Ploidy as a leaky reproductive barrier: mechanisms, rates and evolutionary significance of interploidy gene flow.

BACKGROUND: Whole genome duplication (polyploidization) is a dominant force in sympatric speciation, particularly in plants. Genome doubling instantly poses a barrier to gene flow owing to the strong crossing incompatibilities between individuals differing in ploidy. The strength of the barrier, however, varies from species to species and recent genetic investigations revealed cases of rampant interploidy introgression in multiple ploidy-variable species. SCOPE: Here, we review novel insights into the frequency of interploidy gene flow in natural systems and summarize the underlying mechanisms promoting interploidy gene flow. Field surveys, occasionally complemented by crossing experiments, suggest frequent opportunities for interploidy gene flow, particularly in the direction from diploid to tetraploid, and between (higher) polyploids. However, a scarcity of accompanying population genetic evidence and a virtual lack of integration of these approaches leave the underlying mechanisms and levels of realized interploidy gene flow in nature largely unknown. Finally, we discuss potential consequences of interploidy genome permeability on polyploid speciation and adaptation and highlight novel avenues that have just recently been opened by the very first genomic studies of ploidy-variable species. Standing in stark contrast with rapidly accumulating evidence for evolutionary importance of homoploid introgression, similar cases in ploidy-variable systems are yet to be documented. CONCLUSIONS: The genomics era provides novel opportunity to re-evaluate the role of interploidy introgression in speciation and adaptation. To achieve this goal, interdisciplinary studies bordering ecology and population genetics and genomics are needed.

Mate-choice for close kin is associated with improved offspring survival in Lodoicea maldivica, the largest-seeded plant in the world.

We studied spatial patterns of kinship in the offspring of the endangered Lodoicea maldivica, a dioecious palm that produces the largest seed of any plant. Previous research has suggested that restricted seed and pollen dispersal in populations resulted in strong spatial genetic structure. We used microsatellites to genotype young plants and their potential parents at four sites across the species' entire natural range. We determined the most likely parents of each young plant based on the spatial separation of each parent pair, their genetic relatedness, and the level of correlated paternity. We identified both parents (43 female, 54 male) for 139 of 493 young plants. Mean distance between parental pairs was 26.8 m. Correlated paternity was low (0.168), indicating that mother trees were often pollinated by several fathers. Parental pairs were more closely related than expected by chance, suggesting outbreeding depression. Our results highlight the apparent strong mate choice for close kin in parent pairs of surviving offspring. We discuss the alternative biological processes that could lead to this, including the potential for break-up of favourable allelic combinations necessary for the development of the palm's very large seed. Management implications include germinating seeds where they naturally fall, using a diverse range of male plants as pollen donors for hand pollination, and protecting the native community of gecko pollinators.

I see you, you see me: the impact of social presence on social interaction processes in autistic and non-autistic people.

Environments that require social interaction are complex, challenging and sometimes experienced as overwhelming by autistic people. However, all too often theories relating to social interaction processes are created, and interventions are proposed, on the basis of data collected from studies that do not involve genuine social encounters nor do they consider the perception of social presence to be a potentially influential factor. In this review, we begin by considering why face-to-face interaction research is important in this field. We then discuss how the perception of social agency and social presence can influence conclusions about social interaction processes. We then outline some insights gained from face-to-face interaction research conducted with both autistic and non-autistic people. We finish by considering the impact of social presence on cognitive processes more broadly, including theory of mind. Overall, we demonstrate that choice of stimuli in studies assessing social interaction processes has the potential to substantially alter conclusions drawn. Ecological validity matters and social presence, in particular, is a critical factor that fundamentally impacts social interaction processes in both autistic and non-autistic people. This article is part of a discussion meeting issue 'Face2face: advancing the science of social interaction'.

Gaze cueing, mental States, and the effect of autistic traits.

The ability to interpret and follow the gaze of our social partners is an integral skill in human communication. Recent research has demonstrated that gaze following behaviour is influenced by theory of mind (ToM) processes. However, it has yet to be determined whether the modulation of gaze cueing by ToM is affected by individual differences, such as autistic traits. The aim of this experiment was to establish whether autistic traits in neurotypical populations affect the mediation of gaze cueing by ToM processes. This study used a gaze cueing paradigm within a change detection task. Participants' perception of a gaze cue was manipulated such that they only believed the cue to be able to 'see' in one condition. The results revealed that participants in the Low Autistic Traits group were significantly influenced by the mental state of the gaze cue and were more accurate on valid trials when they believed the cue could 'see'. By contrast, participants in the High Autistic Traits group were also more accurate on valid trials, but this was not influenced by the mental state of the gaze cue. This study therefore provides evidence that autistic traits influence the extent to which mental state attributions modulate social attention in neurotypical adults.

The Effect of Social Presence on Mentalizing Behavior.

Our behavior is frequently influenced by those around us. However, the majority of social cognition research is conducted using socially isolated paradigms, without the presence of real people (i.e., without a "social presence"). The current study aimed to test the influence of social presence upon a measure of mentalizing behavior in adults. Study 1 used a first-order theory of mind task; and study 2 used a second-order theory of mind task. Both studies included two conditions: live, where the task protagonists were physically present acting out the task, or recorded, where the same task protagonists demonstrated the task in a video recording. In both experiments, participants were affected by the social presence and demonstrated significantly different patterns of behavior in response to the presence of real people. This study, therefore, highlights the critical importance of understanding the effect of a social presence in mentalizing research, and suggests that the inclusion of a social presence needs to be given strong consideration across social cognition paradigms.

Disentangling the components of triploid block and its fitness consequences in natural diploid-tetraploid contact zones of Arabidopsis arenosa.

Hybrid seed inviability (HSI) is an important mechanism of reproductive isolation and speciation. HSI varies in strength among populations of diploid species but it remains to be tested whether similar processes affect natural variation in HSI within ploidy-variable species (triploid block). Here we used extensive endosperm, seed and F1 -hybrid phenotyping to explore HSI variation within a diploid-autotetraploid species. By leveraging 12 population pairs from three ploidy contact zones, we tested for the effect of interploidy crossing direction (parent of origin), ploidy divergence and spatial arrangement in shaping reproductive barriers in a naturally relevant context. We detected strong parent-of-origin effects on endosperm development, F1 germination and survival, which was also reflected in the rates of triploid formation in the field. Endosperm cellularization failure was least severe and F1 -hybrid performance was slightly better in the primary contact zone, with genetically closest diploid and tetraploid lineages. We demonstrated overall strong parent-of-origin effects on HSI in a ploidy variable species, which translate to fitness effects and contribute to interploidy reproductive isolation in a natural context. Subtle intraspecific variation in these traits suggests the fitness consequences of HSI are predominantly a constitutive property of the species regardless of the evolutionary background of its populations.

Sensitivity to Social Agency in Autistic Adults.

The presence of other people, whether real or implied, can have a profound impact on our behaviour. However, it is argued that autistic individuals show decreased interest in social phenomena, which leads to an absence of these effects. In this study, the agency of a cue was manipulated such that the cue was either described as representing a computer program or the eye movements of another participant. Both neurotypical and autistic participants demonstrated a social facilitation effect and were significantly more accurate on a prediction task when they believed the cue represented another participant. This demonstrates that whilst autistic adults may show difficulties in interpreting social behaviour this does not necessarily arise from a lack of sensitivity to social agency.

Niche similarity in diploid-autotetraploid contact zones of Arabidopsis arenosa across spatial scales.

PREMISE: Whole genome duplication is a major evolutionary event, but its role in ecological divergence remains equivocal. When populations of different ploidy (cytotypes) overlap in space, "contact zones" are formed, allowing the study of evolutionary mechanisms contributing toward ploidy divergence. Multiple contact zones per species' range are often described but rarely leveraged as natural replicates. We explored whether the strength of niche differentiation of diploid and autotetraploid Arabidopsis arenosa varies over distinct contact zones and if the frequency of triploids decreases from seedling to adult stage. METHODS: We characterized ploidy composition and habitat preferences in 264 populations across three contact zones using climatic niche modeling. Ecological differences of cytotypes were also assessed using local vegetation surveys at 110 populations within two contact zones, and at the finer scale within five mixed-ploidy sites. This was complemented by flow cytometry of seedlings. RESULTS: We found no niche differences between diploid and tetraploid populations within contact zones for either climatic or local environmental variables. Comparisons of cytotypes within mixed-ploidy sites found weak niche differences that were inconsistent in direction. Triploid individuals were virtually absent (0.14%) in the field, and they were at a similarly low frequency (0.2%) in ex situ germinated seedlings. CONCLUSIONS: This study demonstrates the strength in investigating different spatial scales across several contact zones when addressing ecological niche differentiation between ploidies. The lack of consistent habitat differentiation of ploidies across the scales and locations supports the recently emerging picture that processes other than ecological differentiation may underlie ploidy coexistence in diploid-autopolyploid systems.

Identification of sex-linked markers in the sexually cryptic coco de mer: are males and females produced in equal proportions?

Lodoicea maldivica (coco de mer) is a long-lived dioecious palm in which male and female plants are visually indistinguishable when immature, only becoming sexually dimorphic as adults, which in natural forest can take as much as 50 years. Most adult populations in the Seychelles exhibit biased sex ratios, but it is unknown whether this is due to different proportions of male and female plants being produced or to differential mortality. In this study, we developed sex-linked markers in Lodoicea using ddRAD sequencing, enabling us to reliably determine the gender of immature individuals. We screened 589 immature individuals to explore sex ratios across life stages in Lodoicea. The two sex-specific markers resulted in the amplification of male-specific bands (Lm123977 at 405 bp and Lm435135 at 130 bp). Our study of four sub-populations of Lodoicea on the islands of Praslin and Curieuse revealed that the two sexes were produced in approximately equal numbers, with no significant deviation from a 1:1 ratio before the adult stage. We conclude that sex in Lodoicea is genetically determined, suggesting that Lodoicea has a chromosomal sex determination system in which males are the heterogametic sex (XY) and females are homogametic (XX). We discuss the potential causes for observed biased sex ratios in adult populations, and the implications of our results for the life history, ecology and conservation management of Lodoicea.

Mental State Attributions Mediate the Gaze Cueing Effect.

Understanding the mental states of our social partners allows us to successfully interact with the world around us. Mental state attributions are argued to underpin social attention, and have been shown to modulate attentional orienting to social cues. However, recent research has disputed this claim, arguing that this effect may arise as an unintentional side effect of study design, rather than through the involvement of mentalising processes. This study therefore aimed to establish whether the mediation of gaze cueing by mental state attributions generalises beyond the specific experimental paradigm used in previous research. The current study used a gaze cueing paradigm within a change detection task, and the gaze cue was manipulated such that participants were aware that the cue-agent was only able to 'see' in one condition. The results revealed that participants were influenced by the mental state of the cue-agent, and were significantly better at identifying if a change had occurred on valid trials when they believed the cue-agent could 'see'. The computation of the cue-agent's mental state therefore mediated the gaze cueing effect, demonstrating that the modulation of gaze cueing by mental state attributions generalises to other experimental paradigms.

Tracing coco de mer's reproductive history: Pollen and nutrient limitations reduce fecundity.

Habitat degradation can reduce or even prevent the reproduction of previously abundant plant species. To develop appropriate management strategies, we need to understand the reasons for reduced recruitment in degraded ecosystems. The dioecious coco de mer palm (Lodoicea maldivica) produces by far the largest seeds of any plant. It is a keystone species in an ancient palm forest that occurs only on two small islands in the Seychelles, yet contemporary rates of seed production are low, especially in fragmented populations. We developed a method to infer the recent reproductive history of female trees from morphological evidence present on their inflorescences. We then applied this method to investigate the effects of habitat disturbance and soil nutrient conditions on flower and fruit production. The 57 female trees in our sample showed a 19.5-fold variation in flower production among individuals over a seven-year period. Only 77.2% of trees bore developing fruits (or had recently shed fruits), with the number per tree ranging from zero to 43. Flower production was positively correlated with concentrations of available soil nitrogen and potassium and did not differ significantly between closed and degraded habitat. Fruiting success was positively correlated with pollen availability, as measured by numbers and distance of neighboring male trees. Fruit set was lower in degraded habitat than in closed forest, while the proportion of abnormal fruits that failed to develop was higher in degraded habitat. Seed size recorded for a large sample of seeds collected by forest wardens varied widely, with fresh weights ranging from 1 to 18 kg. Synthesis: Shortages of both nutrients and pollen appear to limit seed production of Lodoicea in its natural habitat, with these factors affecting different stages of the reproductive process. Flower production varies widely amongst trees, while seed production is especially low in degraded habitat. The size of seeds is also very variable. We discuss the implications of these findings for managing this ecologically and economically important species.

Development of 12 polymorphic microsatellite loci for the endangered Seychelles palm Lodoicea maldivica (Arecaceae).

PREMISE OF THE STUDY: The evolutionarily and ecologically distinct coco de mer palm Lodoicea maldivica (Arecaceae) is endemic to two islands in the Seychelles. Before colonization of the islands by man, the endangered palm formed large monodominant stands, but its natural range is now restricted to four main populations and several patches of isolated individuals. Microsatellite markers were designed to investigate the genetic structure of the remaining natural populations of L. maldivica. METHODS AND RESULTS: We developed 12 polymorphic and three monomorphic microsatellite markers for this species, with a mean number of alleles per locus of 13.2 (range 5-21) and expected heterozygosity values ranging from 0.31-0.91 for the polymorphic loci. CONCLUSIONS: These markers enable us to study the patterns of genetic diversity, contemporary seed dispersal, and the fine-scale spatial genetic structure of this important conservation flagship species.

The role of the oculomotor system in covert social attention.

Observing a change in gaze direction triggers a reflexive shift of attention and appears to engage the eye-movement system. However, the functional relationship between social attention and this oculomotor activation is unclear. One extremely influential hypothesis is that the preparation of a saccadic eye movement is necessary and sufficient for a covert, reflexive shift of attention (the premotor theory of attention; Rizzolatti et al., 1994). Surprisingly, this theory has not been directly tested with respect to reflexive gaze cueing. In order to address this issue, gaze cueing, peripheral cueing, and arrow cueing were examined under conditions in which some stimuli appeared at locations that could not become the goal of a saccadic eye movement. It was observed that peripheral cues failed to elicit reflexive attentional orienting when targets appeared beyond the range of eye movements. Similarly, nonpredictive arrow cues were ineffective when targets could not become the goal of a saccade. In contrast, significant gaze-cueing effects were still observed when targets were beyond the range of eye movements. These data demonstrate that the mechanisms involved in gaze cueing are dissociated from those involved in exogenous orienting to peripheral or arrow cues. Furthermore, the findings suggest that, unlike peripheral cueing and reflexive arrow cueing, gaze cueing is independent of oculomotor control. We conclude that the premotor theory does not offer a compelling explanation for gaze cueing.