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Bird song, ecology and speciation.

The study of bird song dialects was once considered the most promising approach for investigating the role of behaviour in reproductive divergence and speciation. However, after a series of studies yielding conflicting results, research in the field slowed significantly. Recent findings, on how ecological factors may lead to divergence in both song and morphology, necessitate a re-examination. We focus primarily on species with learned song, examine conflicting results in the literature and propose some potential new directions for future studies. We believe an integrative approach, including an examination of the role of ecology in divergent selection, is essential for gaining insight into the role of song in the evolution of assortative mating. Habitat-dependent selection on both song and fitness-related characteristics can lead to parallel divergence in these traits. Song may, therefore, provide females with acoustic cues to find males that are most fit for a particular habitat. In analysing the role of song learning in reproductive divergence, we focus on post-dispersal plasticity in a conceptual framework. We argue that song learning may initially constrain reproductive divergence, while in the later stages of population divergence it may promote speciation.

Animals↗

Estimation of genetic variability and selection response for clutch length in dwarf brown-egg layers carrying or not the naked neck gene.

In order to investigate the possibility of using the dwarf gene for egg production, two dwarf brown-egg laying lines were selected for 16 generations on average clutch length; one line (L1) was normally feathered and the other (L2) was homozygous for the naked neck gene NA. A control line from the same base population, dwarf and segregating for the NA gene, was maintained during the selection experiment under random mating. The average clutch length was normalized using a Box-Cox transformation. Genetic variability and selection response were estimated either with the mixed model methodology, or with the classical methods for calculating genetic gain, as the deviation from the control line, and the realized heritability, as the ratio of the selection response on cumulative selection differentials. Heritability of average clutch length was estimated to be 0.42 +/- 0.02, with a multiple trait animal model, whereas the estimates of the realized heritability were lower, being 0.28 and 0.22 in lines L1 and L2, respectively. REML estimates of heritability were found to decline with generations of selection, suggesting a departure from the infinitesimal model, either because a limited number of genes was involved, or their frequencies were changed. The yearly genetic gains in average clutch length, after normalization, were estimated to be 0.37 +/- 0.02 and 0.33 +/- 0.04 with the classical methods, 0.46 +/- 0.02 and 0.43 +/- 0.01 with animal model methodology, for lines L1 and L2 respectively, which represented about 30% of the genetic standard deviation on the transformed scale. Selection response appeared to be faster in line L2, homozygous for the NA gene, but the final cumulated selection response for clutch length was not different between the L1 and L2 lines at generation 16.

Animals↗

The use of a sociobiological questionnaire (SQ) for the assessment of sexual dimorphism.

A sociobiology questionnaire (SQ) (Thiessen and Kittrell, 1980) was administered to 527 undergraduate males and females. Twenty-seven predictions were tested. Sixteen of the predictions were statistically confirmed, and eight were in the expected direction. Overall, females appeared to be more interested in their mate's success than their own and were more selective in the choice of mates. Males gave more evidence of competitiveness and tended to mate polygamously. The results are compatible with theoretical expectations about mating strategies (Buss, 1987; Rushton et al., 1984; Thiessen and Gregg, 1980). A number of problems with regard to the application of sociobiological and sociocultural theories to human behavior are discussed.

Adolescent↗

The cryptochrome (cry) gene and a mating isolation mechanism in tephritid fruit flies.

Two sibling species of tephritid fruit fly, Bactrocera tryoni and Bactrocera neohumeralis, are differentiated by their time of mating, which is genetically determined and requires interactions between the endogenous circadian clock and light intensity. The cryptochrome (cry) gene, a light-sensitive component of the circadian clock, was isolated in the two Bactrocera species. The putative amino acid sequence is identical in the two species. In the brain, in situ hybridization showed that cry is expressed in the lateral and dorsal regions of the central brain where PER immunostaining was also observed and in a peripheral cell cluster of the antennal lobes. Levels of cry mRNA were analyzed in whole head, brain, and antennae. In whole head, cry is abundantly and constantly expressed. However, in brain and antennae the transcript cycles in abundance, with higher levels during the day than at night, and cry transcripts are more abundant in the brain and antennae of B. neohumeralis than in that of B. tryoni. Strikingly, these results are duplicated in hybrid lines, generated by rare mating between B. tryoni and B. neohumeralis and then selected on the basis of mating time, suggesting a role for the cry gene in the mating isolation mechanism that differentiates the species.

Amino Acid Sequence↗

[Effect of treatment with p-fluorophenylalanine on the mating capacity of industrial polyploid yeasts].

Brewing yeast strains of the genus Saccharomyces uvarum (carlsbergensis) were grown on media containing p-fluorophenylalanine (p-FPA). After the treatment, non-sporulation colonies were selected, and these were mated with haploid strains of flocculent and amylolytic yeasts of the genus Saccharomyces. The selected hybrids, which carried the greater part of the parental genetic markers and produced asci containing 2,3 and 4 spores per ascus, were placed on sporulation medium. Some aspects of the probable action of p-FPA are discussed.

Industrial Microbiology↗

Genetics of nonspecific immunity: I. Bidirectional selective breeding of lines of mice endowed with maximal or minimal inflammatory responsiveness.

The genetic regulation of acute inflammatory reaction (AIR) was studied by the method of bidirectional selective breeding, used to produce a line of mice giving the maximal and a line of mice giving the minimal inflammatory reaction (AIR max and AIR min, respectively). The AIR was triggered by subcutaneous injection of a neutral substrate (suspension of polyacrylamide microbeads), and measured by the leukocyte and serum protein accumulation in the exudate. The two parameters are positively correlated and present a normal frequency distribution. The highly genetically heterogeneous foundation population was produced by the equipoised intercrossing of eight inbred strains of mice, and selective breeding carried out by assortative matings of extreme phenotypes. The response to selection in 11 consecutive generations was highly asymmetrical: a marked AIR increase in the AIR max and no change in the AIR min line occurred. The mean value of realized heritability in the AIR max line was 0.26 and 0.18 for cell and protein concentrations, respectively. The response to selection must have resulted from the interaction of seven to nine independent gene loci endowed with additive effects. The lack of response to selection of the AIR min line is discussed. The large inter-line difference opens new possibilities for studying the biochemistry and molecular genetics of inflammation, and also for investigating the beneficial or detrimental effect of inflammatory responses.

Acute Disease↗

The recovery of nocardial recombinants from broth cultures.

Broth culture media were examined for their ability to support growth and recombination between compatible strains of Nocardia erythropolis. Nutrient(Nut) and peptone-yeast-extract (PY) broths supported the production of recombinants after 36 h of incubation with a maximum recovery of about 6.0 times 10(-7) CFU/ml. Cells mated in trypticase broth (TB) yielded the highest incidence of recombinants (1.0 times 10(-2) CFU/ml) in the absence of parental cell growth. From a chemically defined mating broth (CD), supplemented with limited amounts of the parental-growth requirements, recombinant recovery reached about 1.0 times 10(-4) after 120 h of incubation. The recombinant class types obtained from Nut- or PT-mated strains were predominantly auxotrophic while TB-mated strains produced stable proteotrophs. The high incidence of recombinant types from TB-mated strains was due to growth of selected prototrophic classes. Studies with strains mated in PY broth indicated that the mating event occurs at very low frequencies between older, stationary-phase cells rather than between actively growing, log-phase cells.

Adenine↗

Sexual selection when fertilization is not guaranteed.

Much of the theory of sexual selection assumes that females do not generally experience difficulties getting their eggs fertilized, yet sperm limitation is occasionally documented. How often does male limitation form a selection for female traits that improve their mating rate? The question is difficult to test, because if such traits evolve to be efficient, sperm limitation will no longer appear to be a problem to females. Here, we suggest that changes in choosiness between populations, and in particular between virgin and mated females, offer an efficient way to test this hypothesis. We model the "wallflower effect," that is, changes in female preferences due to time and mortality costs of remaining unmated (for at least some time). We show that these costs cause adaptive reductions of female choice, even if mate encounter rates appear high and females only rarely end their lives unfertilized. We also consider the population consequences of plastic or fixed mate preferences at different mate encounter rates. If mate choice is plastic, we confirm earlier verbal models that virgins should mate relatively indiscriminately, but plastic increase of choosiness in later matings can compensate and intensify sexual selection on the male trait, particularly if there is last male sperm precedence. Plastic populations will cope well with unusual conditions: eagerness of virgins leads to high reproductive output and a relaxation of sexual selection at low population densities. If females lack such plasticity, however, population-wide reproductive output may be severely reduced, whereas sexual selection on male traits remains strong.

Animals↗

Sensitivity of neurons in the auditory midbrain of the grassfrog to temporal characteristics of sound. III. Stimulation with natural and synthetic mating calls.

Using an ensemble of natural and synthetic mating calls we studied the single unit (N = 189) and small neuronal group (91 unit pairs) responsiveness in the auditory midbrain of the grassfrog. We compared this responsiveness to that obtained for toneburst and sine-modulated stimuli in order to reveal the presence of so-called mating call detectors. Under the set of stimuli used 5% of the single units appeared to respond selectively to the natural mating call. Simultaneously recorded units appeared to have in about half of the cases identical properties, in the other half even completely complementary properties were found. These properties are related to the organizational structure of the torus semicircularis. A mechanism of feature extraction based on near coincident firings in simultaneously recorded neurons is presented.

Acoustic Stimulation↗

Distribution of allelic forms of erythrocyte H1 histones in Japanese quail populations divergently selected for amount of weight loss after transient starvation.

Three polymorphic subtypes of erythrocyte histone H1 (H1.a, H1.b, and H1.z) were analyzed using a sodium dodecyl sulfate polyacrylamide gel in quail populations divergently selected for a high (line 1) or low (line 2) reduction in body mass following temporary food withdrawal. Both H1.b and H1.z histone alleles were found to be differently distributed in these populations during the selection period. The frequency of b1 in line 2 was approximately 1.9-2.8 times lower than in line 1 and approached the values in line 1 when the selection was suspended. Similarly, the frequency of allele z2 at locus H1.z increased significantly (about 1.6-2.3 times) in line 2 during selection and returned to the initial values when selection was stopped. On the other hand, allele a0 at locus H1.a was kept at relatively low levels (usually below 0.05) in both lines during selection. At that time its level was approximately three to four times lower than in a random mating control population. When selection was suspended, the frequency of a0 in line 1 increased significantly, approaching the values in the control line, and remained essentially unchanged in line 2. Thus, all three polymorphic histone H1 loci in quail responded through changes in allele frequencies to the breeding selection, which was directed at the amount of body weight loss upon transient starvation. It seems that either H1 histone locus could be linked to loci controlling the rate of body weight reduction following starvation or weight loss during fasting might be influenced by a panel of H1 histone alleles that can contribute to functional differences in avian chromatin.

Alleles↗

A kin-selection approach to the resolution of sex-ratio conflict between mates.

We investigate an instance of conflict between mates over the sex ratio of their brood. We construct a kin-selection model for the evolution of the sex ratio assuming local resource competition (LRC) among females. We explore two basic scenarios: (a) the case where parents make simultaneous sex-ratio decisions (the simultaneous allocation model); and (b) the case where parental sex-ratio decisions occur one after the other (the sequential allocation model). In the simultaneous investment model, resolution of the conflict between mates depends on the extent to which relative paternal contribution influences the brood sex ratio. In the sequential allocation model, fathers determine primary sex-ratio through fertilization bias; then mothers modify the paternal sex-ratio decision by adjusting the level of investment of some resource that contributes to offspring survival. Under the sequential model, a compromise is always achieved; however this compromise favours one perspective or the other, depending on the extent to which maternal investment influences offspring survival.

Animals↗

Predicting rates of inbreeding in populations undergoing selection.

Tractable forms of predicting rates of inbreeding (DeltaF) in selected populations with general indices, nonrandom mating, and overlapping generations were developed, with the principal results assuming a period of equilibrium in the selection process. An existing theorem concerning the relationship between squared long-term genetic contributions and rates of inbreeding was extended to nonrandom mating and to overlapping generations. DeltaF was shown to be approximately (1)/(4)(1 - omega) times the expected sum of squared lifetime contributions, where omega is the deviation from Hardy-Weinberg proportions. This relationship cannot be used for prediction since it is based upon observed quantities. Therefore, the relationship was further developed to express DeltaF in terms of expected long-term contributions that are conditional on a set of selective advantages that relate the selection processes in two consecutive generations and are predictable quantities. With random mating, if selected family sizes are assumed to be independent Poisson variables then the expected long-term contribution could be substituted for the observed, providing (1)/(4) (since omega = 0) was increased to (1)/(2). Established theory was used to provide a correction term to account for deviations from the Poisson assumptions. The equations were successfully applied, using simple linear models, to the problem of predicting DeltaF with sib indices in discrete generations since previously published solutions had proved complex.

Inbreeding↗

Disruptive sexual selection against hybrids contributes to speciation between Heliconius cydno and Heliconius melpomene.

Understanding the fate of hybrids in wild populations is fundamental to understanding speciation. Here we provide evidence for disruptive sexual selection against hybrids between Heliconius cydno and Heliconius melpomene. The two species are sympatric across most of Central and Andean South America, and coexist despite a low level of hybridization. No-choice mating experiments show strong assortative mating between the species. Hybrids mate readily with one another, but both sexes show a reduction in mating success of over 50% with the parental species. Mating preference is associated with a shift in the adult colour pattern, which is involved in predator defence through Müllerian mimicry, but also strongly affects male courtship probability. The hybrids, which lie outside the curve of protection afforded by mimetic resemblance to the parental species, are also largely outside the curves of parental mating preference. Disruptive sexual selection against F(1) hybrids therefore forms an additional post-mating barrier to gene flow, blurring the distinction between pre-mating and post-mating isolation, and helping to maintain the distinctness of these hybridizing species.

Animals↗

Mating patterns of different Adh genotypes of Drosophila melanogaster. II. Testing rare-male mating advantage.

In this paper we analyze the rare-male mating advantage at the Adh locus in Drosophila melanogaster by three different statistical methods: chi 2 test [classically applied for this kind of experiment (Ehrman, 1966; Spiess and Schwer, 1978; Markow et al., 1980)], linear regression (Adams-Duncan's test), and estimation of relative male mating success (Anderson-Browns's test). With the chi 2 test, a rare-male SS mating advantage could be speculated in the FF vs. SS comparison, but when the other two statistical methods were applied, this effect was not evident. However, in the FF vs. SS comparison, FF males show a much greater mating success than SS males in experiments with a 1:1 or 1:9 ratio. In the FF vs. FS and FS vs. SS comparisons, differences in mating success and frequency dependent selection were not found. The fact that a rare-male mating advantage is not detected for any of the Adh genotypes seems to indicate that this type of frequency-dependent selection is not a major component of the mechanism to maintain polymorphism at the Adh locus.

Animals↗

A comparison of the reproductive and growth performances of offspring from broiler breeder males selected for early growth rate using artificial insemination and unselected males kept on deep litter.

Broiler breeder males were selected for early growth rate at 5 weeks of age (average weight + 0.5 SD). The reproductive and growth performances of the offspring of these males using artificial insemination with stored semen was compared with those from non-selected males kept on deep litter under conditions of natural mating. On three different occasions (31, 41 and 51 weeks of age), the eggs of the two treatments were incubated and, although the reproductive performances showed little difference over the entire period, hatchability of eggs set was significantly (p less than 0.01) higher for the field hens on the first occasion. The opposite was true after 50 weeks of age due to a serious decline in fertility in natural mating. At all periods, the selected males produced significantly (p less than 0.01) heavier offspring at six weeks of age.

Animals↗

Theoretical influence of female mating status and remating propensity on male sperm allocation patterns.

Theoretical models predict that males should allocate more sperm in matings where the immediate risk of sperm competition is high. It has therefore often been argued that males should invest less sperm in matings with virgin females compared with matings with already mated females. However, with relatively polyandrous females, high sperm competition risk will covary with high sperm competition intensity leading to more unpredictable conditions, as high competition intensity should favour smaller ejaculates. With the use of a genetic algorithm, we found that males should allocate more sperm in matings with virgin females when female mating frequency is relatively high, whereas low remating rates will select for higher effort in matings with nonvirgin females. At higher remating rates, first male sperm precedence favours larger ejaculates in matings with virgin females and second male precedence favours the reverse. These results shed some light on several findings that have been difficult to explain adaptively by the hitherto developed theory on sperm allocation.

Algorithms↗

Why do female migratory birds arrive later than males?

1. In migratory birds males tend to arrive first on breeding grounds, except in sex-role reversed species. The two most common explanations are the rank advantage hypothesis, in which male-male competition for breeding sites drives stronger selection for early arrival in males than females, and the mate opportunity hypothesis, which relies on sexual selection, as early arrival improves prospects of mate acquisition more for males than for females. 2. To date, theoretical work has focused on selection for early arrival within a single sex, usually male. However, if fitness depends on territory quality, selection for early arrival should operate on both sexes. Here we use two independent modelling approaches to explore the evolution of protandry (male-first arrival) and protogyny (female-first arrival) under the rank advantage and mate opportunity hypotheses. 3. The rank advantage hypothesis, when operating alone, fails to produce consistent patterns of protandry, despite our assumption that males must occupy territories before females. This is because an individual of either sex benefits if it out-competes same-sex competitors. Rather than promoting protandry, the rank advantage mechanism can sometimes result in protogyny. Female-female competition is stronger than male-male competition early in the season, if females compete for a resource (territories occupied by males) that is initially less common than the resource of interest to males (unoccupied territories). 4. Our results support the mate opportunity hypothesis as an explanation of why protandry is the norm in migratory systems. Male-biased adult sex ratios and high levels of sperm competition (modelled as extra-pair young: EPY) both produce protandry as a result of sexual selection. Protogyny is only observed in our models with female-biased sex ratios and low EPY production. 5. We also show that the effects of sex ratio biases are much stronger than those of EPY production, explore the evidence for sex ratio biases and extra-pair paternity in migratory species and suggest future research directions.

Animal Migration↗