Transcriptional profiles in Strongyloides stercoralis males reveal deviations from the Caenorhabditis sex determination model.

The human and canine parasitic nematode Strongyloides stercoralis utilizes an XX/XO sex determination system, with parasitic females reproducing by mitotic parthenogenesis and free-living males and females reproducing sexually. However, the genes controlling S. stercoralis sex determination and male development are unknown. We observed precocious development of rhabditiform males in permissive hosts treated with corticosteroids, suggesting that steroid hormones can regulate male development. To examine differences in transcript abundance between free-living adult males and other developmental stages, we utilized RNA-Seq. We found two clusters of S. stercoralis-specific genes encoding predicted transmembrane proteins that are only expressed in free-living males. We additionally identified homologs of several genes important for sex determination in Caenorhabditis species, including mab-3, tra-1, fem-2, and sex-1, which may have similar functions. However, we identified three paralogs of gld-1; Ss-qki-1 transcripts were highly abundant in adult males, while Ss-qki-2 and Ss-qki-3 transcripts were highly abundant in adult females. We also identified paralogs of pumilio domain-containing proteins with sex-specific transcripts. Intriguingly, her-1 appears to have been lost in several parasite lineages, and we were unable to identify homologs of tra-2 outside of Caenorhabditis species. Together, our data suggest that different mechanisms control male development in S. stercoralis and Caenorhabditis species.

The impact of asexual and sexual reproduction in spatial genetic structure within and between populations of the dioecious plant Marchantia inflexa (Marchantiaceae).

Background and Aims: In dioecious plants, sexual reproduction requires close proximity to potential mates, but clonal growth can increase this distance and, therefore, reduce the probability of mating. Reduction in sexual propagules can lead to decreased dispersal and gene flow between populations. Gene flow and clonal growth may be further influenced by the size of the habitat patch. The effects of habitat size and reproductive mode (sexual or asexual reproduction) on spatial genetic structure and segregation of the sexes were tested by quantifying the distributions of genotypes and the sexes using the dioecious liverwort Marchantia inflexa. Methods: Plants were sampled from five pairs of small-large habitat patches to identify within- and among-population spatial genetic structure using 12 microsatellite markers. Spatial distributions were calculated as the likelihood that pairs of individuals were the same sex or genotype, and it was determined how that likelihood was affected by habitat patch size (small/large). Key Results: Asexual reproduction dominates within populations, and asexual dispersal also occurred across populations. Spatial segregation of the sexes was observed within populations; males were more likely to be near individuals of the same sex than were females. Although the likelihood of both sexes being near members of the same sex was similarly greater on small habitat patches, on large habitat patches male genotypes were almost 15 % more likely to be near clonemates than were female genotypes. Conclusions: The results show a sex difference in clonal clumping that was dependent upon habitat size, suggesting differential colonization and/or survival between males and females. The sexes and genotypes being structured differently within and among populations have implications for the persistence of populations and the interactions between them. This study demonstrates that studying only the sexes and not their genotypes (or vice versa) can limit our understanding of the extent to which reproductive modes (sexual or asexual) influence genetic structure both within and between populations.

Moving forward in circles: challenges and opportunities in modelling population cycles.

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer-resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.

The Effects of Plant Compensatory Regrowth and Induced Resistance on Herbivore Population Dynamics.

Outbreaks of herbivorous insects are detrimental to natural and agricultural systems, but the mechanisms driving outbreaks are not well understood. Plant responses to herbivory have the potential to produce outbreaks, but long-term effects of plant responses on herbivore dynamics are understudied. To quantify these effects, we analyze mathematical models of univoltine herbivores consuming annual plants with two responses: (1) compensatory regrowth, which affects herbivore survival in food-limited situations by increasing the amount of food available to the herbivore; and (2) induced resistance, which reduces herbivore survival proportional to the strength of the response. Compensatory regrowth includes tolerance, where plants replace some or all of the consumed biomass, and overcompensation, where plants produce more biomass than was consumed. We found that overcompensation can cause bounded fluctuations in the herbivore density (called outbreaks here) by itself, whereas neither tolerance nor induced resistance can cause an outbreak on its own. Food limitation and induced resistance can also drive outbreaks when they act simultaneously. Tolerance damps these outbreaks, but overcompensation, by contrast, qualitatively changes the conditions under which the outbreaks occur. Not properly accounting for these interactions may explain why it has been difficult to document plant-driven insect outbreaks and could undermine efforts to control herbivore populations in agricultural systems.

The dispersal process of asexual propagules and the contribution to population persistence in Marchantia (Marchantiaceae).

PREMISE OF THE STUDY: The dispersal process involves emigration from a focal source, dispersal through the landscape, and immigration into a new population or habitat. Despite the fact that dispersal is vital for the long-term persistence of a species, key stages of the process are unknown or understudied for many species, including the importance and contribution of asexual reproduction. Focusing only on a single stage in the dispersal process may give an incomplete and potentially flawed picture of the effects of asexual reproduction on metapopulation dynamics in plant species. METHODS: Using a multifaceted approach that combines laboratory experiments, field studies, and mathematical models, we quantify the production, dispersal, and survival of immigrants of water-dispersed asexual offspring (gemmae) of the clonal liverwort Marchantia inflexa. KEY RESULTS: Compared to female plants, male plants of Marchantia inflexa produce gemmae more quickly and in higher numbers, but due to desiccation have lower gemmae survival rates. Gemmae move up to 20 cm per minute in light rain, suggesting they can leave the source population. Long distance dispersal of gemmae is supported by the mathematical analysis of unisexual metapopulations. Upon reaching the new habitat, gemmae survival is high if they stay moist. CONCLUSIONS: By integrating multiple experiments to quantify the effects of gemmae on metapopulation dynamics, we found that different stages of dispersal can lead to different conclusions on which sex has an advantage. Gemmae are critical for the maintenance of both sexes, the persistence of single-sex metapopulations and species, and the invasibility of clonal organisms.

Sex-specific plant responses to light intensity and canopy openness: implications for spatial segregation of the sexes.

In seed plants, the proximate causes of spatial segregation of the sexes (SSS) and its association with environmental variation are thought to be linked to sex-specific morphological and physiological variation. To address the general question of linkage among SSS, plant traits and environmental gradients, Marchantia inflexa was used, for which male plants are found under more open tree canopy than females. We hypothesized that males are adapted to higher light intensity and are better able to tolerate water stress than females, as is the case with seed plants. We tested for sex-specific habitat and trait relationships by quantifying plant traits (morphological and physiological) and estimates of the light conditions (percent canopy openness and light intensity) in the field. Using path analysis, we found that edge pore density in both sexes was negatively correlated with canopy openness, while in males, edge pore density had a weak but positive relationship to light intensity. These responses suggest that canopy openness and light intensity have opposing effects on edge pore density in males and that males might be more responsive to water stress than females. Additionally, the greater importance of female support tissue, which functions as storage, in explaining and being explained by other variables in the path analysis, relative to male support tissue, may reflect sex-specific allocation differences related to resources needed for female function.

Overgrowth competition, fragmentation and sex-ratio dynamics: a spatially explicit, sub-individual-based model.

Sessile organisms that compete for access to resources by overgrowing each other may risk the local elimination of one sex or the other, as frequently happens within clumps of the dioecious liverwort Marchantia inflexa. A multi-stage, spatially implicit differential-equation model of M. inflexa growing in an isolated patch, analysed in a previous study, indicated that long-term coexistence of the sexes within such patches may be only temporary. Here we derive a spatially explicit, sub-individual-based model to reconsider this interpretation when much more ecological realism is taken into account, including the process of fragmentation. The model tracks temporally discrete growth increments in continuous space, representing growth architecture and the overgrowth process in significant geometric detail. Results remain generally consistent with the absence of long-term coexistence of the sexes in individual patches of Marchantia. Dynamics of sex-specific growth qualitatively resemble those generated by differential-equation models, suggesting that this much simpler framework may be adequate for multi-patch metapopulation models. Direct competition between fragmenting and non-fragmenting clones demonstrates the importance of fragmentation in overgrowth competition. The results emphasize the need for empirical work on mechanisms of overgrowth and for modeling and empirical studies of life history tradeoffs and sex-ratio dynamics in multi-patch systems.