Brakefield, P.M.

Leiden University Scholary Publications

Phenotypic PlasticityBeyond Nature and NurtureMASSIMO PIGLIUCCITHE JOHNS HOPKINS UNIVERSITY PRESSBALTIMORE AND LONDONContentsSeries Editor's Foreword, by Samuel M. Scheiner ixForeword, by Paul Brakefield xiPreface xiii1 What Is Phenotypic Plasticity? 12 Studying and Understanding Plasticity 293 A Brief (Conceptual) History of Phenotypic Plasticity 474 The Genetics of Phenotypic Plasticity 675 The Molecular Biology of Phenotypic Plasticity 1096 The Developmental Biology of Phenotypic Plasticity 1337 The Ecology of Phenotypic Plasticity 1568 Behavior and Phenotypic Plasticity 1829 Evolution of and by Phenotypic Plasticity 19710 The Theoretical Biology of Phenotypic Plasticity 21611 Phenotypic Plasticity as a Central Conceptin Evolutionary Biology 237Epilogue: Beyond Nature and Nurture 253Notes 263References 267Author Index 305Subject Index 315VIIForewordResearch on phenotypic plasticity has been at the forefront of evolutionarythinking for the last two decades or so. Diversity generated by the ability ofany particular genotype to adjust its development and, therefore, phenotypein response to the environment can be dramatic. However, few books onthis subject have appeared, which makes this offering a most timely one.Without doubt this book demonstrates why phenotypic plasticity has be-come so fascinating to many biologists, especially to those who share a benttoward multidisciplinary approaches. Like-minded researchers are most ex-cited by those topics that reflect intersecting areas of biology. In my opin-ion these are precisely where the opportunities to achieve the most innova-tive and important insights lie. Phenotypic plasticity must be one of themore striking phenomena in need of integration across different areas ofbiology. As such it represents both a formidable challenge and an excit-ing opportunity for research. To study plasticity effectively one needs to bea biologist of many colors: a geneticist, a physiologist, a developmental bi-ologist, an ecologist, and an evolutionary biologist. I think the challengingnature of the topic is also evident in a healthy level of controversy, espe-cially with regard to the nature of genes for plasticity. This book rises ad-mirably to the challenge.Historically, attempts to integrate ecology and genetics have sought tostudy the genetic basis of phenotypic variation in natural populations andthe ways in which such variability is influenced by natural selection. It can,however, be very difficult to identify the specific targets of selection. Someexamples of plasticity can prove excellent tools for exploring natural se-lection in the wild and addressing the question of to what extent plasticityis the result of adaptive evolution. The other side of the coin—that plastic-ity can reflect constraints on the ability of organisms to develop optimalphenotypes across environments—is also receiving avid attention.XIXÜ FOREWORDThe ways in which genes map onto phenotypes in natural populationshave represented something of a black box in the history of evolutionaryecology. Once again, phenotypic plasticity can provide wonderful materialfor opening this box. How are developmental pathways within an individ-ual organism modulated to produce the alternative phenotypes? The pres-ent opportunities for identifying genes to match up with the developmentaland physiological mechanisms are endless. To do this in the context of phe-notypes of evolutionary significance in natural populations is an excitinggoal indeed. Functional genomics will only provide yet more tools and tech-nologies for further exploration of the developmental and physiologicalmechanisms underlying the ability to express phenotypic plasticity.The environment of an individual of a given genotype not only is crit-ical in shaping how it is influenced by natural selection but also may pro-foundly affect the phenotype it will produce during development. Again thedifferent levels of interaction between genotype, environment, and pheno-type are at the heart of thinking about the evolution of phenotypic plastic-ity, and such complexity is part of its challenge. Those interactions betweengenotype and environment that are the basis for the evolution of plasticityshould provide a cautionary tale to those applying functional genomics:Gene expression may be highly dependent on both external and internalenvironment.At the same time that phenotypic plasticity presents such opportunitiesto the empiricist to explore fundamental issues in biology, the field as awhole has also benefited greatly from contributions from theoretical biol-ogy. These have provided both extremely useful frameworks for the em-piricists as well as unique insights into how organisms can adapt to spatialor temporal heterogeneity in their environments.All these issues and many more are explored in this timely and wide-ranging book. It shows what has been achieved but also sets the scene forthe many exciting discoveries that the burgeoning molecular world will in-creasingly open up. Fortunately, the topic of phenotypic plasticity will con-tinue to draw together biologists working with organisms in nature, in thelaboratory, and in the test tube.Paul BrakefieldLeiden University, The Netherlands

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