Species phylogeny, ecology, and root traits as predictors of root exudate composition.

Root traits including root exudates are key factors affecting plant interactions with soil and thus play an important role in determining ecosystem processes. The drivers of their variation, however, remain poorly understood. We determined the relative importance of phylogeny and species ecology in determining root traits and analyzed the extent to which root exudate composition can be predicted by other root traits. We measured different root morphological and biochemical traits (including exudate profiles) of 65 plant species grown in a controlled system. We tested phylogenetic conservatism in traits and disentangled the individual and overlapping effects of phylogeny and species ecology on traits. We also predicted root exudate composition using other root traits. Phylogenetic signal differed greatly among root traits, with the strongest signal in phenol content in plant tissues. Interspecific variation in root traits was partly explained by species ecology, but phylogeny was more important in most cases. Species exudate composition could be partly predicted by specific root length, root dry matter content, root biomass, and root diameter, but a large part of variation remained unexplained. In conclusion, root exudation cannot be easily predicted based on other root traits and more comparative data on root exudation are needed to understand their diversity.

The role of plant-soil feedback in long-term species coexistence cannot be predicted from its effects on plant performance.

BACKGROUND: Despite many studies on the importance of competition and plants' associations with mutualists and pathogens on plant performance and community organization, the joint effects of these two factors remain largely unexplored. Even less is known about how these joint effects vary through a plant's life in different environmental conditions and how they contribute to the long-term coexistence of species. METHODS: We investigated the role of plant-soil feedback (PSF) in intra- and interspecific competition, using two co-occurring dry grassland species as models. A two-phase PSF experiment was used. In the first phase, soil was conditioned by the two plant species. In the second, we assessed the effect of soil conditioning, competition and drought stress on seedling establishment, plant growth in the first and second vegetation season, and fruit production. We also estimated effects of different treatments on overall population growth rates and predicted the species' potential coexistence. RESULTS: Soil conditioning played a more important role in the early stages of the plants' life (seedling establishment and early growth) than competition. Specifically, we found strong negative intraspecific PSF for biomass production in the first year in both species. Although the effects of soil conditioning persisted in later stages of plant's life, competition and drought stress became more important. Surprisingly, models predicting species coexistence contrasted with the effects on individual life stages, showing that our model species benefit from their self-conditioned soil in the long run. CONCLUSIONS: We provide evidence that the effects of PSF vary through plants' life stages. Our study suggests that we cannot easily predict the effects of soil conditioning on long-term coexistence of species using data only on performance at a single time as commonly done in PSF studies. We also show the importance of using as realistic environmental conditions as possible (such as drought stress experienced in dry grasslands) to draw reasonable conclusions on species coexistence.

Pre- and postzygotic mechanisms preventing hybridization in co-occurring species of the Impatiens purpureoviolacea complex.

In the species-rich genus Impatiens, few natural hybrids are known, even though closely related species often occur sympatrically. In this study, we aim to bridge the gap between micro- and macro-evolution to disentangle pre- and postzygotic mechanisms that may prevent hybridization in the Impatiens purpureoviolacea complex from Central Africa. We analyzed habitat types, species distribution, pollination syndromes, pollinator dependency, genome sizes, and chromosome numbers of seven out of the ten species of the complex as well as of one natural hybrid and reconstructed the ancestral chromosome numbers of the complex. Several species of the complex occur in sympatry or geographically very close to each other. All of them are characterized by pre- and/or postzygotic mechanisms potentially preventing hybridization. We found four independent polyploidization events within the complex. The only known natural hybrid always appears as single individual and is self-fertile. But the plants resulting from self-pollinated seeds often die shortly after first flowering. These results indicate that the investigated mechanisms in combination may effectively but not absolutely prevent hybridization in Impatiens and probably occur in other genera with sympatric species as well.

Plant palatability and trait responses to experimental warming.

Climate warming is expected to significantly affect plant-herbivore interactions. Even though direct effects of temperature on herbivores were extensively studied, indirect effects of temperature (acting via changes in host plant quality) on herbivore performance have rarely been addressed. We conducted multiple-choice feeding experiments with generalist herbivore Schistocerca gregaria feeding on six species of genus Impatiens cultivated at three different temperatures in growth chambers and a common garden. We also studied changes in leaf morphology and chemistry. We tested effects of temperature on plant palatability and assessed whether the effects could be explained by changes in the leaf traits. The leaves of most Impatiens species experienced the highest herbivory when cultivated at the warmest temperature. Traits related to leaf morphology (specific leaf area, leaf dry matter content and leaf area), but not to leaf chemistry, partly mediated the effects of temperature on plant palatability. Herbivores preferred smaller leaves with lower specific leaf area and higher leaf dry matter content. Our study suggests that elevated temperature will lead to changes in leaf traits and increase their palatability. This might further enhance the levels of herbivory under the increased herbivore pressure, which is forecasted as a consequence of climate warming.

Plant Origin, but Not Phylogeny, Drive Species Ecophysiological Response to Projected Climate.

Knowledge of the relationship between environmental conditions and species traits is an important prerequisite for understanding determinants of community composition and predicting species response to novel climatic conditions. Despite increasing number of studies on this topic, our knowledge on importance of genetic differentiation, plasticity and their interactions along larger sets of species is still limited especially for traits related to plant ecophysiology. We studied variation in traits related to growth, leaf chemistry, contents of photosynthetic pigments and activity of antioxidative enzymes, stomata morphology and photosynthetic activity across eight Impatiens species growing along altitudinal gradients in Himalayas cultivated in three different temperature regimes and explored effects of among species phylogenetic relationships on the results. Original and target climatic conditions determine trait values in our system. The traits are either highly plastic (e.g., APX, CAT, plant size, neoxanthin, ß-carotene, chlorophyll a/b, DEPSC) or are highly differentiated among populations (stomata density, lutein production). Many traits show strong among population differentiation in degree of plasticity and direction in response to environmental changes. Most traits indicate that the species will profit from the expected warming. This suggests that different processes determine the values of the different traits and separating the importance of genetic differentiation and plasticity is crucial for our ability to predict species response to future climate changes. The results also indicate that evolution of the traits is not phylogenetically constrained but including phylogenetic information into the analysis may improve our understanding of the trait-environment relationships as was apparent from the analysis of SLA.

Altitude, habitat type and herbivore damage interact in their effects on plant population dynamics.

Insects represent one of the most abundant groups of herbivores, and many of them have significant impacts on the dynamics of plant populations. As insects are very sensitive to changes in climatic conditions, we hypothesize that their effects on plant population dynamics will depend on climatic conditions. Knowledge of the variation in herbivore effects on plant population dynamics is, however, still rather sparse. We studied population dynamics and herbivore damage at the individual plant level of Salvia nubicola along a wide altitudinal gradient representing a range of climatic conditions. Using integral projection models, we estimated the effect of changes in herbivore pressure on plant populations in different climates and habitat types. Since we recorded large differences in the extent of herbivore damage along the altitudinal gradient, we expected that the performance of plants from different altitudes would be affected to different degrees by herbivores. Indeed, we found that populations from low altitudes were better able to withstand increased herbivore damage, while populations from high altitudes were suppressed by herbivores. However, the pattern described above was evident only in populations from open habitats. In forest habitats, the differences in population dynamics between low and high altitudes were largely diminished. The effects of herbivores on plants from different altitudes were thus largely habitat specific. Our results indicate potential problems for plant populations from high altitudes in open habitats because of increased herbivore damage. However, forest habitats may provide refuges for the plants at these high altitudes.

Arbuscular mycorrhizal fungi and associated microbial communities from dry grassland do not improve plant growth on abandoned field soil.

After abandonment of agricultural fields, some grassland plant species colonize these sites with a frequency equivalent to dry grasslands (generalists) while others are missing or underrepresented in abandoned fields (specialists). We aimed to understand the inability of specialists to spread on abandoned fields by exploring whether performance of generalists and specialists depended on soil abiotic and/or biotic legacy. We performed a greenhouse experiment with 12 species, six specialists and six generalists. The plants were grown in sterile soil from dry grassland or abandoned field inoculated with microbial communities from one or the other site. Plant growth, abundance of mycorrhizal structures and plant response to inoculation were evaluated. We focused on arbuscular mycorrhizal fungi (AMF), one of the most important parts of soil communities affecting plant performance. The abandoned field soil negatively affected plant growth, but positively affected plant response to inoculation. The AMF community from both sites differed in infectivity and taxa frequencies. The lower AMF taxa frequency in the dry grassland soil suggested a lack of functional complementarity. Despite the fact that dry grassland AMF produced more arbuscules, the dry grassland inoculum did not improve phosphorus nutrition of specialists contrary to the abandoned field inoculum. Inoculum origin did not affect phosphorus nutrition of generalists. The lower effectiveness of the dry grassland microbial community toward plant performance excludes its inoculation in the abandoned field soil as a solution to allow settlement of specialists. Still, the distinct response of specialists and generalists to inoculation suggested that they differ in AMF responsiveness.

Duration of the conditioning phase affects the results of plant-soil feedback experiments via soil chemical properties.

Plant-soil feedback (PSF) is a fundamental mechanism explaining plant community composition. Two-phase experiments, i.e., conditioning and feedback, represent a common methodology to study PSF. The duration of the conditioning phase varies among studies and the PSF observed is often explained by its biotic component. Little is known about the temporal variation of PSF and its abiotic component. As early life stages are crucial for plant establishment, we grew Rorippa austriaca in soil conditioned over 2, 4, 6 or 8 weeks by a conspecific or a co-occurring species, Agrostis capillaris. For each conditioning duration, we analysed the soil chemical properties and the direction and intensity of intra- or inter-specific feedbacks. With increasing duration, the negative intra- and inter-specific feedbacks became stronger and weaker, respectively. The inter-specific feedback was more negative than the intra-specific feedback at 2 weeks and this reversed thereafter. The Mg content decreased with conditioning duration whatever the conditioning species was. With increasing duration, conditioning by R. austriaca strongly decreased pH, while A. capillaris did not affect pH. The K and P contents were not affected by the conditioning duration and were higher in R. austriaca soil than in A. capillaris soil. Our results suggest that not only conditioning species but also duration of conditioning phase may affect the magnitude of PSF. The changes in soil chemical properties linked to the conditioning species or the conditioning phase duration may drive the feedbacks by affecting plant growth directly or via the interacting microbial communities.

Evolutionary responses to climate change in a range expanding plant.

To understand the biological effects of climate change, it is essential to take into account species' evolutionary responses to their changing environments. Ongoing climate change is resulting in species shifting their geographical distribution ranges poleward. We tested whether a successful range expanding plant has rapidly adapted to the regional conditions in its novel range, and whether adaptation could be driven by herbivores. Furthermore, we investigated if enemy release occurred in the newly colonized areas and whether plant origins differed in herbivore resistance. Plants were cloned and reciprocally transplanted between three experimental sites across the range. Effects of herbivores on plant performance were tested by individually caging plants with either open or closed cages. There was no indication of (regional) adaptation to abiotic conditions. Plants originating from the novel range were always larger than plants from the core distribution at all experimental sites, with or without herbivory. Herbivore damage was highest and not lowest at the experimental sites in the novel range, suggesting no release from enemy impact. Genotypes from the core were more damaged compared to genotypes from newly colonized areas at the most northern site in the novel range, which was dominated by generalist slug herbivory. We also detected subtle shifts in chemical defenses between the plant origins. Genotypes from the novel range had more inducible defenses. Our results suggest that plants that are expanding their range with climate change may evolve increased vigor and altered herbivore resistance in their new range, analogous to invasive plants.

Trade-off among different anti-herbivore defence strategies along an altitudinal gradient.

The type and intensity of plant-herbivore interactions are likely to be altered under climate change as a consequence of differential dispersal rates of plants and their herbivores. Here, we studied variation in herbivore damage on Salvia nubicola in the field and compared its growth and defence strategies against herbivores in controlled conditions using seeds from populations along a broad altitudinal gradient. Our work is one of the first studies to simultaneously measure complex intraspecific variation in plant growth, direct and indirect defences as well as plant tolerance (ability to regrow) as a consequence of herbivore attack simulated by clipping. In the field, we found that plants experienced higher herbivore pressure in lower altitudes. In the greenhouse, plants grown from seeds collected in lower-altitude populations grew better and produced a higher content of phenolic compounds (direct defence) and volatile organic compounds (indirect defence) in response to simulated herbivory. However, there were no differences in tolerance and effect of S. nubicola extracts on the model generalist herbivore Spodoptera littoralis (direct defence) along the altitudinal gradient. Although we found that S. nubicola developed a range of defence strategies, the strategies do not seem to be used simultaneously in all populations even though most of them are correlated with altitudinal gradient. Our finding is in agreement with the current knowledge that co-expression of multiple defences might be costly for a plant, since investment in defensive traits is assumed to reduce the resource availability for growth and reproduction. Our study thus shows the importance of simultaneous study of different defence strategies since understanding these trade-offs could be necessary for detecting the mechanisms by which plants are able to cope with future climate change.

The effect of AMF suppression on plant species composition in a nutrient-poor dry grassland.

Arbuscular mycorrhizal fungi (AMF) are expected to be one of the key drivers determining the diversity of natural plant communities, especially in nutrient-poor and dry habitats. Several previous studies have explored the importance of AMF for the composition of plant communities in various types of habitats. Surprisingly, studies of the role of AMF in nutrient-poor dry grassland communities dominated by less mycotrophic plant species are still relatively rare. We present the results of a 3-year study in which a plant community in a species-rich dry grassland was subjected to the fungicide carbendazim to suppress AMF colonization. We tested the effect of the fungicide on the following parameters: the plant species composition; the number of plant species; the cover of the rare, highly mycorrhiza-dependent species Aster amellus; the cover of the dominant, less mycorrhiza-dependent species Brachypodium pinnatum; and the cover of graminoids and perennial forbs. In addition, we examined the mycorrhizal inoculation potential of the soil. We found that the suppression of AMF with fungicide resulted in substantial changes in plant species composition and significant decrease in species richness, the cover of A. amellus and the cover of perennial forbs. In contrast the species increasing their cover after fungicide application were graminoids--the C3 grasses B. pinnatum and Bromus erectus and the sedge Carex flacca. These species appear to be less mycorrhiza dependent. Moreover, due to their clonal growth and efficient nutrient usage, they are, most likely, better competitors than perennial forbs under fungicide application. Our results thus suggest that AMF are an essential part of the soil communities supporting a high diversity of plant species in species-rich dry grasslands in nutrient-poor habitats. The AMF are especially important for the maintenance of the populations of perennial forbs, many of which are rare and endangered in the area.