A Central Concept in Biology

1st Edition - June 24, 2005

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  • Editors: Benedikt Hallgrímsson, Brian Hall
  • eBook ISBN: 9780080454467
  • Hardcover ISBN: 9780120887774

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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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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
    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


Product details

  • No. of pages: 592
  • Language: English
  • Copyright: © Academic Press 2005
  • Published: June 24, 2005
  • Imprint: Academic Press
  • eBook ISBN: 9780080454467
  • Hardcover ISBN: 9780120887774

About the Editors

Benedikt Hallgrímsson

Affiliations and Expertise

University of Calgary, Alberta, Canada

Brian Hall

Affiliations and Expertise

Dalhousie University, Halifax, Nova Scotia, Canada

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