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Monogenic inheritance of apomixis in two Hieracium species with distinct developmental mechanisms.

Apomixis, the asexual formation of seed, has been known in angiosperms for more than a century yet the genetic mechanisms that control this trait remain poorly understood. Most members of the genus Hieracium are apomicts, forming predominantly asexual seed. Some purely sexual forms, however, also exist. In this paper we present a study of the inheritance of apomixis using two apomictic species of Hieracium which utilize very different forms of megagametogenesis. In both cases the progeny inherited apomixis as a monogenic, dominant trait that could be transferred by both haploid and diploid male gametes. In common with observations from other systems, no diploid apomictic progeny were recovered from these crosses. This appears to have been caused by selection against the survival of diploid zygotes, rather than against the mediation of haploid gametes as has been noted in other systems. Crosses between the two apomicts showed that the dominant determinants in the two forms examined were closely linked, possibly allelic. The significance of these data is discussed with respect to current theories on the associative link between gametophytic apomixis and polyploidy.

Alleles↗

Intratetrad mating, heterozygosity, and the maintenance of deleterious alleles in Microbotryum violaceum (=Ustilago violacea).

The mating system of Microbotryum violaceum was investigated in populations that are polymorphic for mating-type bias, where individuals produce viable haploids of only one of the two required mating types. The cause of mating-type bias was identified as deleterious recessive alleles linked to mating type. Maintenance of the deleterious alleles was promoted by early conjugation among products of single meioses, such that the duration of the free-living haploid stage is minimized. This development was also observed in nonbiased isolates. As a consequence, the mating system tends toward mating within the tetrad, which might be expected to reduce heterozygosity. However, complete centromere linkage of mating type ensures conjugation between first division meiotic products, such that mating in M. violaceum is analogous to forms of meiotic parthenogenesis with first division restitution (i.e. automixis with central fusion). This fungus was used to test the prediction that this mating system would maintain heterozygosity in regions of the genome linked to centromeres. Therefore, populations were screened for additional heterozygous lethal recessive alleles linked to centromeres, and several examples were found. Furthermore, the occurrence of intratetrad mating in M. violaceum provides an explanation for low variation among individuals within populations, inconsistent estimates of outcrossing rates, low levels of mating between tetrads of one diploid individual, and high frequencies of haplo-lethal alleles in natural populations.

Alleles↗

Outcrossing and paternity analysis of Pinus densiflora (Japanese red pine) by microsatellite polymorphism.

This study employed microsatellite loci to analyse outcrossing rate and pollen dispersal in Japanese red pine (Pinus densiflora) in an isolated stand. The average offspring outcrossing rate for 29 cones was 0.955. Significant differences in outcrossing rates between offspring groups on individual branches that extended in different directions at different heights were not detected. Male parents of 874 offspring collected from the maternal tree were assessed by exclusion using polymorphisms at three microsatellite loci. Paternity analysis indicated that at least 31% of the offspring were fertilized by pollen from trees outside the stand. The average distance of pollen migration within the study stand was 68 m, with a maximum value of 325 m. There was excess mating with nearby P. densiflora trees, of which only a few were predominant pollen donors. In addition, a weakly directional bias in P. densiflora pollination was also detected in the study stand, suggesting that female strobili on a branch of the maternal tree were more easily fertilized by pollen from trees in that direction.

Alleles↗

Intraindividual and intraspecies variability of ITS1 sequences in the ancient asexual Darwinula stevensoni (Crustacea: Ostracoda).

The lack of variability in ITS regions within individuals and within species has been explained as the result of concerted evolution. In fact, many examples of intraindividual variation in the ITS regions have been reported. Here we report evidence of within-individual variation of the ITS1 region in the obligate parthenogenetic species Darwinula stevensoni. We analysed 46 clones obtained from 12 individuals of D. stevensoni, from three Italian sites and one site in Luxembourg. Seven nucleotides out of 366 were variable. Most variability (80%) was found among clones within individuals, and the remainder of the variability was observed among individuals. No difference was found among populations or between habitats. The low intraspecific variability and the observation of recombinant molecules are evaluated in light of the relevant literature. The high percentage of variation within individuals and the occurrence of recombination without meiosis are discussed by considering the ancient asexual "status" of the species.

Analysis of Variance↗

Social structure of the mound-building mouse Mus spicilegus revealed by genetic analysis with microsatellites.

The Mound-building mouse Mus spicilegus possesses a unique behaviour amongst mice. It constructs large earthen mounds and associated nesting chambers which serve to store food for immature individuals during the winter nesting period. We have used genetic analysis of four autosomal and four X-linked microsatellite loci to determine relationships between individuals inhabiting 40 mounds in Bulgaria. We show that, in almost all cases, individuals in a mound are the product of multiple parentage. We estimate the minimum number of males and female parents contributing offspring to each mound and demonstrate that at least two male and two female parents contribute offspring to a minimum of seven mounds. Analyses of relatedness coefficients and allele sharing values demonstrate that parents of different sibships within mounds are more related than if they had been chosen at random from the population and suggest that it is the female parents that contribute this excess relatedness. These results suggest that the mechanism by which individuals congregate to build mounds is kin-based and that the evolution of mound building and communal nesting in M. spicilegus is due in part to kin selection. This study represents a novel approach to the study of mammalian behavioural ecology. We have used a genetic dataset to construct an outline of social structure in the absence of behavioural data. These inferences can now be used to direct further work on this species.

Alleles↗

Microsatellites reveal clonal structure of populations of the thelytokous ant platythyrea punctata (F. Smith) (Hymenoptera; formicidae)

Parthenogenesis is often thought to constitute an evolutionary dead end as compared with sexual reproduction because genetic recombination is limited or nonexistent in parthenogenetic populations. Yet there are many species to demonstrate that parthenogenesis can initially be extremely successful under certain environmental conditions. In this study we used microsatellite markers to investigate the genetic structure of four natural populations of the neotropical thelytokous parthenogenetic ant Platythyrea punctata. Ten dinucleotide microsatellites were isolated from a partial genomic library of P. punctata. Five of these were found to be polymorphic. In a subsequent analysis of 314 workers taken from 51 colonies, we detected low intraspecific levels of variation at all loci, expressed both in the number of alleles detected and heterozygosities observed. Surprisingly, we found almost no differentiation within populations. Populations rather had a clonal structure, with all individuals from all colonies usually sharing the same genotype. Only in one colony from Puerto Rico did some workers have an additional genotype. This low level of genotypic diversity probably reflects the predominance of thelytoky in P. punctata, together with genetic bottlenecks and founder effects. Cross-species amplification of all 10 loci in 29 ant species comprising four different subfamilies yielded positive amplification products in only a limited number of species.

Journal Article↗

Multilocus sequence typing suggests the chytrid pathogen of amphibians is a recently emerged clone.

Chytridiomycosis is a recently identified fungal disease associated with global population declines of frogs. Although the fungus, Batrachochytrium dendrobatidis, is considered an emerging pathogen, little is known about its population genetics, including the origin of the current epidemic and how this relates to the dispersal ability of the fungus. In this study, we use multilocus sequence typing to examine genetic diversity and relationships among 35 fungal strains from North America, Africa and Australia. Only five variable nucleotide positions were detected among 10 loci (5918 bp). This low level of genetic variation is consistent with the description of B. dendrobatidis as a recently emerged disease agent. Fixed (i.e. 100%) or nearly fixed frequencies of heterozygous genotypes at two loci suggested that B. dendrobatidis is diploid and primarily reproduces clonally. In contrast to the lack of nucleotide polymorphism, electrophoretic karyotyping of multiple strains demonstrated a number of chromosome length polymorphisms.

Amphibians↗

Sexual stages in trypanosomes and implications.

The basis for concluding that a system of genetic exchange exists in African trypanosomes of the Trypanosoma brucei group is outlined by the accumulated data from variation in natural parasite populations and from laboratory crossing experiments. The results indicate that these parasites can complete their life cycle either with or without genetic exchange. This sexual event involves meiosis and syngamy, but the order of these processes and the sexual stages involved are not yet known.

Animals↗

Female investment is retarded pending reception of allosperm in a hermaphroditic colonial invertebrate.

Young colonies of the bryozoan Celleporella hyalina are capable of acquiring water-borne allosperm and of using it to fertilize ova for a period of 3-6 weeks after reaching female sexual maturity. In these simultaneous hermaphrodites, early allocation to female modules, but not male, is greatly enhanced by the acquisition of allosperm. The degree of enhancement is inversely proportional to coancestry of the recipient and donor colonies, thus promoting outcrossing. This apparently novel mechanism of adjusting operational sex ratio depends on the uptake and storage of sperm by nonreproductive (somatic) modules and subsequent translocation to females.

Animals↗

Structure and organization of transposable mating type cassettes in Saccharomyces yeasts.

Cell type in Saccharomyces yeasts is regulated by two transposable blocks of DNA, the a and alpha cassettes. There are three loci where either cassette can exist. At the HML and HMR loci the cassettes are not expressed. The cassette at the MAT locus is expressed and controls the cell type. Changes of cell type involve transposition-substitution of cassettes from HML or HMR into MAT. We recently reported the molecular cloning of the alpha cassette at the HML locus, HML alpha, and showed that it contained sequences homologous to HMR and MAT. Using HML alpha as a hybridization probe, we have isolated HMLa, HMR alpha, HMRa, MAT alpha, and MATa. Heteroduplex analysis and restriction endonuclease mapping studies indicate that the a and alpha cassettes differ by a substitution corresponding to about 750 base pairs in alpha and about 600 base pairs in a. The HML, HMR, and MAT loci have regions of homology flanking the position of the a versus alpha substitution. We have used specific chromosome rearrangements fusing MAT and HML and MAT with HMR to orient the cloned sequences on the genetic map and have found that all three genes have the same left-to-right polarity on the chromosome.

Alleles↗

Cytotype polymorphism of the P-M system in two wild populations of Drosophila melanogaster.

In Drosophila melanogaster the interactions of the P-M system generate germ-line aberrations (e.g., sterility, mutations) found in certain interstrain hybrids. Two wild populations, from France and Tunisia, were examined in order to determine the distribution of the chromosomal P factor and the extrachromosomal cytotypes. No P factors active for potential GD sterility were found in these populations. But search for the M cytotype, which causes susceptibility to the P factors, and for the P cytotype, which causes resistance to the P factors, showed that both populations are polymorphic for the cytotypes. Such a polymorphism seems to be stable in the wild, at least over a 2-year period. Mutator activity (measured by generated mutations at the sn and ras loci) was found to be present. Some possible interactions between cytotype polymorphism, mutator activity, and the structure of Drosophila populations are discussed.

Animals↗

Regulation of yeast mating-type interconversion: feedback control of HO gene expression by the mating-type locus.

The ultimate product of yeast mating-type interconversion is a stable a/alpha diploid cell. A haploid cell carrying the HO gene gives rise to a diploid cell in a two-step process: first, the cell switches mating type as a result of genetic rearrangement (cassette substitution) catalyzed by HO; then, cells of opposite type mate to form a/alpha diploids. Mating-type interconversion does not occur in a/alpha diploids despite the presence of the HO gene. We have identified a plasmid carrying the HO gene by screening a yeast clone bank (constructed in vector YEp13) for plasmids that allow mating-type switching by ho cells. The yeast segment responsible for mating-type interconversion integrates by homology at the ho locus, thus confirming that it carries HO. Using the HO gene as a probe, we find that strains with an active mating-type interconversion system produce HO RNA, whereas a/alpha HO/HO cells do not and that this inhibition requires products of both the MATa1 and MATa2 genes. Thus, mating-type interconversion does not occur in a/alpha HO/HO cells because the HO gene product is not synthesized. These results demonstrate the following: (i) The mating-type locus, proposed on genetic grounds to be a regulatory locus, controls expression of an unlinked gene (HO) at the level of RNA production. (ii) The HO gene is under negative feedback control: its expression is inhibited after successful completion of diploidization (formation of a/alpha diploids).

DNA, Fungal↗

Ancient and recent patterns of geographic speciation in the oyster mushroom Pleurotus revealed by phylogenetic analysis of ribosomal DNA sequences.

Evidence from molecular systematic studies suggests that many mushroom species may be quite ancient. Gene phylogenies were developed to examine the relationship between reproductive isolation, genetic divergence, and biogeography in oyster mushrooms (Pleurotus). Sequence data were obtained for two regions of DNA from populations belonging to eight intersterility groups (biological species). Phylogenetic analysis of sequences from the 5' portion of the nuclear encoded large subunit rDNA demonstrates an ancient origin for four intersterility groups of broad geographic distribution (world-wide), with a more recent radiation of several intersterility groups that are restricted to the Northern Hemisphere. An expanded analysis using sequence data from the more variable rDNA internal transcribed spacer region also reveals a phylogenetically based pattern of genetic divergence associated with allopatric speciation among populations from different continents in the Northern Hemisphere. The ability of rDNA sequences to resolve phylogenetic relationships among geographically isolated populations within intersterility groups illustrates the importance of biogeography for understanding speciation in Pleurotus. Patterns of geographic distribution among intersterility groups suggest that several species lineages evolved quite early, with recently evolved groups restricted to the Northern Hemisphere and older lineages occurring throughout the world. Based on phylogenetic evidence, analysis of historical biogeography using area cladograms shows that multiple dispersal and vicariance events are responsible for patterns of speciation observed.

Base Sequence↗