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In the Light of Evolution: Volume 1. Adaptation and Complex Design (2007)
National Academy of Sciences (NAS)

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. "5 The Frailty of Adaptive Hypotheses for the Origins of Organismal Complexity--MICHAEL LYNCH." In the Light of Evolution: Volume 1. Adaptation and Complex Design. Washington, DC: The National Academies Press, 2007.

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In the Light of Evolution, Volume I: Adaptation and Complex Design

hand, standard theory (Kimura, 1983) shows that the fixation probability of a mutation to A is es times that for a mutation to a, where S = 2 Ngs is twice the ratio of the power of selection to the power of random genetic drift (1/Ng). Because the population-level rate of transition from one allelic type to another is equal to the product of the mutation rate and the fixation rate, the ratio of probabilities of being in states A versus a at selection-mutation-drift equilibrium is simply mes (Fig. 5.1). This simple expression leads to two general conclusions: (i) regardless of the strength of selection, if 1/Ng >> |s| the population will be driven to a state expected under mutation pressure alone; and (ii) the equilibrium composition of a population depends not on the absolute power of mutation, but on the relative rates of forward and reverse mutations (the mutation bias).

FIGURE 5.1 The long-term probability that an allele residing at a biallelic locus will be of the selectively advantageous type, given a selective advantage s, an effective population number of gene copies of Ng, and a mutation rate to the beneficial allele m times that in the reverse direction. The lowest curve (2 Ngs = 0.0) denotes neutrality, whereas the upper line (2Ngs = ∞) denotes an effectively infinite population, such that genetic drift is a negligible force.

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Front Matter (R1-R18)
Part I: INTRODUCTORY ESSAY (1-2)
1 Darwin's Greatest Discovery: Design Without Designer--FRANCISCO J. AYALA (3-22)
Part II: EPISTEMOLOGICAL APPROACHES TO BIOCOMPLEXITY ASSESSMENT (23-24)
2 Functional Information and the Emergence of Biocomplexity--ROBERT M. HAZEN, PATRICK L. GRIFFIN, JAMES M. CAROTHERS, and JACK W. SZOSTAK (25-44)
3 The Theory of Facilitated Variation--JOHN GERHART and MARC KIRSCHNER (45-64)
4 Between ‘‘Design'' and ‘‘Bricolage'': Genetic Networks, Levels of Selection, and Adaptive Evolution--ADAM S. WILKINS (65-82)
5 The Frailty of Adaptive Hypotheses for the Origins of Organismal Complexity--MICHAEL LYNCH (83-104)
Part III: FROM INDIVIDUAL ONTOGENY TO SYMBIOSIS: A HIERARCHY OF COMPLEXITY (105-108)
6 Emerging Principles of Regulatory Evolution--BENJAMIN PRUD'HOMME, NICOLAS GOMPEL, and SEAN B. CARROLL (109-128)
7 Evolution of Individuality During the Transition from Unicellular to Multicellular Life--RICHARD E. MICHOD (129-144)
8 Insect Societies as Divided Organisms: The Complexities of Purpose and Cross-Purpose--JOAN E. STRASSMANN and DAVID C. QUELLER (145-164)
9 Symbiosis as an Adaptive Process and Source of Phenotypic Complexity--NANCY A. MORAN (165-182)
Part IV: CASE STUDIES: DISSECTING COMPLEX PHENOTYPES (183-186)
10 Adaptive Evolution of Color Vision as Seen Through the Eyes of Butterflies--FRANCESCA D. FRENTIU, GARY D. BERNARD, CRISTINA I. CUEVAS, MARILOU P. SISON-MANGUS, KATHLEEN L. PRUDIC, and ADRIANA D. BRISCOE (187-204)
11 Plant Domestication, a Unique Opportunity to Identify the Genetic Basis of Adaptation--JEFFREY ROSS-IBARRA, PETER L. MORRELL, and BRANDON S. GAUT (205-224)
12 An Experimental Test of Evolutionary Trade-Offs During Temperature Adaptation--ALBERT F. BENNETT and RICHARD E. LENSKI (225-238)
13 Two Routes to Functional Adaptation: Tibetan and Andean High-Altitude Natives--CYNTHIA M. BEALL (239-256)
14 On the Origin and Evolutionary Diversification of Beetle Horns--DOUGLAS J. EMLEN, LAURA CORLEY LAVINE, and BEN EWEN-CAMPEN (257-282)
Part V: CONCLUDING ESSAY (283-284)
15 Biological Design in Science Classrooms--EUGENIE C. SCOTT and NICHOLAS J. MATZKE (285-304)
References (305-344)
Index (345-360)