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.     

The role of sexual and asexual reproduction in structuring high latitude populations of the reef coral Pocillopora damicornis

The genotypic composition of populations of the asexually viviparous coral Pocillopora damicornis varies in a manner that challenges classical models of the roles of sexual and asexual reproduction. On the geographically isolated Hawaiian reefs and high latitude reefs in Western Australia, P. damicornis populations are highly clonal although it has been argued that sexual reproduction via broadcast spawning generates widely dispersed colonists. In contrast, on eastern Australia's tropical Great Barrier Reef populations show little evidence of clonality. Here, we compare the genotypic diversity of adult and juvenile colonies of P. damicornis at seven sites on eastern Australia's high latitude Lord Howe Island reefs to determine if levels of clonality vary with habitat heterogeneity and age of colonies (as predicted by theory) or alternatively if clonality is again always high as for other isolated reef systems. We found 55-100% of the genotypic diversity expected for random mating at all seven sites and little evidence of asexual recruitment irrespective of habitat heterogeneity (sheltered versus wave exposed) or colony age. We found reduced levels of genetic diversity compared with tropical reefs (2.75 versus 4 alleles/locus), which supports earlier findings that Lord Howe Island is an isolated reef system. Furthermore, heterozygote deficits coupled with significant genetic subdivision among sites (FST=0.102+/-0.03) is typical of populations that have limited larval connections and are inbred. We conclude that the genetic structure of P. damicornis at Lord Howe Island reflects populations that are maintained through localised recruitment of sexually produced larvae.     

Limits to physiological plasticity of the coral Pocillopora verrucosa from the central Red Sea

Many coral species display changing distribution patterns across coral reef depths. While changes in the underwater light field and the ability to associate with different photosynthetic symbionts of the genus Symbiodinium explain some of the variation, the limits to physiological plasticity are unknown for most corals. In the central Red Sea, colonies of the branching coral Pocillopora verrucosa are most abundant in shallow high light environments and become less abundant in water depths below 10 m. To further understand what determines this narrow distribution, we conducted a cross-depths transplant experiment looking at physiological plasticity and acclimation in regard to depth. Colonies from 5, 10, and 20 m were collected, transplanted to all depths, and re-investigated after 30 and 210 d. All coral colonies transplanted downward from shallow to deep water displayed an increase in photosynthetic light-harvesting pigments, which resulted in higher photosynthetic efficiency. Shallow-water specimens transplanted to deeper water showed a significant decrease in total protein content after 30 and 210 d under low light conditions compared to specimens transplanted to shallow and medium depths. Stable isotope data suggest that heterotrophic input of carbon was not increased under low light, and consequently, decreasing protein levels were symptomatic of decreasing photosynthetic rates that could not be compensated for through higher light-harvesting efficiency. Our results provide insights into the physiological plasticity of P. verrucosa in changing light regimes and explain the observed depth distribution pattern. Despite its high abundance in shallow reef waters, P. verrucosa possesses limited heterotrophic acclimation potential, i.e., the ability to support its mainly photoautotrophic diet through heterotrophic feeding. We conclude that P. verrucosa might be a species vulnerable to sudden changes in underwater light fields resulting from processes such as increased turbidity caused by coastal development along the Saudi Arabian Red Sea coast.     

Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

Trade-offs between sexual and asexual reproduction in the genus Mimulus

Summary Six estimates of resource allocation to sexual reproduction (nectar concentration, nectar volume, sugar production, pollinator visitation rates, fruit-set, and seed production) and a single estimate of resource allocation to asexual reproduction (the rate of rooted-branch production) were examined for five species of Mimulus. There were significant interspecific differences for all paramaters. With the exception of nectar concentration, there was 1) a consistent positive correlation among the parameters measuring allocation to sexual reproduction, and 2) a significant negative correlation between parameters measuring allocation to sexual reproduction and the rate of rooted-branch production (asexual reproduction). The results indicate that Mimulus species which produce the highest volumes of nectar 1) receive the most pollinator visits, 2) have the highest fruit-set, 3) produce the most seeds, and 4) produce the fewest rooted branches.     

Dominance of Endozoicomonas bacteria throughout coral bleaching and mortality suggests structural inflexibility of the Pocillopora verrucosa microbiome

The importance of Symbiodinium algal endosymbionts and a diverse suite of bacteria for coral holobiont health and functioning are widely acknowledged. Yet, we know surprisingly little about microbial community dynamics and the stability of host‐microbe associations under adverse environmental conditions. To gain insight into the stability of coral host‐microbe associations and holobiont structure, we assessed changes in the community structure of Symbiodinium and bacteria associated with the coral Pocillopora verrucosa under excess organic nutrient conditions. Pocillopora‐associated microbial communities were monitored over 14 days in two independent experiments. We assessed the effect of excess dissolved organic nitrogen (DON) and excess dissolved organic carbon (DOC). Exposure to excess nutrients rapidly affected coral health, resulting in two distinct stress phenotypes: coral bleaching under excess DOC and severe tissue sloughing (>90% tissue loss resulting in host mortality) under excess DON. These phenotypes were accompanied by structural changes in the Symbiodinium community. In contrast, the associated bacterial community remained remarkably stable and was dominated by two Endozoicomonas phylotypes, comprising on average 90% of 16S rRNA gene sequences. This dominance of Endozoicomonas even under conditions of coral bleaching and mortality suggests the bacterial community of P. verrucosa may be rather inflexible and thereby unable to respond or acclimatize to rapid changes in the environment, contrary to what was previously observed in other corals. In this light, our results suggest that coral holobionts might occupy structural landscapes ranging from a highly flexible to a rather inflexible composition with consequences for their ability to respond to environmental change.     

Genetic variation in organisms with sexual and asexual reproduction

The genetic variation in a partially asexual organism is investigated by two models suited for different time scales. Only selectively neutral variation is considered. Model 1 shows, by the use of a coalescence argument, that three sexually derived individuals per generation are sufficient to give a population the same pattern of allelic variation as found in fully sexually reproducing organisms. With less than one sexual event every third generation, the characteristic pattern expected for asexual organisms appear, with strong allelic divergence between the gene copies in individuals. At intermediary levels of sexuality, a complex situation reigns. The pair-wise allelic divergence under partial sexuality exceeds, however, always the corresponding value under full sexuality. These results apply to large populations with stable reproductive systems. In a more general framework, Model 2 shows that a small number of sexual individuals per generation is sufficient to make an apparently asexual population highly genotypically variable. The time scale in terms of generations needed to produce this effect is given by the population size and the inverse of the rate of sexuality.     

Risky business: sexual and asexual reproduction in variable environments

Patterns of reproductive uncertainty can have an important influence on population dynamics. There is a crucial distinction between what we describe here as aggregate uncertainty (in which reproductive output in each generation is correlated among the individuals in a population) and idiosyncratic risk (in which reproductive output is independent across individuals). All else being equal, populations experiencing idiosyncratic risk enjoy a higher asymptotic growth rate than do those experiencing aggregate uncertainty. Therefore individuals in populations of the former type will have a competitive advantage over individuals in populations of the latter type. Applying this distinction to models of randomly fluctuating environments, we point out that genetic variation among offspring can serve to reduce aggregate uncertainty, transforming it into a more idiosyncratic form of risk. We show that this transformation underlies the dynamics observed in several previous models of the role of outcrossing in the evolution of sex.     

Recovery and succession in a multi-species tropical seagrass meadow following experimental disturbance: the role of sexual and asexual reproduction

Recolonisation and succession in a multi-species tropical seagrass meadow was examined by creating gaps (50×50 cm) in the meadow and manipulating the supply of sexual and asexual propagules. Measurements of leaf shoot density and estimates of above-ground biomass were conducted monthly to measure recovery of gaps between September 1995 and November 1997. Measurements of the seeds stored in the sediment (seed bank) and horizontal rhizome growth of colonising species were also conducted to determine their role in the recovery process.Asexual colonisation through horizontal rhizome growth from the surrounding meadow was the main mechanism for colonisation of gaps created in the meadow. The seed bank played no role in recolonisation of cleared plots. Total shoot density and above-ground biomass (all species pooled) of cleared plots recovered asexually to the level of the undisturbed controls in 10 and 7 months, respectively. There was some sexual recruitment into cleared plots where asexual colonisation was prevented but seagrass abundance (shoot density and biomass) did not reach the level of unmanipulated controls. Seagrass species did not appear to form seed banks despite some species being capable of producing long-lived seeds.The species composition of cleared plots remained different to the undisturbed controls throughout the 26-month experiment. Syringodium isoetifolium was a rapid asexual coloniser of disturbed plots and remained at higher abundances than in the control treatments for the duration of the study. S. isoetifolium had the fastest horizontal rhizome growth of species asexually colonising cleared plots (6.9 mm day⁻¹). Halophila ovalis was the most successful sexual coloniser but was displaced by asexually colonising species. H. ovalis was the only species observed to produce fruits during the study.Small disturbances in the meadow led to long-term (>2 years) changes in community composition. This study demonstrated that succession in tropical seagrass communities was not a deterministic process. Variations in recovery observed for different tropical seagrass communities highlighted the importance of understanding life history characteristics of species within individual communities to effectively predict their response to disturbance. A reproductive strategy involving clonal growth and production of long-lived, locally dispersed seeds is suggested which may provide an evolutionary advantage to plants growing in tropical environments subject to temporally unpredictable major disturbances such as cyclones.     

Existence of two sexual races in the planarian species switching between asexual and sexual reproduction

In certain planarian species that are able to switch between asexual and sexual reproduction, determining whether a sexual has the ability to switch to the asexual state is problematic, which renders the definition of sexuals controversial. We experimentally show the existence of two sexual races, acquired and innate, in the planarian Dugesia ryukyuensis. Acquired sexuals used in this study were experimentally switched from asexuals. Inbreeding of acquired sexuals produced both innate sexuals and asexuals, but inbreeding of innate sexuals produced innate sexuals only and no asexuals. Acquired sexuals, but not innate sexuals, were forced to become asexuals by ablation and regeneration (asexual induction). This suggests that acquired sexuals somehow retain asexual potential, while innate sexuals do not. We also found that acquired sexuals have the potential to develop hyperplastic and supernumerary ovaries, while innate sexuals do not. In this regard, acquired sexuals were more prolific than innate sexuals. The differences between acquired and innate sexuals will provide a structure for examining the mechanism underlying asexual and sexual reproduction in planarians.     

Character compatibility of molecular markers to distinguish asexual and sexual reproduction

Particularly in polyploids, the potential of the high variability of dominant markers such as random amplified polymorphic DNA fragments (RAPDs) and amplified fragment length polymorphisms (AFLPs) in population genetic studies and analysis of breeding systems is reduced due to their dominant nature. In contrast, the criterion of character compatibility is hindered neither by dominance nor by polyploidy as allelic interpretation is not necessary. Character compatibility, which can be used to detect events of genetic exchange (or recombination), is particularly informative if these events are expected to be rare such as in taxa with extensive vegetative reproduction or apomixis. Binary unordered characters such as presence and absence of anonymous DNA markers are incompatible if all four pairwise combinations of character states are present among the individuals studied. Because incompatible character state distributions defy any progenitor–derivative relationship among individuals, they provide strong evidence for genetic exchange. Both the absolute number of incompatible character combinations and the probability of compatibility can be used as a measure of incompatibility. Although these measures may not directly relate to the frequency of genetic exchange, they provide a useful tool to heuristically explore data sets. The most commonly used input for multivariate analyses and analysis of molecular variance in population genetic studies of (dis)similarity of marker distributions are amalgamates of mutation and recombination. Character compatibility can be used to complement these traditional methods of analysis. Advantages and disadvantages of character incompatibility relative to multilocus analysis of modes of reproduction and population genetics are demonstrated with data from RAPDs, isozymes, and restriction fragment length polymorphisms (RFLPs) of the nuclear ribosomal and chloroplast genome.     

The importance of sexual and asexual reproduction in the recent evolution of Allium vineale

In the weedy plant species Allium vineale (wild garlic), individuals may simultaneously produce sexually and asexually derived offspring, by seed and bulbils, respectively. In this study, genetic and genotypic diversity was determined in samples from 14 European A. vineale populations using nuclear (RAPD) and cytoplasmic (PCR-RFLP of cpDNA) markers to investigate the importance of the different reproductive modes. In the whole sample, 77 nuclear multilocus genotypes and four chloroplast haplotypes (chlorotypes) were found. Populations exhibited a high degree of subdivision for nuclear and cytoplasmic markers as estimated from hierarchical F-statistics; at the same time, identical chlorotypes could be found in populations separated by large distances. Genotypic diversity was significantly lower than expected under free recombination in almost all populations, indicating that recruitment into populations is mostly by asexually produced offspring. Nevertheless, within each chlorotype, the distribution of markers from pairs of nuclear loci was incompatible with a purely clonal structure, suggesting that many multilocus genotypes have originated by sexual recombination rather than by mutation within asexual lineages. It is argued that the weedy habit of A. vineale is likely to have favored bulbil reproduction, whereas sexually generated genotypes may have facilitated local adaptation during the species' expansion across Europe.     

Assessing the effects of iron enrichment across holobiont compartments reveals reduced microbial nitrogen fixation in the Red Sea coral Pocillopora verrucosa

The productivity of coral reefs in oligotrophic tropical waters is sustained by an efficient uptake and recycling of nutrients. In reef‐building corals, the engineers of these ecosystems, this nutrient recycling is facilitated by a constant exchange of nutrients between the animal host and endosymbiotic photosynthetic dinoflagellates (zooxanthellae), bacteria, and other microbes. Due to the complex interactions in this so‐called coral holobiont, it has proven difficult to understand the environmental limitations of productivity in corals. Among others, the micronutrient iron has been proposed to limit primary productivity due to its essential role in photosynthesis and bacterial processes. Here, we tested the effect of iron enrichment on the physiology of the coral Pocillopora verrucosa from the central Red Sea during a 12‐day experiment. Contrary to previous reports, we did not see an increase in zooxanthellae population density or gross photosynthesis. Conversely, respiration rates were significantly increased, and microbial nitrogen fixation was significantly decreased. Taken together, our data suggest that iron is not a limiting factor of primary productivity in Red Sea corals. Rather, increased metabolic demands in response to iron enrichment, as evidenced by increased respiration rates, may reduce carbon (i.e., energy) availability in the coral holobiont, resulting in reduced microbial nitrogen fixation. This decrease in nitrogen supply in turn may exacerbate the limitation of other nutrients, creating a negative feedback loop. Thereby, our results highlight that the effects of iron enrichment appear to be strongly dependent on local environmental conditions and ultimately may depend on the availability of other nutrients.     

Two patterns of energy allocation for growth,reproduction and lipid storage in the scleractinian coral Pocillopora damicornis

A population of the coral Pocillopora damicornis was examined for 12 months to investigate whether different reproductive activities in the corals affected patcerns of energy allocation. Growth, the amount of lipid in the tissue, and reproductive activity were measured simultaneously at fortnightly intervals during the summer months when production of planulae was expected, and monthly for the cooler period of the year. Two group of corals could be identified: those which produced brooded larvae and formed ova and sperm during gametogenesis, and those which did not brood larvae and produced only sperm. Although they exhibited no morphological differences, these two groups displayed different patterns of energy allocation to growth, reproduction and lipid storage. Corals which did not produce brooded larvae grew one and a half times faster and had more lipid for storage after reproduction than those which produced brooded larvae. Corals that brooded larvae had more structural lipid before release of larvae than those without brooded larvae, reflecting a greater volume of reproductive material rich in lipid. These results demonstrate a substantial cost of reproduction since corals which did not produce brooded larvae and ova put more resources into grwoth and storage of lipid.     

The effects of sexual and asexual reproduction on geographic variation in the sea anemone Actinia tenebrosa

Allelic and genotypic frequencies were determined for samples from 35 widely distributed Australasian colonies of Actinia tenebrosa and 2 South African colonies of A. equina. These data provided no evidence of gene flow between Australisian and South African Actinia colonies and indicated that there may be some restriction of gene flow between widely separated Australasian colonies.Both species are viviparous, and brooded A. tenebrosa are known to be produced asexually. The present data indicate that, within both species, almost all genotypic diversity is generated by sexual reproduction with recombination. Sexually produced juveniles appear to be widely dispersed and panmixis may occur over thousands of kilometres. However, successful sexual recruitment must be episodic or rare. Colonies on stable shores displayed relatively low levels of genotypic diversity, as compared with expectations for sexually reproducing populations, indicating strong local effects of asexual recruitment. Clonal genotypes may be spread over hundreds of metres of shore, but are typically restricted to discrete colonies. Asexual recruitment is highly localised and asexual dispersal appears to be limited by lengths of shore (500 m) which are unsuitable for colonization. Colonies on unstable shores are significantly more diverse genotypically and show little evidence of clonal proliferation.     

Variations in sexual and asexual reproduction of Scirpus mariqueter along an elevational gradient

In order to assess the importance of sexual and asexual reproduction during the life history of Scirpus mariqueter, its reproductive and growth characters were concurrently examined along an elevational gradient (from low elevation to high elevation). The proportions of flowering shoot and inflorescence mass, seed : flower ratio and seed weight were used to quantify the investment in sexual reproduction. The proportions of current-year shoot and rhizome mass were used to quantify the investment in asexual reproduction, and the proportion of corm mass was used for growth, respectively. It was found that vegetative propagation predominated at low elevation, whereas sexual reproduction predominated at high elevation; and that sexual reproduction increased with declining asexual reproduction along the gradient. The results suggest that asexual reproduction is relatively favored in the early life stage, whereas sexual reproduction is favored when the population becomes mature and aged, probably because of the functional differentiation between the two reproductive types. Sexual productive characters (i.e. the proportions of flowering shoot and inflorescence mass) were negatively correlated to both growth and asexual reproductive characters along the gradient, indicating there might exist some trade-offs among growth, sexual and asexual reproduction during the life history. However, no obvious pattern was found between asexual reproductive characters and growth characters along the elevational gradient, possibly because of the varied relationships between them at different life stages. The variations in sexual and asexual reproduction in the species and the relationship between them are thought to be of great significance for local population growth, species persistence and evolution.     

The cost of switching from asexual to sexual reproduction in an aphid

The fecundity of asexual and sexual morphs of Sitobion avenae (F.) is compared. The cost of sex is quantified in terms of the reduction in numbers of offspring produced. The cost is much greater than the halving of fecundity that is usually assumed to be associated with the production of males.

---

Résumé Les fécondités des virginipares ailés, des gynopares ailés et des ovipares d'un même clone de Sitobion avenae ont été déterminées à 12°C. Bien que les types qui ayant donné naissance à des sexués aient eu une fécondité initiale plus élevée que celle de la lignée asexuée, celle-ci a été significativement plus féconde après 15 jours de reproduction.Les résultats du laboratoire ont été utilisés pour évaluer la fécondité de lignées sexuées et asexuées pendant quatre générations à 12°C. La reproduction parthénogénétique a présenté un avantage numérique sur la reproductions sexuée dans le rapport 320/1. La reproduction sexuée a été accompagnée d'une diminution significative de la fécondité potentielle. Bien que la reproduction sexuée de S. avenae soit avantageuse dans les régions où les hivers sont durs, parce que les oeufs sont résistants au froid, les lignées asexuées présentent un avantage considérable en ce qui concerne la fécondité potentielle pour l'augmentation de la population en automne et elles conservent de plus la capacité d'un accroissement numérique ultérieur si l'hiver est doux.

     

The balance between sexual and asexual reproduction in plants living in variable environments

The balance between sexual and asexual propagule production is studied in an evolutionary model where plants produce the two kinds of propagules in genetically determined proportions. The male function of plants producing asexual propagules can be varied, and the sexual and asexual propagules carry different probabilities to turn into new reproductive individuals. These fitnesses may vary over years. The evolution of the population’s reproductive system is studied assuming modifier alleles with small effects. In this setting a balanced, mixed reproductive system can evolve, but only if the difference in fitness between the sexual and asexual propagules varies over years. When the two kinds of propagules are very similar to each other, as is often the case with sexual and asexual seed formation, evolution will tend towards a state dominated by the one or the other reproductive system.