Richard Borowsky

Bio: Richard Borowsky is an academic researcher from New York University. The author has contributed to research in topics: Cavefish & Xiphophorus. The author has an hindex of 33, co-authored 77 publications receiving 4436 citations. Previous affiliations of Richard Borowsky include Wildlife Conservation Society.

Genetic analysis of cavefish reveals molecular convergence in the evolution of albinism

Abstract: The genetic basis of vertebrate morphological evolution has traditionally been very difficult to examine in naturally occurring populations. Here we describe the generation of a genome-wide linkage map to allow quantitative trait analysis of evolutionarily derived morphologies in the Mexican cave tetra, a species that has, in a series of independent caves, repeatedly evolved specialized characteristics adapted to a unique and well-studied ecological environment. We focused on the trait of albinism and discovered that it is linked to Oca2, a known pigmentation gene, in two cave populations. We found different deletions in Oca2 in each population and, using a cell-based assay, showed that both cause loss of function of the corresponding protein, OCA2. Thus, the two cave populations evolved albinism independently, through similar mutational events.

Cryptic variation in morphological evolution: HSP90 as a capacitor for loss of eyes in cavefish.

Abstract: In the process of morphological evolution, the extent to which cryptic, preexisting variation provides a substrate for natural selection has been controversial. We provide evidence that heat shock protein 90 (HSP90) phenotypically masks standing eye-size variation in surface populations of the cavefish Astyanax mexicanus. This variation is exposed by HSP90 inhibition and can be selected for, ultimately yielding a reduced-eye phenotype even in the presence of full HSP90 activity. Raising surface fish under conditions found in caves taxes the HSP90 system, unmasking the same phenotypic variation as does direct inhibition of HSP90. These results suggest that cryptic variation played a role in the evolution of eye loss in cavefish and provide the first evidence for HSP90 as a capacitor for morphological evolution in a natural setting.

A Novel Role for Mc1r in the Parallel Evolution of Depigmentation in Independent Populations of the Cavefish Astyanax mexicanus

Abstract: The evolution of degenerate characteristics remains a poorly understood phenomenon. Only recently has the identification of mutations underlying regressive phenotypes become accessible through the use of genetic analyses. Focusing on the Mexican cave tetra Astyanax mexicanus, we describe, here, an analysis of the brown mutation, which was first described in the literature nearly 40 years ago. This phenotype causes reduced melanin content, decreased melanophore number, and brownish eyes in convergent cave forms of A. mexicanus. Crosses demonstrate non-complementation of the brown phenotype in F2 individuals derived from two independent cave populations: Pachon and the linked Yerbaniz and Japones caves, indicating the same locus is responsible for reduced pigmentation in these fish. While the brown mutant phenotype arose prior to the fixation of albinism in Pachon cave individuals, it is unclear whether the brown mutation arose before or after the fixation of albinism in the linked Yerbaniz/Japones caves. Using a QTL approach combined with sequence and functional analyses, we have discovered that two distinct genetic alterations in the coding sequence of the gene Mc1r cause reduced pigmentation associated with the brown mutant phenotype in these caves. Our analysis identifies a novel role for Mc1r in the evolution of degenerative phenotypes in blind Mexican cavefish. Further, the brown phenotype has arisen independently in geographically separate caves, mediated through different mutations of the same gene. This example of parallelism indicates that certain genes are frequent targets of mutation in the repeated evolution of regressive phenotypes in cave-adapted species.

Cryptic Variation in Morphological Evolution: HSP90 as a Capacitor for Loss of Eyes in Cavefish

Abstract: In the process of morphological evolution, the extent to which cryptic, preexisting variation provides a substrate for natural selection has been controversial. We provide evidence that heat shock protein 90 (HSP90) phenotypically masks standing eye-size variation in surface populations of the cavefish Astyanax mexicanus. This variation is exposed by HSP90 inhibition and can be selected for, ultimately yielding a reduced-eye phenotype even in the presence of full HSP90 activity. Raising surface fish under conditions found in caves taxes the HSP90 system, unmasking the same phenotypic variation as does direct inhibition of HSP90. These results suggest that cryptic variation played a role in the evolution of eye loss in cavefish and provide the first evidence for HSP90 as a capacitor for morphological evolution in a natural setting.

The Ecology of Individuals: Incidence and Implications of Individual Specialization

Abstract: Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species dis- tributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can some- times comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, be- havioral, and ecological mechanisms that can generate intrapopu- lation variation. Finally, individual specialization has potentially im- portant ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.

Phenotypic Plasticity and the Origins of Diversity

Abstract: Phenotypic plasticity is the ability of a single genotype to produce more than one alternative form of morphology, physiological state, and/or behavior in response to environmental conditions. "Plasticity" and "development" are related terms that are becoming increasingly common in evolutionary biology and ecology. Both phenomena have passed through a period of neglect. In the 1960s Wigglesworth (228, p. 107) described some geneticists as being "apologetic" about environmentally cued polymorphisms, which they considered examples of unfortunate defects in the delicate genetic apparatus: "As R. A. Fisher once said to me, it is not surprising that such elaborate machinery should sometimes go wrong." And Bradshaw (19, p. 148) noted that botanists were carefully avoiding any mention of plasticity; environmental effects in experiments were considered "only an embarrassment." Until recently, genetic considerations have predominated in discussions of evolution and selection. Compared to the enormous progress made in genetics, there has been relatively little systematic effort to analyze environmental effects on the phenotype, and their evolutionary consequences. The plastic phenotype, stigmatized by poorly understood environmental influences and the ghost of Lamarck, has sometimes been lost from view as the focus of selection (e.g. 46; but see 48, 49). Much recent progress has been made toward integrating developmental and evolutionary biology, especially in vertebrate morphology (2, 12, 16, 216), developmental genetics (16, 163, 164), and molecular biology (103; also see 10, 111). "Developmental constraints" is a term symptomatic of this progress, though an unfortunate one because it seems to imply that the main effect of