Mixed asexual and sexual reproduction in the Indo‐Pacific reef coral Pocillopora damicornis

Pocillopora damicornis is one of the best studied reef‐building corals, yet it's somewhat unique reproductive strategy remains poorly understood. Genetic studies indicate that P. damicornis larvae are produced almost exclusively parthenogenetically, and yet population genetic surveys suggest frequent sexual reproduction. Using microsatellite data from over 580 larvae from 13 colonies, we demonstrate that P. damicornis displays a mixed reproductive strategy where sexual and asexual larvae are produced simultaneously within the same colony. The majority of larvae were parthenogenetic (94%), but most colonies (10 of the 13) produced a subset of their larvae sexually. Logistic regression indicates that the proportion of sexual larvae varied significantly with colony size, cycle day, and calendar day. In particular, the decrease in sexual larvae with colony size suggests that the mixed reproductive strategy changes across the life of the coral. This unique shift in reproductive strategy leads to increasingly asexual replications of successful genotypes, which (in contrast to exclusive parthenogens) have already contributed to the recombinant gene pool.     

Male offspring production by asexual Potamopyrgus antipodarum, a New Zealand snail

As only females contribute directly to population growth, sexual females investing equally in sons and daughters experience a two-fold cost relative to asexuals producing only daughters. Typically, researchers have focused on benefits of sex that can counter this 'cost of males' and thus explain its predominance. Here, we instead ask whether asexuals might also pay a cost of males by quantifying the rate of son production in 45 experimental populations ('lineages') founded by obligately asexual female Potamopyrgus antipodarum. This New Zealand snail is a powerful model for studying sex because phenotypically similar sexual and asexual forms often coexist, allowing direct comparisons between sexuals and asexuals. After 2 years of culture, 23 of the 45 lineages had produced males, demonstrating that asexual P. antipodarum can make sons. We used maximum-likelihood analysis of a model of male production in which only some lineages can produce males to estimate that ~50% of lineages have the ability to produce males and that ~5% of the offspring of male-producing lineages are male. Lineages producing males in the first year of the experiment were more likely to make males in the second, suggesting that some asexual lineages might pay a cost of males relative to other asexual lineages. Finally, we used a simple deterministic model of population dynamics to evaluate how male production affects the rate of invasion of an asexual lineage into a sexual population, and found that the estimated rate of male production by asexual P. antipodarum is too low to influence invasion dynamics.     

Sex is not a solution for reproduction: the libertine bubble theory

Here, I propose a new hypothesis: sex originated from an archaic gene transfer process among prebiotic bubbles without the prerequisite for reproduction. This de‐coupling from reproduction might make the thorny problem of accounting for the evolution of sex, despite the apparent advantages of parthenogenicity, more tractable.     

Habitat heterogeneity favors asexual reproduction in natural populations of grassthrips

Explaining the overwhelming success of sex among eukaryotes is difficult given the obvious costs of sex relative to asexuality. Different studies have shown that sex can provide benefits in spatially heterogeneous environments under specific conditions, but whether spatial heterogeneity commonly contributes to the maintenance of sex in natural populations remains unknown. We experimentally manipulated habitat heterogeneity for sexual and asexual thrips lineages in natural populations and under seminatural mesocosm conditions by varying the number of hostplants available to these herbivorous insects. Asexual lineages rapidly replaced the sexual ones, independently of the level of habitat heterogeneity in mesocosms. In natural populations, the success of sexual thrips decreased with increasing habitat heterogeneity, with sexual thrips apparently only persisting in certain types of hostplant communities. Our results illustrate how genetic diversity‐based mechanisms can favor asexuality instead of sex when sexual lineages co‐occur with genetically variable asexual lineages.     

Morphological characterization of the asexual reproduction in the acorn worm Balanoglossus simodensis

The acorn worm Balanoglossus simodensis reproduces asexually by fragmentation and subsequent regeneration from the body fragments. We examined the morphogenesis of its asexual reproduction. At first, we collected asexually reproducing specimens and observed their morphogenesis. Then, we succeeded in inducing the asexual reproduction artificially by cutting the worm at the end of the genital region. The process of morphogenesis is completely the same between naturally collected and artificially induced specimens. The stages during morphogenesis were established on the basis of the external features of the asexually reproducing fragments. The internal features of the fragments were also examined at each stage. In a separate phase of the study, the capacity for regeneration of some body parts was also examined by dividing intact worms into about 10 fragments. Although the capacity for regeneration varied among the different body parts, some fragments regenerated into complete individuals in 1 month. The process of regeneration was the same as that in the asexually produced fragments.     

Expression and function of myc during asexual reproduction of the budding ascidian Polyandrocarpa misakiensis

The budding ascidian Polyandrocarpa misakiensis proliferates asexually by budding. The atrial epithelium is a multipotent but differentiated tissue, which transdifferentiates into various tissues and organs after the bud separates from the parental body. We isolated cDNA clones homologous to the myc proto-oncogene from P. misakiensis. The cDNA, named Pm-myc, encoded a polypeptide of 639 amino acid residues, containing Myc-specific functional motifs, Myc box I and Myc box II, and the basic helix-loop-helix domain. Expression of Pm-myc was observed in the atrial epithelium in the organ-forming region of the developing bud, where the epithelial cells dedifferentiate and re-enter the cell cycle. The expression was also observed in fibroblast-like cells, which are known to participate in the organogenesis together with the epithelial cells. Unexpectedly, the atrial epithelium expressed Pm-myc more than one day before the dedifferentiation. The organogenesis was disturbed by Pm-myc-specific double-stranded RNA. In situ hybridization revealed that Pm-myc-positive fibroblast-like cells disappeared around the organ primordium of the dsRNA-treated bud. The results suggest that the mesenchymal-epithelial transition of fibroblast-like cells is important for the organogenesis in this budding ascidian species.     

Food limitation of asexual reproduction in a spionid polychaete

Summary

The rate of asexual fragmentation in the spionid polychaete Pygospio elegans is shown to increase in the presence of augmented food levels and is a density-dependent response. Asexual reproduction does not occur when the animals are reproducing sexually.     

Locomotion,asexual reproduction,and killing of corals by the corallimorpharian Corynactis californica

The rates at which some sedentary coelenterates move across substrates, spread by asexual reproduction, and competitively damage their neighbors may greatly affect their ability to survive on hard marine substrates. However, almost nothing is known about these processes in the Corallimorpharia. In this study, locomotory rates varied widely between members of a single clone of the corallimorpharian Corynactis californica. Polyp movement rates (0–14 mm month^–1) were slower than those of many Actiniaria, possibly due to the lack of basilar muscles in corallimorpharians. Rates of asexual reproduction by C. californica polyps in the laboratory varied 10-fold among 8 clones. The minimal time to double the number of polyps per clone (2 months) was intermediate to doubling times known for Actiniaria and Scleractinia. The rate at which polyps killed individuals of the scleractinian Astrangia lajollaensis also varied widely within and between clones. Asexual reproductive rate correlated positively with coral killing rate.     

Multipotent epithelial cells in the process of regeneration and asexual reproduction in colonial tunicates

The cellular and molecular features of multipotent epithelial cells during regeneration and asexual reproduction in colonial tunicates are described in the present study. The epicardium has been regarded as the endodermal tissue-forming epithelium in the order Enterogona, because only body fragments having the epicardium exhibit the regenerative potential. Epicardial cells in Polycitor proliferus have two peculiar features; they always accompany coelomic undifferentiated cells, and they contain various kinds of organelles in the cytoplasm. During strobilation a large amount of organelles are discarded in the lumen, and then, each tissue-forming cell takes an undifferentiated configuration. Septum cells in the stolon are also multipotent in Enterogona. Free cells with a similar configuration to the septum inhabit the hemocoel. They may provide a pool for epithelial septum cells. At the distal tip of the stolon, septum cells are columnar in shape and apparently undifferentiated. They are the precursor of the stolonial bud. In Pleurogona, the atrial epithelium of endodermal origin is multipotent. In Polyandrocarpa misakiensis, it consists of pigmented squamous cells. The cells have ultrastructurally fine granules in the cytoplasm. During budding, coelomic cells with similar morphology become associated with the atrial epithelium. Then, cells of organ placodes undergo dedifferentiation, enter a cell division cycle, and commence morphogenesis. Retinoic acid-related molecules are involved in this dedifferentiation process of multipotent cells. We conclude that in colonial tunicates two systems support the flexibility of tissue remodeling during regeneration and asexual reproduction; dedifferentiation of epithelial cells and epithelial transformation of coelomic free cells.     

Sexual and asexual reproduction in geographically separated populations of the fissiparous asteroid Coscinasterias calamaria (Gray)

Sexual reproduction and asexual reproduction by fission were studied in four populations of Coscinasterias calamaria (Gray), two in Otago Harbour in the South Island of New Zealand and two in the North Island near Auckland. The annual reproductive cycle in both islands of New Zealand is clearly defined with a spawning season between November and January. In both sites the pyloric caeca index was approximately inverse to the gonad index cycle as found in other forcipulate asteroids. There are substantial differences in the sex ratios of mature starfish at each site studied, with 1:1 ratios in two populations, one population heavily biased towards females and the fourth consisting almost entirely of males. Morphometric variation in arm number due to splitting was studied and the frequency of splitting varied considerably between the four populations. Generally speaking sublittoral C. calamaria divide less frequently than intertidal starfish and populations in which food is less abundant or of poorer quality are more fissiparous and put less energy into sexual reproduction, than populations with plentiful readily available food in the form of mussels. The significance of the different reproductive patterns in C. calamaria is discussed.     

Soft selection reduces loss of heterozygosity in asexual reproduction

The adaptive value of sexual reproduction is still debated in evolutionary theory. It has been proposed that the advantage of sexual reproduction over asexual reproduction is to promote genetic diversity, to prevent the accumulation of harmful mutations or to preserve heterozygosity. Since these hypothetical advantages depend on the type of asexual reproduction, understanding how selection affects the taxonomic distribution of each type could help us discriminate between existing hypotheses. Here, I argue that soft selection, competition among embryos or offspring in selection arenas prior to the hard selection of the adult phase, reduces loss of heterozygosity in certain types of asexual reproduction. Since loss of heterozygosity leads to the unmasking of recessive deleterious mutations in the progeny of asexual individuals, soft selection facilitates the evolution of these types of asexual reproduction. Using a population genetics model, I calculate how loss of heterozygosity affects fitness for different types of apomixis and automixis, and I show that soft selection significantly reduces loss of heterozygosity, hence increases fitness, in apomixis with suppression of the first meiotic division and in automixis with central fusion, the most common types of asexual reproduction. Therefore, if sexual reproduction evolved to preserve heterozygosity, soft selection should be associated with these types of asexual reproduction. I discuss the evidence for this prediction and how this and other observations on the distribution of different types of asexual reproduction in nature is consistent with the heterozygosity hypothesis.     

Sediment type and the clonal size greatly affect the asexual reproduction,productivity, and nutrient absorption of Vallisneria natans

Most aquatic vegetation restorations involve the transplantation of submerged macrophytes. Sediment type and the clonal size are of great significance as they determine the fate of submerged macrophytes. In order to ensure successful restoration, a simulation experiment was conducted using aquarium mesocosms to investigate the response of stolon propagation capacity, the morphological features and productivity of Vallisneria natans for four types of sediment (lake mud [L], lake mud + sand [L + S, 50:50, v/v mixture], sand [S], clay [C]), and three types of clonal sizes. Results showed that sediment types significantly affected V. natans biomass accumulation, stolon propagation ability, ramet morphological characteristics, and productivity, where the asexual reproduction ability and productivity ranked as L > L + S > S > C in four sediment types. Total biomass, maximum net production, number of ramets, root diameter, number of stolons, and stolon propagation rate were all highest in L. In L and L + S, the plant chlorophyll content was higher than in S and C. The root diameter and the ratio of aboveground/underground biomass in S were the smallest among the four sediments. Moreover, when more V. natans seedlings were linked, more ramets and biomass were produced. The stolon propagation rate was ranked as the stolon with single seedling greater than the stolon with two‐linked seedlings greater than the stolon with three‐linked seedlings in L and L + S. The concentration of total nitrogen, total phosphorus, and NO₃^?‐N in water was remarkably reduced in four aquariums. Findings provide a scientific basis for restoring submerged macrophytes in different sediment settings.     

Parthenogenetic female populations in the brown alga Scytosiphon lomentaria (Scytosiphonaceae,Ectocarpales): decay of a sexual trait and acquisition of asexual traits

In isogamous brown algae, the sexuality of populations needs to be tested by laboratory crossing experiments, as the sexes of gametophytes are morphologically indistinguishable. In some cases, gamete fusion is not observed and the precise reproductive mode of the populations is unknown. In the isogamous brown alga Scytosiphon lomentaria in Japan, both asexual (gamete fusion is unobservable) and sexual populations (gamete fusion is observable) have been reported. In order to elucidate the reproductive mode of asexual populations in this species, we used PCR‐based sex markers to investigate the sex ratio of three asexual and two sexual field populations. The markers indicated that the asexual populations consisted only of female individuals, whereas sexual populations are composed of both males and females. In culture, female gametes of most strains from asexual populations were able to fuse with male gametes; however, they had little to no detectable sexual pheromones, significantly larger cell sizes, and more rapid parthenogenetic development compared to female/male gametes from sexual populations. Investigations of sporophytic stages in the field indicated that alternation of gametophytic and parthenosporophytic stages occur in an asexual population. These results indicate that the S. lomentaria asexual populations are female populations that lack sexual reproduction and reproduce parthenogenetically. It is likely that females in the asexual populations have reduced a sexual trait (pheromone production) and have acquired asexual traits (larger gamete sizes and rapid parthenogenetic development).     

Automixis: its distribution and status

Biologists have conclusively failed to arrive at a generally acceptable definition of sexual reproduction. Because of this, several reproductive processes are seen as sexual by some authors but as asexual by others. Included among these are automictic methods of reproduction. Automixis describes several reproductive processes whereby a new individual derives from a product or products of a single meiotically dividing cell. Several forms involve an episode of nuclear fusion and it is argued that, because of this, they should be seen as sexual processes irrespective of whether the fusing bodies are differentiated as gametes or are simply meiotic tetrad nuclei. Other forms involve no episode of nuclear fusion and it is argued that, because of this, they should be seen as asexual processes. These latter forms involve the generation of diploid eggs either by restitutional meioses, or by an endomitotic event preceding or following a reductional meiosis, or involve the generation of a diploid embryo by the fusion of cleavage division nuclei in a haploid embryo; in each case the egg develops parthenogenetically. In addition to the disagreement that exists over the reproductive status of automixis, considerable confusion exists over its taxonomic distribution. It is often described as being restricted to a few species of insects, where it is parthenogenetic, but in factde range of taxa, including both isogamous and anisogamous plants and fungi, where it may be either parthenogenetic or non-parthenogenetic. This confusion results both from a failure of many biologists writing on this subject to adequately consider the variation in life-cycles existing between major taxa and from a general failure by botanists and mycologists to distinguish between automixis and autogamous forms of self-fertilization (in which the fusing nuclei derive from different meioses). It is further compounded by a proliferation of synonyms for automictic processes. Thus in a number of publications automictic processes are variously described as being matromorphic, thelytokous, parthenogamic, autogamic or apomictic rather than as being automictic.     

Origin and maintenance of sex: the evolutionary joys of self sex

Sex is generally thought of as meiosis, conjugation, and syngamy, with the primary function of sex believed to be genetic mixing. However, conjugation does not occur with complete automixis, whereas syngamy does not occur with restitutional automixis. Self sex in the forms of automixis and autogamy does not include genetic mixing. Yet sex, including self sex, is necessary for most eukaryotic lineages. What is the purpose of sex without genetic mixing? Obligate self sex is not an evolutionary dead end, but holds the key to understanding the evolutionary origin, function, maintenance, and ubiquity of sex. We extend the rejuvenescence hypothesis that sex provides a necessary developmental reset for multicellular eukaryotes and even many unicellular eukaryotes. Sex reduces additive genetic variance of epigenetic signals, especially cytosine methylation, and of ploidy levels. Furthermore, we argue that syngamy is a modified form of meiosis that maintains ploidy and resets epigenetic signals. Epigenetic resetting is consistent with sex being induced by starvation or desiccation. Diminution of additive genetic variance is consistent with the origin and maintenance of an adaptive trait, sex, that has been present for approximately two billion years. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 707–728.     

Ant queens adjust egg fertilization to benefit from both sexual and asexual reproduction

An enduring problem in evolutionary biology is the near ubiquity of sexual reproduction despite the inherent cost of transmitting only half the parent's genes to progeny. Queens of some ant species circumvent this cost by using selectively both sexual reproduction and parthenogenesis: workers arise from fertilized eggs, while new queens are produced by parthenogenesis. We show that queens of the ant Cataglyphis cursor maximize the transmission rate of their genes by regulating the proportion of fertilized and parthenogenetic eggs laid over time. Parthenogenetic offspring are produced in early spring, when workers raise the brood into sexuals. After the mating period, queens lay mostly fertilized eggs that will be reared as the non-reproductive caste.     

Effects of low dissolved oxygen on survival and asexual reproduction of scyphozoan polyps (Chrysaora quinquecirrha)

Hypoxic conditions are common in many coastal environments such as Chesapeake Bay. While medusae appear to be quite tolerant of low dissolved oxygen (DO) concentrations, the effects of hypoxia on the benthic polyp stages are unknown. Chrysaora quinquecirrha (DeSor) polyps, and were subjected to 5 DO treatments (air-saturated [control], 3.5, 2.5, 1.5 and 0.5 mg l^–1) in the laboratory. Polyp survival and development were documented over 24 d. Virtually no mortality occurred in any treatment during the first 5 d. Total polyp mortality after 24 d was 59.3% at the lowest DO concentration, whereas <3% mortality was observed in the air-saturated treatment. Formation of stolons and strobilae occurred in all treatments, however, the proportions of polyps undergoing stolonation and strobilation were significantly greater in all DO concentrations above 0.5 mg l^–1. Polyp encystment was not observed in any treatment over the course of the 24 d experiment. These results indicate that polyps can survive and asexually propagate even during prolonged exposure to hypoxic conditions.     

A novel invertebrate trophic factor related to invertebrate neurotrophins is involved in planarian body regional survival and asexual reproduction

Trophic factors are a heterogeneous group of molecules that promote cell growth and survival. In freshwater planarians, the small secreted protein TCEN49 is linked to the regional maintenance of the planarian central body region. To investigate its function in vivo, we performed loss-of-function and gain-of-function experiments by RNA interference and by the implantation of microbeads soaked in TCEN49, respectively. We show that TCEN49 behaves as a trophic factor involved in central body region neuron survival. In planarian tail regenerates, tcen49 expression inhibition by double-stranded RNA interference causes extensive apoptosis in various cell types, including nerve cells. This phenotype is rescued by the implantation of microbeads soaked in TCEN49 after RNA interference. On the other hand, in organisms committed to asexual reproduction, both tcen49 mRNA and its protein are detected not only in the central body region but also in the posterior region, expanding from cells close to the ventral nerve chords. In some cases, the implantation of microbeads soaked in TCEN49 in the posterior body region drives organisms to reproduce asexually, and the inhibition of tcen49 expression obstructs this process, suggesting a link between the central nervous system, TCEN49, regional induction, and asexual reproduction. Finally, the distribution of TCEN49 cysteine and tyrosine residues also points to a common evolutionary origin for TCEN49 and molluscan neurotrophins.