THE GENETIC BASIS OF SEXUAL ISOLATION BETWEEN DROSOPHILA MELANOGASTER AND D. SIMULANS

The genetic analysis of sexual isolation between the closely-related species Drosophila melanogaster and Drosophila simulans involved two experiments with no-choice tests. The efficiency of sexual isolation was measured by the frequency of courtship initiation and interspecific mating. We first surveyed the variation in sexual isolation between D. melanogaster strains and D. simulans strains of different geographic origin. Then, to investigate variation in sexual isolation within strains, we made F₁ diallel sets of reciprocal crosses within strains of D. melanogaster and D. simulans. The F₁ diallel progeny of one sex were paired with the opposite sex of the other species. The first experiment showed significant differences in the frequency of interspecific mating between geographic strains. There were more matings between D. simulans females and D. melanogaster males than between D. melanogaster females and D. simulans males. The second experiment uncovered that the male genotypes in the D. melanogaster diallel significantly differed in interspecific mating frequency, but not in courtship initiation frequency. The female genotypes in the D. simulans diallel were not significantly different in courtship initiation and interspecific mating frequency. Genetic analysis reveals that in D. melanogaster males sexual isolation was not affected by either maternal cytoplasmic effects, sex-linked effects, or epistatic interaction. The main genetic components were directional dominance and overdominance. The F₁ males achieved more matings with D. simulans females than the inbred males. The genetic architecture of sexual isolation in D. melanogaster males argues for a history of weak or no selection for lower interspecific mating propensity. The behavioral causes of variation in sexual isolation between the two species are discussed.     

Ecology meets molecular genetics in Arabidopsis

 Arabidopsis thaliana has been used as a model plant for molecular genetics studies. Recently, Arabidopsis has been utilized in an increasing number of ecological and evolutionary studies. These studies are beginning to explain three “black boxes” of ecology: the nature of the mutations responsible for phenotypic variation, female–male interactions in the pistil, and the genetic basis of speciation. Among the advances are the inclusive fitness of haploid gametophytes, the testing of the arms-race model, morphological diversity, and reproductive isolation. Important methods include quantitative trait locus (QTL) mapping and molecular population genetics. Most significantly, natural variations of various aspects are now available via the taxonomic revision of its closely related species, and by the worldwide collection of accessions. Some conspicuous works using maize, Drosophila, and other species will be also discussed. Received: June 1, 2002 / Accepted: October 15, 2002 Acknowledgments I would like to express my deepest gratitude to Prof. K. Okada for supporting this research. I thank the organizers of the symposium for fruitful discussion, T. Yahara and M. Kanaoka for critical reading, T. Araki and T. Kenta for valuable discussions, and members of the Okada laboratory for technical assistance and discussion. The work was supported by JSPS Research Fellowships for Young Scientists.     

Sexual Isolation Between Drosophila Melanogaster, D. Simulans and D. Mauritiana: Sex and Species Specific Discrimination

The sexual isolation among the related species Drosophila melanogaster, D. simulans and D. mauritiana is asymmetrical. While D. mauritiana males mate well with both D. melanogaster and D. simulans females, females of D. mauritiana discriminate strongly against males of these two species. Similarly, D. simulans males mate with D. melanogaster females but the reciprocal cross is difficult. Interspecific crosses between several populations of the three species were performed to determine if (i) males and females of the same species share a common sexual isolation genetic system, and (ii) males (or females) use the same genetic system to discriminate against females (or males) of the other two species. Results indicate that although differences in male and female isolation depend on the populations tested, the isolation behaviour between a pair of species is highly correlated despite the variations. However, the rank order of the isolation level along the populations was not correlated in both sexes, which suggests that different genes act in male and female sexual isolation. Neither for males nor for females, the isolation behaviour of one species was paralleled in the other two species, which indicates that the genetic systems involved in this trait are species-pair specific. The implications of these results are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Ecological diversity and speciation in land snails of the genus Mandarina from the Bonin Islands

 Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an example of habitat and character divergence among closely related species. The molecular phylogenies of Mandarina show that a divergence of arboreal, semiarboreal, and ground-dwelling species has occurred repeatedly in different times, areas, and lineages. Ecological diversification is suggested to be important for the coexistence of Mandarina species based on the facts that sympatric species are typically highly differentiated ecologically and morphologically, and that species occupying similar habitats do not coexist. The ecological diversification of Mandarina has occurred without much genetic divergence compared with that of its mainland relatives. This suggests that morphological and ecological diversifications are accelerated in depauperate environments where there are fewer competitors and predators. Although the details of the reproductive isolation mechanisms are not understood and further examination is needed, the rapid evolution of prezygotic isolation is the main cause of speciation in Mandarina. In particular, ecological diversification may be an effective barrier to gene exchange between two species. Because of incomplete postmating isolation and the lower genetic divergence among species of Mandarina, breakdowns of reproductive isolation have frequently occurred as a result of habitat change. It is important to estimate the effect of hybridization on species diversification in future studies. Received: February 7, 2002 / Accepted: October 22, 2002 Acknowledgments I express my sincere thanks to A. Davison, B.C. Clarke, A. Guiller, D. Thomaz, K. Tomiyama, I. Hayami, and K. Tanabe for helpful advice and assistance. This study was supported by grants from the Japan Society for the Promotion of Science and the Nippon Life Insurance Foundation.     

Pleosporales in Japan (1): the genus <Emphasis Type="Italic">Lophiostoma</Emphasis>

 Seven species of the genus Lophiostoma were the subject of this study. Among these, Lophiostoma mucosum is described and illustrated as a new species. All other species, L. macrostomum, L. semiliberum, L. arundinis, L. caulium, L. caudatum, and L. winteri, are reported for the first time in Japan. A key to the species of Lophiostoma in Japan is given. Received: August 5, 2002 / Accepted: November 28, 2002 Acknowledgments We are grateful to Dr. Hideki Takahashi (curator of SAPA) for the loan of fungal material. Correspondence to:Y. Harada     

Fine-scale genetic analysis of species-specific female preference in Drosophila simulans

Behavioural differences are thought to be the first components to contribute to species isolation, yet the precise genetic basis of behavioural isolation remains poorly understood. Here, we used a combination of behaviour assays and genetic mapping to provide the first refined map locating candidate genes for interspecific female preference isolating Drosophila simulans from D. melanogaster. First, we tested whether two genes identified as affecting D. melanogaster female intraspecific mate choice also affect interspecific mate choice; neither of these genes was found to contribute to species‐specific female preference. Next, we used deficiency mapping to locate genes on the right arm of the third chromosome for species‐specific female preference and identified five small significant regions that contain candidate genes contributing to behavioural isolation. All five regions were located in areas that would have low interspecific recombination, which mirrors the results of other behavioural isolation studies that used quantitative trait locus (QTL) mapping, but without the potential concern of bias towards regions of low recombination that QTL mapping may have. As this model system may be refined to the individual gene level using the same methodology, this initial map we provide may potentially serve as a ready template for the identification and characterization of the first behavioural isolation genes.     

High divergence of reproductive tract proteins and their association with postzygotic reproductive isolation in Drosophila melanogaster and Drosophila virilis group species

The possible association between gonadal protein divergence and postzygotic reproductive isolation was investigated among species of the Drosophila melanogaster and D. virilis groups. Protein divergence was scored by high-resolution two-dimensional electrophoresis (2DE). Close to 500 protein spots from gonadal tissues (testis and ovary) and nongonadal tissues (malpighian tubules and brain) were analyzed and protein divergence was calculated based on presence vs absence. Both testis and ovary proteins showed higher divergence than nongonadal proteins, and also a highly significant positive correlation with postzygotic reproductive isolation but a weaker correlation with prezygotic reproductive isolation. Particularly, a positive and significant correlation was found between proteins expressed in the testis and postzygotic reproductive isolation among closely related species such as the within-phylad species in the D. virilis group. The high levels of male-reproductive-tract protein divergence between species might be associated with F₁ hybrid male sterility among closely related species. If so, a lower level of ovary protein divergence should be expected on the basis that F₁ female hybrids are fully fertile. However, this is not necessarily true if relatively few genes are responsible for the reproductive isolation observed between closely related species, as recent studies seem to suggest. We suggest that the faster rate of evolution of gonadal proteins in comparison to nongonadal proteins and the association of that rate with postzygotic reproductive isolation may be the result of episodic and/or sexual selection on male and female molecular traits. Correspondence to: A. Civetta     

Genetics of a difference in pigmentation between Drosophila yakuba and Drosophila santomea

Abstract.— Drosophila yakuba is a species widespread in Africa, whereas D. santomea, its newly discovered sister species, is endemic to the volcanic island of São Tomé in the Gulf of Guinea. Drosophila santomea probably formed after colonization of the island by its common ancestor with D. yakuba. The two species differ strikingly in pigmentation: D. santomea, unlike the other eight species in the D. melanogaster subgroup, almost completely lacks dark abdominal pigmentation. D. yakuba shows the sexually dimorphic pigmentation typical of the group: both sexes have melanic patterns on the abdomen, but males are much darker than females. A genetic analysis of this species difference using morphological markers shows that the X chromosome accounts for nearly 90% of the species difference in the area of abdomen that is pigmented and that at least three genes (one on each major chromosome) are involved in each sex. The order of chromosome effects on pigmentation area are the same in males and females, suggesting that loss of pigmentation in D. santomea may have involved the same genes in both sexes. Further genetic analysis of the interspecific difference between males in pigmentation area and intensity using molecular markers shows that at least five genes are responsible, with no single locus having an overwhelming effect on the trait. The species difference is thus oligogenic or polygenic. Different chromosomal regions from each of the two species influenced pigmentation in the same direction, suggesting that the species difference (at least in males) is due to natural or sexual selection and not genetic drift. Measurements of sexual isolation between the species in both light and dark conditions show no difference, suggesting that the pigmentation difference is not an important cue for interspecific mate discrimination. Using DNA sequence differences in nine noncoding regions, we estimate that D. santomea and D. yakuba diverged about 400,000 years ago, a time similar to the divergences between two other well‐studied pair of species in the subgroup, both of which also involved island colonization.     

The origins of postmating reproductive isolation: testing hypotheses in the grasshopper <Emphasis Type="Italic">Chorthippus parallelus</Emphasis>

 Although there are several well-established hypotheses for the origins of postmating isolation during allopatric divergence, there have been very few attempts to determine their relative importance in nature. We have developed an approach based on knowledge of the differing evolutionary histories of populations within species that allows systematic comparison of the predictions of these hypotheses. In previous work, we have applied this methodology to mating signal variation and premating reproductive isolation between populations of the meadow grasshopper Chorthippus parallelus. Here we review the principles behind our approach and report a study measuring postmating isolation in the same set of populations. The populations have known and differing evolutionary histories and relationships resulting from the colonization of northern Europe following the last glaciation. We use a maximum-likelihood analysis to compare the observed pattern of postmating isolation with the predictions of the hypotheses that isolation primarily evolves either as a result of gradual accumulation of mutations in allopatry, or through processes associated with colonization, such as founder events. We also quantify the extent to which degree of postmating isolation can be predicted by genetic distance. Our results suggest that although there is only a weak correlation between genetic distance and postmating isolation, long periods of allopatry do lead to postmating isolation. In contrast to the pattern of premating isolation described in our previous study, colonization does not seem to be associated with increased postmating isolation. Received: January 24, 2002 / Accepted: July 26, 2002 Acknowledgments Numerous people helped with collecting and rearing grasshoppers. We are grateful to NERC for funding. Correspondence to:R.K. Butlin     

Pleosporales in Japan (2): the genus <Emphasis Type="Italic">Lophiotrema</Emphasis>

 Five species of the genus Lophiotrema are reported in this article. Of these, Lophiotrema vitigenum is described and illustrated as a new species. Three species, L. nucula, L. neohysterioides, and L. vagabundum, are newly added to the Japanease mycoflora. Lophiosphaera orientalis, Nodulosphaeria araucariae, and N. rosae, which had been recorded in Japan, are treated as synonyms of Lophiotrema fuckelii. A key to the species of Lophiotrema in Japan is given. Received: October 9, 2002 / Accepted: January 20, 2003 Acknowledgments We are grateful to Drs. Shuhei Tanaka (curator of YAM) and Ken Katumoto for the loan of fungal material. Correspondence to:K. Tanaka     

Identification of One Intron Loss and Phylogenetic Evolution of Dfak Gene in the Drosophila melanogaster Species Group

Intron loss and its evolutionary significance have been noted in Drosophila. The current study provides another example of intron loss within a single-copy Dfak gene in Drosophila. By using polymerase chain reaction (PCR), we amplified about 1.3 kb fragment spanning intron 5–10, located in the position of Tyr kinase (TyK) domain of Dfak gene from Drosophila melanogaster species group, and observed size difference among the amplified DNA fragments from different species. Further sequencing analysis revealed that D. melanogaster and D. simulans deleted an about 60 bp of DNA fragment relative to other 7 Drosophila species, such as D. elegans, D. ficusphila, D. biarmipes, D. takahashii, D. jambulina, D. prostipennis and D. pseudoobscura, and the deleted fragment located precisely in the position of one intron. The data suggested that intron loss might have occurred in the Dfak gene evolutionary process of D. melanogaster and D. simulans of Drosophila melanogaster species group. In addition, the constructed phylogenetic tree based on the Dfak TyK domains clearly revealed the evolutionary relationships between subgroups of Drosophila melanogaster species group, and the intron loss identified from D. melanogaster and D. simulans provides a unique diagnostic tool for taxonomic classification of the melanogaster subgroup from other group of genus Drosophila.     

Fungi on submerged wood in the Koito River,Japan

 Woody substrates were collected from the Koito River in Japan, and the biodiversity of fungi on these substrates was investigated. Twenty-eight species were identified, comprising 12 ascomycete and 16 anamorphic taxa. The common fungi included Chaetosphaeria sp., Ophioceras commune, Pseudohalonectria lignicola, and Savoryella lignicola. The occurrence of fungi on submerged wood is discussed, and three interesting taxa – Pseudohalonectria lignicola, Bactrodesmium arnaudii, and B. pallidum – are described and illustrated. Received: August 19, 2002 / Accepted: November 25, 2002 Acknowledgments We are grateful to the financial support of the Special Coordination Fund for Promoting Science and Technology from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Profs. K. Nishimura and M. Miyaji are thanked for scientific invitation and assistance in Chiba, Japan. Dr. Norio Hayashi from the Natural History Museum and Institute, Chiba, is thanked for providing physicochemical data for the Koito River. Y.M. Leung and M.H. Ng are thanked for technical assistance. Correspondence to:C.K.M. Tsui     

The magnitude of behavioral isolation is affected by characteristics of the mating community

Gene exchange between species occurs in areas of secondary contact, where two species have the opportunity to hybridize. If heterospecific males are more common than conspecific males, females will experience more encounters with males of other species. These encounters might increase the likelihood of heterospecific matings, and lead to the production of hybrid progeny. I studied the mating behavior of two pairs of sibling species endemic to Africa: Drosophila yakuba/Drosophila santomea and Drosophila simulans/Drosophila sechellia. Drosophila yakuba and D. simulans are cosmopolitan species widely distributed in the African continent, while D. santomea and D. sechellia are island endemics. These pairs of species hybridize in nature and have the potential to exchange genes in natural conditions. I used these two pairs of Drosophila species, and constructed mating communities of different size and different heterospecific:conspecific composition. I found that both the total number of potential mates and the relative frequency of conspecific versus heterospecific males affect female mating decisions in the cosmopolitan species but not in the island endemics. These results suggest that the population characteristics, in which mating occurs, may affect the magnitude of premating isolation. Community composition might thus facilitate, or impair, gene flow between species.     

Three new species of Annulatascus (Ascomycetes) from Hong Kong freshwater habitats

 Annulatascus joannae, A. lactus, and A. tropicalis are described and illustrated from decaying woody substrata in freshwater habitats in Hong Kong. Annulatascus joannae is distinguished by ellipsoidal and thick-walled ascospores whereas A. lacteus has milky ascomata and A. tropicalis has relatively large, fusiform, 1–3-septate ascospores. Annulatascus biatriisporus is reported as a new record in Hong Kong. A key to and a synoptic table of Annulatascus species are provided. Received: December 14, 2001 / Accepted: July 5, 2002 Acknowledgments C.K.M. Tsui and V.M. Ranghoo are grateful to The University of Hong Kong for the award of postgraduate studentships. A.Y.P. Lee, M.H. Ng, and M.Y. Chan are thanked for photographic and technical assistance. Correspondence to:C.K.M. Tsui     

Chlamydospore formation of <Emphasis Type="Italic">Entoloma clypeatum</Emphasis> f. <Emphasis Type="Italic">hybridum </Emphasis>on mycorrhizas and rhizomorphs associated with <Emphasis Type="Italic">Rosa multiflora</Emphasis>

 Chlamydospores of Entoloma clypeatum f. hybridum were described on the mycorrhizas and rhizomorphs associated with Rosa multiflora. Their developmental pattern seems to be the Nyctalis type. This is the first report on chlamydospore formation on the mycorrhizae in entolomatoid fungi. Received: January 17, 2002 / Accepted: November 5, 2002 Acknowledgments K.H. is grateful to Emeritus Professor N. Sagara in Kyoto University, in whose laboratory part of this study was undertaken. Thanks are due to Mr. D. Sakuma for allowing the specimens to be kept in Osaka Museum of Natural History. Correspondence to:H. Kobayashi     

Hybrid lethal systems in theDrosophila melanogaster species complex

Lethal phases of the hybrids betweenDrosophila melanogaster and its sibling species,D. simulans are classified into three types: (1) embryonic lethality in hybrids carryingD. simulans cytoplasm andD. melanogaster X chromosome, (2) larval lethality in hybrids not carryingD. simulans X, and (3) temperature-sensitive pupal lethality in hybrids carryingD. simulans X. The same lethal phases are also observed when either of the two other sibling species,D. mauritiana orD. sechellia, is employed for hybridization withD. melanogaster. Here, we describe genetic analyses of each hybrid lethality, and demonstrate that these three types of lethality are independent phenomena. We then propose two models to interpret the mechanisms of each hybrid lethality. The first model is a modification of the conventional X/autosome imbalance hypothesis assuming a lethal gene and a suppressor gene are involved in the larval lethality, while the second model is for embryonic lethality assuming an interaction between a maternal-effect lethal gene and a suppressor gene.