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L1 Domestication
Where members of one species assume control of the feeding, breeding, and general care of another species
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Why study behaviour?
- 1. Fitness value of exploiting other species
- a) Obtaining food
- b) Efficiently exploiting labour
- c) Insight via research
- 2. Coping with the natural environment
- a) avoiding predation
- b) pest control
- 3. Conservation
- 4. Curiosity about the natural world
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Anecdotal
Descriptions of the behaviour of a single individual on a single occasion
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Anthropomorphic
Ascribing human attributes, particularly emotions, to non-human animals (without scientific evidence
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Teleological
Assuming conscious intent of part of animal
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L2 Ethology
- Study of instinct
- Training: Zoology
- Location: Europe
- Emphasis: evolutionary approach to the study of instinct
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Instinct
Behaviour that is not amenable to experiential modification
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Ethology reasoning
- - Natural selection can only act on traits having a genetic basis
- - Therefore focus should be on inherited instinctual traits
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Ethology - Subjects and Methods
- - Varied
- - Field studies:
- Comparative: behaviour contrasted among closely-related species to explore evolution
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Sexual cannibalism
Apparent sacrifice consistent with the Central Principle of behavioural biology
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Central Principle of Behavioural Biology
Animals are expected to behave in a manner that maximizes the propagation of like copies of their copies of their genes.
A simple consequence of natural selection resulting from differential reproductive success
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Benefits of nuptial gifts
- Female:
- Nutritional benefit with no cost or risk associated with prey capture
- Allows assessment of male quality
- Male:
- Increase duration of copulation increasing sperm transfer
- Prevents female assessment
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Comparative Psychology
- Training: Psychology
- Location: NA
- Emphasis: The study of learning
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Learning
The developmental modification of behaviour through experience
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Comparative Psychology Reasoning
- - Learning plays the paramount role in determining behaviour
- - Therefore, everything worth knowing about behaviour could be determined by studying learning
- - The principles of learning are universal
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Comparative Psychology Subjects and Methods
- - Norway Rats
- - Laboratory experiments:
- Permits control of extraneous variables that would otherwise confound results
- In particular, allows control of prior experience of subjects
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Ethology: Whitman and Heinroth did what?
- Examined behaviour of birds
- Concluded that displays are relatively constant within species, and similar in closely-realted species.
- Suggested that displays could be used to classify species just as morphological characters had been used in the past
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Ethology: Wallace Craig
- Constituents of Instinct - 2 types: Appetitive and consummatory behaviour
- Pursued study of avian displays, examining behavioural
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Ethology - Craig - Appetitve
- At the beginning of a behavioural sequence, behavioural acts are highly variable
- Variable behaviours brings the animal into "the appropriate stimulus situation to release a simpler, more stereotyped response.
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Ethology - Craig - Consummatory
- Once released, the stereotyped behaviour that completes the behavioural sequence.
- Repeated without variation (stereotyped)
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Craig Consummatory -> FAP (Lorenz)
7 characteristics of FAP (Lorenz)
- Stereotyped
- Self-exhausting
- Specie specific
- Released
- Triggered
- Complex
- Independent of experience
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Stereotyped
Motor patterns that occur in rigid, predictable, highly-structured sequences
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Complex
Are patterns of behaviour, not simple reflexes
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Species characteristic
Performed by all normal members of a species, or at least all members of a given sex, at the appropriate age that have attained a certain physiological condition.
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Released
Appear in response to certain simple, yet highly effective stimuli
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Self-Exhausting
Occurrence of a FAP reduces ease of re-release
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Triggered
Once released, continue to completion regardless of changes in environment
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Independent of experience
Not learned; complete upon first release
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L3 FAP
All produced in response to certain extrinsic stimuli (releasers and sign stimuli)
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Releasers
Stimuli emanating from members of one's own species
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Sign stimuli
If from other elements of the environment
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Umwelt
- An animal's perceptual world.
- The stimuli tat elicit behaviour in one species are not the same as those the elicit response in another species
- Each kind of animal lives in a species-characteristic perceptual world that differs from other kinds of animals
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Supernormal Stimulus
- Exaggerated characteristics of an effective stimulus (relevant cues)
- More effective at releasing a response than the natural stimulus
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Exceptions to classic FAP rules
- Reduced response threshold
- Vacuum activity
- Displacement behaviour
- Graded response
- Intention movements
- Redirection
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Reduced response threshold
Over time, FAPs become progressively easier to release
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Vacuum activity (Craig with doves)
eventually, FAPs are expressed in the absence of the sign stimulus or releaser
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Displacement behaviour
FAPs may be performed that are irrelevant to the stimulus present
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Graded response
FAPS may not be triggered, but rather vary in intensity of expression
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Conceptual models
promote understanding of complex phenomena by presenting essential components and relationships in simpler terms
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Lorenz's Psychohydraulic model- Ethology
- Reduced response threshold: build up of ASE placing greater pressure on IRM (innate releasing mechanism)
- Vacuum activity: eventual buildup of ASE forces action of IRM
- Displacement behaviour: overflow of ASE into centers controlling other FAPS
- Graded response: Rate of ASE release proportionate to stimulus intensity
- Suggest that mechanisms are hard-wird into nervous system that act like a flush toilet.
- Normally inhibit bahaviour
- Respond to simuli to release behaviour like a lock and key
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Intention movements
- Signals that suggest preparedness to attack
- Incomplete FAP
- Preparatory movements
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Redirection
Where an FAP is directed at an individual or object other than that which released the response
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Thorndike
- Puzzle-box learning
- - trial and error learning
- - operant/instrumental conditioning
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Pavlov
- Associative learning
- - Classical conditiioning
- UCR: normal response. In response to UCS
- UCS: normal stimulus
- CS: Stimulus not initially associated with UCR (novel)
- CR: If the animal begins to respond to CS alone, we have a CR
CR is conditional upon CS predicting the arrival of UCS
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Effective Classical Conditioning 3#
- 1. CS presented before UCS (adaptive learning)
- 2. Latency between presentation of CS and UCS should be minimal
- 3. The CS should ideally be unique to a single UCS
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the adaptive nature of learning
- animals are pre-disposed to learn stimuli that predict significant events
- animals that attended to such signals and made associations enjoyed enhanced success, and thereofre became characteristic of population (evolutionary)
- Signals of little value after the fact
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Extinction
Unlearning of an association that no longer have any predictive validity
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L4 Treatment of Eneuresis
- Non bed wetters: UCS of a full bladder produces UCR of waking up, and subjects goes to the toilet
- Eneuresis sufferers: Full bladder does not act as a UCS promoting UCR
- Therefore we must condition a response (CR - waking up) in response to the CS of a full bladder
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Employing Apparatus
- Alarm acts as a UCS producing the UCR of waking up
- Repeated presentation of full bladder (CS) before alarm (UCS) ultimately leads to waking (CR) to full bladder because of CS predicting the alarm (UCS) and that UCS already waking the subject (UCR)
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Why doesn't the repeated presentation of the full bladder alone (CS) lead to extinction of the CR.
- Effects of operant conditioning appear
- - Parental praise
- - Waking up in a dry bed
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operant
learning is contingent upon operations the animal performs in its environment
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Law of Effect - Thorndike
Responses that are followed by a satisfying state of affairs tend to be repeated
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Classical versus Operant Conditioning
- Nature of association:
- Stimulus-stimulus (CS-UCS)
- Stimulus-response (Reward-Bhv.)
- Genesis of Response:
- Elicited: Subject doesn't control sequence of events
- Emitted: Subject's behaviour controls events
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Thorndike - Principle of Intellectual Continuity
Although species may differ in what they learn, or how rapidly they learn, the process of learning must be the same in all species
Behaviorism
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Behaviorism - Watson
Psychology as a behaviorist views it
Called for elimination from psychology all concepts that made reference to the mind.
Reflects earlier notion put forward by Morgan
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Morgan's Canon
Actions of animals should not be attributed to the exercise of a higher psychical faculty if they can be interpreted as the outcome of one that stands lower on a psychological scale
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Behaviorists
- Ejected mental events from the scientific study of psychology: Such events neither observable nor measurable -> Watershed event leading to conflict with ethologists
- Rejection of mental events also led to comparative psychologists rejecting instinct
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behaviorists focused on:
Skinner dominant figure among behaviourists
- Stimuli that elicit response
- Rewards and punishments that maintain them
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reinforcer
- an object or event that increases the probability of expression of a given behaviour
- Positive (addition of desirable stimulus) and negative (removal of a noxious stimuli) reinforcers
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punishment
- Objects or events that decrease the probability of expression of a given behaviour
- adding noxious stimulus
- removing a desirable stimulus
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Shaping
Production of desired behaviour by rewarding successive approximations to that behaviour
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Click and treat system
- combines elements of operant and classical conditioning
- Eventually click alone reinforces behaviour (secondary reinforcer/CS)
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Continuous Reinforcement schedule
- every performance of desired behaviour is rewarded
- promotes rapid acquisition
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Fixed ratio reinforcement schedule
- set number of responses required for reward
- establishes high response rates
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Variable ratio reinforcement schedule
- Random variation in number of responses before reward
- Highly resistant to extinction
- Also produces high response rate
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L5 Two other movements existed other than behaviourists
- Physiological psychologists
- Social Behaviorists
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Physiological psychology
def. and people
- Recognizes the importance of the nervous system and other physiological elements to behaviour
- Lashley and Beach
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Lashley
- Trained rats in mazes or on discrimination tasks
- Then selectively ablated area of cortex to test for influence on memory
- Also study neural substrates of vision and emotion by similar techniques
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Beach
- Pioneering work on role of nervous system and hormones in sexual behaviour of many animals
- Ablated parts of nervous, endocrine and reproductive systems to determine their normal role
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Social Behaviour
- Recognized that behaviour involved groups as well as individuals
- Focused on the expression and control of social behaviour
- - Robert Yerkes
- - Carpenter
- - Schneirla
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Yerkes, Clarence Carpenter, and theodore Schneirla
- Studied social interactions among primates in a field setting
- Studied a variety of different primate species in the field (mating systems and communication)
- Army ants. Lab and field studies on social behaviour
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beach, Schneirla, and Aronson formed group of animal psychologists called:
- Behavioural epigeneticists
- Emphasized complex interactions between genes and environment in development of behaviour
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Criticisms of Ethology
- 1. Lack of control and experimental rigor in field-based observational studies
- 2. Lack of quantification and failure to use inferential statistics
- 3. Abstract conceptual model: Suggesting that such models provided no useful infor about functioning of actual nervous system
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Criticisms of Comparative Psychology
- 1. Lack of natural context in lab studies - Unsuitable for expression of adaptive behaviour
- 2. Types of studied - suggested that such behaviours had no importance to rats in the natural context
- 3. Preoccupation with rats as subjects: suggesting that years of laboratory breeding removed any natural element of behaviour
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Nature/Nurture Controversy
- The relative importance of inheritance versus experience in producing behaviour
- Ethologists: behaviour inherited ina form suited to the life circumstace of the animal bc it is the product of natural selection
- Comparative Psychologists: Learning is the ultimate basis of behaviour, so instinct and innate tendencies are perpheral, or in fact are irrelevant
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Watson - Tabula rasa
Individuals are born with a black slate
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Challenges to Ethology
Behaviour that ethologists considered innate bc they were stereotyped and specie characteristic could be accounted for by experience e.g. Parasitoid wasps/ host preference via experience
Neither sterreotypy nor the observation that a beaviour is specie characteristic means that is independent of experience ESSENTIALLY IMPOSSIBLE TO PROVE
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Behavioural epigeneticists pointed out...
that even socially isolated animals may not be isolated from extrinsic stimuli that normally contribute to the development of their behaviour -> developing animal may provide the stimulus themselves
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Peking ducklings
individuals provide their stimulus
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Challenges to Comparative Psychology
Innate predispositions interfere with conditioned response
- Breland's banking raccoons (students)
- - No element of experience known to enter rubbing (FAP)
- - Operant associated with food and food prompts rubbing
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Breland's Conclusion
The behaviour of animal cannot be adequately understood, predicted, or controlled without knowledge of its instinctive patterns, evolutionary history, and ecological niche
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Next blow to Comparative Psychological ->
Principle of Equivalence of Associability
- All stimuli have an equal ability to become associated with a response
- NOT THE CASE!
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Garcia and Koelling's Experiment disputes what? How?
- Principle of Equivalence of Associability
- Conditioned rats to avoid sweetend using punishment of electric shock and x-ray irradiation
- -> Answer requires an understanding of natural history of rat feeding: pre-disposed to associate sickness with food
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Conflict between ethologists and Comparative Psychologists was essentially unfounded why?
- Tinbergen
- They were asking fundamentally different questions
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Tenbergen's 4 questions
- Q of Immediate Causation
- Q of Development
- Q of Function
- Q of Evolution
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Question of Immediate causation
- Deal with factor that elicit and control behaviour over the short run
- - The effects of various stimuli, sensory,a dn perceptual processes and the physiological mechanisms underlying behaviour
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Q of Development
- Deal with factors that control how behaviour changes within the organism's life
- - genetic, physiological and experiential factors that cause age-related changes in behaviour
- - Longer time frame than Q of immediate causation
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Wilson did what?
- Pooled Tinbergen's questions together
- Proximate question (how)
- Ultimate question (why)
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Q of function
- Deal with role behaviour plays in adapting organism to its environment
- - How does the behaviour in question enhance survival or propagation of the organisms genes?
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Q of evolution
- Deal with the way in which behaviour can be seen to change as natural selection operates within a species or in the appearance of new species.
- Unlike others, deals with time frame longer than the life of the organism
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Positive Aspects for ethologists
- 1. Came to realize that learning can play a role in expression of behaviour
- 2. Learned the value of the laboratory environment
- 3. Became more sophisticated in their application of the experimental method
- 4. Began to propose models in terms of actual physiology of the organism
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Positive aspects for CP
- 1. Came to realize that some elements of behaviour are under genetic control
- 2. Learned the value of the natural context for understanding behaviour and the potential dangers of examining behaviour only in the lab.
- 3. Gained renewed appreciation of the value of studying many different species
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How do we know that there is a genetic basis to behaviour?
Evidence for genetic basis of behaviour?
- Logical argument: Genetic code for structure; structure produces behaviour
- Artificial selection: Dogs; Tolman bright and dull rats
- Hybridization: Rothenbuhler Hygienic behaviour of honey bees; crickets
- Inbreeding: Avoidance learning in mice Bovet
- Molecular Genetics
- A. Genetic similarity detection: Sherman and Holmes with Belding's Ground Squirrels
- B. Genetic manipulation: Konopka and Benzer Patterns of emergence in Fruit flies
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Articial Selection
- Humans decide which animals will reproduce based on desirable traits.
- Such selective breeding can result in structural change.
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Tolman
- Examined maze navigation in rats
- - Because rearing conditions were held constant throughout, selection resulted in discrete groups, maze navigation behaviour is under genetic control
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Hybridization
- Cross individuals with different expressions of the same underlying behavioural traits
- - Control crosses can provide insight into the genetics of behaviour
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Codominance
- Crickets
- Hybrid males produce calls that are int. to parentals in both refquency and duration
- Hybrid females preferentially orient to int. calls
- Genetic control with codominance or incomplete dominance
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Inbreeding - Bovet
- Selective mating of closely related individuals
- Reduces heterozygosity
- Individuals within inbred lines are virtually genetically identical
- Different alleles become fixed in different inbred lines: Comparison of individuals from different lines reared in common environment reveals genetic contribution
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Avoidance learning in mice - Bovet
- Used shuttle box learning paradigm using different inbred mice lines and an outbred Swiss strain
- Because environment constant, differences in learning are genetically controlled
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Don't forget the environmental influences interact with genes in producing behaviour
- Holding environment constant allows assessment of genetic contributions to behaviour
- Holding genotype constant allows assessment of environmental contribution
- Inbred strains also provide a convenient source of genetically similar individuals with which to examine environmental effects
- - inbred strains provides a larger sample size than identical twins
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