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Use of cobalt as a mitochondrial vital stain to study cytoplasmic exchange in matings of the basidiomycete Schizophyllum commune.

Differential labeling of mates, based on the selective uptake of cobalt by mitochondria of one of the partners, has been used to determine visually by means of phase-contrast microscopy whether transfer of mitochondria occurs after hyphal fusion. Compatible and incompatible matings of the tetrapolar basidiomycete, Schizophyllum commune, were studied. Transfer was detectable in common-A, common-AB, and fully compatible matings. It was not detectable in common-B matings

Basidiomycota↗

Directional bias during mating type switching in Saccharomyces is independent of chromosomal architecture.

Haploid Saccharomyces cells have the remarkable potential to change mating type as often as every generation, a process accomplished by an intrachromosomal gene conversion between an expressor locus MAT and one of two repositories of mating type information, HML or HMR. The particular locus selected as donor is dictated by the mating type of the cell, a bias that ensures productive mating type interconversion. Here we use green fluorescent protein tagging of the expressor and donor loci on chromosome III to show that this preference for donor locus does not result from a predetermined organization of chromosome III: HML and MAT as well as HMR and MAT remain separated in cells of both mating types. In fact, cells in which the inappropriate donor locus is artificially tethered to MAT still predominantly select the correct donor. We find, though, that initiation of switching leads to a rapid association of the correct donor locus with MAT. Thus, in mating type switching in Saccharomyces, donor preference is imposed at commitment to recombination rather than at physical contact of interacting DNA strands.

Chromosomes, Fungal↗

Use of repeated matings to estimate environmental and genetic trends and effects of relaxing selection in a selected strain of Leghorn chickens.

Response from index selection for egg production traits in a Leghorn strain over 7 years was partitioned into estimates of total (delta T), temporary (delta R), and net genetic (delta G) response, by producing progeny from selected (A) and unselected (D) first year breeders and from second year breeders (B) and first year full-sisters of previous years A breeders (C). In A, B, C, and D populations, average yearly number of full-sib families was 121, 47, 37, and about 50 and of birds with complete records was 824, 287, 258, and 298. Restricting A to repeated B families added positive bias in delta G for viability, age at first egg, and rate of lay. Estimates of delta G and delta R from B and C populations were similar. Yearly environmental change (delta E) was over three times the phenotypic change (delta P) for egg numbers, emphasizing necessity for intra-year estimates of delta G and delta R. Temporary response (delta R) approached or exceeded total response (delta T) for mortality and accounted for 40% of delta T for rate of lay and 60% for economic index, suggesting adverse maternal environmental effects from relaxing selection (D). Net yearly genetic change (delta G) was 1.5% for rate of lay, $.18 per bird, and in desired direction for most traits. Estimated net yearly response for rate of lay exceeded that predicted from .64 sigma male and .92 sigma female index selection differentials and sire-component genetic parameters (.6%) but agreed ($.16) for economic index. Efficiency and bias for the repeat-mating design are handicapped by mortality and health-related maternal effects.

Animals↗

Male heroin addicts and their female mates: impact on disorder and recovery.

A pilot investigation was undertaken to determine whether the social relationships between men who currently used heroin with their female mates differed from those of men who formerly used heroin, but were currently abstinent, with their female mates. Six couples were selected for study via the representative case method. Men currently abstinent from heroin had begun drug use later, had a family history of affective disorders, and had initiated their conjugal relationship after cessation of heroin use. Female mates of males currently abstinent from heroin had never used heroin with their mates in conjunction with sexual activity, had significantly positive ratings of their mates' performance of social roles, and were striving to obtain formal training to improve their employment skills. In contrast, the couples including actively using heroin addicts used opiates together, especially to enhance their sexual activity. The relationships of these couples were not as mutually supportive, and the male's role was less responsible and attracted less respect than that of the abstinent male. The females were depressed and had little positive aspects to their lives. Partners in all couples experienced a large number of stressful events, but current heroin use increased stress and hampered coping efforts. Interaction analysis of dynamics in relationships of men currently abstinent from heroin revealed that participation in mates' family life served to reshape their behaviors into more socially acceptable roles.

Adult↗

Mating system of Microcebus murinus.

Microcebus murinus, a small nocturnal lemur from Madagascar, has retained features of ancient primates. Based on these ancestral traits, its social organization has often been used as a model for early primate social systems. In captivity it breeds polygynously, i.e., one male mates with several females, while females usually copulate only with the dominant male. The present project tested whether or not sexual size dimorphism, spatial distribution, and relative testis size of M. murinus correspond with predictions of the sexual selection theory concerning polygynous mating systems. The study was combined with a mark-recapture study and radio tracking of 12 animals in 1993 in a dry deciduous forest of western Madagascar at the end of the dry season. Large overlapping home ranges in males, lack of sexual size dimorphism, and relatively large testes suggest a multi-male mating system, i.e., one that is promiscuous rather than polygynous.

Adaptation, Biological↗

Theoretical modifications of reciprocal recurrent selection.

Reciprocal recurrent selection (RRS), which assumes overdominant loci to be important, alters two genetically different populations to improve their crossbred mean. Individual plants from two populations (A and B) are selfed and also crossed with plants from the reciprocal female tester population (B and A, respectively). Selection is based on the mean of crossbred families, and the selected individuals are randomly mated within A and B to form new populations.--We propose two alternatives to RRS. The first (RRS-I) uses, as the tester of population A, a population (LB) that is derived from population B by family selection for low yield. The second (RRS-II) is similar to RRS-I, but also uses, as the tester of B, a population (LA) that is derived from population A by family selection for low yield.--The expected crossbred means of RRS, RRS-I, and RRS-II were compared, assuming equal sigmaP, at several cycles of selection for incomplete and complete dominance, and for several cases of overdominance (depending on the gene frequencies in A and B, and on the equilibrium gene frequency).--The choice of selection method depends on the importance of the effects of overdominant loci compared to loci exhibiting incomplete or complete dominance. If overdominance is unimportant, RRS-II is the best selection method, followed by RRS-I and RRS. If overdominance is important, both RRS and RRS-I are superior to RRS-II; RRS is preferred to RRS-I if the effects of overdominant loci are sufficiently important. If the genetic model is a mixture of levels of dominance at different loci, a combination of selection systems is suggested.

Crosses, Genetic↗

Sympatric speciation by sexual selection: a critical reevaluation.

Several empirical studies put forward sexual selection as an important driving force of sympatric speciation. This idea agrees with recent models suggesting that speciation may proceed by means of divergent Fisherian runaway processes within a single population. Notwithstanding this, the models so far have not been able to demonstrate that sympatric speciation can unfold as a fully adaptive process driven by sexual selection alone. Implicitly or explicitly, most models rely on nonselective factors to initiate speciation. In fact, they do not provide a selective explanation for the considerable variation in female preferences required to trigger divergent runaway processes. We argue that such variation can arise by disruptive selection but only when selection on female preferences is frequency dependent. Adaptive speciation is therefore unattainable in traditional female choice models, which assume selection on female preferences to be frequency independent. However, when frequency-dependent sexual selection processes act alongside mate choice, truly adaptive sympatric speciation becomes feasible. Speciation is then initiated independently of nonadaptive processes and does not suffer from the theoretical weaknesses associated with the current Fisherian runaway model of speciation. However, adaptive speciation requires the simultaneous action of multiple mechanisms, and therefore it occurs under conditions far more restrictive than earlier models of sympatric speciation by sexual selection appear to suggest.

Animals↗

The influence of sexual dimorphism in body size and mouth morphology on diet selection and sexual segregation in cervids.

In mammals patterns of food resource distribution influence female distribution, leading to aggregation and favouring the evolution of a polygynous mating system. Under polygyny, sexual selection favours an increase of the male body size, since larger bodied males have competitive advantage in fights for mates. As a result, sexual body size dimorphism is a general rule in polygynous artiodactyls and is correlated with the degree of polygyny. Sex differences in body size lead to differences in energy requirements and food selection between the sexes. This has led to the sexual size dimorphism hypothesis being used to explain sexual segregation in ungulates, although from the available studies, it is not possible to deduce a consistent pattern between sexes in the use of forage of different abundance or quality. Two other groups of hypotheses have been put forward to explain sexual segregation in ungulates. These are based on reproductive strategy and social factors, both of which are independent of body size. The mechanistic explanation for differences in food selection ability and intake rate between animals of different body size and how this can lead to an understanding of the sex differences in diet and sexual segregation, both of which are intimately linked, is discussed.

Animal Nutritional Physiological Phenomena↗

Development of two isogenic sweet corn hybrids differing for glycinebetaine content.

A hybrid of sweet corn, Zea mays L. (;1720'; Rogers Brothers Seed Co.), was found to be comprised of glycinebetaine-positive and glycinebetaine-deficient individuals in a 1:1 mixture. This phenomenon was traced to segregation for a single, nuclear, dominant gene determining leaf glycinebetaine content within the female inbred parent of this hybrid. Selection for homozygous recessive (glycinebetaine-deficient) and homozygous dominant (glycinebetaine-positive) genotypes of the female inbred parent enabled production of two isogenic versions of hybrid ;1720' differing with respect to a single copy of the dominant allele, by mating these female parent selections with the common homozygous recessive (glycinebetaine-deficient) male parent. These two isogenic hybrids are shown to differ by a factor of 300- to 400-fold in glycinebetaine titer of young expanding leaves of salinized plants, but exhibit no striking differences in the levels of free amino acids or the level of N-methylnicotinic acid (nicotinic acid betaine; trigonelline). The only significant difference between the two hybrids in terms of amino acid composition was found to be in the level of alanine under nonsalinized conditions. The betaine-deficient hybrid exhibited a 14% lower alanine level than the betaine-positive hybrid. Betaine deficiency was not associated with altered stress-induced accumulation of amino acids such as proline, serine, and asparagine plus aspartate, attesting to the high specificity of the genetic difference between these isogenic hybrids with respect to betaine accumulation. This germplasm offers unique opportunities to test whether a single dominant allele determining stress-induced betaine accumulation capacity influences stress resistance in maize.

Journal Article↗

Natural and sexual selection on many loci.

A method is developed that describes the effects on an arbitrary number of autosomal loci of selection on haploid and diploid stages, of nonrandom mating between haploid individuals, and of recombination. We provide exact recursions for the dynamics of allele frequencies and linkage disequilibria (nonrandom associations of alleles across loci). When selection is weak relative to recombination, our recursions provide simple approximations for the linkage disequilibria among arbitrary combinations of loci. We show how previous models of sex-independent natural selection on diploids, assortative mating between haploids, and sexual selection on haploids can be analyzed in this framework. Using our weak-selection approximations, we derive new results concerning the coevolution of male traits and female preferences under natural and sexual selection. In particular, we provide general expressions for the intensity of linkage-disequilibrium induced selection experienced by loci that contribute to female preferences for specific male traits. Our general results support the previous observation that these indirect selection forces are so weak that they are unlikely to dominate the evolution of preference-producing loci.

Animals↗

Frequency dependence in matings with water-borne sperm.

Negative frequency-dependent mating success--the rare male effect--is a potentially powerful evolutionary force, but disagreement exists as to whether previous work, focusing on copulating species, has robustly demonstrated this phenomenon. Noncopulating sessile organisms that release male gametes into the environment but retain their eggs for fertilization may routinely receive unequal mixtures of sperm. Although promiscuity seems unavoidable it does not follow that the resulting paternity obeys 'fair raffle' expectations. This study investigates frequency dependence in the mating of one such species, the colonial ascidian Diplosoma listerianum. In competition with an alternative sperm source males fathered more progeny if previously mated to a particular female than if no mating history existed. This suggests positive frequency-dependent selection, but may simply result from a mate order effect involving sperm storage. With fewer acclimation matings, separated by longer intervals, this pattern was not found. When, in a different experimental design, virgin females were given simultaneous mixtures of gametes at widely divergent concentrations, sperm at the lower frequency consistently achieved a greater than expected share of paternity--a rare male effect. A convincing argument as to why D. listerianum should favour rare sperm has not been identified, as sperm rarity is expected to correlate very poorly with ecological or genetic male characteristics in this pattern of mating. The existence of nongenetic female preferences at the level of colony modules, analogous in effect to fixed female preferences, is proposed. If visible to selection, indirect benefits from increasing the genetic diversity of a sibship appear the only likely explanation of the rare male effect in this system as the life history presents virtually no costs to multiple mating, and a near absence of direct (resource) benefits, whereas less controversial hypotheses of female promiscuity (e.g. trade up, genetic incompatibility) do not seem appropriate.

Animals↗

Reinforcement of genetic coherence: a single-locus model.

Genetic coherence and genetic separation are the outcome of evolutionary mechanisms which maintain genetic variation within populations through recombination on the one hand, and which divide this variation via speciation between reproductively (recombinatorically) more or less isolated populations on the other. While mechanisms of speciation have received considerable attention in biology, their counterpart, mechanisms of genetic coherence, are addressed only implicitly, if at all. Usually, genetic coherence is intuitively associated with the forces maintaining genetic polymorphisms and thus potential for flexible adaptational reaction of populations. However, so far no models seem to exist which explain the evolution of genetic coherence as the natural counterpart of genetic separation or speciation. In this paper a single-locus model is analyzed, in which a mutant allele is introduced into a resident stable diallelic polymorphism, and where this allele is equivalent to one of the resident alleles in all respects with the exception of mating relations. The conditions for replacement of the resident allele by its selectively equivalent mutant are obtained with reference to the associated mating relations. It turned out that for heterozygote advantage the mutant replaces the selectively equivalent resident allele if it increases the mating preferences for carriers of other alleles. The evolution of lower such preferences requires heterozygote inferiority, which confirms the Wallace effect of speciation (by reinforcement). It is argued that this observation suggests that non-selective constituents of the mating system form the section of the genetic system that is responsible for moderating the genetic load implied by adapting selection while simultaneously securing the adaptational potential embodied in the resident allelic variation. Mating systems thus serve the preservation of adaptability.

Adaptation, Physiological↗

The contribution of male-sterility mutations to inbreeding depression in Mimulus guttatus.

The magnitude of inbreeding depression can influence many aspects of a population's ecology and evolution, including the nature of selection acting on the mating system and the chances that the population will go extinct during periods of small population size. If inbreeding depression is caused primarily by recessive mutations of large effect on fitness, such as lethals or steriles, then it is expected to be purged rapidly by selection with moderate amounts of inbreeding. In contrast, inbreeding depression primarily caused by many genes with mild effects on fitness will not be rapidly purged with inbreeding, so it should be a more resilient barrier to the evolution of self-fertilization and a more significant threat to the survival of endangered species. Here I show that recessive male-sterility alleles at individual loci are common in a primarily outcrossing population of the plant Mimulus guttatus. Despite the high frequency of these major mutations, most of the inbreeding depression for male fertility and cumulative measures of lifetime fitness results from more mildly deleterious alleles. Male-sterility alleles contribute to 31% of the inbreeding depression for the fraction of viable pollen grains, and to 26% of the inbreeding depression for total fitness. These results suggest that most of the inbreeding depression for male fertility in this population would not be purged, in the short term, with moderate inbreeding.

Alleles↗

The dynamics of two- and three-way sexual conflicts over mating.

We consider mathematical models describing the evolutionary consequences of antagonistic interactions between male offence, male defence and female reproductive tract and physiology in controlling female mating rate. Overall, the models support previous verbal arguments about the possibility of continuous coevolutionary chase between the sexes driven by two-way (e.g. between male offence and female traits) and three-way (e.g. between male offence, male defence and female traits) inter-sexual antagonistic interactions. At the same time, the models clarify these arguments by identifying various additional potential evolutionary dynamics and important parameters (e.g. genetic variances, female optimum mating rates, strength of selection in females and the relative contributions of first and second males into offspring) and emphasizing the importance of initial conditions. Models also show that sexual conflict can result in the evolution of monandry in an initially polyandrous species and in the evolution of random mating in a population initially exhibiting non-random mating.

Animals↗

Strong balancing selection at HLA loci: evidence from segregation in South Amerindian families.

The genotypic proportions for major histocompatibility complex loci, HLA-A and HLA-B, of progeny in families in 23 South Amerindian tribes in which segregation for homozygotes and heterozygotes could occur are examined. Overall, there is a large deficiency of homozygotes compared with Mendelian expectations (for HLA-A, 114 observed and 155.50 expected and for HLA-B 110 observed and 144.75 expected), consistent with strong balancing selection favoring heterozygotes. There is no evidence that these deficiencies were associated with particular alleles or with the age of the individuals sampled. When these families were divided into four mating types, there was strong selection against homozygotes, averaging 0.462 for three of the mating types over the two loci. For the other mating type in which the female parent is homozygous and shares one allele with the heterozygous male parent, there was no evidence of selection against homozygotes. A theoretical model incorporating these findings surprisingly does not result in a stable polymorphism for two alleles but does result in an excess of heterozygotes and a minimum fitness at intermediate allele frequencies. However, for more than two alleles, balancing selection does occur and the model approaches the qualities of the symmetrical heterozygote advantage model as the number of alleles increases.

Alleles↗

Background selection and population differentiation.

A general analytical formula is derived, which predicts the effects of background selection on population differentiation at a neutral locus as a result of its linkage with selected loci of deleterious mutations. The theory is based on the assumptions of random mating, multiplicative fitness, and weak selection in hermaphrodite plants in the island model of population structure. The analytical results show that Fst at the neutral locus increases as a result of the effects of background selection, regardless of the dependence or independence among linked background selective loci. The increment in Fst is closely related to the magnitude of linkage disequilibria between the neutral locus and selected loci, and can be estimated by the ratio of Fst with background selection to Fst without background selection minus one. The steady-state linkage disequilibrium between a neutral locus and a selected locus in subpopulations, primarily attained by gene flow, decreases with the recombination rate, and can be enhanced when there are dependence among linked selected loci. Monte Carlo computer simulations with two- and three-locus models show that the analytical formulae perform well under general conditions. Application of the present theory may aid in analyzing the genome-wide mapping of the effect of background selection in terms of Fst.

Computer Simulation↗

Is size-assortative mating important for rapid pigment differentiation in a freshwater isopod?

Identifying mechanisms behind assortative mating is central to the understanding of ecological divergence and speciation. Recent studies show that populations of the freshwater isopod Asellus aquaticus can rapidly become locally differentiated when submerged Chara vegetation expands in lakes. In the novel Chara habitat, isopods have become lighter pigmented and smaller than in ancestral reed stands. In this study, we used a laboratory multiple-choice experiment to investigate assortative mating as a possible prezygotic reproductive barrier between Chara and reed isopods. Mating was assortative when Chara isopods were experimentally mixed with isopods from an adjacent reed site with large-size individuals, suggesting a partial prezygotic reproductive barrier. No deviation from random mating could, however, be detected when Chara isopods were mixed with smaller sized isopods from another reed site. In both experiments, assortative mating was apparently based on size, as Chara isopods were larger and reed isopods smaller in mixed pairs than in assortative pairs. Pigmentation did not have any clear influence on mating. We suggest that divergence in pigmentation evolved through natural selection in conjunction with size-assortative mating indirectly causing assortative mating between Chara and reed isopods. Size-assortative mating is likely a by-product of natural selection, but its importance may hypothetically be transient, if selection erodes the correlation between pigmentation and size over time.

Animals↗

Selection, generalized transmission and the evolution of modifier genes. I. The reduction principle.

Modifier gene models are used to explore the evolution of features of organisms, such as the genetic system, that are not directly involved in the determination of fitness. Recent work has shown that a general "reduction principle" holds in models of selectively neutral modifiers of recombination, mutation, and migration. Here we present a framework for models of modifier genes that shows these reduction results to be part of a more general theory, for which recombination and mutation are special cases. The deterministic forces that affect the genetic composition of a population can be partitioned into two categories: selection and transmission. Selection includes differential viabilities, fertilities, and mating success. Imperfect transmission occurs as a result of such phenomena as recombination, mutation and migration, meiosis, gene conversion, and meiotic drive. Selectively neutral modifier genes affect transmission, and a neutral modifier gene can evolve only by generating association with selected genes whose transmission it affects. We show that, in randomly mating populations at equilibrium, imperfect transmission of selected genes allows a variance in their marginal fitnesses to be maintained. This variance in the marginal fitnesses of selected genes is what drives the evolution of neutral modifier genes. Populations with a variance in marginal fitnesses at equilibrium are always subject to invasion by modifier genes that bring about perfect transmission of the selected genes. It is also found, within certain constraints, that for modifier genes producing what we call "linear variation" in the transmission processes, a new modifier allele can invade a population at equilibrium if it reduces the level of imperfect transmission acting on the selected genes, and will be expelled if it increases the level of imperfect transmission. Moreover, the strength of the induced selection on the modifier gene is shown to range up to the order of the departure of the genetic system from perfect transmission.

Biological Evolution↗