SummaryPlants adapt phenotypically to different conditions of light and nutrient supply, supposedly in order to achieve colimitation of these resources. Their key variable of adjustment is the ratio of leaf area to root length, which relies on plant biomass allocation and organ morphology.We recorded phenotypic differences in leaf and root mass fractions (LMF, RMF), specific leaf area (SLA) and specific root length (SRL) of 12 herbaceous species grown in factorial combinations of high/low irradiance and fertilization treatments.Leaf area and root length ratios, and their components, were influenced by nonadditive effects between light and nutrient supply, and differences in the strength of plant responses were partly explained by Ellenberg's species values representing ecological optima. Changes in allocation were critical in plant responses to nutrient availability, as the RMF contribution to changes in root length was 2.59 that of the SRL. Contrastingly, morphological adjustments (SLA rather than LMF) made up the bulk of plant response to light availability.Our results suggest largely predictable differences in responses of species and groups of species to environmental change. Nevertheless, they stress the critical need to account for adjustments in below-ground mass allocation to understand the assembly and responses of communities in changing environments.
Mating partners often have conflicting interests when copulating. One of the major agents affecting female mating partners is seminal fluid, transferred along with sperm. The role of seminal fluid proteins (SFPs) in reproductive success is well studied in separate-sexed animals but is much less so in simultaneous hermaphrodites. The latter potentially have a unique target to exploit for the sperm donor's own benefit: the male function of their mating partners. Here we show that, in the great pond snail Lymnaea stagnalis, receipt of specific SFPs reduces both sperm transfer and paternity success in a subsequent insemination event. Lowering investment in the mating partner's male function constitutes a novel role for SFPs. This demonstrates for the first time that hermaphrodites alter their mates' male as well as female reproductive output. Although it remains to be tested whether this represents mate manipulation or an adaptive response of recipients, our findings identify male investment as a new target for postcopulatory sexual selection.
Lay summaryQuite a few animals are male and female at the same time, so they can choose to mate either as male or female on copulation. The decision to perform either sex role was known to be highly flexible depending on various, but often confounding, factors. For the pond snail, we report that young and small snails tend to mate as males first, though old and large snails do not seem to be better females.
In internally fertilizing animals, seminal fluid is usually added to the spermatozoa, together forming the semen or ejaculate. Besides nourishing and activating sperm, the components in the seminal fluid can also influence female physiology to augment fertilization success of the sperm donor. While many studies have reported such effects in species with separate sexes, few studies have addressed this in simultaneously hermaphroditic animals. This video protocol presents a method to study effects of seminal fluid in gastropods, using a simultaneously hermaphroditic freshwater snail, the great pond snail Lymnaea stagnalis, as model organism. While the procedure is shown using complete prostate gland extracts, individual components (i.e., proteins, peptides, and other compounds) of the seminal fluid can be tested in the same way. Effects of the receipt of ejaculate components on egg laying can be quantified in terms of frequency of egg laying and more subtle estimates of female reproductive performance such as egg numbers within each egg masses. Results show that seminal fluid proteins affect female reproductive output in this simultaneous hermaphrodite, highlighting their importance for sexual selection.
Sexual conflicts often arise between mating partners because each sex tries to maximize its own reproductive success. One major male strategy to influence a partner's resource allocation is the transfer of accessory gland proteins. This has been shown to occur in simultaneous hermaphrodites as well as in organisms with separate sexes. Although accessory gland proteins affect the investment of resources in both male and female function, we here specifically focus on female investment. In the great pond snail, Lymnaea stagnalis, previous studies found that the accessory gland protein ovipostatin reduced female fecundity by suppressing egg laying in the partner in the short term (days). To investigate whether this reduction in egg laying is a commonly found effect of mating in freshwater snails, we compared egg output for evidence of suppression in isolated and paired snails of eight pulmonate species. Furthermore, we determined whether the suppression of egg laying caused a shift in resource allocation to the eggs. We found that in five of the eight species egg laying was suppressed, with fewer and lighter egg masses being laid when they had access to a mating partner. In mated pairs of L. stagnalis and Biomphalaria alexandrina, allocation of resources to the eggs was altered in opposite ways: individuals of L. stagnalis laid fewer but larger and heavier eggs; individuals of B. alexandrina laid smaller and lighter eggs, with no change in egg numbers. Such changes in the female function are most likely the result of combined effects of receiving accessory gland proteins, and the cost of mating in both male and female roles. Thus, effects of the maternal environment, including the receipt of accessory gland proteins, on offspring investment are not restricted to species with separate sexes.
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