Relative contribution of canopy and soil effects between plants with different metal tolerance along a metal pollution gradient.

Multiple effects, operating either on the long-term (soil-engineering effects) or on the short-term during plant life (microclimate modification or resources pre-emption), can act simultaneously and determine the outcome of plant-plant interactions. These diverse effects have not been disentangled along a gradient of metal/metalloid pollution, although this is crucial for understanding the dominant species turnover along the gradient, and thus the driving processes of facilitation recurrently found in metalliferous ecosystems, which could help improving ecological restoration of these degraded ecosystems. Here, we experimentally assessed different short-term effects of two dominant forbs of highly polluted habitats (Hutchinsia alpina and Arenaria multicaulis, tolerant to metal stress) and two grasses of less polluted habitats (Agrostis capillaris and Festuca rubra, less tolerant to metal stress) on target plant species (the same as the dominant species mentioned above) transplanted along a large metal pollution gradient. Additionally, in highly polluted environments, we differentiated short- from long-term effects of the two metallicolous forbs, which had different abilities to concentrate metals in their leaves. In line with other studies along metal gradients, variation of short-term interactions appeared to follow the Stress Gradient Hypothesis for plants less adapted to metal pollution (p = 0.030), with positive interactions dominating in most severe areas. Regarding long-term effects, the species with highest leaf metal-accumulation showed no negative effect contrary to the Elemental allelopathy Hypothesis. Long-term effects of the species with lower leaf-metal accumulation could not be determined because of the occurrence of an unexpected difference in micro-habitat conditions (soil depth and humidity) for this species along the metal pollution gradient. Increasing short-term facilitation along metal pollution gradients, which confirmed previous studies, is promising for improving conditions and restoring the most polluted environments. However, long-term results stressed the difficulty to quantify these effects given that these areas are highly fragmented and heterogeneous.

The role of ecotypic variation for plant facilitation in a metal-polluted system: Stress-intolerant target ecotypes are the best beneficiaries and stress-tolerant nurse ecotypes the best benefactors.

Disentangling competitive-response and -effect abilities has strongly improved our understanding of the role of competition for the diversity and composition of plant communities. Much less is known about the relative importance of facilitative-effect and -response abilities in harsh ecosystems. Here, we aim to fill this gap by simultaneously assessing the facilitative-response and -effect abilities of different species and ecotypes in former mining sites in the French Pyrenees, both in naturally occurring communities and in a common-garden designed on a slag heap. The response of two ecotypes of the target species Festuca rubra with contrasting metal-stress tolerances and the facilitative effects of two ecotypes with contrasting metal-stress tolerances of four different metallicolous nurse species were assessed. The results revealed that the response of the Festuca ecotype with lower metal-stress tolerance shifted from competitive (RII = -0.24) to facilitative (RII = 0.29) as pollution increased, consistently with the stress-gradient-hypothesis. The Festuca ecotype with high metal-stress tolerance did not show any facilitative response. Regarding facilitative effect ability assessed in the common-garden, nurse ecotypes from highly polluted habitats (RII = 0.04) had a significantly higher facilitative effects than ecotypes from less polluted habitats (RII = -0.05). Metal-intolerant target ecotypes of Festuca rubra were the most sensitive to the positive effects of neighbours, while metal-tolerant nurse ecotypes were the best benefactors. Facilitative-response ability appeared to be driven by a trade-off between stress-tolerance and facilitative response of target ecotypes. In contrast, facilitative-effect ability was positively correlated to the stress-tolerance of nurse plants. The results of this study show that the highest restoration success of highly metal-stressed systems should be found when highly stress-tolerant nurse ecotypes are associated with less stress-tolerant target ecotypes.

Disentangling the effects of biomass and productivity in plant competition.

The relationship between competition and productivity in plant communities is unclear, and this is likely to be due to (1) a confusion in the literature between productivity and biomass, (2) the lack of studies assessing variation in competition in all combinations of biomass and productivity. We assessed the outcome of plant-plant interactions by removing the neighbors around five focal species in 14 herbaceous communities with contrasting biomasses and productivities: meadows with high biomass and productivity, heathlands with high biomass and low productivity, understory communities of deciduous forests with low biomass and high productivity and calcareous grasslands with low biomass and low productivity. Competition intensity was quantified with the relative interaction index (RII) calculated for both survival and growth of the transplanted targets assessed with the increase in leaf number. To examine which traits better explain variation in competition and what drives variation in diversity, we also quantified litter decomposition rate, species composition and diversity and six morphological traits related to plant size and growth rate for eight dominant species of each community. Our main questions were: (1) Is competition mostly related to biomass or productivity? (2) Which traits of the community dominants better explain variation in competition? (3) Is variation in competition and related traits correlated with variation in diversity? Competition for survival significantly increased with increasing community biomass (but not productivity). In addition, competition for survival increased with the size traits and competitive effects of the dominant species of the communities, whereas diversity decreased. Competition for growth also increased with increasing productivity, but only for high-biomass communities. Additionally, the increase in competition for growth with increasing soil fertility, as measured with litter decomposition rate, was only due to an increase in target growth in plots without neighbors and was unrelated to community competitive effects and species diversity. The results of our study illustrate how the confusion between productivity and biomass could have contributed to the long-standing debate on variation in competition along productivity gradients and its consequence for diversity.

Biomass partitioning of plants under soil pollution stress.

Polluted sites are ubiquitous worldwide but how plant partition their biomass between different organs in this context is unclear. Here, we identified three possible drivers of biomass partitioning in our controlled study along pollution gradients: plant size reduction (pollution effect) combined with allometric scaling between organs; early deficit in root surfaces (pollution effect) inducing a decreased water uptake; increased biomass allocation to roots to compensate for lower soil resource acquisition consistent with the optimal partitioning theory (plant response). A complementary meta-analysis showed variation in biomass partitioning across published studies, with grass and woody species having distinct modifications of their root: shoot ratio. However, the modelling of biomass partitioning drivers showed that single harvest experiments performed in previous studies prevent identifying the main drivers at stake. The proposed distinction between pollution effects and plant response will help to improve our knowledge of plant allocation strategies in the context of pollution.

Status of Dieldrin in vegetable growing soils across a peri-urban agricultural area according to an adapted sampling strategy.

Since the fifties, organochlorine pesticides (OCPs) had been used in agriculture to protect vegetables. Two decades after their ban by the Stockholm convention in 2001, OCPs are still present in agricultural soils inducing vegetable contamination with concentrations above Maximum Residue Level (MRL). This is a major concern for a 5 km2 peri-urban vegetable growing valley located in the south west of France. In the present work, the sampling method was developed to clarify the spatial distribution of one OCP, Dieldrin, and its relationship with soil properties at the scale of study area. A total of 99 soil samples was collected for physicochemical analyses and Dieldrin concentrations. Results show Dieldrin concentrations in soils up to 204 µg kg-1. The horizontal distribution of this pesticide is heterogeneous at the study area scale but homogeneous in each reference plot studied. About 85% of the contamination was located in the top soil layers (0-40 cm depth), but Dieldrin may still be quantified at a depth of 80 cm. Among all soil physicochemical parameters analysed, SOM was the most significantly related (P < 10-4) with Dieldrin concentrations, once different grain size fractions were considered. Moreover, results indicate a 33 times higher Dieldrin concentration and/or extractability for coarse sand than for other grain size fractions. These results show that the developed sampling method is adapted for the study area scale as it helps understanding the factors influencing the spatial distribution of Dieldrin. Historical amendments are the predominant factor for the horizontal contamination and deep ploughing for the vertical contamination. Also, the variations of coarse sand repartition in soils prevents identification of relationships between SOM and Dieldrin contamination in bulk soil. Further investigation is required to explain these relationships but these results highlight why no clear relationship between OCPs and SOM was previously identified.

Explaining the larger seed bank of an invasive shrub in non-native versus native environments by differences in seed predation and plant size.

BACKGROUND AND AIMS: Large, persistent seed banks contribute to the invasiveness of non-native plants, and maternal plant size is an important contributory factor. We explored the relationships between plant vegetative size (V) and soil seed bank size (S) for the invasive shrub Ulex europaeus in its native range and in non-native populations, and identified which other factors may contribute to seed bank variation between native and invaded regions. METHODS: We compared the native region (France) with two regions where Ulex is invasive, one with seed predators introduced for biological control (New Zealand) and another where seed predators are absent (La Réunion). We quantified seed bank size, plant dimensions, seed predation and soil fertility for six stands in each of the three regions. KEY RESULTS: Seed banks were 9-14 times larger in the two invaded regions compared to native France. We found a positive relationship between current seed bank size and actual plant size, and that any deviation from this relationship was probably due to large differences in seed predation and/or soil fertility. We further identified three possible factors explaining larger seed banks in non-native environments: larger maternal plant size, lower activity of seed predators and higher soil fertility. CONCLUSIONS: In highlighting a positive relationship between maternal plant size and seed bank size, and identifying additional factors that regulate soil seed bank dynamics in non-native ranges, our data offer a number of opportunities for invasive weed control. For non-native Ulex populations specifically, management focusing on 'S' (i.e. the reduction of the seed bank by stimulating germination, or the introduction of seed predators as biological control agents) and/or on 'V' (i.e. by cutting mature stands to reduce maternal plant biomass) offers the most probable combination of effective control options.

A multi-site approach to investigate the role of toxicity and confounding factors on plant bioassay results.

Development of organisms that live on contaminated soils depends on toxicity as well as several physical and chemical soil properties. We aimed to identify plant bioassays most responsive to contaminants and not to confounding factors due to soil type differences. We implemented a multi-site approach in seven contaminated sites and used different ordinary plant bioassays (fourteen-day-shoot biomass and five-day-root and shoot elongation). Most of the sites were contaminated with polycyclic aromatic hydrocarbons (PAHs), and soils were sampled from areas of both high and low contamination. Bioassays were performed on ninety soil samples and were carried out with six model species. We performed analyses of regulatory PAHs and their derivatives content in the samples. Fourteen-day-shoot biomass responses depended on the site's origin, with an intricate response of plants that faced contrasted soil pH and organic matter content and various contaminant levels. Five-day-shoot and root lengths were informative when considering the most heavily PAH-contaminated site, since both measures exhibited a close dose-dependent response to PAHs but not to soil pH or organic matter content. For the other sites, elongation tests revealed tenuous effects somehow related to the presence of PAHs or their derivatives. We propose that tests based on plant development during their autotrophic phase (the fourteen-day-shoot biomass test in this study) are likely more sensitive to environmental stressors but less specific for contaminant-induced effects. Comparatively, tests based on early and heterotrophic plant development could be particularly more specific for soil contaminants, but the associated responses may be of low sensitivity.

Competition along productivity gradients: news from heathlands.

The importance of competition in low productive habitats is still debated. Studies which simultaneously evaluate preemption of resources and consequences for population dynamics are needed for a comprehensive view of competitive outcomes. We cultivated two emblematic species of European heathlands (Calluna vulgaris and Molinia caerulea) in a nursery for 2 years at two fertility levels, reproducing the productivity gradient found in phosphorus (P)-depleted heathlands in southwest France. The second year, we planted Ulex europaeus seedlings, a ubiquitous heathland species, under the cover of the two species to evaluate its ability to regenerate. Half of the seedlings were placed in tubes for exclusion of competitor roots. We measured the development of the competitors aboveground and belowground and their interception of resources (light, water, inorganic P). Ulex seedlings' growth and survival were also measured. Our results on resources interception were consistent with species distribution in heathlands. Molinia, which dominates rich heathlands, was the strongest competitor for light and water in the rich soil. Calluna, which dominates poor heathlands, increased its root allocation in the poor soil, decreasing water and inorganic P availability. However, the impact of total competition and root competition on Ulex seedlings decreased in the poor soil. Other mechanisms, especially decrease of water stress under neighbouring plant cover, appeared to have more influence on the seedlings' response. We found no formal contradiction between Tilman and Grime's theories. Root competition has a primary role in acquisition of soil resources in poor habitats. However, the importance of competition decreases with decreasing fertility.

Phenotypic Plasticity in Reproductive Traits of the Perennial Shrub Ulex europaeus in Response to Shading: A Multi-Year Monitoring of Cultivated Clones.

Phenotypic plasticity may be advantageous for plants to be able to rapidly cope with new and changing environments associated with climate change or during biological invasions. This is especially true for perennial plants, as they may need a longer period to respond genetically to selective pressures than annuals, and also because they are more likely to experience environmental changes during their lifespan. However, few studies have explored the plasticity of the reproductive life history traits of woody perennial species. This study focuses on a woody shrub, Ulex europaeus (common gorse), and on the response of its reproductive traits to one important environmental factor, shading. The study was performed on clones originating from western France (within the native range of this invasive species) and grown for seven years. We compared traits of plants grown in a shade treatment (with two successive shade levels) vs. full natural light. The traits monitored included flowering onset, pod production and seed predation. All traits studied responded to shading, exhibiting various levels of plasticity. In particular, dense shade induced a radical but reversible decrease in flower and pod production, while moderate shade had little effect on reproductive traits. The magnitude of the response to dense shade depended on the genotype, showing a genetically based polymorphism of plasticity. The level of plasticity also showed substantial variations between years, and the effect of environmental variations was cumulative over time. This suggests that plasticity can influence the lifetime fitness of U. Europaeus and is involved in the capacity of the species to grow under contrasting environmental conditions.

Weak Evidence of Regeneration Habitat but Strong Evidence of Regeneration Niche for a Leguminous Shrub.

The identification of an ecological niche specific to the regeneration phase has mobilised significant attention. However, the importance of the regeneration niche concept remains unclear. Our main objective was to study the existence of such a regeneration niche for a leguminous shrub, Ulex europaeus. This study was carried out in southwest France in the context of water and nutrient stresses (mainly phosphorus limitation) due to the presence of nutrient-poor sandy soils. We analysed the regeneration of the species from the germination of seeds and emergence of new seedlings until the seedlings reached young shrub size. Our design included a P fertilisation treatment. We also investigated microsite characteristics (micro-topography and vegetation development) as they can interact with meteorological conditions and determine water availability for seeds and seedlings. We found that P availability controlled seedling growth and the time necessary to reach young shrub size. Water availability appeared to impact the species germination and seedlings survival. We also found that P and water availability depended on the interactions between microsite characteristics and climatic variations. Finally we found evidence that P and water availability are important ecological factors shaping the regeneration niche of the species, but we found weak evidence that any microsite would be appropriate for the regeneration of the species in the long term. Future studies regarding regeneration niches need to distinguish more clearly the ecological factors important for regeneration (the regeneration niche per se) and the physical world where the seedlings appear and develop (the regeneration habitat).