Why are small seeds dispersed through animal guts: large numbers or seed size per se?

Long-distance dispersal of seeds is an important process in metapopulation dynamics and in plant migrations, but at the same time extremely difficult to observe or quantify directly. If seed dispersal ability were related to attributes of seeds or motherplants, long-distance seed dispersal would be predictable by indirect approximation using easy-to-measure traits. Seed size has been suggested to be such a key trait in seed dispersal ability. However, having smaller seeds also implies having more numerous seeds per plant individual (given equal reproductive effort), and consequently increases the probability of seeds being ingested accidentally. The question is whether small-seeded species are more abundant in herbivore dung because smaller seed size increases survival rate during gut passage or because they are produced (and ingested) in greater numbers than larger seeds. We investigated endozoochorous seed dispersal via cattle grazing a meadow, and related seed abundance in dung samples to seed attributes. We found that seeds were ingested and passed through the bovine intestinal tract in proportion to the numbers produced per unit area in the grazed vegetation. In contrast, no relationship could be found between endozoochorous dispersal potential (measured as abundance of seeds in dung samples corrected for seed output in the grazed vegetation) and seed attributes such as seed mass, seed shape (roundness), and thickness of the seed coat. This finding underlines the importance of seed number in plant dispersal ability. In addition, it shows that grazing mammals may constitute an important dispersal vector for many plant species conventionally classified as 'unspecialised'.     

Plant response to disturbance in a Mediterranean grassland: How many functional groups?

Abstract. Data referring to changes in vegetation composition resulting from cattle exclosure and ploughing in a Portuguese pasture dominated by annuals were used to test hypotheses regarding the biology of species favoured or eliminated by disturbance in semi-natural herbaceous communities. These hypotheses were tested in two ways. First we compared the distribution of six a priori groups – grasses, small rosettes, large rosettes, small species with leafy stems, large species with leafy stems, legumes – across grazed, ploughed and undisturbed plots. In a second set of analyses we examined changes in the frequencies of individual biological attributes in response to grazing and ploughing. These analyses were carried out separately for grasses and dicot forbs. Overall, the species composition showed little response to either grazing or ploughing, though species dominance changed. This lack of responsiveness of species composition was attributed to the long history of intensive land use which has resulted in the loss of disturbance-intolerant species over entire landscapes. When considering a priori groups, small rosettes were indifferent to disturbance. grazing and ploughing showed that dominated. Large rosettes, large species with leafy stems and legumes were generally intolerant to both grazing and ploughing, though individual species may increase in response to disturbance. Small species with leafy stems were the only group favoured by grazing whereas ploughing favoured grasses. As to individual traits, grazing excluded large grass species with heavy seeds and promoted a flat rosette canopy structure and a small size, along with a moderate dormancy and protected inflorescences. In forbs, grazing favoured small species, as expected, while it excluded tall species, and, in contrast to earlier results, a rosette canopy. These attributes were consistent with responses of the a priori groups, though it would not have been possible to reconstruct groups directly from the attribute list. Ploughing had no effect on any of the forb traits. As to grass traits, flat- and short-statured species increased and heavy-seeded species decreased. Our analysis revealed two advantages of establishing plant functional classifications within life forms. Subgroups within forbs had contrasting types of behaviour. For the same trait patterns could differ within the grass group from within the forb group. Finally, this analysis emphasizes the need for plant functional classifications aiming at the identification of syndromes of co-occurring attributes rather than of lists of isolated traits of which actual combinations are not specified.     

Size traits and site conditions determine changes in seed bank structure caused by grazing exclusion in semiarid annual plant communities

1. Contrasting patterns of change in the seed bank of natural grasslands are frequently found in response to grazing by domestic herbivores. Here, we studied the hypotheses that a) patterns of change in seed bank density and composition in response to grazing depend on spatial variation in resource availability and productivity, and b) that variation among species in patterns of seed bank response to grazing is linked to differences in species size traits (i.e. size of plant, dispersal unit and seed). 2. Effects of sheep grazing exclusion on the seed bank were followed during five years in a semiarid Mediterranean annual plant community in Israel. Seed bank density and composition were measured in autumn, before the rainy season, inside and outside fenced exclosures in four neighboring topographic sites differing in vegetation characteristics, soil resources and primary productivity: Wadi (dry stream terraces, high productive site), Hilltop, South‐ and North‐facing slopes (less productive sites). 3. Topographic sites differed in seed density (range ca 2500–18000 seed m^?2) and in seed bank response to grazing exclusion. Fencing increased seed density by 78, 51 and 18% in the Wadi, South‐ and North‐facing slopes, respectively, but had no effect in the Hilltop. At the species level, grazing exclusion interacted with site conditions in determining species seed bank density, with larger or opposite changes in the high productive Wadi compared to the other less productive sites. 4. Changes in seed bank structure after grazing exclusion were strongly related to species size traits. Grazing exclusion favored species with large size traits in all sites, while seed density of tiny species decreased strongly in the high productive Wadi. Species with medium and small size traits showed lesser or no responses. 5. The size of plants, dispersal units and seeds were strongly correlated to each other, thus confounding the evaluation of the relative importance of each trait in the response of species to grazing and site conditions. We propose that the relative importance of plant size vs seed size in the response to grazing changes with productivity level.     

Severely degraded dunes of the southern Kalahari: local extinction,persistence and natural re‐establishment of plants

This study aimed to quantify and understand the impact of severe land degradation on plant attributes and diversity on dunes of the southern Kalahari. Heavy grazing pressure in particular resulted in a significant decline of canopy cover and species number in annual and perennial life forms; forb and graminoid growth forms; erect and prostrate habits; and leafy stem, tussock and stoloniferous architectures. However, no significant change was found in shrub and tree forms which persisted without apparent new recruitment. Under these conditions, species diversity dropped sharply and a number of species, mainly graminoids, became apparently locally extinct. The perennial shrub, Crotalaria cf. spartioides, showed the converse with a high frequency of establishing seedlings on the degraded dunes. Changes in relative dominance show that the extreme treatment favours perennial over annual, woody over graminoid and forb, erect over prostrate and leafy stem over stoloniferous and tussock. Some of these results and certain species and soil responses differ from those reported from grazing studies elsewhere, and are possibly no longer directly related to the impact of the primary grazing pressure but to the secondary effect of subsequent instability of the dunes.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

Flowering phenology and reproduction of a forest understorey plant species in response to the local environment

Temperate forest understorey plants are subjected to a strong seasonality in their optimal growing conditions. In winter and early spring, low temperatures are suboptimal for plant growth while light becomes limited later in spring season. We can thus expect that differences in plant phenology in relation to spatiotemporal environmental variation will lead to differences in reproductive output, and hence selection. We specifically studied whether early flowering, a paradoxical pattern that is observed in many plant species, is an adaptive strategy, and whether selection for early flowering was confounded with selection for flower duration or was attributable to environmental variables. We used Geum urbanum as a study species to investigate the effect of relevant environmental factors on the species’ flowering phenology and the consequences for plant reproductive output. We monitored the phenology of four to six plants in each of ten locations in a temperate deciduous forest (Belgium). We first quantified variation in flowering time within individuals and related this temporal variation to individual flower reproductive output. Then, we studied inter-individual variation here-in and linked this to reproduction at the plant level, hence studying the selection differential. We found that flowering within individual plants of Geum urbanum was spread over a long period from June to October. Reproductive output of individual flowers, measured as total seed mass per flower, declined during the season. We found no indication for selection for early flowering but rather for longer flower duration. Larger plants had an earlier flowering onset and a higher seed mass, which suggests that these factors covary and are condition dependent. None of the studied environmental variables could explain plant size, although soil pH and to a lesser extent light availability had a positive direct effect on seed mass per plant. Finally, we suggest that the high intra-individual variation in flowering time, which might be a risk spreading strategy of the plant in the presence of seed predation, limits the potential for selection on flowering phenology.

     

Phenological responses to nitrogen and water addition are linked to plant growth patterns in a desert herbaceous community

Increases in nitrogen (N) deposition and variation in precipitation have been occurring in temperate deserts; however, little information is available regarding plant phenological responses to environmental cues and their relationships with plant growth pattern in desert ecosystems. In this study, plant phenology and growth of six annuals in response to N and water addition were monitored throughout two consecutive growing seasons in 2011 and 2012 in a temperate desert in northwestern China. The effects of N and water addition on reproductive phenology differed among plant species. N and water addition consistently advanced the flowering onset time and fruiting time of four spring ephemerals; however, their effects on two spring‐summer annuals were inconsistent, with advances being noted in one species and delays in another. N and water addition alone increased plant height, relative growth rate, leaf number, flower number, and individual biomass, while their combinative effects on plant growth and reproductive phenology were dependent on species. Multiple regression analysis showed that flowering onset time was negatively correlated with relative growth rate of two species, and negatively correlated with maximum plant height of the other four species. Our study demonstrates that phenological responses to increasing precipitation and N deposition varied in annuals with different life histories, whereby the effects of climate change on plant growth rate were related to reproductive phenology. Desert annuals that were able to accelerate growth rate under increasing soil resource availability tended to advance their flowering onset time to escape drought later in the growing season. This study promotes our understanding of the responses of temperate desert annuals to increasing precipitation and N deposition in this desert.     

Plant phenology,leaf traits and leaf litterfall of contrasting life forms in the arid Patagonian Monte,Argentina

Question: Do coexisting plant life forms differ in overall phenology, leaf traits and patterns of leaf litterfall? Location: Patagonian Monte, Chubut Province, Argentina. Methods: We assessed phenology, traits of green and senesced leaves and the pattern of leaf litterfall in 12 species of coexisting life forms (perennial grasses, deciduous shrubs, evergreen shrubs). Results: We did not identify differences in phenology, leaf traits and patterns of leaf litterfall among life forms but these attributes contrasted among species. Independent of the life form, the maintenance of green leaves or vegetative growth during the dry season was mostly associated with leaves with high leaf mass per area (LMA) and high concentration of secondary compounds. Low LMA species produced low litterfall mass with low concentration of secondary compounds, and high N concentration. High LMA species produced the largest mass of leaf litterfall. Accordingly, species were distributed along two main dimensions of ecological variation, the dimension secondary compounds in leaves ‐ length and timing of the vegetative growth period (SC ‐ VGP) and the dimension leaf mass per area ‐ leaf litterfall mass (LMA ‐ LLM). Conclusions: Phenology, leaf traits and leaf litterfall varied among species and overlapped among life forms. The two dimensions of ecological variation among species (SC ‐ VGP, LMA ‐ LLM) represent distinct combinations of plant traits or strategies related to resource acquisition and drought tolerance which are reflected in the patterns of leaf litterfall.     

Body size and measurement of species diversity in large grazing mammals

Species are by definition different from each other. This fact favours ranking rather than additive indices. However, ecologists have measured species diversity in terms of species richness, or by combining species richness with the relative abundance of species within an area. Both methods raise problems: species richness treats all species equally, while relative abundance is not a fixed property of species but varies widely temporally and spatially, and requires a massive sampling effort. The functional aspect of species diversity measurement may be strengthened by incorporating differences between species such as body size as a component of diversity. An index of diversity derived from a measure of variation in body size among species is proposed for large grazing mammals. The proposed diversity index related positively to species abundance, indicating that the use of body size as a surrogate for diversity is adequate. Because the proposed index is based on presence or absence data, the expensive and time consuming counting of individuals per species in each sampling unit is not necessary.     

Determining suitable grazing time for Puccinella distans Parl. based on its phenology in West Azerbaijan Province of Iran

Studying the phenology, which is an important issue of plants, can be used for determining the best time of all kinds of utilizations, and also conservation and stability of plants’ growth and breeding will result. Therefore, this study was performed in Tezkharab rangelands, where Puccinella distans is an important vegetative element that is consumed by livestock, to determine the suitable grazing time for this plant based on its phenology, for about a 4-year period (2007–2010). For this purpose, plant phenology and changes in its growth characteristics were investigated weekly. Moreover, during this period, all phenological stages were adjusted to the monthly precipitation and temperature. The results indicated that the time of phenological stages had fluctuations which were related to environmental conditions. The best time to start grazing is early May and the best time for collecting the seeds of this species is late July to early August.     

Introduced and native herbivores have different effects on plant composition in low productivity ecosystems

Questions

Understanding how livestock grazing alters plant composition in low productivity environments is critical to managing livestock sustainably alongside native and introduced wild herbivore populations. We asked four questions: (1) does recent livestock and rabbit grazing reduce some plant attributes more strongly than others; (2) does grazing by introduced herbivores (i.e. livestock and rabbits) affect plants more strongly than native herbivores (i.e. kangaroos); (3) do the effects of recent livestock grazing differ from the legacy effects of livestock grazing; and (4) does the probability of occurrence of exotic plants increase with increasing net primary productivity (NPP)?

Location

South‐eastern Australia.

Methods

We measured the recent grazing activity of co‐occurring livestock (cattle, sheep, goats), rabbits and kangaroos by counting faecal pellets; historic grazing activity by measuring livestock tracks; and derived NPP from satellite imagery. We used a hierarchical GLMM to simultaneously model the presence or absence (i.e. probability of occurrence) of all plant species as a function of their attributes (growth form, lifespan and origin) to assess their average response to recent grazing, historic grazing and productivity in a broad‐scale regional study.

Results

Recent and historic livestock grazing, rabbit grazing and increasing NPP reduced the average probability of occurrence of plant species, although responses varied among plant attributes. Both recent and historic livestock grazing strongly reduced the average probability of occurrence of native species, and forbs and geophytes, but differed in their relative effects on other growth forms. Recent livestock grazing, rabbit grazing and NPP had similar effects, strongly reducing native species and forbs, geophytes, shrubs and sub‐shrubs. The overall effects of recent kangaroo grazing were relatively weak, with no clear trends for any given plant attribute.

Conclusion

Our results highlight the complex nature of grazing by introduced herbivores compared with native herbivores on different plant attributes. Land managers need to be aware that domestic European livestock, rabbits and other free‐ranging introduced livestock such as goats have detrimental impacts on native plant communities. Our results also show that kangaroo grazing has a relatively benign effect on plant occurrence.     

ENVIRONMENTAL AND GENETIC DETERMINANTS OF PLANT SIZE IN VIOLA SORORIA

Survivorship and fecundity in the forest herb, Viola sororia, are size-dependent. The basis of size variation among individuals of Viola sororia was investigated with a uniform environment experiment. Plants collected from natural populations were vegetatively reproduced and grown under two light regimes in a greenhouse. Analysis of quantitative variation showed: 1) significant differences between light treatments for characters related to plant shape and relative growth rate; 2) significant among-genet variation for plant size, plant shape and relative growth rate but none for physiological characters; and 3) a size threshold for cleistogamous seed production and rhizome production. Heritability estimates for the characters associated with plant size and shape ranged from 0.09 to 0.39, indicating significant genetic determination for these traits. In addition, among-genet differences in relative growth rate were substantial. The results of this study suggest that the size variation found in natural populations is not solely a function of environmental heterogeneity but is significantly influenced by the genotypes composing the population.     

Plant functional traits under cattle grazing and fallow age scenarios in a tropical dry forest of Northwestern Mexico

Evaluating plant functional traits helps to understand how plants respond to changing environmental conditions and resource availability associated with disturbance events. Livestock production is one of the primary drivers of tropical forest loss and degradation. Livestock alter environmental conditions within the forest by grazing, trampling and nutrient inputs, which in turn can influence species composition and functional traits of species. Understanding how livestock influence functional traits along a successional gradient is poorly understood. Here, we studied the effect of cattle grazing and fallow age on plant functional traits and soil nutrients in secondary and old-growth tropical dry forests. We analyzed plant functional traits of the most important species in successional and old-growth forest communities in both cattle present and cattle excluded plots. Our results showed the effects of cattle grazing and fallow age on plant functional traits, with fallow age explaining more variation than cattle grazing. In early succession, functional traits were associated with water conservation (thicker leaves, lower specific leaf area), and in later successional they were linked with sunlight conservation (larger height, higher specific leaf area). The presence of large fruits and seeds in advanced successional sites suggests high resource availability, which may help plants to successfully reproduce. Moreover, under cattle grazing some functional traits are associated with herbivory defense (high foliar dry weight and thick leaves). Even though N and C increased as succession advanced, the sites with cattle grazing had higher NH4 and NO3 concentrations as a result of fecal deposition. Plant functional traits responded to fallow age than to cattle grazing. Our study showed that cattle exclusion, as a management and biodiversity conservation strategy, contributes positively to soil nutrition. Thus, fallow age and cattle exclusion facilitate soil recovery and allows establishing species with suitable functional attributes for overcoming environmental filters in abandoned cattle fields.     

How do host plant use and seasonal life cycle relate to insect body size: A case study on European geometrid moths (Lepidoptera: Geometridae)

We used European geometrid moths (>630 species) as a model group to investigate how life history traits linked to larval host plant use (i.e., diet breadth and host-plant growth form) and seasonal life cycle (i.e., voltinism, overwintering stage and caterpillar phenology) are related to adult body size in holometabolous insect herbivores. To do so, we applied phylogenetic comparative methods to account for shared evolutionary history among herbivore species. We further categorized larval diet breadth based on the phylogenetic structure of utilized host plant genera. Our results indicate that species associated with woody plants are, on average, larger than herb feeders and increase in size with increasing diet breadth. Obligatorily univoltine species are larger than multivoltine species, and attain larger sizes when their larvae occur exclusively in the early season. Furthermore, the adult body size is significantly smaller in species that overwinter in the pupal stage compared to those that overwinter as eggs or caterpillars. In summary, our results indicate that the ecological niche of holometabolous insect herbivores is strongly interrelated with body size at maturity.     

Differences in species richness and life-history traits between grazed and abandoned grasslands in southern Sweden

Disturbance has profound effects on plant community composition. This paper deals with the influence of grazing on species richness and proportions of life‐history attributes of grassland vegetation at six spatial scales (0.001–1000 m²) in two provinces of southern Sweden. The study comprised 33 dry grassland sites, including 22 grazed and 11 abandoned localities, and 28 sites of coastal brackish meadows, divided into five management types (from “heavily grazed” to “abandoned since long time”).
In general grazed sites were species‐richer than abandoned sites, especially at small plot sizes. However, there was a steeper increase in species number towards larger plot sizes in the abandoned sites. Heavy grazing in the coastal meadows resulted in a comparatively low number of species, corroborating the intermediate disturbance hypothesis.
The analysis of life‐history traits indicated the importance of taxonomic group, canopy structure, height, regenerative strategy and, in particular, life form. Leaf anatomy and seed dispersal seemed to be less important. The responses to grazing as regards species traits differed somewhat between grassland types. Grazed sites generally had high proportions of legumes, therophytes, species with basal position of leaves and with regeneration by means of a persistent seed bank. Abandonment of grazing favoured monocots, geophytes, species with vegetative regeneration and (partly) leafy canopy structure. Some differences between grazed and abandoned sites were confined to either the smallest or largest plot sizes, indicating different responses of matrix and interstitial species. Various positive associations (attribute syndromes) or negative associations between individual traits were identified. There was, for example, a positive link between the attributes “geophytes” and “ability of vegetative regeneration”. The recognition of such links is important to avoid misinterpreting certain attributes as functional adaptations to grazing while they are only positively correlated to other attributes of larger significance.     

Contrasting Effects of Intraspecific Trait Variation on Trait-Based Niches and Performance of Legumes in Plant Mixtures

Niche differentiation, assumed to be a key mechanism of species coexistence, requires that species differ in their functional traits. So far it remains unclear to which extent trait plasticity leads to niche shifts of species at higher plant diversity, thereby increasing or decreasing niche overlap between species. To analyse this question it is convenient to measure niches indirectly via the variation in resource-uptake traits rather than directly via the resources used. We provisionally call these indirectly measured niches trait-based niches. We studied shoot- and leaf-morphological characteristics in seven legume species in monoculture and multi-species mixture in experimental grassland. Legume species varied in the extent of trait variation in response to plant diversity. Trait plasticity led to significant shifts in species niches in multiple dimensions. Single-species niches in several traits associated with height growth and filling of canopy space were expanded, while other niche dimensions were compressed or did not change with plant diversity. Niche separation among legumes decreased in dimensions related to height growth and space filling, but increased in dimensions related to leaf size and morphology. The total extent of occupied niche space was larger in mixture than in the combined monocultures for dimensions related to leaf morphology and smaller for dimensions related to whole-plant architecture. Taller growth, greater space filling and greater plasticity in shoot height were positively, while larger values and greater plasticity in specific leaf area were negatively related with increased performance of species in mixture. Our study shows that trait variation in response to plant diversity shifts species niches along trait axes. Plastically increased niche differentiation is restricted to niche dimensions that are apparently not related to size-dependent differences between species, but functional equivalence (convergence in height growth) rather than complementarity (divergence in traits associated with light acquisition) explains increased performance of legumes in mixture.     

Nurse plant effects on plant species richness in drylands: The role of grazing,rainfall and species specificity

The outcome of plant–plant interactions depends on environmental (e.g. grazing and climatic conditions) and species-specific attributes (e.g. life strategy and dispersal mode of the species involved). However, the joint effects of such factors on pairwise plant–plant interactions, and how they modulate the role of these interactions at the community level, have not been addressed before. We assessed how these species-specific (life strategy and dispersal) and environmental (grazing and rainfall) factors affected the co-occurrence of 681 plant species pairs on open woodlands in south-eastern Australia. Species-specific attributes affected the co-occurrence of most species pairs, with higher co-occurrence levels dominating for drought-intolerant species. The dispersal mechanism only affected drought-tolerant beneficiaries, with more positive co-occurrences for vertebrate-dispersed species. Conversely, the percentage of facilitated species at the community scale declined under higher rainfall availabilities. A significant grazing × rainfall interaction on the percentage of facilitated species suggests that grazing-mediated protection was important under low to moderate, but not high, rainfall availabilities. This study improves our ability to predict changes in plant–plant interactions along environmental gradients, and their effect on community species richness, by highlighting that: (1) species-specific factors were more important than environmental conditions as drivers of a large amount (～37%) of the pairwise co-occurrences evaluated; (2) grazing and rainfall interaction drive the co-occurrence among different species in the studied communities, and (3) the effect of nurse plants on plant species richness will depend on the relative dominance of particular dispersal mechanisms or life strategies prone to be facilitated.     

Genome size and DNA base composition of geophytes: the mirror of phenology and ecology?

Background and Aims

Genome size is known to affect various plant traits such as stomatal size, seed mass, and flower or shoot phenology. However, these associations are not well understood for species with very large genomes, which are laregly represented by geophytic plants. No detailed associations are known between DNA base composition and genome size or species ecology.

Methods

Genome sizes and GC contents were measured in 219 geophytes together with tentative morpho-anatomical and ecological traits.

Key Results

Increased genome size was associated with earliness of flowering and tendency to grow in humid conditions, and there was a positive correlation between an increase in stomatal size in species with extremely large genomes. Seed mass of geophytes was closely related to their ecology, but not to genomic parameters. Genomic DNA GC content showed a unimodal relationship with genome size but no relationship with species ecology.

Conclusions

Evolution of genome size in geophytes is closely related to their ecology and phenology and is also associated with remarkable changes in DNA base composition. Although geophytism together with producing larger cells appears to be an advantageous strategy for fast development of an organism in seasonal habitats, the drought sensitivity of large stomata may restrict the occurrence of geophytes with very large genomes to regions not subject to water stress.     

Plasticity to wind is modular and genetically variable in Arabidopsis thaliana

Thigmomorphogenesis, the characteristic phenotypic changes by which plants react to mechanical stress, is a widespread and probably adaptive type of phenotypic plasticity. However, little is known about its genetic basis and population variation. Here, we examine genetic variation for thigmomorphogenesis within and among natural populations of the model system Arabidopsis thaliana. Offspring from 17 field-collected European populations was subjected to three levels of mechanical stress exerted by wind. Overall, plants were remarkably tolerant to mechanical stress. Even high wind speed did not significantly alter the correlation structure among phenotypic traits. However, wind significantly affected plant growth and phenology, and there was genetic variation for some aspects of plasticity to wind among A. thaliana populations. Our most interesting finding was that phenotypic traits were organized into three distinct and to a large degree statistically independent covariance modules associated with plant size, phenology, and growth form, respectively. These phenotypic modules differed in their responsiveness to wind, in the degree of genetic variability for plasticity, and in the extent to which plasticity affected fitness. It is likely, therefore, that thigmomorphogenesis in this species evolves quasi-independently in different phenotypic modules.