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Molecular evolution of the GapC gene family in Amsinckia spectabilis populations that differ in outcrossing rate.

Molecular evolutionary analysis of the glyceraldehyde 3-phosphate dehydrogenase (GapC) gene family was conducted in the plant genus Amsinckia (Boraginaceae), a group that exhibits marked variation in the mating system. GapC genes in this group differ from those of Arabidopsis thaliana in terms of both intron size and number. Phylogenetic and Southern hybridization analyses suggest the presence of multiple GapC loci, each defined by a set of base substitutions that are in strong linkage disequilibrium. One species of Amsinckia, A. spectabilis, was studied in some detail. This species consists of selfing (A. s. spectabilis) and outcrossing (A. s. microcarpa) varieties. Two selfing populations and one outcrossing population sample were analyzed in detail for variation at one of the members of this gene family. GapC3. A reduction in number of GapC3 haplotypes and level of genetic diversity was observed in the selfing populations of A. spectabilis. GapC3 in the outcrossing population (but not the two selfing populations) exhibited a significant departure from neutrality in the direction of an excess of singletons. These results are discussed in the context of forces acting on sequence evolution in populations with different mating systems.

Amsinckia↗

Pollen discounting and the evolution of selfing in Arenaria uniflora (caryophyllaceae).

Although most models of mating system evolution assign a central role to the male transmission advantage of selfing genotypes, empirical data on the male fitness consequences of increased self-pollination are still uncommon. Here, I use measures of pollen import and export by focal plants in genotyped arrays to investigate the effects of floral morphology and pollination environment on self and outcross male function. Plants from an autogamous population of Arenaria uniflora (Caryophyllaceae) exhibit complete pollen discounting relative to closely related outcrossers, as do morphologically intermediate F1 hybrids between the two populations. However, the low cumulative male fitness of hybrids probably results from reduced pollen number or competitive ability, rather than a nonlinear relationship with floral morphology. When surrounded by selfers, plants from the outcrosser population self-fertilize at nearly the same rate as selfers (>80%), but have much lower self male fitness due to reduced fruit set. Because outcross siring success is also extremely low (<8%) in this treatment, these mate-limited outcrossers are at male fitness disadvantage to both pseudocleistogamous selfers and nonlimited outcrossers. The relative male fitness of plants with different mating systems appears dependent on the ecological context, as well as on morphological trade-offs.

Biological Evolution↗

Genetic evidence for female-biased dispersal and gene flow in a polygynous primate.

Many models of sex-biased dispersal predict that the direction of sex-bias depends upon a species' mating system. In agreement with this, almost all polygynous mammals show male-biased dispersal whereas largely monogamous birds show female-biased dispersal (FBD). The hamadryas baboon (Papio hamadryas hamadryas) is polygynous and so dispersal is predicted to be male biased, as is found in all other baboon subspecies, but there are conflicting field data showing both female and male dispersal. Using 19 autosomal genetic markers genotyped in baboons from four Saudi Arabian populations, we found strong evidence for FBD in post-dispersal adults but not, as expected, in pre-dispersal infants and young juveniles, when we compared male and female: population structure (F(st)), inbreeding (F(is)), relatedness (r), and the mean assignment index (mAIc). Furthermore, we found evidence for female-biased gene flow as population genetic structure (F(st)), was about four times higher for the paternally inherited Y, than for either autosomal markers or for maternally inherited mtDNA. These results contradict the direction of sex-bias predicted by the mating system and show that FBD has evolved recently from an ancestral state of male-biased dispersal. We suggest that the cost-benefit balance of dispersal to males and females is tightly linked to the unique hierarchical social structure of hamadryas baboons and that dispersal and social organization have coevolved.

Animal Migration↗

Genetic diversity and geographic differentiation in Tacca chantrieri (Taccaceae): an autonomous selfing plant with showy floral display.

BACKGROUND AND AIMS: Despite considerable investment in elaborate floral displays, Tacca chantrieri populations are predominantly selfing. It is hypothesized that this species might possess considerable spatial or temporal variation in outcrossing rates among populations. To test this hypothesis, genetic variability and genetic differentiation within and among T. chantrieri populations were investigated to find out if they are in agreement with expectations based on a predominantly inbred mating system. METHODS: Genetic diversity was quantified using inter-simple sequence repeats (ISSR) in 303 individuals from 13 populations taken from known locations of T. chantrieri in China, and from one population in Thailand. KEY RESULTS: Of the 113 primers screened, 24 produced highly reproducible ISSR bands. Using these primers, 160 discernible DNA fragments were generated, of which 145 (90.62 %) were polymorphic. This indicated considerable genetic variation at the species level. However, there were relatively low levels of polymorphism at population levels, with percentages of polymorphic bands (PPB) ranging from 8.75 % to 55 %. A high level of genetic differentiation among populations was detected based on different measures (Nei's genetic diversity analysis: G(ST) = 0.5835; AMOVA analysis: F(ST) = 0.6989). Furthermore, based on levels of genetic differentiation, the 14 populations clustered into two distinct groups separated by the Tanaka Line. CONCLUSIONS: High levels of differentiation among populations and low levels of diversity within populations at large spatial scales are consistent with earlier small-scale studies of mating patterns detected by allozymes which showed that T. chantrieri populations are predominantly selfing. However, it appears that T. chantrieri has a mixed-mating system in which self-fertilization predominates, but there is occasional outcrossing. Significant genetic differences between the two distinct regions might be attributed to vicariance along the Tanaka Line. Finally, possible mechanisms of geographic patterns based on genetic differentiation of T. chantrieri are discussed.

Dioscoreaceae↗

Regulatory elements required for development of caenorhabditis elegans hermaphrodites are conserved in the tra-2 homologue of C. remanei, a male/female sister species.

The Caenorhabditis elegans hermaphrodite is essentially a female that produces sperm. In C. elegans, tra-2 promotes female fates and must be repressed to achieve hermaphrodite spermatogenesis. In an effort to learn how mating systems evolve, we have cloned tra-2 from C. remanei, the closest gonochoristic relative of C. elegans. We found its structure to be similar to that of Ce-tra-2 but its sequence to be divergent. RNA interference demonstrates that Cr-tra-2 promotes female fates. Two sites of tra-2 regulation are required for the onset of hermaphrodite spermatogenesis in C. elegans. One, the MX region of TRA-2, is as well conserved in C. remanei as it is in C. briggsae (another male/hermaphrodite species), suggesting that this control is not unique to hermaphrodites. Another, the DRE/TGE element of the tra-2 3' UTR, was not detected by sequence analysis. However, gel-shift assays demonstrate that a factor in C. remanei can bind specifically to the Cr-tra-2 3' UTR, suggesting that this translational control is also conserved. We propose that both controls are general and do not constitute a novel "switch" that enables sexual mosaicism in hermaphrodites. However, subtle quantitative or qualitative differences in their employment may underlie differences in mating system seen in Caenorhabditis.

3' Untranslated Regions↗

Genetic versus phenotypic models of selection: can genetics be neglected in a long-term perspective?

Game theoretical concepts in evolutionary biology have been criticized by populations geneticists, because they neglect such crucial aspects as the mating system or the mode of inheritance. In fact, the dynamics of natural selection does not necessarily lead to a fitness maximum or an ESS if genetic constraints are taken into account. Yet, it may be premature to conclude that game theoretical concepts do not have a dynamical justification. The new paradigm of long-term evolution postulates that genetic constraints, which may be dominant in a short-term perspective, will in the long run disappear in the face of the ongoing influx of mutations. Two basic results (see Hammerstein; this issue) seem to reconcile the dynamical approach of long-term population genetics with the static approach of evolutionary game theory: (1) only populations at local fitness optima (Nash strategies) can be long-term stable; and (2) in monomorphic populations, evolutionary stability is necessary and sufficient to ensure long-term dynamic stability. The present paper has a double purpose. On the one hand, it is demonstrated by fairly general arguments that the scope of the results mentioned above extends to non-linear frequency dependent selection, to multiple loci, and to quite general mating systems. On the other hand, some limitations of the theory of long-term evolution will also be stressed: (1) there is little hope for a game theoretical characterization of stability in polymorphic populations; (2) many interesting systems do not admit long-term stable equilibria; and (3) even if a long-term stable equilibrium exists, it is not at all clear whether and how it is attainable by a series of gene substitution events.

Biological Evolution↗

Two-, three-, and four-breed rotational crossbreeding of beef cattle: carcass traits.

Carcass data from 1,494 straightbred and rotational crossbred steers were collected over four generations. Mating systems included straightbreds (Angus [A], Brahman [B], Charolais [C], and Hereford [H]); two-breed rotations (A-B, C-B, and H-B); three-breed rotations (A-B-C, A-B-H, and B-C-H); and a four-breed rotation (A-B-C-H). Steers were randomly allocated to one of four postweaning treatments that varied in length of grazing and feeding periods. Treatment and breed group (four straightbreds and seven rotational combinations) significantly influenced hot carcass weight (HCWT), retail yield (RY), longissimus muscle area (LM), fat thickness (FT), marbling score (MS), USDA quality grade (QG), and Warner-Bratzler shear force (WBS). Feeding for longer periods resulted in greater (P < .05) HCWT, RY, FT, and MS, higher QG, and lower (P < .05) WBS values. Among the straightbreds, C was heavier and larger for HCWT, RY, and LM (P < .01), whereas A and H had greater (P < .01) FT and MS. The B was similar to H for HCWT, RY, and LM and to C for FT but ranked last (P < .01) for MS and WBS. Three- and four-breed rotational mating systems were superior (P < .05) to the two-breed rotation for HCWT, RY, and LM but were similar for FT, MS, and WBS. Rotational combinations exceeded (P < .05) the straightbreds for all carcass traits except MS.

Adipose Tissue↗

Genetic population structure of the Greater Bilby Macrotis lagotis, a marsupial in decline.

The Greater Bilby has shown a rapid decline in range during this century and now occupies only a small isolated area in south-western Queensland (QLD) and a larger, but mostly low-density area in the north-western deserts of the Northern Territory (NT) and Western Australia (WA). We have examined variation in the control region of mitochondrial DNA (mtDNA) and at nine microsatellite loci in order to investigate the extent of current and historical subdivision across the species range, and to provide a preliminary assessment of genetic structuring and mating system on a finer scale within the QLD population. Both mtDNA and microsatellite loci had substantial variation within and among populations, with mtDNA divergence being greater between QLD and NT than between NT and WA. The QLD population had two unique and divergent mtDNA lineages, but there was no evidence for strong phylogeographical structure across the range. The available evidence suggests that the bilby should be considered as a single Evolutionarily Significant Unit consisting of multiple Management Units. Augmentation of the remnant QLD population from the NT does not appear necessary at this stage, at least not on genetic grounds. Finer-scale analysis of microsatellite variation for two QLD colonies revealed a deficiency of heterozygotes and significantly greater relatedness within than between colonies. However, structuring was observed only for males; relatedness values for females did not depart from those expected under panmixia. Parentage exclusion analysis for one colony allowed the construction of a partial pedigree which indicated strong polygyny, with one male fathering all but one of the eight offspring assigned. The extent to which fine-scale genetic structuring and differences between sexes is due to sex-biased dispersal vs. effects of mating system remain to be determined.

Animals↗

Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae).

Previous studies have demonstrated genetic variation for resistance to insect herbivores and host plant quality. The effect of plant mating system, an important determinant of the distribution of genetic variation, on host plant characteristics has received almost no attention. This study used a controlled greenhouse experiment to examine the effect of self- and cross-pollination in Mimulus guttatus (Scrophulariaceae) on resistance to and host plant quality for the xylem-feeding spittlebug Philaenus spumarius (Homoptera: Cercopidae). Spittlebugs were found to have a negative effect on two important fitness components in M. guttatus, flower production and above ground biomass. One of two M. guttatus populations examined showed a significant interaction between the pollination and herbivore treatments. In this case, the detrimental effects of herbivores on biomass and flower production were much more pronounced in inbred (self) plants. The presence of spittlebug nymphs increased inbreeding depression by as much as three times. Pollination treatments also had significant effects on important components of herbivore fitness, but these effects were in opposite directions in our two host plant populations. Spittlebug nymphs maturing on self plants emerged as significantly larger adults in one of our host plant populations, indicating that inbreeding increased host plant quality. In our second host plant population, spittlebug nymphs took significantly longer to develop to adulthood on self plants, indicating that inbreeding decreased host plant quality. Taken together these results suggest that the degree of inbreeding in host plant populations can have important and perhaps complex effects on the dynamics of plant-herbivore interactions and on mating-system evolution in the host.

Animals↗

Pollination in the marine realm: microsatellites reveal high outcrossing rates and multiple paternity in eelgrass Zostera marina.

The mating system was examined in two annual populations of eelgrass (Zostera marina), a marine angiosperm displaying subaqueous pollination. Multilocus genotyping using microsatellite DNA markers allowed the assessment of the pollen source based on single progeny as units of observation. Outcrossing was detectable by the presence of non-maternal alleles at one or more of the loci. In outcrossing cases, three microsatellite alleles were present in unripe seeds, consisting of both maternal alleles and the paternal allele composing the triploid primary endosperm. In ripe seeds, only the diploid embryonal genotype was amplifiable by PCR. Two intertidal populations situated in the German Wadden Sea were almost entirely outcrossing (t +/- SE 0.96 +/- 0.03, N=60 and 0.97 +/- 0.029, N=37). Because of the high polymorphism displayed by the eight chosen microsatellites, representing a total of 69 and 76 alleles, the likelihood of erroneously inferring selfing was small (alpha=0.0026 and 0.0007). In order to study the correlation of paternity, the coefficient of relatedness was determined within sibships. Relatedness (r +/- SE) was calculated as 0.357 +/- 0.059 and 0.343 +/- 0.037, indicating multiple paternities within inflorescences. Small amounts of tissue (< or = 0.1 mg) such as the developing seeds of recently fertilized ovules, were sufficient for PCR-amplification. Hence, PCR-based methods, such as multilocus microsatellite genotyping, allow the detection of pollen origin early in the development of progeny. They will be useful to distinguish postfertilization processes such as selective abortion and germination from other prefertilization determinants of plant mating systems.

Crosses, Genetic↗

Mating communication systems of four Plusiinae species distributed in Japan: identification of the sex pheromones and field evaluation.

The sex pheromones of four Plusiinae species in the family Noctuidae, Ctenoplusia albostriata (CA), Macdunnoughia purissima (MP), Syngrapha ain (SA), and Diachrysia stenochrysis (DS) were identified by GC-EAD and GC-MS analyses. These were subsequently evaluated by a series of field trials in different ecological habitats. CA females produced (Z)-5-decenyl acetate (I), (Z)-7-dodecenyl acetate (II), and (Z)-7-dodecen-1-ol (III) in a ratio of 2:100:13. While the antennae of CA males responded to all three components, neither I nor III was essential to capture males. The MP females produced II, III, and (Z)-5-dodecenyl acetate (IV) in a ratio of 100:80:20, and this blend captured the most MP males. Compounds II and III were also identified from SA females, and mixtures ranging from 4:1 to 2:3 were equally attractive to males. DS females only produced one active compound, (Z)-7-decenyl acetate (V), and DS is the first Plusiinae species identified as using only the omega3-compound and none of omega5-compounds, such as II and III, which are common components of Plusiinae pheromones. Using previously published chemical and biosynthetic data, together with the finding from this study, we propose a classification of the Plusiinae sex pheromones.

Acetates↗

Drosophila sex-peptide stimulates female innate immune system after mating via the Toll and Imd pathways.

Insect immune defense is mainly based on humoral factors like antimicrobial peptides (AMPs) that kill the pathogens directly or on cellular processes involving phagocytosis and encapsulation by hemocytes. In Drosophila, the Toll pathway (activated by fungi and gram-positive bacteria) and the Imd pathway (activated by gram-negative bacteria) lead to the synthesis of AMPs. But AMP genes are also regulated without pathogenic challenge, e.g., by aging, circadian rhythms, and mating. Here, we show that AMP genes are differentially expressed in mated females. Metchnikowin (Mtk) expression is strongly stimulated in the first 6 hr after mating. Sex-peptide (SP), a male seminal peptide transferred during copulation, is the major agent eliciting transcription of Mtk and of other AMP genes. Both pathways are needed for Mtk induction by SP. Furthermore, SP induces additional AMP genes via the Toll (Drosomycin) and the Imd (Diptericin) pathways. SP affects the Toll pathway at or upstream of the gene spätzle, the Imd pathway at or upstream of the gene imd. Mating may physically damage females and pathogens may be transferred. Thus, endogenous stimulation of AMP transcription by SP at mating might be considered as a preventive step to encounter putative immunogenic attacks.

Animals↗

Low reproductive success in territorial male Antarctic fur seals (Arctocephalus gazella) suggests the existence of alternative mating strategies.

Microsatellites were used to conduct an analysis of paternity of Antarctic fur seals (Arctocephalus gazella) from Bird Island, South Georgia. At most, only 28% of pups at our study site could be assigned a father, even though the majority (approximately 90%) of candidate males within this colony were sampled. The behavioural and genetic evidence from this study suggests that a number of alternative mating strategies may exist within this fur seal population. Holding a land-based territory conferred an advantage to male reproductive success. However, this advantage was much smaller than expected from behavioural observations. At least 70% of fur seal pups born at our study site in a given year are not fathered by males who held a territory or were observed copulating with females in the previous year, implying that there exists a pool of males that seldom venture ashore at this site. To explain this discrepancy we suggest that female choice is an integral component of the Antarctic fur seal mating system and that aquatic mating may play a much larger role in the Antarctic fur seal than previously thought.

Animals↗

Multiple pheromone system controlling mating in Drosophila melanogaster.

The signals essential to Drosophila melanogaster courtship include pheromones emitted by the female which stimulate the male to court and pheromones emitted by the courting male which stimulate the female to accept. Genetic variation among these phermones is a common (if not universal) requirement for stimulation of either sex. The signal from the courting male to the female involves both a volatile and a nonvolatile component. The volatile component is associated with loci on the second and/or third chromosomes, while the monvolatile component is associated with the X and/or fourth chromosomes. This widespread distribution in the genome of loci controlling various components in the communication network inevitably results in linkage associations with other loci. The genetic array of gametes was limited. When combined with the negative assortitative mating pattern produced by the stimulation by dissimilar pheromones, linkage disequilibrium creates a strong counterforce to inbreeding during population bottlenecks.

Animals↗

Isolation of a Cryptococcus neoformans serotype A MATa strain from the Italian environment.

Cryptococcus neoformans is a heterothallic basidiomycete which possesses a bipolar mating system based on two mating type alleles, MATa and MATalpha. In the type variety, C. neoformans var. neoformans, both mating types have been found among strains of one serotype, serotype D, whereas only MATalpha was identified after extensive survey of serotype A strains. Serotype A MA Ta appeared to be extinct or to exist only in a vestigial, non-functional form. We report the isolation of a C. n. var. neoformans serotype A MATa strain from the Italian environment. The strain was serotyped by slide agglutination test, genotyped by polymerase chain reaction (PCR) fingerprinting using the (GACA)4 primer, and its haploid state was determined by flow cytometry. The mating type was identified by PCR amplification of the pheromone a gene. In addition, the amplification of the four STE20 alleles, specific for the mating type of serotypes A and D, showed that the strain contains only the MATa locus. By crossing experiments the strain was found to be fertile. The interest in the finding of this fertile isolate is related to the possibility to construct a congenic pair of serotype A MATa/MATalpha strains to be used in genetic and pathogenesis studies.

Cryptococcus neoformans↗