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ANIMAL COGNITION (I/II) – LEARNING
WHAT IS LEARNING?
Knowledge sources = innate + learned by experience + cognitive (not independent of other 2)
Learning = inferred change in organism’s mental state…results from experience…influences, relatively
permanently, the organism’s potential for subsequent adaptive behaviour
-

Need to separate from the following 3 phenomena that appear to overlap in some cases

Learning vs performance
-

Learning vs
...
g
...
maturation
Learning depends on special experience
Unlearned sexual responses appear at maturity, regardless of experience

Learning vs motivation
-

Learning vs
...
= reversible

TYPES OF LEARNING
Non-associative
Responding to repetition of a single stimulus
-

Habituation – decreased innate responding to consequence-less stimulus
Sensitisation - ^ responding

Associative
Association of 2+ events
-

-

Classical conditioning
o Associations between external events
o Pavlov => dog and bell
Operant conditioning
o Associations between own behaviour and outcomes
o Thorndike/Skinner => Skinner box

CONDITIONS FOR ASSOCIATIVE LEARNING
Contiguity
= shorter interval between CS & US
Appropriateness
= easier recognition of relevance of CSs for USs
-

E
...
smell to taste easy…smell to electric shock less easy

Contingency
= correlation between CS + US = most important
-

i
...
how often does CS precede US

-

CS can…
o Predict US presence (excitatory)
o Predict US absence (inhibitory)
o Be non-predictive

E
...
rats: tones + electric shocks
-

P = 0
...
1, 0
...
3, 0
...
g
...
OPERANT
-

Associations between external events vs
...
g
...
g
...
g
...
g
...
“different”
E
...
Fagot et al (2001): baboons vs
...
g
...
g
...
g
...


WHAT IS TOOL USE?
-

E
...
termite fishing in chimpanzees?
E
...
bait-fishing in herons using bread?
E
...
F palm cockatoos attentively watch Ms drumming display?
E
...
octopus carrying coconut to retreat into for protection?

Definitions
-

Pioneering (Goodall) + broad + narrow and/or anthropocentric + …
BETTER…Shumaker, Walkup, Beck (2011)…
o Externally employs unattached/manoeuvrable attached object
o To alter form/position/condition of another object/organism/user itself
o Tool is held/directly manipulated prior to/during use
o ‘responsible’ for proper and effective orientation of tool

WHEN DOES IT EVOLVE?
-

Facilitates access to scarce and important resources
Enables colonisation of new habitats
Improves ‘niche’ requirements e
...
diet
By-product of complex cognition

4 main, non-mutually
exclusive hypotheses

DOES IT INDICATE INTELLIGENCE?
-

Tool use in humans much more common than in non-humans => tool use requires intelligence
BUT…
o Can be acquired w/ associative learning and/or SL
o Can be relatively genetically fixed
o Is linked to morphology (i
...
need appropriate appendages to use tools)

NEW CALEDONIAN CROWS
-

Key behaviours discovered are:
o Hook making: distal, proximal, unbending e
...
Betty – tool modification
o Sequential tool use e
...
use short stick to obtain longer stick etc
...
g
...
)?

Tool use  physical cognition?
Study 1: assessed performance on tasks testing for causal understanding
-

Could always improve via trial and error
No evidence they were forming a mental representation of the problem
=> NO CAUSAL UNDERSTANDING

Study 2: comparison of 2 tool-using spp
...

-

No diff
...

NC crows > hooded crows
=> INCONCLUSIVE

MORPHOLOGICAL RESTRICTIONS
-

-

Kea (hooked bill) vs
...
g
...
“ANIMAL CONSTRUCTION”
-

Behaviours like nest-building excluded from most tool use discussions

-

Assumed that animals FAP’d (followed Fixed Action Patterns)
…BUT weaving suggests otherwise…
FAP = rigid, heritable motor programs that involve
o Variable
no learning/innovation/cognitive complexity
o Improves w/ experience
o Individual signatures

Why?

Nest building and learning in zebra finches
Study 1: is the choice of materials genetically predetermined?
-

-

2 options…
o Flexible string
o Stiffer string
After building w/ flexible, preferred stiffer
o Choose material based on structural properties
BUT if raise chicks successfully w/ flexible, continue w/ it
o Type + amount of experience influences choice

Study 2: is there a role for adult experience?
-

-

Method…
o Ms tested for colour preference
o Allowed to build nest w/ preferred/unpreferred colour
o Eggs removed/left
o Provided w/ both colours and allowed to build again
Result…
o Forced to use unpreferred material…
▪ Successful => same colour again
▪ Unsuccessful => neutral preference

An alternative approach
-

Tool use isn’t rare because of cognitive complexity
Tool use is rare because tools are rarely useful/used
o E
...
some “tool-using” chimps only use tools for < 1% of feeding
In order to study physical cognition…
o Tools should be examined in wider context of construction behaviours
▪ More widespread + more important

ANIMAL COGNITION (IV) – PHYSICAL INTELLIGENCE II: INNOVATION &
PROBLEM SOLVING
-

Traits displayed regularly w/I sp
...
g
...
g
...
g
...
g
...


WHAT CAN PROBLEM-SOLVING TELL US ABOUT COGNITION?
Goffin’s cockatoos vs
...
(more likely in larger flocks)
o All birds in larger flocks benefited
Conclusion…
o “Empirical evidence for “pool of competence” hypothesis” <= subsequently challenged

COMPARATIVE METHODS
Do innovators have bigger brains?
-

Foraging innovations reported in 800 bird spp
...
appears to be positively correlated to…
o Tool use freq
...
American bird spp
...
g
...
g
...

▪ Rabbit < wildebeest in terms of size…
▪ Rabbit > wildebeest in terms of benefit…relatively more important if rabbit
effectively saves lives in times of starvation
• Conflict between memory of prey size vs
...
brain…what should animals use…what do animals use?
Working for an outcome ^ its value
E
...
starlings
-

Starlings prefer stimulus encountered after HARD trials…despite EASY yielding same food reward

Putative valuation mechanisms
-

Low state of reserves => ^ benefit from reward
Several psychological processes could drive preferences:
o ABSOLUTE MAGNITUDE (M)
o VALUE (V)
o MARGINAL VALUE (m)
o DISTORTED PERCEPTION
▪ Memory of physical properties of reward
distorted => costly outcomes valued more

E
...
starlings
-

Is preference between delayed rewards state-dependent?
o State at time of choice => irrelevant
o State during training => strong effect
▪ …therefore, state-dependent VALUE
drives choice
▪ VALUE + DISTORTED PERCEPTION
accommodate results

-

Does memory of time delays depend on outcome?
o # pellets = reward magnitude
o Peck harder for more pellets
o Perceive timing accurately
▪ VALUE accommodates results

Summary so far
-

Starlings prefer stimuli associated w/ more work/hunger
Preference sensitive to state + size of outcome
Distorted representation of outcomes unlikely
Surviving hypothesis = preference reflects utility gains during learning
Can cause paradoxical choices: they “know” what they’re choosing, but can choose a worse option
State-dependent valuation = adaptive or costly constraint?
If remote spp
...

In Mexican cave fish…
-

Y-maze w/ coloured arms
o Preference for HUNGRY option
o State during testing = irrelevant

Functional reflection
-

Taxonomic ubiquity suggests convergence
o Convergence suggests adaptive advantage
What is the advantage???
o Easier to have common currency to compare
rewards in diff dimensions (food vs water etc)?

RELATIVE VS
...
B (10s) vs
...
A (5s) vs
...
B (10s) vs
...
? Wouldn’t it take too long to evaluate every possibility?
o E
...
chess masters asked to think out loud
▪ Only considered 3-4 options and usually went w/ 1st
▪ Unconscious board examination and discarding of unworthwhile avenues

Are starlings like Darwin?
What is a choice?
-

2+ options followed by decision
o E
...
caterpillar (preferred) vs
...
e
...
SCM predicts choice trial preferences using no choice trial latencies; ToW ignores no-choice trials
- Choices accurately predicted by latencies in no-choice trials => support for SCM
2
...
e
...
trinary tests are equal
o => supports rationality via IIA

ANIMAL COGNITION (VII) – MECAHNISMS AND EVOLUTION OF SONG
LEARNING IN BIRDS
-

Cultural trait in many spp
...
g
...
g
...
g
...
Sensory phase (memorisation)
2
...
Song crystallisation phase (own and tutor’s song compared)

-

Evidence consolidated

Deafened/raised in acoustic isolation => abnormal song (2002)
o E
...
zebra finch

Innate song

Adult song

Suboscines…

Tutored plastic song

-

No dialects
Simple songs
Song @ crystallisation
Evidence…
o Eastern phoebe
Isolation
▪ Deafened/isolated => normal song
o Alder flycatcher
▪ Tutored by heterospecifics => normal song

Tracheophones…
-

Similar evidence ^^^

Genetic control
-

Song learning in anterior forebrain (IMAN, Area X)
o Evidence = suboscines lack well-defined IMANs
Gene products change in abundance during song development
FOXP2…
o Expressed in Area X
o When inhibited in zebra finches => unable to copy tutors

Differences in learning
-

-

Inter-sexual…
o M oscines => larger song nuclei w/ more neurons + brain connections, despite uniform
brain size
o Crystallisation + production controlled by testosterone
▪ Ms injected w/ test => produce song out of season
▪ Castrated Ms => fail to produce normal adult song
W/I-sp…
o Photoperiod diffs => variable breeding period duration => variable timing + duration of 3
stages

-

Between spp…
o Age-limited OR open-ended

Innate learning preferences
-

Conspecific > heterospecific
Live heterospecific > recorded conspecific
Live heterospecific father > live conspecific w/ no visual contact
Variant > invariant song
Sensory period longer w/o tutor

Visual cues important

Innate song properties
-

Aspects develop normally even if isolated (genetically determined):
o Duration/note shape/frequency range/# of songs
Other aspects do not (culturally determined):
o Syllables
▪ E
...
swamp sparrows
• Of c
...
Complexity
2
...
^ development costs

All about maximising transmission
and minimising degradation

Why learn to sing?
1
...
g
...
g
...
INTRASEXUAL SELECTION (rivals)
- Song mediates same-sex resource comp
- Learning => more efficient communication w/ rivals
o Young Ms preferentially learn songs of older, established neighbours
o Territorial Ms respond more aggressively to matched songs
o Young Ms w/ matched songs => ^er RHP
o Song and repertoire matching = signal of aggressive intent
o Song sharing => ^ RHP
o Song sharing only in stable communities
3
...
g
...
g
...
g
...

o No diff in O2 consumption when singing w/ large vs
...
s develop complexity => reliable indicator of ind
...

singing-related neural pathways
o Experimentally stressed juveniles => shorter song bouts +
smaller adult repertoires

WHY STUDY BIRDSONG?
-

-

Model system for understanding learning
o Birds vocally active from early age
o Song dev
...
early on
o Insights into interaction of nature + nurture
o Insights into adult neurogenesis + behaviour
Integrates knowledge from behaviour + genetics + physiology + neurobiology

Like humans

ANIMAL COGNITION (VIII) – SOCIAL LEARNING
-

-

Animals can learn from others in various ways:
o From products of their activity
o Observing what object/part of environment they attend to
o Copying actions and/or outcomes of activities
SL = cognitively “special” OR associative learning in a social context…open debate
Does imitation require specialised cognitive skills?
Teaching is rare

DEFINIITION & E
...
S
-

= learning from others’ behaviour
o This simple definition shirks some big questions:
▪ Is learning from others diff in mechanism/function to learning from own experience
▪ SL  special intelligence?
▪ When is SL preferable?
▪ How and when does SL result in culture?

Classic e
...

-

E
...
blue tits milk bottle opening in early 1900’s Britain
o Distribution = patchy occurrence w/ local hotspots => indicated learning
o Numerous potential explanations…
▪ Imitation/emulation/stimulus enhancement/local enhancement/social facilitation

Pre-existent behaviour e
...

-

E
...
blackbird mobbing response
o Juveniles learn to target mobbing through influence of
others = OBSERVATIONAL CONDITIONING
o Pre-programmed = the response + certain targets
o SL = new targets

Novel behaviour e
...

-

E
...
bee string pulling
o More social observers learnt than individual innovators => SL
important
o Non-social observers failed => conspecific observation important
o Analysis of observers’ behaviour => learnt by attending to:
▪ Local enhancement (demonstrator’s activity location)
▪ Stimulus enhancement (string position)

…examples above relatively complex, but not as cognitively demanding as true imitation…
IMITATION DEBATE
-

-

-

Correspondence problem = projecting someone else’s body/actions onto one’s own
body…COGNITIVELY DEMANDING
Who is capable?
o Humans => expert imitators from young age
o Non-humans => some anecdotal evidence but few clear experimental demonstrations
What is the value?
o Allows rapid behaviour transmission
o Allows ^ fidelity behaviour transmission => “ratcheting”
Any negatives?

o May allow non-adaptive behaviours to persist
Testing: the “two-action” task
-

Demonstrators-to-be trained to solve task by 1 of 2 actions
Naïve individuals observe proficient demonstrators before being allowed to try the task
=> do previously naïve individuals solve task in manner they observed????
o Yes => attention paid to actions involved…rather than objects/location/outcome

E
...
quails
-

Observed demonstrator getting food via pedal operation: pecking at it OR stepping on it
Results allude to true imitation…
o …BUT maybe quails have innate “mirror peck” + “mirror step” neurons that trigger when
they see the action performed i
...
need not be
“cognitive perspective-taking”

To imitate or to emulate?
E
...
chimps + humans box-opening test
-

-

Openings on top and front
o Only front opening connected to reward location
Transparent or opaque
o Reward location only evident when transparent
Subjects shown both relevant + irrelevant actions for reward
attainment
SURPRISING RESULTS…
o Transparent => chimps omit irrelevant action,
therefore switching to emulation…humans don’t!!!
▪ => OVER-IMITATION…
• Same experiment as above…relevant
+ irrelevant actions shown to 3-year-olds + 5-year-olds + adults
o => tendency to over-imitate ^ w/ age!!!
Problems w/ comparing imitative abilities in humans vs
...
e
...
early on) on non-human primates => human demonstrator
▪ Non-conspecific = inconsistent

IDENTIFYING SL IN THE WILD
How can we distinguish between social transmission and series of repeated individual inventions?
-

Increasingly sophisticated network-based analyses being developed

Analysis of order of acquisition

E
...
spread of “moss sponging” (for drinking water) in wild chimps
-

Camera trap caught alpha M using moss instead of leaf in presence of alpha F, who watched intently
Spread locally…most parsimonious explanation = SL

SL STRATEGIES
What to copy?
-

Majority behaviour?
Behaviour most recently observed?

Whom to copy?

Arrow = observation
Dashed arrow = picked up
a dropped moss sponge

-

Those who achieve a reward?
Those who have proven ability?
Those who are dominant?
Those who are kin?

When to copy?
-

When uncertain in one’s own info?
When environment is rapidly changing?

TEACHING
If A is teaching B…
1
...
A incurs cost/derives no immediate benefit from instruction
3
...
g
...
g
...
g
...
g
...


How to investigate it?
-

Assess how individuals manipulate others’ behaviour

DECEPTION
Types
1
...
Tactical deception
- Acts from w/I normal repertoire…deployed in a way that benefits agent by making another ind
...
Intentional deception

-

Intent to deceive by altering target’s mental state

Nesting plovers – intentional?
-

-

-

Distracts enemy
from nest by
running &
acting wounded

3 hypotheses:
o 0th => behaviour automatically triggered by predator
o 1st => plover intends to lead predator away
o 2nd => plover intends predator to believe she’s easy prey
Results of experiment…
o 87% of human approaches => moved so as to lead away from nest
o ‘dangerous’ intruders => more arousal than ‘non-threatening’
o Intensified/re-approached intruders if they stopped following
Conclusion…
o Observed flexibility => intentionality?
▪ Assumes non-intentioanl systems to be rigid + inflexible
▪ BUT associative learning + inherited predispositions can lead to subtle behaviour
• Innate anti-predator response, made more flexible by learning
Lecture mantra: don’t
assume intentionality where
simpler explanations possible

Tactical/intentional deception in the wild
-

Whiten & Byrne (1988) => collated apparently
Only included acts:
deceptive acts performed by wild primates
- Which seemed deliberate
o 117 cases of tactical
- Which seemed to deceive target
o 18 potential cases of intentional
- Performed by spp
...
explicit understanding

GAZE TRACKING
-

Understanding that others see suggests implicit understanding
Humans use others’ gaze to work out what they’re attending to
o Can non-humans do the same?
In chimps…
o Able to follow gaze direction (also ravens + goats)
o Understand that barriers block gaze

‘SEEING’
-

On the contrary, DOGS do preferentially
beg from ‘attentive’ humans… feasible
that this behaviour is learnt however

Is there an understanding that others see???

In chimps
Study 1: choose to beg for food from experimenters who can/can’t see
them
-

Results…
o Mostly random

Study 2: subordinate chooses between hidden or visible food
-

Results…
o Hidden food approached first in 73% of trials
Conclusion…
o Contradicts results from study 1 and perhaps suggest
chimps do know what others see

Does ‘seeing’ imply knowing?
Do chimps understand what dominant sees or just react to dominant as stimulus?
-

4 conditions:

Competitive feeding paradigm

-

-

o Informed – dominant witnessed baiting
o Uninformed – dominant did not witness baiting
o Misinformed – food moved post-baiting
o Switched – dominant changed for another chimp post-baiting
Results…
o Uninformed/misinformed => subordinate less inhibited to approach food
o Switched => subordinate retrieved more food
So… do chimps have ToM?
o Not necessarily
▪ Behaviour reading
▪ More fearful if dominant has been there for longer

…NEED TESTS WHERE INDIVIDUALS HAVE TO PROJECT THEIR OWN EXPERIENCES…
PROJECTING ONE’S EXPERIENCE ONTO OTHERS
Chimps (‘goggles experiment’ paradigm)
-

Gaze-following experiment => no effect
Competitive feeding experiment => weak effect

Jays (‘pilfering experience’ paradigm)
-

In wild…
o Cache food
o Remember ‘what’ + ‘where’ + ‘when’
o Sometimes pilfer others’ caches
o Re-cache food hidden in presence of conspecifics
▪ => understand others’ mental states?

Study: Hand-raised jays allowed to cache waxworms…in private OR in presence of conspecific…later, given
opportunity to re-cache in private
-

-

-

2 groups:
o Never pilfered
o Have pilfered
Results…
o Re-cached when observed caching
o Re-cached in new sites
o Re-caching only performed by those w/ prior pilfering experience
So…do jays project pilfering experience onto others?
o MAYBE
▪ Needs replicating
▪ Behaviour-reading still possible
o Difficult to distinguish between cognitivist + sophisticated behaviourist accounts of
apparently intentional behaviour

ANIMAL COGNITION (X) – CONSCIOUSNESS
-

Consciousness remains ‘hard problem’
Main method for inferring animal consciousness is by analogy w/ ourselves
Search for Neural Correlates of Consciousness (NCC) in humans has proved difficult
Perception + emotional responses in humans can occur w/o consciousness

WHY IS IT SUCH A PROBLEM?
Definition
We all think we know what it is, so why is defining it so hard?
-

= a wide range of states in which there is an immediate awareness of thought/image/sensation

Different forms
-

PHENOMENAL (aka ‘qualia’/’raw feels’)
Memory
o Subjective experience of seeing/hearing/feeling pain etc
...
Why doesn’t all
this information processing go on ‘in the dark’”

‘easy problem’ vs
...
g
...


-

How do physical processes in
brain => subjective experiences?

Easy => can be studied by known scientific methods
Hard => no idea how this happens…no idea what to look for in other spp
...
materialism
Is consciousness explained by physical laws?
-

DUALISM (Rene Descartes c
...
by-product
Is consciousness an adaptation or by-product?
-

-

ADAPTATION
o Consciousness must do something advantageous
▪ By doing, must intrude into workings of brain
• By intruding, must be by magic or physical laws…bloody dualism
BY-PRODUCT
o Avoids dualism problems whilst still acknowledging hardness of ‘hard problem’
o Occurs as a result of body mechanism working…no power to modify working

The Behaviourism debate
Can consciousness be studied scientifically?
-

= only observable events can be studied scientifically
Hindered consciousness study until recently
o Psychology + Animal Behaviour avoided the topic for majority of 1900s
o Watson (1924) => “States of consciousness…are not objectively verifiable and for that
reason can never become data for science”

Revolt against behaviourism…
-

-

Griffin (1976) => The Questions of Animal Awareness
Griffin (1991) => Animal Minds
o “The taboo against considering subjective mental experiences of non-human animals has
become a serious impediment to scientific investigation”
Has the revolt gone too far?
o No harm necessarily in sticking to behaviourism, whilst appreciating the issue’s complexity

The language debate
Is language necessary for consciousness?
-

-

Reports of experiences = 1 of main tests of human consciousness
Consciousness only occurs when capable of Higher Order Thoughts
o Such thoughts require language
o Most animals don’t have language, thus not sentient?
Live debate…

WHICH ANIMALS HAVE IT?
-

To infer human consciousness…
o Reporting
o ‘ARGUMENT FROM ANALOGY’
▪ E
...
pain
• I know I experience pain…
• When experiencing pain, I show signs that can be observed by others…
• Other people react similarly…
• Therefore, infer that others also consciously experience pains

Argument from analogy in animals
-

-

-

E
...
rats’ thermal comfort
o Learn to press a lever to alter temperature of stimulus applied to skin if too hot/cold
o Humans behave similarly
o Humans report very cold/hot stimuli as unpleasant
o => BY ANALOGY, rats consciously experience thermal comfort
Which animals feel pain?
o NOCIOCEPTION (detection of + response to potentially tissue-damaging stimulus)
…or…PAIN (unpleasant conscious feeling associated w/ nocioception)
o Elwood (2009) on pain…
▪ Not reflexive
▪ Motivational trade-off
▪ Remembered + avoided
o E
...
hermit crabs
Crabs leave shells upon
▪ Fewer crabs evacuated preferred shells
mild electric shock
• Not reflexive + motivational trade-off
▪ Avoided old shell + entered new shell w/ less exploration
• Remembered + avoided
Use brain imaging to study:
o Vegetative humans
o Humans/animals coming round from anaesthesia
o Specific kinds of cortical activity that indicate consciousness
▪ …BUT consciousness signs remain difficult to interpret

Problems w/ applying argument from analogy to other spp
...
‘Killjoy’ explanations
- = explaining complex human-like behaviour via simple processes rather than consciousness
- E
...
jays recaching
o More likely to recache when they’ve been observed by another bird
o BUT ‘virtual birds’ also recache when observed…
▪ => recaching related to ‘stress’
- E
...
monkey’s memory
o Appear to report on how reliable their own memories are
o BUT self-monitoring can be achieved w/o consciousness
▪ E
...
trivial to program a computer to act differently depending on strength of
memory trace
2
...

o Mammals + birds = conscious
o Apes + humans = conscious
▪ Parts of human brain responsible for consciousness are phylogenetically young
▪ Right neocortex + prefrontal cortex = prerequisites for emotional experiences…
absent from other animals
3
...
performed via conscious or unconscious routes
- Blindsight
o = seeing w/o being conscious of seeing
o E
...
emotions can be influenced by stimuli that we’re unaware of…
▪ Brief presentation of happy/sad/angry face affect interpretation of neural
ideographs, despite subjects saying they haven’t seen any faces
o E
...
patient w/ damage to visual cortex => cortically blind in left visual field
▪ More accurate than chance when guessing what objects and where they’

ANIMAL COGNITION (XI) – INDIVIDUAL DIFFERENCES IN BEHAVIOUR
-

-

Introduce current work on causes/consequences of I
...
-dependence important for understanding maintenance
o Continuous variation often described as ‘personalities’
Introduce recent work on fitness/broader social consequences of I
...
in cognition/SL in wild

EXPLAINING PHENOTYPIC VARIATION
-

Extensive for some characters, v limited for others
Given a sample from a pop…
o How much of variation about mean is non-adaptive vs
...
s produce all Ps?
o Are I
...
-dependence often needed to maintain var
...
theory because 1
occurs and is rapid

strategy should prevail unless exactly
equal fitness OR fitness adv
...
I
...
Alternative behaviours => equivalent fitness @ ESS…BUT sel
...
-dependent
Foraging behavioural game: producer-scrounger division
-

Predicts how frequently food parasitism should occur
-ve freq
...
g
...
s demonstrated required plasticity

Conditional strategies
1
...
Blue-gill sunfish M DEVELOPMENT tactics
i)
Parental
ii)
Sneaker, before growing into…
iii)
Satellite (F mimic? – what’s benefit of this?)
3
...
e -ve freq/-dependence
CONTINUOUS TRAITS
-

Vary continuously w/I pop
I
...
variation in personality
-

Personality/behavioural syndrome = suites of behaviours that correlate across contexts
Recent explosion of interest in animal personalities

E
...
3-spined sticklebacks
-

-

5 behaviours measured
In 2 environments:
o Normal
▪ Strong correlations between behaviours across
contexts
▪ Variation applies to multiple behaviours
▪ Often-documented syndrome between aggressiveness
+ activity + exploration found
o Predator-free
▪ Lower freq
...
g
...
g
...

in pop density => var
...
in personalities
Which personality does best depends on:
o Environmental conditions
o Gender

E
...
Western Bluebirds/Mountain Bluebirds invasion front
-

Western Bluebird range expanding…displacing Mountain Bluebird
o Biased dispersal of highly aggressive M Westerns to the invasion front
=> displacement
o Mountain exclusion => aggression of Westerns decreased
rapidly…local selection on highly heritable aggressive behaviour

E
...
spiders
-

-

-

3 colony mixtures assessed for % agggressive I
...
s
In 2 diff state sites:
o High-resource
o Low-resource
Results…
o All pops returned to something akin to natural state => selection dependent on environment

o

Sites consistent in selecting for particular group-size-dependent personality combos

Naturally occurring mixture

Initial experimental mixture

Surviving experimental mixtures

Colony size

Why are there personalities at all?
Why isn’t behaviour completely flexible? What are the costs of flexibility?
1
...
behaviour dependent on inherently stable states (e
...
sex) => time consistency of state differences
reflected in time-consistent behaviour
2
...
g
...
Non-state-related models
- Social interactions => consistent and correlated behaviour can evolve in absence of state differences

INDIVIDUAL VARIATION IN COGNITIVE PERFORMANCE
-

Cognitive differences between spp
...
variation in captivity
-

-

-

2 tasks:
o Lever-pulling => food
o String-pulling => food
Simple tasks…require diff motor actions…ecologically relevant
Results…
o I
...


Study 2: did I
...
role in driving evo
...
CONCEALMENT (this lecture)
2
...
ARMOUR
2
...
TOXINS
4
...
Countershading
i
...
g
...
Shape scrambling
i
...
g
...
g
...
Disruptive colouration
i
...
Does shape disruption affect predator’s ability to recognise outline?
o YES
o Cardboard moths w/ mealworm on them pinned to trees & exposed to
bird predation
o 5 designs
▪ Those w/ disruptive colourations experienced ^ survival
o
o
o
o

iii
...
g
...
E
...
vizcacha – uses bands to disguise eyes
d
...
Colour matching w/ background
CRYPSIS IN DEPTH
-

E
...
Biston betularia typical vs melanic morphs
o Each less susceptible when in matched background

Matching the background
-

-

General colour resemblance…
Background selection…
o Non-uniformity of backgrounds + prey usually ranging across several types = difficulties
Developmental switches + seasonal phases…
o E
...
willow ptarmigan plumage
▪ Winter => white
▪ Summer => dappled, largely brown
Active colour change…
o E
...
chameleons
▪ Relative expansion/contraction of skin melanophores

Special resemblance
-

Confusing recognition…
o E
...
leaf mimic moths

-

Freq
...
g
...
g
...
American moths
▪ ^ crypsis values on available backgrounds than unavailable
▪ Jays take longer to attack moths w/ ^ crypsis values

RECEIVER PSYCHOLOGY OF CONCEALMENT
-

Predator response to novel cryptic prey = sigmoidal
o As abundance ^ become adjusted => ^ capture rate

-

Achieved by:
o Search image improvement
▪ TRANSIENT improvement in detection ability for characters of single/small prey set
o Search rate adjustment
▪ TRANSIENT reduction in rate ^ detection of more cryptic prey

ANIMAL COGNITION (XIII) – RECEIVER PSYCHOLOGY: EVOLUTION OF
DEFENSIVE ADVERTISEMENT
ADVERTISEMENT MECHANISMS
-

-

Flash colouration
o Overlaps with mechanisms of concealment
o Camo til predator is close…once spotted, no point trying to stay concealed so flash helps escape
o DISPLAY TO CONCEAL…fool predator attention mechanisms
Startle
o DISPLAY TO CONFUSE…confound predator expectations
▪ Frightening
▪ Eye spots…may exploit subliminal reactions
Least effective - arbitrary
Intermediate - circles
Most effective – asymmetric sub-circles

-

-

Deflection
o DIVERT ATTENTION from body/towards less crucial body part
▪ Inhibits detection of cryptic cues
• E
...
insect leg flags
o No tests actually performed…appears to be only feasible hypothesis
WARNING
o Conspicuous colours => unpalatability
o APOSEMATISM = warning signal is associated w/ unprofitability of prey
o Gregariousness ^ learning
o Multicomponent displays

WARNING DISPLAYS
Aposematism: the role of conspicuousness
-

-

Conspicuous => ^ chance of detection
Might assume distinctive but as inconspicuous as possible would be ideal…avoids naïve predation
Does it work?
o YES – Initial rate of consumption will be greater BUT will asymptote @ lower level
Hypotheses based on receiver psychology:
o More memorable
o More discriminable
o Reduces recognition errors
A more strategic hypothesis:
o Conspicuousness = handicap
▪ Only genuinely unpalatable prey can afford it

Gregariousness enhances learning
-

-

Predators…
o Learn quicker
o Forget slower
Enhancement is visual (not chemical)
o Green line matches red => visual aggregation is important

Discrimination
ratio

Multicomponent signals
-

Warning odours may potentiate colour aversion learning
o E
...
domestic chicks
▪ Hidden colour aversion triggered by pyrazine odours of insect warning displays
▪ Pyrazine interacts with red and yellow to induce strong aversions to these aposematic
colours that are not shown in the absence of the odour

MIMICRY
= predators confuse the mimic for the model due to evolved visual resemblance
2 types
-

-

Batesian – advergent
o Only works when relatively rare
o E
...
Papillo dardanus
▪ Uses sex-limited polymorphic Batesian mimicry as
response to problem of freq
...
g
...
America
▪ Geographical polymorphism…unusual for Mullerian mimics
▪ 1 gene (Optix) controls the extreme colour variation
in independent convergent events

Evolution
-

Zone of protection important
Predators generalise responses to similar stimuli due to stimuli variability in nature
o Generalisation provides opportunity for mimicry…mimics only need to get so close
Initial large mutation followed by smaller refinements

Classic e
...
= Monarch + Viceroy
What type of mimic is Viceroy?
-

-

-

May be Mullerian
Batesian vs
...
e
...
Arctic tern trans-global migration
2
...
Desert ant direct return post-foraging
- Keeps running vector of tortuous movements
4
...
Shearwater long-distance migration
- ~10,000km in c
...
11,000 birds
o Adults => compensation
o Juveniles => maintained innate vector course
Genetically based vector orientation e
...
Blackcaps
o Migratory restlessness (directional tendency pre-migration) in
orientation cage
o Migratory restlessness in Emlen funnel
o Conclusions…
▪ 1st-time migrators showed directional tendencies consistent w/ their intended popspecific migratory direction
• E
...
SW for German pop normally migrating to Africa
▪ Also genetically encoded duration => clock-and-compass navigation (Kramer, 1953)
• Hybrids => intermediate tendencies

WHAT DO ANIMALS USE AS A COMPASS?
Compass info – constant geographic direction regardless of viewer’s position
1
...
e
...
g
...
POLARIZED LIGHT
- E
...
Cataglyphis
o In ‘colour and polarisation vision’ lecture
o Retinal polarisation analysers
3
...
g
...
MAGNETIC
- Earth-s geomagnetic field = ubiquitous cue
- E
...
European robin
o Computerised registration cage inside large Helmholz coil
o Sensitive to angle of dip NOT polarity => ineffective at equator
- E
...
garden warbler
o = transequatorial migrant

-

o Reverses compass after a few days in a horizontal field…avoids issue of responding to angle
of dip instead of polarity
Magnetoreceptor…
o Electroreceptors
▪ E
...
sharks and rays (elasmobranchs)
o Biogenic magnetite crystals?
▪ Found in many animals (e
...
salmonids) sensitive to magnetic field
o Light-dependent magnetoreception
▪ E
...
red-spotted newts
• When trained under red spectrum light, fail to orient under full spectrum
▪ E
...
birds? – robins
• Trigeminal nerve (innervates magnetite in beak) sectioning => ineffectual wrt
magnetic orientation…NOT MAGNETITE-RELATED
• Lesioning ‘night-vision’ region => magnetic disorientation (no effect on sun
or star)…APPEARS LIGHT-DEPENDENT
▪ Cryptochromes = likely receptor molecules
o Overwhelming evidence suggests magnetoreception is light-dependent in most organisms,
therefore not magnetite-based but probably a radical pair system

MULTIPLE COMPASSES
-

Compasses can be used to calibrate each other
o E
...
savannah sparrows
▪ Use polarised light to calibrate magnetic compass

ANIMAL COGNITION (XV) – NAVIGATION: TRUE NAVIGATION
TRUE NAVIGATION
“after displacement to a location where they have never been, they can determine their position relative to a goal without relying on
familiar surroundings, cues that emanate from the destination, or information collected during the outward journey”
-

-

Positional sense + compass sense = true navigation capability
o Many animals, both vertebrate and invertebrate, possess diverse compasses
o Few animals (all vertebrates) known to be able to determine position relative to a goal after
displacement to unfamiliar areas when prevented from monitoring the outward journey
Invertebrates that have been carefully studied return to target areas using
path integration, landmark recognition, compass orientation etc
...
g
...
g
...
5 of expected
Sun compass orientation predicts 90 degree anti-clockwise shift
o During route learning, deviated anti-clockwise
o Once faithfully established, robust to clock-shifts
▪ Although still some minimal residual effect

Clock-shift = internal
diurnal clock is re-entrained
to an experimentally altered
light:dark cycle… leads to
predictable deviations in
the time-compensated sun
compass, therefore can be
used to determine what
role the compass plays in
orientation

Anticlockwise deviation (degrees)

Filled circles = high-familiarity birds
Empty circles = low-familiarity birds

Distance from release (m)
Evidence suggests the pigeon’s familiar area map is a pilotage map:
-

Learn individually idiosyncratic routes home (top right)
Routes often follow distinctive and linear landscape features
Attracted back to their routes if released off-route (bottom left)

BUT…before routes are formed birds seem to use their sun compass to encode the relationship of places to
home (mosaic map)…this effect persists to an extent even as birds become experienced w/ landscape
UNDERSTANDING SPATIAL RELATIONSHIPS: COGNITIVE MAPS? GEOMETRIC
MODULES?
Can novel routes be computed from foundation of spatial knowledge?
E
...
Honey bees
-

-

Gould (1986), yes
o Novel shortcuts after displacement to a familiar site when on route to another site
Dyer (1991), no
o Behaviour of all but 1 of the bees implied vector map
▪ The 1 was released from same point as initial experiment implying that something
was fundamentally wrong with experimental design…able to see landmarks
associated with target site
Learn efficient “traplines” (stable multi-location routes) w/o need for understanding complex spatial
relationships
o Use simple search and snapshot memories (retinotopic snapshots)…experience gradually
reduces travel distances

E
...
Pigeons
-

Very local shortcuts only
o Explained by sighting of familiar landmarks on route ahead
o Gradually ^ efficiency over each rep

E
...
Rodents
-

Many expts have suggested they can take novel shortcuts
o Can often be explained by simpler local rules
Do animals learn geometry of familiar space or use simple rules?

E
...
Ants
-

-

Can constantly update homeward vectors…sophisticated path
integration
Simple visual centring response to landmarks
o In cluttered environments, tend to run along midlines of alleys
formed by adjacent shrubs
o No response to heterogeneous stationary/moving black/white
gratings
o Only respond to wall height…vertical angles subtended by the walls must be identical
Learn simple relationships with landmarks
o Stay certain distance away



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