Abstract
Variability contrasts with variation in that variability describes the potential for variation, not simply the expressed variation. The power of studying variability lies in creating a conceptual framework around which the relationship between the genotype and phenotype can be understood. Here, we attempt to demonstrate the importance of phenotypic variability, how it structures variation, and how fundamental developmental processes structure variability. Given the broad scope of this topic, we focus on three widely studied properties of variability: canalization, developmental stability and morphological integration. We have organized the paper to emphasize the importance of differentiating between the theory surrounding these components of phenotypic variability, their measurement and the biological factors surrounding their expression. First, we define these properties of variability, how they relate to each other and to variability as a whole. Second, we summarize the common methods of measurement for canalization, developmental stability and morphological integration and the reasoning behind these methods. Finally, we focus on jaw development as an example of how the basic processes of development affect variability and the resultant variation, with emphasis on how processes at all levels of the organismal hierarchy interact with one another and contribute to phenotypic variability.
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Acknowledgements
We thank Miriam Zelditch, David Polly and an anonymous reviewer for their insightful comments on a previous draft of this paper. We also thank Benedikt Hallgrímsson and the members of the Richtsmeier laboratory for their willingness to discuss these issues at great length and for their many helpful suggestions. This work was supported by the National Science Foundation grant BCS-0523637, as well as the William H. Davies Student Fellowship, Medical Science Graduate Research Fellowship, University of Calgary Graduate Studies (to KW).
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Glossary
- Buffering
-
any mechanism that limits the phenotypic effects of a perturbation whether it is adaptive or not.
- Canalization
-
refers to the property of an organism that ensures similarity of phenotypic expression by buffering development against both environmental and genetic perturbations.
- Developmental instability
-
the inverse of developmental stability.
- Developmental noise
-
minor environmentally induced developmental accidents.
- Developmental stability
-
the property of an organism that buffers variation of micro-environmental origin. That is, developmental stability ensures consistent phenotypic expression within individuals, or within a given genotype and environment.
- Environmental canalization
-
refers to the limitation of phenotypic effect caused by exposure to environmental change.
- Epigenetics
-
heritable changes in gene expression that are not due to changes in DNA sequence.
- Fluctuating asymmetry (FA)
-
random, non-directional deviations from symmetry of bilaterally symmetric traits such as limbs, wings or antennae. FA measures the balance between developmental noise and developmental stability.
- Genetic architecture
-
refers to the number of genes and the interactions between them whose products govern a particular trait.
- Genetic canalization
-
refers to the robustness of the phenotype to the effects of mutation.
- Module
-
a character, or a set of characters that are more tightly integrated internally than they are with other characters.
- Morphological integration
-
refers to the inter-relatedness between morphological structures often due to common developmental origins or functional demands.
- Norm of reaction
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the range of expected phenotypes under a given set of environmental and genetic conditions.
- Perturbation
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used here as any disruption to development. Could be genetic, developmental, environmental, or adverse interactions between these factors.
- Phenotypic plasticity
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phenotypic variation induced by the environement.
- Pleiotropy
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the control of one gene on multiple traits.
- Stabilizing selection
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most common form of natural selection, selects for the mean phenotype.
- Variability
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a dispositional term like solubility describing the tendency or propensity to vary.
- Variation
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observed phenotypic differences.
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Willmore, K.E., Young, N.M. & Richtsmeier, J.T. Phenotypic Variability: Its Components, Measurement and Underlying Developmental Processes. Evol Biol 34, 99–120 (2007). https://doi.org/10.1007/s11692-007-9008-1
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DOI: https://doi.org/10.1007/s11692-007-9008-1