Nature, Nurture, and Behaviour

1. Nature, Nurture, and Behaviour

2. Behavioural Genetics • Genetic contribution to behaviour • But also environmental role • Interaction • Could just as well say BG studies the role of environment on behaviour as that of genetics…

3. Polygenic Traits • Single gene effects (relatively) easy to study and understand • Polygenic effects • Complex behaviour traits • Environmental interactions for both single and polygenic effects, but much more complex for the latter

4. Heredity • Obviously important (and easily seen) in single gene effects • More difficult to assess heredity’s role in complex behaviour traits

5. Synthesis • Quantitative genetics • Estimates degree to which differences among individuals due to genetic and environmental differences • Doesn’t identify specific genes or environmental factors • Molecular genetics • Identifies specific genes in behaviours

6. Humans • Limited experimental designs • Ethics • Twin studies, adoption studies, naturally existing polymorphisms

7. Non-human Animal Studies • Much more empirical control • Induce polymorphisms, transgenics, etc. • Even simpler: directed selective breeding studies and inbred strain studies

8. Selected Breeding • Variability in individuals’ behaviour due to genetic variance • Variability can be exploited to show role of genetics on behaviour • Pick specific trait, cross-breed individuals with that trait for multiple generations • Can also study environmental effects this way • E.g., cross-fostering, different nutrition levels, training protocols, etc.

9. Inbred Strain Studies • Brothers and sisters mated for at least 20 generations • Result is individuals that are virtual clones of all other members of strain • Any differences therefore due to environmental effects • Prenatal and postnatal environmental factors • Also, genetic factors, of course

10. Techniques • Cross different strains to look at changes in behaviour across subsequent generations • Diallel design • Compares several inbred strains and all possible F1 crosses • Cross fostering • F1 crosses to mother from one strain or the other • Assess role of postnatal care • Environmental manipulation • Alter specific environmental factors to examine genotype-environmental interactions

11. Scott & Fuller (1965) • 13 year study of hundreds of crossbreed and purebreed dogs • Beagle, Shetland sheep dog, African basenji, wire-haired fox terrier, American cocker spaniel • Behaviour patterns • Dogs perhaps the oldest domesticated species • High chromosomal similarity across breeds, but high behavioural differences; easily picked out by selective breeding (Borgaonkar et al., 1968)

12. Critical Developmental Periods • Heightened sensitivity to specific environmental stimuli • Time critical • Gene-environmental interaction

13. Puppy Toileting • Born blind and deaf, but excellent sense of smell • Mother needs to stimulate them to pass feces and urine • Mother consumes puppies’ waste; genetic programming to keep nest clean and free of smells that would attract predators • Activated by having litter; otherwise dogs not normally coprophagic

14. After 4 weeks, pups mobile • Nursing mother trains pups not to toilet near nest • Disciplining behaviour can appear harsh • Pups separated from mother too early don’t learn proper toileting • Subsequently have difficulties learning to be housebroken; critical period missed

15. First Fear Period • Fear/hazard avoidance starts in pups at about 5 weeks, peaks between 8 to 10 weeks • Typically when people take puppy home • Easier to condition fear responses at this time • Fear of cars, fear of vet clinic, etc. • (Rawlinson, 2005)

16. Farm-Fox Experiment • Domestication • “Self-domestication” from colonization of new ecological niches? (Morey 1994) • Intentional decisions by human beings? • Trut (1999): 40 year selective breeding study of 45,000 Russian silver foxes <http://i.pbase.com/o4/29/336929/1/58820261. T1333SilverFoxFaceOn.jpg>

17. Patterns of Domestication • Darwin recognized similar changes in appearance and behaviour across species Appearance of dwarf and giant varieties All Piebald coat colour All Wavy or curly hair Sheep, poodles, donkeys, horses, pigs, goats, mice, guinea pigs Rolled tails Dogs, pigs Shortened tails, fewer vertebrae Dogs, cats, sheep Floppy ears Dogs, cats, pigs, horses, sheep, goats, cattle Changes in reproductive cycle All except sheep

18. Selection • Dmitry Belyaev • Believed behaviour, not size or reproduction, was key factor in changes from domestication • Specifically, selection for amenability (tamability) • In 1959 started selected breeding of Vulpes vulpes for tamability

19. One Change to Many • Balance of hormones and neurotransmitters at level of whole organism • Slight changes to regulatory genes for some trait (e.g., amenability to domestication) can have wide-ranging effects on endocrine system • Cascading effects on animal development, physiologically and behaviourally

20. Belyaev’s Hypothesis • Two conditions required • Variations in tamability must be partly determined by genes • Domestication places animal under strong selective pressure <http://sybilsden.com/images/imgfox /darla-2006/07-2006/08darla14.jpg>

21. Cross-Breeding Studies • Cross-bred foxes of different behaviour • Cross-fostered newborns • Transplanted embryos between donor and host mothers with different tamability levels • About 35% of variation in foxes’ defense response to people is genetically determined (i.e., heritability)

22. Selective Pressure? • Attempted domestication of other wild-type animals • River otters (Lutra lutra) and gray rats (Rattus norvegicus) • Only 14-16% of wild-caught individuals showing weak defensive reactions produce offspring that survive to adulthood • Domestication puts animals under extreme stress and selective pressure

23. The Experiment • 30 male and 100 female foxes, mostly from commercial fur farms in Estonia • So, already tamer than wild relatives • Selected for tameness alone • Only about 4-5% of male and 20% of female offspring allowed to breed • No training of foxes, minimal human contact • Ensures tameness due to genetic selection

24. Behavioural Classes • Class III • Flee from experimenters or try to bite when handled • Class II • Let themselves be handled but show no emotionally friendly response • Class I • Friendly to experimenters, wagging tails and whining • Class IE • Domesticated “elite” • Eager for human contact, whimper to attract attention, sniff & lick experimenters like dogs • Appeared in sixth generation; by 10th gen. 18% of pups; by 20th gen., 35%; by 35th gen., 75%

25. Physical Changes • Domesticated fox pups respond to sounds two days earlier and open eyes one day earlier than wild type • Fear response starts at 9 weeks in domesticated foxes (instead of 6 weeks in wild type) • More time to become incorporated into human social environment • Appearance: • From 10th gen., changes in coat colour • From 10th - 15th gen., floppy ears seen • From 15th - 20th gen., tails and legs shortening • These are still unusual traits, but more common than in wild-type

26. Traits Days Weeks 11 12 13 14 15 16 17 18 19 4 6 8 10 12 Farmed Foxes Domesticated Foxes Dogs Fear of unknown Response to sound Eyes fully open Window of socialization

27. Ontogenetic Changes • Changes in mechanisms regulating development • Effects on normal timing of embryonic and post-embryonic development

28. Animals per 100,000 with trait Increase Characteristic Domesticated Nondomesticated in frequency Population Population (percent) Depigmentation (Star) 12,400 710 +1,646 Brown mottling 450 86 +423 Gray hairs 500 100 +400 Floppy ears 230 170 +35 Short tail 140 2 +6,900 Tail curled up 9,400 830 +1,033 Incidence of Change

29. Inbreeding? • Maybe changes due to recessive alleles • Would manifest from long-term inbreeding • Study designed to prevent this • Outbreeding with commercial fox farm animals • Inbreeding coefficients of 0.02-0.07 • Some of new traits are dominant, not recessive • Couldn’t have been hidden carriers in original population

30. Star Gene • White patch on head • Affects migration rate of melanoblasts (embryonic precursors of pigment cells) • Melanoblasts form in embryonic neural crest, normally moving to embryo’s epidermis between 28-31 days after conception • Foxes with single Star gene • Melanoblasts’ movement delayed two days • Leads to death of “tardy” melanoblasts • Alters pigment pattern, producing star pattern

31. Corticosteroids • Changes in plasma levels of corticosteroids, hormones linked to adaptation to stress • Adrenal glands producing less adrenalin • Hence, lowered stress response, and more tamable • Adrenalin also on biochemical pathway, linked to, among other things, melanin (pigment) • Changes in coat colour • Different hormone levels have far-reaching effects • Cascade of changes to behaviour and physiology • Polygenic and environmental interaction • Video

32. Human Behaviour Studies • Essentially “observational” in nature • Limited to naturally occurring genetics and environmental variation • Adoption designs • Twin designs • Combination designs

33. Adoption Designs • Traits running in families can be due to nature and/or nurture • Adoption studies can separate heredity and environmental factors

34. Parents and Offspring • Genetically related family • Shared genes and environment • Adoption produces “genetic” (i.e., birth) and “environmental” (i.e., adoptive) parents • Comparing genetic parents and offspring assesses genetic contribution to parent-offspring similarity • Comparing environmental parents and offspring assesses post-natal environmental contribution to parent-offspring similarity

35. Siblings • Similarly, genetic and environmental siblings • Again, comparisons made to assess genetic and post-natal environmental effects

36. Designs • Adoptees’ study method • Examine incidence of trait in adopted away offspring of parents who possess the trait • Adoptees’ family method • Compare adopted offspring who possess trait and lack the trait of interest (control offspring) • Assess incidence of trait in the biological and adoptive families • Genetic influence if trait greater for biological relatives of affected adoptees than biological relatives of controls • Environmental influence if incidence greater for adoptive relatives of affected adoptees than biological relatives of controls

37. Representative? • Are adopted parents and adopted children representative of general population? • Could be biases related to those parents who adopt and children who are adoptees • Some studies suggest no bias with respect to population, others raise questions

38. Prenatal Environment • Can be difficult to assess this • Sometimes simply hard to get access to information about prenatal conditions of adopted children post-hoc • Various environmental factors could be different between mothers (and fathers) who keep, as opposed to give up, their offspring

39. Selective Placement • Placement of adoptive children may not be random • E.g., certainly isn’t based on appearance • Preference for like characteristics between adoptive parents and offspring, inflates apparent correlation between biological parents and adopted-away children • Inflates effect of genes on behaviour

40. Selective Placement (Adjusted) • Matches biological and adoptive parents • Genetic influences can increase correlations b/t adoptive parents and their adopted children • Environmental influence can increase correlations b/t biological parents and their adopted-away children

41. Twin Designs • Monozygotic (MZ) • From single fertilized egg • Genotypically identical • Can be environmental differences, though • Dizygotic (DZ) twins • From separately fertilized eggs • Just siblings born at the same time • Genotypically different; share 50% genes • Allows examination of genetic and environmental variance of a behaviour (a phenotype)

42. Relations • First degree relatives • 50% genetically related, r=0.5 (“coefficient of relatedness”) • E.g., parents and children, full non-MZ siblings • Second degree relatives • 25% genetically related, r=0.25 • E.g., grandparents and grandchildren, half-siblings • Third degree relatives • 12.5% genetically related, r=0.125 • E.g., cousins

43. Equal Environment Assumption • Common to twin design methods • Assumes environmentally caused similarity about the same for both MZ and DZ twins • If MZ twins experienced more similar environments than DZ twins, then assumption would be violated • Studies show the assumption probably holds for most traits

44. Prenatal Environment • Interestingly, MZ twins experience greater prenatal environmental differences than DZ twins • Competition for resources hypothesis • MZ twins identical, so require exactly same resources • Limited prenatal resources; competition produces prenatal stress

45. Are Twins Representative? • Twins not representative of population • Often born prematurely, uterine environment more adverse • E.g., twins usually 30% lower in birth weight than singletons, slower language development, poorer performance on early verbal and IQ tests • However, most differences typically disappear by midchildhood • Shared and unique postnatal environmental effects

46. Combination Designs • Combine family, adoptive, and twin methods • Twin and family methods • MZ twins have r=1.0 • Their children (i.e., cousins) will be r=0.25, not typical r=0.125 of cousins • Nephew of MZ male twins shares same r-value (0.5) with his father as his uncle, but there are different environmental factors in place • Step-family analysis • Get half-siblings (r=0.25) and full siblings (r=0.5) in same family • Different environmental effects for different children

47. Heritability • Proportion of phenotypic variation in a population attributable to genetic variation among individuals • Variation among individuals may be due to genetic and/or environmental factors

48. A Warning • Heritability measures always refer to degree of variation between individuals in a population • Can not be applied at level of individual • E.g., if heritability index of trait is 0.4 it is incorrect to say that 40% of this trait is from the parents and 60% is from the environment • In clones the heritability index for all traits is zero (all variability between individuals due to environment).

49. Statistical Significance • Is similarity between genetically related individuals significantly greater than random? • Depends on sample size • Note that some significant effects really have very little “real” effect • I.e., account for little variance in a trait

50. Effect Size • Extent to which individual differences for a trait in the population can be accounted for by the genetic difference among individuals • Effect of heritable genetic disability can be huge for an individual, but have little effect on population • E.g., if trait only exists in 1 in 10,000 individuals, little overall variation in trait for the entire population • Many statistically significant environmental effects in behavioural sciences have small effects in population