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PubMed2003

Mate choice theory and the mode of selection in sexual populations.

Indirect new data imply that mate and/or gamete choice are major selective forces driving genetic change in sexual populations. The system dictates nonrandom mating, an evolutionary process requiring both revised genetic theory and new data on heritability of characters underlying Darwinian fitness. Successfully reproducing individuals represent rare selections from among vigorous, competing survivors of preadult natural selection. Nonrandom mating has correlated demographic effects: reduced effective population size, inbreeding, low gene flow, and emphasis on deme structure. Characters involved in choice behavior at reproduction appear based on quantitative trait loci. This variability serves selection for fitness within the population, having only an incidental relationship to the origin of genetically based reproductive isolation between populations. The claim that extensive hybridization experiments with Drosophila indicate that selection favors a gradual progression of "isolating mechanisms" is flawed, because intra-group random mating is assumed. Over deep time, local sexual populations are strong, independent genetic systems that use rich fields of variable polygenic components of fitness. The sexual reproduction system thus particularizes, in small subspecific populations, the genetic basis of the grand adaptive sweep of selective evolutionary change, much as Darwin proposed.

Animals↗
PubMed2002

A test of evolutionary theories of aging.

Senescence is a nearly universal feature of multicellular organisms, and understanding why it occurs is a long-standing problem in biology. The two leading theories posit that aging is due to (i) pleiotropic genes with beneficial early-life effects but deleterious late-life effects ("antagonistic pleiotropy") or (ii) mutations with purely deleterious late-life effects ("mutation accumulation"). Previous attempts to distinguish these theories have been inconclusive because of a lack of unambiguous, contrasting predictions. We conducted experiments with Drosophila based on recent population-genetic models that yield contrasting predictions. Genetic variation and inbreeding effects increased dramatically with age, as predicted by the mutation theory. This increase occurs because genes with deleterious effects with a late age of onset are unopposed by natural selection. Our findings provide the strongest support yet for the mutation theory.

Aging↗
PubMed1969

Polymorphism resulting from the mating advantage of rare male genotypes.

An algebraic model is presented for selection by the mating advantage of rare genotypes. In accordance with the results of recent experiments, the selective advantage is formulated as inversely proportional to genotype frequency. The model applies to any number of alleles at an autosomal locus or at a sex-linked locus, and it provides for a different intensity of selection for each genotype in each sex. The frequency of an allele under this selection proceeds to a stable, nontrivial equilibrium when any one of the genotypes carrying the allele and any one of those carrying another allele possess some degree of the advantage which varies inversely with genotype frequency. The changes in gene frequency are not affected by the mating system, being, for instance, the same with random mating as with inbreeding. With low values for the selection parameters, genetic polymorphisms could be maintained with little selection apparent except at rather low or rather high gene frequencies. The rare genotype advantage may be a significant element among the mechanisms which maintain the extensive genetic polymorphisms of natural populations.

Animals↗
PubMed1972

Evidence for coadaptation in Avena barbata.

Two populations of Avena barbata have been sampled intensively and allelic frequencies have been determined on a fine grid for six enzyme loci. The distribution of alleles among microniches within the location occupied by one of these populations (CSA-1) was correlated with environment following patterns of macro- and microdifferentiation previously described in California. Multilocus analyses revealed striking gameticphase disequilibrium for both linked and unlinked loci, and also that two balanced five-locus gametic types, one associated with mesic conditions and the other with more xeric conditions, are in substantial excess at the CSA-1 location. Evidence is presented that these two gametic types are maintained by selection acting on two coadapted combinations of alleles, held together by restriction of recombination resulting from linkage and/or inbreeding. Gametic-phase disequilibrium featuring different allelic complexes were found in each of three microniches in the second population (Geyserville), which occupies a site that is environmentally atypical for A. barbata.

Adaptation, Physiological↗
PubMed1975

Genetic variation as a test of natural selection.

Allozymic variation encoded by 26 loci was analyzed electrophoretically in 507 specimens representing 12 populations of green toads, Bufo viridis, in Israel and the Vis Adriatic Island. Genetic variation in Bufo viridis is higher than in any vertebrate yet studied. Mean heterozygosity per locus per individual (H) is 0.133 (range, 0.105 to 0.159). H is higher in central populations as compared with isolates, and varies among four major protein classes, being highest in transferases and hydrolases and lowest in oxidoreductases and nonenzymatic proteins. Differential gene frequencies among polymorphisms was tested as an indicator of natural selection. Significant heterogeneity between loci in their apparent inbreeding coefficients Fe=S-2p/P(1-p) was found for all alleles and for each of the four major classes of proteins tested, which may be taken as evidence of selection. Both uniform and diversifying selection are suggested by the low and high Fe values, respectively. The general pattern of high heterozygosity in Bufo viridis is best explained as an adaptive strategy in heterogeneous environments.

Alleles↗
PubMed1994

A role for molecular genetics in biological conservation.

The recognition of recent accelerated depletion of species as a consequence of human industrial development has spawned a wide interest in identifying threats to endangered species. In addition to ecological and demographic perils, it has become clear that small populations that narrowly survive demographic contraction may undergo close inbreeding, genetic drift, and loss of overall genomic variation due to allelic loss or reduction to homozygosity. I review here the consequences of such genetic depletion revealed by applying molecular population genetic analysis to four endangered mammals: African cheetah, lion, Florida panther, and humpback whale. The accumulated genetic results, combined with physiological, ecological, and ethological data, provide a multifaceted perspective of the process of species diminution. An emerging role of population genetics, phylogenetics, and phylogeography as indicators of a population's natural history and its future prognosis provides valuable data of use in the development of conservation management plans for endangered species.

Africa↗
PubMed2005

Plant population dynamics, pollinator foraging, and the selection of self-fertilization.

Many flowering plants rely on pollinators, self-fertilization, or both for reproduction. We model the consequences of these features for plant population dynamics and mating system evolution. Our mating systems-based population dynamics model includes an Allee effect. This often leads to an extinction threshold, defined as a density below which population densities decrease. Reliance on generalist pollinators who primarily visit higher density plant species increases the extinction threshold, whereas autonomous modes of selfing decrease and can eliminate the threshold. Generalist pollinators visiting higher density plant species coupled with autonomous selfing may introduce an effect where populations decreasing in density below the extinction threshold may nonetheless persist through selfing. The extinction threshold and selfing at low density result in populations where individuals adopting a single reproductive strategy exhibit mating systems that depend on population density. The ecological and evolutionary analyses provide a mechanism where prior selfing evolves even though inbreeding depression is greater than one-half. Simultaneous consideration of ecological and evolutionary dynamics confirms unusual features (e.g., evolution into extinction or abrupt increases in population density) implicit in our separate consideration of ecological and evolutionary scenarios. Our analysis has consequences for understanding pollen limitation, reproductive assurance, and the evolution of mating systems.

Animals↗
PubMed2004

The Causes of self-sterility in natural populations of the relictual angiosperm, Illicium floridanum (Illiciaceae).

BACKGROUND AND AIMS: Illicium floridanum, a species belonging to the basal extant angiosperm taxon Illiciaceae, reportedly exhibits self-incompatibility (SI). To date, the site and timing of SI within the carpel of this species remains unidentified. Thus, the objective of this research was to determine the cellular and temporal aspects of SI in I. floridanum. METHODS: Following controlled application of cross- and self-pollen in natural populations of I. floridanum, embryo sac development and temporal aspects of stigma receptivity, as well as pollen tube growth, fertilization, and embryo and endosperm development, were investigated with the aid of light and fluorescence microscopy. KEY RESULTS: Flowers of I. floridanum exhibited complete dichogamy whereby stigmas only supported cross- and self-pollen tube growth prior to anther dehiscence. In contrast to earlier reports of SI in this species, a prezygotic SI resulting in rejection of self-pollen tube growth at the stigma was absent and there were no significant differences between cross- versus self-pollen germination and pollen tube growth within the style and ovary during the first 5 d after pollination. Structural development of the four-celled embryo sac was not differentially influenced by pollen type as noted to occur in other angiosperms with late-acting ovarian SI. The ovule micropyle and embryo sac were penetrated equally by cross- and self-pollen tubes. In addition, there were no statistically significant differences in cross- versus self-fertilization. A resting zygote and multicellular endosperm at a variety of developmental stages was present by 30 d after application of cross- or self-pollen. CONCLUSIONS: In the clear absence of a prezygotic SI that was previously reported to result in differential self-pollen tube growth at the stigma, self- sterility in I. floridanum is likely due to early-acting inbreeding depression, although late-acting post-zygotic ovarian SI cannot be ruled out.

Crosses, Genetic↗
PubMed2006

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae).

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.

Animals↗
PubMed2003

PedNavigator: a pedigree drawing servlet for large and inbred populations.

PedNavigator is a pedigree drawing application for large and complex pedigrees. It has been developed especially for genetic and epidemiological studies of isolated populations characterized by high inbreeding and multiple matrimonies. PedNavigator is written in Java and is intended as a server-side web application, allowing researchers to 'walk' through family ties by point-and-clicking on person's symbols. The application is able to enrich the pedigree drawings with genotypic and phenotypic information taken from the underlying relational database.

Algorithms↗
PubMed2005

IRILmap: linkage map distance correction for intermated recombinant inbred lines/advanced recombinant inbred strains.

Intermated Recombinant Inbred Lines (IRILs) in plants, or Advanced Recombinant Inbred Strains in animals, are constructed by carrying out generations of intermating between F2 individuals before starting recurrent inbreeding generations by selfing or sib-mating. IRILs are powerful for high-resolution genetic mapping because they have undergone more recombination than usual Recombinant Inbred Lines (RILs). However, there is no mapping software able to generate actual centiMorgan distances from the segregation data obtained with IRILs. IRILmap software converts genetic distances computed with any linkage mapping program designed for RILs, so that IRIL-derived data can be used to get actual centiMorgan distances, directly comparable to F2, backcross or RIL-derived maps.

Algorithms↗
PubMed1986

Possible derivation of the laboratory mouse genome from multiple wild Mus species.

Laboratory strains of mice are thought to be derived from wild populations of Mus domesticus. Many instances of non-domesticus genetic information fixed in these strains have been described, however, and the amount of strain-to-strain genetic variation exceeds that found in wild domesticus populations. In order to estimate the extent of the non-domesticus contribution to laboratory mouse genomes, and to determine whether it could account for observed variation, we have used computer simulations to investigate the properties of genetically marked chromosomal segments and the distribution of residual allogenicity at various times during inbreeding. A locus or chromosomal segment is allogenic if it is unfixed within a lineage at a given time. The odds of fixation of a foreign chromosome segment are predicted to be an exponentially decreasing function of its length. The median segment length is predicted to be 17 centimorgans. Available data for markers of chromosomes 1, 9 and 12 in recombinant inbred strain sets conform to these predictions. Together, the results suggest that introgression of non-domesticus chromosomes and segregation of residual allogenicity are sufficient to account for the genetic diversity observed among inbred mouse strains and substrains.

Alleles↗
PubMed1987

Definition and estimation of higher-order gene fixation indices.

Fixation indices summarize the associations between genes that arise from the joint effects of inbreeding and selection. In this paper, fixation indices are derived for pairs, triplets and quadruplets of genes at a single multiallelic locus. The fixation indices are obtained by dividing cumulants by constants; the cumulants describe the statistical distribution of alleles and the constants are functions of gene frequency. The use of cumulants instead of moments is necessary only for four-gene indices, when the fourth cumulant is used. A second type of four-gene index is also required, and this index is based upon the covariation of second-order cumulants. At multiallelic loci, a large number of indices is possible. If alleles are selectively neutral, the number of indices is reduced and the relationship between gene identity and gene cumulants is shown.--Two-gene indices can always be estimated from genotypic frequency data at a single polymorphic locus. Three-gene indices are also estimable except when allele frequency equals one-half. Four-gene indices are not estimable unless selection is assumed to have an equal effect upon each allele (such as under selective neutrality) and the locus contains at least three alleles of unequal frequency. For diallelic or selected loci, an alternative four-gene fixation index is proposed. This index incorporates both types of four-gene associations but cannot be related to gene identity.

Alleles↗
PubMed1989

Gustave Malécot and the transition from classical to modern population genetics.

The contributions of Gustave Malécot to theoretical population genetics are described, discussed, and put into perspective relative to earlier and later work. In this context, certain aspects of the theory of inbreeding, the correlation between relatives, the evolution of finite panmictic populations, and (in more depth) spatial variation are reviewed. A brief biographical sketch of Malécot is also presented.

France↗
PubMed1994

A reevaluation of data from competitive tests shows high levels of heterosis in Drosophila melanogaster.

We have analyzed the results from a range of procedures designed to measure the fitness under competitive conditions of inbred strains of Drosophila melanogaster, specifically strains which are homozygous for chromosome 2. All methods show a substantial reduction in fitness, ranging from an estimated 70-80% for single generation competition tests to 80-90% for a multiple generation population cage procedure. Furthermore, inbreeding through brother-sister mating reduces fitness by a comparable amount when allowance is made for the expected degree of homozygosity.

Animals↗
PubMed1994

Effective population sizes with multiple paternity.

While the concept of effective population size is of obvious applicability to many questions in population genetics and conservation biology, its utility has suffered due to a lack of agreement among its various formulations. Often, mathematical formulations for effective sizes apply restrictive assumptions that limit their applicability. Herein, expressions for effective sizes of populations that account for mating tactics, biases in sex ratios, and differential dispersal rates (among other parameters) are developed. Of primary interest is the influence of multiple paternity on the maintenance of genetic variation in a population. In addition to the standard inbreeding and variance effective sizes, intragroup (coancestral) and intergroup effective sizes also are developed. Expressions for effective sizes are developed for the beginning of nonrandom gene exchanges (initial effective sizes), the transition of gene correlations (instantaneous effective sizes), and the steady-state (asymptotic effective size). Results indicate that systems of mating that incorporate more than one male mate per female increase all effective sizes above those expected from polygyny and monogamy. Instantaneous and asymptotic sizes can be expressed relative to the fixation indices. The parameters presented herein can be utilized in models of effective sizes for the study of evolutionary biology and conservation genetics.

Algorithms↗
PubMed1995

Some effects of selection when there is partial full-sib mating.

If there is selection only for viability between zygote formation and adulthood, the frequency of a particular allele changes between these two stages of life. With complete mating this is all that happens, but if there is a positive probability that full sibs mate, there is an extra change between adulthood and the appearance of zygotes in the next generation. This occurs because there are then correlated frequencies of the alleles carried by the mates. An expression for the change in the frequency of an allele, which incorporates these two effects, is derived, and the result is found to be consistent with earlier work by the author on the probability of survival of a rare allele in a large population. The result is inconsistent with the usual expression for the change in frequency of an allele when there is partial inbreeding because that expression does not incorporate the second change in frequency within one generation.

Gene Frequency↗
PubMed1998

Heterosis for viability, fecundity, and male fertility in Drosophila melanogaster: comparison of mutational and standing variation.

If genetic variation for fitness traits in natural populations ("standing" variation) is maintained by recurrent mutation, then quantitative-genetic properties of standing variation should resemble those of newly arisen mutations. One well-known property of standing variation for fitness traits is inbreeding depression, with its converse of heterosis or hybrid vigor. We measured heterosis for three fitness traits, pre-adult viability, female fecundity, and male fertility, among a set of inbred Drosophilia melanogaster lines recently derived from the wild, and also among a set of lines that had been allowed to accumulate spontaneous mutations for over 200 generations. The inbred lines but not the mutation-accumulation (MA) lines showed heterosis for pre-adult viability. Both sets of lines showed heterosis for female fecundity, but heterosis for male fertility was weak or absent. Crosses among a subset of the MA lines showed that they were strongly differentiated for male fertility, with the differences inherited in autosomal fashion; the absence of heterosis for male fertility among the MA lines was therefore not caused by an absence of mutations affecting this trait. Crosses among the inbred lines also gave some, albeit equivocal, evidence for male fertility variation. The contrast between the results for female fecundity and those for male fertility suggests that mutations affecting different fitness traits may differ in their average dominance properties, and that such differences may be reflected in properties of standing variation. The strong differentiation among the MA lines in male fertility further suggests that mutations affecting this trait occur at a high rate.

Alleles↗