The genetic isolation between some populations of Proasellus coxalis

We tested the reroductive compatibility existing between four geographically different populations of Proaselis coxalis (Crustacea Isopoda) collected in the river Sarno (near Naples) and on three Sicilian sites: San Domenico; Villa Grazia di Carini and Enna. By means of laboratory hybridization experiments (“no-choice” method) we investigated and characterized the reproductive isolation established between these populations under allopatric conditions. The populations San Domenico and Enna are reproductively isolated from Villa Grazia di Carini and Sarno, so that they may be considered as belonging to different species. The mechanism of reproductive isolation are of postzygotic nature only: embronic death, hbrid inviability and sterility of the surviving hybrids. These results agree with tie theory of aioatric speciation, according to which in the first phase of speciation only post-zygotic mechanisms occur as an accidental product of genetic divergence.     

Tests of reproductive isolation among species in the <Emphasis Type="Italic">Fundulus notatus</Emphasis> (Cyprinodontiformes: Fundulidae) species complex

We assessed prezygotic (probability of spawning) and postzygotic (hatching success) reproductive isolation among the three ecologically and morphologically similar species in the Fundulus notatus species complex. We employed a multi-generation breeding experiment to test the hypotheses that karyotypic differences, body size differences, or geographic isolation among populations will increase pre or postzygotic reproductive barriers. Overall, prezygotic barriers were strong and postzygotic barriers weak in crosses of non-hybrid heterospecifics (F1 hybrid crosses) while prezygotic barriers were weaker and postzygotic barriers stronger in crosses involving hybrid individuals (F2 hybrid crosses and backcrosses). Prezygotic barriers among the two smaller species (Fundulus notatus and F. euryzonus) broke down rapidly; first generation hybrids spawned (F2 hybrid crosses and backcrosses) as frequently as parental forms in intraspecific crosses. There was no increase in postzygotic barriers among species with cytogenetic differences. There were increased prezygotic, but not postzygotic, barriers among geographically isolated populations of one species. While pure males and females were just as likely to spawn with hybrids, some types of hybrid females suffered from increased sterility, but not inviability, over hybrid males. Female sterility was only seen in hybrids with a Fundulus euryzonus parent, while other female hybrids produced viable eggs.     

Analysis of Early Embryogenesis in Rainbow and Banded Darters (Percidae: Etheostoma) Reveals Asymmetric Postmating Barrier

This study examines early embryogenesis in two species of darters, Etheostoma caeruleum and E. zonale (Teleostei: Percidae), and their hybrids. Results document the course of ontogeny from fertilization until the onset of pigmented eyes. Comparing fertilization and developmental success of conspecific versus heterospecific crosses revealed an asymmetric postmating reproductive barrier. E. caeruleum eggs treated with E. zonale sperm exhibited fertilization and developmental success similar to that of conspecific crosses. In contrast, E. zonale eggs treated with E. caeruleum sperm exhibited reduced fertilization relative to conspecific crosses and abnormal development. Development in this latter cross was compromised at all stages, but appeared to be concentrated around epiboly, or cell migration. As epiboly represents the stage of ontogeny when zygotic genes of both species jointly contribute to embryogenesis, results provide insight into the genetic mechanisms underlying postmating barriers in Etheostoma. Finally, the observed asymmetry in fertilization success is consistent with predictions based on the behavioral ecology of these species. Etheostoma zonale spawn in the open water column, whereas E.␣caeruleum bury their eggs under the substrate. The observed fertilization barrier may have therefore resulted from selection favoring increased fertilization specificity in E. zonale.     

石荠苎属的遗传分化与种间关系分析

杂交实验结果表明,石荠苎属(唇形科)中国产的7个种,即小花荠苎Mosla cavaleriei、石香薷M．chinensis、小鱼仙草M．dianthera、杭州石荠苎M．hangchouensis、疏花荠苎M．pauciflora、石荠苎M．scabra和苏州石荠苎M．soochouensis,彼此之间生殖隔离。这7个种中,疏花荠苎是四倍体,由杂交伴随多倍化产生。根据21个代表居群在15个酶系统的28个位点的等位酶资料分析,其余6个二倍体种组成3个亲缘关系密切的种对。杭州石荠苎和石香薷的亲缘关系最近,石香薷可能由于涉及繁育系统类型的大突变而与杭州石荠苎分化开来。小花荠苎与小鱼仙草比较接近,但它们的遗传分化很显著,除了突变之外,染色体结构也有大的变异,最明显的表现是随体的位置不同,小花荠苎的随体位于长臂上,而小鱼仙草与其它种一样,随体位于短臂上。石荠苎和苏州石荠苎在酶基因位点上的分化不大,但生境隔离、花期隔离和繁育系统的差别等能有效地保持种的独立性。不同的生境要求、不同的花期、花的形态结构的分化、繁育系统类型的不同、杂种F_l代高度不育、染色体数目和核型的差异等均为石荠苎属下种间有效的隔离机制,而且近缘种间常是多种机制共同起作用。     

作物远缘杂交育种的途径及其实质

作物远缘杂交的育种可操作性及效果多年来颇有争议,科学家对物种起源与进化的研究恰恰是指导作物远缘杂交育种的理论基础。物种形成理论研究表明生命的共同起源是远缘杂交的理论基础,生物多样性是远缘杂交的物质基础。生物种间的繁殖隔离机制是远缘杂交不亲和性障碍的根源所在,而物种形成方式又为克服远缘杂交的不亲和性提供了理论依据。其中异域性物种形成方式下的生殖隔离具有不彻底性,是克服远缘杂交受精前不亲和性的理论根据;同域性物种形成方式中多倍体化的方式对远缘杂交受精后不亲和性的克服具有较强的指导意义。本文在通过对以上方面的阐述,剖析了远缘杂交的障碍来源、克服途径及实质,为作物远缘杂交育种工作提供参考。     

The role of host-specificity in the reproductive isolation of Epichloë endophytes revealed by reciprocal infections

Speciation in sexually reproducing organisms hinges on reproductive barriers that reduce gene flow between species or preclude the formation of hybrids. Here, we studied potential reproductive barriers in four members of the Epichloë typhina (Ascomycota, Clavicipitaceae) complex, i.e. Epichloë typhina infecting Dactylis glomerata, E. typhina subsp. clarkii infecting Holcus lanatus, E. typhina subsp. poae infecting Poa nemoralis and E. typhina infecting P. trivialis. Reciprocal inoculation tests showed that these endophytes are host-specific. This suggests that reproductive isolation among Epichloë strains may be the result of specialization to one host, on which mating between different individuals occurs. Furthermore, significantly lower infection frequencies of F1 progeny from crosses between host-strains compared to parental strains and within host-strain progeny suggest that host-dependent effects upon hybrid fitness exist, which would conform to an extrinsic postzygotic isolation barrier. Our results may explain, why members of the E. typhina complex remain genetically differentiated in natural populations.     

Speciation in the large-spored Metschnikowia clade and establishment of a new species, Metschnikowia borealis comb. nov

The reproductive boundaries among species in the large-spored Metschnikowia clade were studied by prototrophic recombinant selection, electrophoretic karyotyping, mitochondrial DNA restriction analysis, and DNA sequence analysis. Inviable ascospores arose from crosses between the two varieties of Metschnikowia continentalis, indicating that they should be recognized as separate species. Prototrophic recombinants were recovered from crosses between auxotrophic mutants of Metschnikowia borealis, M. continentalis, Metschnikowia lochheadii, Metschnikowia sp. UWO(PS)00-154.1, and Candida ipomoeae, showing that some genetic exchange is possible in spite of the sterility of the asci formed in interspecific crosses. Metschnikowia hawaiiensis, although capable of ascus formation when its h(-) mating type is crossed with the h(+) mating type of the other species, did not give rise to recombinants. In the other species, some recombinants acquired the ability to form asci directly from single cells. These often contained the chromosomes of both parents, suggesting formation of allodiploid hybrids. Other recombinants behaved as haploids and were similar to one parent except for having inherited the selectable wild-type allele from the other parent. In most, but not all cases, inheritance of the mitochondrial genome was uniparental and correlated with the inheritance of the nuclear chromosome complement. In some cases, what appeared to be a recombinant mitochondrial genome was observed. Phylogenies derived from the sequences of various DNA regions were not congruent, indicating that hybridization may have taken place in nature as the large-spored species diverged from their common ancestor. Further evidence that C. ipomoeae arose from a natural recombination event was obtained, but a pair of Metschnikowia species that might represent derived forms of the parents could not be identified conclusively. C. ipomoeae and most of its closely related Metschnikowia species contained a group-II intron in the mitochondrial small-subunit ribosomal gene. The intron was absent in M. borealis, M. hawaiiensis, and other species in the genus Metschnikowia.     

Natural selection and divergence in mate preference during speciation

Sexual isolation can evolve due to natural selection against hybrids (reinforcement). However, many different forms of hybrid dysfunction, and selective processes that do not involve hybrids, can contribute to the evolution of sexual isolation. Here we review how different selective processes affect the evolution of sexual isolation, describe approaches for distinguishing among them, and assess how they contribute to variation in sexual isolation among populations of Timema cristinae stick-insects. Pairs of allopatric populations of T. cristinae living on different host-plant species exhibit greater sexual isolation than those on the same host, indicating that some sexual isolation has evolved due to host adaptation. Sexual isolation is strongest in regions where populations on different hosts are in geographic contact, a pattern of reproductive character displacement that is indicative of reinforcement. Ecological costs to hybridization do occur but traits under ecological selection (predation) do not co-vary strongly with the probability of between-population mating such that selection on ecological traits is not predicted to produce a strong correlated evolutionary response in mate preference. Moreover, F1 hybrid egg inviability is lacking and the factors contributing to reproductive character displacement require further study. Finally, we show that sexual isolation involves, at least in part, olfactory communication. Our results illustrate how understanding of the evolution of sexual isolation can be enhanced by isolating the roles of diverse ecological and evolutionary processes.     

Female and male visually based mate preferences are consistent with reproductive isolation between populations of the Lake Malawi endemic Labeotropheus fuelleborni

Sexual selection via female mate choice is thought to have played a key role in the speciation of haplochromine cichlids, but a dominant role for visual signals in such processes has lately been called into question. In addition, the possible role of male mating preferences in haplochromine speciation has been little studied. We studied patterns of both female and male mate choice, based exclusively on visual signals, in order to evaluate potential reproductive isolation between two populations of the Lake Malawi haplochromine Labeotropheus fuelleborni. In the first experiment, females were allowed to choose between two males, one from the same population and the other allopatric with respect to the female. Females in this experiment responded more frequently to males from their own population. Similarly, the males in these trials displayed more frequently when presented with females of their own population. In the second experiment, a female was allowed to choose between two males, either both from her own population or both allopatric. In these trials, both males and females from the Katale population interacted significantly more frequently in settings in which all three individuals were from the same population (same-population trios), and those from the Chipoka population showed a similar trend. Thus, patterns in both male and female courtship behavior suggest that visual signals contribute to at least incipient reproductive isolation between populations of L. fuelleborni [Current Zoology 56 (1): 65-72 2010].     

No evidence for faster male hybrid sterility in population crosses of an intertidal copepod (Tigriopus californicus)

Two different forces are thought to contribute to the rapid accumulation of hybrid male sterility that has been observed in many inter-specific crosses, namely the faster male and the dominance theories. For male heterogametic taxa, both faster male and dominance would work in the same direction to cause the rapid evolution of male sterility; however, for taxa lacking differentiated sex chromosomes only the faster male theory would explain the rapid evolution of male hybrid sterility. It is currently unknown what causes the faster evolution of male sterility, but increased sexual selection on males and the sensitivity of genes involved in male reproduction are two hypotheses that could explain the observation. Here, patterns of hybrid sterility in crosses of genetically divergent copepod populations are examined to test potential mechanisms of faster male evolution. The study species, Tigriopus californicus, lacks differentiated, hemizygous sex chromosomes and appears to have low levels of divergence caused by sexual selection acting upon males. Hybrid sterility does not accumulate more rapidly in males than females in these crosses suggesting that in this taxon male reproductive genes are not inherently more prone to disruption in hybrids.     

Associations between floral specialization and species diversity: cause, effect, or correlation?

It has been proposed frequently, from Darwin’s time onwards, that specialized pollination increases speciation rates and thus the diversity of plant species (i.e. clade species richness). We suggest here that the correlation between clade species richness and floral specialization is real, but that clade species richness is frequently the cause, not the result of floral specialization. We urge a broader, variance-partitioning perspective for assessing the causes of this correlation by suggesting four models of how the diversity-specialization correlation might come about: (1) floral specialization promotes initial reproductive isolation (“Initial-RI” model), (2) floral specialization promotes reinforcement of reproductive isolation upon secondary contact (“Reinforcement” model), (3) floral specialization reduces the extinction rate by promoting tighter species packing (“Extinction” model), (4) floral specialization is the result of high clade species richness, which increases the number of related species in communities, and thus selects for floral character displacement (“Character-Displacement” model). These hypotheses are evaluated by comparing the relationships between species richness, speciation mechanisms, and pollination precision, accuracy, and specialization in the broader literature and, more specifically, in four study systems: Dalechampia (Euphorbiaceae), Collinsia (Plantaginaceae), Burmeistera (Campanulaceae), and Stylidium (Stylidiaceae). These systems provide stronger support for the character-displacement hypothesis, wherein local species diversity drives the evolution of specialized pollination. Although the two reproductive-isolation hypotheses may hold for plants like orchids, with extremely precise pollination systems, the reproductive character-displacement hypothesis seems likely to be more important for plant groups with less precise pollination systems.     

Conflicting selection pressures on reproductive functions and speciation in plants

Recent developments in the field of genetic divergence and speciation focus more on diversifying processes than on geographic mode of speciation (i.e. allopatric versus sympatric). Some of these new theories concern speciation driven by conflicts between the sexes. Even though it is well known that the two reproductive functions in plants can have different selective optima, sexual selection in plants is by many assumed to be weak or non-existent. Here we outline potential sexual conflicts in plants and discuss how selection pressures generated by such conflicts may influence genetic divergence. There is opportunity for conflicting selection pressures between individuals, such as manipulative pollen traits that enhance male reproductive success at the expense of the female reproductive function. Within individual plants, fitness of the male function (pollen export) and fitness of the female function (pollen import) may be optimised by different traits, leading to conflicting selection pressures in relation to pollen transfer. This may affect selection for floral specialisation versus floral generalisation in animal-pollinated species. We believe that selection pressures generated by sexual conflict need to be appreciated in order to fully understand microevolutionary processes which may lead to genetic divergence and speciation in plants.     

Speciation and biogeography of the<Emphasis Type="Italic"> Caloglossa leprieurii</Emphasis> complex (Delesseriaceae,Rhodophyta)

Studies on the morphology, reproductive compatibility, life cycle and molecular phylogeny of the euryhaline red alga Caloglossa provide insights into the speciation events and biogeographic patterns. The C. leprieurii complex is separated into three morphotypes based on the number of cell rows at nodes and the blade width. The three morphotypes are reproductively incompatible with each other, and furthermore many mating groups are recognized within the morphotypes. Incomplete reproductive isolation is occasionally seen between geographically distant mating groups, whereas no sexual compatibility occurs between sympatrically or parapatrically distributed mating groups. In the molecular phylogenetic analyses, the C. leprieurii complex is resolved as two clusters that phenotypically correspond to the single and multiple cell row types, respectively. The strains belonging to the same mating group are closely related to each other, without exception, while the mating groups showing incomplete reproductive reactions do not always make a clade. The genetic distance is generally not correlative to the geographic distance, and this is also suggested by the morphological data and crossability. These results indicate that allopatric speciation has frequently occurred in this species complex, although there is some evidence of long-distance dispersal.M. Kamiya is the recipient of the Botanical Society Award for Young Scientist, 2002.     

Hybrid male sterility is caused by mitochondrial DNA deletion

Although it is known that the hybrid male mouse is sterile just like any other animal’s heterogametic sex, the reason why only the male germ cells are impaired has yet to be discovered. TdT-mediated dUTP nick end labeling assay using a confocal fluorescence microscope and DNA fragmentation assay of hybrid testis indicated destruction of the mitochondrial DNA (mtDNA) rather than the nuclear DNA. Previously we reported that maternal mtDNA inheritance is through selective sperm mtDNA elimination based on the sperm factor and two egg factors, and expression of these three factors was recognized in the hybrid testis. It was thereby assumed that mtDNA destruction caused by the expression of maternal mtDNA inheritance system in male germ cells is implicated in the hybrid male sterility of mice.     

Fungal species and their boundaries matter – Definitions,mechanisms and practical implications

Recent scientific and technological advances have improved and streamlined our ability to recognise and describe fungal species. Detailed comparative genomics studies have also expanded our understanding of species boundaries. Against this background, we explore the nature of fungal species and consider how this impacts our understanding of their genetics and evolution. The current body of evidence suggests that fungal species are unique evolutionary units that are separated from one another by boundaries that are “porous” under certain conditions (“semipermeable” in analogy to the differential permeability of membranes). Overall, the penetrability of these boundaries depends on the relatedness between donor and recipient species, the spatial proximity of related species to one another during their evolution, and the evolutionary potential associated with the breach of a boundary. Furthermore, the semipermeable nature of species boundaries fundamentally affects the population genetics of a species, with potentially profound effects on its overall evolution and biology. This also influences the methodologies used in taxonomy, because some species appear capable of maintaining their genetic isolation despite extensive penetrability of their boundaries. Most analytical procedures are also not able to distinguish the signals of species boundary permeability from those associated with incomplete lineage sorting or intraspecific diversity. Collectively, these issues greatly complicate how we study and name fungi. An awareness of the nature of species, their boundaries and the biological and genomic signatures of boundary breaches, will enhance our ability to identify them and, perhaps more importantly, to develop realistic strategies to manage and manipulate their growth and distribution.     

Emerging directions in the study of the ecology and evolution of plant-animal mutualistic networks: a review

The study of mutualistic plant and animal networks is an emerging field of ecological research. We reviewed progress in this field over the past 30 years. While earlier studies mostly focused on network structure, stability, and biodiversity maintenance, recent studies have investigated the conservation implications of mutualistic networks, specifically the influence of invasive species and how networks respond to habitat loss. Current research has also focused on evolutionary questions including phylogenetic signal in networks, impact of networks on the coevolution of interacting partners, and network influences on the evolution of interacting species. We outline some directions for future research, particularly the evolution of specialization in mutualistic networks, and provide concrete recommendations for environmental managers.