Animal Behaviour and

Chronobiology
Krishnendu Sinha
Assistant Professor
Department of Zoology
Jhargram Raj College

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Unit 1: Introduction to Animal Behaviour
1. Origin and history of Ethology
2. Brief profiles of Karl Von Frish, Ivan Pavlov, Konrad Lorenz, Niko Tinbergen
3. Proximate and ultimate causes of behaviour
4. Methods and recording of a behaviour

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 What animal does is called animal behaviour

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Unit 1: Introduction to Animal Behaviour
1. Origin and history of Ethology
2. Brief profiles of Karl Von Frish, Ivan Pavlov, Konrad Lorenz, Niko Tinbergen
3. Proximate and ultimate causes of behaviour
4. Methods and recording of a behaviour

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 The Nobel Laureates of Ethology

Ivan Pavlov
(1849-1936)

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•Ivan Petrovich Pavlov was a Russian physiologist known primarily for his work in classical conditioning. Pavlov won the Nobel Prize for
Physiology or Medicine in 1904.

•Karl Ritter[a] von Frisch, was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along
with Nikolaas Tinbergen and Konrad Lorenz. His work centered on investigations of the sensory perceptions of the honey bee and he
was one of the first to translate the meaning of the waggle dance.

•Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or
Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarded as one of the founders of modern ethology, the study of
animal behaviour. He developed an approach that began with an earlier generation, including his teacher Oskar Heinroth. Lorenz
studied instinctive behavior in animals, especially in greylag geese and jackdaws.

•Nikolaas "Niko" Tinbergen was a Dutch biologist and ornithologist who shared the 1973 Nobel Prize in Physiology or
Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning organization and elicitation of individual and social
behavior patterns in animals. He is regarded as one of the founders of modern ethology.

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Unit 1: Introduction to Animal Behaviour
1. Origin and history of Ethology
2. Brief profiles of Karl Von Frish, Ivan Pavlov, Konrad Lorenz, Niko Tinbergen
3. Proximate and ultimate causes of behaviour
4. Methods and recording of a behaviour

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Niko Tinbergen’s 4 Types of questions
Niko Tinbergen outlined in a classic paper entitled “On the Aims and Methods of Ethology” (N. Tinbergen, 1963). These questions centre
on:

• Mechanism—What stimuli elicit behavior? What sort of neurobiological and hormonal changes occur in response to, or in
anticipation of, such stimuli?

• Development—How does behavior change as an animal matures? How does behavior change with the ontogeny, or
development, of an organism? How does developmental variation affect behavior later in life?

• Survival value—How does behavior affect survival and reproduction?

• Evolutionary history—How does behavior vary as a function of the evolutionary history, or phylogeny, of the animal being
studied? When did a behavior first appear in the evolutionary history of the species under study?

Tinbergen’s four questions can be captured in two different kinds

of analyses—proximate analysis and ultimate analysis
Proximate analysis is focuses on immediate causes, whereas
Ultimate analysis is defined in terms of the evolutionary forces that have shaped a trait over time.

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THREE FOUNDATIONS OF ANIMAL BEHAVIOUR

1.The force of natural selection

2.The ability of animals to learn (individual learning), and
3.The power of transmitting learned information to others
(cultural transmission)

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 Hawaiian island, Kauai

Natural Selection
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 Natural Selection
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 Individual Learning
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Individual Learning
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 Cultural Transmission
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 • Kin selection
•Direct fitness
•Indirect fitness
•Inclusive fitness

Conceptual Approach
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 • OFT
•Game theory

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Theoretical Approach
 • Observational
•Experimental

Empirical Approach
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Unit 1: Introduction to Animal Behaviour
1. Origin and history of Ethology
2. Brief profiles of Karl Von Frish, Ivan Pavlov, Konrad Lorenz, Niko Tinbergen
3. Proximate and ultimate causes of behaviour
4. Methods and recording of a behaviour

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Basic consideration
Approaches in behavioural studies
1. observational
2. experimental
3. ethological

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Will discuss at practical…

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Unit 2: Patterns of Behaviour
1. Stereotyped Behaviours (Orientation, Reflexes)
2. Individual Behavioural patterns
3. Instinct vs. Learnt Behaviour
4. Associative learning, classical and operant conditioning, Habituation,
Imprinting

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Stereotyped response
 Unlearned behavioral reaction of an organism to some environmental stimulus
It is an adaptive mechanism and may be expressed in a variety of ways
All living organisms exhibit one or more types of stereotyped response
Stereotyped responses are the result of a continuing process of evolutionary modification and
refinement
Those actions that most successfully aid the animal or plant in its basic drives (e.g., reproduction,
search for nourishment, escape from predators) are the ones most likely to be retained in succeeding
generations
As environmental conditions change, inherently determined responses also become modified in
order to ensure continuation of the species
Types: Orientation (Taxis and kinesis) & Reflexes
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Unit 2: Patterns of Behaviour
1. Stereotyped Behaviours (Orientation, Reflexes)
2. Individual Behavioural patterns
3. Instinct vs. Learnt Behaviour
4. Associative learning, classical and operant conditioning, Habituation,
Imprinting

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Types of behaviours
1. Taxis and kinesis
2. innate and learned
3. egocentric and altruistic

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 Maintaenance
Body surface maintenance Comfort movement Anting
• Body care • Active
• Rubbing
 Bathing • Passive
• Rutting
 Water bathing
 Mud bathing • Scratching
 Sand bathing
 Fire bathing • Shaking
 Powdering • Stretching
 Cleaning • Niblling

 Moving tail • Yawning

 Grooming
• Panting
• Sub bathing/ basking
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Unit 2: Patterns of Behaviour
1. Stereotyped Behaviours (Orientation, Reflexes)
2. Individual Behavioural patterns
3. Instinct vs. Learnt Behaviour
4. Associative learning, classical and operant conditioning, Habituation,
Imprinting

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Motivation and drive
 What compels an animal to do what it does is motivation
 All the internal and external factors which cause an organism to behave in a goal seeking way is motivation
 Motivation leads to build-up of drives for the animal to perform appropriate behaviour (appetitive and
consummatory)
 Motivation helps to optimize stimulation to the homeostasis
 There is no behaviour without motivation except strict Pavlovian reflex
 Instincts are important to understand motivation
 Instincts are unlearned behaviours and responses which help them to survive and reproduce
 Animals have needs which motivate it to perform a behaviour

 Drives are internal state of arousal (e.g. thirst or hunger etc.) which animals tried to reduce and re-
establish the homeostasis

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 Hierarchy of drives

1. First level drive

(reproductive drive)

2. Second level drive

(territoriality, mating, nest
building, parental care)

3. Third level drive

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Sign stimulus
Specific external stimuli that triggered stereotyped
responses from conspecifics, usually called fixed action
pattern

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Unit 2: Patterns of Behaviour
1. Stereotyped Behaviours (Orientation, Reflexes)
2. Individual Behavioural patterns
3. Instinct vs. Learnt Behaviour
4. Associative learning, classical and operant conditioning, Habituation,
Imprinting

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 HOW ANIMALS LEARN?
This discussion of how animals learn follows an outline developed by Cecilia Heyes,
both because of its conciseness and its attempt to tie how animals learn to why they
learn (Heyes, 1994)

Heyes notes that there are three commonly recognized types of experience that can
lead to learning—namely,
1. Single stimulus
2. Stimulus-stimulus (Pavlovian (classical) conditioning), and
3. Response-reinforce (Operant conditioning)— each of which facilitates certain
forms of learning.
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Single-stimulus learning
Sensitization and habituation are two simple single-stimulus forms of learning

Habituation and sensitization:

Numerous times each day, a blue stick is placed in a
rat’s cage. If the rat takes less and less notice of the
stick, habituation has occurred. If the rat pays more
attention to the blue stick over time, sensitization has
taken place.

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 Habituation as an problem

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Pavlovian (classical) conditioning

Paired stimuli
Five seconds after a blue stick (stimulus 1)
is placed in a rat’s cage, the odor of a cat
(stimulus 2) is sprayed in as well. The
question then becomes: Will the rat pair
the blue stick with danger (cat odor)?

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In nature, US=UR
Upon pairing, US+NS=UR/R
When, NS=R
Then, NS=CS & R=CR

Conditioned response
If the rat pairs the blue stick (CS) and the cat odor (US), it will
hide under the chips when the blue stick alone is presented.
Such hiding represents a conditioned response (CR)

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Ivan Pavlov and classical conditioning. Pavlov watches a classical
conditioning experiment as it is being conducted in his laboratory. In the
experiment, a device to measure salivation has been attached to the dog’s
cheek, the unconditioned stimulus is a dish containing meat powder, and the
conditioned stimulus is a light. (Photo credit: Sovfoto/Eastfoto)
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Second-order conditioning The rat learns to respond to a
second CS—the yellow light—with the conditioned response.

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Pavlovian conditioning affects not only behavior
per se but also what is referred to as
learnability (that is, the ability to learn under
certain conditions).

We will explore three types of learnability:

overshadowing, blocking, and latent inhibition.
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Overshadowing
The process of overshadowing is shown in two groups of rats.
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 Blocking
Learning can be slowed down
depending on prior association or
lack of association between
stimuli

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Finally
 Let’s consider a fourth group of rats that is initially exposed to a blue stick,
but no cat odor, for a long period of time
We then try to pair the blue stick with cat odor at some subsequent point
in time
If we find that the rats in group 4 have more difficulty learning than the
rats in group 1 (where standard Pavlovian pairing has occurred), then we
would say that latent inhibition is responsible for this lack of learning in
group 4
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Instrumental (operant) conditioning
 Instrumental conditioning, also known as operant or goal-directed learning, occurs when
the response that is made by an animal is reinforced (increased) by the presentation of a
reward or the termination of an aversive stimulus, or when the response is suppressed
(decreased) by the presentation of aversive stimulus or the termination of a reward.

 One of the most fundamental differences between Pavlovian and instrumental learning is
that, in instrumental learning, the animal must undertake some action or response in
order for the conditioning process to produce learning.

 Thorndike’s law of effect: This law states that if a response in the presence of a stimulus is
followed by a positive event, the association between the stimulus and the response will
be strengthened. Conversely, if the response is followed by an aversive event, the
association will be weakened.

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Rats in a Skinner box
To test various theories of animal learning, rats are often placed in “Skinner
boxes,” where they have to take an action (here, pressing a button) to get a
reward of food or water. (Photo credit: Walter Dawn/Science Photo
Library/Photo Researchers, Inc.)
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Extinction In learning, the decrease of response
rate or magnitude of response with lack of
reinforcement in a learning situation

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 Imprinting is a form of learning in which a very young animal fixes its attention on the first object with
which it has visual, auditory, or tactile experience and thereafter follows that object

In nature the object is almost invariably a parent; in experiments, other animals and inanimate objects
have been used

Imprinting has been intensively studied only in birds, especially chickens, ducks, and geese, but a
comparable form of learning apparently occurs in the young of many mammals and some fishes and
insects

In mallard ducklings and domestic chicks, imprinting can be accomplished in a few hours, but receptivity
to imprinting stimuli vanishes at the age of about 30 hours.

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Unit 2: Social and Sexual Behavior
1. Social Behaviour: Concept of Society, Altruism (Insects’ society with
Honey bee as example, Foraging in honey bee and advantages of the
waggle dance)
2. Communication and the senses
3. Sexual Behaviour: Asymmetry of sex, Sexual dimorphism, Mate choice,
Intra-sexual selection (male rivalry), Inter-sexual selection (female
choice), Sexual conflict in parental care

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 Special Topic

Theory of Decision Making

Game theory is a branch of applied mathematics
that provides tools for analysing situations in which
parties, called players, make decisions that are
interdependent. This interdependence causes each
player to consider the other player’s possible
decisions, or strategies, in formulating his own
strategy.
Game theory was originally developed by…

John von Neumann Oskar Morgenstern

Hungarian-born American mathematician German-born American economist
Types of games…

1. One-person, two-person, or n-person game (based on number of players)

2. Games of perfect information vs. game of imperfect information (based

on available information about the opponent’s strategies, e.g. Chess vs.
Poker)

3. Constant-sum games (game of pure competition e.g. poker) vs. variable-

sum game (based on the distribution of resource)

4. Variable-sum games can be further distinguished as being either

cooperative or non-cooperative.
 Prisoner’s Dilemma

Albert W. Tucker
(Canadian mathematician) John F. Nash
American Mathematician
T = “Temptation to cheat” payoff,
R = “Reward for mutual cooperation” payoff,
P = “Punishment for mutual defection” payoff, and
S = “Suckers” payoff.
For the matrix to qualify as a prisoner’s dilemma game, it must be true that T > R > P > S

PD Matrix Suspect 2

Cooperate Cheat

Suspect 1 Cooperate Suspect 1: Suspect 1:

R = 5 year in jail S = 15 years in jail
Suspect 2: Suspect 2:
R = 5 year in jail T = 0 years in jail

Cheat Suspect 1: Suspect 1:

T = 0 years in jail P = 10 years in jail
Suspect 2: Suspect 2:
S = 15 years in jail P = 10 years in jail
What should be the Evolutionary Stable Strategy
(or the Nash Equilibrium)?

• If cooperation adopted by most then

cheating penetrate

• If cheating adopted by most then system

collapses and cooperation penetrate

So…Tit for Tat

Among us…

Tit for Tat

Generous
Cheating Tit for Tat
 The Hawk-Dove game

John Maynard Smith George Price

British theoretical and American geneticist
mathematical evolutionary
biologist
c>v leads to hawk-dove
v>c leads to PD

(we have them in this room!!!)

What should be the Evolutionary Stable Strategy
(or the Nash Equilibrium)?

A mixture of both
(can be calculated mathematically)
which is dynamic in nature
and
the third bourgeois behaviour
Unit 3: Social and Sexual Behaviour
1. Social Behaviour: Concept of Society
2. Communication and the senses
3. Altruism: Insects’ society with Honey bee as example, Foraging in honey
bee and advantages of the waggle dance
4. Sexual Behaviour: Asymmetry of sex, Sexual dimorphism, Mate choice,
Intra-sexual selection (male rivalry), Inter-sexual selection (female
choice), Sexual conflict in parental care

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Unit 3: Social and Sexual Behaviour
1. Social Behaviour: Concept of Society
2. Communication and the senses
3. Altruism: Insects’ society with Honey bee as example, Foraging in honey
bee and advantages of the waggle dance
4. Sexual Behaviour: Asymmetry of sex, Sexual dimorphism, Mate choice,
Intra-sexual selection (male rivalry), Inter-sexual selection (female
choice), Sexual conflict in parental care

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 Communication is the transfer of information from a signaler to a receiver
 It modulate the behavior of the receiver

The vervet monkey story of Amboseli National Park, Southern Kenya

Cough-Eagle Barking-Leopard Chutter-Snakes

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Types and Evolution of Behaviour
one might expect from a salesman attempting to convince
Signaler Receiver
a prospective buyer that his product is the top of the line
+ve -ve
+ve True Communication Manipulation signaling often involves some costs—for example, energy
Conspirational whispers Exaggerated signals costs or drawing attention from predators—natural
selection should favor minimizing these costs during
-ve Exploitation Ignorance conspirational whispers, thus reducing the
Krebs and Dawkins, 1978
conspicuousness

 When what is good for the signaler is not good for the recipient, natural selection will favor signalers that send
signals in whatever way best increases the fitness of the signaler, even if that means manipulating recipients
 But natural selection also favor recipients with the ability to unscramble what is honest and what isn’t, so it can act
in ways that maximize its own fitness. Krebs and Dawkins refer to recipients as “mind readers” and describe an
“arms race” between signaler and recipient
 in which the signaler is selected to better manipulate the receiver, which then is selected to better filter out only
that information that benefi ts it, and so on KS_JRCZOO
Types and Evolution of Behaviour

Signaler Receiver
+ve -ve
+ve True Communication Manipulation
Conspirational whispers Exaggerated signals

-ve Exploitation Ignorance

Krebs and Dawkins, 1978

 When what is good for the signaler is not good for the recipient, natural selection will favor signalers that send
signals in whatever way best increases the fitness of the signaler, even if that means manipulating recipients
 But natural selection also favor recipients with the ability to unscramble what is honest and what isn’t, so it can act
in ways that maximize its own fitness. Krebs and Dawkins refer to recipients as “mind readers” and describe an
“arms race” between signaler and recipient
 In ‘arms race’ which the signaler is selected to better manipulate the receiver, which then is selected to better filter
out only that information that benefits it, and so on KS_JRCZOO
Types and Evolution of Behaviour

Signaler Receiver
+ve -ve
+ve True Communication Manipulation
Conspirational whispers Exaggerated signals

-ve Exploitation Ignorance

Krebs and Dawkins, 1978

 When what is good for the signaler is not good for the recipient, natural selection will favor signalers that send
signals in whatever way best increases the fitness of the signaler, even if that means manipulating recipients
 But natural selection also favor recipients with the ability to unscramble what is honest and what isn’t, so it can act
in ways that maximize its own fitness. Krebs and Dawkins refer to recipients as “mind readers” and describe an
“arms race” between signaler and recipient
 In ‘arms race’ which the signaler is selected to better manipulate the receiver, which then is selected to better filter
out only that information that benefits it, and so on KS_JRCZOO
Types and Evolution of Behaviour

Signaler Receiver
+ve -ve
+ve True Communication Manipulation
Conspirational whispers Exaggerated signals

-ve Exploitation Ignorance

Krebs and Dawkins, 1978

 When what is good for the signaler is not good for the recipient, natural selection will favor signalers that send
signals in whatever way best increases the fitness of the signaler, even if that means manipulating recipients
 But natural selection also favor recipients with the ability to unscramble what is honest and what isn’t, so it can act
in ways that maximize its own fitness. Krebs and Dawkins refer to recipients as “mind readers” and describe an
“arms race” between signaler and recipient
 In ‘arms race’ which the signaler is selected to better manipulate the receiver, which then is selected to better filter
out only that information that benefits it, and so on KS_JRCZOO
Even within the manipulator/mind-reader view of communication, honesty might evolve:

 Signals being sent are impossible Under Zahavi’s handicap principle

 Or at least, very difficult to fake
But…  Honesty is also possible when traits are not impossible but
This appears to be the case in toads: merely very costly to fake
Deep croaks and physiological advantage
 Honest communication may be an outcome even when
deception is possible in principle, as long as deception is
costly

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Even within the manipulator/mind-reader view of communication, honesty might evolve:

 Signals being sent are impossible Under Zahavi’s handicap principle

 Or at least, very difficult to fake
But…  Honesty is also possible when traits are not impossible but
This appears to be the case in toads: merely very costly to fake
Deep croaks and physiological advantage
 Honest communication may be an outcome even when
deception is possible in principle, as long as deception is
costly

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This thought leads Amotz Zahavi to Handicap principle

Competitive altruism: In Arabian babbler the helping-at-the-nest behavior often occurs

among unrelated individuals- thus cannot be explained by kin selection-The altruistic act
is costly to the donor, but may improve its attractiveness to potential mates

Obesity, a sign of ability to procure or afford plenty of food, comes at the expense of health, agility, and in more
advanced cases, even strength; in cultures that experience food scarcity, this exhibits "fitness" to the opposite sex and
is also often associated with wealth

Peacock to peahen 'I have survived in spite of this huge tail;

hence I am fitter and more attractive than others‘
Response to male cowbirds song by female:
Each point represents the copulation-solicitation displays (CSDs) of one female

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 Deceptive Alarm Calls!!?
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 Honey Bee Dance Languages

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 Modification of Behavior Due to Anthropogenic Disturbance

Cock-of-the-Rock (South America/Peru)

Altering forest canopy in or near lek (traditional place where males assemble during the mating season
and engage in competitive displays that attract females) leads to…

(1) stop males from displaying at all because of increased exposure to predators,
(2) induce males to display so often that they become energetically drained, or
(3) lead to females no longer being able to assess male quality accurately
References

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