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Organism Life Histories BIOL400 9 November 2015
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Energy Allocation An organism assimilates a finite amount of energy, which it can devote to: Growth Growth Reproduction Reproduction Tissue maintenance Tissue maintenance Storage for later (see above) Storage for later (see above)
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Fitness Measure of survival likelihood and reproductive output Natural selection, based on the physical and biotic environment, determines the life history a species has The one that maximizes fitness The one that maximizes fitness
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Fig. 2.8 p. 25 Selection for optimum clutch size of 11 in blue tits
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Fig. 2.9 p. 25 Experimental demonstration that selection has not optimized clutch size in the house wren (unless there is a life-history trade-off…)
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Key Life-History Attributes Growth rate Age at maturity Size at maturity Reproductive frequency Annual Annual Lifetime Lifetime Fecundity Propagule size
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Life-History Trade-Offs Stearns: “Linkages between traits that constrain the simultaneous evolution of two or more traits” Increased allocation toward A decreases possible allocation toward B
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Present Reproduction vs. Survival and Future Reproduction
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House wren?? Red deer Kenyan Lobelia Beech trees Fig. 2.9 p. 25 Fig. 8.20 p. 137 Fig. 8.19 p. 136 HANDOUT
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Maturation age vs. fecundity and/or propagule size and survival HANDOUT
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Egg Size vs. Clutch Size Trade-off may select for point at which increasing clutch size leads to lower fitness by reducing offspring size, and increasing offspring size leads to lower fitness by reducing clutch size Hence, an optimal egg size
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Optimal Egg Size Vs. Anatomical Constraints on Egg Size If selection optimizes egg size, egg size should not correlate with female body size However, anatomical contraints may cause eggs of small females to be smaller than optimum Egg size increases with female body size Egg size increases with female body size
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HANDOUT Congdon and Gibbons 1987
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HANDOUT—Doughty 1997
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Human Menopause Trading off future reproduction and its increased risks against helping of grandchildren?
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Life-History Invariants
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Invariant … …ratios (“dimensionless numbers”) …ratios (“dimensionless numbers”) …X-Y relationships with set slopes …X-Y relationships with set slopes Demonstrate the nature of life-history trade-offs
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HANDOUTS—Charnov 1993
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Genotype and Phenotype Is variation in life history… …genetic (induced by DNA)? …phenotypic (induced by environment)? …phenotypic (induced by environment)?
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Fig. 6.11 p. 91 Common-garden experiments demonstrate genotypic effect—adaptation to local conditions? All grown in identical greenhouse conditions Yarrow
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Reaction Norm Stearns’ definition: "The mapping of the genotype onto the phenotype as a function of the environment—expressed as a plot of phenotypic values [Y] against environmental values [X]. The reaction norm of a genotype is the full set of phenotypes that the genotype will express in interaction with the full set of environments in which it can survive."
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Countergradient Variation Seemingly good evidence that much of the variation in organism life histories must be genetic and adaptive Genetic basis verified in common-garden laboratory experiments
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HANDOUT—Conover and Present 1990
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Categorizing Life-History Strategies
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r- and K-selection Pianka (1970) Name denotes r and K in logistic growth equation r is intrinsic rate of increaser is intrinsic rate of increase K is karrying kapacityK is karrying kapacity
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Table 10.2 p. 180
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Fig. 10.20 p. 181 Grimes (1979) Ruderal, competitive, and tolerant plant life histories
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Salisbury (1942) The Reproductive Capacity of Plants Anticipated the r-selection/K-selection dichotomy in plants, measuring the mass in mg of various plants' seeds: Open habitats: 120 mg Open habitats: 120 mg Semi-closed: 220 mg Semi-closed: 220 mg Meadows: 490 mg Meadows: 490 mg Wood margins: 440 mg Wood margins: 440 mg Shaded habitats:1400 mg Shaded habitats:1400 mg
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Fig. 10.21 p. 182 Larger seeds have higher survival rates
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Bet Hedging Some species spread reproductive effort over long lifespan May have unpredictably variable juvenile mortality that is often high Hence no advantage to investing heavily in reproduction in any year—trade reduced reproductive effort off against increased adult survival Opposite of “big bang” reproducers Opposite of “big bang” reproducers
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p. 138 Bet hedgers in lower left Big-bang reproducers in upper right
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