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Units of selection Or distinguishing correlation from causation in evolutionary biology Adaptation A trait that has become prevalent because of a selective advantage


  1. Units ¡of ¡selection Or ¡distinguishing ¡correlation ¡from ¡causation ¡in ¡evolutionary ¡biology

  2. Adaptation A ¡trait ¡that ¡has ¡become ¡prevalent ¡because ¡of ¡a ¡selective ¡ advantage due ¡to ¡an ¡improvement ¡of ¡some ¡function

  3. At ¡what ¡level ¡do ¡adaptations ¡act? • genes • chromosomes • genomes • individuals • groups • species • species ¡community • ecosystem • biosphere ¡(-­‑> ¡“Gaia”)

  4. At ¡what ¡level ¡do ¡we ¡recognize ¡adaptations? Herd ¡of ¡fast ¡deer vs. Fast ¡herd ¡of ¡deer? • Is ¡speed ¡the ¡adaptation ¡of ¡the ¡herd ¡or ¡the ¡deer?

  5. The distinction between natural selection ACTING ON vs. SELECTING FOR A famous toy analogy (Sober 1989; Futuyma 1998)

  6. The distinction between natural selection ACTING ON vs. SELECTING FOR turn toy over; what happens?

  7. The distinction between natural selection ACTING ON vs. SELECTING FOR return toy; what happens?

  8. The distinction between natural selection ACTING ON vs. SELECTING FOR What does selection ACT ON? What is SELECTED FOR?

  9. The distinction between natural selection ACTING ON vs. SELECTING FOR The more realistic gene gene-phenotype gene relationship

  10. The distinction between natural selection ACTING ON vs. SELECTING FOR gene gene phenotype (= vehicle) built by genes

  11. The distinction between natural selection ACTING ON vs. SELECTING FOR gene gene phenotype (= vehicle) gene traveling built by genes inside vehicle

  12. The distinction between natural selection ACTING ON vs. SELECTING FOR gene gene gene gene blue vehicle built red vehicle built by blue genes by red genes

  13. The distinction between natural selection ACTING ON vs. SELECTING FOR gene gene gene gene gene gene

  14. The distinction between natural selection ACTING ON vs. SELECTING FOR gene gene gene gene gene gene gene gene gene gene gene gene gene gene a population of vehicles built by genes

  15. The distinction between natural selection ACTING ON vs. SELECTING FOR population gene gene gene gene gene gene gene before selection gene gene gene gene gene gene gene

  16. The distinction between natural selection ACTING ON vs. SELECTING FOR population gene gene gene gene gene gene gene before selection gene gene gene gene gene gene gene selection (differential survival & reproduction) population gene gene gene gene gene gene gene after selection gene gene gene gene gene gene gene

  17. The distinction between natural selection ACTING ON vs. SELECTING FOR population gene gene gene gene gene gene gene before selection gene gene gene gene gene gene gene selection (differential survival & reproduction) population gene gene gene gene gene gene gene after selection gene gene gene gene gene gene gene What does selection ACT ON? What is SELECTED FOR?

  18. The distinction between natural selection ACTING ON vs. SELECTING FOR population gene gene gene gene gene gene gene before selection gene gene gene gene gene gene gene selection (differential survival & reproduction) population gene gene gene gene gene gene gene after selection gene gene gene gene gene gene gene Natural selection: ACTS ON phenotype (= “vehicle” = unit of selective interaction) SELECTS FOR genes (= “replicators” = unit of inheritance) that build the phenotype / vehicle

  19. Selfish ¡genes • Transposons • Homing ¡endonucleases • Segregation ¡distortion • Sex-­‑ratio ¡distorters • Altruism ¡by ¡kin ¡selection

  20. Transposons

  21. Homing ¡endonuclease

  22. Fun ¡aside: ¡domestication ¡of ¡a ¡selfish ¡element

  23. SELF-INTERESTED (SELFISH) GENES: Segregation Distorters e.g., meiotic drive in Drosophila Germ cell Line Gametes normal gamete production

  24. SELF-INTERESTED (SELFISH) GENES: Segregation Distorters e.g., meiotic drive in Drosophila Germ cell Line Gametes Meiotic drive : “blue” allele eliminates gametes carrying “red” allele => all suriving gametes carry “blue”

  25. SELF-INTERESTED (SELFISH) GENES: Sex-Ratio Distorters e.g., psr -gene in Nasonia jewel-wasp Nasonia vitripennis

  26. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants)

  27. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants) diploid haploid 2n 1n

  28. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants) diploid haploid 2n 1n meiosis no meiosis in female in male (= males make one type of sperm)

  29. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants) diploid haploid 2n 1n meiosis no meiosis in female in male (= males make one type of sperm) unfertilized egg fertilized egg haploid (1n) diploid (2n)

  30. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants) diploid haploid 2n 1n meiosis no meiosis in female in male (= males make one type of sperm) unfertilized egg fertilized egg haploid (1n) diploid (2n) => =>

  31. Need to understand first: Haplodiploid Sex Determination in the Hymenoptera (wasps, bees, ants) diploid haploid 2n 1n meiosis no meiosis in female in male (= males make one type of sperm) unfertilized egg fertilized egg haploid (1n) diploid (2n) => => Only females have sons & daughters; Males do not have sons, only daughters

  32. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males psr chromosome

  33. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males psr chromosome

  34. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males - sperm containing psr causes paternal set of chromosome to condense (=> elimination), whereas psr survives psr chromosome Elimination of all paternal chromosomes (except psr )

  35. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males - sperm containing psr causes paternal set of chromosome to condense (=> elimination), whereas psr survives psr chromosome Elimination of all paternal chromosomes (except psr) => haploidy => male

  36. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males - sperm containing psr causes paternal set of chromosome to condense (=> elimination), whereas psr survives psr chromosome Elimination of all paternal chromosomes (except psr) => haploidy => male - psr causes chromosomal “fratricide”, thus causing haploidy (=> offspring of psr -male are all male)

  37. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males - sperm containing psr causes paternal set of chromosome to condense (=> elimination), whereas psr survives psr = “paternal sex ratio” psr chromosome because sex-ratio is determined by male in Elimination of all paternal some matings chromosomes (except psr) => haploidy => male - psr causes chromosomal “fratricide”, thus causing haploidy (=> offspring of psr -male are all male)

  38. SELFISH GENES: Sex-Ratio Distorters psr -gene in Nasonia vitripennis - supernumary chromosome ( psr ); found only in males - sperm containing psr causes paternal set of chromosome to condense (=> elimination), whereas psr survives psr chromosome Elimination of all paternal chromosomes (except psr) => haploidy => male - psr causes chromosomal “fratricide”, thus causing haploidy (=> offspring of psr -male are all male) => psr doubles its rate of reproduction, because it circumvents 50% chance of elimination during meiosis (i.e., psr manages to avoid the cost of meiosis)

  39. SELF-INTERESTED (SELFISH) GENES: Natural selection can select for genes against the interest of the individual For example: - Repetitive DNA - Spread of transposons in a genome - Segregation distorters (e.g., meiotic drive in Drosophila ) - Sex-ratio distorters (e.g., psr -gene in the jewel wasp Nasonia vitripennis )

  40. SELF-INTERESTED (SELFISH) GENES: Natural selection can select for genes against the interest of the individual For example: - Repetitive DNA - Spread of transposons in a genome - Segregation distorters (e.g., meiotic drive in Drosophila ) - Sex-ratio distorters (e.g., psr -gene in the jewel wasp Nasonia vitripennis ) - Altruism and kin-selection

  41. Helping / Altruism is widespread among animals e.g., alarm calling in Belding’s Ground Squirrel

  42. Helping / Altruism is widespread among animals e.g., helping in Florida scrub jays

  43. Altruism = fitness-reducing behavior that benefits another individual

  44. Altruism = fitness-reducing behavior that benefits another individual Evolution of altruism / helping can be understood in terms of indirect fitness effects (effects on relatives)

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