An accumulating body of evidence indicates that natural plant populations harbour a large diversity of transposable elements (TEs). TEs provide genetic and epigenetic variation that can substantially translate into changes in plant phenotypes. Despite the wealth of data on the ecological and evolutionary effects of TEs on plant individuals, we have virtually no information on the role of TEs on populations and ecosystem functioning. On the example of Arabidopsis thaliana, we demonstrate that TE-generated variation creates differentiation in ecologically important functional traits. In particular, we show that Arabidopsis populations with increasing diversity of individuals differing in copy numbers of the ONSEN retrotransposon had higher phenotypic and functional diversity. Moreover, increased diversity enhanced population productivity and reduced performance of interspecific competitors. We conclude that TE-generated diversity can have similar effects on ecosystem as usually documented for other biological diversity effects.
The efficient conservation of vulnerable ecosystems in the face of global change requires a complete understanding of how plant communities respond to various environmental factors. We aim to demonstrate that a combined use of different approaches, traits, and indices representing each of the taxonomic and functional characteristics of plant communities will give complementary information on the factors driving vegetation assembly patterns. We analyzed variation across an environmental gradient in taxonomic and functional composition, richness, and diversity of the herb-layer of a temperate beech-oak forest that was located in northern Spain. We measured species cover and four functional traits: leaf dry matter content (LDMC), specific leaf area (SLA), leaf size, and plant height. We found that light is the most limiting resource influencing herb-layer vegetation. Taxonomic changes in richness are followed by equivalent functional changes in the diversity of leaf size but by opposite responses in the richness of SLA. Each functional index is related to different environmental factors even within a single trait (particularly for LDMC and leaf size). To conclude, each characteristic of a plant community is influenced by different and even contrasting factors or processes. Combining different approaches, traits, and indices simultaneously will help us understand how plant communities work.
Phenotypic plasticity is essential for organisms to adapt to local ecological conditions. It is expected that mutualistic interactions, such as arbuscular mycorrhizal (AM) symbiosis, mediate plant phenotypic plasticity, although it is not clear to what extent this plasticity may be heritable (i.e. transgenerational plasticity). We tested for plant plasticity within‐ and across‐generations in response to AM symbiosis and varying water availability in a full factorial experiment over two generations, using a genetically uniform line of a perennial apomictic herb, Taraxacum brevicorniculatum. We examined changes in phenotype, performance and AM fungal colonization of the offspring throughout plant development. AM symbiosis and water availability triggered phenotypic changes during the life cycle of plants. Additionally, both factors triggered adaptive transgenerational effects especially detectable during the juvenile stage of the offspring. Water deficit and absence of AM fungi caused concordant plant phenotypic modifications towards a ‘stress‐coping phenotype’, both within and across generations. Parental environment also affected AM fungal colonization of the offspring. Juvenile offspring of amply watered parents and adut offspring of mycorrhizal parents had increasing length of root colonized by AM fungi when they were under water stress. Synthesis. We show that, in addition to providing beneficial transgenerational effects in offspring traits proxies of fitness (such as increased biomass, survival or nutritional status), AM symbiosis can trigger transgenerational plasticity in anatomical and physiological traits related to resource‐use acquisition, and further influence offspring AM fungal colonization. Our results show that AM symbiosis could improve plants' ability to cope with environmental stress, not only within, but also across generations.
Population ecologists develop theoretical and pragmatic knowledge of how and why populations change or remain stable, how life histories evolve and devise management strategies for populations of concern. However, forecasting the effects of global change or recommending management strategies is often urgent, requiring ecologists to work without detailed local evidence while using data and models from outside the focal location or species. Here we explore how the comparative ecology of populations, population macroecology, can be used to develop generalisations within and between species across different scales, using available demographic, environmental, life history, occurrence and trait data. We outline the strengths and weaknesses of using broad climatic variables and suitability inferred from probability of occupancy models to represent environmental variation in comparative analyses. We evaluate the contributions of traits, environment and their interaction as drivers of life history strategy. We propose that insights from life history theory, together with the adaptive capacity of populations and individuals, can inform on 'persist in place' vs 'shift in space' responses to changing conditions. As demographic data accumulate at landscape and regional scales for single species, and throughout plant phylogenies, we will have new opportunities for testing macroecological generalities within and across species.
Premise of research. Root traits of annual plants are known to vary across environmentally manipulated conditions in controlled settings. Roots absorb nutrients and water, essential for individual function and survival. Yet how ruderal plant populations respond belowground to spatial and temporal variation in field conditions is largely unknown. The present study examines the natural variation of root traits and survival among populations of the annual Arabidopsis thaliana across gardens along a fertility gradient.Methodology. We established common gardens in field settings and in raised beds in the plant’s native and nonnative ranges using populations of A. thaliana from western/central Spain. Survival through the growing season and root traits were investigated for garden and population influences and for the relationship between survival and root biomass.Pivotal results. We demonstrate differences in survival among gardens, populations, and years. Survival was highest in the higher-fertility gardens in the native and nonnative ranges. We detected plasticity (among gardens and between years) of root length, lateral root production, primary root length, and biomass. Roots were substantially larger when grown in the more fertile gardens, where water and nutrients were consistently available. Yet lateral root production was also high in a low-fertility field garden. We detected substantial among-population variation in root traits in the higher-fertility gardens.Conclusions. This study demonstrates among-population variation in root traits in field gardens and substantial root plasticity measured across locations. Evidence of local adaptation was indicated by native local populations showing the highest survival and root growth in the home native garden. Such differences in survival and root functional trait responses inform how plants respond to dramatic changes in environmental conditions, particularly in novel sites. We advocate for additional studies of belowground traits to further uncover the extent of natural variation in root phenotypic responses in the field.
Background and Aims The observed positive diversity effect on ecosystem functioning has rarely been assessed in terms of intraspecific trait variability within populations. Intraspecific phenotypic variability could stem both from underlying genetic diversity and from plasticity in response to environmental cues. The latter might derive from modifications to a plant's epigenome and potentially last multiple generations in response to previous environmental conditions. We experimentally disentangled the role of genetic diversity and diversity of parental environments on population productivity, resistance against environmental fluctuations and intraspecific phenotypic variation. Methods A glasshouse experiment was conducted in which different types of Arabidopsis thaliana populations were established: one population type with differing levels of genetic diversity and another type, genetically identical, but with varying diversity levels of the parental environments (parents grown in the same or different environments). The latter population type was further combined, or not, with experimental demethylation to reduce the potential epigenetic diversity produced by the diversity of parental environments. Furthermore, all populations were each grown under different environmental conditions (control, fertilization and waterlogging). Mortality, productivity and trait variability were measured in each population. Key Results Parental environments triggered phenotypic modifications in the offspring, which translated into more functionally diverse populations when offspring from parents grown under different conditions were brought together in mixtures. In general, neither the increase in genetic diversity nor the increase in diversity of parental environments had a remarkable effect on productivity or resistance to environmental fluctuations. However, when the epigenetic variation was reduced via demethylation, mixtures were less productive than monocultures (i.e. negative net diversity effect), caused by the reduction of phenotypic differences between different parental origins. Conclusions A diversity of environmental parental origins within a population could ameliorate the negative effect of competition between coexisting individuals by increasing intraspecific phenotypic variation. A diversity of parental environments could thus have comparable effects to genetic diversity. Disentangling the effect of genetic diversity and that of parental environments appears to be an important step in understanding the effect of intraspecific trait variability on coexistence and ecosystem functioning.
Competitive exclusion is to be expected between phylogenetically similar species that share traits and resources. However, species may overcome this, either through differentiation of their responses to biotic and abiotic conditions, or by trait differentiation, thus enabling their coexistence. We identified differences in phenotypic traits between seven coexisting Carex species and their responses to competition and fertilization in pot experiments, before using long-term field experiments to generate responses of the Carex species to fertilization and mowing and to illustrate temporal variability between species. Finally, we assessed how effective the results of the pot experiment were at predicting species responses in the field. In pot experiments, we found that species responded more to competition than to fertilization. Notably, all species showed similar responses to these factors in the pot experiments. Fertilization decreased the root:shoot ratio, whilst competition decreased growth-related characteristics such as total biomass, irrespective of the species. Differences among species were only found in their clonal response to competition, namely rhizome production and generation rate of new ramets. These findings support the idea that different clonal growth strategies may facilitate niche partitioning of Carex species. Species responses measured from pot experiments were poor predictors of their responses in the field experiment. Nevertheless, we confirmed the prediction that, over time, Carex species with lower growth rates in pot experiments showed more stable biomass production than in the field. We suggest that differences in clonal traits and temporal dynamics support the ability of Carex species to avoid competitive exclusion, enabling their coexistence.
Phenotypic plasticity, within and across generations (transgenerational plasticity), allows organisms and their progeny to adapt to the environment without modification of the underlying DNA. Recent findings suggest that epigenetic modifications are important mediators of such plasticity. However, empirical studies have, so far, mainly focused on plasticity in response to abiotic factors, overlooking the response to competition. We tested for within-generation and transgenerational phenotypic plasticity triggered by plant-plant competition intensity, and we tested whether it was mediated via DNA methylation, using the perennial, apomictic herb Taraxacum brevicorniculatum in four coordinated experiments. We then tested the consequences of transgenerational plasticity affecting competitive interactions of the offspring and ecosystem processes, such as decomposition. We found that, by promoting differences in DNA methylation, offspring of plants under stronger competition developed faster and presented more resource-conservative phenotypes. Further, these adjustments associated with less degradable leaves, which have the potential to reduce nutrient turnover and might, in turn, favour plants with more conservative traits. Greater parental competition enhanced competitive abilities of the offspring, by triggering adaptive phenotypic plasticity, and decreased offspring leaf decomposability. Our results suggest that competition-induced transgenerational effects could promote rapid adaptations and species coexistence and feed back on biodiversity assembly and nutrient cycling.
Disturbance is an omnipresent selective factor that shapes plant strategies. While annual species that rely on rapid generative reproduction dominate in habitats frequently affected by severe disturbance, long-lived woody species occupy habitats where the effects of disturbance are weak. These are, however, the extremes in the disturbance gradient. Habitats under intermediate disturbance are occupied by the whole spectrum of plant strategies. In this study, we hypothesized that clonal herbs are better adapted to intermediate disturbance than non-clonal herbs because the characteristics of their bud bank and related belowground carbohydrate storage promote vegetative regeneration. In a greenhouse experiment, we tested the effects of disturbance on the survival and performance of 17 congeneric pairs of clonal and non-clonal herbs cultivated from seed. In addition to a non-disturbed control, we applied five disturbance treatments: biomass removal 2 cm above the soil surface, removal of 70 % of the aboveground biomass, late spring frost, flooding, and belowground injury. Mortality was negligible except the flooding treatment, where it was higher for non-clonal than for clonal plants. Total biomass was reduced more by the disturbance treatments for clonal than for non-clonal plants, suggesting that clonal species suffer more than non-clonal species. On the other hand, clonal plants had significantly greater total belowground biomass and R:S ratios than non-clonal plants, suggesting a strong investment in belowground organs by clonal plants. In response to disturbance, the SLA and SRL values increased in clonal plants but decreased in non-clonal plants, indicating that clonal and non-clonal plants differ in how they cope with disturbance. Although clonality is considered to be an adaptation to disturbance, our results indicate that, during the first year of life, clonal herbs are more sensitive to disturbance than non-clonal herbs, likely due to preferential investment in belowground biomass.
Functional and phylogenetic diversity (FD and PD respectively) of the resident community are expected to exert a key role in community resistance to colonization by surrounding species, and their establishment success. However, few studies have explored this topic experimentally or evaluated the interactive effects of these diversity measures. We implemented a diversity experiment to disentangle the role of FD and PD by sowing mixtures of 6 species, drawn from a pool of 19 species naturally coexisting in central European mesic meadows. The mixtures were designed to cover four independent combinations of high and low FD and PD. Species covers were estimated in spring and late summer over two growing seasons. We then assessed the establishment success of colonizers as a function of their mean traits and phylogenetic distance to the resident (i.e. sown) communities, as well as the resistance of the resident communities to natural colonizers as a function of their functional and phylogenetic structure. Results generally indicated a temporal shift regarding which trait values made a colonizer successful, from an acquisitive strategy in early stages to a more conservative trait syndrome in later stages. FD decreased community resistance to natural colonization. However, PD tempered this effect: with high PD, FD was not significant, suggesting complementary information between these two components of biodiversity. On average, colonizing species were more functionally distant from the resident species in sown communities with high functional diversity, i.e. those that were more colonized. Synthesis. Our results confirm an interplay between FD and PD during community assembly processes, namely resistance to colonizers, suggesting that these two descriptors of biodiversity only partially overlap in their contribution to the overall ecological structure of a community. The hypothesis that higher FD increases resistance through a more complete use of resources was challenged. Results rather suggested that greater FD could provide an unsaturated functional trait space allowing functionally unique species to occupy it.
Experimental demethylation of plant DNA enables testing for epigenetic effects in a simple and straightforward way without the use of expensive and laborious DNA sequencing. Plants are commonly demethylated during their germination with the application of agents such as 5-azacytidine (5-azaC). However, this approach can cause unwanted effects such as underdeveloped root systems and high mortality of treated plants, hindering a full comparison with untreated plants, and can be applied only on plant reproducing by seeds. Here we test a simple alternative method of plant demethylation designed to overcome the shortcomings of the germinating method. We compared a novel method of demethylating plants, based on periodical spraying of 5-azaC aqueous solution on established seedlings, with the previous method in which seeds were germinated directly in 5-azaC solution. We quantified the amount of methylated DNA and measured various aspects of plant performance. Also, we demonstrated its applicability in ecological epigenetic experiments by testing transgenerational effects of plant-plant competition. We found that the spray application had similar DNA-demethylating efficiency than the germination method, particularly in the earlier phases of plant development, but without unwanted effects. The spray application method did not reduce plant growth and performance compared to untreated plants, as opposed to the traditional method which showed reduced growth. Also, the spray application method equalized the epigenetically modified plant features of seedlings coming from plants grown under competition and plants growing without competition, demonstrating its application in ecological epigenetic experiments. We conclude that regular spraying of 5-azaC solution onto established seedlings surpassed the germination-in-solution method in terms of vigour and fitness of treated plants. This novel method could thus be better suited for experimental studies seeking valuable insights into ecological epigenetics. Furthermore, the spray method can be suitable for clonal species reproducing asexually, and most importantly, it opens the possibility of community-level experimental demethylation of plants.