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Shared forces of sex chromosome evolution in haploid-mating and diploid-mating organisms: Microbotryum violaceum and other model organisms.

It is usually posited that the most important factors contributing to sex chromosome evolution in diploids are the suppression of meiotic recombination and the asymmetry that results from one chromosome (the Y) being permanently heterozygous and the other (the X) being homozygous in half of the individuals involved in mating. To distinguish between the roles of these two factors, it would be valuable to compare sex chromosomes in diploid-mating organisms and organisms where mating compatibility is determined in the haploid stage. In this latter group, no such asymmetry occurs because the sex chromosomes are equally heterozygous. Here we show in the fungus Microbotryum violaceum that the chromosomes carrying the mating-type locus, and thus determining haploid-mating compatibility, are rich in transposable elements, dimorphic in size, and carry unequal densities of functional genes. Through analysis of available complete genomes, we also show that M. violaceum is, remarkably, more similar to humans and mice than to yeast, nematodes, or fruit flies with regard to the differential accumulation of transposable elements in the chromosomes determining mating compatibility vs. the autosomes. We conclude that restricted recombination, rather than asymmetrical sheltering, hemizygosity, or dosage compensation, is sufficient to account for the common sex chromosome characteristics.

Base Sequence↗

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↗

Genotypic differences in mating success and the maintenance of the alcohol dehydrogenase polymorphism in Drosophila melanogaster: No evidence for overdominance or rare genotype mating advantage.

Strains of Drosophila melanogaster having different alcohol dehydrogenase (Adh) genotypes (FF, FS, or SS) were assembled in a mating chamber in varying ratios, and the mating successes were recorded. In experiments with a 25:25 ratio, the FF males succeeded in mating more than did FS and SS males, while the FS males surpassed the SS males. As for the females, FF also surpassed SS. In experiments with a 5:45 or 45:5 ratio, some differences from the 25:25 ratio occurred, but in these cases the rare genotypes were at a disadvantage. In one case, female genotypes (FF vs. SS) displayed a difference in mating latency time, but male genotypes did not. The findings did not suggest that rare genotype mating advantage and overdominance in mating success play a role in the maintenance of the Adh polymorphism.

Alcohol Oxidoreductases↗

Common regulation of feeding and mating in Aplysia fasciata: pheromones released by mating and by egg cordons increase feeding behavior.

We examined whether pheromones released by reproductive behaviors (mating and egg-laying) affect feeding behavior. A preliminary experiment demonstrated that the quantity of food eaten can be used to measure the effects of pheromones on feeding. Using this measure, we then showed that Aplysia that were prevented from mating, but that were in the same aquarium as mating conspecifics, eat more food than do Aplysia in a medium lacking mating animals. Mating and feeding were not temporally correlated, indicating that pheromones released by mating probably do not initiate feeding, but rather modulate feeding after it has begun. Aplysia that were in the same aquarium as freshly deposited egg cordons also ate more than did animals in a medium lacking eggs.

Animals↗

Underground mating in the fiddler crab Uca tetragonon: the association between female life history traits and male mating tactics.

Brood size and other life-history traits of females affect male investment in mating. Female Uca tetragonon, producing relatively small broods, were attracted to the burrows of males for underground mating (UM) while carrying eggs. Most UM females released larvae and ovulated new broods during the pairing, averaging 3.9 days. While a female was incubating one brood, another brood was developing within the ovaries because the females were feeding adequately during incubation. These findings suggest that in U. tetragonon, a small-brood species, females increase the total number of broods produced by breeding continually. In contrast, in large-brood species, feeding by ovigerous females is relatively brief and not enough to prepare the next brood during incubation, inducing temporal separation between incubation and brood production. Unlike females in other ocypodids where females with large broods remain in the breeding burrows of males, most female U. tetragonon left the male after UM. Wandering in female U. tetragonon after the pairs separate may occur because their small broods are adequately protected by an abdominal flap. Relative brood size probably determines the vulnerability of the incubated broods to the females' surface behavior. Hence, male reproductive success in large-brood species may decrease greatly if males expel their mates after ovulation, although this is not necessarily so in small-brood species. Whether the male drives away the female or not may depend on which behavior within either small- or large-brood species yields the greater male reproductive success. In U. tetragonon some females extruded eggs in their own burrows after surface mating as well as in males' burrows after UM. It was unclear whether females chose a male with a larger burrow as an UM mate unlike several large-brood species. Burrows of both UM males and ovigerous females in U. tetragonon were relatively smaller than those in some large-brood species, indicating that incubation of small broods does not require large burrows. Rather than benefits of UM by female choice, wandering resulting from intersexual conflict, and sperm competition may explain why some females mate in males' burrows in this small-brood species.

Journal Article↗

Male mating success in an aquatically mating pinniped, the harbour seal (Phoca vitulina), assessed by microsatellite DNA markers.

Similar to many other pinniped species, harbour seals (Phoca vitulina) mate exclusively at sea. Here we present the first attempt to measure male mating success in an aquatically mating pinniped. Male mating success was estimated by paternity analysis in two cohorts of pups born at Sable Island, Nova Scotia, Canada, using microsatellite DNA markers. The genotypes of 275 pups born in 1994 and 1995 were compared to those of 90 candidate males at six microsatellite loci using a likelihood approach to resolve paternity. Paternity could be assigned for two, 22, 40 and 85 pups at confidence levels of 95, 80, 65 and 50%, respectively. Most successful males were assigned the paternity of a single offspring, suggesting a low variance in male mating success relative to most pinniped species. The proportion of paternal half sibs within cohorts and between maternally related sibs estimated by maximum likelihood were not significantly different from zero. It is thus unlikely that most offspring were sired by a small number of highly successful unsampled males, and that female harbour seals do not usually exhibit fidelity to the same male in sequential breeding seasons. A low level of polygyny in Sable Island harbour seals is consistent with predictions based on their breeding ecology, as females are highly mobile and widely dispersed in the aquatic mating environment at Sable Island.

Animals↗

Parental investment, mate choice, and mate quality.

Current theory in sexual selection is extended to predict within-sex variability with regard to selectivity towards mates in different mating systems. Generally, the sex that invests more in the care of each offspring should be more selective of mates than the sex investing less. Within each sex, individuals of low male quality should be less selective than individuals of high quality, but there should be less variation in selectivity among individuals of the sex investing more. When only one sex contributes parental care, however, individuals of that sex should be uniformly selective, while the other sex is expected to mate indiscriminately. Using feral pigeons (Columba livia), these hypotheses are tested for the case in which both sexes contribute substantial parental care, but in which females contribute more than males. As predicted, females were found to be more selective of mates than males were. On certain criteria, males of lower quality were less selective of mates than males of higher quality.

Animals↗

A suppressor of mating-type locus mutations in Saccharomyces cerevisiae: evidence for and identification of cryptic mating-type loci.

A mutation has been identified that suppresses the mating and sporulation defects of all mutations in the mating-type loci of S. cerevisiae. This suppressor, sir1-1, restores mating ability to mat alpha 1 and mat alpha 2 mutants and restores sporulation ability to mat alpha 2 and mata1 mutants. MATa sir1-1 strains exhibit a polar budding pattern and have reduced sensitivity to alpha-factor, both properties of a/alpha diploids. Furthermore, sir1-1 allows MATa/MATa, mat alpha 1/mat alpha/, and MAT alpha/MAT alpha strains to sporulate efficiently. All actions of sir1-1 are recessive to SIR1. The ability of sir1-1 to supply all functions necessary for mating and sporulation and its effects in a cells are explained by proposing that sir1-1 allows expression of mating type loci which are ordinarily not expressed. The ability of sir1-1 to suppress the mat alpha 1-5 mutation is dependent on the HMa gene, previously identified as required for switching of mating types from a to alpha. Thus, as predicted by the cassette model, HMa is functionally equivalent to MAT alpha since it supplies functions of MAT alpha. We propose that sir1-1 is defective in a function. Sir ("Silent-information regulator"), whose role may be to regulate expression of HMa and HM alpha.

Chromosome Mapping↗

Mate choice assays and mating propensity differences in natural yeast populations.

In sexual microbes, mating occurs by fusion of individual cells. This complete fitness investment suggests that cell behaviour could potentially mediate prezygotic isolation between microbial species, a topic about which very little is known. To investigate this possibility, we conducted individual cell mate choice trials and mass-culture mating propensity assays with isolates from sympatric natural populations of the closely related yeasts Saccharomyces cerevisiae and Saccharomyces paradoxus. Although we found no evidence for active species recognition in mate choice, we observed a marked difference in mating propensity between these two species. We briefly discuss the possibility that this mating propensity difference may contribute to reproductive isolation between S. cerevisiae and S. paradoxus in nature.

Reproduction↗

Variable assortative mating in replicate mating trials using Drosophila melanogaster populations derived from contrasting opposing slopes of 'Evolution Canyon', Israel.

Significant assortative mating in laboratory studies has been previously shown between two populations of Drosophila melanogaster collected from micro-climactically contrasting and opposing slopes of 'Evolution Canyon' (Lower Nahal Oren, Israel; Korol et al., 2000). Coupled with evidence that the two populations are adapted to their respective environments, this has been suggested as a rare example of ongoing behaviourally mediated speciation occurring in the face of gene flow. Reproductive isolation between these populations, however, has never been confirmed by replicate experiments in an independent laboratory. For this reason, we tested recent collections of these populations for premating isolation in both the original (Haifa) and a new (Burnaby) laboratory under a variety of experimental protocols. Although non-random mating was found in the majority of trials conducted in Haifa, we were unable to replicate these strong results in Burnaby. Most notably, we failed to detect assortative mating in four separate double choice experiments. Significant non-random mating was detected, however, in three of six single choice experiments in Burnaby, suggesting that the populations are behaviourally differentiated in some manner. Why nonrandom mating was weaker in Burnaby than Haifa is not understood, but suggests that assortative mating may be sensitive to unknown environmental factors.

Animals↗

Courtship Disruptions and Male Mating Strategies: Examples from Female-Defense Mating Systems.

Males frequently interrupt the copulation attempts of other males, and these courtship disruptions may limit the extent to which a few males are able to monopolize mating access to females. Males actively defend sexually receptive females in many species in which females form dense aggregations during the breeding season. Across and within such species there is considerable variation in the mating tactics adopted by males, with males in some cases defending groups of females and in other cases sequentially consorting with individual females. Colonial blackbirds have been central to studying this mating system, and we develop a conceptual model for how courtship disruption may account for variation in male mating tactics in this group. Our model assumes that the frequency of disruptions increases with greater colony size. As a consequence, successful copulations are less likely to occur at large colonies than at small colonies, and males are expected to switch from defending multiple females at the colony to consorting individual females away from it. Results from two species of blackbird support the basic assumptions of this model. In one species, the Montezuma oropendola, disruptions occur rarely and males defend groups of females, whereas in the other species, the yellow-rumped cacique, disruptions are frequent and males defend single females. Moreover, consistent with a key prediction, within each species, males associated with small colonies remain at the colony and defend groups of females, whereas males spend little time defending groups of females at large colonies and rarely attempt copulations there. This model has the potential to explain variation in male mating strategies and female monopolization for other taxa in which females form breeding aggregations.

Icteridae↗

Species-specific and mating type-specific DNA regions adjacent to mating type idiomorphs in the genus Neurospora.

Mating type idiomorphs control mating and subsequent sexual development in Neurospora crassa and were previously shown to be well conserved in other Neurospora species. The centromere-proximal flanks of the A and a idiomorphs, but not the distal flanks from representative heterothallic, pseudohomothallic, and homothallic Neurospora species contain apparent species-specific and/or mating type-specific sequences adjacent to the well-conserved idiomorphs. The variable flank is bordered by regions that are highly homologous in all species. The sequence of approximately 1 kb immediately flanking the conserved idiomorphs of each species was determined. Sequence identity between species ranged from 20% (essentially unrelated) to > 90%. By contrast, the mt-A1 gene shows 88-98% identity. Sequence and hybridization data also show that the centromere-proximal flanks are very different between the two mating types for N. intermedia, N. discreta, and N. tetrasperma, but not for N. sitophila and N. crassa. The data suggest a close evolutionary relationship between several of the species; this is suppported by phylogenetic analysis of their respective mt-A1 genes. The origin of the variable regions adjacent to the evolutionarily conserved mating type idiomorphs is unknown.

Base Sequence↗

Incipient sexual isolation in the nasuta-albomicans complex of Drosophila: mating preference in male-, female- and multiple-choice mating experiments.

Interracial divergence is an important facet of speciation. The nasuta-albomicans complex of Drosophila with sixteen morphologically identical, karyotypically different but cross-fertile races is an excellent system to study a few dimensions of raciation. Drosophila nasuta nasuta, Drosophila nasuta albomicans, Cytorace 1, Cytorace 2, Cytorace 3 and Cytorace 4 of this subgroup have been subjected to male-, female- and multiple-choice mating experiments. Out of 8456 crosses conducted, 7185 had successful matings. The overall impression is that mating is far from random amongst these six closely related races of the nasuta-albomicans complex. The males of D. n. albomicans, Cytorace 1 and Cytorace 4 in male-choice, the females of Cytorace 1 and Cytorace 2 in female-choice, and the males and females of D. n. nasuta, D. n. albomicans, Cytorace 1 and Cytorace 4 against the males and females of Cytorace 2 in multiple-choice experiments, had significantly more homogamic matings than expected. Thus in this study of evolutionary experimentation on raciation under laboratory conditions, we have documented the initiation of preference for conspecific matings among closely related and independently evolving members of the nasuta-albomicans complex of Drosophila.

Animals↗

Mating-type loci segregate aberrantly in Phytophthora infestans but normally in Phytophthora parasitica: implications for models of mating-type determination.

In the oomycete, Phytophthora infestans, mating type is determined by a locus that segregates in a non-Mendelian manner consistent with its linkage to a system of balanced lethals. The significance of this unusual phenomenon was addressed by studying the segregation patterns of DNA markers linked to mating type in the related species, P. parasitica. This was done using loci identified by either RAPD analysis of P. parasitica crosses or by cross-hybridization with RFLP markers linked to mating type in P. infestans. The resulting data revealed that, unlike P. infestans, mating type in P. parasitica was regulated by a locus displaying Mendelian segregation. An improved model for mating-type determination in Phytophthora is presented.

Chromosome Mapping↗

Cortisol response in heifers to artificial insemination, natural mating, and no mating at estrus.

Eight mixed-breed dairy heifers were used in a random block design to study the stress-producing effect of various types of mating on heifers. Treatments consisted of artificial insemination, natural mating, or not mating at estrus. Stress was evaluated by measuring blood plasma cortisol. Blood samples were collected by indwelling jugular catheters at 60 and 30 min before treatment; immediately before treatment (0 min); and 5, 15, 30, 60, 120 and 180 min after treatment. Means for cortisol were not significant for treatment or treatment across time. Mean cortisol response to the artificial insemination, natural mating and no mating treatments were 5.36, 8.05 and 5.56 ng/ml, respectively. These results suggest that the use of artificial insemination does not impose and added stress at estrus.

Journal Article↗

Healing of mat mutations and control of mating type interconversion by the mating type locus in Saccharomyces cerevisiae.

Homothallic yeasts switch cell types (mating types a and alpha) at high frequency by changing the alleles of the mating type locus, MATa and MATalpha. We have proposed in the cassette model that yeast cells contain silent MATa and MATalpha blocs ("cassettes"), copies of which can be substituted at the mating type locus for the resident information. The existence of silent cassettes was originally proposed to explain efficient switching of a defective MATalpha locus (matalpha) to a functional MATalpha locus. We report here that this "healing" of mat mutations is a general property of the mating type interconversion system and is not specific to the class of matalpha mutations studied earlier: a defective MATa (mata1) switches readily to MATa and various matalpha loci switch readily to MATalpha. These observations satisfy the prediction of the cassette model that all mutations within MATa and MATalpha be healed. These studies also identify MAT functions that control the switching process: the same functions known to promote sporulation and prevent mating in a/alpha cells also inhibit the switching system in a/alpha cells. Finally, we present additional characterization of a natural variant of MATalpha, MATalpha-inc [Takano, I., Kusumi, T. & Oshima, Y. (1973) Mol. Gen. Genet. 126, 19-28] that is insensitive to switching. Our observation that MATalpha-inc acts in cis suggests that it may be altered in a site concerned with excision of MATalpha-inc or its replacement by another cassette.

Journal Article↗

Degradation of mating factor by alpha-mating type cells of Saccharomyces cerevisiae.

The change of the mating factor activity during the culture of Saccharomyces cerevisiae X-2180 1B, an alpha-mating type haploid strain, were followed. The activity increased rapidly during the exponential phase of growth, reached a maximum during the early stationary phase and then decreased. Oligopeptides comprising partial sequences of the mating factor were isolated from the culture fluids at various phases of cell growth. We concluded that the mating factor, a tridecapeptide, was degraded during culture into two peptides, Trp-His-Trp-Leu-Gln-Leu and Lys-Pro-Gly-Gln-Pro-Met-Tyr, by cleavage of the peptide bond between Leu-6 and Lys-7 of the mating factor. A dodecapeptide lacking the N-terminal Trp residue was not detected at any stage of cell growth examined.

Amino Acid Sequence↗