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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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. "3 The Theory of Facilitated Variation--JOHN GERHART and MARC KIRSCHNER." 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

of conserved components and processes to give phenotypic variations and selected traits. Regulatory change acts on the repertoire of unchanging core processes to select subsets, which are then externally selected upon. The burden of creativity in evolution, down to minute details, does not rest on selection alone. Through its ancient repertoire of core processes, the current phenotype of the animal determines the kind, amount, and viability of phenotypic variation the animal can produce in response to regulatory change. Thanks to the nature of the processes, the range of possible anatomical and physiological variations is enormous, and many are likely nonlethal, in part simply because the processes have been providing “useful” function since pre-Cambrian times. Phenotypic plasticities, both those evokable by environmental change and those developmental adaptabilities not evocable, are rich sources and favored paths of variation requiring little regulatory change.

These views are not at all Lamarckian, nor does facilitated phenotypic variation require selection for future good. Such facilitation arose, we think, as a by-product of the evolution of the special properties of the core processes, namely, of their robustness, adaptability, modularity, exploratory behavior, and capacity for weak regulatory linkage. These properties were probably selected at the level of the individual, simply for their capacity to make core processes work effectively under fluctuating external and internal conditions (Gerhart and Kirschner, 1997; Kirschner and Gerhart, 2005). In this way, the same molecular features that facilitate physiological and developmental change in an organism’s lifetime also facilitate evolutionary change in the long run, as regulatory changes become genetically fixed.

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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)