Stefan C. Dekker

Divergent composition but similar function of soil food webs of individual plants: plant species and community effects

Soils are extremely rich in biodiversity, and soil organisms play pivotal roles in supporting terrestrial life, but the role that individual plants and plant communities play in influencing the diversity and functioning of soil food webs remains highly debated. Plants, as primary producers and providers of resources to the soil food web, are of vital importance for the composition, structure, and functioning of soil communities. However, whether natural soil food webs that are completely open to immigration and emigration differ underneath individual plants remains unknown. In a biodiversity restoration experiment we first compared the soil nematode communities of 228 individual plants belonging to eight herbaceous species. We included grass, leguminous, and non-leguminous species. Each individual plant grew intermingled with other species, but all plant species had a different nematode community. Moreover, nematode communities were more similar when plant individuals were growing in the same as compared to different plant communities, and these effects were most apparent for the groups of bacterivorous, carnivorous, and omnivorous nematodes. Subsequently, we analyzed the composition, structure, and functioning of the complete soil food webs of 58 individual plants, belonging to two of the plant species, Lotus corniculatus (Fabaceae) and Plantago lanceolata (Plantaginaceae). We isolated and identified more than 150 taxa/groups of soil organisms. The soil community composition and structure of the entire food webs were influenced both by the species identity of the plant individual and the surrounding plant community. Unexpectedly, plant identity had the strongest effects on decomposing soil organisms, widely believed to be generalist feeders. In contrast, quantitative food web modeling showed that the composition of the plant community influenced nitrogen mineralization under individual plants, but that plant species identity did not affect nitrogen or carbon mineralization or food web stability. Hence, the composition and structure of entire soil food webs vary at the scale of individual plants and are strongly influenced by the species identity of the plant. However, the ecosystem functions these food webs provide are determined by the identity of the entire plant community.

A Putative Mechanism for Bog Patterning

The surface of bogs commonly shows various spatial vegetation patterning. Typical are “string patterns” consisting of regular densely vegetated bands oriented perpendicular to the slope. Here, we report on regular “maze patterns” on flat ground, consisting of bands densely vegetated by vascular plants in a more sparsely vegetated matrix of nonvascular plant communities. We present a model reproducing these maze and string patterns, describing how nutrient‐limited vascular plants are controlled by, and in turn control, both hydrology and solute transport. We propose that the patterns are self‐organized and originate from a nutrient accumulation mechanism. In the model, this is caused by the convective transport of nutrients in the groundwater toward areas with higher vascular plant biomass, driven by differences in transpiration rate. In a numerical bifurcation analysis we show how the maze patterns originate from the spatially homogeneous equilibrium and how this is affected by changes in rainfall, nutrient input, and plant properties. Our results confirm earlier model results, showing that redistribution of a limiting resource may lead to fine‐scale facilitative and coarse‐scale competitive plant interactions in different ecosystems. Self‐organization in ecosystems may be a more general phenomenon than previously thought, which can be mechanistically linked to scale‐dependent facilitation and competition.

Global CO2 rise leads to reduced maximum stomatal conductance in Florida vegetation

A principle response of C3 plants to increasing concentrations of atmospheric CO2 (CO2) is to reduce transpirational water loss by decreasing stomatal conductance (gs) and simultaneously increase assimilation rates. Via this adaptation, vegetation has the ability to alter hydrology and climate. Therefore, it is important to determine the adaptation of vegetation to the expected anthropogenic rise in CO2. Short-term stomatal opening–closing responses of vegetation to increasing CO2 are described by free-air carbon enrichments growth experiments, and evolutionary adaptations are known from the geological record. However, to date the effects of decadal to centennial CO2 perturbations on stomatal conductance are still largely unknown. Here we reconstruct a 34% (±12%) reduction in maximum stomatal conductance (gsmax) per 100 ppm CO2 increase as a result of the adaptation in stomatal density (D) and pore size at maximal stomatal opening (amax) of nine common species from Florida over the past 150 y. The species-specific gsmax values are determined by different evolutionary development, whereby the angiosperms sampled generally have numerous small stomata and high gsmax, and the conifers and fern have few large stomata and lower gsmax. Although angiosperms and conifers use different D and amax adaptation strategies, our data show a coherent response in gsmax to CO2 rise of the past century. Understanding these adaptations of C3 plants to rising CO2 after decadal to centennial environmental changes is essential for quantification of plant physiological forcing at timescales relevant for global warming, and they are likely to continue until the limits of their phenotypic plasticity are reached.

Climate forcing due to optimization of maximal leaf conductance in subtropical vegetation under rising CO2

Plant physiological adaptation to the global rise in atmospheric CO2 concentration (CO2) is identified as a crucial climatic forcing. To optimize functioning under rising CO2, plants reduce the diffusive stomatal conductance of their leaves (gs) dynamically by closing stomata and structurally by growing leaves with altered stomatal densities and pore sizes. The structural adaptations reduce maximal stomatal conductance (gsmax) and constrain the dynamic responses of gs. Here, we develop and validate models that simulate structural stomatal adaptations based on diffusion of CO2 and water vapor through stomata, photosynthesis, and optimization of carbon gain under the constraint of a plant physiological cost of water loss. We propose that the ongoing optimization of gsmax is eventually limited by species-specific limits to phenotypic plasticity. Our model reproduces observed structural stomatal adaptations and predicts that adaptation will continue beyond double CO2. Owing to their distinct stomatal dimensions, angiosperms reach their phenotypic response limits on average at 740 ppm and conifers on average at 1,250 ppm CO2. Further, our simulations predict that doubling todays CO2 will decrease the annual transpiration flux of subtropical vegetation in Florida by ≈60 W·m−2. We conclude that plant adaptation to rising CO2 is altering the freshwater cycle and climate and will continue to do so throughout this century.

Application of parallel computing to stochastic parameter estimation in environmental models

Parameter estimation or model calibration is a common problem in many areas of process modeling, both in on-line applications such as real-time flood forecasting, and in off-line applications such as the modeling of reaction kinetics and phase equilibrium. The goal is to determine values of model parameters that provide the best fit to measured data, generally based on some type of least-squares or maximum likelihood criterion. Usually, this requires the solution of a non-linear and frequently non-convex optimization problem. In this paper we describe a user-friendly, computationally efficient parallel implementation of the Shuffled Complex Evolution Metropolis (SCEM-UA) global optimization algorithm for stochastic estimation of parameters in environmental models. Our parallel implementation takes better advantage of the computational power of a distributed computer system. Three case studies of increasing complexity demonstrate that parallel parameter estimation results in a considerable time savings when compared with traditional sequential optimization runs. The proposed method therefore provides an ideal means to solve complex optimization problems.

A critical transition in leaf evolution facilitated the Cretaceous angiosperm revolution

The revolutionary rise of broad-leaved (flowering) angiosperm plant species during the Cretaceous initiated a global ecological transformation towards modern biodiversity. Still, the mechanisms involved in this angiosperm radiation remain enigmatic. Here we show that the period of rapid angiosperm evolution initiated after the leaf interior (post venous) transport path length for water was reduced beyond the leaf interior transport path length for CO2 at a critical leaf vein density of 2.5–5 mm mm−2. Data and our modelling approaches indicate that surpassing this critical vein density was a pivotal moment in leaf evolution that enabled evolving angiosperms to profit from developing leaves with more and smaller stomata in terms of higher carbon returns from equal water loss. Surpassing the critical vein density may therefore have facilitated evolving angiosperms to develop leaves with higher gas exchange capacities required to adapt to the Cretaceous CO2 decline and outcompete previously dominant coniferous species in the upper canopy.

Islands of fertility induce co-occurring negative and positive plant-soil feedbacks promoting coexistence

Positive plant-soil feedback by “ecosystem engineers” is an important driver for the structuring and organization of resource-limited ecosystems. Although ample evidence demonstrates that plant-soil feedbacks can range from positive to strongly negative, their co-occurrence in plant communities have not yet been investigated. We test the hypothesis that the plant-soil feedback generated by the nitrogen-fixer shrub Medicago marina during primary succession in a sand dune community has a positive effect on the coexisting grass Lophochloa pubescens and a negative effect on the shrub species itself. We conducted field measurements and laboratory bioassays to evaluate (1) the effects of islands of fertility on the recruitment and growth of its ecosystem engineer and on the performance of a coexisting species and (2) the mechanisms involved that can explain the opposite effects of islands of fertility on coexisting species. Islands of fertility were present under Medicago crowns evidenced by higher available nitrogen, extractable phosphorus and potassium, organic matter, microbial activity, water holding capacity, soil humidity, and lower salt concentrations. The effects of these islands of fertility were clearly species-specific, with a facilitative impact on Lophochloa and a negative effect on Medicago recruitment. Lophochloa was denser and produced more biomass when rooted inside as compared to outside the crown area of the shrub. Contrarily, the number of seedlings of Medicago was lower inside, despite the higher seed abundance, and higher outside the crown area of adult shrubs as compared to predictions based on random distribution, thus showing a Janzen-Connell distribution. Laboratory experiments demonstrate the occurrence of Medicago negative plant-soil feedback, and that the auto-toxicity of the aboveground senescent plant material is a potentially important underlying mechanism explaining this negative feedback and the resulting Janzen-Connell distribution in the field.

Negative Plant–Soil Feedback and Positive Species Interaction in a Herbaceous Plant Community

Increasing evidence shows that facilitative interaction and negative plant–soil feedback are driving factors of plant population dynamics and community processes. We studied the intensity and the relative impact of negative feedback on clonal growth and seed germination of Scirpus holoschoenus, a ‘ring’ forming sedge dominant in grazed grassland, and the consequences for species coexistence. The structure of aboveground tussocks was described. A Lithium tracer assessed belowground distribution of functional roots. Seed rain and seedling emergence were compared for different positions in relation to Scirpus tussocks. Soil bioassays were used to compare growth on soil taken from inside and outside Scirpus tussocks of four coexisting species (Mentha acquatica, Pulicaria dysenterica, Scirpus holoschoenus and Dittrichia viscosa). We also compared plant performance of dominant plant species inside and outside Scirpus tussocks in the field. The ‘ring’ shaped tussocks of S. holoschoenus were generated by centrifugal rhizome development. Roots were functional and abundant under the tillers and extending outside the tussocks. The large roots mats that were present in the inner tussock zone were almost all dead. Seedling emergence and growth both showed a strong negative feedback of Scirpus in the inner tussock zone. Scirpus clonal development strongly reduced grass biomass. In the degenerated tussock zone, Pulicaria and Mentha mortality was lower, and biomass of individual plants and seed production were higher. This positive indirect interaction could be related to species-specific affinity to soil conditions generated by Scirpus, and interspecific competitive release in the degenerated tussock zone. We conclude that Scirpus negative feedback affects its seedling emergence and growth contributing to the development of the degenerated inner tussock zone. Moreover, this enhances species coexistence through facilitative interaction because the colonization of the inner tussock zone is highly species-specific.

Optimal allocation of leaf epidermal area for gas exchange

Summary A long‐standing research focus in phytology has been to understand how plants allocate leaf epidermal space to stomata in order to achieve an economic balance between the plants carbon needs and water use. Here, we present a quantitative theoretical framework to predict allometric relationships between morphological stomatal traits in relation to leaf gas exchange and the required allocation of epidermal area to stomata. Our theoretical framework was derived from first principles of diffusion and geometry based on the hypothesis that selection for higher anatomical maximum stomatal conductance (g smax) involves a trade‐off to minimize the fraction of the epidermis that is allocated to stomata. Predicted allometric relationships between stomatal traits were tested with a comprehensive compilation of published and unpublished data on 1057 species from all major clades. In support of our theoretical framework, stomatal traits of this phylogenetically diverse sample reflect spatially optimal allometry that minimizes investment in the allocation of epidermal area when plants evolve towards higher g smax. Our results specifically highlight that the stomatal morphology of angiosperms evolved along spatially optimal allometric relationships. We propose that the resulting wide range of viable stomatal trait combinations equips angiosperms with developmental and evolutionary flexibility in leaf gas exchange unrivalled by gymnosperms and pteridophytes.

Analysing forest transpiration model errors with artificial neural networks

Abstract A Single Big Leaf (SBL) forest transpiration model was calibrated on half-hourly eddy correlation measurements. The SBL model is based on the Penman–Monteith equation with a canopy conductance controlled by environmental variables. The model has eight calibration parameters, which determine the shape of the response functions. After calibration, residuals between measurements and model results exhibit complex patterns and contain random and systematic errors. Artificial Neural Networks (ANNs) were used to analyse these residuals for any systematic relations with environmental variables that may improve the SBL model. Different sub-sets of data were used to calibrate and validate the ANNs. Both wind direction and wind speed turned out to improve the model results. ANNs were able to find the source area of the fluxes of the Douglas fir stand within a larger heterogeneous forest without using a priori knowledge of the forest structure. With ANNs, improvements were also found in the shape and parameterisation of the response functions. Systematic errors in the original SBL model, caused by interdependencies between environmental variables, were not found anymore with the new parameterisation. After the ANNs analyses, about 80% of the residuals can be attributed to random errors of eddy correlation measurements. It is finally concluded that ANNs are able to find systematic trends even in very noisy residuals if applied properly.