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Course Animal behavior by Girma Mengesha (PhD)

Introduction

What is Animal Behavior?

Animal behavior is the scientific study of everything animals do, whether the animals are single-celled

organisms, invertebrates, fish, amphibians, reptiles, birds, or mammals. It involves investigating the relationship

of animals to their physical environment as well as to other organisms, and includes such topics as how animals

find and defend resources, avoid predators,

choose mates and reproduce, and care for their young.

Who studies animal behavior? Etiologists or Psychologists?

Ethology from Greek, ethos, "character"; and -, -logia, "the study of" is the scientific study of animal behavior,

and a sub-topic of zoology.

 Ethology is a combination of laboratory and field science, with a strong relation to certain other disciplines —

e.g., neuroanatomy, ecology and evolution. 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.

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Thus the biological study of animal behavior is Ethology. It is the natural studies of animal behavior. Interest in

behavior received its most important boost from the writing of Charles Darwin since he included the chapter on

“instinct in the Origin of species” and also wrote a book specifically about behavior called The Expression of

Emotion in Man and Animals. However, in half –century after Darwin, there was little work on the animal

behavior. Therefore, it was in the 1930s that compressive theory of animal behavior began to emerge through

the writings of Knord Lorenz and latter Nikio Tinbergen. In those years, ethology can truly be said to have been

born. Then in 1972, it came of age as a science when Lorenze and Tinbergen won noble Prize for Physiology

which they shared with Karl von Fricsh who discovered the remarkable dance of honey bee. The dance of

honey bee enabled the forgers to tell the place for good food sources.

Observing and describing exactly what animals do is festinating study by its own right and in the study of

animal behavior and essential prelude to more scientific analysis of animal behavior. As a result, money

ethologists spend long hours patiently watching their animals. Through careful description, ethologists able to

make inventory, ethogram, and the behavior pattern possessed by each species that they study. For the casual

observer, it might seem that different species of birds and fish behave in much the same way. It might seem that

the behavior of each animals is a highly varied business, not easy to split into a particular type. Fortunately,

these are not true for most animals species. Each species tends to have an array of stereotyped behavioral

patterns some which may be shared with related species but others of which are unique to itself.

According to Niko Tinbergen there are four sorts of questions that could be answered related to animal behavior

These are 1.Causation, 2. Function 3. Development 4. Evolution of behavior

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1. Causation

The causes of behavior include both the external stimuli that affect behavior, and the internal hormonal and

neural mechanisms that control behavior. For instance the question why starlings, Sturnus vulgaris, sing in

spring could b answered by internal and external causes. The internal factor could be the increase in day length

that triggered change in hormone levels in the body or and external factor could be air flow through the syrnix

and set up membrane vibrations.

2. Function
The functions of behavior include its immediate effects on animals and its adaptive value in helping

animals to survive or reproduce successfully in a particular environment

This is related to survival value of behavior, Starling sings to attract mates

3. Development
The development of behavior pertains to the ways in which behavior changes over the lifetime of an

animal, and how these changes are affected by both genes and experience. Starlings sing because they have

learned the songs from their parents and neighbors

4. Evolution

The evolution of behavior relates to the origins of behavior patterns and how these change over generations.

The answer would be about how song had evolved in starlings from their avian ancestors. The most

primitive living birds make very simple sounds, so it is reasonable to assume that the complex songs of

starling and other song birds have evolved from simple ancestral calls.
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Reproductive behavior in Lions (Pnthera leo)

In the Serengeti National Park, Tanzania Lions live in prides consisting of between 3 and 12 adult, females

from 1 to 6 adult males and several cubs. The group defends territory in which it hunts for prey, specially

gazelle and zebra. In the pride all the females are related they are sisters, mothers, daughters’ cousins and etc.

All were born and reared in the pride and all stay here to breed. Females reproduce from 4 to 18 years and so

enjoy large reproductive life.

For the males, life is very different. When they are 3 years old, young related males (sometimes brothers leave

their natal pride). After a couple of years as a nomad, they attempt to take over another pride from old and week

males. After successful take over, they stay in the pride for 2 to 3 years before they in turn are driven out by

new males. The male reproductive life is therefore short.

The lion pride thus consists of a permanent group of closely relatively species, females and smallest group of

separately interrelated males present for short period of time.

Based on Bertram (1975), we consider three interesting observations about the reproductive behavior of a lion

1. Lions may breed throughout the year but although different prides may breed at different times, within
a pride all females tends to come into oestrus at about the same time. The causal explanation may be

influence of individual’s pheromones on the oestrus cycles of other females in the pride. Similar

situation may explain why girls in school at dormitory synchronize their menstrual cycle, perhaps due

to the effect of pheromones. The function of synchrony in lionesses is that the different litters in the

pride are born at the same time and cubs born synchronously survive better. This is because communal
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suckling, with all the females lactating together a cub may suckle form another female whenever it

mother out for hunting. Moreover, synchrony birth increases chance that a young male will have a

companion when it reaches the ages at which it leaves the pride which is helpful for successful

takeover of another pride.

2. A lioness comes into heat every month or so when she is not pregnant. She is on heat for 2 to 4 days
during which time she copulates once every 15 minutes throughout the day and night. Despite the rate

of copulation, the birth rate is low. Even for those cubs that are born only 20% will survive to

adulthood. The causal explanation why lion mating are unsuccessful is not the failure of the male to

ejaculate but the high probability of ovulation failure by the female or high rate of abortion.

One hypothesis is that it is a for the female to be receptive when conception is unlikely because each

copulation is devalued

For male since there is only 1:3000 chance of that a given copulation will produce surviving cub, it is

not worth fighting with other pride over single mating opportunity.

3. When a new male, or a group of males, takeover the pride they sometimes kill the cubs already present.
The causal explanation may be unfamiliar odor of the cubs which induces male to destroy them. A

similar effect in rodents known as Bruce effect occurs in rodents where the presence of strange male

prevent implantation of fertilized egg or induce abortion.

The advantage of infanticide for the male that takeover the pride is that killing the cubs fathered by a

pervious male brings the female into reproductive condition again quicker and hastens the day that he

can father his own offspring. If the cubs were left intact the female couldn’t come into oestrus again

for about 25 months. By killing the cubs she becomes ready after 9 months. Remember the male’s
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reproductive life is short so any individuals that practice infanticide after taking over the pride will

father more offspring and therefore tendency to commit infanticide will spread by natural selection.

Assignment read a book, Introduction to Ethnology By P.J.B.Slater (1985). And also Introduction to

behavioral ecology by Krebs and Davies, 1993.third edition

Study the case study on introduction to ethnology and answer question about cause, function development and

evolution of animal behavior using the example of three- spin Stickleback

What are the two schools of thought of animal behavior? Discuss their similarities and differences?

Zigzag dance….swollen belly on intruder (external), male sex hormone (internal) presence of nest

Head down threat posture……male sex hormone, red belly on intruder

Fanning…..male sex hormone, eggs in nest, presence of nest

The two schools of thought the two group…the two group shared an interest in the behavior of animals.

American school of comparative psychology…..

Tend to study very few spp., such as rats and pigeons, interested to look general law of behavior regardless of

the spp. being studied preferably apply to man .their reputation has for rigorous experimental work in carefully

controlled laboratory condition. Put their animals in the small box and peers into what it is doing.

American school of Ethology

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Have respect for evolution, interested in a wide variety of species, and the different ways in which they

behaved, put himself in a small box and looks out at what the animals round about are up to.

Motivation

When confronted with an appropriate stimulus, an animal does not allow the same response. For instance A

male stickleback on his own tank and shown a model of female for a brief period everyday may sometimes

court with its great vigor, yet on other occasion, it ignores it. Therefore, any one studying animal behavior soon

come to realize that it varies enormously from time to time, and this is frustrating if one hopes that animals will

always behave in the same way. But on the other hand, discovering just what is that leads them to behave

differently from one time to the other presents some very interesting problems which are well worth studying by

its own right. These are the problems of motivation, or in other words, the mechanism leading animals to do

what they do when they do it.

What motivate animals is from within is a much more difficult subject than from the outside world which

affects its behavior , and is can be studied in a very different ways. One is to look inside and see what is

happening there, for example by stimulating nerves or recording from them. The hope is to find pathway

between the sense organs and the muscles and the physiological factors which affect them and influence

behavior. The other approach for the ethologist is to treat the animals as a ‘black box’, changing the input to it

in various ways. For instance by depriving it of food or introducing rivals to it and see how its behavior alters

rather than looking inside what mechanisms are involved. Just like one need not understand how a computer
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works to program it and understand how the output it provides, a lot can be found out about motivation without

any knowledge of the hardware responsible for it inside the animal. In thinking about motivation, it is worth

considering, two different aspects separately. One is the tendency to behave in a particular way fluctuates

over time, and these involves looking at different type of behavior and the factor affecting them without

much concern for other activity. For example, one might study how eating is patterned in time and how it

is affected by period of food deprivation. The second is to examine how the animal decides which of the

many behavior patterns at its disposal it will perform. In this case one might look to see whether a rat

which is hungry and also thirsty decides to eat or drink first. To examine let see using simple model

Model of motivation

Since we experience hunger at some time and not at others, we realize that urge to eat is not constant. The sight

of delicious meals is one in some occasion mouth watering and on the others, no interest at all. Konard Lorenze

has put this in the form of a model. The word model refers to a simple proposed scheme that work in a way

similar to real system. This was called as Lorenze’s ‘Pschohydrolic’ model or Lorenze’s water closet.

The model is not something to look inside but it is what is called an’ as if’ model, animal may behave as if it

had such a system for organizing its behavior within it. Lornze supposed that different actions depended for

their appearance on a supply of ‘action specific energy’ which accumulated with time since the animals last

behaved that way and was used up as it preformed the act. He visualized this as water accumulating in a tank

out of which it could only escape through the valve at the bottom. The valve was however, a rather strange

one. It could open either by water pushing within it or by string attached to the scale pan pulling from outside.

To Lorenze weights on this scale-pan were the equivalent of stimuli leading the behavior, the more appropriate
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the stimulation, the heavier the weight and the more likely the valve was to be opened. Lorenz’s model has

some interesting properties which can be compared with real behavior. First, for the longer, since the behavior

was last preformed the more action specific energy will have accumulated and the more likely it is that the

behaviors appear. Secondly the model suggest the accumulation of action specific energy lead the behavior to

occur even if the stimuli present are slight ( the weight on the scale-pan are light). Ultimately, Lorenze argued

that behavior an animal has long been deprived will appear as ‘vacuum activity’ with no stimuli present at all.

These are external stimulation. The idea of vacuum activity; when there was an excess of action specific energy.

At the other was exhaustion which occurred when an activity had run out of this energy. The way the valve

work suggests the a particular relationship between internal factors and external stimuli; provided that there are

action specific energy, the push of this and the pull of external stimuli will add up to give rise the behavior

rather than being multiplied together or related in some complicated way.

There is no doubt that certain aspects of behavior do become more likely the longer the gap since they last

appeared. Feeding and drinking is the most obvious example as well as we all known from our experiences.

There are very good reasons for these nutrients are used up by the body and water is lost constantly form it, so

both must be replenished and the need to do so will rise with interval since the last meal or last drink. However,

since the many activities an animal perform are not concerned with the regulation of their physiology, there is

no reason to think in advance that they might be more likely with time.

Many behavior patterns, such as singing, in birds, mating in fish or exploration in rodents, could be used to

illustrate this point. However, aggression is appropriate. Although Lorenz did not mention in his Pschhydrolic

model, in his book on Aggression, he suggested that aggression is innate drive which rises with time and
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somehow be expended, his view was that innate behavior is inevitable. Example, a mouse place in isolation in a

cage in its own. The more it has been placed in isolation, the more it fights another mouse which is put within it.

Many factors affect aggressive behavior such as hormones and shortage of food from the inside and presence of

rival and contested resources on the outside. Furthermore, aggression is complex behavior which may be caused

by quite different factors though they seem superficially similar ( see Figure 4.3. and 4.4 on Pages 56 and 57).

Will aggressive animal have to perform a certain kind of fight or cease after its rivals repelled?

Animals have a tendency to regulate their behavior as they go along in the light of its consequences: a process

called feedback. Behavioral pattern lies on both the external and internal stimuli and the extent and influences

of each of them vary depending on species. For example, a rat only drink if it is thirsty and presented with water

and eat if hungry and presented with food but consider two other example the exploration by Mice and song by

birds, mice’s exploration is mainly influenced by the external environment and song of birds by internal factor (

Page 48).

Deciding what to do

Animals generally do one thing at a time ,yet they often have the need to do several. For instance, a caged zebra

finch walking up after a 12 hour night must have long list priorities. The birds normally eat and drink about

every half an hour, sing and fly around the cage for a period after this and groom and rest till the next meal. The

male zebra finch thus shows sequences of organized behavioral patterns.

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How they decide between them? Yet no definite answer. Internal and external factor relevant to do different

activities.

The animal must make decision. it must have some means of weighing against each other.

Psychologist== action specific energy==the more==the priority

Ethologist==instinct==brain ==Tinbergen hierarchal model

Tinbergen hierarchical model= a center in the brain controlled each major instinct. Energy flowed down to

lower centers when a gat was opened by the presence of appropriate releasers.

Current view of motivation does not see animals as endowed with a serious of separate drives or instincts. But

influenced by iternal and external factors.

Displacement activities

Watching a pair of courting birds, the observer might be surprised to see one of them break off to preen briefly

and hurriedly before carrying on with its courtship. Similarly the fish in the middle of threatening a rival may

suddenly swim down and start digging in the sand as if switching to nest building or cock might interrupt

fighting to take few picks of food. Such actions that seem irrelevant to those who saw them are labeled as

displacement activities. It arose when the animal unable to perform the normal activity of the more

relevant behavior. The finding is that the casual factors that affect the normal behavior also affects it i.e.

displacement activities. For example the funning by stickle back is motivated by the concentration of carbon
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dioxide in the water. Birds in fight show grooming the behavior when their feathers have become weter. the

process is called disinhibition.

fighting = grooming (disheveled feather=enhance stimuli to groom)

Function of animal behavior

The ward function has different meanings. However; biologists give a very specific and precise meaning. The

function of a behavior pattern is its selective advantage or survival value; this is the reason why individuals

possessing it are thought to do better. Why do birds sing? If one asks different people, one would be likely to

get very different replies; some may say that it was because of sunny weather or because they are male and have

appropriate sex hormone circulating in the blood. These are however, casual explanation rather than function.

Other people might the say the birds sing to attract mate or drive a way rival. These are advantageous songs

may have and are possible functional explanation that is based on consequence the behavior rather than cause.

Nevertheless, all, behaviors has both cause and function and its understanding also involves knowing both.

Some functions are easy to understand, without the need of a lot of argument or for complicated experiments.

The functional significant of escaping a predators or producing egg is obvious. It is for survival and

reproduction and are clearly of selective advantage Perhaps bird’s song does exclude rivals, but what is the

function of that? Less competition for food may be one reason, that is, it helps the animals to live, surviving
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young which favors the animal. But it takes several links in the chain of argument (fig 7:1). Understanding the

functional significance of behavior involves discovering how it increases the transmission of individual’s gene

into the next generation. This is the ultimate function of behaviour. (more of own young fledge).

An important point is that behavior patterns don’t necessarily have only one function. Indeed any

consequences that they have which lead their processors to more successful may help to maintain them in the

population. Bird’s songs exclude rivals, attract females, and stimulate reproductive system of females: ovaries

grow when male songs and eventually lay eggs even the in the absence of male. Song may also have quite a few

disadvantages.

Birds may be sitting target for a hawk, spend time. The most important point is for behaviour pattern is that

its advantage and must outweigh the various costs. Communication may have both advantage and disadvantage

but the former must outweigh the later if it is to be favored by selection.

Experiment and Observation

A problem with studding selective and adaptive significance of behaviour is that it has evolved to function in a

particular environment so it can only be understood in relation to that environment.

In the study of behavior much more information comes from observation where experiment’s not possible.

Observations are most fruitful where they also involve comparisons between different species or how one

species behaves and how one species behaves and how theory predicts that.

Experiment
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The classic example of an experiment on function was done by Niko Tinbergen and his colleagues on egg shale

removal in black headed gules. Short after a chick hatches, these birds remove the broken shell from their nest

and carry it away. Tinbergen asked for the function of this behaviour, Perhaps the chicks were sometimes cut

by the broken egg shell, or predators might be attracted by the conspicuous white inside the broken egg shall.

To test the last idea, Tinbergen laid a number of artificial nest containing gull’s eggs some with broken shell.

He later found out that the presence of shell indeed led to the greatly increased egg loss by predators. John

Krebs also came out with similar result great tit where one of the functions of song is “to keep out signal to

male great tit function of which is to advertise ownership of territories & repel rivals.

Student of Niko Tinbergen, Esther Cullen, did work for her doctorate perched on cliff watching kittiwakes,

small gulls nest in dens.

Their behavior is related to cliff-nesting habit. The Galepage swallow tailed gull similar has behavioral. Tim

Clutton-Brok studied on two closely related east African monkeys- the black and white colobus & red colobus

overlap in their range and both live in forest where their diet primarily consists of foliage (fig 7:6). However,

they are different in many other ways. The red colobus is restated to wet forest where it lives in troops of 40 or

more ranging over about square kilometer they feed on flower, fruits, shoots & leaves of marry different types

of trees. The balk and white by contrast lives only on dry forest where it feeds on mature Leaves of few tree

species. It is troops size usually is less and its range is much more restricted. The reason for the difference may

be the red requires more varied diet where some trees are in fruit all times of the year. They also require feed on

many species, for efficient resource use with large range etc.
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Development of animal behaviour

This is the most contentious in the whole field of animal behaviour. Ethologists stressed and used the word’

innate’ ‘inherited’ and ‘instinctive’ to describe the behaviour patterns they studied. Thus, the German word for

fixed action pattern Erbkooordination is to mean ‘inherited coordination’. The sensory routes through which

these actions are elicited where termed innate releasing mechanisms and the motivational system controlling

behaviour from within is referred to as instincts. All these implied great rigidity in behaviour and its

development with no scope for environment to have any influence and as if the actions are inevitable.

According to this the whole form of development is simply to readout genetic instructions. This indicates the

springing of fully formed behaviour from animal at the time that it is required without involving complicated

development processes that involves interaction with the environment. However, it contrasted with the

‘Behaviourist ‘school of psychologists in America which stressed the prime importance of learning in the

development of behaviour. For many years the main issue in the development of animal behaviour is the

controversy between learning and instinct

5.1. Psychologists and learning

The major figure in the behaviourist school of psychology was J.B.Watson who followed John Locke in

believing that innate mind is ‘tabula rasa’, a blank slate on which subsequent experiences was written. Watson

studied learning in animals such as rats and was impressed by the flexibility of their behaviour and by how

learning could lead them to behave in adaptive manner. He believed that the animals built up connection in their
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brains due to perceiving associations in the outside world so that they associate a particular place with food run

through a maze to get there. Watson’s views were partly stimulated by the work of Ivan Pavlov, who studied

conditioned reflex in dogs. In Pavlov’s experiment associations were built between the stimulus which normally

stimulate make animal to elicit reflexes (food in dogs) and another stimuli bell ringing. This sort of training if

referred to as conditioned and the response is called conditioned response.

The main stress on Pavlov and Watson was not on reward: the dog did not need to eat the food to form the

connection. Nevertheless, later ideas suggested that great deal of learning does rely on reward, the most

exponent of this view being B.F.Skinner. Skinner also believed that the great majority of behaviour, with the

exception of simple reflexes, results from experiences. But he stressed the way in which the response of animals

comes to be linked to particular stimuli as a result of reward. Animals tend to repeat rewarded actions and

reward can thus be used to alter their behaviour. The dove in the Skinner box soon learns that pecking the right-

hand key gives it water while the left-hand key yields food. Based on ideas such as these, elaborate theories of

animal learning have been built up which stress the role of the environment and of experience rather than

genetics and inheritance in the development of behaviour, At another extreme such psychologists have

suggested that learning in all animals is subject to the same rules and that animals do not have predispositions

about the sort of things they will learn but will build up any associations with equal ease provided that their

sensory and motor systems allow them to do so .Obviously one would not attempt to train a sparrow to fly in

the dark or a tortoise to walk on its hind legs.

The following famous assertion of Watson’s illustrates the favour of his belief that the environment rather

than hereditary was the major determinant of behaviour. Give me a dozen of healthy infants, well-formed and
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my own specified world to bring them up in and I’ll guarantee to take any one at random and train him to

become any type of specialist I might suggest- doctor, lawyer, artist, merchant-chief and, yet, even beggar-man

and thief, regardless of his talents, penchants, tendencies, abilities, vocations, race of his ancestors. He went on

to admit that he was overstating his case, but insisted he was doing so as a counter to the sweeping claims of his

opponents.

Both views such as this and those of the etiologists could not be right, but to some extent they existed

because their adherents were studying different things. Psychologists were interested in learning and in

behaviour patterns, like the lever pressing of a rat in a skinner box which could be modified by it so leading an

animal to behave differently from other members of its species? By contrast, the ethologists were interested in

more fixed behaviour such as courtship displays that were common to all members of a species. Why did they

consider such behaviour to be innate and how does the evidence they used hold up to detailed examinations?

5.2. Ethnologists and instinct

In 1828 a boy of about 16 was found in the market-place at Nuremberg in Germany. He could not speak and his

behaviour was like that of a little child. So he was labelled the wild boy. His name was Kaspar Hauser and

when he was able to communicate, he explained that he had been brought up entirely in isolation by a man who

had kept him and cared for him all by himself in a hole. Kaspar Hauser may well have been a fraud, but he has

given his name to the deprivation experiments which have often been used by ethologists in an attempt to

understand whether or not the behaviour they were studying was innate. The idea was to deprive an animal of
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relevant aspects of experience and if the behaviour appeared despite this it could be regarded as inherited

whereas, if it did not appear of developed abnormally it could be assumed that the missing experience was

important.

There are many striking examples of behaviour appearing apparently normally in animals whose experience

is distinctly abnormal. A case in point was shown by experiments carried out by Janet Kear on ducklings which

she had reared by hand from the egg. Most duck species nest on the ground but some of them, such as the

wood duck, do so in holes far up in trees from which the fledglings have to leap to the ground beneath

.Fortunately they are light and fluffy when they do so and as a result, they bounce and run off rather than

suffering multiple fractures. Kear tested chicks of various species on a visual cliff such as that shown in Figure

5.4. In this apparatus the animal is placed in the centre and can move either to none side where there is a drop

looking like a cliff beneath the glass or to the other where the floor is immediately under the glass so that it

looks shallow. The behaviour of the ducklings was appropriate to their normal nesting place, Tree-nesters did

not avoid the deep side of the cliff but, if they moved that way they would leap as if casting themselves into

space. On the other hand, the ground- nesters tended to move to the shallow side rather than the deep one

suggesting that they avoided heights. Interestingly, if they did move to the deep side their behaviour was quite

different: they pushed off with both feet as they would when moving out from the edge of a pond! Being hand

reared these young birds had had no opportunity to learn, the actions they showed from other or from earlier

experiences with ponds or with cliffs. Many ethologists have used such evidences to argue that behaviour is

innate because it develops despite deprivation of opportunities for learning, and the main motivation of making

such experiment has often been to discover whether behaviour is ‘innate’ or ‘learned’. Though, ducklings were
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reared in a very impoverished environment their behaviour develops normally. Another good example is the

hording behaviour or squirrels where they bury nuts underground to form store which they eat later when food

is scarce, even in captivity. By contrast to this experiment others have shown behaviour patterns to be radically

altered unless particular behaviours are available. For example if young dog is reared in total isolation from

others, in a chamber in which it can be fed and cared without contact with even its human caretaker it turns into

strange animal, apparently careless of its own welfare. It will put its paw in a fire and sting it and it repeatedly

approach an object that gives it an electric shock. This is quite unlike the normal puppy of the same age which

hastily withdraws from such painful experiences. Furthermore, the normal puppy will yelp when hurt where as

the one develops in isolation behaves as if it doesn’t even feel pain. This clearly showed the deprivation has led

to drastic alteration in the way its behaviour developed. Another example is the case of chaffinch reared out of

earshot of songs of all of birds of its own species which is becoming simple and unstructured and unlike the

adult normal chaffinch.

In general the Kasper Houser experiments vary enormously in what the actually deny the animals.

5.3. A false Dichotomy

The deprivation experiments by ethologists were the main evidence in deciding whether or not particular

behaviour is innate. But the above experiment pointed out problems in interpreting such experiments. American

psychologist Danny Lehrman raised the following points for discussion.

1. He argued that deprivation could show where behaviour was important but not where it was not. Thus
the chaffinch which sings abnormally after being isolated shows that the hearing is essential for normal
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song development. But the birds that build its nest the first time it encounters hay had many experiences

from grooming its own feathers to husking seeds which could have contributed to its nest building

capacity. It rush to assume that simple deprivations experiments will remove all the experiences that

may be relevant.

2. Viewing behaviour either/or ie. Either innate or not

Konard Lorenz believed that behaviour is fixed and sow as ‘blueprint on the genes’. However Lehrman pointed

out that such expression gave false impression on the development of animals behaviour. By depriving of

certain behavioural development, one can show that they are unnecessary for its behaviour to develop normally

but where should we stop? One may predict specific cases where particular behaviour develops through

experiences. For example nest building development through experiences with net materials and seeing another

animals building nest where the experiences during feeding and grooming may be important which is similar to

manipulating thing in the mouth.

Case study Song development in birds read on page 88.

Learning ….chaffinches, parrot, hill mynah and innate…..dove (cooing)…cocks (crowing)

Recognising prey and predators

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Animals should distinguish what eats them and what they eat. This is important if the animal specializes on few

types of food or if eaten by few predators. On the other hand animals like rat and human being eat varies of

food and must distinguish those that are nutritious or not. Threats also vary. Killer whale, polar deer and human

with gun are lethal to a seal. Food preferences differ in different species. For example Newborn snake strong

species difference in their response to various food items which are in line with that of the adults. On the other

hand preferences can be acquired to adjust their feeding to much availability. Example rats seem to learn about

food by themselves or from others picking information

5.4.3 Social development.

Most animals do not have an entirely solitary existence, but must develop social relationships with other

individuals of the species. Initially this may be mother or father to whom an attachment grows early in life

ensuring that the young animals stays close by and doesn’t wonder into danger. Example, Rhesus, monkey. in

birds, it is called imprinting.

5. Evolution of animal Behaviour

Evolution acts because of the difference in individuals; those doing better leave more of their genes than those

doing worse so that fit genes spread. Evolution of behaviour is in part based on genetic differences among

individuals up on which it acts. The effects of genes on behaviour have taken several forms. Many genes affect
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each behavioural patterns and each gene influences many actions. Therefore, there are plenty of genetic

variability up on which selection could act. Studies using artificial selection confirmed this point.

The most satisfactory examination of the evolution of animal behaviour is the study of displays. Consequently

comparisons of displays have helped especially to reconstruct the course of evolution on animals (see on the

separate PDF documents).

Reproductive behaviour

In most species all individuals of one sex adopt a similar reproductive strategy and thinking about cost and

benefit why they follow this one rather than any of the other possibilities. In some cases it is clear from this that

one breed system is indeed the best for all members of a particular species so that no animal that happened to

adapt another one could do better that if it conformed. In the terminology by John Maynard Smith, there is in

this case a single ‘evolutionary sable strategy or ESS which it benefit all individuals to follow. In other cases

however, a mixed ESS can occur it pays some individuals to do one thing and other another. The foregoing of

bumble-bees is a good care here, not from point of view of reproduction. The follower it is the best for a bee to

visit are those being least exploited by others, so the bees end up with different strategies. The male

reproductive behaviour of the male green tree frogs provides another instance. These animals form assemblies

besides ponds and set up calling choruses to which the females are attracted. However, calling is dangerous

and energy consuming as well as making it hard to keep a look out for the females that approach. Some males

avoid this disadvantage by adopting ‘satellite’ status instead of calling. They sit and wait near calling males and

watch out for females, intercepting them as they arrive and often succeeding in mating with them before they

reach the caller to whom they are attracted. Not all males can become satellites, however, because then the pond
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would be silent and no females would come, so the stable situation is a population consisting a mixture of two

sorts of frog. These examples drawn from the breeding behaviour of animals ,show just how powerfully modern

evolutionary theory can shed light on various aspects of behaviour of animals .

Selfishness and altruism

So far we have seen that natural selection designs individuals to behave in their own selfish interest and not for

the good of the species or for the good of the group in which they live. For example observed clutch size,

foraging behaviour and mating patterns are what could be expected if selection optimized behaviour and life

history strategies so as to maximize an individual’s reproductive success. However, it will be obvious to

naturalist that animals do not behave selfishly all the time. Often individuals apparently co-operate with others.

For example, several lions may co-prate to hunt prey, in many species of birds and mammals, individuals give

an alarm call to warn others the approach of a predator, sometimes individuals may help others to produce

offspring rather than have young itself. Before 1960 co-operation did not demand special attention but since

1960s people have appreciated Darwin’s statement that in evolution there is struggle between individuals to

outcompete others in the population. How can we account the evolution of co-operative behaviour in terms of

advantage and disadvantage to individuals? If natural selection favours individuals who do the best and have the

most surviving young, how can behaviour evolve which involves helping others to survive and have young at a

cost to the helper’s own chance of doing so? As of E.O Wilson (1975), the central problem of socio-biology

is how altruism evolves? Altruism is defined as acting to increase another individual’s lifetime number

offspring at a cost to its own survival and reproduction. Example young of many bird species including

European long tailed tit help parents in feeding their brood

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Kin selection

Assisting close kin may actually help an animal to get more of its genes into the next generation, though in

more roundabout way than by breeding itself. This theory is referred to as ‘kin selection’, because it explains

why animals are prepared to behave in a way that involves sacrifice on their own part but advantage to their kin.

This just maximize not their own fitness but what Hamilton refers to as their ‘inclusive fitness’

Communication

Acts of communication between animals tend to be very fixed in form, to act as releasers for other actions and

to develop very similarly in all members of the species. Whenever, we see animals that are brightly colored or

boldly patterned, or one that is making a loud or obvious noise we can be certain that communication is taking

place. Most often communication is between members of the same species and the risk come when predators

cue on this as a means of finding a meal. Sometimes however, different species communicate with one another,

there is often prey animals that forsake their camouflage and communicate directly with the predators that

might eat them

Social Organization in animals

In the earlier chapters we have emphasized that natural selection acts on individuals, favoring those that are well

adapted and whose genes thus spread at the expense of those others but not selecting for one group than other.

Societies have thus not evolved through the action of natural selection up them, but they have emerged as a

result of the way selection has affected the behavior of their individual member. Nevertheless, different animal
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species occur in very different social structures and though each may strive selfishly, to maximize his inclusive

fattiness, elaborate societies may result from the way in which such individual interact with each other. In this

chapter, we will explore future of theses society.

Why animals live in group?

Some animals such as tigers and other solitary hunters spend most of their time on their own though they must

come together in pairs of reproduction. Nevertheless, their mode of food finding is one which the presence of

others would interfere and their life is thus essentially a lone one, wandering widely over a range on which they

hunt. Sometimes such ‘home ranges’ overlap so that different individuals may use the same area and, and if

those of males habitually overlap with those of females, they may meet for mating without either leaving their

patch. Sometimes, however, animals live as individuals, pairs or larger group on’ exclusive territories’ from

which they repel others members of their species, at least theose of the same sex as themselves. The boundary

between such areas can be fiercely contested and often end up with quite sharp. A nice example is that the

borderline that can be seen in which the small fish mudskipper, the fish has the capacity to live out of water for

a period when the tide is out. The males are highly territorial and build mud walls between the areas they

occupy, so that by the time the tide returns, the mud is a mosaic of areas, each fenced off like a suburban

garden. When the tide is out, the walls preserve some water within the territory and this helps the animal to

respire, but the main function of the territory seems to be as a display area for the males leap in the air and this

attracts females to mate with them.

There are several possible reasons why animals might defend territories. The two most common ones are that

enable them to court and mate without interferences from others and that allows them to sequester them food
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supply which they can use systematically without others raiding on it. The males of many small songbirds set

up such territories, and this are usually large enough to provide enough food both for themselves and their

mates and offspring. Defending a territory as a source is food mainly a practicable propositions if the food

supply is rather constant and evenly spread. If this is not so, then the area required to feed the animal or family

if it is during the breeding season, maybe so great that it is impossible to patrol its boundaries and ensure that

others are kept out. The distribution of food and the amount of it that an individual can defend can have a

profound influence on social structures. For example, it has been argued that polygyny, a single male having

more than mate, may have evolved in territorial birds where some territories are very rich in food than others.

As a result, a female may breed more successfully by being the second one of a male on a rich territory than the

sole partner of a male whose territory is poor. If food is patchy in time and space, it may also have

implication on social structure. European badger lives in group of 2-12, on range which vary enormously in

size. Their food is largely earth worms, and the abundance of these varies from place to place and also with time

in the same place. The range size in particular area is such as to ensure that there is always a productive patch

somewhere within it, and the group is related how good a supply of worms is here overall within the range.

Therefore, probably it is the food availability that determines whether the animal maintain territories or live in

loose home ranges. Generally, the numbers of individuals that occupy the territory or range vary from one to a

sizable group. Why should it benefit selfish individuals to live in such group? Most of the reasons that have

been suggested relating to the way in which grouping helps are concerned with food finding and defense

against predators.
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Animal in groups gain a number of advantages related to defiance against predators. While a group animals are

bigger than a single one, and so can be spotted off, if the predators eat only one prey at a time it will take longer

to find a meal if the pry are clumped than they are evenly spread out. Furthermore, there are more pairs of eyes

in group, so the predators are less likely to creep up undetected. To some extent his is offset by the fact that

animals in groups are able to be less vigilant than solitary individuals. For instance, Long gees raise their heads

much more frequently while cropping the grass than they do when feeding in groups. Because here are pairs of

eyes they can afford to keep their heads down and feed or longer periods with the security that others will raise

a warning if danger threatens (See Figure 9.2 on page) .

A very good example is the adaptation of weaver birds in Africa in which some species are solitary and other

live in group.