Variational properties

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In evolutionary biology, the variational properties of an organism are those properties relating to the production of variation among its offspring. In a broader sense variational properties include phenotypic plasticity. [1] [2] [3] Variational properties contrast with functional properties. While the functional properties of an organism determine is level of adaptedness to its environment, it is the variational properties of the organisms in a species that chiefly determine its evolvability and genetic robustness.

Variational properties group together many classical and more recent concepts of evolutionary biology. It includes the classical concepts of pleiotropy, canalization, developmental constraints, developmental bias, morphological integration, developmental homeostasis and later concepts such as robustness, neutral networks, modularity, the G-matrix and distribution of fitness effects.

Variational properties also include the production of DNA sequence variation, epigenetic variation, and phenotypic variation. While the genome is typically thought of as the storehouse of information that generates the organism, it can also be seen as the set of heritable degrees of freedom for varying the organism. DNA thus has both a generative role in the organism, and variational role in the lineage.

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<span class="mw-page-title-main">Phenotype</span> Composite of the organisms observable characteristics or traits

In genetics, the phenotype is the set of observable characteristics or traits of an organism. The term covers the organism's morphology, its developmental processes, its biochemical and physiological properties, its behavior, and the products of behavior. An organism's phenotype results from two basic factors: the expression of an organism's genetic code and the influence of environmental factors. Both factors may interact, further affecting the phenotype. When two or more clearly different phenotypes exist in the same population of a species, the species is called polymorphic. A well-documented example of polymorphism is Labrador Retriever coloring; while the coat color depends on many genes, it is clearly seen in the environment as yellow, black, and brown. Richard Dawkins in 1978 and then again in his 1982 book The Extended Phenotype suggested that one can regard bird nests and other built structures such as caddisfly larva cases and beaver dams as "extended phenotypes".

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Evolutionary biology is the subfield of biology that studies the evolutionary processes that produced the diversity of life on Earth. It is also defined as the study of the history of life forms on Earth. Evolution holds that all species are related and gradually change over generations. In a population, the genetic variations affect the phenotypes of an organism. These changes in the phenotypes will be an advantage to some organisms, which will then be passed on to their offspring. Some examples of evolution in species over many generations are the peppered moth and flightless birds. In the 1930s, the discipline of evolutionary biology emerged through what Julian Huxley called the modern synthesis of understanding, from previously unrelated fields of biological research, such as genetics and ecology, systematics, and paleontology.

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Genetic architecture is the underlying genetic basis of a phenotypic trait and its variational properties. Phenotypic variation for quantitative traits is, at the most basic level, the result of the segregation of alleles at quantitative trait loci (QTL). Environmental factors and other external influences can also play a role in phenotypic variation. Genetic architecture is a broad term that can be described for any given individual based on information regarding gene and allele number, the distribution of allelic and mutational effects, and patterns of pleiotropy, dominance, and epistasis.

<span class="mw-page-title-main">Facilitated variation</span>

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<span class="mw-page-title-main">Phenotypic integration</span>

Phenotypic Integration is a metric for measuring the correlation of multiple functionally-related traits to each other. Complex phenotypes often require multiple traits working together in order to function properly. Phenotypic integration is significant because it provides an explanation as to how phenotypes are sustained by relationships between traits. Every organism's phenotype is integrated, organized, and a functional whole. Integration is also associated with functional modules. Modules are complex character units that are tightly associated, such as a flower. It is hypothesized that organisms with high correlations between traits in a module have the most efficient functions. The fitness of a particular value for one phenotypic trait frequently depends on the value of the other phenotypic traits, making it important for those traits evolve together. One trait can have a direct effect on fitness, and it has been shown that the correlations among traits can also change fitness, causing these correlations to be adaptive, rather than solely genetic. Integration can be involved in multiple aspects of life, not just at the genetic level, but during development, or simply at a functional level.

<span class="mw-page-title-main">Outline of evolution</span>

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In biology, constructive development refers to the hypothesis that organisms shape their own developmental trajectory by constantly responding to, and causing, changes in both their internal state and their external environment. Constructive development can be contrasted with programmed development, the hypothesis that organisms develop according to a genetic program or blueprint. The constructivist perspective is found in philosophy, most notably developmental systems theory, and in the biological and social sciences, including developmental psychobiology and key themes of the extended evolutionary synthesis. Constructive development may be important to evolution because it enables organisms to produce functional phenotypes in response to genetic or environmental perturbation, and thereby contributes to adaptation and diversification.

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In evolutionary biology, developmental bias refers to the production against or towards certain ontogenetic trajectories which ultimately influence the direction and outcome of evolutionary change by affecting the rates, magnitudes, directions and limits of trait evolution. Historically, the term was synonymous with developmental constraint, however, the latter has been more recently interpreted as referring solely to the negative role of development in evolution.

Lee Altenberg is an American theoretical biologist. He is on the faculty of the Departments of Information and Computer Sciences and of Mathematics at the University of Hawaiʻi at Mānoa. He is best known for his work that helped establish the evolution of evolvability and modularity in the genotype–phenotype map as areas of investigation in evolutionary biology, for moving theoretical concepts between the fields of evolutionary biology and evolutionary computation, and for his mathematical unification and generalization of modifier gene models for the evolution of biological information transmission, putting under a single mathematical framework the evolution of mutation rates, recombination rates, sexual reproduction rates, and dispersal rates.

References

  1. Altenberg, L. 1994. The evolution of evolvability in genetic programming. Chapter 3 in Advances in Genetic Programming, ed. Kenneth Kinnear. MIT Press, pp. 47-74.
  2. Altenberg, L. 1995. Genome Growth and the Evolution of the Genotype-Phenotype Map In Evolution and Biocomputation: Computational Models of Evolution. Wolfgang Banzhaf and Frank H. Eeckman (Eds.). pp. 205-259. Berlin: Springer.
  3. Wagner, G. P. and Altenberg, L. 1996. Complex adaptations and the evolution of evolvability. Evolution 50 (3): 967-976.