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Direct response in yield and correlated response in components accompanying selection for milk yield in Jerseys.

In 1967, the Jersey herd at the Dairy Experiment Station, Lewisburg, TN was divided into two groups on the basis of ancestry, type, and breeding value for milk as part of a project to determine effects of single trait selection for high milk yield on yield and correlated traits. Control group was mated randomly to 20 unproven young sires selected randomly from those available from breeding studs in 1967. Selection group was mated to sires selected solely on the basis of their high transmitting ability for milk. Selection sires were selected at intervals and used for 4 yr. By the end of the project (1984), lactation information was available on 672 daughters (520 selection and 152 control) of 37 bulls (17 selection and 20 control). Differences in breeding values for milk, fat, and fat test as calculated from the PTA reported in the July 1989 USDA genetic evaluations and differences in first lactation mature equivalent production of milk, fat, fat test, and 4% FCM were examined. Linear mixed models were used for all analyses and contained the fixed effects group, generation within group, and year. Sires were random, nested within group, and used to test for group differences. Groups differed for all traits. Selection was superior to control in breeding value for milk and fat (828 and 31 kg) and for production of milk, fat, and 4% FCM (1066, 42, and 1061 kg). Control was superior to selection in breeding value and production fat test (.15 and .12%). Group differences existed within generation class for all yield traits but not for fat percentage. Realized response closely matched or exceeded expected response as estimated from pedigree information.

Animals↗

Divergent selection and the evolution of signal traits and mating preferences.

Mating preferences are common in natural populations, and their divergence among populations is considered an important source of reproductive isolation during speciation. Although mechanisms for the divergence of mating preferences have received substantial theoretical treatment, complementary experimental tests are lacking. We conducted a laboratory evolution experiment, using the fruit fly Drosophila serrata, to explore the role of divergent selection between environments in the evolution of female mating preferences. Replicate populations of D. serrata were derived from a common ancestor and propagated in one of three resource environments: two novel environments and the ancestral laboratory environment. Adaptation to both novel environments involved changes in cuticular hydrocarbons, traits that predict mating success in these populations. Furthermore, female mating preferences for these cuticular hydrocarbons also diverged among populations. A component of this divergence occurred among treatment environments, accounting for at least 17.4% of the among-population divergence in linear mating preferences and 17.2% of the among-population divergence in nonlinear mating preferences. The divergence of mating preferences in correlation with environment is consistent with the classic by-product model of speciation in which premating isolation evolves as a side effect of divergent selection adapting populations to their different environments.

Adaptation, Biological↗

Socially transmitted mate preferences in a monogamous bird: a non-genetic mechanism of sexual selection.

There is increasing evidence that animals can acquire mate preferences through the use of public information, notably by observing (and copying) the mate preferences of others in the population. If females acquire preferences through social mechanisms, sexual selection could act very rapidly to spread the preference and drive elaboration of the preferred trait(s). Although there are reports of 'mate-choice copying' in polygynous species, there is no clear evidence for this process in monogamous species. Here, we investigated whether adult female zebra finches Taeniopygia guttata can socially acquire sexual preferences for individual males and, in a separate study, for a generalized trait (coloured leg bands) of males. In both studies, test females observed males in two simultaneous conditions: a ('chosen') mixed-sex situation in which a male was paired with a (model) female, and a ('unchosen') same-sex situation in which a male was paired with another male. In the first experiment, after two weeks of females observing males, test females significantly preferred individual males who had been paired with another female (i.e. chosen males). In the second experiment, test females significantly preferred novel males that were wearing the same leg band colour as the apparently chosen males. Our findings are consistent with the conclusion that female zebra finches' mate preferences are altered by public information. Our study implies that mate preferences can spread rapidly through populations by social mechanisms, affecting the strength of sexual selection in a monogamous species.

Animals↗

Predicting response to selection on a quantitative trait: a comparison between models for mixed-mating populations.

Two different theoretical frameworks have been developed to predict response to selection in a mixed mating population (in which reproduction occurs by a mixture of outcrossing and self-fertilization). The genotypic covariance model (GCM) and the structured linear model (SLM) rely on the same assumptions regarding quantitative trait inheritance, but use different genetic summary statistics. Here, we demonstrate the algebraic relationships between the various genetic metrics used in each theory. This is accomplished by reformulating the GCM in terms of the Wright-Kempthorne equation. We use stochastic simulations to investigate the relative accuracy of each theory for a range of selfing rates. The SLM is generally more accurate than the GCM, the most pronounced differences emerging in simulations with inbreeding depression for fitness. In fact, with strong inbreeding depression and high selfing rates, evolution can occur opposite the direction predicted by the GCM. The simulations also indicate that direct application of random mating models to partially selfing populations can produce very inaccurate predictions if quantitative trait loci exhibit dominance.

Animals↗

Female choice selects for a viability-based male trait in pheasants.

Recent theory on sexual selection suggests that females in species without paternal care choose mates by their secondary sexual characters because these indicate genotypic quality which will be transmitted to the offspring. These ideas are not yet empirically supported as data quantifying the relationship between female mate choice and female reproductive success are lacking. Only in one case, in Colias butterflies, has it been demonstrated unequivocally that females choose 'good genotypes' as mates and there is only one study, on Drosophila, demonstrating that mate choice increases one component of offspring fitness. Spur length of male pheasants (Phasianus colchicus) correlates with various fitness-related properties. We here present the first experimental field data showing that female pheasants select mates on the basis of male spur length and that female mate choice correlates with female reproductive success.

Animals↗

Two-way selection for daily gain and feed conversion in a composite rabbit population.

We conducted a two-way selection experiment in a composite rabbit population to investigate the responses to selection for postweaning ADG and feed conversion (FC). Two generations of crossing, followed by four generations of random pair matings, preceded three generations of selection. Selection was practiced within four lines: high-feed conversion (HFC), low-feed conversion (LFC), high gain (HG), and low gain (LG). Data on 1,446 rabbits from the random mating and selection generations were fitted to an animal model to estimate heritabilities of and the genetic correlation between ADG and FC. The two-trait model included rabbit and common litter random effects and line, generation, and sex fixed effects. Estimates of heritability of ADG and FC were .48 and .29, respectively, and the genetic correlation between them was -.82. Common litter environmental effects accounted for a proportion of .11 and .13 of the phenotypic variation of the two traits, respectively. For ADG (in g/d) the regressions of mean breeding values on generation number during the selection period were 1.23 +/- .12 (P < .01) in the HG line and -.86 +/- .12 (P < .01) in the LG line; the regressions for FC (in g feed/g gain) were -.07 +/- .01 (P < .01) in the HFC line and .03 +/- .01 (P < .05) in the LFC line. Selection for ADG was effective in improving ADG and FC.

Animals↗

Genetics of immunological tolerance: I. Bidirectional selective breeding of mice for oral tolerance.

The polygenic nature of oral tolerance regulation was elucidated by the method of bidirectional selective breeding of mouse strains for tolerance susceptibility (TS) and resistance (TR) starting from a genetically heterogeneous population achieved by the equilibrated intercrossing of eight inbred mouse strains (A/J, DBA/2J, P/J, SWR/J, SJL/J, CBA/J, BALB/cJ and C57BL/6J). Seven days after intragastric administration of 5 mg OVA or BSA, mice were intraperitoneally immunized with 100 microg of the corresponding antigen. The individual antibody titres were measured by haemagglutination. The phenotypes at the highest and lowest extremes were selected for assortative mating, avoiding consanguinity. The second litter of each mating couple was intraperitoneally immunized only to evaluate the immunocompetence of the corresponding generation and to ascertain the non-selection of non-responder mice. A normal distribution of agglutinin titres ranging from 4 to 14 log2 was observed in the F0 population. In the F12 generation, TR and TS strains showed highly significant differences for agglutinin titres (TR=15.06+/-1.80 and TS=8.35+/-2.44), and IgG responses by ELISA. Up to the F12 generation, the mean realized heritability was 0.14+/-0.02. The response to the selection was 0.43 log2 and the selection differential 3.10 log2/generation. A provisional estimation indicated that oral tolerance may be influenced by eight or nine independent loci.

Administration, Oral↗

[Hybridization of cells of the same mating type in Saccharomyces yeasts].

The problem of mating-type switches in heterothallic yeast cells was investigated. 93% of non-mating hybrids were obtained in a X a crosses. The hybrids obtained in alpha X alpha crosses expressed alpha-mating type predominantly. Hybrids with no major rearrangements or loss of chromosome III were detected among these hybrids. In the selective system for cytoduction in a X a crosses the significant part of all cytoductants were alpha-maters, i.e. those originated through a----alpha switches. In alpha X alpha crosses alpha cytoductants were predominantly obtained either spontaneously or after UV-irradiation, though the frequency of cytoductants after UV-irradiation exceeded the control value several times. So, we developed the method for selection of mating-type "switchers" (a in equilibrium alpha), avoiding the diploid stage, and demonstrated the possibility of hybridization among the alpha-cells without hereditary changes at the MAT locus.

Alleles↗

Female resistance to male harm evolves in response to manipulation of sexual conflict.

The interests of males and females over reproduction rarely coincide and conflicts between the sexes over mate choice, mating frequency, reproductive investment, and parental care are common in many taxa. In Drosophila melanogaster, the optimum mating frequency is higher for males than it is for females. Furthermore, females that mate at high frequencies suffer significant mating costs due to the actions of male seminal fluid proteins. Sexual conflict is predicted to lead to sexually antagonistic coevolution, in which selection for adaptations that benefit males but harm females is balanced by counterselection in females to minimize the extent of male-induced harm. We tested the prediction that elevated sexual conflict should select for increased female resistance to male-induced harm and vice versa. We manipulated the intensity of sexual conflict by experimentally altering adult sex ratio. We created replicated lines of D. melanogaster in which the adult sex ratio was male biased (high conflict lines), equal (intermediate conflict lines), or female biased (low conflict lines). As predicted, females from high sexual conflict lines lived significantly longer in the presence of males than did females from low conflict lines. Our conclusion that the evolutionary response in females was to the level of male-induced harm is supported by the finding that there were no female longevity differences in the absence of males. Differences between males in female harming ability were not detected. This suggests that the response in females was to differences between selection treatments in mating frequency, and not to differences in male harmfulness.

Animals↗

Inheritance of stress-induced analgesia in mice. Selective breeding study.

Random mated Swiss mice swam for 3 min at 20 degrees C. The animals with less than or equal to 10 s and greater than or equal to 50 s postswim latencies on the hot plate (56 degrees C) were selected as progenitors of low (LA) and high (HA) analgesia lines, respectively. Gradual divergence of latency distributions and postswim hind paw flick latencies between the two lines was observed in successive generations. As shown by the tail flick test applied to the sixth offspring generation, postswim analgesia in the HA line was not only more pronounced but also lasted longer compared to the LA line and the unselected controls. Possible differentiation of the endorphin system activity in the course of selection is discussed.

Animals↗

Reproductive isolation driven by the combined effects of ecological adaptation and reinforcement.

Recent years have seen a resurgence of interest in the process of speciation but few studies have elucidated the mechanisms either driving or constraining the evolution of reproductive isolation. In theory, the direct effects of reinforcing selection for increased mating discrimination where interbreeding produces hybrid offspring with low fitness and the indirect effects of adaptation to different environments can both promote speciation. Conversely, high levels of homogenizing gene flow can counteract the forces of selection. We demonstrate the opposing effects of reinforcing selection and gene flow in Timema cristinae walking-stick insects. The magnitude of female mating discrimination against males from other populations is greatest when migration rates between populations adapted to alternate host plants are high enough to allow the evolution of reinforcement, but low enough to prevent gene flow from eroding adaptive divergence in mate choice. Moreover, reproductive isolation is strongest under the combined effects of reinforcement and adaptation to alternate host plants. Our findings demonstrate the joint effects of reinforcement, ecological adaptation and gene flow on progress towards speciation in the wild.

Adaptation, Biological↗

Spontaneous mutation for a quantitative trait in Drosophila melanogaster. II. Distribution of mutant effects on the trait and fitness.

Starting from a completely homozygous population of Drosophila melanogaster, lines have been derived and subjected to 47 generations of divergent selection for abdominal bristle number (20 lines selected in each direction) or to 60-67 generations of inbreeding (100 B lines maintained by a single brother-sister mating, 100 C lines maintained by two double first cousin matings). In the selected lines, 25 were identified carrying at least 30 mutations affecting bristle number. A large fraction of these mutations (42%) were lethals. Non-lethal mutations had smaller effects on the trait, were predominantly additive and had no detectable pleiotropic effects on fitness. In the inbred lines, 21 mutations affecting bristles were individually analysed. Deleterious mutations had the largest effects on the trait (irrespective of sign) and showed recessive gene action (complete or incomplete). The rest were predominantly additive and had smaller effects. Thus, both procedures identify a quasi-neutral class of additive mutations which should be close to that responsible for standing variation in natural populations. Moreover, the results indicate a leptokurtic distribution of mutant effects, consistent with a model of natural selection acting on bristles through pleiotropic effects of pertinent loci on fitness. Consequently, neutral additive alleles of considerable effect can be found segregating at intermediate frequencies in natural populations.

Abdomen↗

Sex difference in the effect of mating on c-fos expression in luteinizing hormone-releasing hormone neurons of the ferret forebrain.

The pulsatile secretion of LH was previously found to rise in female ferrets after receipt of an intromission, whereas in males that achieved an intromission, both LH and testosterone secretion were either reduced or unchanged. We sought to determine whether this sexually dimorphic pattern of LH secretion reflects a sex difference in the effect of mating on the activity of forebrain neurons that secrete LHRH. Immunocytochemical methods were used to localize the nuclear protein product of the immediate early gene, c-fos, as an index of increased neuronal activity after mating. Nuclear FOS immunoreactivity (FOS-IR) was monitored in LHRH-IR neurons as well as other non-LHRH forebrain neurons. In confirmation of previous reports, LHRH-IR perikarya in ferrets of both sexes were located medially along the base of the brain at rostral, medial, and caudal levels of the preoptic-hypothalamic continuum. In each of these regions a significantly higher percentage of LHRH-IR neurons was colabeled with nuclear FOS-IR in mated than in unpaired females. By contrast, an equivalent low percentage of LHRH-IR neurons was colabeled with FOS-IR in mated and unpaired male ferrets. Significantly more FOS-IR neurons (not colabeled with LHRH) were detected in the bed nucleus of the stria terminalis, the medial preoptic area, the dorsal-medial hypothalamus, and the medial amygdala (MA) of mated vs. unpaired females. By contrast, mating significantly augmented FOS-IR only in the MA of male ferrets. The results suggest that the sexually dimorphic pattern of LH secretion that occurs in ferrets after mating reflects a selective activation of LHRH neurons in the female forebrain. This sex-specific increase in the responsiveness of LHRH neurons to mating may depend on input from a limbic circuit which includes the medial amygdala, bed nucleus of the stria terminalis, and medial preoptic area.

Animals↗

Selection for weaning weight in Targhee sheep in two environments. I. Direct response.

In 1961, selection for 120-d weight was initiated in two flocks from a common base population of grade Targhee sheep. At Davis, sheep were maintained on a good plane of nutrition, on irrigated pasture or in drylot. At Hopland, sheep grazed annual grassland range, with supplementary feeding only at mating and lambing. Selected (DW) and control (DC) lines were maintained at Davis from 1961 through 1977. A selected (HW) line, replicate control (HC1 and HC2) lines and a line (DH) mated to the Davis DW rams were maintained at Hopland from 1961 through 1980, with the exception that HC2 was terminated in 1977. Multiplicative factors were used to adjust weights for effects of age of dam, sex and type of birth and rearing. Response to selection was estimated as the difference between selected and control line linear regression coefficients of adjusted line means on year. The Hopland replicate controls did not differ significantly from each other (HC1 - HC2 = .004 +/- .056 kg/yr), and the control line data were pooled (HC). The overall control line mean 120-d weights on a female, single, mature-dam basis were 33.2 and 30.4 kg at Davis and Hopland, respectively. Direct response was greater at Davis than at Hopland: DW - DC = .524 +/- .073 kg/yr (P less than .001); HW - HC = .151 +/- .034 kg/yr (P less than .001). Corresponding realized heritabilities were .17 and .06. Direct response for the DH line was DH - HC = .226 +/- .036 (P less than .001); realized heritability was .08. Response in the DH line was greater (P less than .05) than that in the HW line: HW - DH = -.075 +/- .037 kg/yr. This indicates that: (1) genetic improvement made on a higher plane of nutrition was expressed, but to a lesser degree, under range conditions and (2) selection under better feed conditions resulted in at least as much improvement in growth rate in a range environment as did selection under range conditions.

Animals↗

The significance of over- and underdominance for the maintenance of genetic polymorphisms. I. Underdominance and stability.

It is pointed out that the standard selection models in population genetics all require some form of heterozygote advantage in fitness in order to guarantee the maintenance or stability of genetic polymorphisms. Even more recent results demonstrating the existence of stable two-locus polymorphisms with marginal underdominance at both loci are based on certain epistatically acting heterosis assumptions. This raises the question as to whether heterozygote advantage in fitness is indeed a generally valid principle of maintaining polymorphisms. To avoid ambiguity in definition of heterozygote advantage (overdominance) as it appears in multiallele or multilocus systems, a one-locus-two-allele model is considered. This model allows for sexually asymmetric selection and random mating. It is shown that the model produces globally stable polymorphisms exhibiting underdominance in fitness for a considerable and biologically reasonable range of selection values. Having thus properly refuted the general validity of the common overdominance principle, a modified version is suggested which covers the classical viability selection model and its extension to arbitrary, sexually asymmetric viability and fertility selection. This modified overdominance principle is based on the notion of fractional fitnesses and relates protectedness of biallelic polymorphisms to the extent to which each genotype reproduces its own type. The fact that the model treated displays frequency dependent fitnesses which may change in ranking while approaching equilibrium is discussed in relation to problems of the evolution of overdominance and underdominance.

Animals↗

Unusually dynamic sex roles in a fish.

Sex roles are typically thought of as being fixed for a given species. In most animals males compete for females, whereas the females are more reluctant to mate. Therefore sexual selection usually acts most strongly on males. This is explained by males having a higher potential reproductive rate than females, leading to more males being sexually active (a male-biased operational sex ratio). However, what determines sex roles and the strength of sexual selection is a controversial and much debated question. In this large-scale field study, we show a striking temporal plasticity in the mating competition of a fish (two-spotted goby, Gobiusculus flavescens). Over the short breeding season fierce male-male competition and intensive courtship behaviour in males were replaced by female-female competition and actively courting females. Hence, sex role reversal occurred rapidly. This is the first time that a shift in sex roles has been shown in a vertebrate. The shift might be explained by a large decline in male abundance, strongly skewing the sex ratio towards females. Notably, the sex role reversal did not occur at an equal operational sex ratio, contrary to established sex role theory.

Animals↗

The robustness of Hamilton's rule with inbreeding and dominance: kin selection and fixation probabilities under partial sib mating.

Assessing the validity of Hamilton's rule when there is both inbreeding and dominance remains difficult. In this article, we provide a general method based on the direct fitness formalism to address this question. We then apply it to the question of the evolution of altruism among diploid full sibs and among haplodiploid sisters under inbreeding resulting from partial sib mating. In both cases, we find that the allele coding for altruism always increases in frequency if a condition of the form rb>c holds, where r depends on the rate of sib mating alpha but not on the frequency of the allele, its phenotypic effects, or the dominance of these effects. In both examples, we derive expressions for the probability of fixation of an allele coding for altruism; comparing these expressions with simulation results allows us to test various approximations often made in kin selection models (weak selection, large population size, large fecundity). Increasing alpha increases the probability of fixation of recessive altruistic alleles (h<1/2), while it can increase or decrease the probability of fixation of dominant altruistic alleles (h>1/2).

Altruism↗

No detectable genetic correlation between male and female mating frequency in the stalk-eyed fly Cyrtodiopsis dalmanni.

There is much interest in explaining why female insects mate multiply. Females of the stalk-eyed fly Cyrtodiopsis dalmanni can mate several times each day in a lifetime which may span several months. There are many adaptive explanations, but one hypothesis that has received little rigorous empirical attention is that female multiple mating has evolved for non-adaptive reasons as a correlated response to selection for high male mating frequency rather than because of direct or indirect benefits accruing to females. We tested this hypothesis in stalk-eyed flies by measuring the mating frequency of females from lines that exhibited a direct response in males to artificial selection for increased ('high') and decreased ('low') male mating frequency. We found that the mating frequency of high-line females did not differ from that of low-line females. Hence, there was no support for a genetic correlation between male and female mating frequency in this species. Our study suggests that the genes which influence remating may not be the same in the sexes, and that females remate frequently in this species to gain as yet unidentified benefits.

Analysis of Variance↗