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The use of genetic drive mechanisms to replace native mosquito genotypes with individuals bearing antipathogen transgenes is a potential strategy for repressing insect transmission of human diseases such as malaria and dengue. Antipathogen transgenes have been developed and tested, but efficient gene drive mechanisms are lacking. Here we theoretically assess the feasibility of introducing antipathogen genes into wild Aedes aegypti populations by using a naturally occurring meiotic drive system. We consider the release of males having both a Y-linked meiotic drive gene and an X-linked drive-insensitive response allele to which an antipathogen gene is linked. We use mathematical models and computer simulations to determine how the post-introduction dynamics of the antipathogen gene are affected by specific genetic characteristics of the system. The results show that when the natural population is uniformly sensitive to the meiotic drive gene, the antipathogen gene may be driven close to fixation if the fitness costs of the drive gene, the insensitive response allele, and the antipathogen gene are low. However, when the natural population has a small proportion of an X-linked insensitive response allele or an autosomal gene that strongly reduces the effect of the drive gene, the antipathogen gene does not spread if it has an associated fitness cost. Our modeling results provide a theoretical foundation for further experimental tests.
Evolvability is a key characteristic of any evolving system, and the concept of evolvability serves as a unifying theme in a wide range of disciplines related to evolutionary theory. The field of quantitative genetics provides a framework for the exploration of evolvability with the promise to produce insights of global importance. With respect to the quantitative genetics of biological systems, the parameters most relevant to evolvability are the G-matrix, which describes the standing additive genetic variances and covariances for a suite of traits, and the M-matrix, which describes the effects of new mutations on genetic variances and covariances. A population's immediate response to selection is governed by the G-matrix. However, evolvability is also concerned with the ability of mutational processes to produce adaptive variants, and consequently the M-matrix is a crucial quantitative genetic parameter. Here, we explore the evolution of evolvability by using analytical theory and simulation-based models to examine the evolution of the mutational correlation, rμ, the key parameter determining the nature of genetic constraints imposed by M. The model uses a diploid, sexually reproducing population of finite size experiencing stabilizing selection on a two-trait phenotype. We assume that the mutational correlation is a third quantitative trait determined by multiple additive loci. An individual's value of the mutational correlation trait determines the correlation between pleiotropic effects of new alleles when they arise in that individual. Our results show that the mutational correlation, despite the fact that it is not involved directly in the specification of an individual's fitness, does evolve in response to selection on the bivariate phenotype. The mutational variance exhibits a weak tendency to evolve to produce alignment of the M-matrix with the adaptive landscape, but is prone to erratic fluctuations as a consequence of genetic drift. The interpretation of this result is that the evolvability of the population is capable of a response to selection, and whether this response results in an increase or decrease in evolvability depends on the way in which the bivariate phenotypic optimum is expected to move. Interestingly, both analytical and simulation results show that the mutational correlation experiences disruptive selection, with local fitness maxima at –1 and 1. Genetic drift counteracts the tendency for the mutational correlation to persist at these extreme values, however. Our results also show that an evolving M-matrix tends to increase stability of the G-matrix under most circumstances. Previous studies of G-matrix stability, which assume nonevolving M-matrices, consequently may overestimate the level of instability of G relative to what might be expected in natural systems. Overall, our results indicate that evolvability can evolve in natural systems in a way that tends to result in alignment of the G-matrix, the M-matrix, and the adaptive landscape, and that such evolution tends to stabilize the G-matrix over evolutionary time.
In this paper, we present results of the first comprehensive study of the introgression of both autosomal and sex-chromosome markers across the central European portion of the hybrid zone between two house mouse subspecies, Mus musculus musculus and M. m. domesticus. More than 1800 individuals sampled from 105 sites were analyzed with a set of allozyme loci (hopefully representing neutral or nearly neutral markers) and X-linked loci (which are assumed to be under selection). The zone center is best modeled as a single straight line independent of fine-scale local geographic or climatic conditions, being maintained by a balance between dispersal and selection against hybrids. The width (w) of the multilocus autosomal cline was estimated as 9.6 km whereas the estimate for the compound X-chromosome cline was about 4.6 km only. As the former estimate is comparable to that of the Danish portion of the zone (assumed to be much younger than the central European one), zone width does not appear to be related to its age. The strength (B) of the central barrier was estimated as about 20 km; with dispersal (σ) of about 1 km/gen1/2, this means effective selection (s*) is approximately 0.06–0.09 for autosomal loci and about 0.25 for X-linked loci. The number of loci under selection was estimated as N = 56–99 for autosomes and about 380 for X-linked loci. Finally, we highlight some potential pitfalls in hybrid zone analyses and in comparisons of different transects. We suggest that conclusions about parts of the mouse genome involved in reproductive isolation and speciation should be drawn with caution and that analytical approaches always providing some estimates should not be used without due care regarding the support or confidence of such estimates, especially if conclusions are based on the difference between these estimates. Finally, we recommend that analysis in two-dimensional space, dense sampling, and rigorous treatment of data, including inspection of likelihood profiles, are essential for hybrid zone studies.
Gamete-recognition proteins often evolve rapidly, but it is not known if their divergence occurs within species and corresponds with the evolution of reproductive isolation, or if divergence typically accumulates between already isolated lineages. We examined the evolution of a candidate gamete-recognition protein in several sympatric and allopatric populations of Mytilus blue mussels, species that hybridize in nature. Within a single species, Mytilus galloprovincialis, we found adaptive divergence of Lysin-M7, a sperm acrosomal protein that dissolves the egg vitelline envelope during fertilization. Mytilus galloprovincialis Lysin-M7 alleles group into two distinct clades (termed G and GD), and individual alleles in these clades are separated from each other by at least three and up to eleven amino-acid substitutions. Maximum-likelihood estimates of selective pressure (dN/dS = ω) implicate selection in the divergence between M. galloprovincialis Lysin-M7 clades, and within the GD clade. Exact tests of population differentiation indicate that the relative frequency of G and GD Lysin-M7 alleles differs significantly among M. galloprovincialis populations. Compared with allopatric Mediterranean samples, Lysin-M7 alleles in the GD clade are found at elevated frequency in samples from the East Atlantic and California, areas of secondary contact and hybridization between Mytilus species, and Australia, an area of unknown species composition. Adaptive divergence between the alleles most common in allopatry and those found at elevated frequency in samples from sympatry suggests that selection pressures acting in hybridizing populations, likely following Pleistocene secondary contact with M. edulis in the East Atlantic, drove the divergence of Lysin-M7 in M. galloprovincialis.
Speciation of plant-feeding insects is typically associated with host-plant shifts, with subsequent divergent selection and adaptation to the ecological conditions associated with the new plant. However, a few insect groups have apparently undergone speciation while remaining on the same host-plant species, and such radiations may provide novel insights into the causes of adaptive radiation. We used mitochondrial and nuclear DNA to infer a phylogeny for 14 species of gall-inducing Asphondylia flies (Diptera: Cecidomyiidae) found on Larrea tridentata (creosote bush), which have been considered to be monophyletic based on morphological evidence. Our phylogenetic analyses provide strong support for extensive within-host plant speciation in this group, and it demonstrates that diversification has involved numerous shifts between different plant organs (leaves, buds, flowers, and stems) of the same host-plant species. Within-plant speciation of Asphondylia is thus apparently facilitated by the opportunity to partition the plant ecologically. One clade exhibits temporal isolation among species, which may have facilitated divergence via allochronic shifts. Using a novel method based on Bayesian reconstruction, we show that the rate of change in an ecomorphological trait, ovipositor length, was significantly higher along branches with inferred shifts between host-plant organs than along branches without such shifts. This finding suggests that Larrea gall midges exhibit close morphological adaptation to specific host-plant parts, which may mediate ecological transitions via disruptive selection.
A substantial body of theory indicates that parasites may mould the population genetic structure of their hosts, but few empirical studies have directly linked parasitism to genetic dynamics. We used molecular markers (allozymes) to investigate genotype frequency changes in a natural population of the crustacean Daphnia magna in relation to an epidemic of the bacterial pathogen Pasteuria ramosa. The population experienced a severe epidemic during the study period in which parasite prevalence reached 100% of the adult portion of the population. The parasite epidemic was associated with genetic change in the host population. Clonal diversity was observed to decrease as parasite prevalence increased in the population, and tests for differences in the clonal composition of the population before, during, and after the epidemic indicated that significant change had occurred. A laboratory infection experiment showed that the genotypes which were more common following the peak of the parasite epidemic were also the most resistant to parasite infection. Thus, this study provides an illustration of parasite-mediated selection in the wild.
Understanding the architecture of genetic variation, that is the number, effect, location, and interaction, of genes responsible for phenotypic variability in nature is important for the understanding of microevolutionary processes. In this study, we have used a quantitative trait loci (QTL) approach to uncover the genetic architecture of fitness-relevant traits associated with reproduction and immune defense in male Bombus terrestris bumblebees. Three male reproductive investment traits, the number and length of the produced sperm and the size of the accessory glands, were studied. Two branches of the insect innate immune system, the activation of the Phenoloxidase-cascade and the hemolymph's antibacterial activity, were investigated. We found that variation in most of the studied traits is based on a network of minor QTLs and epistatic interactions. Unexpectedly, there was no evidence for phenotypic or genetic trade-offs between the presumably costly investment in immune defense and reproductive effort in this population for the measured traits. In fact, we found a positive correlation, both, in phenotype and genetic architecture for the number of produced sperm and antibacterial activity against an insect pathogen. A major finding for all traits analyzed was that the epistatic interactions accounted for a major proportion of the explained phenotypic variance. Especially for traits involved in immune defense, this pattern highlights the possible role of parasites in the evolution and maintenance of recombination and sexual reproduction.
Spermatozoa exhibit taxonomically widespread patterns of divergent morphological evolution. However, the adaptive significance of variation in sperm morphology remains unclear. In this study we examine the role of natural variation in sperm length on fertilization success in the dung beetle Onthophagus taurus. We conducted sperm competition trials between males that differed in the length of their sperm and determined the paternity of resulting offspring using amplified fragment length polymorphism (AFLP) markers. We also quantified variation in the size and shape of the female's sperm storage organ to determine whether female morphology influenced the competitiveness of different sperm morphologies. We found that fertilization success was biased toward males with relatively shorter sperm, but that selection on sperm length was dependent on female tract morphology; selection was directional for reduced sperm length across most of the spermathecal size range, but stabilizing in females with the smallest spermathecae. Our data provide empirical support for the theory that sperm competition should favor the evolution of numerous tiny sperm. Moreover, because sperm length is both heritable and genetically correlated with condition, our results are consistent with a process by which females can accrue genetic benefits for their offspring from the incitement of sperm competition and/or cryptic female choice, as proposed by the “sexy sperm” and “good sperm” models for the evolution of polyandry.
Male genital morphology in insects and arachnids is characterized by static hypoallometry and low intrapopulational levels of phenotypic variation relative to other male traits. The one-size-fits-all model of genital evolution attributes these patterns to stabilizing sexual selection. This model relies on the assumption that the observed patterns of variation and allometry reflect the form of sexual selection acting these traits. We test this by examining the patterns of scaling and trait variation for a set of genitalic and somatic morphological traits in male water striders (Aquarius remigis). This suite of traits is of particular interest because previous work has shown that the genitalic traits are under strong directional selection whereas the somatic traits are under either weak directional or stabilizing selection. Because the selection regime for these traits is known, we can, for the first time, test the purported relationship between trait variation, scaling, and the form of sexual selection. We show that the patterns of variation and scaling of these traits differ sharply from those predicted for traits experiencing strong directional sexual selection. Specifically, the male genital structures show static hypoallometry and low intrapopulational levels of phenotypic variation relative to other male traits, in spite of consistent, strong, directional sexual selection. These scaling relationships and levels of variation are typical of genital traits in other insect species, where they have been presumed to reflect stabilizing sexual selection. Our data clearly refute the assumption of the one-size-fits-all hypothesis that hypoallometric scaling of genitalic traits implies stabilizing selection. We discuss the implications of this finding and propose future directions for improving our current understanding of genital evolution in arthropods.
One of the most pervasive ideas in the sexual selection literature is the belief that sexually selected traits almost universally exhibit positive static allometries (i.e., within a sample of conspecific adults, larger individuals have disproportionally larger traits). In this review, I show that this idea is contradicted by empirical evidence and theory. Although positive allometry is a typical attribute of some sexual traits in certain groups, the preponderance of positively allometric sexual traits in the empirical literature apparently results from a sampling bias reflecting a fascination with unusually exaggerated (bizarre) traits. I review empirical examples from a broad range of taxa illustrating the diversity of allometric patterns exhibited by signal, weapon, clasping and genital traits, as well as nonsexual traits. This evidence suggests that positive allometry may be the exception rather than the rule in sexual traits, that directional sexual selection does not necessarily lead to the evolution of positive allometry and, conversely, that positive allometry is not necessarily a consequence of sexual selection, and that many sexual traits exhibit sex differences in allometric intercept rather than slope. Such diversity in the allometries of secondary sexual traits is to be expected, given that optimal allometry should reflect resource allocation trade-offs, and patterns of sexual and viability selection on both trait size and body size. An unbiased empirical assessment of the relation between sexual selection and allometry is an essential step towards an understanding of this diversity.
Both song and color patterns in birds are thought to evolve rapidly and exhibit high levels of homoplasy, yet few previous studies have compared the evolution of these traits systematically using the same taxa. Here we reconstruct the evolution of song in the New World orioles (Icterus) and compare patterns of vocal evolution to previously reconstructed patterns of change in plumage evolution in this clade. Individual vocal characters exhibit high levels of homoplasy, reflected in a low overall consistency index (CI = 0.27) and retention index (RI = 0.35). Levels of lability in song are comparable to those found for oriole plumage patterns using the same taxa (CI = 0.31, RI = 0.63), but are strikingly dissimilar to the conservative patterns of change seen in the songs of oropendolas (Psarocolius, Ocyalus; CI = 0.82, RI = 0.87), a group closely related to the orioles. Oriole song is also similar to oriole plumage in exhibiting repeated convergence in overall patterns, with some distantly related taxa sounding remarkably similar. Thus, both song and plumage in orioles show repeated convergence in individual elements and in overall patterns across the clade, suggesting that both of these character classes are highly labile between taxa yet highly conserved within the genus. Our results provide new insights into the tempo and mode of evolution in sexually selected traits within and across clades.
We construct a species-level phylogeny for the Pentaschistis clade based on chloroplast DNA, from the following regions: trnL-F, trnT-L, atpB-rbcL, rpL16, and trnD-psbA. The clade comprises 82 species in three genera, Pentaschistis, Pentameris, and Prionanthium. We demonstrate that Prionanthium is nested in Pentaschistis and that this clade is sister to a clade of Pentameris plus Pentaschistis tysonii. Forty-three of the species in the Pentaschistis clade have multicellular glands and we use ancestral character state reconstruction to show that they have been gained twice or possibly once, and lost several times. We suggest that the maintenance, absence, loss, and gain of glands are correlated with leaf anatomy type, and additionally that there is a difference in the degree of diversification of lineages that have these different character combinations. We propose that both glands and sclerophyllous leaves act as defense systems against herbivory, and build a cost/benefit model in which multicellular glands or sclerophyllous leaves are lost when the alternative defense system evolves. We also investigate the association between leaf anatomy type and soil nutrient type on which species grow. There is little phylogenetic constraint in soil nutrient type on members of the Pentaschistis clade, with numerous transitions between oligotrophic and eutrophic soils. However, only orthophyllous-leaved species diversify on eutrophic soils. We suggest that the presence of these glands enables the persistence of orthophyllous lineages and therefore diversification of the Pentaschistis clade on eutrophic as well as oligotrophic soils.
Lineage persistence is as central to biology as evolutionary change. Important questions regarding persistence include: why do some lineages outlive their relatives, neither becoming extinct nor evolving into separate lineages? Do these long-duration lineages have distinctive ecological or morphological traits that correlate with their geologic durations and potentially aid their survival? In this paper, I test the hypothesis that lineages (species and higher taxa) with longer geologic durations have morphologies that are more average than expected by chance alone. I evaluate this hypothesis for both individual lineages with longer durations and groups of lineages with longer durations, using more than 60 published datasets of animals with adequate fossil records. Analyses presented here show that groups of lineages with longer durations fall empirically into one of three theoretically possible scenarios, namely: (1) the morphology of groups of longer duration lineages is closer to the grand average of their inclusive group, that is, their relative morphological distance is smaller than expected by chance alone, when compared with rarified samples of their shorter duration relatives (a negative group morpho-duration distribution); (2) the relative morphological distance of groups of longer duration lineages is no different from rarified samples of their shorter duration relatives (a null group morpho-duration distribution); and (3) the relative morphological distance of groups of longer duration lineages is greater than expected when compared with rarified samples of their shorter duration relatives (a positive group morpho-duration distribution). Datasets exhibiting negative group morpho-duration distributions predominate. However, lineages with higher ranks in the Linnean hierarchy demonstrate positive morpho-duration distributions more frequently. The relative morphological distance of individual longer duration lineages is no different from that of rarified samples of their shorter duration relatives (a null individual morpho-duration distribution) for the majority of datasets studied. Contrary to the common idea that very persistent lineages are special or unique in some significant way, both the results from analyses of long-duration lineages as groups and individuals show that they are morphologically average. Persistent lineages often arise early in a group's history, even though there is no prior expectation for this tendency in datasets of extinct groups. The implications of these results for diversification histories and niche preemption are discussed.
Evolutionary constraint results from the interaction between the distribution of available genetic variation and the position of selective optima. The availability of genetic variance in multitrait systems, as described by the additive genetic variance–covariance matrix (G), has been the subject of recent attempts to assess the prevalence of genetic constraints. However, evolutionary constraints have not yet been considered from the perspective of the phenotypes available to multivariate selection, and whether genetic variance is present in all phenotypes potentially under selection. Determining the rank of the phenotypic variance–covariance matrix (P) to characterize the phenotypes available to selection, and contrasting it with the rank of G, may provide a general approach to determining the prevalence of genetic constraints. In a study of a laboratory population of Drosophila bunnanda from northern Australia we applied factor-analytic modeling to repeated measures of individual wing phenotypes to determine the dimensionality of the phenotypic space described by P. The phenotypic space spanned by the 10 wing traits had 10 statistically supported dimensions. In contrast, factor-analytic modeling of G estimated for the same 10 traits from a paternal half-sibling breeding design suggested G had fewer dimensions than traits. Statistical support was found for only five and two genetic dimensions, describing a total of 99% and 72% of genetic variance in wing morphology in females and males, respectively. The observed mismatch in dimensionality between P and G suggests that although selection might act to shift the intragenerational population mean toward any trait combination, evolution may be restricted to fewer dimensions.
We investigate multilocus patterns of differentiation between parental populations of two swallowtail butterfly species that differ at a number of ecologically important sex-linked traits. Using a new coalescent-based approach, we show that there is significant heterogeneity in estimated divergence times among five Z-linked markers, rejecting a purely allopatric speciation model. We infer that the Z chromosome is a mosaic of regions that differ in the extent of historical gene flow, potentially due to isolating barriers that prevent the introgression of species-specific traits that result in hybrid incompatibilities. Surprisingly, a candidate region for a strong barrier to introgression, Ldh, does not show a significantly deeper divergence time than other markers on the Z chromosome. Our approach can be used to test alternative models of speciation and can potentially assign chronological order to the appearance of factors contributing to reproductive isolation between species.
The history of repeated inter- or transoceanic invasions in bivalve mollusks of the circumpolar Macoma balthica complex was assessed from mtDNA COIII sequences. The data suggest that four independent trans-Arctic invasions, from the Pacific, gave rise to the current lineage diversity in the North Atlantic. Unlike in many other prominent North Atlantic littoral taxa, no evidence for (postinvasion) trans-Atlantic connections was found in the M. balthica complex. The earliest branch of the mtDNA tree is represented by the temperate-boreal North American populations (= Macoma petalum), separated from the M. balthica complex proper in the Early Pliocene at latest. The ensuing trans-Arctic invasions established the North European M. b. rubra, which now prevails on the North Sea and northeast Atlantic coasts, about two million years ago, and the currently northwest Atlantic M. balthica lineage in the Canadian Maritimes, in the Middle Pleistocene. The final reinvasion(s) introduced a lineage that now prevails in a number of North European marginal seas and is still hardly distinguishable from North Pacific mtDNA (M. b. balthica). We used coalescence simulation analyses to assess the age of the latest invasion from the Pacific to the northeast Atlantic. The results refute the hypothesis of recent, human-mediated reintroductions between northeast Pacific and the North European marginal seas in historical times. Yet they also poorly fit the alternative hypotheses of an early postglacial trans-Arctic invasion (< 11 thousand years ago), or an invasion during the previous Eemian interglacial (120 thousand years ago). Divergence time estimates rather fall in the Middle Weichselian before the Last Glacial Maximum, in conflict with the conventional thinking of trans-Arctic biogeographical connections; an early Holocene reinvasion may still be regarded as the most plausible scenario. Today, the most recently invaded Pacific mtDNA lineage is found admixed with the earlier established European Atlantic “rubra” lineage in the Baltic Sea and in Barents Sea populations east of the Varanger peninsula, and it is practically exclusive in the White and Pechora seas. Yet mtDNA does not always constitute an unequivocal taxonomic marker at individual level; the marginal populations represent hybrid swarms of the Atlantic and Pacific lineages in their nuclear genes.
Theory predicts that biogeographic factors should play a central role in promoting population divergence and speciation. Previous empirical studies into biogeography and diversification have been relatively restricted in terms of the geographical area, phylogenetic scope, and the range of biogeographic factors considered. Here we present a global analysis of allopatric phenotypic divergence (measured as subspecies richness) across more than 9600 bird species. The main aim of this study was to examine the extent to which biogeographical factors can explain patterns of phenotypic divergence. Analysis of the taxonomic distribution of subspecies among species suggests that subspecies formation and extinction have occurred at a considerably faster rate than has species formation. However, the observed distribution departs from the expectation under a random birth–death model of diversification. Across 19 phylogenetic trees, we find no significant linear relationship between species age and subspecies richness, implying that species age is a poor predictor of subspecies richness. Both subspecies richness and subspecies diversification rate are found to exhibit low phylogenetic signal, meaning that closely related species do not tend to possess similar numbers of subspecies. As predicted by theory, high subspecies richness was associated with large breeding range size, island dwelling, inhabitation of montane regions, habitat heterogeneity, and low latitude. Of these factors, breeding range size was the variable that explained the most variation. Unravelling whether species that have invaded previously glacial areas have more or fewer subspecies than expected proves to be complicated due to a covariation between the postglacial colonization, latitude, geographic range size, and subspecies richness. However, the effect of postglacial colonization on subspecies richness appears to be small. Mapping the distribution of species' subspecies richness globally reveals geographical patterns that correspond to many of the predictions of the statistical models, but may also reflect geographical variation in taxonomic practice. Overall, we demonstrate that biogeographic models can explain about 30% of the global variation in subspecies richness in birds.
Two issues that have captured the attention of tropical plant evolutionary biologists in recent years are the relative role of long distance dispersal (LDD) over vicariance in determining plant distributions and debate about the extent that Quaternary climatic changes affected tropical species. Propagules of some mangrove species are assumed to be capable of LDD due to their ability to float and survive for long periods of time in salt water. Mangrove species responded to glaciations with a contraction of their range. Thus, widespread mangrove species are an ideal system to study LDD and recolonization in the tropics. We present phylogenetic and phylogeographic analyses based on internal transcribed spacers region (ITS) sequences, chloroplast DNA (cpDNA), and amplified fragment length polymorphisms (AFLPs) of genomic DNA that demonstrate recent LDD across the Atlantic, rejecting the hypothesis of vicariance for the widespread distribution of the black mangrove (Avicennia germinans). Northern latitude populations likely became extinct during the late Quaternary due to frosts and aridification; these locations were recolonized afterward from southern populations. In some low latitude regions populations went extinct or were drastically reduced during the Quaternary because of lack of suitable habitat as sea levels changed. Our analyses show that low latitude Pacific populations of A. germinans harbor more diversity and reveal deeper divergence than Atlantic populations. Implications for our understanding of phylogeography of tropical species are discussed.
A method for estimating the number of founding chromosomes in an isolated population is introduced. The method assumes that n/2 diploid individuals are sampled from a population and that alleles are identified at L unlinked loci. The population is assumed to have been founded T generations in the past by individuals carrying c chromosomes drawn randomly from a known source population, which has also been sampled. If c is small and the population grew rapidly after it was founded, accurate estimates of c can be obtained and those estimates are not sensitive to details of the history of population sizes. If c is larger or the population remained small after it was founded, then estimates of c depend on the history of population sizes. We test the performance of our method on simulated data and demonstrate its use on data from a rainbow trout (Oncorhynchus mykiss) population.
A new hypothesis for the evolution of overproduction of ovules within flowers is proposed: overproduction is a counter-strategy of female seed production in the conflict with males and/or offspring. It is advantageous for females to produce a uniform size of seeds, whereas it is advantageous for fertilized ovules to absorb more resources than this size. If there is a variance in resource absorption ability among fertilized ovules, nonuniform seeds are produced. Then, by overproducing ovules, females should select fertilized ovules with similar resource absorption rates, resulting in seeds of uniform size. A model analysis confirmed that this hypothesis works. In the model, the fertilized ovules of a plant consist of two genotypes that differ in resource absorption rate. I found that overproduction of ovules and selective abortion is advantageous if the difference in the resource absorption rates of the genotypes is large. The new hypothesis is different from the selective abortion hypothesis in that selecting ovules is advantageous even if there are no differences in the genetic quality of resulting seeds.
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