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Foreword | |
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Contributors | |
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Variation and Variability: Central Concepts in Biology | |
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Variation from Darwin to the Modern Synthesis | |
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Introduction | |
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Variation before Darwin | |
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Darwin and Variation | |
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Alternative Theories of Variation and Evolution | |
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Neo-Darwinism | |
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The Evolutionary Synthesis | |
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Conclusions | |
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References | |
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The Statistics of Variation | |
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Abstract | |
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Introduction | |
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Absolute Variation: Univariate Case | |
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Levene's Test | |
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Smith's Test | |
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Jackknifing | |
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Absolute Variation: Multivariate Case | |
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Relative Variation: Univariate Case | |
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Relative Variation: Multivariate Case | |
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Dimensionality of Variation | |
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Tightness | |
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Measurement Error and Single Specimens | |
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References | |
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Landmark Morphometrics and the Analysis of Variation | |
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Introduction | |
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Coordinate Data and the Coordinate System | |
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The General Perturbation Model for Landmark Variation | |
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Proper Elimination of Nuisance Parameters Using a Coordinate System Invariant Method of Estimation | |
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Adding Assumptions to the Perturbation Model | |
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Model 0: Isotropic Error Model | |
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Model 1: [Sigma subscript K]=Independent Local Variation | |
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Model 2: [Sigma subscript K]=Independent Modules | |
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Model 3: [Sigma subscript K]=Co-dependent Modules | |
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Conclusions | |
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Acknowledgments | |
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References | |
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Variation in Ontogeny | |
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Introduction | |
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Measuring Variation: A Case Study | |
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Data Analyzed | |
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Levels of Variation in Data on Growth and Protein Malnutrition | |
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Measuring within Individual Variation | |
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Measuring among Individual Variation | |
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Testing for Variation between Treatment Groups | |
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Factor Differences for within Individual Variation | |
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Factor Differences for among Individual Variation | |
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Implications for Studies of Variation | |
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Utility of Standard Laboratory Animal Models | |
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Within Individual Variation | |
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Between or among Individual Variation | |
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Variation across Hierarchical Levels | |
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Conclusions | |
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Acknowledgments | |
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References | |
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Constraints on Variation from Genotype through Phenotype to Fitness | |
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Introduction | |
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RNA Evolutionary Model | |
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Evolving Constraints on Variation in RNA | |
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Mechanistic Constraints | |
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The Spectrum of Mutational Constraints | |
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The Evolution of Mutational Constraints | |
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Epistatic Constraints | |
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The Spectrum of Epistatic Constraints | |
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The Evolution of Epistatic Constraints | |
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Viability Constraints | |
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Modularity: A Way Out of the Constraints | |
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Acknowledgments | |
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References | |
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Developmental Origins of Variation | |
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Introduction | |
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Does Intrinsic Developmental Variation Exist? | |
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Intrinsic Variation in Different Environments | |
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Potential Origins of Intrinsic Developmental Variation | |
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Noise | |
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An Example of Noise in Eukaryotic Transcription | |
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Noisy bicoid Gene Expression in Fruit Flies | |
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Noise in Asymmetry Production | |
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Noisy Implication for Evolution | |
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Networks | |
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Morphogenetic Fields: A Potential Source of Variation | |
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Implications | |
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Summary | |
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Acknowledgments | |
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References | |
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Canalization, Cryptic Variation, and Developmental Buffering: A Critical Examination and Analytical Perspective | |
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Introduction | |
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A Review of the Reviews | |
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Empirical Concerns for the Study of Canalization | |
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The Amount of Genetic Variation must be Controlled between Lines/Populations | |
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The Need for Multiple, Independent Samples (Across Genotypes, Not Individuals) | |
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Genetic Background must be Controlled for Comparisons between Treatments | |
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Definitions of Canalization | |
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Reaction Norm of the Mean (RxNM) Definition of Canalization | |
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The Variation Approach to Canalization | |
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Partitioning Sources of Variation | |
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Variation within Individual (V[subscript WI]) | |
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Variation between Individuals, within Genotype (V[subscript BI]) | |
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Between-Line (Genetic) Variation (V[subscript G]) | |
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Inferring Canalization: When is a Trait Canalized? | |
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What are the Appropriate Tests for Making Statistical Inferences about Canalization? | |
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In the Interim... | |
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Analysis for the RxNM Approach | |
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The Analysis of Cryptic Genetic Variation | |
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Mapping Cryptic Genetic Variants | |
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Is the Genetic Architecture of Cryptic Genetic Variation Different from that of Other Genetic Variation Involved with Trait Expression? | |
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Now that I have All of this Cryptic Genetic Variation, What do I do with It? | |
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The Future for Studies of Canalization | |
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Acknowledgments | |
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References | |
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Mutation and Phenotypic Variation: Where is the Connection? Capacitators, Stressors, Phenotypic Variability, and Evolutionary Change | |
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Abstract | |
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Introduction: Variability and Limits | |
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Mutators, Recombinators, Stressors, and Genetic Variability | |
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Mutation | |
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Recombination | |
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The Impact of New Mutants and Recombinants: Canalization and Capacitators | |
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In Search of Capacitators: Genes that Influence Developmental Stability and Canalization | |
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Capacitators, Stressors, and Quantitative Variation | |
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Do We Need Variability Generators? | |
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Concluding Remarks: Experimental Programs for Defining the Role of Variability Generators | |
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References | |
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Within Individual Variation: Developmental Noise versus Developmental Stability | |
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Introduction | |
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Causes of Developmental Noise | |
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Causes of Developmental Noise at the Molecular Level | |
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Causes of Developmental Noise at the Developmental Systems Level | |
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Causes of Developmental Noise at the Organismal Level | |
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Mechanisms of Developmental Stability | |
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Mechanisms for Developmental Stability at the Molecular Level | |
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Mechanisms for Developmental Stability at the Developmental Systems Level | |
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Mechanisms for Developmental Stability at the Organismal Level | |
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Implications | |
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References | |
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Developmental Constraints, Modules, and Evolvability | |
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Abstract | |
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Introduction | |
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Evolvability and Constraints | |
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Integration and Modularity | |
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Developmental Origins of Covariation among Traits | |
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Developmental Interactions and Pleiotropy | |
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Evolution of Pleiotropy and Developmental Interactions | |
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Modularity of Pleiotropic Effects: Inherent in Developmental Systems or Evolved Property? | |
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From Pleiotropic Gene Effects to G Matrices | |
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G Matrices, Constraints, and Evolutionary Dynamics | |
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Perspective: Developmental Processes and Evolutionary Constraints | |
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Acknowledgments | |
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References | |
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Developmental Regulation of Variability | |
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Introduction | |
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Empirical Patterns | |
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The Ontogeny of Variation in Male Norway Rat Cranial Shape | |
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Biological Patterns versus Artifacts | |
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Morphological Sampling | |
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Life History/Developmental Rate | |
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Mechanisms Generating and Regulating Craniofacial Shape Variance | |
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Targeted Growth | |
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Organismal Developmental Timing | |
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Variation in Relative Developmental Timing of Modules | |
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Neural Regulation of Musculoskeletal Interactions | |
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Canalized Shape as an Epiphenomenon | |
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Acknowledgments | |
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References | |
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Role of Stress in Evolution: From Individual Adaptability to Evolutionary Adaptation | |
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Introduction | |
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Evolution of Response to Stress | |
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Detection and Avoidance | |
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Stress-Avoidance Strategies | |
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Evolutionary Consequences of Stress | |
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Stress-Induced Variation | |
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Buffering, Accommodating, and Directing Stress-Induced Variation | |
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Inheritance | |
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Evolutionary Adaptation | |
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Stress-Induced Evolution versus Stress-Induced Stasis | |
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Conclusions | |
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Acknowledgments | |
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References | |
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Environmentally Contingent Variation: Phenotypic Plasticity and Norms of Reaction | |
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Introduction | |
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Plasticity Concepts | |
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Specific Types of Plasticity | |
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Reaction Norms | |
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Parental Effect Reaction Norms (Cross-Generational Plasticity) | |
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Imprinted Reaction Norms | |
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Iterated Reaction Norms | |
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Dynamic Reaction Norms | |
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The Genetic and Developmental Basis of Phenotypic Plasticity | |
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Photomorphogenetic Plasticity in Plants | |
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Adaptive Plasticity for Timing of Amphibian Metamorphosis | |
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Mediation of Phenotypic Expression | |
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Genetic Variation and the Evolution of Plasticity | |
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How Plasticity Interacts with Conserved Developmental Patterns | |
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Genetic Causation and the Butterfly Wing: A More Complicated Picture | |
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The Same Networks May Give Rise to Both Plasticity and Constraint | |
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What Effects Does Plasticity Have on Populations and Communities? | |
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Research Agenda | |
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Acknowledgments | |
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References | |
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Variation and Life-History Evolution | |
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Introduction | |
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Phenotypic Variation in a Constant Environment | |
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Mutation-Selection Balance | |
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Heterozygous Advantage | |
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Antagonistic Pleiotropy | |
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Frequency-Dependent Selection | |
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Phenotypic Variation in a Stochastic Environment | |
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Temporal Variation | |
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Spatial Variation | |
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Spatial and Temporal Variation | |
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Predictable Environments | |
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Temporal Variation | |
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Spatial Variation | |
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Concluding Comments | |
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References | |
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Antisymmetry | |
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Introduction | |
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Asymmetry Terminology | |
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Terms for Subtle Asymmetries | |
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Terms for Conspicuous Asymmetry in an Individual | |
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Terms for the Orientation of Bilateral or Spiral Asymmetries | |
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Terms for Conspicuous Asymmetries in a Population or Species | |
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The History of Antisymmetry | |
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Taxomonic Distribution and Functional Significance of Antisymmetry | |
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Plants | |
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Cnidaria | |
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Mollusca | |
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Annelida | |
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Arthropoda: Chelicerata | |
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Arthropoda: Crustacea | |
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Arthropoda: Insecta | |
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Brachiopoda | |
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Bryozoa | |
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Echinodermata | |
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Chordata | |
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Development and Regeneration of Asymmetry in Antisymmetric Species | |
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Ontogeny | |
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Regeneration of Missing Antimeres | |
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Inheritance of Direction in Antisymmetric Species | |
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Inheritance of Direction in Directionally Asymmetric Species | |
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Evolutionary Significance of Antisymmetry | |
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What Next? | |
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Acknowledgments | |
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Appendix | |
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References | |
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Variation in Structure and Its Relationship to Function: Correlation, Explanation, and Extrapolation | |
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Abstract | |
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Introduction | |
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Background | |
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Approaches to the Study of Structural Variation | |
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Variation as an Observable Phenomenon | |
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Variation and Taxonomic Utility | |
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Variation Associated with Developmental Plasticity | |
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Geographically Based Variation | |
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In Situ Correlational Studies of the Relationship between Structural Variation and Functional Attributes | |
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Trophic Polymorphism and Environmental Fluctuation | |
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Clinal Variation | |
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Microgeographic Variation | |
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Ex Situ Studies of the Relationship between Structural Variation and Performance | |
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Variation in Trophic Performance | |
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Locomotor Performance | |
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Fluctuating Asymmetry and Variation in Performance | |
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Selection Experiments and the Investigation of the Limits of Variability | |
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Other Measures of Structural and Functional Variation | |
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Concluding Remarks | |
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References | |
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A Universal Generative Tendency toward Increased Organismal Complexity | |
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Introduction | |
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Internal Variance as Complexity | |
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Three Simple Models | |
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Model 1 | |
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Model 2 | |
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Model 3 | |
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The Effect of Increased Dimensionality | |
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Apparent Difficulties | |
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Is There an Upward Bias in Real Lineages? | |
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If So, the Principle Is Supported | |
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If Not, Why Not? | |
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Testing the Principle | |
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A Reversal of Intuition | |
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References | |
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Variation and Versatility in Macroevolution | |
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Principles | |
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To Vary is Easy | |
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Evolvability and Versatility | |
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Examples | |
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Elephantid Teeth | |
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Disparity and Versatility in Sciuridae | |
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Overview and Conclusion | |
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Acknowledgments | |
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References | |
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Variation and Developmental Biology: Prospects for the Future | |
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Introduction | |
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Model Organisms: Expanding the Fold | |
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Ecologically Significant Differences in Form between Species | |
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How Many Ways to Make a Phenotype: Developmental Variation and Morphological Similarity | |
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Intraspecific Developmental Variation: Canalization and Developmental Plasticity | |
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Conclusions | |
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References | |
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Phenogenetics: Genotypes, Phenotypes, and Variation | |
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Introduction | |
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Mechanism versus Variation | |
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From Genotype to Phenotype: Mechanism | |
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A Quick Digression Concerning DNA Sequence: Arbitrary and Saturated | |
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Pretranscriptional Mechanisms | |
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Posttranscriptional Mechanisms | |
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From Genotype to Phenotype: Variation | |
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A Lexicographer's Nightmare: Canalization, Robustness, Plasticity, Polyphenism... | |
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Developmental Process: Patterning Repeated Traits | |
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Gene Regulation and the Evolution of Phenotypes | |
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Phenogenetic Drift: The Role of Chance in the Evolution of Genotype-Phenotype Relationships | |
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Summary | |
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Acknowledgments | |
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References | |
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The Study of Phenotypic Variability: An Emerging Research Agenda for Understanding the Developmental-Genetic Architecture Underlying Phenotypic Variation | |
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Introduction | |
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Variability and the Biological Hierarchy | |
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Components of Variability | |
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Current Approaches to Understanding the Development-Genetic Architecture of Variability | |
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Pattern-Based Approaches | |
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Perturbation-Based Approaches | |
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Model-Driven Approaches | |
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A Developmental Systems Approach to Phenotypic Variability | |
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The Regulation of Form in the Mouse Mandible | |
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The Regulation of Outgrowth of the Facial Prominences in Mice | |
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Conclusion | |
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References | |
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Index | |