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Perspective: Reproductive isolation caused by natural selection against immigrants from divergent habitats.

The classification of reproductive isolating barriers laid out by Dobzhansky and Mayr has motivated and structured decades of research on speciation. We argue, however, that this classification is incomplete and that the unique contributions of a major source of reproductive isolation have often been overlooked. Here, we describe reproductive barriers that derive from the reduced survival of immigrants upon reaching foreign habitats that are ecologically divergent from their native habitat. This selection against immigrants reduces encounters and thus mating opportunities between individuals from divergently adapted populations. It also reduces the likelihood that successfully mated immigrant females will survive long enough to produce their hybrid offspring. Thus, natural selection against immigrants results in distinctive elements of premating and postmating reproductive isolation that we hereby dub "immigrant inviability." We quantify the contributions of immigrant inviability to total reproductive isolation by examining study systems where multiple components of reproductive isolation have been measured and demonstrate that these contributions are frequently greater than those of traditionally recognized reproductive barriers. The relevance of immigrant inviability is further illustrated by a consideration of population-genetic theory, a review of selection against immigrant alleles in hybrid zone studies, and an examination of its participation in feedback loops that influence the evolution of additional reproductive barriers. Because some degree of immigrant inviability will commonly exist between populations that exhibit adaptive ecological divergence, we emphasize that these barriers play critical roles in ecological modes of speciation. We hope that the formal recognition of immigrant inviability and our demonstration of its evolutionary importance will stimulate more explicit empirical studies of its contributions to speciation.

Animals↗

Broad-sense sexual selection, sex gene pool evolution, and speciation.

Studies of sexual selection have traditionally focused on explaining the extreme sexual dimorphism in male secondary sexual traits and elaborate mating behaviors displayed by males during courtship. In recent years, two aspects of sexual selection have received considerable attention in the literature: an extension of the sexual selection concept to other traits (i.e., postcopulatory behaviors, external and internal genital morphology, gametes, molecules), and alternative mechanistic explanations of the sexual selection process (i.e., coevolutionary runaway, good-genes, sexual conflicts). This article focuses on the need for an extension of sexual selection as a mechanism of change for courtship and (or) mating male characters (i.e., narrow-sense sexual selection) to all components of sexuality not necessarily related to courtship or mating (i.e., broad-sense sexual selection). We bring together evidence from a wide variety of organisms to show that sex-related genes evolve at a fast rate, and discuss the potential role of broad-sense sexual selection as an alternative to models that limit speciation to strict demographic conditions or treat it simply as an epiphenomenon of adaptive evolution.

Animals↗

Antibiotic selection pressure resulting in multiple antibiotic resistance and localization of resistance determinants to conjugative plasmids in streptococci.

The plasmid content, antibiotic susceptibilities, and biochemical traits of fecal streptococci isolated from pigs during a tylosin feeding experiment were analyzed. The presence of tylosin in the feed did not select for a particular streptococcal strain or biochemical type but did select for multiple antibiotic resistance. A DNA probe obtained from Escherichia coli and containing a cloned macrolide-lincosamide-streptogramin B (MLS) resistance determinant from a streptococcal strain of human origin hybridized primarily to large plasmids among MLS-resistant streptococci from antibiotic-fed pigs. Patterns of hybridization were similar among MLS-resistant clinical isolates from human infections and isolates from other antibiotic-fed animals. The results of mating experiments indicated that tylosin selected for localization of the MLS determinants to large conjugative plasmids. These data established (1) a strong sequence homology between MLS resistance genes in isolates from humans and those from farm animals and (2) an increased potential for dissemination of MLS resistance as a consequence of tylosin feeding.

Animal Feed↗

Mutations in a gene encoding the alpha subunit of a Saccharomyces cerevisiae G protein indicate a role in mating pheromone signaling.

Mutations which allowed conjugation by Saccharomyces cerevisiae cells lacking a mating pheromone receptor gene were selected. One of the genes defined by such mutations was isolated from a yeast genomic library by complementation of a temperature-sensitive mutation and is identical to the gene GPA1 (also known as SCG1), recently shown to be highly homologous to genes encoding the alpha subunits of mammalian G proteins. Physiological analysis of temperature-sensitive gpa1 mutations suggests that the encoded G protein is involved in signaling in response to mating pheromones. Mutational disruption of G-protein activity causes cell-cycle arrest in G1, deposition of mating-specific cell surface agglutinins, and induction of pheromone-specific mRNAs, all of which are responses to pheromone in wild-type cells. In addition, mutants can conjugate without the benefit of mating pheromone or pheromone receptor. A model is presented where the activated G protein has a negative impact on a constitutive signal which normally keeps the pheromone response repressed.

Agglutinins↗

A mating type-linked gene cluster expressed in Chlamydomonas zygotes participates in the uniparental inheritance of the chloroplast genome.

A characteristic feature of early zygote development in Chlamydomonas is the selective degradation of chloroplast DNA from the mating type minus parent. The zygote-specific gene cluster ezy-1 is linked to the mating type locus and is transcribed almost immediately upon zygote formation. We show here that the acidic Ezy-1 polypeptide is rapidly transported to both the plus and minus chloroplasts, where it interacts with each chloroplast nucleoid. Expression of ezy-1 is selectively inhibited when plus, but not minus, gametes are briefly ultraviolet irradiated just prior to mating, a treatment known to disrupt the uniparental inheritance of chloroplast traits. We propose that the Ezy-1 polypeptide participates in the destruction of the minus chloroplast DNA in zygotes and thus the uniparental inheritance of chloroplast traits. The ezy-1 gene represents a valuable molecular probe for dissecting mechanisms underlying organelle inheritance.

Amino Acid Sequence↗

Methods for evaluating and developing commercial chicken strains free of endogenous subgroup E avian leukosis virus.

The genome of nearly all chickens contains various DNA proviral insertions of retroviruses of subgroup E avian leukosis virus (ALVE). However, the elimination or control of ALVE gene expression is desirable to improve productivity, to improve resistance to avian leukosis virus (ALV)-induced tumours, and to develop safer live virus vaccines in chick embryos and cultured chick cells. Restriction fragment length polymorphism and polymerase chain reaction methods are used to define the presence of ALVE genes; and the expression of ALVE in chicken plasma or on cells, and the susceptibility of cells to ALVE is determined by flow cytometry using a specific (R2) antibody. ADOL line 0 chickens have been selected to be free of ALVE genes, while being resistant (i.e. lack receptors to ALVE), but susceptible to exogenous ALV (i.e. ALVA, ALVB, ALVC and ALVJ). To develop improved line 0-type chickens, ADOL line 0 was outcrossed to a commercial line that had one ALVE gene and evidence for ALVE resistance. Rous sarcoma virus (RSV) challenge was used to confirm resistance of F1 chickens to ALVE, and susceptibility of F2 breeders to ALVA and ALVB using test chicks produced by matings to line 7(2). Selected F2 breeders were resistant to ALVE, but susceptible to exogenous ALVA, ALVB, ALVC and ALVJ, based on challenge tests of progeny chick cells using an enzyme-linked immunosorbent assay. The new line, 0(1), has evidence for improved egg size, productivity, fertility and hatchability. Similar procedures may be used for development of productive ALVE free chicken lines with preferred ALV susceptibility traits.

Animals↗

Optimization of sire selection based on maximization of guaranteed income and risk associated with sire merit.

A method based on discounted income and risk assessment was developed to aid in the selection of dairy sires. The discounted profit generated from milk production of daughters was proposed as the suitable composite selection criterion to combine estimates of predicted transmitting ability (PTA) for yields of milk, fat, and protein and estimates of sire evaluations for dystocia or expected progeny difference. Steps are described to derive discounted profit (defined as expected income) for a sire with known PTA and evaluation for dystocia. The derivation of profit considered costs for semen, dystocia, heifer raising, production and maintenance of the daughter, and income from milk. Variance of income from a sire depended on the reliability of his PTA and evaluation for dystocia. Total variance from a selected set of sires was defined as the risk. A quadratic programming procedure was developed to identify the best set of sires from a given pool of sires that met a desired expected income goal with minimum risk. Combinations of sires with minimum risk for all possible levels of expected income were defined by the expected income variance frontier. The set of sires at the maximum lower boundary for 95% confidence of the expected income variance frontier was defined as the optimum set of sires to be selected; the optimum set maximized the 95% guaranteed expected income. The quadratic programming procedure provided the optimum percentage of cows to be mated to each sire in the selected set.

Animals↗

SELECTION OF A STRAIN OF ALBINO MICE REFRACTORY TO PLASMODIUM BERGHEI INFECTION.

Some features of the natural history of malaria in holoendemic areas can be explained by the presence of acquired immunity. There is also evidence that transmission of humoral antibodies in utero from the immune mother plays some part in the transient protection of the offspring. Little is known about the importance of a genetic factor.The possibility of inheritance of immunity to malaria was investigated on six generations of a strain of white mice originating from 50 parents fully susceptible to infection with Plasmodium berghei. These mice were interbred in such a way that each generation was exposed to an infection attenuated by chloroquine. The survivors were mated with a view to selecting a strain with a decreased susceptibility to infection. Some 16% of mice in the sixth generation and one animal out of four in the seventh were able to resist the initial inoculation with 0.1 million parasites. By the sixth generation, there was evidence of reduction of the period of primary parasitaemia and lengthening of the prepatent period, suggesting that some selection of animals with greater tolerance had occurred. A decrease of fertility, particularly obvious in the seventh generation, limited the number of surviving animals and the experiment had to be discontinued.

Animals↗

Chromosomal polymorphism, morphological traits and male mating success in Leptysma argentina (Orthoptera).

Leptysma argentina (Acrididae: Orthoptera) is polymorphic for a centric fusion between pairs 3 and 6 of its basic chromosomal complement. With the aim of carrying out an analysis of selection components in this species, male mating success, in relation to karyotype and three morphometric traits, was assessed by using experimental contests with four males and one female per mating cage. Karyotype was assessed in terms of fused chromosome 3/6 dosage and coded as "0", "1" and "2" for unfused homozygotes, heterozygotes and fusion homozygotes, respectively. The individual ANOVA showed significant differences between "successful" and "unsuccessful" males in all four measured characters. Fusion carriers showed a higher relative fitness. Since the phenotypic correlations among traits were significant, the selective effects were assessed by means of a multivariate analysis, which demonstrated that selection acted significantly on femur length only, selection on karyotype being indirect. Given that the fusion frequency has been constant since 1985, the presence of selective trade-offs cannot be ruled out.

Animals↗

Intrasexual competition and canine dimorphism in anthropoid primates.

A number of factors, including sexual selection, body weight, body-weight dimorphism, predation, diet, and phylogenetic inertia have been proposed as influences on the evolution of canine dimorphism in anthropoid primates. Although these factors are not mutually exclusive, opinions vary as to which is the most important. The role of sexual selection has been questioned because mating system, which should reflect its strength, poorly predicts variation in canine dimorphism, particularly among polygynous species. Kay et al. (1988) demonstrate that a more refined estimate of intermale competition explains a large proportion of the variation in canine dimorphism in platyrrhine primates. We expand their analysis, developing a more generalized measure of intermale competition based on the frequency and intensity of male-male agonism. We examine the relative influences of predation (inferred by substrate use), female body weight, body-weight dimorphism, diet, and sexual selection on the evolution of anthropoid canine dimorphism. Intermale competition is very strongly associated with canine dimorphism. Predation also has a marked effect on canine dimorphism, in that savanna-dwelling species consistently show greater canine dimorphism than other species, all other factors being held equal. Body-weight dimorphism is also strongly associated with canine dimorphism, though apparently through a common selective basis, rather than through allometric effects. Body weight seems to play only a minor, indirect role in the evolution of canine dimorphism. Diet plays no role. Likewise, we find little evidence that phylogenetic inertia is a constraint on the evolution of canine dimorphism.

Analysis of Variance↗

Male Drosophila melanogaster have higher mating success when adapted to their thermal environment.

Adaptation to new environments is a well-documented phenomenon. Individuals from populations maintained in a particular environment for multiple generations tend to be better able to survive and/or reproduce in that environment than their ancestors or other individuals adapted to alternative environments. A third major component of fitness, mating success, has not been well studied in replicated populations under selection in divergent environments. In this study, we used mating trials to compare the mating success of male Drosophila melanogaster adapted for 10 years to two different temperatures, 18 and 25 degrees C. In competition for female partners, males had significantly higher mating success at their adapted temperature compared with males adapted to a different temperature. These results are consistent with the notion that those mutations favoured by natural selection also tend to be favoured by sexual selection.

Adaptation, Physiological↗

The evolution of multiple mating behavior by honey bee queens (Apis mellifera L.).

A model is presented showing that natural selection operating at the individual level can adequately explain the evolution of multiple mating behavior by honey bee queens. Group selection need not be invoked. The fitness of a given female genotype is a function of the number of sex alleles in the population, the number of matings by an individual female and the specific parameters that determine the relationship of brood viability to individual fitness. Even though the exact relationship is not known, it is almost certainly not linear. A nonlinear relationship between worker brood viability and fitness and a significant genetic load associated with the sex-determination system in honey bees are the essential components of this model.

Animals↗

Screening for highly heterozygous chickens in outbred commercial broiler lines to increase detection power for mapping quantitative trait loci.

Two commercial broiler lines were used in an experiment to map quantitative trait loci (QTL) affecting disease resistance. Chickens from these lines were genotyped with 27 microsatellite markers to estimate heterozygosity and polymorphism information content (PIC), the probability that one parent is heterozygous at a marker locus and the other has a different genotype. Heterozygosity estimated from allelic frequencies was 0.52 for the two lines; however, heterozygosity calculated from actual counts of heterozygous loci was much lower (0.36) than the estimated heterozygosity. The PIC was 0.45 in these lines, and average allele number per marker locus was about 3.5. Twenty-five males produced from a cross between these two lines were screened with the DNA markers to select birds with high heterozygosity at marker loci. Mating simulation showed that uninformative matings could be reduced by about 5% if 12 pairs of males and females with the highest heterozygosity at marker loci were selected, which was about a 25% reduction in total uninformative matings. This experiment demonstrated that the heterozygosity and PIC in commercial broiler lines were low and selection for the birds with high heterozygosity at marker loci could increase informative content in chickens used in the experiments of QTL mapping, thus increasing detection power for QTL mapping.

Animal Husbandry↗

Genetic benefits of a female mating preference in gray tree frogs are context-dependent.

"Good genes" models of sexual selection predict that male courtship displays can advertise genetic quality and that, by mating with males with extreme displays, females can obtain genetic benefits for their offspring. However, because the relative performance of different genotypes can vary across environments, these genetic benefits may depend on the environmental context; in which case, static mating preferences may not be adaptive. To better understand how selection acts on the preference that female gray tree frogs (Hyla versicolor) express for long advertisement calls, I tested for genetic benefits in two realistic natural environments, by comparing the performance of half-sibling offspring sired by males with long versus short calls. Tadpoles from twelve such maternal half-sibships were raised in enclosures in their natal pond at two densities. In the low-density treatment, offspring of long-call males were larger at metamorphosis than were offspring of short-call males, whereas in the high-density treatment, offspring of males with long calls tended to metamorphose later than offspring of males with short calls. Thus, although the genes indicated by long calls were advantageous under low-density conditions, they were not beneficial under all conditions, suggesting that a static preference for long calls may not be adaptive in all environments. Such a genotype-by-environment interaction in the genetic consequences of mate choice predicts that when the environment is variable, selection may favor plasticity in female preferences or female selectivity among environments to control the conditions experienced by the offspring.

Animals↗

Palliating the impact of fixation of a major gene on the genetic variation of artificially selected polygenes.

Selective sweeps of variation caused by fixation of major genes may have a dramatic impact on the genetic gain from background polygenic variation, particularly in the genome regions closely linked to the major gene. The response to selection can be restrained because of the reduced selection intensity and the reduced effective population size caused by the increase in frequency of the major gene. In the context of a selected population where fixation of a known major gene is desired, the question arises as to which is the optimal path of increase in frequency of the gene so that the selective sweep of variation resulting from its fixation is minimized. Using basic theoretical arguments we propose a frequency path that maximizes simultaneously the effective population size applicable to the selected background and the selection intensity on the polygenic variation by minimizing the average squared selection intensity on the major gene over generations up to a given fixation time. We also propose the use of mating between carriers and non-carriers of the major gene, in order to promote the effective recombination between the major gene and its linked polygenic background. Using a locus-based computer simulation assuming different degrees of linkage, we show that the path proposed is more effective than a similar path recently published, and that the combination of the selection and mating methods provides an efficient way to palliate the negative effects of a selective sweep.

Computer Simulation↗

The conflict theory of genomic imprinting: how much can be explained?

In some mammalian genes, paternally and maternally derived alleles are expressed differently: this phenomenon is called genomic imprinting. Several-explanations have been proposed for the observed patterns of genomic imprinting, but the most successful explanation is the genetic conflict hypothesis--natural selection operating on the gene expression produces the parental origin-dependent gene expression--because the paternally derived allele tends to be less related to the siblings of the same mother than the maternal allele and hence the paternal allele should evolve to be more aggressive in obtaining maternal resources. The successes and failures of this argument have been examined in explaining the observed patterns of genomic imprinting in mammals. After a brief summary of the observations with some examples, a quantitative genetic model describing the evolution of the cis-regulating element of a gene affecting the maternal resource acquisition was presented. The model supports the verbal argument that the growth enhancer should evolve to show imprinting with the paternal allele expressed and the maternal allele inactive, whereas a growth suppressor gene tends to have an inactive paternal allele and an active maternal allele. There are four major problems of the genetic conflict hypothesis. (1) Some genes affect embryonic growth but are not imprinted (e.g., Igf1), which can be explained by considering recessive, deleterious mutations on the coding regions, (2) A gene exists that shows the pattern that is a perfect reversal (Mash2), which is needed for placental growth, and yet has an active maternal allele and an inactive paternal allele. This can be explained if the overproduction of this gene causes dose-sensitive abortion to occur in early gestation. (3) Paternal disomies are sometimes smaller than normal embryos. This is a likely outcome of evolution if imprinted genes control the allocation between placenta and embryo by modifying the cell developmental fate. (4) Genes on X chromosomes do not follow the predictions of the genetic conflict hypothesis. For genes on X chromosomes, two additional forces of natural selection (sex differentiation and dosage compensation) cause genomic imprinting, possibly in the opposite direction. Available evidence suggests that these processes are stronger than the natural selection caused by female multiple mating. Finally, the same formalism of evolution can handle an alternative nonconflict hypothesis: genomic imprinting might have evolved because it reduces the risk of the spontaneous development of parthenogenetic embryo, causing a serious threat to the life of the mother (ovarian time bomb hypothesis). This hypothesis can also explain major patterns of genomic imprinting. In conclusion, the genetic conflict hypothesis is very successful in explaining the observed patterns of imprinting for autosomal genes and probably is the most likely evolutionary explanation for them. However, for genes on X chromosomes, other processes of natural selection are more important. Considering that a nonconflict hypothesis can also explain the patterns in principle, we need a quantitative estimate of various parameters, such as the rate of dose-dependent abortion, the degree of female promiscuity, and the rate of spontaneous development of the parthenogenetic embryo, in order to make judgments on the relative importance of different forces of natural selection to form genomic imprinting.

Abortion, Veterinary↗

Genetic quality and sexual selection: an integrated framework for good genes and compatible genes.

Why are females so choosy when it comes to mating? This question has puzzled and marveled evolutionary and behavioral ecologists for decades. In mating systems in which males provide direct benefits to the female or her offspring, such as food or shelter, the answer seems straightforward--females should prefer to mate with males that are able to provide more resources. The answer is less clear in other mating systems in which males provide no resources (other than sperm) to females. Theoretical models that account for the evolution of mate choice in such nonresource-based mating systems require that females obtain a genetic benefit through increased offspring fitness from their choice. Empirical studies of nonresource-based mating systems that are characterized by strong female choice for males with elaborate sexual traits (like the large tail of peacocks) suggest that additive genetic benefits can explain only a small percentage of the variation in fitness. Other research on genetic benefits has examined nonadditive effects as another source of genetic variation in fitness and a potential benefit to female mate choice. In this paper, we review the sexual selection literature on genetic quality to address five objectives. First, we attempt to provide an integrated framework for discussing genetic quality. We propose that the term 'good gene' be used exclusively to refer to additive genetic variation in fitness, 'compatible gene' be used to refer to nonadditive genetic variation in fitness, and 'genetic quality' be defined as the sum of the two effects. Second, we review empirical approaches used to calculate the effect size of genetic quality and discuss these approaches in the context of measuring benefits from good genes, compatible genes and both types of genes. Third, we discuss biological mechanisms for acquiring and promoting offspring genetic quality and categorize these into three stages during breeding: (i) precopulatory (mate choice); (ii) postcopulatory, prefertilization (sperm utilization); and (iii) postcopulatory, postfertilization (differential investment). Fourth, we present a verbal model of the effect of good genes sexual selection and compatible genes sexual selection on population genetic variation in fitness, and discuss the potential trade-offs that might exist between mate choice for good genes and mate choice for compatible genes. Fifth, we discuss some future directions for research on genetic quality and sexual selection.

Animals↗

Harnessing the biology of the oviduct for the benefit of artificial insemination.

Spermatozoa fulfil a single role, namely achieving syngamy by transporting the haploid genome to their counterpart gamete, the oocyte. Simple as this may seem, it is fraught with many difficulties, especially in the face of biological processes that enable females to select spermatozoa after they have mated multiply with several males. Conversely, the female reproductive tract sequesters a privileged sperm subpopulation in the oviductal isthmus for variable periods of time, releasing them when the time is opportune for fertilisation. Recent studies of sperm transport in the female reproductive tract suggest that these phenomena involve signalling dialogues between spermatozoa and the female reproductive tract environment. Opportunities for mutual signalling are immense but have received relatively little attention. The oviduct is an organ of crucial significance in modulating sperm function and may be one of the most important sites for determining many aspects of sperm selection and competition. The oviductal environment possesses the potential for enhancing sperm survival, suppressing and activating sperm motility as required, and responds to the arrival of spermatozoa by producing novel proteins. While the biological nature of the sperm-oviduct dialogue is interesting for its own sake, the mechanisms that govern these processes offer opportunities for the improvement of artificial insemination procedures. If oviductal proteins enhance sperm survival, they offer opportunities for the development of long-life semen diluents. Conversely, if we understood the basis of sperm selection we may be able to concentrate on identifying and using only the best sperm subpopulations for improved animal breeding efficiency.

Animals↗