SPERM-MEDIATED GENE FLOW AND THE GENETIC STRUCTURE OF A POPULATION OF THE COLONIAL ASCIDIAN BOTRYLLUS SCHLOSSERI

The genetic structure of populations of sessile and sedentary organisms is often characterized by microgeographic differentiation in gene frequencies and deviations from panmixia. In many terrestrial botanical systems, restricted gene flow via seed and pollen dispersal may have an important role in promoting such genetic patterns. Until recently, however, limited dispersal of the sexual propagules of benthic invertebrates has not been considered to play a comparable role in aquatic systems. Based on paternity analyses in the field using rare allozyme markers, it appears that concentrations of sibling sperm of the sessile, colonial ascidian Botryllus schlosseri decline rapidly within 50 cm of a source colony. In combination with spatially restricted dispersal of brooded larvae, limited dispersal of sperm should enhance the potential for genetic diversification and inbreeding. However, analysis of allelic and genotypic frequencies at three independent, polymorphic allozyme loci using F-statistics provides little evidence for microgeographic variation in gene frequencies. This lack of differentiation can be explained in terms of the absolute number (rather than concentration) of gametes and larvae dispersing from a point source, which—depending on diffusion and geometric assumptions—may actually increase with distance. In contrast to the absence of differentiation, levels of inbreeding are high, even within the confines of 25 times 25–cm quadrats. The absence of genetic diversification and presence of inbreeding caution against inferring levels and causes of gene flow from indirect analysis of genetic structure and, conversely, making predictions about genetic and breeding structure based solely on direct analysis of gene flow.     

LIMITED DISPERSAL AND PROXIMITY-DEPENDENT MATING SUCCESS IN THE COLONIAL ASCIDIAN BOTRYLLUS SCHLOSSERI

Although the propagules of many sessile organisms have the capacity to disperse over large distances, dispersal is often spatially restricted. In this paper, I document, using a combination of mark/recapture techniques and histocompatibility assays, dispersal distance of the planktonic larvae of the sessile, colonial sea squirt Botryllus schlosseri. Both of these methods indicate that most larvae remain within a meter of their birthplace. Such limited dispersal should lead to increased matings among relatives, and the potential for inbreeding depression. However, the success of: 1) fertilization, 2) embryogenesis, and 3) larval metamorphosis all decrease as distance between mated colonies increases. The spatial scale over which this decrease in mating success occurs is concordant with the estimates of dispersal distance based on the larval mark/recapture data and histocompatibility assays. Taken together, these results imply that inbreeding depression is not a necessary consequence of limited dispersal and consanguineous matings in B. schlosseri.     

GENETIC AND ENVIRONMENTAL VARIATION IN LIFE-HISTORY TRAITS OF A MONOCARPIC PERENNIAL: A DECADE-LONG FIELD EXPERIMENT

Directional and stabilizing selection tend to deplete additive genetic variance. On the other hand, genetic variance in traits related to fitness could be retained through polygenic mutation, spatially varying selection, genotype-environment interaction, or antagonistic pleiotropy. Most estimates of genetic variance in fitness-related traits have come from laboratory studies, with few estimates of heritability made under natural conditions, particularly for longer lived organisms. Here I estimated additive genetic variance in life-history characters of a monocarpic herb, Ipomopsis aggregata, that lives for up to a decade. Experimental crosses yielded 229 full-sibships nested within 32 paternal half-sibships. More than 5000 offspring were planted as seeds into natural field sites and were followed in most cases through their entire life cycle. Survival showed substantial additive genetic variance (genetic coefficient of variation ≈ 54%). Small differences at seedling emergence were magnified over time, such that the genetic variability in survival was only detectable by tracking the success of offspring for several years starting from seed. In contrast to survival, reproductive traits such as flower number, seeds per flower, and age at flowering showed little or no genetic variability. Despite relatively high levels of additive genetic variation for some life-history characters, high environmental variance in survival resulted in very low heritabilities (0–9%) for all of these characters. Maternal effects were evident in seed mass and remained strong throughout the lengthy vegetative period. No negative genetic correlations between major components of female fitness were detected. Mean corolla width for a paternal family was, however, negatively correlated with the finite rate of increase based on female fitness. That negative correlation could help to maintain additive genetic variance in the face of strong selection through male function for wide corollas.     

MUTATION,SELECTION, AND THE MAINTENANCE OF LIFE-HISTORY VARIATION IN A NATURAL POPULATION

In an effort to provide insight into the role of mutation in the maintenance of genetic variance for life-history traits, we accumulated spontaneous mutations in 10 sets of clonal replicates of Daphnia pulex for approximately 30 generations and compared the variance generated by mutation with the standing level of variation in the wild population. Mutations for quantitative traits appear to arise at a fairly high rate in this species, on the order of at least 0.6 per character per generation, but have relatively small heterozygous effects, changing the phenotype by less than 2.5% of the mean. The mean persistence time of a new mutation affecting life-history/body-size traits is approximately 40 generations in the natural population, which requires an average selection coefficient against new mutations of approximately 3% in the heterozygous state. These data are consistent with the idea that the vast majority of standing genetic variance for life-history characters may be largely a consequence of the recurrent introduction of transient cohorts of mutations that are at least conditionally deleterious and raise issues about the meaning of conventional measures of standing levels of variation for fitness-related traits.     

THE POTENTIAL FOR COEVOLUTION IN A HOST-PARASITOID SYSTEM. I. GENETIC VARIATION WITHIN AN APHID POPULATION IN SUSCEPTIBILITY TO A PARASITIC WASP

For coevolution to occur, there must be genetic variation in each species for traits relevant to their interaction. Here, statistically significant variation in susceptibility to a parasitic wasp was found among pea-aphid clones collected from a single population. In a subset of clones that was tested further, wasps were found to oviposit in aphids from both resistant and susceptible lines, but eggs failed to develop in resistant hosts. Significant genetic variance in susceptibility provides evidence that this aphid population has the potential to evolve resistance in response to selection by one of its major natural enemies. Predictions of an expected response to selection based on the experimental measures of variation and field parasitism rates suggested that there should be a detectable change in susceptibility over the course of a season. However, an experimental comparison of mean susceptibility of clones collected early and late in the summer, a period of several generations, revealed no response to selection by the wasps. Aphids collected late in the season were as susceptible, on the average, as those collected early in the summer. Possible constraints on the response of the aphids to selection by this natural enemy are considered.     

QUANTITATIVE GENETICS OF SIZE,SHAPE, LIFE-HISTORY,AND FRUIT CHARACTERISTICS OF THE SEED-HETEROMORPHIC COMPOSITE HETEROSPERMA PINNATUM. I. VARIATION WITHIN AND AMONG POPULATIONS

We document phenotypic and genetic variation within and among populations of the seed heteromorphic species Heterosperma pinnatum Cav. (Compositae) in the production of seed morphs and in a variety of life-history and morphological characteristics that might be correlated with seed and head traits. Each trait is found to have significant genetic variance in most or, usually, all populations. Significant among-population genetic variation exists for all traits except number of achenes per head and seedling shape, although some traits have much less genetic variation among than within populations. Number and percentage of intermediate achenes per head, total number of achenes per head, and lengths of central and peripheral achenes had little among-population genetic variation compared to within-population variation. Most traits had slightly less genetic variation among than within populations; however, some traits (percentage of central achenes, length of awns, date that the first flowering head opened, date that the first fruiting head opened, and death date) had more among-population genetic variation. The proportions of achene morphs produced had high broad-sense heritabilities and high among-population genetic variance, except in the case of intermediate achenes. All phenological variables had high among-population genetic variation. Within-population heritabilities were high for dates of first flowering head and fruiting head but low for death date and reproductive interval. Family and population means measured in the greenhouse for traits having high broad-sense heritability or among-population genetic variance were closely correlated with field means for the corresponding families or populations. The amounts of phenotypic variation were similar for traits that were measured in both the field and the greenhouse. These lines of evidence suggest that greenhouse results provide reasonable estimates of genetic variation in the field for this species. Numerous studies have reported variation in the proportion of seed morphs for different heteromorphic-seeded species and have discussed adaptive scenarios for the evolution of seed proportions; however, our investigation is one of only a few that have documented the amount of phenotypic and genetic variation within and among populations.     

ENVIRONMENTAL AND GENETIC CONTROL OF BRAIN AND SONG STRUCTURE IN THE ZEBRA FINCH

Birdsong is a classic example of a learned trait with cultural inheritance, with selection acting on trait expression. To understand how song responds to selection, it is vital to determine the extent to which variation in song learning and neuroanatomy is attributable to genetic variation, environmental conditions, or their interactions. Using a partial cross fostering design with an experimental stressor, we quantified the heritability of song structure and key brain nuclei in the song control system of the zebra finch and the genotype‐by‐environment (G × E) interactions. Neuroanatomy and song structure both showed low levels of heritability and are unlikely to be under selection as indicators of genetic quality. HVC, in particular, was almost entirely under environmental control. G × E interaction was important for brain development and may provide a mechanism by which additive genetic variation is maintained, which in turn may promote sexual selection through female choice. Our study suggests that selection may act on the genes determining vocal learning, rather than directly on the underlying neuroanatomy, and emphasizes the fundamental importance of environmental conditions for vocal learning and neural development in songbirds.     

THE POTENTIAL FOR COEVOLUTION IN A HOST-PARASITOID SYSTEM. II. GENETIC VARIATION WITHIN A POPULATION OF WASPS IN THE ABILITY TO PARASITIZE AN APHID HOST

Much of the study of coevolution has focused on the adaptations that have resulted from interactions between species. For reciprocal evolution to occur, there must be genetic variation in each species for traits that directly affect their interaction. Here I report evidence of significant additive genetic variance within a population of parasitic wasps in the ability to successfully parasitize an aphid host. These data, combined with companion work documenting clonal variation in a population of aphids from the same site, provide evidence that within the same population both a host and its parasitoid have the potential for specific and reciprocal genetic interactions.     

PAEDOMORPHOSIS IN AMBYSTOMA TALPOIDEUM: MAINTENANCE OF POPULATION VARIATION AND ALTERNATIVE LIFE-HISTORY PATHWAYS

A “common garden” experiment using artifical ponds was performed to test if differences in frequency of paedomorphosis and metamorphosis among six natural populations of the salamander Ambystoma talpoideum resulted from the drying regime of the aquatic habitat acting as an agent of selection. Our experiment supports the hypothesis of genetic differentiation in the propensity to metamorphose among the populations, but gave mixed evidence that pond-drying regime is the selective force directing evolution of this trait. Some populations appear to have evolved phenotypic plasticity whereas others may have a genetic polymorphism in their propensity to metamorphose as ponds dry.     

UNUSUAL POPULATION GENETICS OF A PARASITIC NEMATODE: mtDNA VARIATION WITHIN AND AMONG POPULATIONS

Very little is known about the distribution of genetic variance within and among populations of parasitic helminths. In this study we used mitochondrial DNA (mtDNA) restriction fragment analysis to describe the population genetic structure of Ostertagia ostertagi, a nematode parasite of cattle, in the United States. Estimates of within-population mtDNA diversity are 5 to 10 times greater than typical estimates reported for species in other taxa. Although populations are genetically differentiated for a key life–history trait, greater than 98% of the total genetic diversity is partitioned within populations, and the geographic distribution of individual mtDNA haplotypes suggests high gene flow among populations.     

SOME POPULATION GENETIC CONSEQUENCES OF COLONY FORMATION AND EXTINCTION: GENETIC CORRELATIONS WITHIN FOUNDING GROUPS

Extinction and recolonization in an island model affects genetic differentiation among subpopulations through a combination of sampling and mixing. We investigate the balance of these forces in a general model of population founding that predicts first the genetic variance among new groups and then the effect of these new groups on the total genetic variance among all populations. We allow for a broad range of types of mixing at the time of colonization and demonstrate the significant effects on differentiation from the probability of common origin of gametes (φ). We further demonstrate that kin-structured founding and inbreeding within populations can have a significant effect on the genetic variance among groups and use these results to make predictions about lineal fission and fusion of populations. These results show that population structure is critically affected by non-equilibrium dynamics and that the properties of new populations, especially founding number, probability of common origin, and kin structure, are vital in our understanding of genetic variation.     

长江下游鳙鱼天然种群的生化遗传变异

近年来,我们在从事鳙鱼同工酶发育遗传学研究的基础上,分析了长江下游鳙鱼天然种群的生化遗传变异性,以便为合理开发和利用鳙鱼天然资源以及建立和保护人工种质资源库及基因库提供理论依据。