PSYC 2721: Intro to Animal Behaviour

Territorial Behaviour

Deciding whether to be territorial or not; can have influence on whether a mate can be found/survival.

Behavioural /developmental plasticity

Ability to switch between behaviours in response to changing conditions, typically harsh environmental conditions

Organizational effects of hormones

Hormones that cause permanent, developmentally significant differences that cannot be changed

Activational effects of hormones

Hormones that cause temporary changes in an individuals morphology, physiology, or behaviour that last until hormone dissipates

Otoacoustic emissions

Evidence of organization effects of hormones in humans; amplitude of weak sounds produced by inner ear when triggered by ‘clicks’ → amplitude correlates with prenatal androgen levels (strong in females, weak in males, intermediate in opposite-sex fraternal twins = dizygotic)

Examples of organizational effects of hormones in humans:

• Otoacoustic emissions

• Finger length ratio

Finger length ratio

An example of the organizational effects of hormones in humans; finger-length ratios (2D:4D) → males = .98, female = 1.00, intermediate in females from opposite sex fraternal twins

Development of behaviour

How behaviour changes over the life on an organism (most behaviours change → varying needs, circumstances, etc. — although some remain unchanged)

Study of chicken hatching behaviour → whether hatching behaviour of an organism changes or not with age

Anne Bekoff & Julie Kauer

Hatching

Stereotyped thrusts of head and legs to break through a shell; only required one in a lifetime.

Chickens were re-shelled into ventilated, glass egg shells following actual hatch (up until 61 days); despite this, all showed stereotypical ‘hatching behaciour’ although, theoretically no practical, real-world use of behaviour → behaviour retained

What happened in Bekoff and Kauer’s chicken hatching experiment?

I. Changes in neural structure

II. Changes in hormonal state

III. Changes in non-neural morphology

IV. Effect of the environment

V. Experience

What are the 5 causes of behavioural change?

Change in neural structure

Change in neural substrate/mechanism that underlies current behaviour; e.g. as the tobacco hornworm transforms from a larvae → pupa, the underlying neural circuits degenerate and adult circuits evolve which shift behaviour from crawling/feeding → adult movement (flying) and reproduction

Changes in hormonal state

Changes in hormonal states = major contributes to behavioural development → can significantly change effects/behaviour by altering organization of NS → Note that not all developmental changes involve the nervous system

Changes in non-neural morphology

Changes in non-neural morphology (body) can lead to behavioral changes that coincide with a newly developed structure. E.g. Larvae = active and selective predators of zooplankton, Adults = passive indiscrimative filter feeders → coincides with appearance of ‘gill rakers in paddlefish.

Effects of the environment on behavioural change

E.g. leopard gecko; temperature dependent sex determination → females at cool/hotter temps (extremes) and varying ratios of males at intermediate temperatures: temperature affects aggressive and sexual behaviour: 30 degree males prefer 34 degree females/30 degree females & 32.5 degree males prefer 30 degree females/34 degree females

Effect of experience on behaviour

Experience may cause temporary or permanent changes in behaviour → effects = most prominent/influential during early life, effects can continue throughout adulthood

Neophobic feeders

Organisms that are cautious of trying new foods for consumption as a protective measure to prevent toxicity; e.g. Norway rats

To glean

Extract information from a source

Performed by Galef et al., the rat feeding study examined pars of rats housed together. One partner was removed from a cage, fed either cocoa or cinnamon flavored diet (‘demonstrator’), and then was returned to its cage. The other partner that remained in the cage (‘observer’) and did not witness feeding process of the ‘demonstrator’. Rats were allowed 15 mins of uninterrupted interaction and then, observers were tested for food preference (offered choice between cocoa and cinnamon flavored diets) → observers chose/preferred diet their demonstrator had been given! Evidence of social learning (via odor) and gleaning abilities of Norway rats

What was the Rat feeding experiment, what was its purpose, and who performed it?

Plexiglass → blocked odor, no learning

Wire screen → allows odor, leaning intact

How was odor determined to be the principle cue?

The return of the demonstrator rat from a ‘foraging’ trip indicates that food = safe; therefore, experience (either personal or observed) affects behaviour

Why would Norway rats show a preference to diets they had not tried themselves, but smelled from cage mates, when neophobic?

the early, variable, “practice” vocalizations of juvenile songbirds, similar to babbling in human infant; highly variable, rambling series of sounds without syllables of adult song

Subsong How do we know odour is the cue?

The developing song of a 3-7 month old male songbird; some recognizable elements

Plastic song

Crystallized song of a near-adult (7-9 month) male songbird

Full adult song

Social isolation results in males singing atypical songs

What is the evidence that song -learning is ‘learning’ and not instinct or genetically predetermined

10-50 days post hatch; Marler →

• if young male songbirds heard adult song b/w 10-50 days post hatch, song developed normally

• If there was playback of songs prior to 10 days or after 50, song was not learned.

What is the critical period for song-learning for a white-crowned sparrow? Who discovered this and what was the reasoning?

Critical period (Lorenz)

Periods of susceptibility to environmental stimuli: i. Brief, ii. Well-defined, iii. Species-characteristic, iv. Irreversible effects on behaviour

• Breif

• Well-defined

• Species-characteristic

• Irreversible effects on behaviour

What are the four components of a critical learning period ?

• Critical = cannot miss it

• Sensitive = easier now, possible later under specific circumstances

What is the difference between a critical period and a sensitive period of learning?

• fairly extended

• Not as well-defined; gradual onset/offset

• Differ induration between individuals

• Effects not irreversible → resistant to change

• Length of period may be dependent on stimulus characteristics (e.g. for white crowned sparrows sensitive period is longer than 50 days if live song tutors are used)

Characteristics of a Sensitive learning period

If young males are deafened before the onset of subsong (aka, bid babbling) → will never sing normally

What is the evidence that hearing an appropriate birdsong model is not the only essential component in birdsong acquisition in white-crowned sparrows?

i. Sensory phase

II. Sensorimotor phase

What are the two phases in the song development of white-crowned sparrows?

Characterized by retrieval and production of birdsong (rehearsal); matching sound to template in memory , through motor stabilization, reaches crystallized song. (Follows end of sensitive period)

What is the sensorimotor phase of white crowned sparrows song learning?

Occurs during sensitive period/phase; acquisition → song learned from ‘tutor’

Storage → song saved in memory

What is the sensory phase of the white-crowned sparrows song-learning process?

The gradual refinement and fixing of the motor patterns needed to produce an adult birdsong

Motor stabilization

True

True or False: the timing of sensory and sensorimotor phases of birdsong learning varies among species

E.g. Brown-headed cowbird: a type of brood parasite species; during the first spring after hatch, the brown-headed cowbird experiences the birdsong of the host species, not their own species. Therefore, sensory period is delayed by a FULL YEAR, when young bird will be surrounded by its own species.

What evidence supports the theory that the timing of sensory and sensory motor phases vary by species?

Product of physiological restraints:

• sensory receptors → tuned to frequencies of conspecifics

• effectors (morphology and musculature) → structurally specialized to produce own species song

• Central neural networks → maximally responsive to own-species’ song/promote production of own species’ song

What ate the reasons why there is own-species bias in song learning?

True: must be closely related to → structurally able to reproduce sound; saliency (importance) of stimulus matters.

**studied in lab

Can one species learn the birdsong of another? T/F?

Böhner → “Böhner’s methods” w/ zebra finch

Who performed the first ‘tutor choice’ experiment in birdsong acquisition?

→ placed two breeding pairs in separated compartments of a 3-chambered cage

→ at hatch, the youth male was placed into the middle compartment between parents and neighbors

→ after 100 days, recorded songs from hatchling male and compared to males of each pair (father & neighbor)

Most biased to learn father’s song

What did Böhner’s experiment with zebra finch tutor choice conclude?

Wanted to know whether the results of Böhner’s study were biased because the male of the same species was related to the young male (father). She instead chose a young male in between pairs of unrelated adult males.

• Male A: song similar to biological father

• Male B: song unlike biological fathers

Noted that the young male showed preference to learn from tutor who’s songs was like his biological father’s

Concluded: young “tuned” to father’s song

What spin did Nicola Clayton add to Böhner’s zebra finch tutor choice experiment?

Imprinting

A form of early learning that occurs during a brief, well defined period that produces irreversible effects on recognition/attachment behaviour; two types of→ filial & sexual

Immelman; fostered zebra finch eggs into clutches of Bengalese finches → foster parents accepted and raised young to independence → young were then isolated until sexual maturity:

• once sexually mature, males offered a choice of a female zebra finch and female bengalese finch _> majority chose bengalese finch

However, not unequivocal evidence of sexual imprinting → A control is necessary

Who conducted the study regarding sexual imprinting in finches?

Filial imprinting

Process by which young animals form a social attachment to their primary caregiver → most common in precocial species

→ Lorenz first to study systematically (e.g. greylag geese)

Williams

• noted that zebra finches breed colonially and synchronously in wild → triggered by rainfall

• Established aviary with 10 breeding pairs and 2 non-breeding males → discovered that sons in the aviary did not copy their father’s song father’s song exclusively; but did show preference

Who conducted an experiment to discern whether or not tutor choice experiments were applicable to what happens in nature?

• Young male birds may be biased towards learning from parents

• Song learning appears sensitive to aspects of social environment

• Results of an experience t are subject to be influenced by the design of an experience t (is it applicable to behaviour in nature/wild)?

What were the lessons learning in tutor choice studies of birdsongs?

I) transmission of complex patterns between generations

II) avoid risk of mutation in genetically controlled behaviour

III) adapt rapidly to physical and social environment (more dynamic)

IV) population regulation (affects mate choice and location)

Why would birds need to ‘learn’ songs? It is critical for mate attraction and territory defense → aka risky. Why learn when could inherit?

I) physiological constraints → most sensitive to/plastic during early life

II) more to gain, less to lose (sooner the better → e.g. anti predator behaviour)

III) Reliability associated with successful members of own species early in life (parents are guaranteed successful models for learning)

Why do birds learn birdsongs early in life?

Once species leave the nest, individuals are no longer exclusively conspecifics; given reliability of association with conspecifics (esp parents) early on, good way to form an image of one’s own species & a way to form the image of a suitable mate.

Why is the reliability of the presence of a successful member of your own species particularly important for species recognition?

Divided clutches of eggs laid by greylag geese into two groups:

I) hatched by mother: followed mother remaining in proximity

II) hatched by incubator: → first moving object was Lorenz → followed Lorenz as ‘mother figure’

Concluded that the geese imprinted to the first moving object in their environment: later noted that ‘following’ action was necessary

What was Lorenz’ experiment with greylag geese? What was its purpose and significance?

to keep young close to parents;

→ ensures young remain in suitable habitat

→ protects young from dangers of other conspecifics and predators

→ places young in situation to learn additional species-typical behaviour

What is the function of filial imprinting?

Sexual Imprinting

Early learning whereby an individual forms an image of what constitutes a suitable mate; typically after filial imprinting, sensitive periods may overlap

Cognition

Mechanisms by which animals acquire, process, store, and act on information in their environment; mechanisms include: perception, learning, memory and decision making

Learning

The acquisition of knowledge or skill through study, experience, or being taught, and is manifest as relatively permanent changes in behaviour.

Knowledge and skill to aid in survival or reproduction

What do animals learn?

B-GEN-PB

• Beneficial (reproduction/survival)

• Genetic basis (genetic heritability to learn)

• Environmental predictability ( ensures benefit from experience)

• Natural selection (nature selects those that have capacity to learn)

• Brain size (often correlate with greater learning ability)

Why do animals learn?

Ecological intelligence hypothesis

Variation in cognitive abilities among species evolve in response to aspects of diet and foraging, including fluctuating or complex spatiotemporal distributions of food and the need to extract food

Social Intelligence hypothesis

Variation in cognitive abilities among species evolve in response tp aspects of social life, including living in large or dynamic groups and the need to keep track of the identities of many individuals, their distinctive roles, and their past interactions.

Concerted brain evolution

Brain size and overall cognition evolves as a whole

Mosaic Brain evolution

The size of a specific region of the brain and its associated cognitive functions, can evolve independently from overall brain.

• individual learning

• Cultural transmission

What are the two ways in which animals learn?

Individual learning

the acquisition of knowledge or skills by a single animal through its own experience, trial and error, or interaction with the environment, without relying on others.

Sensitization

When an animal becomes more likely to respond to a stimulus after repeated exposure

Habituation

When an animal becomes less likely to respond to a stimulus after repeated exposure

Classical conditioning

When an animal forms an association between two stimuli

Operant learning

When an animal forms an association between its expression of a particular behaviour and subsequent reinforcement or punishment

Non-associative or higher order learning

Any cognitive performance that is not based on single-stimulus experience or associative leaning and which typically involves goal-oriented behaviours (e.g. emoe the macaque)

• Path integration

• Insight

• Innovation

• Reasoning

• Higher-order representations

What are the 5 types of higher-order/non associative learning ?

I) cognitive flexibility

II) self control

III) working memory

What three executive functions comprise higher-order learning and facilitate goal-oriented behaviour?

Cognitive flexibility

Ability to switch attention and adjust behaviour; tested using: a. Novel problem-solving tasks, b. Reversal learning tasks

Self-control

Ability to pause and override habitual responses that do not work

Working memory

Using temporary information that is no longer perceptually present; active recollection of past experience/knowledge

• Single-Stimulus experience

• Classical conditioning

• Operant conditioning

• Non-associative/Higher-order leaning

How many aspects of individual learning in animals are there?

• cognitive flexibility

• Self control

• Working memory

What are the three components of non-associative/higher-order learning?

Novel problem solving task

A task in which an animal or human is presented with a completely new situation or problem that they have not encountered before; quantifying time is super important to determine whether higher-order learning is occurring.

Reversal learning task

Train animal to associate food

with a colour via operant

conditioning, then switch the

food association to a different

colour to test if animal can

reverse its preference

Detour-reaching task or cylinder task; shortest path to reward first try

What test is used to evaluate self control in animals?

Object permanence task; e.g. hide reward under cloth → will animal remove cloth to access reward? Can alter difficulty via introducing delay before covering reward and giving animal access to covered reward

How do we test working memory in animals?

Cultural Transmission

When an individual’s behaviour(s) is/are influenced by the behaviour of another through ‘social learning’ or ‘teaching’; individuals can be from the same generation of from different generations. E.g. Emoe the Japanese macaque and her sweet potatoes

A. Provides a mechanism for spreading information among individuals = much more efficient

B. Allows learned behaviours to persist in a population even after original ‘learner’ or ‘innovator’ dies; mechanism so behaviour persists in populations for long periods of time.

Also, alternative mechanism of how behaviour can be transmitted and how evolution acts on this.

What are the benefits of cultural transmission in comparison to individual learning?

Traditions

Socially learned and taught behaviours that spread through a group and are stable over time (i.e. across generations)

Social learning

Occurs when one individual known as an ‘observer’ learns a behaviour from another individual, known as a demonstrator.

Social facilitation

When an animal is attracted to a particular location because a conspecific is present there. They do not learn anything from conspecific, but, once in location, they may be more likely to experience individual learning

Social facilitation

What is the exception to social learning?

Copying and imitating

What are the two variations of social learning?

Imitation

acquisition of a topographically novel response through the observation of a demonstrator making that response

Copying

When an observer retreats what it has seen a demonstrator do (behaviour does not have to be novel)

An individual is teaching if it modifies its

behaviour in the presence of a naïve student, without

obtaining an immediate benefit for itself. The teacher’s

behaviour encourages or punishes the student’s behaviour,

or provides the student with experience, or sets an

example for the student. As a result, the student acquires

knowledge or learns a skill earlier in life or more efficiently

than it might otherwise do

What constitutes ‘teaching’?

‘If it modifies its behaviour in the presence of a naïve student’

What distinguishes teaching from social facilitation, imitation and copying?

‘Without obtaining an immediate benefit for itself’

What distinguishes teaching from instances of social aggression?

“As a result, the student acquires knowledge or learns a skill earlier in life or more efficiently than it might have otherwise done”

What distinguishes teaching from commonly observed social interactions (e.g. dominance interactions)?

Meerkats siblings teaching pups how to forage for scorpions (dangerous); as young pups age, siblings, aka ‘helpers’ gradually adjust behaviour (dead → injured → live → hunting expeditions)

What is an example of teaching in animal communities?

• Vertical

• Oblique

• Horizontal

What are the three ways in which cultural transmission can occur?

Vertical cultural transmission

Information is transmitted from parent → offspring (most common)

Oblique cultural transmission

Information is transmitted from one generation to the next, but does not involve parent and offspring; group members, subordinates, etc.

Horizontal Cultural Transmission

Information is transmitted among peers in the same generation (siblings)

Personality

A phenomenon where individual behavioural differences are consistent over time and/or across situations

• Shyness-boldness

• Exploration-avoidance

• Activity

• Aggressiveness

• Sociability

(SEAAS)

What are the “Big 5” personality traits?