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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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. "4 Between ‘‘Design'' and ‘‘Bricolage'': Genetic Networks, Levels of Selection, and Adaptive Evolution--ADAM S. WILKINS." 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

hierarchy, then the nature of the gene tinkering process is seen to be much less haphazard than the process connoted by the term “bricolage,” having more built-in molecular constraints yet, at the same time, lacking the goal-directed nature of a process that is implied by the term “design.”

BUT IS A NETWORK PERSPECTIVE TRULY NECESSARY TO UNDERSTAND ADAPTIVE EVOLUTIONARY CHANGES?

All of the above statements can be accepted, however, without embracing the idea that the network perspective is needed for experimental research in evolution. A case can be made that a sufficient understanding of the genetic basis of adaptive evolutionary changes emerges from classic quantitative trait locus and molecular single gene-based experimental perspectives and that neither the concept of networks nor detailed knowledge of particular networks is needed for actual progress. This position is seemingly bolstered by recent success in understanding the genetic foundations of several adaptive traits, work that has underlined the key importance of a small number of specific genes. These cases involve finch beak dimensions in Darwin’s finches, characteristics related directly to specific feeding adaptations (Abzhanov et al., 2004, 2006); the adaptive evolution of bat wings for flight (Sears et al., 2006; Weatherbee et al., 2006); and the adaptive loss of pelvic armature in freshwater sticklebacks (Shapiro et al., 2004; Colosimo et al., 2004).).

The adaptive radiation of Darwin’s finches, with their different kinds of beaks suitable for different feeding adaptations, is one of the classic instances of evolutionary divergence due to natural selection (Lack, 1947). Abzhanov et al. (2004) have identified two key gene activities that are associated, respectively, with beak depth and width, on one hand and beak length on the other. The first characteristic, beak depth and width, is correlated with and evidently determined by an elevated level of activity of bone morphogenetic protein 4 (BMP4) during a critical phase of beak development. In contrast, finch beak length is evidently linked to a specific elevation of calmodulin activity during beak development (Abzhanov et al., 2006).).

The analysis of the genetic basis of bat wing evolution bears some similarity. A key component in the evolution of the distinctive wings of bats is the elevation in activity during a key phase of forelimb development in the embryo of another TGF-β activity, also a member of the BMP family, BMP2, which promotes the selective growth of the metacarpals to extend the key digits (Sears et al., 2006). Making a batwing, however, involves more than just exaggerated digit length; it also involves suppression of the waves of apoptosis that eliminate interdigital material in tetrapods with distinct digits. In the case of bats, the maintenance of

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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)