Phenotypic integration

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Phenotypic integration is a metric for measuring the correlation of multiple functionally-related traits to each other. [1] 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. [2] It is hypothesized that organisms with high correlations between traits in a module have the most efficient functions. [3] 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. [4] Integration can be involved in multiple aspects of life, not just at the genetic level, but during development, or simply at a functional level.

Contents

Integration can be caused by genetic, developmental, environmental, or physiological relationships among characters. [5] Environmental conditions can alter or cause integration, i.e. they may be plastic. [6] Correlational selection, a form of natural selection can also produce integration. At the genetic level, integration can be caused by pleiotropy, close linkage, or linkage disequilibrium among unlinked genes. [7] At the developmental level it can be due to cell-cell signaling such as in the development of the ectopic eyes in Drosophila. It is believed that the patterns of genetic covariance helped distinguish certain species. [8] It can create variation among certain phenotypes, and can facilitate efficiency. This is significant because integration may play a huge role in phenotypic evolution. Phenotypic integration and its evolution can not only create large amounts of variety among phenotypes which can cause variation among species. For example, the color patterns on Garter snakes range widely and are caused by the covariance among multiple phenotypes.

Phenotypic Integration is important for many traits. One specifically is the development of fewer bones in the neurocranium. Cranial bones en.svg
Phenotypic Integration is important for many traits. One specifically is the development of fewer bones in the neurocranium.

Origins

Shortly after the structure of DNA was uncovered, Everett C. Olson and Robert L. Miller (1958) wrote the first book regarding the topic of phenotypic integration. [9] The term integration was first used in reference to genetics by Olson and Miller, referring to correlations among characters that are influenced by selection. [10] Following Olson and Miller, botanical studies on coherence between characters were done spanning over many years. [11] Its first expansion was in the construction of a morphological integration genetic model constructed by Russell Lande (1980). However, the term "Phenotypic Integration" was first coined by Massimo Pigliucci and Katherine Preston, in their book, Phenotypic Integration, which helped elucidate the observed laws of correlation and some theoretical issues regarding the topic. [12]

Natural Selection and Phenotypic Integration

Phenotypic Integration can be favorable or unfavorable with respect to natural selection. It has been shown that certain combinations of correlated traits can be unfavorable to an organism. In an ontogenetic study of laboratory rats, certain covariances among developmental characters which produced differing functions in the skull and limb were less favorable than another set that contributed to skull and limb structure. [13] The most common form of selection on phenotypic integration is correlational selection. Correlational selection is a form of natural selection that favors certain combinations of traits (phenotypic integration). It can promote both genetic correlations and high levels of genetic variation. It has even been found that correlational selection may be the most common form of natural selection. [14] Occasionally, this form of selection will favor a group of traits at the expense of others and if it does favor a particular set of traits it will include the most used traits whose functional effectiveness is essential for their ability to work together, and whose successful interaction is needed for the fitness of the individual. [15] [16] Phenotypic integration may be the adaptive product of correlational selection. An example of natural selection favoring integration is in the color patterns and escape mechanisms of the Garter snake, Thamnophis Ordinoides. [17] Another example is in plants that have highly-specific pollinators, natural selection favors plants that have highly specialized flowering to pair with the specific pollinators, and therefore high floral integration. [18]

Northwestern Garter Snake's color patterns and escape mechanisms are integrated. Thamnophis ordinoides.jpg
Northwestern Garter Snake's color patterns and escape mechanisms are integrated.

Evolution of Phenotypic Integration

Integration can be found at the genetic level due to genetic linkage. Genetic linkage involves multiple genes being inherited together during meiosis because they are close to each other on the same chromosome. Alleles at different loci can be inherited together if they are tightly linked. Large genetic correlations can only be upheld if the loci that influence different characters are tightly linked, or if high levels of inbreeding in the population occur. Even if selection favors the correlations, it will not be maintained unless those conditions are met. Selection will favor tight linkage because it is maintained better. Poorly linked genetic correlations will not last. [19] Transposition allows the loci at different locations on the chromosome to move so that they can become close to each other and be inherited together. This is significant to understanding the relationship between phenotypic integration and evolution because it is one of the mechanisms of how multiple traits that are connected to each other to evolve and change together. For instance, the Papilio dardanus butterflies come in three different forms, each mimicking a different distasteful butterfly species. [20] Multiple loci contribute to these different forms, and a butterfly with alleles for form A at one locus and B at another locus would have poor fitness. However, the multiple loci are tightly linked, so they are inherited together as a single allele. Through transposition, these multiple loci ended up close to each other. [21]

Mutations among these linked genes are the nonadaptive fuel which can create evolution. Evolution may also occur because the integration may have an adaptive advantage in a particular environment for an organism. It is also important to recognize that not only can the traits be inherited together, but inherited separately and selected together. Another important example of phenotypic integration evolving over time is the relationship between the neurocranium and the brain. Over the last 150 million years the number of bones in the brain has decreased while the size of the brain in mammals has changed. Integration between the brain and the skull has evolved over this time period to reduce the number of bones in the cranium, while increasing the size of the brain. This relationship between correlated traits has played an important role in the evolution of mammalian cranium structure and brain size. [22] Finally, development is another crucial cause of phenotypic integration that has evolved over time. Cell-signaling pathways which utilize integration in the form of complex interactions among specific cells in the pathway are crucial to proper development in many organisms. The interactions among the cells in the pathway, and the interaction of the pathways with other pathways have evolved over time to create complex structures. [23]

Examples

Aposematism in poison dart frogs has also shown that phenotypic integration may be involved. Aposematism is the use of warning colors to deter predators because it often conveys the organism being poisonous, and this study found that diet specialization, and chemical defense are integrated and help affect aposematism. [24]

The Poison Dart Frog is an example of a species that illustrates phenotypic integration. Korreldragende-gifkikker-3.jpg
The Poison Dart Frog is an example of a species that illustrates phenotypic integration.

In another study regarding the relationship of sexual ornaments and phenotypic integration, there seems to be a paradox where sexual traits are expected to be both less integrated for greater expression, and more integrated to better indicate physiological quality. However, in the case of the house finch, the female house finches select for males based on their likelihood to be a good parent. The females base their choice of male parental behaviors on the elaboration of the male's sexual ornamentation. Thus, female choice favors hormonally controlled integration of male sexual behaviors and male sexual ornamentation. [25]

Phylogenetically consistent patterns of phenotypic integration have also been recently reported in leaves, floral morphology, and dry fruits as well as in the morphology of some animal organs. [26] [27]

Future work

Understanding phenotypic integration will continue as more research and understanding is done with regards to genetic, developmental, and physiological mechanisms, and learn more about the relationship of selection and complex phenotypes. [28] Research of this topic can even be beneficial to modern biomedicine. [29]

Related Research Articles

<span class="mw-page-title-main">Heredity</span> Passing of traits to offspring from the species parents or ancestor

Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic information of their parents. Through heredity, variations between individuals can accumulate and cause species to evolve by natural selection. The study of heredity in biology is genetics.

<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".

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative phenotypes, in the population of a species. To be classified as such, morphs must occupy the same habitat at the same time and belong to a panmictic population.

A quantitative trait locus (QTL) is a locus that correlates with variation of a quantitative trait in the phenotype of a population of organisms. QTLs are mapped by identifying which molecular markers correlate with an observed trait. This is often an early step in identifying the actual genes that cause the trait variation.

Evolvability is defined as the capacity of a system for adaptive evolution. Evolvability is the ability of a population of organisms to not merely generate genetic diversity, but to generate adaptive genetic diversity, and thereby evolve through natural selection.

<span class="mw-page-title-main">Baldwin effect</span> Effect of learned behavior on evolution

In evolutionary biology, the Baldwin effect describes an effect of learned behaviour on evolution. James Mark Baldwin and others suggested that an organism's ability to learn new behaviours will affect its reproductive success and will therefore have an effect on the genetic makeup of its species through natural selection. It posits that subsequent selection might reinforce the originally learned behaviors, if adaptive, into more in-born, instinctive ones. Though this process appears similar to Lamarckism, that view proposes that living things inherited their parents' acquired characteristics. The Baldwin effect only posits that learning ability, which is genetically based, is another variable in / contributor to environmental adaptation. First proposed during the Eclipse of Darwinism in the late 19th century, this effect has been independently proposed several times, and today it is generally recognized as part of the modern synthesis.

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">Pleiotropy</span> Influence of a single gene on multiple phenotypic traits

Pleiotropy occurs when one gene influences two or more seemingly unrelated phenotypic traits. Such a gene that exhibits multiple phenotypic expression is called a pleiotropic gene. Mutation in a pleiotropic gene may have an effect on several traits simultaneously, due to the gene coding for a product used by a myriad of cells or different targets that have the same signaling function.

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

The theory of facilitated variation demonstrates how seemingly complex biological systems can arise through a limited number of regulatory genetic changes, through the differential re-use of pre-existing developmental components. The theory was presented in 2005 by Marc W. Kirschner and John C. Gerhart.

<span class="mw-page-title-main">Canalisation (genetics)</span> Measure of the ability of a population to produce the same phenotype

Canalisation is a measure of the ability of a population to produce the same phenotype regardless of variability of its environment or genotype. It is a form of evolutionary robustness. The term was coined in 1942 by C. H. Waddington to capture the fact that "developmental reactions, as they occur in organisms submitted to natural selection...are adjusted so as to bring about one definite end-result regardless of minor variations in conditions during the course of the reaction". He used this word rather than robustness to consider that biological systems are not robust in quite the same way as, for example, engineered systems.

An evolutionary landscape is a metaphor or a construct used to think about and visualize the processes of evolution acting on a biological entity. This entity can be viewed as searching or moving through a search space. For example, the search space of a gene would be all possible nucleotide sequences. The search space is only part of an evolutionary landscape. The final component is the "y-axis", which is usually fitness. Each value along the search space can result in a high or low fitness for the entity. If small movements through search space cause changes in fitness that are relatively small, then the landscape is considered smooth. Smooth landscapes happen when most fixed mutations have little to no effect on fitness, which is what one would expect with the neutral theory of molecular evolution. In contrast, if small movements result in large changes in fitness, then the landscape is said to be rugged. In either case, movement tends to be toward areas of higher fitness, though usually not the global optima.

Genetic assimilation is a process described by Conrad H. Waddington by which a phenotype originally produced in response to an environmental condition, such as exposure to a teratogen, later becomes genetically encoded via artificial selection or natural selection. Despite superficial appearances, this does not require the (Lamarckian) inheritance of acquired characters, although epigenetic inheritance could potentially influence the result. Waddington stated that genetic assimilation overcomes the barrier to selection imposed by what he called canalization of developmental pathways; he supposed that the organism's genetics evolved to ensure that development proceeded in a certain way regardless of normal environmental variations.

<span class="mw-page-title-main">Evolutionary physiology</span> Study of changes in physiological characteristics

Evolutionary physiology is the study of the biological evolution of physiological structures and processes; that is, the manner in which the functional characteristics of individuals in a population of organisms have responded to natural selection across multiple generations during the history of the population. It is a sub-discipline of both physiology and evolutionary biology. Practitioners in the field come from a variety of backgrounds, including physiology, evolutionary biology, ecology, and genetics.

In multivariate quantitative genetics, a genetic correlation is the proportion of variance that two traits share due to genetic causes, the correlation between the genetic influences on a trait and the genetic influences on a different trait estimating the degree of pleiotropy or causal overlap. A genetic correlation of 0 implies that the genetic effects on one trait are independent of the other, while a correlation of 1 implies that all of the genetic influences on the two traits are identical. The bivariate genetic correlation can be generalized to inferring genetic latent variable factors across > 2 traits using factor analysis. Genetic correlation models were introduced into behavioral genetics in the 1970s–1980s.

<span class="mw-page-title-main">Genetic variance</span> Biological concept

Genetic variance is a concept outlined by the English biologist and statistician Ronald Fisher in his fundamental theorem of natural selection. In his 1930 book The Genetical Theory of Natural Selection, Fisher postulates that the rate of change of biological fitness can be calculated by the genetic variance of the fitness itself. Fisher tried to give a statistical formula about how the change of fitness in a population can be attributed to changes in the allele frequency. Fisher made no restrictive assumptions in his formula concerning fitness parameters, mate choices or the number of alleles and loci involved.

The Extended Evolutionary Synthesis (EES) consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis was called for in the 1950s by C. H. Waddington, argued for on the basis of punctuated equilibrium by Stephen Jay Gould and Niles Eldredge in the 1980s, and was reconceptualized in 2007 by Massimo Pigliucci and Gerd B. Müller.

A human disease modifier gene is a modifier gene that alters expression of a human gene at another locus that in turn causes a genetic disease. Whereas medical genetics has tended to distinguish between monogenic traits, governed by simple, Mendelian inheritance, and quantitative traits, with cumulative, multifactorial causes, increasing evidence suggests that human diseases exist on a continuous spectrum between the two.

<span class="mw-page-title-main">Epistasis</span> Dependence of a gene mutations phenotype on mutations in other genes

Epistasis is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is dependent on the genetic background in which it appears. Epistatic mutations therefore have different effects on their own than when they occur together. Originally, the term epistasis specifically meant that the effect of a gene variant is masked by that of different gene.

This glossary of genetics and evolutionary biology is a list of definitions of terms and concepts used in the study of genetics and evolutionary biology, as well as sub-disciplines and related fields, with an emphasis on classical genetics, quantitative genetics, population biology, phylogenetics, speciation, and systematics. Overlapping and related terms can be found in Glossary of cellular and molecular biology, Glossary of ecology, and Glossary of biology.

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.

References

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