1. Genetic Determinism and Heritability Gene selectionism All instances of selection are instances of selection acting on genes (rather than individuals or groups) How can selection act on genes?

2. Genetic Determinism and Heritability How can selection act on genes? It selects genes via their phenotypic effects What is the relation between genes and their effects? Received view: Genes are major causes of development and traits Many traits have a genetic basis are inherited are under genetic control are genetically determined

3. Genetic determinism the ultrabithorax mutation causes flies to have a second pair of wings

4. Genetic determinism G determines trait X – what does this mean? (1) Having G is the only cause for developing X Rejected by todays interactionist consensus: All genes interact with environmental factors to produce any trait (2) Any organism with G develops X Counter-example: many organisms have G without developing X – all those G-individuals that die before X is being expressed (e.g. flower petals)

5. Genetic determinism G determines trait X – what does this mean? (3) G has a flat norm of reaction Change in environment makes no difference to phenotype

6. Developmental Systems Theory Interactionism is an inadequate thesis about the causal role of genes in development and inheritance: it still privileges genes Genes are just one of many causal components DST comes in many versions; many, but not all, defend the following more specific theses 1.Organisms inherit more than genes 2.The notion of gene for X is incoherent 3.The parity thesis

7. 2.The notion of gene for X is incoherent Gene for X requires that the genes causal contribution can be isolated from that of the environment But their contributions are inseparable and incommensurable Lewontins brick and mortar metaphor One builder lays bricks, the other applies mortar Which is more important to build a house? Genes and environment lack a uniform kind of effect by which their contributions could be compared Contrast: two people pushing a car (both exert a force)

8. 2.The notion of gene for X is incoherent Defending genes for (Kitcher) Allele A is for root proliferation iff AB-trees produce more roots than BB-trees, given a standard environment and keeping other things constant A makes a phenotypic difference in standard environments Standard environment statistically normal (frequently encountered by species) does not reduce mean population fitness

9. 2.The notion of gene for X is incoherent The Acorn objection (Griffiths, Gray) In standard environments for oak trees most acorns rot Therefore, an acorns having A will not make a phenotypic difference in standard environment: AB offspring will on average produce as many roots as BB, i.e. none

10. 2.The notion of gene for X is incoherent Refinement (Kitcher) Allele A is for root proliferation iff AB-trees produce more or equal number of roots than BB- trees, given a standard environment and keeping other things constant Prematurely dying ABs produce equal number (0) Surviving ABs produce more

11. Heritability Heritability analysis is a statistical procedure to measure the causal contribution of genes 1) Heritability as resemblance (non-standard, e.g. Sober) Degree of parent-offspring similarity; independent of the degree to which genes are responsible for similarity 2) Heritability in the sense of heritability analysis Degree to which phenotypic variation is due to genetic differences – measured as a fraction of total variance broad heritability:H 2 = V G / V P narrow heritability:h 2 = V A / V P

12. Heritability Genotypic difference makes 3-unit difference on phenotype Environmental difference also makes 3-unit difference H 2 = 50%

13. Heritability Genotypic difference makes 3-unit difference on phenotype Environmental difference just makes 1-unit difference H 2 >> 50%

14. Heritability An important assumption for heritability analyses: Additivity = lack of statistical gene-environment interaction Genotype makes the same difference in all environments, i.e. there is an overall genotypic effect

15. Heritability Statistical gene-environment interaction: Genotype does not make the same difference in all environments E1: +3 E2: 0 E3: -3 There is no overall genotypic effect

16. Objection to Heritability Studies (Lewontin) Heritability studies are useless for tracing causal relations 1. Additivity rarely obtains: different populations live in very different environments, leading to different phenotypic effects for any given genotype

17. Objection to Heritability Studies (Lewontin) Heritability studies are useless for tracing causal relations 1. Additivity rarely obtains: different populations live in very different environments, leading to different phenotypic effects for any given genotype C1. Therefore, heritability estimates are local: a given heritability value may be different outside the narrow range of environments that hold in the study population it may not be extrapolated C2. Therefore, heritability cannot provide evidence of general relation between cause and effect

18. Response (Sesardic) Ad 1) Additivity rarely obtains Empirical data are few and inconclusive Most philosophers simply take Lewontins word for it Ad C2) Heritability does not trace causation Accepts that we cannot make global claims across all or even a wide range of not investigated environments But it does not follow that the local claims are not causal: Provided additivity holds in a given context, heritability does quantify the causal contribution of genotypes within that context