Variation, 1st Edition

A Central Concept in Biology

Variation, 1st Edition,Benedikt Hallgrímsson,Brian Hall,ISBN9780120887774

Hallgrímsson   &   Hall   

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This book expands upon Darwin's theory of natural selection, a central concept in evolutionary biology.

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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


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. This volume situates the role of variability within this broad framework, bringing variation back to the center of the evolutionary stage.


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

Benedikt Hallgrímsson

Affiliations and Expertise

University of Calgary, Alberta, Canada

Brian Hall

Affiliations and Expertise

Dalhousie University, Halifax, Nova Scotia, Canada

View additional works by Brian K. Hall

Variation, 1st Edition

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


Quotes and reviews

"...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
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