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Biomedical subjects

Trevor D Price

Publications and source records attributed to Trevor D Price.

8 recordsLinked to original sources

Phenotypic plasticity, sexual selection and the evolution of colour patterns.

When a population comes to occupy a new environment, phenotypically plastic responses alter the distribution of phenotypes, and hence affect both the direction and the intensity of selection. Rates of evolution can be accelerated or retarded compared to what would happen in the absence of plasticity. Plastic responses in one trait result in novel selection pressures on other traits, and this can lead to evolution in completely different directions than predicted in the absence of plasticity. In this paper I use the concept of the adaptive surface in order to identify conditions under which the various different outcomes are expected. I then discuss differences between sexually and naturally selected traits. Sexually selected traits are often expected to be plastic in their expression, with individuals in high condition developing greater elaboration. As examples of sexually selected traits I review the evolution of colour patterns in birds with a view to assessing the magnitude of plastic responses in their development, and to ask how such responses may have influenced genetic evolution. The common colour pigments in birds are carotenoids and melanins. Both are used in social signaling, and consequently are expected to evolve to be phenotypically plastic indicators of an individual's quality. Perhaps partly because they are condition indicators, the quantity of carotenoids in the plumage can be strongly influenced by diet. Examples are described where alterations of carotenoids in the diet are thought to have altered the phenotype, driving genetic evolution in novel directions. Melanin patterns seem to be less affected by diet, but the intensity of melanization after moult is affected by social interactions during the moult and by raising birds in humid conditions. Hormonal manipulations can have dramatic effects on both the kinds of melanin produced (eumelanin or phaeomelanin) as well as the patterns they form. Differences between species in melanin patterns resemble differences produced by environmental manipulations, as well as those produced by simple modulations of parameters in computer simulations of pattern formation. While phenotypic plasticity is one way that genetic change in plumage patterns (and other traits) could be driven, there are others, including the appearance of major mutations and selection on standing variation whose distribution is not altered in the new environment. I consider some evidence for the different alternatives, and ask when they might lead to qualitatively different evolutionary outcomes.

Adaptation, Physiological↗

Correcting for regression to the mean in behavior and ecology.

If two successive trait measurements have a less-than-perfect correlation, individuals or populations will, on average, tend to be closer to the mean on the second measurement (the so-called regression effect). Thus, there is a negative correlation between an individual's state at time 1 and the change in state from time 1 to time 2. In addition, whenever groups differ in their initial mean values, the expected change in the mean value from time 1 to time 2 will differ among the groups. For example, birds feeding nestlings lose weight, but initially heavier birds lose more weight than lighter birds, a result expected from the regression effect. In sexual selection, males who remain unmated in the first year are, on average, less attractive than mated males. The regression effect predicts that these males will increase their attractiveness in the second year more than mated males. In well-designed experiments, changes in the experimental and control groups would be compared. In observational studies, however, no such comparison is available, and expected differential effects must be accounted for before they can be attributed to external causes. We describe methods to correct for the regression effect and assess alternative causal explanations.

Animals↗

Speciation by distance in a ring species.

Ring species, which consist of two reproductively isolated forms connected by a chain of intergrading populations, have often been described as examples of speciation despite gene flow between populations, but this has never been demonstrated. We used amplified fragment length polymorphism (AFLP) markers to study gene flow in greenish warblers (Phylloscopus trochiloides). These genetic markers show distinct differences between two reproductively isolated forms but gradual change through the ring connecting these forms. These findings provide the strongest evidence yet for "speciation by force of distance" in the face of ongoing gene flow.

Adaptation, Biological↗

Adaptive phenotypic plasticity and the successful colonization of a novel environment.

Behavior and other forms of phenotypic plasticity potentially enable individuals to deal with novel situations. This implies that establishment of a population in a new environment is aided by plastic responses, as first suggested by Baldwin (1896). In the early 1980s, a small population of dark-eyed juncos from a temperate, montane environment became established in a Mediterranean climate in coastal San Diego. The breeding season of coastal juncos is more than twice as long as that of the ancestral population, and they fledge approximately twice as many young. We investigated the adaptive significance of the longer breeding season and its consequences for population persistence. Within the coastal population, individuals with longer breeding seasons have higher offspring production and recruitment, with no measured detrimental effects such as higher mortality or lower reproductive success the following year. Population size has remained approximately constant during the 6 years of study (1998-2003). The increase in reproductive effort in the coastal population contributes substantially to the persistence of this population because there is no evidence of density-dependent recruitment, which would otherwise negate the effects of increased fledgling production. These results provide the first quantitative support of Baldwin's proposition that plasticity can be crucial for population persistence during the early stages of colonization.

Adaptation, Physiological↗

Comparative methods based on species mean values.

Comparative methods that use simple linear regression based on species mean values introduce three difficulties with respect to the standard regression model. First, species values may not be independent because they form part of a hierarchically structured phylogeny. Second, variation about the regression line includes two sources of error: 'biological error' due to deviations of the true species mean values from the regression line and sampling error associated with the estimation of these mean values [B. Riska, Am. Natural. 138 (1991) 283]. Third, sampling error in the independent variable results in an attenuated estimate of the regression slope. We consider estimation and hypothesis testing using two statistical models which explicitly justify the use of the species mean values, without the need to account for phylogenetic relationships. The first (random-effects) is based on an evolutionary model whereby species evolve to fill a bivariate normal niche space, and the second (fixed-effects) is concerned with describing a relationship among the particular species included in a study, where the only source of error is in the estimation of species mean values. We use a modification of the maximum-likelihood method to obtain an unbiased estimate of the regression slope. For three real datasets we find a close correspondence between this slope and that obtained by simply regressing the species mean values on each other. In the random effects model, the P-value also approximates that based on the regression of species mean values. In the fixed effects model, the P-value is typically much lower. Simulated examples illustrate that the maximum-likelihood approach is useful when the accuracy in estimating the species mean values is low, but the traditional method based on a regression of the species mean values may often be justified provided that the evolutionary model can be justified.

Acer↗

The role of phenotypic plasticity in driving genetic evolution.

Models of population divergence and speciation are often based on the assumption that differences between populations are due to genetic factors, and that phenotypic change is due to natural selection. It is equally plausible that some of the differences among populations are due to phenotypic plasticity. We use the metaphor of the adaptive landscape to review the role of phenotypic plasticity in driving genetic evolution. Moderate levels of phenotypic plasticity are optimal in permitting population survival in a new environment and in bringing populations into the realm of attraction of an adaptive peak. High levels of plasticity may increase the probability of population persistence but reduce the likelihood of genetic change, because the plastic response itself places the population close to a peak. Moderate levels of plasticity arise whenever multiple traits, some of which are plastic and others not, form a composite trait involved in the adaptive response. For example, altered behaviours may drive selection on morphology and physiology. Because there is likely to be a considerable element of chance in which behaviours become established, behavioural change followed by morphological and physiological evolution may be a potent force in driving evolution in novel directions. We assess the role of phenotypic plasticity in stimulating evolution by considering two examples from birds: (i) the evolution of red and yellow plumage coloration due to carotenoid consumption; and (ii) the evolution of foraging behaviours on islands. Phenotypic plasticity is widespread in nature and may speed up, slow down, or have little effect on evolutionary change. Moderate levels of plasticity may often facilitate genetic evolution but careful analyses of individual cases are needed to ascertain whether plasticity has been essential or merely incidental to population differentiation.

Adaptation, Biological↗

The evolution of F1 postzygotic incompatibilities in birds.

We analyzed the rate at which postzygotic incompatibilities accumulate in birds. Our purposes were to assess the role of intrinsic F1 hybrid infertility and inviability in the speciation process, and to compare rates of loss of fertility and viability between the sexes. Among our sample more than half the crosses between species in the same genus produce fertile hybrids. Complete loss of F1 hybrid fertility takes on the order of millions of years. Loss of F1 hybrid viability occurs over longer timescales than fertility: some viable hybrids have been produced between taxa that appear to have been separated for more than 55 my. There is strong support for Haldane's rule, with very few examples where the male has lower fitness than the female. However, in contrast to Drosophila, fertility of the homogametic sex in the F1 appears to be lost before viability of the heterogametic sex in the F1. We conclude that the time span of loss of intrinsic hybrid fertility and viability is often, but not always, longer than the time to speciation. Premating isolation is an important mechanism maintaining reproductive isolation in birds. In addition, other factors causing postzygotic reproductive isolation such as ecological causes of hybrid unfitness, reduced mating success of hybrids, and genetic incompatibilities in the F2s and backcrosses may often be involved in the speciation process.

Animals↗

Domesticated birds as a model for the genetics of speciation by sexual selection.

In theory, even populations occupying identical environments can diverge in sexually selected traits, as a consequence of different mutational input. I evaluate the potential of this process by comparing the genetics of breeds of domesticated birds to what is known about the genetics of differences among species. Within domesticated species there is a strong correlation of time since domestication with the number of breeds. Descendants of the rock dove, Columba livia (the oldest domesticate) show differences in courtship, vocalizations, body shape, feather ornaments (crests and tails) and colors and color patterns. When nine other domesticated species are included there is a striking hierarchy, with more recent domesticates having a nested subset of these traits: the youngest domesticated species have breeds distinguished only by color. This suggests that selection of new, visible, mutations is driving the process of breed diversification, with mutations that appeal to the breeder happening the most frequently in color. In crosses among related species, color, feather ornaments and many vocalizations and displays show both intermediate dominance and pure dominance. Although the number of loci affecting each of these traits is typically unknown, limited evidence of the genetics of species' differences suggests that some differences are due to the substitution of single genes of major effect. While neither the genetics of breeds nor the genetics of species provide a perfect model for the genetics of speciation, similarities between the two are sufficiently striking to infer that major, visible, mutations can provide the impetus underlying new directions of sexual selection.

Acoustic Stimulation↗