Biology subjects

Gilbert, K. J.

Publications and source records attributed to Gilbert, K. J..

16 recordsLinked to original sources

bioRxiv2026

Shedding light on fenestrations in carnivorous pitcher plants: a test for convergent function

A longstanding question in evolution is if similar morphologies shared by distantly related organisms facing similar environmental pressures belie similar functions. The carnivorous pitcher form has evolved in three distinct plant lineages and encompasses a suite of convergent morphological traits, providing a natural experiment to test hypothesized links between morphology and function. Multiple pitcher plant species possess fenestrations (i.e., small windows) that let light pass through the pitcher leaf. While research on one species, Nepenthes aristolochioides, demonstrates that fenestrations manipulate light to create false exits and increase prey capture, whether this convergent morphology serves this convergent function in other pitcher species remains unresolved. We explored the function of fenestrations in two additional species of pitcher plants, Nepenthes klossii and Cephalotus follicularis, where the morphology and function of fenestration were previously unexplored. We inspected micromorphology of pitcher structure, conducted an insect bioassay with live plants to test for links between morphology and function, and quantified traits associated with light manipulation from herbarium specimens. Both species exhibited fenestration and related pitcher morphology similar to N. aristolochioides. Yet our experiments did not reveal a significant effect of light passing through fenestrations on fly capture for these two species. Key differences between these species and the previously investigated species may be the presence of wax crystals and the absence of viscoelastic fluid. While future studies may uncover weak to moderate light manipulation function not detected in our study, our data do not support convergent function in these distantly related, morphologically similar species.

evolutionary biology↗

bioRxiv2026

A continuum of CAM phenotypes in the carnivorous plant genus Pinguicula (Lentibulariaceae)

PremisePinguicula are carnivorous plants occupying a wide range of habitats, from wetlands to barren rock cliffs, where CAM photosynthesis was recently discovered in several species. This ecological and physiological diversity makes the genus a promising system for exploring the environmental drivers and evolutionary transitions underlying variation in CAM. Here, we present a physiological survey of CAM with a particular focus on the Mexican Clade, which contains the recently identified CAM species. MethodsWe conducted a carbon isotope survey of live and herbarium specimens to identify candidate CAM species. We then measured diurnal gas-exchange patterns and changes in leaf titratable acidity to quantify CAM activity. ResultsPinguicula species exhibited a variety of photosynthetic phenotypes. Pinguicula vulgaris showed no detectable evidence of CAM, whereas P. cyclosecta and P. moranensis demonstrated a C3-CAM physiology with most net assimilation via C3. Pinguicula martinezii also showed the same C3-CAM physiology under well-watered conditions but had strong facultative induction of CAM with drought. Pinguicula agnata displayed net assimilation via CAM. Titratable acidity had a positive correlation with {delta}13C within the species we sampled. ConclusionsOur results support the existence of a CAM physiological continuum within Pinguicula. Notably, closely related species encompass physiologies that span the large variation of CAM. This diversity provides a rare opportunity for future studies to investigate the evolution of CAM in closely related species, including differences and similarities between differing states across the continuum.

plant biology↗

bioRxiv2026

Host-mediated pH influences microbiome assembly and function on the phylloplane

Plant leaves harbor diverse microbial communities influenced by environmental inputs and host traits, yet it remains unclear whether leaves act as passive substrates or active ecological filters that reorganize microbial functional capacity. Phylloplane pH regulation is one hostplant trait that has been traditionally underexplored. We used metatranscriptomics to examine microbial gene expression on the phylloplane and within whole leaves of five plant species spanning the extremes of baseline phylloplane pH, including hyperalkalinizing Gossypium species, weakly buffering Beta vulgaris, and hyperacidifying Nepenthes species. Young leaves were inoculated with a common soil-derived microbial community to quantify host-associated restructuring of taxonomic and functional profiles, and short-term pH perturbations were applied to test the effect of transient abiotic stress. Across both phylloplane and whole-leaf datasets, host species identity was the primary axis structuring microbial taxonomic composition and expressed functional repertoires. Leaf-associated communities diverged from the source inoculum, but retained a substantial shared functional backbone enriched for central biosynthetic and core metabolic pathways. Host-associated differentiation reflected selective retention and redistribution of reactions within this shared environmental pool rather than acquisition of novel metabolic capacity. Enriched pathway subsets were metabolically coherent and taxonomically distributed across multiple bacterial orders, consistent with functional redundancy and trait-based assembly. Among hosts, Gossypium exhibited the strongest restructuring relative to inoculum, suggesting comparatively stronger host-associated filtering. In contrast, short-term pH manipulation did not induce consistent community-wide functional reorganization. Microbial physiological responses to the phylloplane environment and external pH were observed at the organismal level. Together, these results position leaves as active ecological filters that reorganize microbial functional landscapes through host-specific trait regimes. This work begins to implicate some role of phylloplane pH regulation in microbial assembly and function.

ecology↗

bioRxiv2026

Convergent evolution of red pigmentation in extrafloral nectaries: global patterns and mechanisms

Convergent traits are a signature of adaptation, where shared selective pressures may lead to the replicated evolution of traits with similar function. Here, we present an integrative investigation of a widespread yet understudied feature of an iconic, convergent mutualistic plant trait - red pigmentation in extrafloral nectaries (EFNs). We surveyed 625 species to describe the distribution of red coloration in EFNs across plant morphology, biogeography, and phylogeny for the first time and propose several adaptive hypotheses for the drivers of their convergence across plants. We then test several of these hypotheses by integrating our global-scale biogeographic and phylogenetic analysis with phylogenetically paired common garden bioassays and a museum collections survey. We find that across the plant phylogeny, cold wet regions and deserts are hotspots of red EFN coloration, and multiple lines of evidence point to a fungal defense role of red pigments in EFNs. Ultimately, our findings highlight red EFNs as a highly convergent phenotype across plants and point to distinct selective pressures underlying the distribution of EFN coloration and EFNs themselves.

evolutionary biology↗

bioRxiv2026

The Frequency of Population and Masting Fluctuations Is Increasing Globally

Substantial anthropogenic changes to the environment have motivated efforts to quantify temporal trends in population dynamics. While most ecological research has focused on the mean and variance of population density and reproduction, the frequency of these fluctuations through time may also be changing. We analyzed 1,563 datasets of population density and 1,456 datasets of plant reproduction (masting) across the globe. The average frequency of fluctuations increased by [~] 0.5 - 3% per decade within each time series, representing a moderate change (Cohens d {approx} 0.4) over a period of 60 years. We tested four hypothesized mediators of this trend: increased temperature, increased frequency of environmental forcing, increased intrinsic growth rate, and increased distance from a saddle at zero density. Although all hypotheses were rejected, changes in the frequency of environmental forcing and intrinsic growth rate exhibited positive correlations with changes in population fluctuation frequency as expected. Our results suggest that successive peaks in population and masting density fluctuations are becoming closer in time, which may reduce the effectiveness of predator satiation, resilience of food-webs, and the risk of critical transitions, such as population extinction. We suggest some alternative hypotheses for what may underlie this surprising global pattern.

ecology↗

bioRxiv2024

Species-specific phyllosphere responses to external pH change

The leaf surface, known as the phylloplane, represents the initial point of contact for plants in their interaction with the aboveground environment. Although prior research has assessed how leaves respond to external pH variations, particularly in the context of acid rain, there remains a limited understanding of the molecular mechanisms through which plants detect, respond to, and mitigate cellular damage. To look at plant responses to external pH changes, we measured the phylloplane pH for five species with variable phylloplane pH that ranged in the dry control. Moreover, we investigated the phylloplane pH in response to three pH treatments (pH 6.5, 4, and 2) and found that plants can modify their phylloplane pH, and this buffering ability is species-specific. Among the species analyzed, only Gossypium displayed a strong buffering ability. For treatments where leaves were exposed to either pH 6.5 or pH 4, Gossypium alkalinized the phylloplane pH slightly higher than the dry control pH. Remarkably, when leaves were exposed to pH 2, Gossypium was able to buffer the pH to 6 within five minutes. Furthermore, our transcriptional analysis indicated that the responses to external pH changes varied among species, highlighting differentially expressed genes associated with calcium (Ca2+) signaling pathways, as well as Ca2+ and H+-ATPases pumps. These findings also suggest that pH stress negatively impacts photosynthesis, and that both wetness and moderate pH shifts may trigger additional abiotic and biotic stress signaling pathways.

plant biology↗

bioRxiv2024

EdgeHOG: fine-grained ancestral gene order inference at tree-of-life scale

Ancestral genomes are essential for studying the diversification of life from the last universal common ancestor to modern organisms. Methods have been proposed to infer ancestral gene order, but they lack scalability, limiting the depth to which gene neighborhood evolution can be traced back. We introduce edgeHOG, a tool designed for accurate ancestral gene order inference with linear time complexity. Validated on various benchmarks, edgeHOG was applied to the entire OMA orthology database, encompassing 2,845 extant genomes across all domains of life. This represents the first tree-of-life scale inference, resulting in 1,133 ancestral genomes. In particular, we reconstructed ancestral contigs for the last common ancestor of eukaryotes, dating back around 1.8 billion years, and observed significant functional association among neighboring genes. The method also dates gene adjacencies, revealing conserved histone clusters and rapid sex chromosome rearrangements, enabling computational inference of these features.

evolutionary biology↗

bioRxiv2024

Using runs of homozygosity and machine learning to disentangle sources of inbreeding and infer self-fertilization rates

Runs of homozygosity (ROHs) are indicative of elevated homozygosity and inbreeding due to mating of closely related individuals. Self-fertilization can be a major source of inbreeding which elevates genomewide homozygosity and thus should also create long ROHs. While ROHs are frequently used to understand inbreeding in the context of conservation and selective breeding, as well as for consanguinity of populations and their demographic history, it remains unclear how ROH characteristics are altered by selfing and if this confounds expected signatures of inbreeding due to demographic change. Using simulations, we study the impact of the mode of reproduction and demographic history on ROHs. We apply random forests to identify unique characteristics of ROHs, indicative of different sources of inbreeding. We pinpoint distinct features of ROHs that can be used to better characterize the type of inbreeding the population was subjected to and to predict outcrossing rates and complex demographic histories. Using additional simulations and four empirical datasets, two from highly selfing species and two from mixedmaters, we predict the selfing rate and validate our estimations. We find that self-fertilization rates are successfully identified even with complex demography. Population genetic summary statistics improve algorithm accuracy particularly in the presence of additional inbreeding, e.g., from population bottlenecks. Our findings highlight the importance of ROHs in disentangling confounding factors related to various sources of inbreeding and demonstrate situations where such sources cannot be differentiated. Additionally, our random forest models provide a novel tool to the community for inferring selfing rates using genomic data.

genetics↗

bioRxiv2023

Subgenome dominance shapes novel gene evolution in the decaploid pitcher plant Nepenthes gracilis

Subgenome dominance after whole-genome duplication generates distinction in gene number and expression at the level of chromosome sets, but it remains unclear how this process may be involved in evolutionary novelty. Here, we generated a chromosome-scale genome assembly of the Asian pitcher plant Nepenthes gracilis to analyze how its novel traits (dioecy and carnivorous pitcher leaves) are linked to genomic evolution. We found a decaploid karyotype with five complete sets of syntenic chromosomes (2n = 10x = 80) yet with a clear indication of subgenome dominance and highly diploidized gene contents. The male-linked and pericentromerically located region on the putative sex chromosome was identified in a recessive subgenome and was found to harbor three transcription factors involved in flower and pollen development, including a likely neofunctionalized LEAFY duplicate. Transcriptomic and syntenic analyses of carnivory-related genes suggested that the paleopolyploidization events seeded genes that subsequently formed tandem clusters in recessive subgenomes with specific expression in the digestive zone of the pitcher, where specialized cells digest prey and absorb derived nutrients. Novel gene evolution in recessive subgenomes is likely to be prevalent because duplicates were enriched with Nepenthes-specific genes with tissue-specific expression, including those expressed in trapping pitchers. Thus, subgenome dominance likely contributed to evolutionary novelty by allowing recessive subgenomes experiencing relaxed purifying selection to serve as a preferred host of novel tissue-specific duplicates. Our results provide insight into how polyploids, which may frequently be evolutionary dead-ends, have given rise to novel traits in exceptionally thriving high-ploidy lineages.

plant biology↗

bioRxiv2022

Purging due to self-fertilization does not prevent accumulation of expansion load

Species range expansions are a common demographic history presenting populations with multiple evolutionary challenges. It is not yet fully understood if self-fertilization, which is often observed at species range edges, may create an evolutionary advantage against these challenges. Selfing provides reproductive reassurance to counter Allee effects and selfing may purge accumulated mutational burden due to founder events (expansion load) by further increasing homozygosity. We study how selfing impacts the accumulation of genetic load during range expansion via purging and/or speed of colonization. Using simulations, we disentangle inbreeding effects due to demography versus due to selfing and find that selfers expand faster, but still accumulate load, regardless of mating system. The severity of variants contributing to this load, however, differs across mating system: higher selfing rates purge large-effect recessive variants leaving a burden of smaller-effect alleles. We compare these predictions to the mixedmating plant Arabis alpina, using whole-genome sequences from refugial outcrossing populations versus expanded selfing populations. Empirical results indicate accumulation of expansion load along with evidence of purging in selfing populations, concordant with our simulations, and suggesting that while purging is a benefit of selfing evolving during range expansions, it is not sufficient to prevent load accumulation due to range expansion. Author SummaryThe geographic space that species occupy, i.e. the species range, is known to fluctuate over time due to changing environmental conditions. Since the most recent glaciation, many species have recolonized available habitat as the ice sheets melted, expanding their range. When populations at species range margins expand into newly available space, they suffer from an accumulation of deleterious alleles due to repeated founder effects. We study whether self-fertilization, which is considered an evolutionary deadend, can be favored under these expanding edge conditions. Selfing has two important effects: allowing for faster expansion due to reproductive assurance and purging recessive deleterious alleles by exposing them to selection as homozygotes. We use simulations to identify the impact of selfing on expanded populations and then compare these results to an empirical dataset to assess whether our predictions are met. We use the mixed-mating plant alpine rock-cress (Arabis alpina) since it has both expanded since the last glaciation and undergone a mating shift to selfing. We find that selfing does not prevent the accumulation of deleterious load, however purging does still act to remove the most severe variants, indicating that selfing provides this benefit during range expansions.

evolutionary biology↗

bioRxiv2022

Chromosomal inversions can limit adaptation to new environments

Chromosomal inversions are often thought to facilitate local adaptation and population divergence because they can link multiple adaptive alleles into non-recombining genomic blocks. Selection should thus be more efficient in driving inversion-linked adaptive alleles to high frequency in a population, particularly in the face of maladaptive gene flow. But, what if ecological conditions and hence selection on inversion-linked alleles change? Reduced recombination within inversions could then constrain the formation of optimal combinations of pre-existing alleles under these new ecological conditions. Here, we outline this idea of inversions limiting adaptation and divergence when ecological conditions change across time or space. We reason that the benefit of inversions for local adaptation and divergence under one set of ecological conditions can come with a concomitant constraint for adaptation to novel sets of ecological conditions. This limitation of inversions to adaptation may also provide one possible explanation for why inversions are often maintained as polymorphisms within species.

evolutionary biology↗

bioRxiv2021

Gene surfing of underdominant alleles promotes formation of hybrid zones

The distribution of genetic diversity over geographic space has long been investigated in population genetics and serves as a useful tool to understand evolution and history of populations. Within some species or across regions of contact between two species, there are instances where there is no apparent ecological determinant of sharp changes in allele frequencies or divergence. To further understand these patterns of spatial genetic structure and potential species divergence, we model the establishment of clines that occur due to the surfing of underdominant alleles during range expansions. We provide analytical approximations for the fixation probability of underdominant alleles at expansion fronts and demonstrate that gene surfing can lead to clines in 1D range expansions. We extend these results to multiple loci via a mixture of analytical theory and individual-based simulations. We study the interaction between the strength of selection against heterozygotes, migration rates, and local recombination rates on the formation of stable hybrid zones. A key result of our study is that clines created by surfing at different loci can attract each other and align after expansion, if they are sufficiently close in space and in terms of recombination distance. Our findings suggest that range expansions can set the stage for parapatric speciation due to the alignment of multiple selective clines, even in the absence of ecologically divergent selection.

evolutionary biology↗

bioRxiv2020

The distribution of mutational effects on fitness in Caenorhabditis elegans inferred from standing genetic variation

The distribution of fitness effects for new mutations is one of the most theoretically important but difficult to estimate properties in population genetics. A crucial challenge to inferring the distribution of fitness effects (DFE) from natural genetic variation is the sensitivity of the site frequency spectrum to factors like population size change, population substructure, and non-random mating. Although inference methods aim to control for population size changes, the influence of non-random mating remains incompletely understood, despite being a common feature of many species. We report the distribution of fitness effects estimated from 326 genomes of Caenorhabditis elegans, a nematode roundworm with a high rate of self-fertilization. We evaluate the robustness of DFE inferences using simulated data that mimics the genomic structure and reproductive life history of C. elegans. Our observations demonstrate how the combined influence of self-fertilization, genome structure, and natural selection can conspire to compromise estimates of the DFE from extant polymorphisms. These factors together tend to bias inferences towards weakly deleterious mutations, making it challenging to have full confidence in the inferred DFE of new mutations as deduced from standing genetic variation in species like C. elegans. Improved methods for inferring the distribution of fitness effects are needed to appropriately handle strong linked selection and selfing. These results highlight the importance of understanding the combined effects of processes that can bias our interpretations of evolution in natural populations.

evolutionary biology↗

bioRxiv2019

Transition from background selection to associative overdominance promotes diversity in regions of low recombination

Linked selection is a major driver of genetic diversity. Selection against deleterious mutations removes linked neutral diversity (background selection, BGS, Charlesworth et al. 1993), creating a positive correlation between recombination rates and genetic diversity. Purifying selection against recessive variants, however, can also lead to associative overdominance (AOD, Ohta 1971, Zhao & Charlesworth, 2016), due to an apparent heterozygote advantage at linked neutral loci that opposes the loss of neutral diversity by BGS. Zhao & Charlesworth (2016) identified the conditions when AOD should dominate over BGS in a single-locus model and suggested that the effect of AOD could become stronger if multiple linked deleterious variants co-segregate. We present a model describing how and under which conditions multi-locus dynamics can amplify the effects of AOD. We derive the conditions for a transition from BGS to AOD due to pseudo-overdominance (Ohta & Kimura 1970), i.e. a form of balancing selection that maintains complementary deleterious haplotypes that mask the effect of recessive deleterious mutations. Simulations confirm these findings and show that multi-locus AOD can increase diversity in low recombination regions much more strongly than previously appreciated. While BGS is known to drive genome-wide diversity in humans (Pouyet et al. 2018), the observation of a resurgence of genetic diversity in regions of very low recombination is indicative of AOD. We identify 21 such regions in the human genome showing clear signals of multi-locus AOD. Our results demonstrate that AOD may play an important role in the evolution of low recombination regions of many species.

evolutionary biology↗

bioRxiv2018

Evolution of dispersal can rescue populations from expansion load

Understanding the causes and consequences of range expansions or range shifts has a long history in evolutionary biology. Recent theoretical, experimental, and empirical work has identified two particularly interesting phenomena in the context of species range expansions: (i) gene surfing and the relaxation of natural selection, and (ii) spatial sorting. The former can lead to an accumulation of deleterious mutations at range edges, causing an expansion load and slowing down expansion. The latter can create gradients in dispersal-related traits along the expansion axis and cause an acceleration of expansion. We present a theoretical framework that treats spatial sorting and gene surfing as spatial versions of natural selection and genetic drift, respectively. This model allows us to study analytically how gene surfing and spatial sorting interact, and to derive the probability of fixation of pleiotropic mutations at the expansion front. We use our results to predict the co-evolution of mean fitness and dispersal rates, taking into account the effects of random genetic drift, natural selection and spatial sorting, as well as correlations between fitnessand dispersal-related traits. We identify a \"rescue effect\" of spatial sorting, where the evolution of higher dispersal rates at the leading edge rescues the population from incurring expansion load.

evolutionary biology↗

bioRxiv2018

Mutation load dynamics during environmentally-driven range shifts

The fitness of spatially expanding species has been shown to decrease over time and space, but specialist species tracking their changing environment and shifting their range accordingly have been little studied. We use individual-based simulations and analytical modeling to compare the impact of range expansions and range shifts on genetic diversity and fitness loss, as well as the ability to recover fitness after either a shift or expansion. We find that the speed of a shift has a strong impact on fitness evolution. Fastest shifts show the strongest fitness loss per generation, but intermediate shift speeds lead to the strongest fitness loss per geographic distance. Range shifting species lose fitness more slowly through time than expanding species, however, their fitness compared at equivalent geographic distances spread can be considerably lower. These counter-intuitive results arise from the combination of time over which selection acts and mutations enter the system. Range shifts also exhibit reduced fitness recovery after a geographic shift and may result in extinction, whereas range expansions can persist from the core of the species range. The complexity of range expansions and range shifts highlights the potential for severe consequences of environmental change on species survival.\n\nAuthor SummaryAs environments change through time across the globe, species must adapt or relocate to survive. Specialized species must track the specific moving environments to which they are adapted, as compared to generalists which can spread widely. During colonization of new habitat, individuals can accumulate deleterious alleles through repeated bottlenecks. We show through simulation and analytic modeling that the process by which these alleles accumulate changes depending upon the speed at which populations spread over a landscape. This is due to the increased efficacy of selection against deleterious variants at slow speeds of range shifts and decreased input of mutations at faster speeds of range shifts. Under some selective circumstances, shifting of a species range leads to extinction of the entire population. This suggests that the rate of environmental change across the globe will play a large role in the survival of specialist species as compared to more generalist species.

evolutionary biology↗