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PubMed2005

Reproduction in Aspergillus fumigatus: sexuality in a supposedly asexual species?

Aspergillus fumigatus has long been considered to reproduce only by asexual means. However, accumulating evidence suggest that a sexual stage for A. fumigatus may yet be identified. We describe results from published and ongoing studies involving population genetic analyses, genome analysis, studies of mating-type gene presence and distribution, expression of sex-related genes, and taxonomic work which support the assertion that A. fumigatus has the potential to reproduce by sexual means. The consequences of sexual reproduction for the population biology and disease management of the species are discussed. The possible mechanisms of evolution of asexuality are then considered. It is proposed that asexual species may arise in one step by mutation or loss of a key gene(s), and/or there may be a 'slow decline' in sexual fertility within the species as a whole. Thus, it is argued that species should not be considered simply as sexual or asexual, but rather as individual isolates being present on a continuum of sexual fertility, with the implications for understanding sexuality/asexuality in A. fumigatus discussed.

Aspergillus fumigatus↗
PubMed2003

Evidence of recombination in putative ancient asexuals.

Ancient asexuals have been considered to be a contradiction of the basic tenets of evolutionary theory. Barred from rearranging genetic variation by recombination, their reduced number of gene arrangements is thought to hamper their response to changing environments. For the same reason, it should be difficult for them to avoid the build-up of deleterious mutations. Several groups of taxonomically diverse organisms are thought to be ancient asexuals, although clear evidence for or against the existence of recombination events is scarce. Several methods have recently been developed for predicting recombination events by analyzing aligned sequences of a given region of DNA that all originate from one species. The methods are based on phylogenetic, substitution, and compatibility analyses. Here we present the results of analyses of sequence data from different loci studied in several groups of evolutionarily distant species that are considered to be ancient asexuals, using seven different types of analysis. The groups of organisms were the arbuscular mycorrhizal fungi (Glomales), Darwinula stevensoni (Darwinuloidea crustacean ostracods) and the bdelloid rotifers (Bdelloidea), which are thought to have been asexual for the last 400, 25-100, and 35-40 Myr, respectively. The seven different analytical methods evaluated the evolutionary relationships among haplotypes, and these methods had previously been shown to be reliable for predicting the occurrence of recombination events. Despite the different degree of genetic variation among the different groups of organisms, at least some evidence for recombination was found in all species groups. In particular, predictions of recombination events in the arbuscular mycorrhizal fungi were frequent. Predictions of recombination were also found for sequence data that have previously been used to infer the absence of recombination in bdelloid rotifers. Although our results have to be taken with some caution because they could signal very ancient recombination events or possibly other genetic variation of nonrecombinant origin, they suggest that some cryptic recombination events may exist in these organisms.

Animals↗
PubMed2003

How ancient are ancient asexuals?

Ancient asexual animal groups, such as bdelloid rotifers and darwinuloid ostracods, are excellent model organisms to study the effects of long-term asexuality. However, the absolute length of time that these groups have been fully asexual is mostly ignored. In the case of the darwinuloid ostracods, the fossil record shows that sexual reproduction disappeared almost completely after the end of Permian mass extinction (ca. 245 Myr ago), although several putative records of males from the Mesozoic obscure the exact time-frame of obligate asexuality in darwinuloids. Here, we re-examine the Mesozoic darwinuloid records, with regard to the reproductive mode of the assemblages. Three criteria to distinguish males in fossil populations (lack of brood pouch, position of muscle scars and size dimorphism) are used here to test for the presence of males in darwinuloid assemblages. A large, well-preserved assemblage of Darwinula leguminella (Forbes 1885) from the latest Jurassic (ca. 145 Myr ago) of England is found to be markedly variable in size and shape, but nevertheless turns out to be an all female assemblage. The exceptional preservation of the material also allows the re-assignment of this species to the extant darwinuloid genus Alicenula. All other putative dimorphic darwinuloid records from the Mesozoic are re-examined using the same criteria. The hypothesis that these assemblages represent bisexual populations is rejected for all post-Triassic (ca. 208 Myr ago) records.

Animals↗
PubMed2002

Switching from asexual to sexual reproduction in the planarian Dugesia ryukyuensis: change of the fissiparous capacity along with the sexualizing process.

Asexual worms of fissiparous strain of the planarian Dugesia ryukyuensis switch from asexual to sexual reproduction, if they are fed with sexually mature worms of Bdellocephala brunnea. This suggests that the sexually mature worms have a sexualizing substance(s) that induces the sexuality in the asexual worms. Here, we found by analysis of the sexualization that the cessation of the fission, namely their asexual reproduction, occurs immediately after the acquisition of sexuality. This result suggests that the downstream mechanisms induced by the putative sexualizing substance in B. brunnea become responsible for the cessation of fission. We also found that the decapitation triggers fission in the worms even after the acquisition of sexuality if they are not sexually mature, while the fully sexualized worms never fission even though they are decapitated. This result suggests that the cessation of fission takes place via at least two steps: (1) the mechanisms associated with the cephalic system; (2) other mechanisms independent of cephalic control.

Animals↗
PubMed1993

Falciparum malaria in naturally infected human patients: I. Ultrastructural differences between malaria pigments in intraerythrocytic asexual and sexual forms.

Venous blood samples were taken from patients naturally infected with the human malaria parasite Plasmodium falciparum. Two types of malaria pigment (MP) particles have been demonstrated in intraerythrocytic asexual forms (trophozoites and schizonts), while a single type was detected in gametocytes. Type I MP particles, found in both asexual and sexual forms, are electron-dense. It is suggested that these are proteinaceous and may be intermediate, utilizable metabolic products that serve as a food reserve during development of the parasite in the human host and also during the growth cycle of the sexual form in the mosquito. In asexual forms, type I particles occur within food vacuoles (FV) containing semidigested hemoglobin (Hg), while they are unenveloped in the cytoplasm of the sexual forms. Type II MP particles, found in electron-lucent residual bodies, are crystalloid and of low electron density. It is suggested that these are the final, waste product of Hg digestion in the asexual forms.

Animals↗
PubMed2000

Modelling the transition of asexual blood stages of Plasmodium falciparum to gametocytes.

In this paper, we investigate the transition of asexual blood stages of P. falciparum to gametocytes. The study is based on daily data, collected from 262 individual courses of parasitaemia. We propose several mathematical models that follow biological reasoning. The models are fitted with maximum likelihood and are compared with each other. The models differ in the assumptions made about the mortality of circulating gametocytes and about the transition rate of the asexual parasites. Gametocyte mortality is modelled as being (i) constant over time, (ii) linearly increasing over time, (iii) linearly increasing over gametocyte age, and (iv) exponentially increasing over gametocyte age, respectively. The transition rate is either kept constant per patient or piecewise constant within intervals that correspond to waves of asexual parasitaemia which are assumed to be caused by different Pf(emp1)-variants. According to likelihood ratio tests, the models with age-dependent mortality rate and wave-dependent transition rates are superior to the models with constant transition rate and/or constant or time-dependent mortality rate. The best fits are reached for models with exponentially increasing (Gompertz-type) mortality. Furthermore, an impact of high asexual parasite densities on the survival of gametocytes, interpreted as a cytokine-mediated effect, is evident in some cases.

Animals↗
PubMed1996

Phylogeography of an asexual marine clam complex, Lasaea, in the northeastern Pacific based on cytochrome oxidase III sequence variation.

Northeastern Pacific populations of the intertidal marine clam Lasaea are exclusively composed of polyploid asexual clones that lack pelagic larval development. We investigated the phylogeographic structure of this clonal assemblage by assaying genetic divergence in 4 populations along 2000 km of the west coast of North America. In each population a 462-nt portion of the mitochondrial Cytochrome Oxidase III gene was amplified and sequenced for 30 individuals. A total of 21 haplotypes were detected and phylogenetic analyses resolved this variation into 5 main branches that differ in sequence composition by 8.9-14.5%. Two of the branches each encompass a single haplotype, the other 3 terminate in clades containing 3-11 lineages and within clade divergence ranges from 0.4 to 2.3%. Obvious geographic structuring was evident in that 19 of the 21 lineages were restricted in their distributions to single populations and 3 of the main phylogenetic branches were limited to specific marine biogeographic provinces. Two of the clades were encountered in multiple, geographically distant populations; however, the absence of shared haplotypes indicates that long distance gene flow is rare among these direct-developing populations. Northeastern Pacific Lasaea clones form a robust monophyletic grouping relative to other studied populations of the cosmopolitan genus. We estimate divergence times among northeastern Pacific clones to range from 3 to 6 myr. If the latest common ancestor of these exclusively asexual, genetically diverse, regionally monophyletic Lasaea lineages was also asexual, this represents a minimal age estimate for asexuality in the genus. The most plausible alternate hypothesis requires that at least 5 northeastern Pacific sexual parental species have recently become extinct.

Animals↗
PubMed1996

An asexual fungus has the potential for sexual development.

The availability of cloned genes that control sexual reproduction (mating type genes) in high fungi has allowed us to consider the causes of failure to mate in asexual fungi. We report here that the asexual fungus Bipolaris sacchari has a homolog of the MAT-2 gene of its sexual ascomycete relative Cochliobolus heterostrophus. The B. sacchari MAT-2 sequence is highly similar to that of C. heterostrophus MAT-2 and, in fact, functions in transgenic C. heterostrophus. Thus, the asexual nature of B. sacchari is not due to absence or mutation of MAT. When either of the C. heterostrophus MAT genes was transformed into B. sacchari, the recipient could neither self nor cross with other B. sacchari strains, in contrast to transgenic C. heterostrophus strains which can do both. Persistent asexuality of B. sacchari, in spite of the presence of complementary functional MAT genes, suggests that this fungus lacks genes other than MAT which are essential for mating. Notably, the transgenic B. sacchari strains were sometimes able to initiate, but not complete, sexual development in interspecific pairings with C. heterostrophus. Transcript analysis showed that the B. sacchari MAT-2 gene is expressed in transgenic C. heterostrophus and that the C. heterostrophus MAT genes are expressed in transgenic B. sacchari. No transcript of the native B. sacchari MAT-2 gene was detected under any growth condition tested.

Amino Acid Sequence↗
PubMed2006

Does an asexual endophyte symbiont alter life stage and long-term survival in a perennial host grass?

Asexual, seedborne endophytic fungi in perennial grasses are often viewed as strong mutualists because fitness of the symbiont and host grass are closely coupled. However, at least for some native grasses, the asexual endophyte, Neotyphodium, acts parasitically, yet remains at high frequencies in natural populations. Most previous studies of Neotyphodium effects on host survival have been short term relative to the long life span of the perennial grass host. We therefore tested the hypothesis that Neotyphodium alters the survival in various life stages and long-term survival of adult native Arizona fescue (Festuca arizonica). To test the former, we planted 40 infected (E+) and 40 uninfected (E-, endophyte removed) seeds from four different maternal plants in the field under ambient conditions. We followed survival of seeds, seedlings, and adult plants over a 5-year period. To test the latter, we determined the infection of 1633 adult plants and followed their survival over the next 5-7 years. E+ seeds did not differ from uninfected seeds in terms of overall survival from seed germination to seedling to adult. However, the shape of the survival curve differed, with E+ plants showing higher mortality in early life stages. E+ adult plants did not differ from E- plants in long-term survival. Survival was generally very high during the study, which included a severe and prolonged drought. Infection by asexual Neotyphodium does not increase survival in early life stages or that of adult plants. Because asexual, vertically transmitted symbionts are predicted by evolutionary theory to be strong mutualists, the persistence of high infection frequencies in natural populations without long-term benefits to the host remains enigmatic. One possible explanation is that the long life span of the perennial host and low seedling recruitment may obscure either the costs or benefits of endophyte infection.

Festuca↗
PubMed2005

Plasmodium berghei alpha-tubulin II: a role in both male gamete formation and asexual blood stages.

Plasmodium falciparum contains two genes encoding different isotypes of alpha-tubulin, alpha-tubulin I and alpha-tubulin II. alpha-Tubulin II is highly expressed in male gametocytes and forms part of the microtubules of the axoneme of male gametes. Here we present the characterization of Plasmodium berghei alpha-tubulin I and alpha-tubulin II that encode proteins of 453 and 450 amino acids, respectively. alpha-Tubulin II lacks the well-conserved three amino acid C-terminal extension including a terminal tyrosine residue present in alpha-tubulin I. Investigation of transcription by Northern analysis and RT-PCR and analysis of promoter activity by GFP tagging showed that alpha-tubulin I is expressed in all blood and mosquito stages. As expected, alpha-tubulin II was highly expressed in the male gametocytes, but transcription was also observed in the asexual blood stages, female gametocytes, ookinetes and oocysts. Gene disruption experiments using standard transfection technologies did not produce viable parasites indicating that both alpha-tubulin isotypes are essential for the asexual blood stages. Targeted modification of alpha-tubulin II by the addition of the three C-terminal amino acids of alpha-tubulin I did not affect either blood stage development nor male gamete formation. Attempts to modify the C-terminal region by adding a TAP tag to the endogenous alpha-tubulin II gene were not successful. Introduction of a transgene, expressing TAP-tagged alpha-tubulin II, next to the endogenous alpha-tubulin II gene, had no effect on the asexual blood stages but strongly impaired formation of male gametes. These results show that alpha-tubulin II not only plays an important role in the male gamete but is also expressed in and essential for asexual blood stage development.

Animals↗
PubMed2003

Why do species exist? Insights from sexuals and asexuals.

Why does life diversify into the more or less discrete entities we recognise as species? Two main explanations have been proposed: i) species are a consequence of adaptation to different ecological niches, ii) species are a consequence of sexual reproduction and reproductive isolation. Phylogenetic studies of case-study groups can provide insights into the relative importance of divergent selection and isolation for speciation, but it can be difficult to infer causes of speciation unambiguously. The example of North American tiger beetles from the genus Cicindela is discussed. An alternative approach is to compare diversification between related sexual and asexual taxa to infer the relative importance of the two explanations. We outline expected patterns of diversification in sexual and asexual lineages under different scenarios using coalescent theory. Whether sexuals or asexuals diversify to a greater extent depends on the balance among various stages of diversification, particularly on the effects of sexual reproduction on rates of adaptive evolution. Rotifers offer a unique system to test these ideas, allowing comparison of patterns of genetic and functional morphological diversification in sexual (bdelloid) and asexual (monogonont) clades.

Journal Article↗
PubMed2026

High levels of mitotic gene conversion are needed to effectively purge deleterious mutations in asexual organisms.

Self-fertilisation and asexual reproduction are both hypothesised to cause long-term extinction due to inefficient selection against deleterious mutations. Self-fertilisation can counter these effects through creating homozygous genotypes and purging deleterious mutations. Although complete asexuality lacks meiotic gene exchange, mitotic gene conversion creates homozygous regions that could limit deleterious mutation accumulation in an analogous manner. We compare mutation accumulation in self-fertilising and facultative sexual populations subject to mitotic gene conversion, and quantify the efficacy of purging in the latter. We first show analytically that purging is most effective with high levels of asexuality and gene conversion, and when deleterious mutations are recessive. We further show using simulations that, when mitotic gene conversion becomes sufficiently high in obligate asexuals, there is a reduction in the mutation count and a jump in homozygosity, reflecting purging. However, this mechanism is not necessarily as efficient at purging under high self-fertilisation, and elevated rates of mitotic gene conversion seem to be needed for widespread purging compared to empirical estimates. If gene conversion rates are allowed to evolve, then elevated rates that increase mean fitness can arise, but only if there is sufficient variance in the gene conversion rate. Conversely, if gene conversion rates are already high and rates are not constrained then they will slightly decrease, reducing mean fitness.

Self-fertilisation↗
PubMed2006

Temporal differentiation and spatial coexistence of sexual and facultative asexual lineages of an aphid species at mating sites.

Cases of coexisting sexual and asexual relatives are puzzling, as evolutionary theory predicts that competition for the same ecological niches should lead to the exclusion of one or the other population. In the cyclically parthenogenetic aphid, Rhopalosiphum padi, sexual and facultative asexual lineages are admixed in space at the time of sexual reproduction. We investigated how the interaction of reproductive mode and environment can lead to temporal niche differentiation. We demonstrated theoretically that differential sensitivity of sexual and facultatively asexual aphids to an environmental parameter (mating host suitability) shapes the two strategies: whereas the sexual lineages switch earlier to the production of sexual forms, the facultative asexual lineages delay and spread out their investment in sexual reproduction. This predicted pattern of niche specialization is in agreement with the temporal structure revealed in natura by demographic and genetic data. We propose that partial loss of sex by one pool of aphids and subsequent reduction in gene flow between lineages may favour temporal specialization through disruptive selection.

Animals↗
PubMed1996

Antibodies to a conserved-motif peptide sequence of the Plasmodium falciparum thrombospondin-related anonymous protein and circumsporozoite protein recognize a 78-kilodalton protein in the asexual blood stages of the parasite and inhibit merozoite invasion in vitro.

Athrombospondin-related anonymous protein (TRAP) of the human malaria parasite Plasmodium falciparum shares highly conserved amino acid sequence motifs with the circumsporozoite protein of all plasmodia sequenced so far, as well as with unrelated proteins like thrombospondin and properdin. Although it was first described as an asexual blood stages protein, there has been some controversy about its expression in these stages. Pursuant to our interest in the conserved sequences within the malaria antigens, we synthesized an 18-residue peptide (18-mer) representing a conserved motif of TRAP and raised polyclonal antibodies against it. In an immunoblot assay in which we probed proteins from the asexual blood stages of the parasite, we found that this antibody recognized predominantly a 78-kDa protein in the whole parasite lysate. Furthermore, in another immunoblot, the recombinant TRAP constructs containing the conserved-motif sequence were distinctly recognized by the antipeptide antibodies, whereas a construct lacking the motif sequence was not, suggesting that the antibodies specifically cross-reacted with a protein which might be a TRAP-like protein present in the asexual blood stages of the parasite. Also, in an immunofluorescence assay, this antibody brightly stained the acetone-fixed trophozoites of the parasite. Most significantly, anti-18-mer immunoglobulin G, as well as antipeptide antibody against a smaller (nonamer) construct representing the most conserved motif within the 18-mer, inhibited the merozoite invasion of erythrocytes in a dose-dependent manner. These results provide evidence of the expression of TRAP or a TRAP-like protein in the asexual blood stages of the parasite and of a possible role of the conserved motifs in the parasite-host cell interaction during the process of invasion.

Amino Acid Sequence↗
PubMed2025

Investigating the Functions of Hox Genes Using Planarian Asexual Reproduction.

Hox genes are highly conserved developmental regulators instrumental to the formation of a wide range of diverse body plans across metazoans. While significant progress in the field of Hox gene research has been made, persistent challenges in unraveling their mechanisms of action and full repertoire of functions remain. To date, investigations of Hox gene function have been primarily conducted in research models belonging to ecdysozoa and vertebrata. Herein we summarize recent findings on Hox genes' roles in the asexual reproduction of the regenerative flatworm planaria, a member of the understudied superphylum Spiralia. We detail our optimized methods for planarian culture, gene perturbation, and induction of asexual reproduction. We aim to provide an experimentally tractable means to dissect Hox gene adult tissue functions underlying planarian asexual reproduction with broader relevance to Hox genes' established and emerging roles in regulating cellular behaviors, developmental patterning, animal behavior, and tissue regeneration.

Animals↗
PubMed2002

Coexistence of sexual and asexual conspecifics: a cellular automaton model.

Most previous models of populations mixed for reproductive mode have omitted important local interactions between sexual and asexual individuals. We propose a cellular automaton model where local rules focus on fertilization and colonization. This model produces rich sets of data which are then studied by means of spatial statistics. Results point to the fixation of one of the two reproductive modes in the landscape. However, some examples of coexistence of sexual and asexual conspecifics over long periods of time are also found. This model is an example of a CA that diverges from its mean field approximation. The formation of sexual and asexual clusters reduces effective colonization rate in the CA and may account for this behaviour.

Biological Evolution↗
PubMed2003

Feeding and asexual reproduction of the jellyfish Sarsia gemmifera in response to resource enrichment.

Jellyfish are ubiquitous predators in marine pelagic environments and can sometimes control their zooplankton prey populations. Recent considerations of the fertilization of entire food webs in coastal areas make it important to investigate the response of jellyfish to resource enrichment. We investigated feeding, assimilation and life history parameters in the hydromedusa species Sarsia gemmifera. S. gemmifera was able to ingest up to 3 micro g carbon per hour, which corresponds to a daily carbon ingestion that exceeds the individual's body weight (carbon). Conversion of ingested carbon into tissue was less than 30%. The assimilated carbon was allocated such that approximately 65% was used for growth and the remainder for asexual reproduction. Carbon from food was allocated to asexually produced offspring within hours. The numerical response of S. gemmifera reached saturation at prey levels of 100 or more copepods per liter. Propagule quality was influenced by maternal effects: higher net production of the mothers in higher food environments resulted in higher carbon content of individual propagules. Starvation resistance of propagules was therefore positively related to food density in the maternal environment. The food concentrations which S. gemmifera normally experiences in the field are much lower than the food levels at which this species had its maximum asexual reproductive output in the laboratory. Therefore, S. gemmifera may potentially benefit from food web perturbations which increase crustacean zooplankton densities.

Adaptation, Physiological↗
PubMed2005

Adaptation in the asexual false spider mite Brevipalpus phoenicis: evidence for frozen niche variation.

Because asexual species lack recombination, they have little opportunity to produce genetically variable offspring and cannot adapt to changes in their environment. However, a number of asexual species are very successful and appear to contradict this general view. One such species is the phytophagous mite Brevipalpus phoenicis (Geijskes), a species that is found in a wide range of environments. There are two general explanations for this pattern, the General Purpose Genotype (GPG) and Frozen Niche Variation (FNV). According to the GPG model, an asexual species consists of clones that can all survive and reproduce in all the different niches. Alternatively, the FNV model postulates that different clones are specialized to different niches. We have performed a test to distinguish between these models in B. phoenicis. Mites from three populations from three different host plant species (citrus, hibiscus and acerola) were transplanted to their own and the two alternative host plants and mite survival and egg production were measured. Additionally, the mite populations were genotyped using microsatellites. Fitness was seriously reduced when mites were transplanted to the alternative host plant species, except when the alternative host was acerola. We concluded that B. phoenicis clones are specialized to different niches and thus the FNV best describes the broad ecological niche of this species but that there is also some evidence for host plant generalization. This conclusion was strengthened by the observations that on each host plant species the native mite population performed better than the introduced ones, and that three microsatellite markers showed that the mite populations are genetically distinct.

Adaptation, Physiological↗