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The advantages of segregation and the evolution of sex.

In diploids, sexual reproduction promotes both the segregation of alleles at the same locus and the recombination of alleles at different loci. This article is the first to investigate the possibility that sex might have evolved and been maintained to promote segregation, using a model that incorporates both a general selection regime and modifier alleles that alter an individual's allocation to sexual vs. asexual reproduction. The fate of different modifier alleles was found to depend strongly on the strength of selection at fitness loci and on the presence of inbreeding among individuals undergoing sexual reproduction. When selection is weak and mating occurs randomly among sexually produced gametes, reductions in the occurrence of sex are favored, but the genome-wide strength of selection is extremely small. In contrast, when selection is weak and some inbreeding occurs among gametes, increased allocation to sexual reproduction is expected as long as deleterious mutations are partially recessive and/or beneficial mutations are partially dominant. Under strong selection, the conditions under which increased allocation to sex evolves are reversed. Because deleterious mutations are typically considered to be partially recessive and weakly selected and because most populations exhibit some degree of inbreeding, this model predicts that higher frequencies of sex would evolve and be maintained as a consequence of the effects of segregation. Even with low levels of inbreeding, selection is stronger on a modifier that promotes segregation than on a modifier that promotes recombination, suggesting that the benefits of segregation are more likely than the benefits of recombination to have driven the evolution of sexual reproduction in diploids.

Alleles↗

Association of somatotropin (BST) gene polymorphism at the 5th exon with selection for milk yield in Holstein cows.

A selection project produced control cows from continuous matings with breed average bulls for predicted transmitting ability for milk (PTA-milk) in 1964 and select cows from matings to four of the highest PTA-milk bulls each year since 1964. Blood samples were collected in 1992 when milk yield difference of select and control line cows exceeded 3,800 kg of milk/305-day lactation. Genomic DNA from control (n = 49) and select (n = 101) cows was analyzed for the presence of variants associated with amino acid position 127 (leucine, AluI[+]; valine, AluI[-]) of bovine somatotropin (bST). Amplification of a 428 base-pair fragment of the bST gene from individual cows, subsequent restriction enzyme (AluI) digestion, and separation resulting fragments indicated three genotypes AluI(+/+), AluI(+/-), and AluI(-/-) in 110, 39, and 1 animal(s), respectively. Gene frequencies of leucine127 and valine 127 alleles were similar for control (0.867m 0.133) and select (0.861, 0.139) animals. United States Department of Agriculture-PTA values were compared between the two genotypes, AluI(+/+) and AluI(+/-). Estimated breeding value for milk (EBV-milk) and average yield deviation for milk (AYD-milk) were not associated with genotype for control animals. However, presence of the valine allele was correlated with decreased EBV-milk (P = 0.03) and AYD-milk (P = 0.16) in select animals and accounted for a decrease of approximately 170 kg of EBV-milk and 240 kg of AYD-milk.

Animals↗

Evolution of interference competition by individual, family, and group selection.

The necessary conditions for the evolution of social behaviors in a population with three levels of biological organization are derived by using a population genetic model (one locus, two alleles, random mating, discrete generations). Total selection on the behavior, Deltaq, is partitioned into the sum of three components: (i) Deltaq(I), selection between individuals within families; (ii) Deltaq(F), selection between families within groups; and (iii) Deltaq(G), selection between groups of families. I show that any level of selection can be made to operate in concert with or in opposition to any other, depending upon the fitness effects of the behavior. The implications of the model are discussed in relation to those adaptive explanations of phenotypic traits that generally consider selection to operate only between individuals.

Journal Article↗

Mating system and the evolution of quantitative traits: an experimental study of Mimulus guttatus.

The mating system of a population profoundly influences its evolution. Inbreeding alters the balance of evolutionary forces that determine the amount of genetic variation within a population. It redistributes that variation among individuals, altering heritabilities and genetic correlations. Inbreeding even changes the basic relationships between these genetic statistics and response to selection. If populations differing only in mating system are exposed to the same selection pressures, will they respond in qualitatively different ways? Here, we address this question by imposing selection on an index of two negatively correlated traits (flower size and development rate) within experimental populations that reproduce entirely by outcrossing, entirely by self-fertilizing, or by a mixture of outcrossing and selfing. Entirely selfing populations responded mainly by evolving larger flowers whereas outcrossing populations also evolved more rapid development. Divergence occurred despite an equivalent selection regime and no direct effect of mating system on fitness. The study provides an experimental demonstration of how the interaction of selection, genetic drift, and mating system can produce dramatic short-term changes in trait means, variances, and covariances.

Crosses, Genetic↗

The adhesin Hwp1 and the first daughter cell localize to the a/a portion of the conjugation bridge during Candida albicans mating.

The cell wall protein Hwp1 was originally demonstrated to be expressed exclusively in hyphae of Candida albicans and cross-linked to human epithelium by mammalian transglutaminase. Hwp1 is expressed on the walls of hyphae formed by a/alpha, a/a, and alpha/alpha cells. Hence, it is expressed on hyphae independently of mating type. However, Hwp1 is selectively expressed on the wall of conjugation tubes formed by a/a cells, but not alpha/alpha cells, in the mating process. This was demonstrated in all possible crosses between four unrelated natural a/a strains and four unrelated alpha/alpha strains. In zygotes, Hwp1 is restricted to that portion of the wall of the conjugation bridge contributed by the a/a parent cell. Hwp1 staining further revealed that the first daughter bud that emerges from the conjugation bridge does so from the a/a-contributed portion. Hwp1 expression and localization during the mating process is, therefore, mating type specific, opaque phase specific, and alpha-pheromone induced. These results indicate that the mating type-specific contributions to the conjugation bridge during the mating process in C. albicans are qualitatively and functionally distinct and that the a/a portion of the bridge, which selectively contains Hwp1, bears the first daughter cell in the mating process.

Blotting, Northern↗

The sexual selection continuum.

The evolution of mate choice for genetic benefits has become the tale of two hypotheses: Fisher's 'run-away' and 'good genes', or viability indicators. These hypotheses are often pitted against each other as alternatives, with evidence that attractive males sire more viable offspring interpreted as support for good genes and with a negative or null relationship between mating success of sons and other components of fitness interpreted as favouring the Fisher process. Here, we build a general model of female choice for indirect benefits that captures the essence of both the 'Fisherian' and 'good-genes' models. All versions of our model point to a single process that favours female preference for males siring offspring of high reproductive value. Enhanced mating success and survival are therefore equally valid genetic benefits of mate choice, but their relative importance varies depending on female choice costs. The relationship between male attractiveness and survival may be positive or negative, depending on life-history trade-offs and mating skew. This relationship can change sign in response to increased costliness of choice or environmental change. Any form of female preference is subject to self-reinforcing evolution, and any relationship (or lack thereof) between male display and offspring survival is inevitably an indicator of offspring reproductive values. Costly female choice can be maintained with or without higher offspring survival.

Alleles↗

On the origin of species by means of assortative mating.

Assortative mating may split a population even in the absence of natural selection. Here, we study when this happens if mating depends on one or two quantitative traits. Not surprisingly, the modes of assortative mating that can cause sympatric speciation without selection are rather strict. However, some of them may occur in nature. Slow elimination of intermediate individuals caused by the gradual tightening of assortative mating, which evolves owing to relatively weak disruptive selection, provides the alternative scenario for sympatric speciation, in addition to fast elimination of intermediate individuals as a result of the direct action of strong disruptive selection under an invariant mode of assortative mating. Even when assortative mating alone cannot split an initially coherent population, it may be able to prevent the merging of species after their secondary contact.

Animals↗

Selection at work in self-incompatible Arabidopsis lyrata: mating patterns in a natural population.

Identification and characterization of the self-incompatibility genes in Brassicaceae species now allow typing of self-incompatibility haplotypes in natural populations. In this study we sampled and mapped all 88 individuals in a small population of Arabidopsis lyrata from Iceland. The self-incompatibility haplotypes at the SRK gene were typed for all the plants and some of their progeny and used to investigate the realized mating patterns in the population. The observed frequencies of haplotypes were found to change considerably from the parent generation to the offspring generation around their deterministic equilibria as determined from the known dominance relations among haplotypes. We provide direct evidence that the incompatibility system discriminates against matings among adjacent individuals. Multiple paternity is very common, causing mate availability among progeny of a single mother to be much larger than expected for single paternity.

Arabidopsis↗

Heritable variation of sex pheromone composition and the potential for evolution of resistance to pheromone-based control of the Indian meal moth, Plodia interpunctella.

The short-term evolutionary effect of pheromone-based mating disruption on the mating ability of the Indian meal moth, Plodia interpunctella, was investigated. Three independent selection lines were established, and the mating ability of moths in plastic tents treated with high doses of pheromone and in control tents was compared for two consecutive generations. In addition, the heritability of the sex pheromone blend, measured as the ratio of two major pheromone components (Z,E)-9,12-tetradecadienyl acetate and (Z,E)-9,12-tetradecadienol, was estimated. Based on a mother-daughter regression analysis including 21 families, the heritability of the pheromone blend was 0.65 +/- 0.14, indicating a potential for evolutionary change of the character. However, no increase in mating ability of females in pheromone-treated tents or alteration of the pheromone blend was observed in any selection line when compared with control lines, indicating no or weak selection on the pheromone blend as well as other traits influencing mating ability of this species under the created mating disruption conditions. Factors contributing to the lack of selection effects are discussed.

Animals↗

Sperm competition experiments between lines of crickets producing different sperm lengths.

Sperm numbers can be important determinants of fertilization success in sperm competition. However, the importance of variation in sperm size is less well understood. Sperm size varies significantly both between and within species and comparative studies have suggested that some of this variance can be explained by sperm competition. In this study we examine whether variation in sperm length has consequences for fertilization precedence using controlled sperm competition experiments in the field cricket Gryllus bimaculatus. This species is an ideal model for such investigations because the mechanism of sperm competition generates complete mixing of different males' spermatozoa in the female (thereby allowing individual sperm to express their own competitive abilities). We successfully bred lines of crickets, the males of which produced short, medium and long sperm types with narrow and non-overlapping distributions. Males of different lines were then sequentially mated with control females in order to create two-male sperm competitions. The paternity outcomes of these competitions were measured after matings using an irradiated male technique (with a full reciprocal design that controls for natural fertility and any irradiation effects on gamete competitiveness) over a 12 day oviposition period. However, having successfully bred diverging sperm length lines and competing males that differed in sperm length, we found no evidence that a male's sperm size explained any of the variation in their relative fertilization success. Males from lines producing longer sperm showed no fertilization advantage over males producing shorter sperm across 97 double matings. There was also no advantage for males producing a sperm length close to the population mean over those competitors whose sperm length had been selectively diverged across 63 matings.

Animals↗

Investigation of breeding strategies to increase the probability that German shepherd dog and Labrador retriever dog guides would attain optimum size.

An optimum-sized dog guide weighs 18 to 32 kg and measures 53 to 64 cm in height at the withers when mature body size is attained. Effects of selection index with and without restrictions, independent trait selection, directional selection, stabilizing selection, and negative assortative mating were modeled using data from German shepherd dogs and Labrador retrievers raised by the Seeing Eye, Inc., Morristown, NJ from 1979 to 1997. The selection goals were to decrease mature weight and mature height in German shepherd dogs and to decrease mature weight and increase mature height in Labrador retrievers. Mature weights were recorded for 1,333 German shepherd dog offspring and their 69 dams and 17 sires, and 1,081 Labrador retriever offspring and their 51 dams and 13 sires. Mature heights also were recorded for offspring and parents, including 871 German shepherd dogs from 70 dams and 15 sires, and 793 Labrador retrievers from 40 dams and 13 sires. Selecting on mature weight alone produced the highest aggregate genetic-economic gain for German shepherd dogs compared with the selection indices with and without restrictions, generating a 2.10-kg decrease in mature weight and a correlated 0.36-cm decrease in mature height. In Labrador retrievers, selecting for mature height alone produced the highest aggregate genetic-economic gain but caused an increase in mature weight. Weighting the two traits equally but in the opposite direction without restrictions was the only index that produced the desired effect of decreasing mature weight and increasing mature height in Labrador retrievers. Response to selection for one generation of directional selection for a single trait included a 0.50-kg decrease in mature weight for German shepherd dogs, a 0.59-kg decrease in mature weight for Labrador retrievers, a 0.18-cm decrease in mature height for German shepherd dogs, and a 0.91-cm increase in mature height for Labrador retrievers. Increasing the percentage of dogs attaining optimum size may decrease the cost of production for the Seeing Eye, Inc., because fewer dogs would need to be raised and trained to provide assistance to the same number of blind individuals.

Animals↗

Polyandry in Lepidoptera: a heritable trait in Spodoptera exigua Hübner.

The genetic basis as well as the mode of inheritance of polyandry in Spodoptera exigua Hübner was studied in the laboratory by using a simple divergent selection experiment followed by F1 reciprocal crosses, F2 and backcrosses. There was an effective response to artificial selection for high (H line) and low (L line) female mating frequency with significant separation of the lines by the second generation of selection. The mean female mating frequency in the parental generation (1.57 matings per female) reached plateaus of 2.50 and 1.25 matings per female in the H and L lines, respectively, after six generations of selection. Selection response becomes saturated at about 90% and 25% levels of polyandry (percentage females re-mating) in the H and L lines, respectively, and consequently mono- and polyandric pure strains were not obtained. Polyandry levels in offspring from the H and L lines and their hybrids in F1, F2 and backcrosses consistently indicate that female mating frequency was more or less proportional to the relative amounts of genes derived from the H and L lines. Such a clear pattern of hybrid responses, together with the gradual selective changes under artificial selection, suggests the involvement of a polygenic system. Female mating frequencies from progeny of the two reciprocal F1 crosses were not significantly different, which suggest that the trait was autosomally inherited. Moreover, female mating frequency of F1 (pooled) progeny was not significantly different from the mid-parental value, which suggest no dominance. The computation of the Cavalli's joint scaling test consistently confirmed these results yielding values of d = 0.51 +/- 0.10 and h = 0.12 +/- 0.21. The broad sense heritability estimate was H2 = 0.73. It is concluded that polyandry in S. exigua is a polygenic, autosomal heritable trait and that additive genetic variance is available for selection for female mating frequency. The implications of the genetic basis of polyandry are briefly discussed in the context of current theories about this crucial insect mating system.

Animals↗

Sexual selection and genital anatomy of male primates.

Correlations between mating system and various aspects of genital anatomy suggest a strong influence of sexual selection on genital morphology. We test the generality of the influence by examining whether primate taxa in which there might be enhanced sexual selection (those with multi-male mating systems) possess, as expected, relatively more spinous penises than do taxa with other mating systems. As most prosimians, but few anthropoids (monkeys and apes), possess penile spines, and because the predominant mating systems of the two taxa differ, taxonomic constraints are taken into account. Sexual selection apparently does not act on penile spines in the same manner as on other aspects of genital anatomy: spinosity is not greatest in multi-male taxa of either prosimians or anthropoids. In some taxa, spines might stimulate reproductive readiness and synchrony in situations in which the sexes live apart and do not have other means of communicating reproductive state (dispersed social systems and 'stolen' extra-pair copulations), but problems exist with the hypothesis, as they do with the idea that spines are involved with scent marking. It seems that either penile spines have several functions, or penile spinosity in primates, and other orders, remains to be explained.

Animals↗

Effect of genetic selection and MHC haplotypes on lymphocyte proliferation and interleukin-2 like activity in chicken lines selected for high and low antibody production against sheep red blood cells.

Chickens from third generation matings of lines of chickens selected for high (HA) and low (LA) antibody production to sheep red blood cells (SRBC) and typed for MHC genotypes B13/13, B13/21, and B21/21 were used in this study. Chickens from both lines carried all the three genotypes B13/13, B13/21, and B21/21. To study T- and B-lymphocytes mitogenic activity, 12-week-old female chickens were injected intravenously with 0.2 ml of 9% SRBC and spleens were collected at 0, 6 h, and 6 day post-antigen injection (pAg). Isolated lymphocytes were incubated with either Concanavalin-A (Con-A) for T-cell activity, or Pokeweed mitogen (PWM) for B-cell activity and thymidine 3H uptakes were measured. To study the Interleukin-2 (IL-2)-like activity in the same lines and genotypes, splenic lymphocytes from 12-week-old chickens were passed through nylon wool columns to enrich the T-cell population. After a 24 h incubation with Con-A, the conditioned media (CM) were collected. The CM were tested for IL-2 like activity by determining whether they altered the proliferation of Con-A stimulated T cells. This proliferation effect was then compared to that of a reference conditioned media (RCM) prepared from K-strain birds and that were used as the standard for the assay. There was no significant difference (p > 0.05) in IL-2 like activity between HA and LA lines, however, the LA was significantly higher than HA (p < 0.05) in T- and B-cell mitogenic activity. The genotype B13/13 had significantly higher (p < 0.05) IL-2 like activity than the B21/21. The genotype B13/13 was also significantly higher (p < 0.05) in T- and B-cell mitogenic activity than the B21/21. At 0 h, pAg T- and B-mitogenic activity was significantly higher (p < 0.05) than 6 h. In summary, our results indicate that although the birds were selected for high antibody production to SRBC, their lymphocyte mitogenic activity was lower than those selected for low antibody production. Hence, humoral and cell-mediated immune responses appear to be under different genetic controls, and that selection for greater humoral response may be at the expense of cellular responses. Our results also suggest differences in IL-2 like activity production between chickens carrying different MHC B-haplotypes, and that genetic control of such activity is possibly linked to the MHC genes.

Animals↗

Mate choice and imprinting in birds studied by cross-fostering in the wild.

Sexual-selection theories generally assume that mating preferences are heritable traits. However, there is substantial evidence that the rearing environment may be important for the development of mating preferences, indicating that they may be learnt, or modified by experience. The relative importance of such sexual imprinting across species remains largely unexplored. Here, we report results of a large-scale cross-fostering experiment in the wild in which nestling birds were raised by parents of a different species. We show that resulting sexual imprinting may have a negative effect on pairing success in one species (the great tit, Parus major), but not in two other species (the blue tit, P. caeruleus and the pied flycatcher, Ficedula hypoleuca). A remarkable variation thus seems to exist, even between species that are congeneric and have similar breeding ecologies. The cross-fostering resulted in heterospecific pairings between the two tit species (female blue tit breeding with male great tit), which has never, to our knowledge, been previously documented. However, the chicks fledging from these nests were all blue tit.

Animals↗

Mating alters gonadotropin-releasing hormone cell number and content.

Female musk shrews (Suncus murinus) are induced ovulators, which lack a behavioral and ovarian estrous cycle. Females mate the first time they are introduced to a male, but a second or third mating, at least 24 h later, is usually required to induce ovulation. Because GnRH-immunoreactive (GnRH-ir) cell numbers increase during and after exposure to a male, we hypothesized that mating promotes synthesis of this important peptide. To test this hypothesis, we examined changes in GnRH-ir cell number and GnRH-ir content at select time points after mating and ovulation. One hour after mating, GnRH-ir cell numbers in olfactory-related regions of the forebrain were increased. By 15 h after mating, just before ovulation, GnRH-ir cell number and content were increased. Twenty-four hours after mating, GnRH-ir cell numbers in the tenia tecta and medial septum/diagonal band were lower in females that ovulated compared with females that did not ovulate. By 40 h postmating, females that ovulated had fewer GnRH-ir neurons and lower GnRH content in the entire brain than females that did not ovulate. In addition, we found significant negative correlations between plasma estradiol concentrations and both GnRH-ir cell numbers and content in the preoptic area of animals killed around the time of ovulation. Interestingly, significantly more GnRH-ir neurons and a greater content of GnRH peptide were observed in several forebrain nuclei of females that did not ovulate 40, compared to 24, h after mating. In contrast, numbers of GnRH-ir neurons in the midbrain declined 40 h postmating in ovulated females. These results suggest that mating stimulates activity in GnRH-ir neurons, and that ovulation is correlated with a decline in GnRH-ir cell number and content. In this species, mating can be used as an external trigger to activate GnRH neurons and examine the regulation and production of GnRH in heterogenic neuronal populations.

Animals↗

Population and behaviour genetics of Drosophila ananassae.

Drosophila ananassae is a cosmopolitan and domestic species. It occupies a unique status among the Drosophila species due to certain peculiarities in its genetic behaviour. The most unusual feature of this species is spontaneous male recombination in appreciable frequency. The present review summarizes the work done on population and behaviour genetics of D. ananassae from India. Population dynamics of three cosmopolitan inversions has been studied in Indian populations of D. ananassae and it is evident from the results that there is a considerable degree of genetic divergence at the level of inversion polymorphism. In general, the populations from south India show more differentiation than those from the north. These three cosmopolitan inversions, which are coextensive with the species, exhibit heterosis. Interracial hybridization does not lead to breakdown of heterosis, which suggests that evidence for coadaptation is lacking in geographic populations of D. ananassae. Heterosis appears to be simple luxuriance rather than populational heterosis (coadaptation). Unlinked inversions occur in random associations, indicating no interchromosomal interactions. However, two inversions of the third chromosome often show strong linkage disequilibrium in laboratory populations, which is due to epistatic gene interaction and suppression of crossing-over. Genetic variations for certain allozyme polymorphism and sternopleural bristle phenotypes in Indian populations of D. ananassae have also been observed. A number of investigations have also been carried out on certain aspects of behaviour genetics of Indian D. ananassae. There is evidence for sexual isolation within D. ananassae. Significant variations in mating propensity of several isofemale strains, inversion karyotypes, the diminishing effects of certain mutations on sexual activity of males and positive response to selection for high and low mating propensity provide evidence for genetic control of sexual behaviour in D. ananassae. Males contribute more to variation and thus are more subject to intra-sexual selection than females. Evidence for rare male mating advantage has also been presented. Geographic strains of D. ananassae show variation with respect to oviposition site preference. The results of studies on pupation site preference, which is an important component of larval behaviour, suggest that larval pupation behaviour in D. ananassae is under polygenic control with a substantial amount of additive genetic variation.

Animals↗

Chronic toxicity, oncogenic potential, and reproductive toxicity of p-nitroaniline in rats.

Dose levels for these studies were selected mainly on the basis of subchronic studies, although consideration was also given to workplace exposure levels and proposed mechanism of tumor formation with structurally similar compounds. For the chronic study, groups of 60 male and 60 female Sprague-Dawley CD (Registered Trademark of Charles River Breeding Laboratories, Portage, MI) rats were given 0, 0.25, 1.5, or 9.0 mg/kg/day paranitroaniline (PNA) by gavage in corn oil for a period of 2 years. Parameters monitored included clinical observations, ophthalmoscopic exams, body weights, food consumption, hematology, clinical chemistry, and urinalysis at regular intervals throughout the study. All gross lesions and over 40 tissues were examined histologically for all control and high-dosage-level animals. Gross lesions, spleens, and livers of low- and mid-dosage groups were also examined histologically. For the reproduction study, groups of 15 male and 30 female rats, designated as F0 generation, were given PNA at the same levels as the chronic study for 14 weeks prior to mating and during mating, gestation, and lactation. Selected groups of 15 male and 30 female rats of the F1 generation received the same dose of PNA for 18 weeks prior to mating and during mating, gestation, and lactation. F2 pups were observed through weaning at which time they were euthanized. Observations made during the study included body weights, food consumption, mating and fertility indices, pup and litter survival indices, and histopathology of selected tissues. In the chronic study, except for a slight decrease in survival of high-dose male rats late in the study, survival in all treated groups was comparable to controls. Blood methemoglobin levels were elevated in the mid- and high-dosage groups, while slight anemia was observed in the high-dosage group also. Spleen weights were significantly increased in the high-dosage groups. An accumulation of brown pigment was observed in the cytoplasm of the sinusoidal macrophages or littoral cells of the liver and in the reticuloendothelial cells of the spleen. No treatment-related increase in tumor incidence was observed. In the reproduction study, no consistent pattern of effect from treatment between the F0 and F1 generation was seen in mating, pregnancy, or fertility indices. Thus, administration of PNA at levels which produced significant methemoglobinemia and low-level anemia in the rat and histological changes in the spleen produced no tumors or reproducible effects on reproductive performance.

Aniline Compounds↗