Clonal Plant Populations Research Articles

Microbe-induced phenotypic variation leads to overyielding in clonal plant populations.

Overyielding, the high productivity of multispecies plant communities, is commonly seen as the result of plant genetic diversity. Here we demonstrate that biodiversity-ecosystem functioning relationships can emerge in clonal plant populations through interaction with microorganisms. Using a model clonal plant species, we found that exposure to volatiles of certain microorganisms led to divergent plant phenotypes. Assembling communities out of plants associated with different microorganisms led to transgressive overyielding in both biomass and seed yield. Our results highlight the importance of belowground microbial diversity in plant biodiversity research and open new avenues for precision ecosystem management.

Unequal carbon and nitrogen translocation between ramets affects sexual reproductive performance of the clonal grass Leymus chinensis under nitrogen addition

Sexual reproduction is crucial for population continuity in clonal plants. The effect of nutrient translocation between ramets on sexual reproduction of clonal plants under nitrogen addition remains unclear. In this study, we focused on clonal fragments of Leymus chinensis reproductive ramets with different number of vegetative ramets connected to tillering nodes. A series of pot experiments was conducted under nitrogen addition, including 13C and 15N bidirectional labelling of vegetative ramets and reproductive ramets at the milk-ripe stage, determination of the 13C and 15N amount translocated, and assessment of the quantitative characteristics, nitrogen and carbon concentrations of reproductive ramets and vegetative ramets. Nitrogen addition promoted the translocation of 13C while inhibiting 15N between vegetative ramets and reproductive ramets. With an increase in the number of connected vegetative ramets, the 13C translocated by reproductive ramets and the 15N translocated by reproductive and vegetative ramets gradually increased. The translocation of 13C and 15N between vegetative and reproductive ramets was bidirectional and unequal. The translocated amount of 13C and 15N from reproductive ramets to vegetative ramets was always higher than that from vegetative ramets to reproductive ramets. Nitrogen addition did not prominently affect the sexual reproductive performance of L. chinensis, whereas the number of connected vegetative ramets both positively and negatively affected sexual reproductive performance. Ramet biomass is an important driver of nutrient acquisition by L. chinensis ramets. We demonstrate for the first time that unequal nutrient translocation between ramets affects sexual reproductive performance in L. chinensis. The findings contribute to an enhanced understanding of the reproductive strategies of clonal plant populations in future environments.

DNA methylation in the wild: epigenetic transgenerational inheritance can mediate adaptation in clones of wild strawberry (Fragaria vesca).

Due to the accelerating climate change, it is crucial to understand how plants adapt to rapid environmental changes. Such adaptation may be mediated by epigenetic mechanisms like DNA methylation, which could heritably alter phenotypes without changing the DNA sequence, especially across clonal generations. However, we are still missing robust evidence of the adaptive potential of DNA methylation in wild clonal populations. Here, we studied genetic, epigenetic and transcriptomic variation of Fragaria vesca, a predominantly clonally reproducing herb. We examined samples from 21 natural populations across three climatically distinct geographic regions, as well as clones of the same individuals grown in a common garden. We found that epigenetic variation was partly associated with climate of origin, particularly in non-CG contexts. Importantly, a large proportion of this variation was heritable across clonal generations. Additionally, a subset of these epigenetic changes affected the expression of genes mainly involved in plant growth and responses to pathogen and abiotic stress. These findings highlight the potential influence of epigenetic changes on phenotypic traits. Our findings indicate that variation in DNA methylation, which can be environmentally inducible and heritable, may enable clonal plant populations to adjust to their environmental conditions even in the absence of genetic adaptation.

Clonal integration promotes the growth of Phragmites australis populations in saline wetlands of the Yellow River Delta.

Estuarine wetlands are highly heterogeneous due to strong interactions between freshwater input and seawater intrusion. However, little is known about how clonal plant populations adapt to heterogeneous salinity in soil environments. In the present study, the effects of clonal integration on Phragmites australis populations under salinity heterogeneity were studied using field experiments with 10 treatments in the Yellow River Delta. Clonal integration significantly increased plant height, aboveground biomass, underground biomass, root-shoot ratio, intercellular CO2 concentration, net photosynthetic rate, stomatal conductance, transpiration rate, and stem Na+ content under homogeneous treatment. Under the heterogeneous salt treatment, clonal integration significantly affected total aboveground and underground biomass, photosynthetic traits, and stem Na+ content under different salt gradients. The increase in salt concentration inhibited the physiological activity and growth of P. australis to varying degrees. Compared with the heterogeneous saline environment, clonal integration was more beneficial to P. australis populations in the homogeneous saline habitat. The results of the present study suggest that P. australis prefers homogeneous saline habitats; however, plants can adapt to heterogeneous salinity conditions via clonal integration.

Soil saline-alkali heterogeneity is an important factor driving the spatial expansion of clonal plant in grassland

Soil salinity is well known heterogeneous and various within natural soil environment. In Songnen grassland of Northeast China, grazing aggravates the saline-alkali heterogeneity in soil habitat, which led to dominant clonal plant species forming a variety of adaptive strategies. However, based on the previous studies of morphological plasticity and clonal integration in clonal plants, there was a lack of mechanism research on the spatial expansion strategy of clonal plants population subjected to clipping in saline-alkali heterogeneity soil patches. To address this knowledge gap, we carried out an experiment by applying different clipping intensities (0%, 35%, 70% of the above-ground biomass removed) to explore the spatial expansion strategy and morphological plasticity of Leymus chinensis and their belowground rhizomes in different heterogeneous saline-alkali patches. We found that, clipping significantly decreased the plant average height and above-ground biomass in homogeneous patches, especially heavy clipping had a significant adverse impact on plant belowground (rhizome, fine root) biomass and leaf area. However, there was no significant difference in the biomass, average height, and leaf area among the clipping treatments in low saline-alkali heterogeneous patches. In addition, the number of leaves, daughter plants and rhizome internode buds were significantly affected by saline-alkali heterogeneity alone, and low saline-alkali heterogeneity had a positive effect on these parameters. Biomass accumulation and rhizome expansion were gradually inhibited by the increasing of clipping intensity in a homogeneous environment, but the inhibitory effect of clipping was not significant in saline-alkali patches soil. We concluded that L. chinensis can respond to grazing or mowing disturbance by increasing tiller and daughter-plant numbers in a saline-alkali heterogeneous environment. The saline-alkali patches promoted clonal plant population spatial expansion more than grazing or mowing disturbance did. This study emphasizes saline-alkali heterogeneity distribution patterns in grassland soil environment were the primary and important driving factor promoting the spatial expansion of L. chinensis.

Soil Moisture Rather than Soil Nutrient Regulates the Belowground Bud Bank of Rhizomatous Species Psammochloa villosa in Arid Sand Dunes

In arid and semi-arid sand dune ecosystems, belowground bud bank plays an important role in population regeneration and vegetation restoration. However, the responses of belowground bud bank size and composition to sand burial and its induced changes in soil environmental factors have been rarely studied. In arid sand dunes of Northwestern China, we investigated belowground bud bank size and composition of the typical rhizomatous psammophyte Psammochloa villosa as well as three key soil environmental factors (soil moisture, total carbon and total nitrogen) under different depths of sand burial. Total buds and rhizome buds increased significantly with increasing burial depth, whereas tiller buds first increased and then decreased, with a peak value at the depth of 20–30 cm. Soil moisture increased significantly with sand burial depth, and was positively correlated with the number of all buds and rhizome buds. Soil total carbon concentration first increased and then decreased with sand burial depth, and total nitrogen concentration was significantly lower under deep sand burial than those at shallow depths, and only the number of tiller buds was positively correlated with soil total nitrogen concentration. These results indicate that soil moisture rather than soil nutrient might regulate the belowground bud bank of P. villosa, and that clonal psammophytes could regulate their belowground bud bank in response to sand burial and the most important environmental stress (i.e., soil moisture). These responses, as the key adaptive strategy, may ensure clonal plant population regeneration and vegetation restoration in arid sand dunes.

Genet dynamics and its variation among genets of a clonal plant Convallaria keiskei

In clonal plant populations, a number of genetically identical ramets form a genet. While coexisting ramets potentially perform independently, their behaviours not only depend on ages and sizes but are also constrained by genetic background. In this study, genet dynamics and its variability among neighbouring genets were investigated based on the ramet demography of each genet in Convallaria keiskei. Genet dynamics were first formulated as a matrix model with the two components of clonal growth (clonal reproduction) and survival‐transitions between ramet size classes. Then, a statistical estimation of the matrix elements was established using three datasets: aboveground demographic censuses, belowground directional rhizome connections and genetic identification of ramets. Finally, genet growth rates reflecting both the changes of clonal growth and ramet size growth were estimated and compared for fundamental demographic elements among genets. Over three years of aboveground annual censuses of a 28 × 2 m plot, 2021 ramets were identified as belonging to 28 genotypes. Belowground excavation detected 515 clonal fragments. Genet growth rate of three dominant genets varied with medians of 1.13, 1.02 and 1.05; 95% credible intervals of the posterior distributions did not overlap between the genet with the largest median and the others. The variation was caused primarily by differences in clonal growth rather than survival‐transitions between size classes. Clonal growth by branching was rarer than at the tips but contributed to the maintenance of the genet. Therefore, both clonal growth frequencies and connecting patterns of ramets caused the variation of genet dynamics and established genets persist for a long time through the positive growth rates, which would contribute to maintain a population. We also conclude that fundamental demographic elements relating to clonal growth traits (the features of individual genets) strongly impact genet dynamics.

Predictable and stable epimutations induced during clonal plant propagation with embryonic transcription factor.

Clonal propagation is frequently used in commercial plant breeding and biotechnology programs because it minimizes genetic variation, yet it is not uncommon to observe clonal plants with stable phenotypic changes, a phenomenon known as somaclonal variation. Several studies have linked epigenetic modifications induced during regeneration with this newly acquired phenotypic variation. However, the factors that determine the extent of somaclonal variation and the molecular changes underpinning this process remain poorly understood. To address this gap in our knowledge, we compared clonally propagated Arabidopsis thaliana plants derived from somatic embryogenesis using two different embryonic transcription factors- RWP-RK DOMAIN-CONTAINING 4 (RKD4) or LEAFY COTYLEDON2 (LEC2) and from two epigenetically distinct founder tissues. We found that both the epi(genetic) status of the explant and the regeneration protocol employed play critical roles in shaping the molecular and phenotypic landscape of clonal plants. Phenotypic variation in regenerated plants can be largely explained by the inheritance of tissue-specific DNA methylation imprints, which are associated with specific transcriptional and metabolic changes in sexual progeny of clonal plants. For instance, regenerants were particularly affected by the inheritance of root-specific epigenetic imprints, which were associated with an increased accumulation of salicylic acid in leaves and accelerated plant senescence. Collectively, our data reveal specific pathways underpinning the phenotypic and molecular variation that arise and accumulate in clonal plant populations.

Inundation Depth Shape Phenotypic Variability of Phragmites australis in Liaohe Estuary Wetland, Northeast China

Inundation shifts plant growth, species diversity and ecosystem stability, but it remains unclear how inundation depth shapes the phenotypic variability of clonal plants in an estuary wetland. To reveal the response of clonal plant populations to inundation depth, we calculated the variation of Phragmites australis using shoot height, leaf length, leaf width, leaf biomass and panicle length in the Liaohe estuary wetland. Reproductive allocation was defined by the ratio of panicle length to shoot height. Linear regression showed that shoot height, leaf length, leaf width, leaf biomass and panicle length were negatively correlated with inundation depth, while the ratio of panicle length to shoot height was negatively correlated with inundation depth (p < 0.0001). Based on data regarding the statistical differences of plant phenotypic traits among P. australis, we recognized populations had generated variation differentiation. Compared with other functional traits, the coefficient of variation of leaf-related traits were at a high level. Therefore, leaf parameters would be the most suitable, and they increased the area and weight to support the action of plants during floods. Multivariate statistical analysis suggested that P. australis populations in the Liaohe estuary wetland were divided into two phenotypic clusters, consistent with geographical distance and morphological similarity. Our results provide a novel perspective on the ecological strategy of cloned plants under inundation change and offer theories for the conservation and restoration of estuarine wetland ecosystems.

Light condition experienced by parent plants influences the response of offspring to light via both parental effects and soil legacy effects

Abstract Environmental conditions experienced by parent plants can influence offspring performance through parental effects induced by DNA methylation. The offspring can also be influenced by environmental conditions experienced by their parents via soil legacy effects due to plant‐mediated changes in the composition of soil microbes. These two effects are likely to act simultaneously, but empirical evidence for combined effects is limited. We conducted a two‐phase experiment with five genotypes of a clonal plant Hydrocotyle vulgaris. In the first phase, we grew parent plants of each genotype under two light conditions (ambient vs. shade) and two DNA demethylation treatments [treated with water vs. 5‐azacytidine (5‐azaC)]. We then collected soils and clonal offspring for each genotype from each of these four treatments and measured soil (a)biotic properties. In the second phase, we grew the offspring from each of the four treatments in the four different soils, under the two light conditions. When grown under ambient light condition and in soil from ambient parents, offspring produced by ambient parents grew larger than offspring produced by shaded parents when the parents were treated with water. This difference was smaller when the parents were treated with 5‐azaC, and disappeared when the offspring were grown in soil from shaded parents. The growth difference was also observed when the offspring were grown under shaded condition and in soil from shaded parents. However, this difference was greater when the parents were treated with 5‐azaC, and disappeared when the offspring were grown in soil from ambient parents. Moreover, offspring growth was associated with fungal composition and total phosphorus of the soil in which the parents had grown. Our results show, for the first time, that light condition experienced by parents can influence offspring responses to light through both parental effects and soil legacies. The parental effects were mediated by changes in DNA methylation and the soil legacies were due to plant‐mediated changes in a combination of soil biotic and abiotic properties. These impacts may eventually influence the ecological and evolutionary trajectories of clonal plant populations. Read the free Plain Language Summary for this article on the Journal blog.

Effects of Flag Leaf and Number of Vegetative Ramets on Sexual Reproductive Performance in the Clonal Grass Leymus chinensis

Sexual reproduction is vital for population adaptation in clonal plants. The flag leaf is considered to be the primary contributor to sexual reproduction in cereal crops, and there is no unified conclusion on the effect of the number of vegetative ramets on grain yield. However, what effects of the flag leaf and the number of vegetative ramets on sexual reproductive performance of clonal grasses are largely unknown. To test this, under field natural conditions, we grew the rhizomatous grass Leymus chinensis in a homogeneous environment and conducted studies concerning the growth, reproduction and physiology of reproductive ramets in clonal populations. We measured the growth characteristics of different aged leaves, dynamically measured the net photosynthetic rate of different aged leaves and organ biomass, measured the sexual reproductive characteristics of reproductive ramets that had different numbers of connecting vegetative ramets, and performed isotope (15N) labeling of ramet pairs at the seed-filling stage. In L. chinensis clonal populations, from the heading stage, the photosynthetic contribution of the functional leaves to seed production was much greater than that of the flag leaf; the photosynthetic capacity of both the functional leaves and the flag leaf all gradually declined. Vegetative ramets translocated their own resources to the connected reproductive ramets, and a large proportion of translocated resources were allocated to the leaf and stem to sustain life activities; increase in the number of connecting vegetative ramets increased floret number, seed number, seed-setting rate, inflorescence biomass, seed biomass, and reproductive allocation of reproductive ramets, and these parameters significantly and positively correlated with the biomass of connecting vegetative ramets. We conclude that the functional leaf rather than the flag leaf of L. chinensis is the primary contributor to seed production. Reproductive ramets adopt a strategy of growth first and reproduction later to allocate the translocated resources between the organs, but vegetative ramets are very advantageous for sexual reproduction under the tillering node connection form in L. chinensis. Overall, our study implies that vegetative ramets not only play an important role in the spatial expansion but also in the sexual reproduction of clonal plant populations.

Effects of Clipping on Bud Bank and Population Regeneration of Triarrhena lutarioriparia in Dongting Lake Wetland, China

The bud bank plays an important role in the regeneration of clonal plants after disturbance. However, specific effects of clipping disturbance on the bud bank and population characteristics of clonal plants remain unclear. Here, no-clipping (CK), low-intensity continuous clipping (LCC), and high-intensity single clipping (HSC) were tested on Triarrhena lutarioriparia, a typical rhizomatous clonal plant population. The results with respect to population characteristics and bud bank responses showed that: (1) Plant density under HSC and LCC was higher than that under CK, and compared with CK, bud density belowground decreased significantly under HSC; (2) The number of basal buds increased significantly under LCC but disappeared under HSC; the number of terminal buds under LCC and HSC decreased significantly; and (3) The proportions of guerrilla ramets under HSC and LCC were significantly lower than those under CK, while the proportions of phalanx ramets under HSC and LCC were significantly higher than those under CK. The bud bank is the main source of the aboveground population after clipping disturbance. Clipping changed bud-sprouting order and bud bank composition, ultimately leading the plant to change from guerrilla to phalanx growth form. This is an effective strategy for recovery from clipping disturbance.

The role of sexual reproduction in the maintenance of established Zostera marina meadows

Abstract For clonal plants, the role of sexual reproduction in the maintenance of populations can vary widely. Some species are dependent on repeated seedling recruitment. For other species, interactions between adults and seedlings within existing populations can affect seedling survival and limit sexual reproduction in existing populations. Genetic studies of seagrass populations increasingly suggest sexual reproduction is important for the resilience and stability of their populations, but as of yet little observational data support these findings. Because seagrass populations provide important ecosystem services and are threatened with increasing anthropogenic impacts, understanding their reliance on sexual reproduction is evolutionarily and ecologically important. The goals of this study were to determine (a) whether seedlings of a marine angiosperm, Zostera marina, establish and recruit within existing Z. marina meadows and (b) whether interactions between seedlings and surrounding adult shoots influence the survival of established seedlings. To meet these goals, surveys estimated seedling establishment and tracked seedling survival within multiple populations. Manipulative experiments then tested the impact of neighbouring adult shoots on seedling survival and the overall trajectory of experimental plots with and without sexual reproduction. A 3‐year survey identified established seedlings within Z. marina meadows each year. Additionally, concurrent seed addition experiments indicated seed supply could influence seedling establishment rates. A survey tracking the survival of tagged seedlings, as well as the height and density of surrounding adult shoots, showed adult shoots may negatively impact seedling survival. Experiments then demonstrated that seedlings without neighbouring shoots survived longer than those with neighbouring shoots. Lastly, two transplant garden experiments comparing the survival of plots with and without seeds highlighted that seedling recruitment is likely most important to maintain bottom cover where disturbances generate gaps in the adult population. Synthesis. This study demonstrates that seedlings do establish within existing seagrass meadows, and that some survive to recruit into the adult population. Competition with existing vegetation, however, can be a factor compromising seedling survival. Sexual reproduction may thus most likely occur in, and be most important for, clonal plant populations that experience seasonal disturbance.

Genet assignment and population structure analysis in a clonal forest-floor herb, Cardamine leucantha, using RAD-seq.

To study the genetic structure of clonal plant populations, genotyping and genet detection using genetic markers are necessary to assign ramets to corresponding genets. Assignment is difficult as it involves setting a robust threshold of genetic distance for genet distinction as neighbouring genets in a plant population are often genetically related. Here, we used restriction site-associated DNA sequencing (RAD-seq) for a rhizomatous clonal herb, Cardamine leucantha [Brassicaceae] to accurately determine genet structure in a natural population. We determined a draft genome sequence of this species for the first time, which resulted in 66 617 scaffolds with N50 = 6086 bp and an estimated genome size of approximately 253 Mbp. Using genetic distances based on the RAD-seq analysis, we successfully distinguished ramets that belonged to distinct genets even from a half-sib family. We applied these methods to 372 samples of C. leucantha collected at 1-m interval grids within a 20 × 20 m plot in a natural population in Hokkaido, Japan. From these samples, we identified 61 genets with high inequality in terms of genet size and patchy distribution. Spatial autocorrelation analyses indicated significant aggregation within 7 and 4 m at ramet and genet levels, respectively. An analysis of parallel DNA microsatellite loci (simple sequence repeats) suggested that RAD-seq can provide data that allows robust genet assignment. It remains unclear whether the large genets identified here became dominant stochastically or deterministically. Precise identification of genets will assist further study and characterization of dominant genets.

Clonal diversity and spatial genetic structure in the long-lived herb, Prairie trillium.

Combining population genetic studies with demographic surveys in long-lived clonal herbs can yield insight into the population dynamics of clonal plant populations. In this study, we assayed clonal diversity and spatial genetic structure in a population of a long-lived understory herb, Trillium recurvatum, that has been the focus of a demographic study spanning 26 years at the Meeman Biological Station in Memphis, Tennessee, USA. Using a set of five newly developed simple sequence repeat markers first reported here, we assessed 1) the extent of clonal diversity within the Meeman site, 2) the degree to which genetic diversity varies with stage class (juvenile, non-flowering, and flowering adults) at this site, 3) whether there is spatial genetic structure at the Meeman site, and 4) how measures of genetic diversity and inbreeding at the Meeman site compare to two additional nearby populations. Along with these analyses, we calculated and compared traditional population genetic metrics with information theory-based diversity indices. Although clonal propagation was present, the focal population displayed moderate levels of clonal diversity comprising 81 genets from the 174 individuals sampled. In the focal site, we also found that genetic diversity was highest in the flowering stage class when compared to the non-flowering and juvenile classes. We report that genets exhibited spatial genetic structure in the focal site exhibiting values for the Sp statistic of 0.00199 for linear distance and 0.0271 for log distance. Measures of unbiased gene diversity and the inbreeding coefficient were comparable across the sampled populations. Our results provide complementary genetic data to previous demographic studies in T. recurvatum, and these findings provide data for future studies aimed at integrating the degree of clonality, genetic variation, and population dynamics in this species. Our findings suggest that T. recurvatum at the focal Meeman site displays higher levels of sexual reproduction than were previously suggested, and spatial genetic structure estimates were comparable to other plant species with mixed and outcrossing mating strategies.

Correlations between genetic, epigenetic and phenotypic variation of an introduced clonal herb.

Heritable epigenetic modifications may occur in response to environmental variation, further altering phenotypes through gene regulation, without genome sequence changes. However, epigenetic variation in wild plant populations and their correlations with genetic and phenotypic variation remain largely unknown, especially for clonal plants. We investigated genetic, epigenetic and phenotypic variation of ten populations of an introduced clonal herb Hydrocotyle vulgaris in China. Populations of H. vulgaris exhibited extremely low genetic diversity with one genotype exclusively dominant, but significantly higher epigenetic diversity. Both intra- and inter-population epigenetic variation were related to genetic variation. But there was no correlation between intra-/inter-population genetic variation and phenotypic variation. When genetic variation was controlled, intra-population epigenetic diversity was related to petiole length, specific leaf area, and leaf area variation, while inter-population epigenetic distance was correlated with leaf area differentiation. Our study provides empirical evidence that even though epigenetic variation is partly under genetic control, it could also independently play a role in shaping plant phenotypes, possibly serving as a pathway to accelerate evolution of clonal plant populations.

Effects of Temperature on Seed Germination of Invasive Plant Rorippa amphbia (L.) Besser

Aims: Global warming and biological invasion are major environmental issues faced in the world. In the study, Rorippa amphibia, a perennial invasive clone plant in northern China, was used as a material to study the germination characteristics of the seeds at different temperatures.
 Study Design: Germination test of R. amphibia seeds at different temperature was studied by means of laboratory culture. The germination percentage, germination index, germination potential, bud height and root length of the seeds were determined.
 Place and Duration of Study: Samples were collected from the west side of the swimming pool of Shenyang Agricultural University of Liaoning Province in August 2017. Experiments were done in the College of Biological Science and Technology, between October 2017 and June 2018.
 Methodology: The petri dish method was used in the experiment. Fifty seeds were randomly selected and soaked in distilled water for 12h. The seeds were placed in a petri dish covered with double filter paper, cultured at 10℃, 15℃, 20℃, 25℃, 30℃, 35℃ and 40℃ in light incubators for 12h darkness and 12h light (4000lux), with 3 repeats per processing. Seed germination was based on embryo root breakthrough seed coat ≥ 1 mm. During the experiment, the numbers of seed germination were recorded every day, and the filter paper was kept moist until there was no new seed germination for 2 consecutive days, which was regarded as the end of germination. The numbers of seed germination should be counted regularly every day, and the beginning and duration of germination should be recorded. The germination rate, daily germination rate, germinating potential, germinating index and vigor index of R. amphibia seeds at different temperature were calculated by measuring bud height and radicle length on the 10th day after germination.
 Results: The temperature range of seed germination of R. amphibia was wide, which could germinate at 15 - 40℃. Lower temperature delayed the peak period of seed germination at some extent and the germination rates of R. amphibia peak at 30 - 35℃, which were 44.67% and 50% respectively. At 35℃, germination potential and germination index were 25.33% and 29.46, reaching the maximum value.
 Conclusion: The reason for the wide temperature range of seed germination and the low germination rate might be the candidate method for clonal plant population establishment in temperate zone. The higher germination rate of high temperature condition suggested that clone invasive plants in temperate regions were more invasive during global warming.

Genet-specific DNA methylation probabilities detected in a spatial epigenetic analysis of a clonal plant population.

In sessile organisms such as plants, spatial genetic structures of populations show long-lasting patterns. These structures have been analyzed across diverse taxa to understand the processes that determine the genetic makeup of organismal populations. For many sessile organisms that mainly propagate via clonal spread, epigenetic status can vary between clonal individuals in the absence of genetic changes. However, fewer previous studies have explored the epigenetic properties in comparison to the genetic properties of natural plant populations. Here, we report the simultaneous evaluation of the spatial structure of genetic and epigenetic variation in a natural population of the clonal plant Cardamine leucantha. We applied a hierarchical Bayesian model to evaluate the effects of membership of a genet (a group of individuals clonally derived from a single seed) and vegetation cover on the epigenetic variation between ramets (clonal plants that are physiologically independent individuals). We sampled 332 ramets in a 20 m × 20 m study plot that contained 137 genets (identified using eight SSR markers). We detected epigenetic variation in DNA methylation at 24 methylation-sensitive amplified fragment length polymorphism (MS-AFLP) loci. There were significant genet effects at all 24 MS-AFLP loci in the distribution of subepiloci. Vegetation cover had no statistically significant effect on variation in the majority of MS-AFLP loci. The spatial aggregation of epigenetic variation is therefore largely explained by the aggregation of ramets that belong to the same genets. By applying hierarchical Bayesian analyses, we successfully identified a number of genet-specific changes in epigenetic status within a natural plant population in a complex context, where genotypes and environmental factors are unevenly distributed. This finding suggests that it requires further studies on the spatial epigenetic structure of natural populations of diverse organisms, particularly for sessile clonal species.

The analysis of an effect of seed propagation on defense strategy against pathogen transmission within clonal plant population using lattice model

Many clonal plants have two breeding systems, vegetative and seed propagation. In vegetative propagation, plants reproduce genetically identical offspring that have lower mortality rates. By contrast, the seed propagated offspring has higher mortality rate, however, the seed propagation acts an important role in maintaining the genetic diversity and reproduce widely. According to the experimental studies, the balance between the breeding systems, vegetative and seed propagation, is determined by several functions, such as resource allocation. The infection and spread of systemic pathogen also affect the optimal balance of the breeding systems.Thus, we examine the effect of invasion of systemic pathogen on the optimal balance of the breeding systems of clonal plant using lattice model in two cases, single population and mixed population. In the analysis, the equilibrium and its local stability were derived using approximation method and numerical simulation in single population. Additionally, two situations were assumed in mixed population, infected and uninfected populations, and the efficacy of seed propagation on the suppression of epidemic infections was examined by comparing the results in the two situations.As a result, seed propagation is an effective defensive behavior against systemic pathogens. In the single population, the plants increase their population by increasing the proportion of seed propagation when the epidemic pathogen has highly infective. In mixed population, the increasing proportion of seed propagation is the optimal breeding strategy to defend against the spread of a systemic pathogen.

The use of Hardy-Weinberg Equilibrium in clonal plant systems.

Traditionally population genetics precludes the use of the same genetic individual more than once in Hardy-Weinberg (HW) based calculations due to the model's explicit assumptions. However, when applied to clonal plant populations this can be difficult to do, and in some circumstances, it may be ecologically informative to use the ramet as the data unit. In fact, ecologists have varied the definition of the individual from a strict adherence to a single data point per genotype to a more inclusive approach of one data point per ramet. With the advent of molecular tools, the list of facultatively clonal plants and the recognition of their ecological relevance grows. There is an important risk of misinterpretation when HW calculations are applied to a clonal plant not recognized as clonal, as well as when the definition of the individual for those calculations is not clearly stated in a known clonal species. Focusing on heterozygosity values, we investigate cases that demonstrate the extreme range of potential modeling outcomes and describe the different contexts where a particular definition could better meet ecological modeling goals. We emphasize that the HW model can be ecologically relevant when applied to clonal plants, but caution is necessary in how it is used, reported, and interpreted. We propose that in known clonal plants, both genotype (GHet) and ramet (RHet) based calculations are reported to define the full range of potential values and better facilitate cross-study comparisons.