Variation - 1st Edition - ISBN: 9780120887774, 9780080454467


1st Edition

A Central Concept in Biology

Editors: Benedikt Hallgrímsson Brian Hall
eBook ISBN: 9780080454467
Hardcover ISBN: 9780120887774
Imprint: Academic Press
Published Date: 24th June 2005
Page Count: 592
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Darwin's theory of evolution by natural selection was based on the observation that there is variation between individuals within the same species. This fundamental observation is a central concept in evolutionary biology. However, variation is only rarely treated directly. It has remained peripheral to the study of mechanisms of evolutionary change. The explosion of knowledge in genetics, developmental biology, and the ongoing synthesis of evolutionary and developmental biology has made it possible for us to study the factors that limit, enhance, or structure variation at the level of an animals' physical appearance and behavior. Knowledge of the significance of variability is crucial to this emerging synthesis. Variation situates the role of variability within this broad framework, bringing variation back to the center of the evolutionary stage.

Key Features

  • Provides an overview of current thinking on variation in evolutionary biology, functional morphology, and evolutionary developmental biology
  • Written by a team of leading scholars specializing on the study of variation
  • Reviews of statistical analysis of variation by leading authorities
  • Key chapters focus on the role of the study of phenotypic variation for evolutionary, developmental, and post-genomic biology


Intended for scholars, advanced undergraduate students, and graduates in evolutionary biology, biological anthropology, paleontology, morphology, developmental biology, genomics and other related disciplines.

Table of Contents

Foreword - Ernst Mayr

CHAPTER 1. Variation and Viability: Central Concepts in Biology- Benedikt Hallgrímsson & Brian K. Hall

CHAPTER 2. Variation from Darwin to the Modern Synthesis, by Peter J. Bowler Introduction I. Variation before Darwin II. Darwin and Variation III. Alternative Theories of Variation and Evolution IV. Neo-Darwinism V. The Evolutionary Synthesis VI. Conclusions

CHAPTER 3. The Statistics of Variation, by Leigh Van Valen Abstract Introduction I. Absolute Variation: Univariate Case II. Absolute Variation: Multivariate Case III. Relative Variation: Univariate Case IV. Relative Variation: Multivariate Case V. Dimensionality of Variation VI. Tightness VII. Measurement Error and Single Specimens

CHAPTER 4. Landmark Morphometrics and the Analysis of Variation, by Joan T. Richtsmeier, Subhash R. Lele and Theodore M. Cole, III Introduction I. Coordinate Data and the Coordinate System II. The General Perturbation Model for Landmark Variation III. Proper Elimination of Nuisance Parameters using a Coordinate System Invariant Method of Estimation IV. Adding Assumptions to the Perturbation Model V. Conclusions

CHAPTER 5. Variation in Ontogeny, by D.C. Jones and R.Z. German Introduction I. Measuring Variation A. Data B. Levels of Variation in Data on Growth and Protein Malnutrition C. Measuring within Individual Variation D. Among Individual Variation
E. Variation Between Treatment Groups II. Results A. Factor Differences for Within Individual Variation B. Factor Differences for Among Individual Variation III. Discussion A. Within Individual Variation B. Between or among Individual Variation C. Variation across hierarchial levels IV. Conclusions

CHAPTER 6. Constraints on Variation from Genotype through Phenotype to Fitness, by Lauren Ancel Meyers Introduction I. RNA Evolutionary Model II. Evolving Constraints on Variation in RNA III. Mechanistic Constraints A. The spectrum of mutational constraints B. The Evolution of Mutational Constraints IV. Epistatic Constraints A. The spectrum of epistatic constraints B. The evolution of epistatic constraints V. Viability Constraints VI. Modularity: A Way out of the Constraints

CHAPTER 7. Developmental Origins of Variation, by Ellen W. Larsen Introduction I. Does Intrinsic Developmental Variation Exist? II. Intrinsic Variation in Different Environments III. Potential Origins of Intrinsic Developmental Variation A. Noise IV. An Example of Noise in Eukaryotic Transcription V. Noisy Bicoid Gene Expression in Fruit Flies VI. Noise in Asymmetry Production VII. Noisy Implication for Evolution VIII. Networks IX. Morphogenetic Fields a Potential Source of Variation X. Implications XI. Summary

CHAPTER 8. Canalization, Cryptic Variation and Developmental Buffering: A Critical Examination and Analytical Perspective, by Ian Dworkin Introduction I. A Review of the Reviews II. Empirical Concerns for the Study of Canalization A. The amount of genetic variation must be controlled between lines/populations B. The need for multiple, independent samples C. Genetic background must be controlled for comparisons between treatments III. Definitions of Canalization IV. Reaction Norm of the Mean (RXNM) Definition of Canalization V. The Variation Approach to Canalization VI. Partitioning Sources of Variation VII. Inferring Canalization: When is a trait Canalized? VIII. What are the appropriate tests for making statistical inferences about Canalization? IX. In the Interim… X. Analysis for the RXNM Approach XI. The Analysis of Cryptic Genetic Variation XII. Mapping Cryptic Genetic Variants XIII. Is the Genetic Architecture of Cryptic Genetic Variation different from that of other Genetic Variation involved with Trait Expression? XIV. Now that I have all of this Cryptic Genetic Variation, what do I do with it? XV. The future for studies of Canalization

CHAPTER 9. Mutation and Phenotypic Variation: Where is the connection Capacitators, Stressors, Phenotypic Variability and Evolutionary Change, by Ary A. Hoffmann and John A. McKenzie Abstract Introduction: Variability and Limits I. Mutators, Recombinators, Stressors and Genetic Variability II. Recombination III. The Impact of New Mutants and Recombinants – Canalization and Capacitators IV. In Search of Capacitators: Genes that influence Developmental Stability and Canalization V. Capacitators, Stressors, and Quantitative Variation VI. Do we need Cariability Generators? VII. Concluding Remarks: Experimental Programs for Defining the Role of Variability Generators

CHAPTER 10. Within Individual Variation: Developmental Noise versus Developmental Stability, by Katherine E. Willmore and Benedikt Hallgrímsson Introduction I. Causes of Developmental Noise A. Causes of Developmental Noise at the Molecular Level B. Causes of Developmental Noise at the Developmental Systems Level C. Causes of Developmental Noise at the Organismal Level II. Mechanisms of Developmental Stability A. Mechanisms of Developmental Stability at the Molecular Level B. Mechanisms of Developmental Stability at the Developmental Systems Level C. Mechanisms of Developmental Stability at the Organismal Level III. Implications

CHAPTER 11. Developmental Constraints, Modules and Evolvability, by Christian Peter Klingenberg Abstract Introduction I. Evolvability and Constraints II. Integration and Modularity III. Developmental Origins of Covariation among Traits IV. Developmental Interactions and Pleiotropy V. Evolution of Pleiotropy and Developmental Interactions VI. Modularity of Pleiotropic Effects: Inherent in Developmental Systems or Evolved Property? VII. From Pleiotropic Gene Effects to G Matrices VIII. G Matrices, Constraints, and Evolutionary Dynamics IX. Perspective: Developmental Processes and Evolutionary Constraints

CHAPTER 12. Developmental Regulation of Variability, by Miriam Zelditch Introduction I. Empirical Patterns II. The Ontogeny of Variation in Male Norway Rat Cranial Shape III. Biological Patterns Versus Artifacts A. Morphological Sampling B. Life-History/Developmental Rate IV. Mechanisms Generating and Regulating Craniofacial Shape Variance V. Targeted Growth VI. Organismal Developmental Timing VII. Variation in Relative Developmental Timing of Modules VIII. Neural Regulation of Musculoskeletal Interactions IX. Canalized Shape as an Epiphenomenon

CHAPTER 13. Role of Stress in Evolution: From Individual Adaptability to Evolutionary Adaptation, by Alexander Badyaev Introduction I. Evolution of Response to Stress A. Detection and Avoidance II. Evolutionary Consequences of Stress A. Stress-induced Variation III. Buffering, Accommodating, and Directing Stress-Induced Variation IV. Inheritance V. Evolutionary Adaptation VI. Conclusions

CHAPTER 14. Environmentally Contingent Variation: Phenotypic Plasticity and Norms of Reaction, by Sonia Sultan and Steve Stearns Introduction I. Plasticity Concepts II. Specific Types of Plasticity III. Reaction Norms IV. Parental Effect Reaction Norms (Cross-Generational Plasticity) V. Imprinted Reaction Norms VI. Iterated Reaction Norms VII. Dynamic Reaction Norms VIII. Photomorphogenetic Plasticity in Plants IX. Adaptive Plasticity for Timing of Amphibian Metamorphosis X. Mediation of Phenotypic Expression XI. Genetic Variation and the Evolution of Plasticity A. How Plasticity interacts with conserved developmental patterns XII. Genetic Causation and the Butterfly Wing: A More complicated picture XIII. The Same Networks may give rise to both Plasticity and Constraint A. What effects does plasticity have on populations and communities? B. Research Agenda

CHAPTER 15. Variation and Life History Evolution, by Derek A. Roff Introduction I. Phenotypic Variation in a Constant Environment A. Heterozygous advantage B. Antagonistic pleiotropy C. Frequency-dependent selection II. Phenotypic Variation in a Stochastic Environment A. Temporal variation B. Spatial variation C. Spatial and temporal variation III. Predictable Environments A. Temporal variation B. Spatial variation IV. Concluding Comments

CHAPTER 16. Antisymmetry, by A. Richard Palmer Introduction I. Asymmetry Terminology A. Terms for Subtle Asymmetries B. Terms for Conspicuous Asymmetry in an Individual C. Terms for the Orientation of Bilateral or Spiral Asymmetries D. Terms for Conspicuous Asymmetries in a Population or Species II. The History of Antisymmetry III. Taxomonic Distribution and Functional Significance of Antisymmetry A. Plants B. Cnidaria C. Mollusca D. Annelida E. Arthropoda-Chelicerata F. Arthropoda-Crustacea G. Arthropoda-Insecta H. Brachiopods I. Bryozoa J. Echinodermata K. Chordata IV. Development and Regeneration of Asymmetry in Antisymmetric Species A. Ontogeny V. Regeneration of Missing Antimeres VI. Inheritance of Direction in Antisymmetric Species VII. Inheritance of Direction in Directionally Asymmetric Species VIII. Evolutionary Significance of Antisymmetry IX. What Next?

CHAPTER 17. Variation in Structure and its Relationship to Function: Correlation, Explanation and Extrapolation, by Anthony P. Russell and Adam M. Bauer Abstract Introduction I. Background II. Approaches to the Study of Structural Variation III. Variation as an Observable Phenomenon A. Variation and Taxonomic Utility B. Variation Associated with Developmental Plasticity C. Geographically-based Variation IV. In Situ Correlational Studies of the Relationship between Structural Variation and Functional Attributes A. Trophic Polymorphism and Environmental Fluctuation B. Clinical Variation C. Microgeographic Variation V. Ex Situ Studies of the Relationship between Structural Variation and Performance A. Variation in Trophic Performance B. Locomotor Performance C. Fluctuating Asymmetry and Variation in Performance D. Selection Experiments and the Investigation of the Limits of Variability E. Other Measures of Structural and Functional Variation V. Concluding Remarks

CHAPTER 18. A Universal Generative Tendency Toward Increased Organismal Complexity, by D. McShea Introduction I. Internal Variance as Complexity II. Three Simple Models III. The Effect of Increased Dimensionality IV. Apparent Difficulties V. Is there an Upward Bias in Real Lineages? VI. If so, the Principle is Supported VII. If not, Why not? VIII. Testing the Principle IX. A Reversal of Intuition

CHAPTER 19. Variation and Versatility in Macroevolution, by V. Louise Roth I. Principles A. To Vary is Easy B. Evolvability and Versatility II. Examples A. Elephantid Teeth B. Disparity and Versatility in Sciurdae III. Overview and Conclusion

CHAPTER 20. Variation and Developmental Biology: Prospects for the Future, by David M. Parichy Introduction I. Model Organisms: Expanding the Fold II. Ecologically Significant Differences in Form Between Species III. How many ways to make a Phenotype: Developmental Variation and Morphological Similarity IV. Intraspecific Developmental Variation: Canalization, and Developmental Plasticity V. Conclusions

CHAPTER 21. Phenogenetics: Genotypes, Phenotypes, and Variation, by Samuel Sholtis and Kenneth Weiss Introduction I. Mechanism versus Variation II. From Genotype to Phenotype: Mechanism A. A quick digression concerning DNA sequence: arbitrary and saturated B. Pre-transcriptional mechanisms C. Post-transcriptional mechanisms III. From Genotype to Phenotype: Variation A. A lexicographer’s nightmare: Canalization, Robustness, Plasticity, Polyphenism… B. Developmental process: patterning repeated traits C. Gene regulation and the evolution of phenotypes D. Phenogenetic drift: the role of chance in the evolution of genotype-phenotype relationships IV. Summary

CHAPTER 22. The Study of Phenotypic Variability, by Benedikt Hallgrímsson and Brian K. Hall Introduction I. Variability and the Biological Hierarchy II. Components of Variability III. Current Approaches to Understanding the Development-Genetic Architecture of Variability A. Pattern Based Approaches B. Perturbation Based Approaches C. Model Driven Approaches IV. A Developmental Systems Approach to Phenotypic Variability A. The Regulation of Form in the Mouse Mandible B. The Regulation of Outgrowth of the Maxillary Process V. Conclusion



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© Academic Press 2005
Academic Press
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About the Editor

Benedikt Hallgrímsson

Affiliations and Expertise

University of Calgary, Alberta, Canada

Brian Hall

Affiliations and Expertise

Dalhousie University, Halifax, Nova Scotia, Canada


"...Variation: A Central Concept in Biology, is sure to spark the interest of nearly all ecologists and evolutionary biologists...How does this variation arise? How do new variants evolve? What contrains variation? The answers are incomplete. However, the chapters of this book provide a glimpse at our current understanding of phenotypic variation." --James A. Fordyce, University of Tenessee, Department of Ecology and Evolutionary Biology, in ECOLOGY

"Variation is certainly a topic of central interest in evolutionary biology and this new book offers an unusually inclusive array of perspectives on the topic. I especially enjoyed the breadth of coverage. Novel features are found that one might not have expected in a book of this nature, such as structural, functional and developmental variation, as contrasted with the expected emphases on genetic variation, canalization and phenotypic plasticity, and their relation to life history evolution. Palmer’s stimulating chapter on antisymmetry is particularly noteworthy for its originality. Another unusual treatment is Badyaev’s focus on the role of stress in evolution, which is examined from a different perspective by Hoffmann and McKenzie. On the whole this is an assemblage of excellent chapters by many of the central figures in the fields covered and will be a welcome addition to my library." --David B. Wake, University of California, Berkeley, U.S.A.

"This comprehensive, diverse and stimulating volume is a must-read for anyone interested in development and evolution. Never has the critical subject of variation been so well treated in terms of how to analyze variation, how developmental processes induce and constrain its properties, and the complex relationships between genotypic, environmental and phenotypic variation. This is a tour-de-force treatment of a critical subject." --Daniel E. Lieberman, Harvard University, U.S.A.

"Where do genetic and phenotypic diversity come from and how are they are maintained? Do the same processes link the differences within species to the stable differences between species? These are the big questions about the diversity of life on Earth and Hallgrimsson and Hall’s book provides the latest views from leading scientists of diversity and form." --Mark Pagel, University of Reading, England

“Variation is the basic material of evolution. At last we have a book that takes a modern approach to variation in all its forms—populational, developmental, genetic, morphological—and links it to the processes that generate the variation itself. This is not a collection of bland reviews, but a vital compendium of novel perspectives that begin to meld the new data of evo-devo with the accepted body of Modern Synthesis work. A must for any evolutionist’s library.” --Kevin Padian, University of California, Berkeley, U.S.A. Listed in NEW TITLES in BIOSCIENCE (April 2006)

"...chapters are authoritative and well written, and graduate students and scientists will find much here that is thought-provoking." --Carl D. Schlichting, Professor, Department of Ecology and Evolutionary Biology, University of Connecticut, in BIOSCIENCE