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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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. "2 Functional Information and the Emergence of Biocomplexity--ROBERT M. HAZEN, PATRICK L. GRIFFIN, JAMES M. CAROTHERS, and JACK W. SZOSTAK." 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 special interest in this regard are biological systems that display complex emergent behavior, for example, through long-range chemical signaling among a collection of cells in social amoebas (Goldbeter, 1996; Brännström and Dieckmann, 2005; Schaap et al., 2006), cooperation among consortia of host organisms and symbionts (Moran, 2007, Chapter 9, this volume), or colonies of social insects (Solé and Goodwin, 2000; Camazine et al., 2001; Strassmann and Queller, 2007, Chapter 8, this volume). We propose that functional information can be applied, at least in principle, to any such emergent system that has the ability to perform a function.

Many emergent systems can be analyzed in terms of their ability to dissipate energy or maximize entropy production (Nicolis and Progogine, 1977; Lorenz, 2003; Whitfield, 2005; Emanuel, 2006). For example, consider the functional information of an assemblage of sand grains subjected to a steady flow of wind or water (e.g., Bagnold, 1988; Hansen et al., 2001). The formation of periodic sand dunes or ripples serves to initiate turbulent flow and thus increase energy dissipation. Functional information of the system can thus be measured as the fraction of all possible sand configurations, F(Ex), that achieve at least the corresponding energy dissipation, Ex. Such a problem might be analyzed with Monte Carlo simulations of numerous gravitationally stable sand configurations. The analytical challenge remains to determine the degree of function of a statistically significant random fraction of all possible configurations of the system so that the relationship between I(Ex) and Ex can be deduced.

CONCLUSIONS

A complexity metric is of little utility unless its conceptual framework and predictive power result in a deeper understanding of the behavior of complex systems. Analysis of complex systems in terms of functional information reveals several characteristics that are important in understanding the behavior of systems composed of many interacting agents. Letter sequences, Avida genomes and biopolymers all display degrees of functions that are not attainable with individual agents (a single letter, machine instruction, or RNA nucleotide, respectively). In all three cases, highly functional configurations comprise only a small fraction of all possible sequences. Furthermore, these three examples reveal that several discrete classes of functional configurations exist, a situation that can lead to distinctive step features in plots of information versus function.

The functional information formalism may also point to key factors in the origin and emergence of biocomplexity. In particular, functional information quantifies the probability that, for a particular system, a configuration with a specified degree of function will emerge. Furthermore, analysis of the relationship between information and function may reveal

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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)