WHY OVERPRODUCTION? • The Insurance Hypothesis: – 1) Replacement – excess offspring are produced to replace any core sib that disappears (e.g. predation, disease) – 2) In a very good year, everyone lives but in a bad year only the older sibs survive – 3) Parents do not intercede to reduce impact of sibling rivalries A THOUGHT • Overproduction of eggs should only be evolutionary advantageous when eggs are cheap to produce AT THE EXTREME • Trophic eggs rarely hatch (or are greatly delayed) and are eaten by earlier hatching sibs when other foods are hard to find – The Icebox Hypothesis TERRIBLY IMPORTANT • The relationship between genotype and phenotype is critical to our understanding of natural selection • Nature selects for phenotypes but genes may cause particular phenotypes to be expressed CLASSIC MENDELIAN GENETICS • PP x pp gives 3:1 ratio in F2 generation based upon 0.25 PP, 0.5 Pp and 0.25 pp PHENOTYPE • Def’n - the realized expression of the genotype that results from the biological activity of the dna molecules! FACTORS EFFECTING PHENOTYPE • • • • • • Multiple alleles Incomplete dominance Epistasis Polygenic traits Pleiotropy Environment IMPORTANT INSIGHT • None of the factors on the previous overhead invalidate the Mendel’s idea of particulate inheritance and random segregation they just complicate it! MULTIPLE ALLELES • E.g. Human blood type alleles A,B,O • This increases the number of possible genotypes in the population: AA, AB, BB, OB, OA, OO = 6 genotypes • The O allele lacks both A and B antigens • Phenotypes = Type A, Type B, Type AB and Type O MULTIPLE ALLELES • E.g. Drosophila wing alleles • Wild type, vestigial, antlered, intermediate • 6 genotypes: – Ww, Wwv, Wwa – wv wv – wa wa – wv wa Wild type Vestigial Antlered Intermediate MULTIPLE ALLELES • E.g. Drosophila wing alleles INCOMPLETE DOMINANCE • A single copy of the dominant allele does not produce the maximum phenotypic effect INCOMPLETE DOMINANCE • .. EPISTASIS • The effect of genes at one locus can effect the expression of a gene at another locus _ _ ee bb E_ B_E_ POLYGENIC TRAITS aaBbCC SKIN TONE AABbCc aaBBCc aaBbCc AaBBCc AaBbCc AabbCc AaBbCC AabbCC AaBbcc AABBcc AaBBcc aabbCC aabbCc AABbCC AAbbCC AAbbCC aaBBcc aaBbcc AABBCC AABBCc aaBBCC AABbcc AAbbcc Aabbcc AaBBCC DARK aabbcc LIGHT POLYGENIC TRAITS • • • • • • • Some examples: Crop Yield Some Plant Disease Resistances Weight Gain in Animals Fat Content of Meat IQ Scores Blood Pressure PLEIOTROPY • A single gene affects many phenotypic characters e.g. sickle cell allele, cystic fibrosis! ANTAGONISTIC PLEIOTROPY • Can it explain finite lifespan and senescence?! 2. 5 LOG MR 2 POW’s 1. 5 1 0. 5 AUSSIE’s 0 0 2 0 4 0 6 0 8 0 ANTAGONISTIC PLEIOTROPY Late Life • George Williams – selection for early life performance traits may select against old age performance – an life history tradeoff due to antagonistic pleiotropy! Early Life A CAUTION Environmental conditions can mask antagonistic pleiotropy. ENVIRONMENT Gene expression often depends upon the environment 2 ENVIRONMENTS YOLK PLANT MATERNAL EFFECT • Def’n - the effect of the environment provided by a parent on characteristics of its offspring! TEMPERATURE EFFECT ON SEX EXPRESSION • Maternal effect in sea turtles 2 TYPES OF MATERNAL EFFECTS • Common environment • Mother-daughter phenotype match 2 TYPES OF MATERNAL EFFECTS • Common environment 2 TYPES OF MATERNAL EFFECTS • Mother-daughter phenotype match A THOUGHT • Since they affect performance in offspring, maternal effects should be under selection EPIGENETICS • The study of heritable changes in gene expression and function that cannot be explained by changes in DNA sequence. EPIGENETIC MECHANISMS • (i) methylation of cytosine in the DNA, (ii) remodelling of chromatin structure of histone proteins and (iii) regulatory processes mediated by small RNA molecules EPIGENETIC IMPACT • Epigenetic changes may get passed on to daughter cells when altered DNA is replicated during mitosis or meiosis CAUSES OF EPIGENETIC CHANGES • Exposure to toxins • Control during morphogenesis to shut down specific genes to ensure that their products are not made in particular cells (e.g. kidney cells only function as kidney cells even though they have the software for all other cells)