Animal Behaviours

What is behaviour?

The observable actions (or inactions) of animals, including how they interact with their environments, conspecifics, and other species

T/F: Not all behaviours are intentional

True - they may also be driven by motivation

What is the Dead Man’s Test?

If a dead man can do it, it is not behaviour

Who emphasized that behaviour was subject to natural selection?

Darwin and Romanes

Romanes is introduced ___________________ focusing on animal intelligence and learning (made more rigorous by Lloyd Morgan)

Comparative psychology

What is comparative psychology?

the scientific study of similarities and differences in the behavior, cognition, and mental processes of different species, including humans and non-human animals

• cross-species comparisons, especially cognition

• focuses on understanding evolutionary, developmental, and functional aspects of behavior

• often applies principles from ethology and biology to explore learning, memory, and social interactions

Who are the founders of behaviourism?

Watson, Pavlov, and Skinner

• Note: Behaviourism = stimulus response learning

Who is associated with classical ethology (studied a wide variety of
behaviour in natural habitats)?

Lorenz, Tinbergen, and von Frisch

When did ethology and comparative psychology begin to merge?

After WWII

What are Darwin’s two fundamental tenets?

1. Behaviour is an adaptation

2. Behaviour resulted from selection pressures operating in the past

Ex. Death feigning in red flour beetles

What is the Farm-Fox Experiment? (Belyaev et al. - silver foxes)

• Foxes selected and sorted according to tameness

• Tame foxes were bred

• Selection proved to change morphological features seen in dog domestication

• Take-Home: can select for tameness and other traits

• Debated as previous experiment (unknown at time) may have impacted results

Describe George Romanes take on early comparative psychology

• advocate for natural selection

• compare animal intelligence across species

• continuity for mental traits, perception, memory, and problem solving

• Used anecdotes and prone to anthropomorphism and over-interpretation: rats forming supply lines, monkey showing a hunter blood on its hand

Describe Lloyd Morgan’s take on early comparative psychology

• continuation of comparative psychology - but emphasized experimentation

• work laid the foundations for ethology

• simple > complex

What is Morgan’s Canon?

“But surely simplicity of an explanation is no necessary criterion of its truth”

• simple > complex

• emphasizes that all else being equal, simpler explanations are preferred

What did Clever Hans (horse that “solves” math) teach us?

• need for strict controls - isolation from handlers

• benefits of blind and double blind experiments

• against researcher expectations that influence results of live subjects

Relevance to bias in artificial intelligence

What is behaviourism?

theory focusing on observable behaviors as learned responses to environmental stimuli, emphasizing that actions are shaped by conditioning (rewards/punishments)

• focus on stimulus response, observable behaviours

What are the key points of behaviourism?

• Lab animals and experiments (to avoid ‘uncontrollable’ chaos of natural environment)

• Emphasized need for systematic, replicable experiments

• Behaviours considered simple reflexes linked by conditioning

• Considered animals ‘black boxes’, ‘tabulae rasa’

• Not interested in ‘why’ an animal was selected to behave

• Interested in mechanisms of learning (conditioning, positive and negative reinforcement)

Describe the work of John Watson

• founder of behaviourism

• mental states are not scientifically measurable

• focuses on learning

• humans are shaped solely by the environment

What was John Watson’s view on mental states?

mental states are not scientifically measurable and behaviour should be the focus

What was the “Little Albert” experiment?

conditioned "Little Albert," to fear a white rat (a neutral stimulus) by
repeatedly pairing it with a loud noise (an unconditioned stimulus) indicating that fear, could be learned

What researcher is associated with the infamous “Little Albert” experiment?

John Watson

What behaviour does behaviourism emphasize?

Stimulus-response learning and conditioning

Describe the work of Ivan Pavlov

• Classical Conditioning

  • Pavlov’s dog (physiology of digestion) - taught dogs to drool at the sound of a bell

• Conditioned and unconditioned stimuli

What is the difference between conditioned and unconditioned stimuli?

An unconditioned stimulus (US) triggers a natural, automatic, unlearned response (unconditioned response), like food causing salivation

A conditioned stimulus (CS) is a previously neutral stimulus that, after being paired with a unconditioned stimulus (US), comes to trigger a learned response (conditioned response), like a bell (CS) causing salivation after being paired with food (US)

Key difference is the involvement of learning: US is innate, while CS requires association

What is Ethology?

the scientific study of animal behavior, focusing on instinct, evolution, and natural habitats, established as a modern discipline by Konrad Lorenz, Nikolaas Tinbergen, and Karl von Frisch.

• Focus on adaptive significance of behaviour in natural environments

• Evolutionary “why”

• how genetics influence behaviour

• the importance of the environment (interactions, umwelt, etc.)

What is umwelt? Provide an example.

sensory-perceptual world of an organism, different species perceive different stimuli as important

Ex. birds react to UV, dogs and olfactory senses to “see”, snakes detect IR, etc.

Who are the 3 people associated with Ethology?

• Konrad Lorenz

• Karl von Frisch

• Niko Tinbergen

What did Konrad Lorenz do?

• Studies instinct

• Imprinting

• Fixed Action Patterns (FAP)

What are Fixed Action Patterns (FAPs)?

stereotyped behaviour triggered by a simple stimulus, runs indefinitely

Ex. goose chasing rolling egg to bring back to nest

Who is Karl von Frisch?

• studied animal sensory processes

  • Ex. Honey bee figure 8 dance as form of communication

• Also looked at hearing in fish and alarm substances

Who is B.F. Skinner and what did he do?

• Experimental studies on animal training and learning theory

• Skinner’s Box — Pigeons and Operant Conditioning

Niko Tinbergen

• employed hypothesis testing and experiments in the field to ask questions about the adaptive significance, or mechanisms about natural animal behaviour

• Tinbegen’s 4 questions

What was Tinbergen’s Beewolf Experiment?

Wasps that use landmarks to find their nests, helped to identify mechanisms of animal behaviour

Sub-disciplines: Behavioural physiology

study of physiological bases of behaviour (e.g., neuroethology, ethnoendocrinology)

Sub-disciplines: Behavioural genetics

inheritance and molecular mechanisms of behaviour

Sub-disciplines: Cognitive Ethology

animal cognition and learning

Sub-disciplines: Behavioural Ecology

how ecology shapes behaviour and survival

Sub-disciplines: Sociobiology

evolutionary functions of social behaviour

What are Tinbergen’s 4 Questions?

• Mechanism (anatomy, physiology, regulation — “how it works”)

• Development/Ontogeny (role of genes and environment — “how the behaviour is developed”)

• Adaptive Value (how does the behavipur promote survival and reproduction)

• Phylogeny (how has it evolved and why it persisted)

What was Skinner’s Pigeon Box Experiment?

taught pigeons to peck red lights in order to obtain food

positive reinforcement, dopamine reinforces neural pathways

operant conditioning

Proximate VS Ultimate Causes

Proximate: how or why an organism behaves (mechanism + ontogeny)

Ultimate: How or why that behaviour evolved (phylogeny + adaptive value)

What is learning?

a relatively permanent change in behaviour resulting from experience

• Changes in neural representation (not temporary)

• Must store recall memories

Ex. birds refining song through practice, rat navigating a maze, etc.

What are the types of learning?

• associative

  • operant

  • classical

• non-associative

  • habituation

  • sensitization

• generalization

• social learning (teaching)

Associative learning

any learning process where organisms form an association between an behaviour and its outcome, or between two stimuli

Operant conditioning

behaviour modified by consequences (reinforcement or punishment — i.e. trial and error)

• voluntary behaviour (exploration, choices, actions)

• controls what happens + core to human/animal learning

• works as info from experience/stimuli are perceived via sensory receptors (reinforced by dopamine)

• Ex. Skinner’s pigeons, learning to navigate a maze, Macaques choosing opposite hand to obtain peanut reward, etc.

Classical Conditioning

learning when a neutral stimulus is paired with an unconditioned stimulus to elicit a reflexive response (no prior learning)

• involuntary, automatic, passive learning that involves reflexes and predicting what will happen

• Ex. Pavlov’s dog (drool at sound of a bell)

What is the Garcia Effect?

animals learn to associate taste with illness (even hours after and after only one experience)

• associated with classical conditioning

• natural selection pairs taste cues with illness

Habituation

decrease in response to a repeated, harmless stimuli (simplest forms of learning - not to waste energy on irrelevant stimuli)

Ex. reduced startle response (moth + chickadee, red-winged blackbird + whistling caterpillar)

Sensitization

increase in strength or likelihood of behavioural response following repeated exposure to strong, novel, or noxious stimulus

• increases an animal’s readiness to response to danger

Ex. sea slug shows progressive gill withdrawal after tail shocks

Occurs immediately, whereas classical conditioning requires repetition

Generalization

learned response to a particular stimulus by novel stimuli that are similar with response strength declining as stimulus similarity decreases

• learning extends beyond exact experiences to shape behaviour in novel situations

Ex. Pigeon identifying hover flies vs wasps

Social learning + teaching

learning from others

Teaching: Active participation of an experienced individual facilitating by a naive conspecific

• Teaching modifies a behaviour, incurs a cost and the pupil learns faster

• Ex. bird song, follower ants

Social learning has a role in cultural spread of innovations (ex. dolphins with tools, chimpanzees + sticks in ears)

Learned Helplessness and Superstition

Learned Responses VS Innate Behaviour

Learned Response: requires environmental input and changes over life history

Innate Behaviour: genetic basis, no prior experiences, changes over evolutionary history

Interaction — imprinting and predator recognition

Movement

any spatial displacement (foraging, commuting)

Search

movement to find a goal (ex. shelter, prey item)

Dispersal

one-way movement with permanent relocation (game, natal, adult)

Homing

movement with the goal of reaching a known area or resource

Migration

repeated, oriented, seasonal movements between distinct areas (usually return, usually long distance)

Zugunruhe

migratory restlessness, a behavioural manifestation of physiological change before migration

Orientation

movement in a given direction (not necessarily a location)

Navigation

guided movement from one location to another, typically using a compass or landmarks

Stopover

prolonged pause in migration for food and shelter (refueling)

Why do insects commonly orbit lamps?

Artificial light interferes with the dorsal light response (DLR) that orients themselves upwards

Movement ecology paradigm

argues that animal and plant movement can only be understood by jointly considering motivation, biomechanics, navigation and environment (rather than movement paths alone)

Levy Flight

random, memoryless walk where step-lengths have a probability distribution that is heavily tailed (extreme>exponential) — outdated

Area Restricted Search (ARS)

cue and expectation driven walks, movement patterns, for searching

Extensive (global) search — fast, directional, explore

Intensive (local) search — slow movement concentrated near a recent resource encounter

What are the major selective drivers of dispersal?

• avoidance of inbreeding (ex. male squirrels disperse while females stay in home range)

• reduction of kin competition

• escape from high densities, parasites, or depleted resources

• colonization of new/empty habitat

How and why does s e x based dispersal vary between mammals and birds?

Birds: Female biased dispersal because social monogamy so males defend territories & benefit from staying

Mammals: Male biased dispersal because there is frequent polygyny so males compete for mates and disperse

Dispersal Morphs

alternative phenotypes within a single species that are specialized for movement

• Developmental plasticity mediated by hormonal/epigenetic mechanisms (genetic background)

What is an example of dispersal for colonization of a habitat?

Spider ballooning where spiders produce silk to catch wind and disperse to a new habitat. This is influenced by atmospheric electric fields

What is hitch-hiking and phoresy?

one organism travels on another for dispersal (commensalism, mutualism, parasitism)

Ex. mites, flies

Describe homing of Rock Doves

Navigate using the sun as a compass with their internal clock to interpret the sun’s position. Use Earth’s magnetic field as a backup compass.

Pigeons use environmental odours for spatial distribution, alongside geomagnetic field, learned visual landmarks, and social cues from more experienced birds.

Examples of homing

• rock doves

• ant odometers — count steps

• spiny lobsters — geomagnetic map

• african dung beetle — starry sky orientation

Inclination

angle of field lines with Earth

Intensity

strength of field

Magnetic coordinates

combined intensity and inclination

What are the mechanisms of migration?

• hormones regulate migratory restlessness (zugunruhe)

• Melatonin, prolactin, glucocorticoids influence timing, fattening, activity

• Energy reserves act as a go/no-go gate

• Circadian and circannual clocks schedule migration

• Photoperiod is a key time cue

• Temperature and food act as modifiers

The tendency to migrate is shaped by…

• ecological opportunity (seasonality, resources, habitat)

• multiple constraints (energetic costs, parasites, mortality, navigation)

Partial migration

not all individuals migrate; balanced fitness for residency vs migration

Seasonal migration — stay and cope or leave and return

** Migration is phylogenetically labile (readily appears and disappears across lineages)**

What are the genetic and social components to migration?

Genetics provides the rules and predispositions (e.g. migratory restlessness, general compass direction, timing).
Social learning and individual experience fine-tune routes and decisions, especially in long-lived and socially migrating species

Example: Whooping Crane Migration (innate timing and direction but social learning improves route accuracy)

What are the benefits of migration?

• food + water availability

• shelter

• reproduction + mating benefits

• reduced competition

• reduced parasitism