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Diet breadth is a key life-history trait influencing range size, evolutionary trajectories, and ecosystem functioning. While diet breadth studies have traditionally been confined to animals, carnivorous plants provide an exciting conceptual extension to existing theory. We examine the possibility of diet breadth expansion in Sarracenia purpurea, a carnivorous pitcher plant, integrating field and greenhouse experiments with CHN analysis to quantify leaf litter consumption. Wild plants captured leaf detritus at levels comparable to insect prey. Pitchers fed leaf biomass in the field exhibited non-significant increases in foliar nitrogen, while the greenhouse experiment showed no effect of leaf litter. These results demonstrate that S. purpurea pitchers capture a substantial amount of leaf litter but show no clear evidence of nitrogen assimilation from this material. We found that short-term nitrogen acquisition from captured litterfall is minimal or absent in S. purpurea pitchers, especially relative to rapid nutrient assimilation from arthropod prey.

Tropical pitcher plants (Nepenthes) are carnivorous plants that trap and digest prey using highly modified fluid-filled leaves known as pitchers. Prey are digested by plant-secreted enzymes and pitcher symbionts. Pitchers exert control over abiotic properties of the digestive fluid such as pH levels that can influence its symbionts. Here we examine natural variation in pH and dissolved mineral concentrations in three sympatric Nepenthes species, assessing correlations between fluid properties and pitcher traits. We use addition experiments to investigate differences in protein digestion/absorption rates between species. Fluid pH and dissolved mineral levels both showed distinct patterns corresponding to pitcher developmental stages in N. gracilis and N. rafflesiana, whereas N. ampullaria differs from its congeners in exhibiting far less variation in fluid pH, as well as less clear evidence of protein depletion. This study further elucidates the properties of pitchers as habitats, revealing ways in which the host plant regulates that habitat.

Plants interface with and modify the external environment across their surfaces, and in so doing, can control or mitigate the impacts of abiotic stresses and also mediate their interactions with other organisms. Botanically, it is known that plant roots have a multi-faceted ability to modify rhizosphere conditions like pH, a factor with a large effect on a plant’s biotic interactions with microbes. But plants can also modify pH levels on the surfaces of their leaves. Plants can neutralize acid rain inputs in a period of hours, and either acidify or alkalinize the pH of neutral water droplets in minutes. The pH of the phylloplane—that is, the outermost surface of the leaf—varies across species, from incredibly acidic (carnivorous plants: as low as pH 1) to exceptionally alkaline (species in the plant family, Malvaceae, up to pH 11). However, most species mildly acidify droplets on the phylloplane by 1.5 orders of magnitude in pH. Just as rhizosphere pH helps shape the plant microbiome and is known to influence belowground interactions, so too can phylloplane pH influence aboveground interactions in plant canopies. In this review, we discuss phylloplane pH regulation from the physiological, molecular, evolutionary, and ecological perspectives and address knowledge gaps and identify future research directions. Plants alter external environmental conditions in many ways. Well-known is the fact that roots alter the pH of the surrounding soil. However, less appreciated is the fact that plants also alter pH levels on their leaf surfaces. Our review explores this little-known topic; we discuss variation in leaf surface pH across a diversity of plants, the physiological ways by which plants can modify leaf surface pH and the importance of this trait to ecological interactions, including those with microbes and insects. This topic presents many unanswered questions awaiting future work, with relevance to agriculture as well as wild ecosystems.

Characteristics of host species can alter how other, interacting species assemble into communities by acting as ecological filters. Pitchers of tropical pitcher plants ( Nepenthes ) host diverse communities of aquatic arthropods and microbes in nature. This plant genus exhibits considerable interspecific diversity in morphology and physiology; for example, different species can actively control the pH of their pitcher fluids and some species produce viscoelastic fluids. Our study investigated the extent to which Nepenthes species differentially regulate pitcher fluid traits under common garden conditions, and the effects that these trait differences had on their associated communities. Sixteen species of Nepenthes were reared together in the controlled environment of a glasshouse using commonly-sourced pH 6.5 water. We analyzed their bacterial and eukaryotic communities using metabarcoding techniques, and found that different plant species differentially altered fluid pH, viscosity, and color, and these had strong effects on the community structure of their microbiota. Nepenthes species can therefore act as ecological filters, cultivating distinctive microbial communities despite similar external conditions, and blurring the conceptual line between biotic and abiotic filters.

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 analyse how its novel traits (dioecy and carnivorous pitcher leaves) are linked to genomic evolution. We found a decaploid karyotype and a clear indication of subgenome dominance. A male-linked and pericentromerically located region on the putative sex chromosome was identified in a recessive subgenome and was found to harbour 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. A genome-scale analysis suggested that subgenome dominance likely contributed to evolutionary innovation by permitting recessive subgenomes to diversify functions of novel tissue-specific duplicates. Our results provide insight into how polyploidy can give rise to novel traits in divergent and successful high-ploidy lineages.The genome of the Asian pitcher plant shows a decaploid structure and highlights the differential contributions of subgenomes to the evolution of novel genes, such as those associated with dioecy and trapping pitcher leaves.

Elevation is an important determinant of ecological community composition. It integrates several abiotic features and leads to strong, repeatable patterns of community structure, including changes in the abundance and richness of numerous taxa. However, the influence of elevational gradients on microbes is understudied relative to plants and animals. To compare the influence of elevation on multiple taxa simultaneously, we sampled phytotelm communities within a tropical pitcher plant (Nepenthes mindanaoensis) along a gradient from 400 to 1200 m a.s.l. We use a combination of metabarcoding and physical counts to assess diversity and richness of bacteria, micro-eukaryotes, and arthropods, and compare the effect of elevation on community structure to that of regulation by a number of plant factors. Patterns of community structure differed between bacteria and eukaryotes, despite their living together in the same aquatic microhabitats. Elevation influences community composition of eukaryotes to a significantly greater degree than it does bacteria. When examining pitcher characteristics, pitcher dimorphism has an effect on eukaryotes but not bacteria, while variation in pH levels strongly influences both taxa. Consistent with previous ecological studies, arthropod abundance in phytotelmata decreases with elevation, but some patterns of abundance differ between living inquilines and prey.

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 (Cohen's d ≈ 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.Competing Interest StatementThe authors have declared no competing interest.Funder Information DeclaredU.S. National Science Foundation, https://ror.org/021nxhr62

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.

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.