Biology subjects

Ramm, S. A.

Publications and source records attributed to Ramm, S. A..

5 recordsLinked to original sources

bioRxiv2026

Purging of inbreeding depression does not eliminate environmental variation in reproductive onset

Many simultaneous hermaphrodites use selfing for reproductive assurance only when outcrossing opportunities are limited, owing to inbreeding depression in selfed progeny. However, scenarios that enforce substantial selfing (such as during recolonisation) can rapidly select for a high selfing propensity, a shift in mating system that is expected to eliminate both inbreeding depression and the delayed reproductive onset under selfing that is typically associated with it. We tested these predictions in the flatworm Macrostomum hystrix, using a line derived from an outcrossing population that had been subjected to enforced selfing for multiple generations followed by several years of relaxed selection. As predicted, isolated individual forced to self and individuals with constant partner access (i.e. outcrossing opportunities) did not differ in reproductive onset nor in inbreeding depression estimated through offspring survival. However, a third treatment group that provided intermittent partner access (to allow outcrossing but minimise potential competition effects) showed a different pattern: no inbreeding depression in offspring but a substantially accelerated reproductive onset. Whilst our results thus support the effective purging of inbreeding depression and increased selfing propensity under enforced selfing, we suggest that cues of an unstable social or physical environment nevertheless exert a major influence on reproductive timing.

evolutionary biology↗

bioRxiv2022

Rapid evolution of sex role specialization in a hermaphrodite under sex-limited selection

The evolution of separate sexes from hermaphroditism is thought to have occurred independently many times, and is linked to the evolution of sex chromosomes. Even though we have a good understanding of the theoretical steps in the evolution of sex chromosomes from a hermaphrodite ancestor, the initial stages are still hard to study because many sex chromosome systems are old. We addressed this problem by experimentally selecting a hermaphrodite via sex-limited experimental evolution for several generations, simulating the early stages in the evolution of a sex chromosome. More specifically, we used a GFP (green fluorescent protein) marker as a proxy for a sex-determining locus, and selected replicate populations of the simultaneously hermaphroditic flatworm Macrostomum lignano for fitness via the male sex role, female sex role, or both (i.e. a control). After 14 generations, a fitness assay revealed clear evidence for incipient sex role specialization, presumably reflecting the release from constraints usually imposed by selection on the other sex role. Importantly, however, this was not simply explained by differential sex allocation in the different selection regimes - insofar as morphological traits reflect the underlying trade-off over resource allocation to the male and female sex functions - because testis and ovary sizes did not diverge among treatments. Our study shows that sex role specialization can occur rapidly as a result of sex-limited selection, which is consistent with genetic constraints between sex-roles, and in line with the first predicted steps towards the evolution of a new sex chromosome system.

evolutionary biology↗

bioRxiv2020

Waiting for love but not forever: modelling the evolution of waiting time to selfing in hermaphrodites

Although mixed mating systems involving both selfing and outcrossing are fairly common in hermaphrodites, it is unclear how they are maintained. In some species, individuals delay self-fertilization while they have not found a mating partner. The waiting time is subject to two opposing selection pressures: waiting helps to avoid inbreeding depression in offspring by increasing the density-dependent probability to encounter a mate, but also increases the risk of dying before reproduction. In some species waiting time can vary between individuals and be heritable. We therefore used an individual-based model to explore how delayed selfing evolves in response to density and density fluctuations. We find that at high density, when individuals meet often, drift drives waiting time; at intermediate densities, strong inbreeding depression causes waiting time to increase; and at low densities, inbreeding depression is purged, and waiting time approaches zero. Positive feedback loops drive the system to either complete selfing or complete outcrossing. Fluctuating density can slow down convergence to these alternative stable states. However, mixed mating, in the sense of either a stable polymorphism in waiting times, or stable intermediate times, was never observed. Thus, additional factors need to be explored to explain the persistence of delayed selfing.

evolutionary biology↗

bioRxiv2019

Effects of two seminal fluid proteins on post-mating behavior in the simultaneously hermaphroditic flatworm Macrostomum lignano

Along with sperm, in many taxa male ejaculates also contain a complex mixture of proteins, peptides and other substances found in seminal fluid. Once seminal fluid proteins (SFPs) are transferred to the mating partner, they play crucial roles in mediating post-mating sexual selection, since they can modulate the partners behavior and physiology in ways that influence the reproductive success of both partners. One way in which sperm donors can maximize their own reproductive success is by changing the partners (sperm recipients) postcopulatory behavior to prevent or delay re-mating, thereby decreasing the likelihood or intensity of sperm competition. We therefore adopted a quantitative genetic approach combining gene expression and behavioral data to identify candidates that could mediate such a response in the simultaneously hermaphroditic flatworm Macrostomum lignano. We identified two putative SFPs - Mlig-pro46 and Mlig-pro63 - that exhibit a negative genetic correlation between transcript expression and mating frequency. Importantly, however, in one of the two different group sizes, differing in their sperm competition level, in which we measured genetic correlations, these same two transcripts are also linked to a second post-mating behavior in M. lignano, namely the suck behavior of recipients in which, upon ejaculate receipt, the worm places its pharynx over its female genital opening and appears to attempt to remove ejaculate components. To therefore investigate directly whether these two candidates manipulate partner behavior, and test whether this impacts on competitive fertilization success, we performed a manipulative experiment using RNA interference-induced knockdown to ask how loss of Mlig-pro46 and Mlig-pro63 expression, singly and in combination, affects mating frequency, partner suck propensity and both defensive and offensive sperm competitive ability (P1 and P2, respectively). None of the knock-down treatments impacted strongly on mating frequency or sperm competitive ability, but the knock-down of Mlig-pro63 resulted in a significantly decreased suck propensity of mating partners. This suggests that Mlig-pro63 may normally act as a cue in the ejaculate to trigger recipient suck behavior, though the functional and adaptive significance of these two seminal proteins from a donor perspective remains enigmatic.

evolutionary biology↗

bioRxiv2019

Genotype-by-environment interactions for seminal fluid expression and sperm competitive ability

Sperm competition commonly occurs whenever females mate multiply, leading to variation in male paternity success. This can be due to variation in the various traits that might affect sperm competitive ability, which itself depends on both genetic and environmental factors, as well as on genotype-by-environment interactions (GEI). Seminal fluid is a major component of the male ejaculate that is often expected to mediate sperm competition, where different genotypes can differ in their seminal fluid expression as a response to different level of sperm competition (i.e., exhibit GEI). We therefore here focussed on testing for GEI in expression of two recently identified seminal fluid transcripts, suckless-1 and suckless-2, which potentially modulate sperm competitive ability in the simultaneously hermaphroditic flatworm Macrostomum lignano via their effects on manipulating post-mating partner behaviour and ultimately the fate of transferred ejaculates. In addition, we sought to test for GEI in sperm competitive ability, to investigate the relationship between natural variation in the expression of these seminal fluid transcripts generated through GEI and relative paternity success. To do so, we manipulated social group size, which has been shown to successfully alter sperm competition level in M. lignano, in a set of inbred lines (genotypes) and then measured both the expression level of suckless-1 and suckless-2 in focal worms together with their relative paternity success in a standardised sperm competition (P1 & P2) assay. We found GEI for the expression level of suckless-1 and suckless-2, as well as for sperm competitive ability. Moreover, we found a positive relation between the expression of suckless-1 and paternity success. This suggests that natural variation in the expression of this seminal fluid transcript indeed can influence sperm competition outcomes in M. lignano.

evolutionary biology↗