prey capture by toad

Moving objects stimulates receptor cells (rods and

cones) of the retinal surface which in turn stimulates the
interneuron. The information is then transmitted to the
ganglia. This Information received by the toad retina is
transmitted to the optic tectum of the brain via the optic
nerve. If a small object moves in front of the toad light
reflected from it on to the retina will focus on a small area
of the retinal surface. This means that there is a good
chance that a greater number of the excitatory retinal cells
will be stimulated, these ganglia will transmit strong
signals, and the toad will see a potential prey item. On the
other hand, a very large object passing in front of the toad
will stimulate a larger area of the retina and will therefore
stimulate both excitatory and inhibitory receptors and either
a weak signal or no signal will reach the brain. This
arrangement will therefore allow the toad to discriminate
between potential prey items (small objects) and nonprey
items (larger objects). If it can discriminate between these

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two classes of stimulus it can behave appropriately towards
each of them.

The feature detectors located in specific areas of the

brain. The first area to fall under investigation was the
thalamic pretectum which contains a class of cells (termed
TH3) that are responsive to visual information. The second
area is the tectum that has a class of neuron termed T5(1)
and T5(2). They are the feature detectors responding most
strongly to worm-like and least strongly to nonworm-like
stimuli. The TH3 and T5(1) neurons are both involved in
the prey detection and capture. The TH3 cells have an
inhibitory effect and T5(1) cells have an excitatory effect
upon T5(2) cells. In response to a worm-like stimulus the
T5(1) cells are able to stimulate the T5(2) cells without
conflicting inhibition from the TH3 cells, and so the net
effect is a strong T5(2) response and the facilitation of the
prey capture reflex. However when a non-wormlike
stimulus is presented the T5(1) cells respond weakly and do
not counter the strong inhibitory effect of the TH3 cells.
Therefore the T5(1) cells do not respond and the stimulus is
ignored.

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 Toad brain

4-communication

Communicating animals have to produce signals, and to

detect signals produced by conspecifics.

E.g.1 Crickets: Male crickets emit sound via stridulation

(rubbing together a portion of the cuticle of one wing
against a series of ridges on the other wing. This sound
serves to attract female.

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Katydids use several different forms of communication. One of these forms
is called stridulation and is characterized by the rubbing together of the
insect's wings to create sound waves. These sound waves convey specific
types of information and are detected by members of the same species.

The neural events involved are:

Impulses from brain pass through interneuron to the

thoracic ganglia then the stimulus transmitted to wing
muscles and muscles contract. Female crickets have
receptors for this sound that transmit stimuli to neurons
then to thoracic ganglia and the female fly to the source of
the sound.

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E.g. 2 : Bird songs

Birds have a specific organ for song production which is

called syrinx. The junction between the 2 bronchi is
modified to syrinx. Muscles surrounding the syrinx control
the tone and the timing of the song. Avian brain has
discrete nuclei responsible for song production.

The syrinx

The neural pathway for song production originates in HVC,

or higher vocal center, and then passes to RA (robustus
archistriatum). The RA sends projection to the dorsomedial
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nucleus DM and to the hypoglossal nerve which innervates
the syringeal muscles.

Neural pathway of bird song

5-Learning and memory

Birds hide hundreds of seeds and then find them by

memory, often long after they have hidden them. A
particular section of the brain called hippocampus and Para
hippocampus is involved in spatial memory. The
hippocampus is important in spatial learning of humans.

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Taxi drivers were used as examples of humans with very
good spatial skills.

The hippocampus

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 II-Hormones

The endocrine glands secrete chemical messengers, or

hormones, into the circulatory system in response to
various internal and external stimuli. Hormones are also
secreted by special neurons (neurosecretory cells) within
the nervous system itself, where they are transmitted along
axons or, again, into the blood. When they are transmitted
via the circulatory system, hormonal messages are much
slower than the electrical messages of the nervous system.
Thus, they are particularly suited to regulate functions,
physiological and behavioural, that are sustained over
minutes, days or even months.

How hormones affect behaviour

The effects of hormones on behaviour can be traced to

four broad areas of influence: the nervous system, sensory
perception, effector systems and development.

1-Effects on the nervous system

Hormones affect many aspects of the nervous system,

including anatomy, biochemistry and impulse transmission.
In some cases, they may be responsible for basic structural
and functional changes within the CNS. Reflex

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connections, for example, are accelerated by high levels of
thyroxin, a hormone secreted by the thyroid gland. Some
sex differences in behaviour in rats are associated with
sexual dimorphism in the anatomy of cells in the neuropile
of the hypothalamus, a dimorphism that appears to be
mediated by neonatal levels of androgen.

A given hormone can have very different effects on

behaviour depending on the region of the CNS on which it
acts. The effects of intracranial implants of testosterone (an
androgen) in male house mice on various aspects of social
and sexual behaviour, including ultrasonic vocalisations,
urine marking, mounting and aggression was studied. They
implanted the hormone in one of four regions of the brain:
the septum, the medial preoptic area, the anterior
hypothalamus or the ventromedial hypothalamus. Implants
in the median preoptic area showed increased ultrasonic
vocalisation,

while those in other regions had little effect. Those in the

medial preoptic or either hypothalamic regions resulted in
more urine marking than empty. Mounting occurred in
testosterone controls and males implanted in the median
preoptic area, while aggression was rare in all males given
brain implants.

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2-Effects on sensory perception

Many studies suggest that hormones affect an animal’s

sensory capabilities. In doing so, they alter the animal’s
perception of its environment and therefore the way it
responds to particular stimuli. The seasonal migration of
three-spined sticklebacks (Gasterosteus aculeatus) provides
a good example. In spring, male sticklebacks migrate from
the sea to their freshwater breeding grounds. In doing so,
they move through water that gradually changes in salinity.

Three-spined sticklebacks

To facilitate this transition, hormones secreted by the

pituitary and thyroid glands, but particularly thyroxin, alter
the fishes’ salinity preference from salt water to freshwater.
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In many female mammals, sensory perception is influenced
by the oestrous cycle. Female rats fluctuate in their ability
to detect certain odours according to circulating levels of
oestrogen and progesterone. Similarly, visual sensitivity in
women varies with the stage of their menstrual cycle (and
thus relative oestrogen and progesterone levels), being most
acute around the time of ovulation, and least acute during
menstruation, a difference that is abolished when taking
oral contraceptives.

3-Effects on effector systems

Animals use a range of appendages and other external

structures in performing different behaviours. Various
hormones affect the development of such structures and
thus their efficacy in performing whatever behaviours
depend on them. In some cases, hormones induce
appropriate muscle development, as in the hypertrophied
brachial musculature used by male frogs in coupling. Sex
differences in call characteristics in the clawed toad,
Xenopus laevis, are also due to hormonal effects on muscle
development. In this case, androgens increase the number
of muscle fibres in the larynx of males as they mature, and

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stimulate the development of more so-called ‘fast twitch’
fibres that produce the characteristic rapid trill of the male
call.

Male frog calls

In women, changing oestrogen and progesterone levels

during the menstrual cycle affect the strength of several
muscle systems, causing fluctuations in physical
performance at different stages of the cycle. Secondary
sexual adornments provide other examples of hormonally
induced effector systems. In newts, prolactin appears to be
important in the development of the enlarged tail fin in

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males, which is used to fan a stream of water at the female
during courtship. While prolactin also influences the vigour
of tail fanning, there is a synergistic effect with growth
hormone in enhancing the overall performance of the
response.

4-Effects on development

Hormones have a profound effect on the development

of young animals and impart characteristic features to their
behaviour as adults. While hormonal effects on behaviour
and associated physiological processes in adults are usually
reversible and independent of age (once adult), those
affecting development are permanent and irreversible, often
limited to clearly defined periods in the developmental
process, and usually manifested later in life rather than at
the time of effect. In guinea pigs, testosterone levels
influence the development of the genitalia, so that female
offspring born of females treated with testosterone during
pregnancy have male-like genitals. If they are then
ovariectomised and treated with gonadal hormones, these
offspring develop more male-like behaviour than controls
whose mothers were not treated with testosterone. If female
rats are given testosterone at around 4 days of age, their
oestrous cycle and sexual behaviour as adults are

35
suppressed and remain unresponsive to ovariectomy and
oestrogen treatment. Studies of a range of species suggest
that hormonal effects on sexual responses such as those
above occur during more or less clear-cut critical periods
in an animal’s development. Sometimes the critical period
is prenatal, sometimes postnatal.

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 III-Biological rhythms

Biological rhythms are fluctuations in the biological

functions among a day, month or a year. This means the
biological functions increase and decrease in cycles that
repeat every day, month or a year. According to the period
length of the rhythm it divides into:

1-Ultradian rhythm

In ultradian rhythm the period length is less than 24 h.

E.g. lug worms feed every 6-8 minutes.

Tidal rhythms

Tide is the result of unequal gravitational forces of the

sun and the moon. Tides exhibit a 12.4 hour period from
one low tide to the next. Crabs inhabit seashore feed when
tide is out and return to burrows when tide flow return.

2-Infradian rhythm

Infradian rhythm with a period length more than 24 h.

E.g. the menstrual cycle repeat every 28 days.

Circannual rhythms are rhythms that repeat each year, e.g.

hibernation and migration before winter.

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3-Circadian rhythm

Circadian rhythm is a rhythm with a period length equal

to 24 h., e.g. secretion of the hormones, melatonin,
cortisone, sleep-wake cycle (in human), and locomotor
activity (in animals).

Circadian rhythm exhibit 3 main characteristics:

1- Persist or free-run with a period length about 24 h.

in the absence of external cues (zeitgebers).
2- They are reset by changes in environmental
conditions, mostly by the light-dark cycle.

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 3- Their period vary only slightly with ambient
temperature.

Evidences indicate the presence of an endogenous clock

generates a biological rhythm.

1- Free-running rhythm

The activity of a flying squirrel measured in constant

darkness for several weeks. The activities of the animal
remain rhythmic with slight change in period length, which
indicate the presence of endogenous clock.

2- Translocation

Honey bees visit feeding sites in Paris at particular time

(when flowers open). If the bees transferred to New York
with different timing, the bees will forage at the same time
as would have been expected had they remained in Paris –
24 hours after their last foraging. The bees gradually
adjusted to local time by foraging later and later each day.

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 The Biolological clock

The biological rhythm is generated by biological clocks

situated in nearly all organs of the body (peripheral clocks).
The pacemaker (central clock) is located in the
suprachiasmatic nuclei (SCN) of the hypothalamus. The
SCN generates the circadian rhythm and synchronize the
rhythms of the peripheral clocks through specific signals.

The central and peripheral clocks

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The molecular mechanism of the circadian clock

The endogenous circadian rhythm is generated by a

molecular mechanism involves nine clock genes. Clock
genes found in nearly all cells. They are named, Period
(Per1, Per2, Per3), chryptochrome (Cry1, Cry2), Bmal1,
Clock, Rev-erb- and Casein kinase 1. CLOCK and
BMAL1 proteins act as a positive feedback limb that
activate the transcription of Per and Cry genes. PER and
CRY proteins act as a negative feedback limb that inhibit
CLOCK BMAL1 actions and inhibit their own
transcription. Casein kinase 1 phosphorylate PER
proteins and the phosphor-PER become degraded. The
negative limb will be inhibited and the cycle repeats. This
cycle repeats every 24h.

Clock genes transcription and translation

41
The entrainment of the circadian clock

Entrainment is a necessary adjustment of circadian

rhythm to the external light-dark cycle. Even if endogenous
period is very close to 24 h, say 24.1 h, if left by itself, it
will gradually diverge from the outside world until phase
differences are inadequate for survival. In our example, the
endogenous phase of the rhythm is shifted by 0.1 h (6 min)
every day, so in 10 days the circadian system of the animal
will be 1 h advanced with respect to a natural cycle. During
the day the endogenous clock must be resilient to photic
cues, since otherwise its phase would constantly change
during the light phase.

Zeitgeber (timegivers)

Zeitgeber are entraining agents (cyclic environmental

cues that reset endogenous clock). Zeitgeber for the central
clock include light-dark cycle, photoperiod, ebb and flow
of tide. Social cues, melatonin, some food and some drugs
also reset the endogenous clock. The peripheral clocks are
entrained by the SCN and the restricted feeding.

42
 Entrainment of the circadian clock

Circadian rhythm abnormalities

The circadian rhythm controls a variety of biological

processes, such as sleep wake cycle, body temperature,
feeding, hormones secretion, glucose homeostasis, and cell-
cycle regulation. The timing of these physiologic rhythms
may become altered, leading to changes in the phase
relationship of rhythms to each other, which can cause
internal desynchronization. This loss of coordination of
rhythms may have negative consequences on rest-activity
cycles and other physiological and behavioral functions.

Circadian rhythm sleep disorders

Circadian rhythm sleep disorders (CRSDs) are chronic

patterns (for at least 1 month) of sleep-wake rhythm

43
disturbances due to alterations of the circadian timing
system or to a misalignment between the timing of the
endogenous circadian rhythm and the sleep wake times
required by school or work schedules.

1-Delayed sleep-phase disorder

Delayed sleep-phase disorder (DSPD) is characterized

by a chronic or recurrent inability to fall asleep and wake
up at socially acceptable times. Bright light (full spectrum
or blue enriched) in the morning for 2 to 3 hours before
natural wake-up time has been shown to successfully
advance circadian rhythms in patients of DSPS.

2-Advanced sleep-phase disorder

Advanced sleep-phase disorder (ASPD) is characterized

by an advance in the phase of the major sleep episode in
relation to the desired or required sleep and wake-up times.
Patients have chronic or recurrent difficulty staying awake
until the desired or socially acceptable bedtime, together
with an earlier than desired wake-up time. The most
commonly used treatment for ASPD is early-evening light
therapy, usually between 7:00 PM and 9:00 PM for
delaying circadian phase.

44
3-Jet-Lag Disorder

Jet-lag disorder results from travel across several time

zones and subsequent misalignment of the internal
circadian clock and the destination’s local time. Symptoms
of jet lag usually emerge within 1 to 2 days after travel.
Main manifestations of jet lag are generalized malaise,
sleep disturbances, impaired daytime alertness, poor
appetite, diminished cognitive performance, depressed
mood, irritability, and anxiety. Adjustment of the bed time,
caffeine and hypnotic medications, have been explored as
options to alleviate jet-lag symptoms.

4-Shift work disorder

Shift work disorder (SWD) is characterized by a history

of chronic (at least 1 month) excessive sleepiness during
the required wake (work) time and/or insomnia symptoms
during the associated required or desired sleep period that
occurs in relation to unconventional work schedules. Shift
work accompanied with the risk of weight gain,
hypertension, and cardiovascular disease, and some studies
suggest an association with breast and endometrial cancer.
In addition to the medical comorbidities, SWD is
accompanied by significant social and economic burdens in

45
the form of accidents, lost days of work, poorer
performance. Treatment includes optimizing the sleep
environment (eg, darkened room, comfortable temperature,
noise reduction), adherence to good sleep habits (eg,
maintain a regular sleep and wake schedule, avoid
excessive caffeine).

Significance of biological timekeeping

1-Ecological adaptation

Physical factors like light, temperature, humidity are

critical factors for organisms that cannot regulate body
temperature and live in areas with sever daily variation
(Temperate and arctic regions).

E.g. Woodlice exhibit circadian rhythm regulated by light-

dark cycle. They are active at night when temperature
decreases and humidity increases. During day they hide in
dark places where the humidity is higher. Endogenous
clock is important to prepare the animal for changes in
environment and the animal get ready to hide before sun
rise.

46
2-Diurnality

Activities of animals during day or night determined by

biotic factors (feeding, reproduction, escaping predator).
For example honey bees foraging only during day light but
they do other activities at any time.

E.g. (2) Parasitoid insects lay their eggs in the eggs or

pupae of other insects. Adult emerge during early morning
to mate and search for new host before dark. The inherited
endogenous clock makes the adult insect emerge at a
proper time during early morning.

47
 The parasitoid insect lie its eggs in an insect

3-Applied aspects of circadian clock

An example of the application of knowledge about

circadian activity rhythm involves attempts to control
rodent population.

E.g. voles and hamsters were given different rodenticides:

1- Zink phosphide, both species exhibited stronger

susceptibility to the poison when it was provided during
light phase of the daily cycle.

2- Crimidine , hamsters were susceptible at the beginning

of the dark phase of the cycle, and there was no circadian
effect for voles. Variations in effectiveness for the

48
rodenticides can be attributed to differences in their routes
of physiological action and to differences in activity phase.
Voles are active in daylight hours whereas hamsters are
nocturnal. Thus an understanding of circadian rhythm as
well as physiological effects of specific poisons are
required by the rodent-control biologist.

4-Hibernation

Some animals hibernate (for example squirrel, jumping

mice) to escape the cold of the winter. Hibernation is
annual rhythm begins from mid- fall to mid- spring.

Before hibernation

1- Animals feed more to form fat deposits which act as

insulator and the animal use it as source of energy
during hibernation.
2- Animals burrow a suitable hibernating chamber.

After hibernation animals mate directly. So animals must

have endogenous clock that tells the animal the proper time
to build burrows and feed before winter. Also to prepare
the animal during hibernation for mating after the end of
hibernating time.

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5-Migration

Birds migrate from cold areas at north in winter and

return in spring and breed in summer. Endogenous clock
must trigger this action. Endogenous clock entrained by
photoperiod (change in day length).Changes in day length
leads to changes in the duration of melatonin secretion
from the pineal gland, so the body feels with day length
changes.

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 IV-The influence of experience on behavior

Learned behavior

The adaptive change of behavior as the result of

experience—usually known as learned behavior or
experience-dependent behavior—may be observed in all
higher vertebrate forms. Several types of such learning in
animals are recognized: habituation, classical conditioning
(CR Type I), trial and error (CR Type II, instrumental
conditioning, or operant conditioning), latent learning,
insight learning, and imprinting.

1-Habituation

Habituation is learning to disregard stimuli that are

without significance to the animal. In many respects it is
the simplest form of learning, and it is sometimes regarded
as a fundamental property of all living matter. Most
animals inherit a response to be frightened by sudden and
strong stimuli such as loud sound, flashes of light, and the
sudden intrusion of anything foreign into the animal's
sensory field. Yet, if an animal reacts nonselectively to all
phenomena such as rustling leaves, thunder, snapping
twigs, and the sudden appearance of harmless animals,
those phenomena that are significant to the animal's well-

51
being will not receive the intensity of response that they
often require. All animals, then, quickly habituate to such
harmless stimuli, but the adaptation is highly specific. An
animal that habituates to one type of sound does not, as a
consequence of this habituation, become habituated to other
sounds. Habituation is distinct from failing to respond to
stimulation as a result of fatigue, sensory adaptation, or
injury. The effects of habituation are generally long lasting.
If an animal is repeatedly exposed to a potentially harmful
stimulus (such as to a predator) without being harmed,
habituation does not generally occur. Responses to
dangerous stimuli often seem to have an inherited
resistance to habituation—a mechanism of obvious survival
value.

2- Classical conditioning

Classical conditioning was studied early in the 20th

century by the Russian physiologist Ivan Pavlov, who
observed that dogs salivate when food is placed in their
mouths. He gave dogs food and, at the same time, provided
another stimulus such as a flashing light or the sound of a
bell. After a few such pairings of stimuli, a dog would
salivate upon seeing the light or hearing the bell but

52
without the presence of food. The dog had learned to
associate the flashing light or the sound of a bell with food.

A previously irrelevant stimulus that assumes significance

as a result of association with a relevant stimulus is called a
conditioned stimulus (CS). Salivation in response to such a
stimulus is termed a conditioned response (CR). Prior to
the learning experience, only the food (unconditioned
stimulus) was effective in producing salivation
(unconditioned response).

Such conditioned responses have been observed in a

wide variety of animals, from lower invertebrates to man.
Birds learn to avoid noxious insects in this manner; the
distasteful monarch butterfly (Danaus plexippus) or a

53
species of stinging wasp provide effective stimuli that
quickly become associated with the appearance of such
insects. This kind of association together with the
conditioned response relevant to it is also the basis for
Müllerian mimicry, in which palatable insects and other
animals evolve to resemble noxious ones, thus enhancing
their chances of survival.

3-Trial and error

In trial-and-error learning, an animal learns to behave in

a particular way by associating something it does with a
desired effect. If a dog's foot is lifted by the experimenter
and food then given, the dog, after a few such trials, will
spontaneously lift its foot in anticipation of food. Both
classical conditioning (CR Type I) and trial-and-error
learning (CR Type II) are termed associative learning,
because in both cases an unconditioned response is
associated with a conditioned stimulus. In natural situations
an animal probably learns to associate certain spontaneous
activity of its own with certain desired results, thus fixing
the conditioned stimulus and response. Learning of this
type often occurs when animals modify their behaviour
during appetitive sequences such as those involving feeding
and mating.

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 A pigeon learning that pecking at a certain button releases food.

4-Latent learning

Latent learning is the association of indifferent stimuli

or situations with one another without reward. The
phenomenon is clearly exemplified in exploratory
behaviour. Animals finding themselves in unfamiliar
environments or among unfamiliar objects, but in familiar
surroundings, show exploratory behaviour. The animal uses
its sense organs on all that is novel, shows much
ambivalence between approach and avoidance, and, finally,
as the hesitancy to approach wanes, will test the novelty. A
mammal may, at this point, sniff, nudge, or handle a
strange object; a bird may peck at it. The first contact often
results in abrupt avoidance, but, typically, the object or
situation is, at last, thoroughly explored and finally
abandoned if neutral. A mouse sniffs and pokes about most
of a new environment. A bird, having sight and hearing,
rather than smell and touch, as dominant sensory modalities

55
does not have to occupy physically as much of a novel
environment. Instead, it places itself successively at several
vantage points and then carefully peers about and listens.

Subsequent behaviours of an animal can reveal that it has

learned much about its environment during such an
exploratory phase. It may learn, for example, the physical
features of the environment and their spatial relationships
to one another, the location of food and water, and the
location of places safe from predators. Animals apparently
learn all of these things early in their exposure to an
environment, even though the information acquired may
become significant only at some time in the future. The
learning takes place without being associated with
immediate reward (unlike that in conditioned response
Types I and II) unless a need to know is postulated as a
kind of immediate self-reward.

5-Insight learning

Insight learning is believed to be an advanced type of

learning. Insight involves the spontaneous combination of a
number of isolated experiences; the result is a new
experience that is effective for gaining a desired result.
Humans are able to exercise insight; it is extremely

56
difficult, however, to identify such behaviour in most other
animals. Animals are believed to use insight when they
solve a problem too rapidly for normal trial and error to
occur. It is possible that such an animal is carrying out
trials in its brain; this implies reasoning ability. The higher
primates are probably capable of insight learning at times,
but further down the phylogenetic scale the evidence of
such learning becomes progressively less conclusive.

Chimpanzees, to get food out of reach, will pile boxes

to make a stand for themselves or will fit sticks together to
knock the food down. These solutions may come quickly,
an obvious result of prior experience (latent learning).
Much trial-and-error learning is demonstrated, however,
when they actually pile boxes or fit sticks together. In
humans, insight is probably often aided by latent learning
and trial and error.

57
chimpanze uses sticks to extract termites from their nest

6-Imprinting

Imprinting, a learning process observed in young birds

and mammals, is the identification of an animal with
another animal. Normally, it is a relationship between
members of the same species, but it can occur, for example,
between a bird and a human. Imprinting can take place only
during a particular period of the animal's development—a
time span that is specific for each species.

In 1935 the Austrian ethologist Konrad Lorenz first

observed the process in ducklings and goslings. After
goslings hatch and become dry, they follow their parents.
The adults provide warmth, safety, and shelter and bring

58
the goslings food. The more uncomfortable a gosling
becomes (e.g., cold, frightened, hungry), the more intensely
it follows. If goslings are reared by a human, they become
imprinted to humans; thus, they ignore geese.

Ducklings follow their mother

The development of this response occurs during a

sensitive period, before and after which the response cannot
be learned; if the response is not acquired during the
sensitive period, it will never occur. Zebra finches that are
isolated from their own species before they are 35 days old
are never able to distinguish males and females of their
own species. This is because their sensitive period for
imprinting occurs before they are 35 days old.

The duration and time of onset of the sensitive period

depend on the species and on the type of behavior involved.
Some animals imprinted to animals of another species will

59
mate with members of their own species but, if given a
choice, will prefer the animal to which they have been
imprinted. Many species refuse social contact with any
animal except the one to which they are imprinted. Male
golden pheasants (Chrysolophus pictus) imprinted to
humans will court females of their own species but
immediately transfer this behavior to a human, should one
appear. The same is true for budgerigars (Melopsittacus
undulatus) and turkeys (Meleagris gallopavo). Mallard
ducks imprinted to humans, on the other hand, will not
associate with members of their own species (conspecifics)
and will continue throughout their lives to treat humans as
conspecifics. Imprinting is fixed for life, in contrast to other
types of learning, in which forgetting is common.
Imprinting of motor patterns, such as birdsong, also occurs.
Exposure to a particular birdsong may be relatively brief
and still be permanently fixed in the bird's memory.
Chaffinches (Fringilla coelebs) learn their songs during the
first 13 months of life, although they do not sing until
nearly a year later. A 12-day-old nightingale (Erithacus
megarhynchos) was kept in the same room with a singing
black-capped warbler (Sylvia atricapilla) for about one
week. The following spring the nightingale sang a typical
black-capped warbler song.

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 Little is known about imprinting in mammals, but
hoofed animals, such as sheep and horses, that are
imprinted to man express this response by following him
about. Dogs from four to six weeks old develop normal
social responses to dogs or to another species such as man.
Imprinting apparently also occurs in humans. An infant
deprived of its mother for a short period during its first year
may develop serious mental retardation. A separation of
several months—particularly during the seventh to the 12th
month—will frequently result in irreparable damage; under
such conditions, death may result.

Ducklings follow a dog as an example of imprinting to different

speceis

Play behavior and curiosity behavior

Play and curiosity are exhibited by many mammals and

by some birds and figure importantly in the learning of
numerous activities. Play is especially characteristic of
young animals, but the adults of many species also engage

61
in it. Spontaneous curiosity, in which the animal actively
seeks out novel situations for exploration, is exhibited by
the young of mammals and some birds; indeed, they seem
to be under the compulsion of some drive to do so.
Carnivores and primates exhibit more curiosity than
rodents, which gnaw novel objects and may hoard them.
Monkeys inspect and manipulate such objects.

The curiosity drive implements the development of new

motor skills and ensures the acquisition of new perceptual
impressions, thereby resulting in new knowledge. The only
reward, however, seems to be the performance of the
activities themselves. Rhesus monkeys will learn a puzzle
game without any reward except its successful solution.
Among rats, it has been observed that the nerve cells in the
lateral hypothalamus and preoptic regions of the brain are
more active in those rats that explore. Electrical stimulation
of these brain areas is rewarding to rats, and they will learn
to press a lever that activates this stimulation.

Such curiosity behavior seems linked to play behavior.

Play is difficult to define; it is usually easy, however, to
distinguish a playing animal from one that is seriously
occupied. An animal plays only when it is satiated and not
preoccupied with other tasks. Play seems not to be dictated

62
by immediate need but is extremely important in behavioral
development. Only animals that spontaneously seek new
situations on their own initiative play in the true sense.
Invertebrates, fish, and amphibians do not seem to play.
The taxonomic distribution of play among mammals and
birds suggests that play is related to learning. Play involves
interactions with the environment; this leads to the
acquisition of knowledge about environmental features,
including information about conspecifics and the animal's
own possibilities of movement. Play behavior occurs only
at particular times; progression to a second play activity
takes place only after a certain level of skill has been
achieved in the first.

Much play appears to be fighting or fleeing behaviour,

and usually it is easily identified as such. An animal that is
play escaping or play attacking does not actually escape or
attack. A rodent play fleeing into a hole, for example,
quickly reappears. If a rodent's flight is truly an effort to
escape, it reappears only after a much longer interval. Play-
fleeing animals often reverse roles quickly, and the pursuer
becomes the pursued. Threat behaviour that is associated
with real attack is missing, and there is strong reluctance to
bite. Play tends to be highly repetitive. A dog may retrieve

63
a stick many times or play fight until it is exhausted or until
a more interesting activity distracts it.

Such play behaviour could mistakenly be postulated as

the performance of immature instinctive activities. In many
instances, however, this is known not to be the case. Much
playful behaviour occurs at a time in an animal's life when
it is fully capable of serious activity. Play also involves the
use of species-typical patterns of behaviour in various
sequences that do not occur in serious activity.

Modification of instinctive behavior by experience

Behaviors based on both instinct and learning are

commonly intercalated into functional wholes. In the
peach-faced lovebird, for example, the cutting of nest-
material strips is partly instinctive and partly the result of
experience. The propensity for cutting is instinctive. This
includes punching holes in the sheet of material from which
the strips are fashioned and a “knowledge” of the proper
width, length, and straightness of strips. The spacing of
punch holes in such a manner as to form a strip is learned
through experience. It is as if the animal has an instinctive
picture in its central nervous system and persistently tries

64
punching holes, in various relationships to one another,
until the right pattern is made. The bird tends to repeat
punching patterns that most closely approach the ideal and,
thus, gradually, through progressively more satisfying
feedbacks, approach the definitive functional technique of
cutting strips. Idiosyncratic techniques develop—some
birds stand on the sheet while cutting, and others stand off
the sheet; some cut to the left, others to the right, and still
others cut in various combinations of these directions.
Birds developing their techniques try all of the directions
and places of standing but gradually act with less and less
variation. They do not need to observe experienced birds in
order to develop a technique. The stimuli necessary for
learning response are intrinsic to the birds' individual
activities.

If a meadowlark (Sturnella magna) is exposed to an

alien song during its sensitive period for learning song, it
will learn the alien song. Meadowlarks, then, learn their
species-typical songs rather than inherit the capacity for
particular melodies. But, if they are exposed during the
sensitive period to alien songs along with meadowlark
songs, they will learn only the normal species-typical song.
Although the song must be learned, the bird instinctively
learns the species-typical song if there is a choice. The
65
bullfinch (Pyrrhula pyrrhula) instinctively learns the male
parent's song rather than the species-typical song.
Bullfinches raised by foster parents of another species will
learn the song of the male foster parent, even though the
normal bullfinch song is also audible to it during the same
period. In both types of song acquisition, learned and
instinctive elements combine in the development of the
species-typical song. Some species, however, do not learn
their songs and do not need the experience of hearing
others sing during their development. Different
vocalizations in the same species are commonly acquired in
more than one way. Some are purely instinctive; others are
learned.

The behavior of an individual animal is the result of a

genotype that has developed over millions of years of
evolution—a genotype that also permits a certain degree of
variability through experience.

66
 Social and non social behaviour

A “random” pattern of distribution, in which animals

wander without regard to each other, brings asocial animals
together at times. An even distribution is much more
common, as in territories of many songbirds in which each
pair occupies its own plot of ground; these creatures are
actually social animals, in the sense that each interacts with
its neighbours so as to keep them at a certain long distance.
“Clumped” distribution of animals is also common; this
situation usually is truly social in the sense that each animal
interacts with its neighbours so as to keep them at a certain
short distance. Regularity of the short or long distance that
an animal keeps from its neighbours is thus as much an
indication of sociality as is grouping; only the theoretical
(and probably nonexistent) animal that completely ignores
its neighbours is truly nonsocial. Time is also a factor in
spacing; truly social animals have a tendency to move to
the correct distances from each other and to maintain those
positions over specifiable time periods, such as for the
morning hours each day.

Dominance hierarchies and division of labour

Social interaction in time and space is sometimes shown

to the ethologist by patterns of following and leadership.

67
 Dominance hierarchies may occur whether or not there
is social behaviour in the usual sense. Among several
species of birds that follow swarms of army ants in Panama
to catch insects flushed by the ants, the big ocellated ant
bird (Phaenostictus mcleannani) is dominant, the medium-
sized bicoloured ant bird (Gymnopithys bicolor) is next,
and the small, spotted ant bird (Hylophylax naevioides) is
chased by all the other members of the flock.

Crowding almost any two solitary animals together

will produce a dominance hierarchy, in which one animal
becomes boss or kills the other. This is a major cause of
deaths in zoos and aquariums, but it is not necessarily
social behaviour. Some biologists even say that dominance
hierarchies are evidence of antisocial rather than social
behaviour and are expressions of inadequacy in
overcrowded social systems. It is certainly true that most
peck orders appear in unnatural situations, such as among
chickens in a hen yard or animals in a cage. In most
animals, the absence of a dominance hierarchy, rather than
the presence of such, in a crowded context is a sign of a
high development of social behaviour.

68
Phaenostictus mcleannani

Gymnopithys bicolor

Hylophylax naevioides

69
A further interaction that the ethologist watches for in
social animals is

division of labour.

Any two animals will, of course, divide up food or any

other resource between them.

Males and females of some woodpecker species forage

in different places in the trees, taking different types of
food. Some animals use the same resource but do different
things to get it. Male and female birds often build nests
together, dividing the labour equally in some cases.

Social insects have more elaborate divisions of labour.

Animals that cooperate by division of labour tend to use
varied resources and to find more uses for them.

Division of labour seems to be a passing phase in the

evolution of most animals other than man. It is evidently of
little advantage to most animal societies. The honeybee, the
leaf-cutter ants, and other animals with division of labour
nearly always occur only where there is little competition
from more specialized animals. Few of the most advanced
insects or other animals show division of labour. In habitats
such as the coral reef and the tropical forest, division of

70
labour tends to be among different species rather than
within a species.

Social interactions of various types are more important in

determining degree of sociality than are most of the above
characteristics.

Altruistic social behaviour

Members of a group of Japanese macaques grooming each other. Grooming is a

type of altruistic behaviour that can extend even to unrelated individuals when the
behaviour is reciprocal and the giver's costs are smaller than the recipient's benefits.

Altruistic social behaviour is often found among

animals. An altruistic animal is one that expends some of
its energy helping another without direct benefit to itself, be

71
it a mother bear protecting her cubs against their hungry
father or a bird giving an alarm call that warns its
neighbours of a hawk. An alarm call may help the bird
itself, of course, by startling the predator or warning it that
this alert bird will be hard to catch. Psychologists have
found that a rat or monkey will slow its rate of pressing a
lever for food if that lever also gives an electric shock to a
nearby rat or monkey. Rats will take turns sitting on a
platform so that others can feed without being interrupted
by electric shock. Rats or pigeons can be trained to
cooperate in getting food.

Since the altruistic animal always loses something by

its behaviour, the question arises why altruism exists. One

answer is that, as the evolutionist Charles Darwin

suggested, when an animal protects its offspring, it helps its
kind to survive the process of natural selection. Altruism,
significantly enough, is usually limited to an animal's
relatives. Under some conditions, the survival of the group
may be more important even than survival of the
individual, as when the honeybee dies defending the hive.
The worker honeybee, which is not able to reproduce, is in
the biological sense not an individual so much as an extra
limb of a collective animal.

72
Reciprocal altruism, in which a benefit is later returned to
the benefactor, need not be between related animals and
may not even seem altruistic. Alarm calls of birds often
alert entirely unrelated kinds of birds, which later may
return the favour.

Individual and group recognition

Individual and group recognition are often important

aspects of social structure. Ovenbirds (Seiurus
aurocapillus) in North America recognize neighbouring
males by their songs and react aggressively mainly to songs
of strange males.

Ants, bees, and termites often attack strangers, or even

members of their own colony that have been
experimentally removed for a few days or washed. Many
kinds of parasitic insects (beetles, flies, butterfly
caterpillars), however, provide food or scents that gain
them entry to a nest, then prey on larvae there.

Age structure, birth rates, and death rates.

A young wasp or termite colony has few old animals, a

mature colony has more, and a declining colony or one that
is producing reproductive forms has few young. The old
colony has a lower percentage of foraging workers than

73
does the young colony, and has a lower birth rate and
higher death rate as a consequence; but only the old colony
produces reproductive forms.

perform movements

Such as nomadism and migration. Army ants wander

nomadically after prey. Wildebeest and locusts of Africa
emigrate to green areas of local rains; flocking or solitary
birds migrate back and forth to escape winter or drought;
anadromous fishes, such as salmon, move to the sea for
food and to rivers to spawn; catadromous ones, such as
eels, do the reverse.

Migrating birds

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Cooperation and competition

Cooperation and competition are major aspects of

animal social behaviour. Social facilitation, as when
yawning spreads through a pride of lions or chickens eat
more rapidly together, shows that cooperative competition
can be social. Social animals, which live close together,
often interfere and fight more with each other, especially in
early stages, than do solitary ones.

A swarm of ants cooperate to collectively move a leaf.

Construct things

A major external characteristic of the more complex

societies is that they construct things, or modify their

75
environments. The elaborate air-conditioned castles of
some termites, the path systems of feral house cats, and the
patterns of singing at dawn among birds of ephemeral or
coarse-grain habitats, all are “structures” created by
animals.

Termites castle

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Castle of termites

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 Types of animal societies

Parental societies

Parental societies are found at all levels, from the cell to

the monkey troupe. All animals provide for their young in
some way. In every animal there is a period when the
young is part of the parent and receives materials from the
parent. Later, the young may partly or completely separate
from the parent; in some animals, the more or less separate
young is then helped by the parent, or helps it.

Parental behaviour among simple organisms

Even some of the simplest organisms show colonial

aggregations of the parental type. Some viruses form
inclusion bodies in the cells they attack; these bodies are
thought to be colonies of daughter viral strands. Other
viruses form ordered arrays.

Bacteria, only a few steps up the evolutionary scale

beyond viruses, also show parent–young colonies. In a
large number of colonial bacteria, the offspring that are
produced by a dividing parent generally stay together for
some length of time.

Protozoa, Under difficult conditions, protozoans commonly

form a protective “cyst” and divide within it. In such
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divided cysts 2, 4, 8, 16, 32, or even more daughter cells
may associate until the cyst “hatches.”

From flatworms to insects

The annelid worm Ctenodrilus is said to be truly

viviparous, the nutrition of the young coming via maternal
blood vessels. Most molluscs take little care of their young.
Most squids release single eggs or chains of eggs, but some
members of the octopus group stay near their eggs and
remove debris from them.

Social insects

The leaf-cutter ants of the Western Hemisphere live in

huge underground colonies. They, along with termites and
a few beetles and moths, are agriculturalists. The leaf-cutter
ants cut strips of green leaves and make a paste of them in
which they grow fungus. Their underground chambers may
reach several yards. It is hard to realize that such huge
colonies are extended families.

Fish as salmon dig spawning nests in the bottoms of

streams. Stickleback males fan their eggs until the young
hatch, making sure there is enough oxygen. Other fish, such
as the black-chinned mouthbreeder, fan their eggs by
keeping them in their mouths

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 Among reptiles the best parental care is in alligators
and some crocodiles, where the female makes a mound of
dead leaves or sand and stays around to protect the eggs, to
release them when they hatch, and to guard the young for
perhaps as long as a year.

Birds are well known for parental care. Most build

nests, incubate eggs, and care for young. The males
commonly help the females in this. Often young birds stay
with their parents a year or more, and numerous examples
are known of the young of one brood helping to feed the
young of later broods. Often the young help the parents
defend a “group territory,” as among Australian bell
magpies (Cracticus) and Mexican jays (Aphelocoma
ultramarina).

The most advanced parental society yet recorded

among birds is that of the ocellated antbird, which follows
swarms of army ants in order to capture insects they flush
in the neotropical forests. This bird's young stay with their
parents for several months, then go and find mates, but
return to their parents periodically for several years. The
young bird and its mate are accepted as part of the extended
family; they are not chased away as often as are unrelated
birds.

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 An even greater organization of parental care is found
in mammals, except that the males seldom help care for the
young. The young are usually nourished before birth by the
placenta of the mother, except in the egg-laying duck-billed
platypus, the spiny echidna, and in marsupials. The young
of marsupials are born prematurely and grow in the pouch
of the mother for long periods.

Parental community in penguin

Commonly, groups develop around a mother and may

be joined by other such groups and by males to form bands,
troops, and herds. Troops of monkeys and apes are

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basically families or grouped families. These and wolf
bands include males, which may help to raise young. The
“extended families” of humans lead to tribes, states, and
nations.

Societies with sexual bonds

Nonparental social relationships fall into two categories,

sexual bonds and nonsexual bonds. Normally, only the
latter can involve members of two or more species. Sexual
bonds lead in many animals to parental bonds, of course,
but differ in that the bond is normally between offspring of
different families. The reason for this is that the main
advantage of sexual union is to combine the good genetic
features of two different lines. Some young, of course, will
have the bad features of both lines and will be eliminated—
wastage tolerated in nature as a necessary expense.

Courtship must basically ensure two things: that the correct

male and female get together at the right time with as little
loss as possible; and that the offspring have the best
possible chance to survive. Ensuring these two things has
led to elaborate courtship patterns in animals. Where
different kinds of animals that look, smell, or feel alike
coexist, each individual must be especially careful not to
hybridize with the wrong species. Otherwise it will waste

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its eggs or sperm in a union that will produce no young, or
young that are malformed or maladapted for the world into
which they emerge.

Animals often develop complicated odours, colours, or

voices as means of identification. There are many species
of fruit flies of the genus Drosophila, and to avoid
mismatings, each species has its own pattern of waving the
wings by the male. The sight and sound of the wrong
pattern of waving is enough to cause a female fruit fly to
flee.

After the golf ball was coated with a female sex attractant pheremone
this male crab proceeded to attempt to mate with it

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 Courtship displays in birds

Where there is only a limited breeding area or a patchy

environment in which the good areas are restricted, strong
male–female differences are advantageous. In these cases
the males have to advertise, often with song, to help
females locate good areas. Males of species whose
courtship displays are performed in groups usually have to
compete more directly with each other and thus tend to
develop large size or striking colours, overpowering scents
or voices, or other exaggerated features. Female domestic
chickens, sage grouse, and baboons tend to copulate mainly
with the lordliest and most dominant males. This is not so
evident in other animals, in which the females tend to mate
with the male holding the best territory. The dominant male

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is likely to be the oldest one, the one that has proved he can
survive and hence is “fittest.”

The success of the “supermasculine” or lordly males in a

few species may be advantageous only to those males. A
few lordly males often usurp the few suitable places to
breed, as in male sea elephants (Mirounga angustirostris)
on Pacific beaches and in male red-winged blackbirds in
North American marshes. If the lordly male blackbirds are
eliminated, however, other males come in and the females
breed with them just as quickly.

The supermasculine males in these species, full of pomp

and strutting, seldom care for their young. Baboon males, it
is true, do at times stop fights between lesser animals and
drive away leopards or other predators. But usually the
female takes care of the young herself. It may even be an
advantage if she is “ladylike” and unobtrusive, so that the
lordly male may draw predators away from the young.
Keeping the male away from the young may also allow the
female and her young more food. If environments are
limited in food, keeping excess males out of the breeding
area clearly is an advantage.

Super masculinity, as well as being correlated with

female care of the young, is also associated with polygamy.

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The mating of several animals of one sex with a single
individual of the other sex tends to be associated in birds
and mammals with great differences between the sexes.
Serial polygamy, or the mating of an animal of one sex
with several of the other sex at different times, may also
occur. Promiscuity, or the mating of each female with
several males and each male with several females, tends in
super masculine animals to resemble serial polygamy.
Monogamy, or the mating of each male with one female,
tends to occur mainly in animals with little difference
between the sexes.

Having many mates does not necessarily mean that an

animal is more social than if it has only one mate. In most
cases, the polygamous male spends much time driving
away other males and little time courting his females. His
females spend little time with him, because they are busy
raising many young—with little care for each. The whole
system of lordly males, ladylike females, polygamy or
serial polygamy, and multiple young tends to occur mainly
in animals with restricted and undependable sources of
food and other necessities. One investigator found that
males of the long-billed marsh wren (Cistothorus palustris)
in Washington, where their marshes varied greatly in
quality, had several mates; females went to males with
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good territories and left neighbouring males, with poor
territories, mateless. Another investigator found that in
Georgia, where the marshes were everywhere about equal
in food supply and nesting cover, the long-billed marsh
wrens were usually monogamous. The correlation
suggested by many recent studies is this: sexual
dimorphism and ditheism (behavioral differences) arise in
animals in which environmental opportunities are restricted
due to undependability or local distribution.

Nonfamilial social bonds

Social behaviour also occurs among animals that are

not necessarily related by parental or sexual bonds. A
“flock” or “band” of animals may be formed of only one
family in some cases, but often several families or
individuals join together.

Individual animal interactions

The most intimate form of interspecific association is

that known as symbiosis, or mutualism, in which dissimilar
organisms live together.

Multicellular animals often live on or beside other animals.

Small fish live in the tentacles of some jellyfish, sea
anemones, and even the Portuguese man-of-war, yet are

87
able to evade or inactivate the stinging cells. Some
hydroids and worms live on tubes of other worms or on the
shells of other invertebrates. The tubes of worms and
shrimp often harbour other worms or fish. At times, the
animal will live only on one kind of shell and is found
nowhere else.

Dynamics of social behavior

Costs and gains

Social behaviour among humans is often regarded as an

end in itself, the expression of a basic drive that has no
necessary purpose. Biologists doubt that any animal has
social tendencies without some adaptive advantage.

The costs

Social behaviour also attracts enemies. Groups of

animals have epidemics, while solitary animals seldom do.
Many disease-carrying parasites spread much more easily
at times when animals are together. Some rabbit fleas are
even adapted to the hormonal cycles of the rabbits, so that
they reproduce at the times of year the rabbits are
reproducing and hence are social.

Predators, like parasites, often have an easier time if

animals are crowded together; the animals are often busy
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reacting to each other and the predator can sneak up
without being observed. Their communicatory systems may
even attract predators. Tuna prey specifically on fish in
schools; a small hawk in tropical America (Accipiter
superciliosus) mainly on mixed bird flocks.

Social behaviour increases the number of interactions

between animals and thus the chances of conflict. The
conflicts may be solved by fighting, by patterns of
dominance and submission (peck orders), or by mutual
avoidance. Mutual fighting and mutual avoidance have the
same result—a partitioning of resources for which the
animals are competing.

The gains

Against these disadvantages of being social, it is

possible to set a number of clear advantages. They fall into
six broad categories, corresponding to the six possible
kinds of animal behaviour. By social behaviour animals
gain: (1) food and other resources, (2) reproductive
advantages, and (3) shelter and space. They are enabled to
avoid (4) physical and other small hazards, (5) competitors,
and (6) predators or other large dangers. The first and third
of these gains are reactions to desirable things of small (1)

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and medium to large size (3) respectively; the fourth and
sixth are reactions to undesirable things of these sizes.

1-Food

The value of being social in getting food is obvious in

the case of hunting bands. Cooperative hunting has been
found among wolves and African hunting dogs, hyenas,
lions, killer whales, porpoises, cormorants, white pelicans,
pairs of eagles and of ravens, tuna when chasing small fish,
army ants, primitive and modern men, and many other
animals. Animals that hunt cooperatively can trap, chase,
and tear apart prey that would otherwise be too fast, strong,
or large for them. In African hunting dogs the chase is run
by the leader of the pack, but the rest keep the antelope or
other prey from dodging left or right and also help fall on it
when the leader catches it. Flocks of wattled starlings
(Creatophora cinerea) fly after African migratory locusts
and surround one group after another, eating every trapped
locust from each group. In army ants, the individuals are
bound to each other by chemical “trail substances” so that
no individual gets far from the group; when one finds prey,
it grabs it and emits an “alarm” chemical that causes nearby
ants to grab, bite, and sting so that the prey is overwhelmed
within seconds. They then tear the prey, usually insects or

90
other arthropods, limb from limb and carry it back to the
nest.

Interspecific groups of birds are sometimes food-

getting societies. Drongos (Dicrurus species) of Africa
flush much food, and other birds follow them to get it.
Honey-guides (Indicator species) of Africa lead honey
badgers or men to bee nests and eat wax after the mammals
break open the nests for honey. Hawks have been known to
follow railroad trains for the same reason, and hornbills and
hawks follow monkeys. The birds, lizards, flies, and other
animals that associate with army ants offer other examples
of interspecific food-providing associations. One animal
may steal food from another, as American widgeons (Anas
americana) steal grass from redheads (Aythya americana).

In addition to hunting and flushing food cooperatively,

animals sometimes lead others to food or teach them to use
it. Parents, especially among mammals, often teach their
young to hunt or lead them to food. Animals that must
migrate or depend upon seasonally available resources
often depend on others to show them what foods are good
and where. Vultures and jackals flock to carcasses on the
African plains. American robins (Turdus migratorius) in
California have been observed learning to use certain

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berries after flocks of cedar waxwings (Bombycilla
cedrorum) came through and started eating the berries.
Tests with a tape recorder show that the recorded calls of
some birds that follow army ants will attract unrelated
kinds of birds that also follow ants. In the laboratory, some
animals learn to push a lever for food by watching others
get food that way and learn to avoid distasteful foods by
watching others cough it up. In studies of Japanese
monkeys (Macaca fuscata), the habit of washing potatoes
before eating spread from the younger to older monkeys of
a troupe. In Britain, a few titmice learned to open milk
bottles and drink cream; the habit spread much too rapidly
to be a genetic change.

2-Reproduction

The reproductive advantages of social behaviour have

mostly been discussed earlier. It was noted that sex is a way
of combining desirable genes from different lines, genes
that otherwise might slowly or never get together. In many
lines of animals, parental behaviour is clearly useful in
protecting or teaching the young. This normally requires
the adult to have fewer young. The careful parent loses in
time and energy and number of offspring but comes to

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prevail in evolution if it has more descendants than does a
careless parent that lets its young die. The careless parent
prevails if it can get more young out by caring for each one
less; some parasites are careless parents because each of the
young needs little care and a large number must be
produced to get to an extremely distant host.

3-Shelter

Social behaviour is often used in habitat selection and

shelter selection, even to the extent of making it possible
for the animal to improve the environment it finds. Male
birds that later will fight with each other over territorial
boundaries gather first at areas where they hear another
bird singing, rather than hunting for a more isolated (and
probably unsuitable) place. Certain beetles that attack pines
put out a scent that attracts other beetles; only as a result of
concerted attack by all beetles can the protective pitch of
the tree be reduced so that all may enter. Movement to a
flock is a good way to find a patch of habitat or a shelter. It
has been suggested that flocking increases the accuracy of
migration, since the average direction taken by a flock is
more correct than the individual directions taken by
individual birds. Small flocks of European starlings
returning to a California roost were less accurate in their

93
direction than large flocks. Cooperative building of
structures is well known in humans, prairie dogs, rats
(whose tunnel systems rival the catacombs in complexity),
beavers, certain weaver finches, wasps, bees, termites, and
many others; symbiotic use of structures occurs in many
animals.

Naked mole rats

4-Hazards

Social behaviour can also help animals avoid small

hazards. This includes avoiding heat or cold and wet or dry
situations as well as preening or grooming to keep off dirt,
parasites, and other small environmental hazards. A goose
cleaving the air for its companions at the front of a V-

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shaped flock, a parent bird brooding its young or sheltering
it from the Sun, a group of creepers roosting together to
help each other survive the cold winter night, and a group
of baboons grooming each other to pick off ticks furnish
other examples.

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 References

Barnard C.: (2004). Animal behaviour, mechanism,

development, function and evolution. Pearson and Prentice
Hall.

Bollinger T. and Schibler U: (2014). Circadian rhythms –

from genes to physiology and disease. Swiss Med
Wkly.144: w13984.

Drickamer L.C., Vessey S.H., Jakob E.M.: (2002).

Animal behavior. McGraw-Hill Companies, Inc.

Scott G.: ( 2005). Essential animal behaviour. Black well

publisher.

Zee P. C., Attarian H., Videnovic A: (2013). Circadian

rhythm abnormalities. Continuum (Minneap Minn).
19(1):132–147.

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