Liesje Mommer

Biomass allocation to leaves, stems and roots: meta analyses of interspecific variation and environmental control

We quantified the biomass allocation patterns to leaves, stems and roots in vegetative plants, and how this is influenced by the growth environment, plant size, evolutionary history and competition. Dose-response curves of allocation were constructed by means of a meta-analysis from a wide array of experimental data. They show that the fraction of whole-plant mass represented by leaves (LMF) increases most strongly with nutrients and decreases most strongly with light. Correction for size-induced allocation patterns diminishes the LMF-response to light, but makes the effect of temperature on LMF more apparent. There is a clear phylogenetic effect on allocation, as eudicots invest relatively more than monocots in leaves, as do gymnosperms compared with woody angiosperms. Plants grown at high densities show a clear increase in the stem fraction. However, in most comparisons across species groups or environmental factors, the variation in LMF is smaller than the variation in one of the other components of the growth analysis equation: the leaf area : leaf mass ratio (SLA). In competitive situations, the stem mass fraction increases to a smaller extent than the specific stem length (stem length : stem mass). Thus, we conclude that plants generally are less able to adjust allocation than to alter organ morphology.

Going underground: root traits as drivers of ecosystem processes

Ecologists are increasingly adopting trait-based approaches to understand how community change influences ecosystem processes. However, most of this research has focussed on aboveground plant traits, whereas it is becoming clear that root traits are important drivers of many ecosystem processes, such as carbon (C) and nutrient cycling, and the formation and structural stability of soil. Here, we synthesise emerging evidence that illustrates how root traits impact ecosystem processes, and propose a pathway to unravel the complex roles of root traits in driving ecosystem processes and their response to global change. Finally, we identify research challenges and novel technologies to address them.

The way forward in biochar research: targeting trade-offs between the potential wins

Biochar application to soil is currently widely advocated for a variety of reasons related to sustainability. Typically, soil amelioration with biochar is presented as a multiple‐‘win’ strategy, although it is also associated with potential risks such as environmental contamination. The most often claimed benefits of biochar (i.e. the ‘wins’) include (i) carbon sequestration; (ii) soil fertility enhancement; (iii) biofuel/bioenergy production; (iv) pollutant immobilization; and (v) waste disposal. However, the vast majority of studies ignore possible trade‐offs between them. For example, there is an obvious trade‐off between maximizing biofuel production and maximizing biochar production. Also, relatively little attention has been paid to mechanisms, as opposed to systems impacts, behind observed biochar effects, often leaving open the question as to whether they reflect truly unique properties of biochar as opposed to being simply the short‐term consequences of a fertilization or liming effect. Here, we provide an outline for the future of soil biochar research. We first identify possible trade‐offs between the potential benefits. Second, to be able to better understand and quantify these trade‐offs, we propose guidelines for robust experimental design and selection of appropriate controls that allow both mechanistic and systems assessment of biochar effects and trade‐offs between the wins. Third, we offer a conceptual framework to guide future experiments and suggest guidelines for the standardized reporting of biochar experiments to allow effective between‐site comparisons to quantify trade‐offs. Such a mechanistic and systems framework is required to allow effective comparisons between experiments, across scales and locations, to guide policy and recommendations concerning biochar application to soil.

A modular concept of plant foraging behaviour: the interplay between local responses and systemic control

In this paper we examined the notion that plant foraging for resources in heterogeneous environments must involve: (1) plasticity at the level of individual modules in reaction to localized environmental signals; and (2) the potential for modification of these responses either by the signals received from connected modules that may be exposed to different conditions, or by the signals reflecting the overall resource status of the plant. A conceptual model is presented to illustrate how plant foraging behaviour is achieved through these processes acting in concert, from the signal reception through signal transduction to morphological or physiological response. Evidence to support the concept is reviewed, using selective root placement under nutritionally heterogeneous conditions and elongation responses of stems and petioles to shade as examples. We discussed how the adoption of this model can promote understanding of the ecological significance of foraging behaviour. We also identified a need to widen the experimental repertoires of both molecular physiology and ecology in order to increase our insight into both the regulation and functioning of foraging responses, and their relationship with the patterns of environmental heterogeneity under which plants have evolved.

Submergence-Induced Morphological, Anatomical, and Biochemical Responses in a Terrestrial Species Affect Gas Diffusion Resistance and Photosynthetic Performance

Gas exchange between the plant and the environment is severely hampered when plants are submerged, leading to oxygen and energy deficits. A straightforward way to reduce these shortages of oxygen and carbohydrates would be continued photosynthesis under water, but this possibility has received only little attention. Here, we combine several techniques to investigate the consequences of anatomical and biochemical responses of the terrestrial species Rumex palustris to submergence for different aspects of photosynthesis under water. The orientation of the chloroplasts in submergence-acclimated leaves was toward the epidermis instead of the intercellular spaces, indicating that underwater CO2 diffuses through the cuticle and epidermis. Interestingly, both the cuticle thickness and the epidermal cell wall thickness were significantly reduced upon submergence, suggesting a considerable decrease in diffusion resistance. This decrease in diffusion resistance greatly facilitated underwater photosynthesis, as indicated by higher underwater photosynthesis rates in submergence-acclimated leaves at all CO2 concentrations investigated. The increased availability of internal CO2 in these “aquatic” leaves reduced photorespiration, and furthermore reduced excitation pressure of the electron transport system and, thus, the risk of photodamage. Acclimation to submergence also altered photosynthesis biochemistry as reduced Rubisco contents were observed in aquatic leaves, indicating a lower carboxylation capacity. Electron transport capacity was also reduced in these leaves but not as strongly as the reduction in Rubisco, indicating a substantial increase of the ratio between electron transport and carboxylation capacity upon submergence. This novel finding suggests that this ratio may be less conservative than previously thought.

Unravelling below-ground plant distributions: a real-time polymerase chain reaction method for quantifying species proportions in mixed root samples

Knowledge on below‐ground plant distributions is almost lacking to date, despite the fact that such information would be very valuable in understanding below‐ground competition and species‐specific interactions, processes that are expected to shape community structure. Methods available so far for below‐ground species determination have drawbacks that we tried to challenge. Some methods make use of differences in the chemical composition between species, but this is highly variable upon environmental factors. DNA‐based techniques — far less dependent on chemical composition — such as polymerase chain reaction on internal transcribed spacer (ITS) primers can so far only determine presence–absence of a species in a mixed root sample. Here, we present a quantitative DNA‐based technique that allows investigation of relative species abundances in experimental mixed root samples. We used quantitative real‐time polymerase chain reaction (PCR) on species‐specific markers obtained from intersimple sequence repeat (ISSR) analyses in root samples. This molecular technique is novel in the field of root ecology and its development overcame three challenges: (i) determination of species‐specific DNA fragments, (ii) development and optimization of the real time PCR protocol, (iii) designing a data treatment method based on a modified delta–delta‐cycle threshold (CT) analysis. The method gained robustness from using relative DNA abundances in species mixtures rather than absolute concentration readings. This requires accurate multispecies reference series as a calibration. Test samples with different known biomass ratios of all species showed proof of concept of this method. The pros and contras of this method are discussed in the light of its contribution to advancing ecological research on below‐ground plant–plant interactions.

Plant species richness promotes soil carbon and nitrogen stocks in grasslands without legumes

Summary 1. The storage of carbon (C) and nitrogen (N) in soil is important ecosystem functions. Grassland biodiversity experiments have shown a positive effect of plant diversity on soil C and N storage. However, these experiments all included legumes, which constitute an important N input through N2-fixation. Indeed, the results of these experiments suggest that N2 fixation by legumes is a major driver of soil C and N storage. 2. We studied whether plant diversity affects soil C and N storage in the absence of legumes. In an 11-year grassland biodiversity experiment without legumes, we measured soil C and N stocks. We further determined above-ground biomass productivity, standing root biomass, soil organic matter decomposition and N mineralization rates to understand the mechanisms underlying the change in soil C and N stocks in relation to plant diversity and their feedbacks to plant productivity. 3. We found that soil C and N stocks increased by 18% and 16% in eight-species mixtures compared to the average of monocultures of the same species, respectively. Increased soil C and N stocks were mainly driven by increased C input and N retention, resulting from enhanced plant productivity, which surpassed enhanced C loss from decomposition. Importantly, higher soil C and N stocks were associated with enhanced soil N mineralization rates, which can explain the strengthening of the positive diversity–productivity relationship observed in the last years of the experiment. 4. Synthesis. We demonstrated that also in the absence of legumes, plant species richness promotes soil carbon (C) and nitrogen (N) stocks via increased plant productivity. In turn, enhanced soil C and N stocks showed a positive feedback to plant productivity via enhanced N mineralization, which could further accelerate soil C and N storage in the long term.

Contrasting root behaviour in two grass species: a test of functionality in dynamic heterogeneous conditions

Root systems are highly plastic as they express a range of responses to acquire patchily distributed nutrients. However, the ecological significance of placing roots selectively in nutrient hotspots is still unclear. Here, we investigate under what conditions selective root placement may be a significant functional trait that determines belowground competitive ability. We studied two grasses differing in root foraging behaviour, Festuca rubra and Anthoxanthum odoratum. The plants were grown in stable and more dynamic heterogeneous environments, by switching nutrient patches halfway through the experiment. A. odoratum was a factor of two less selective in placing its roots into nutrient-rich patches than F. rubra. A. odoratum produced overall higher root length densities with higher specific root length than F. rubra and acquired more nutrients. A. odoratum appeared to be the superior competitor, irrespective of the nutrient dynamics. Our results suggest that root behaviour consisting of producing high root length densities at relatively low biomass investments can be a more effective foraging strategy than placing roots selectively in nutrient hotspots. When understanding the functionality of root traits among different species, specific root length may play a key role.

Independent variations of plant and soil mixtures reveal soil feedback effects on plant community overyielding

* Recent studies have shown that the positive relationship between plant diversity and plant biomass (‘overyielding’) can be explained by soil pathogens depressing productivity more in low than in high diverse plant communities. However, tests of such soil effects in field studies were constrained by experimental limitations to manipulate soil community composition independent of plant community composition. Here, we report of an experiment where feedback effects to plants were tested for both plant and soil monocultures and mixtures. * Our results demonstrate that overyielding is the result of plant species in mixture being more growth-limited by ‘own’ soil biota than by soil biota of other plant species. This effect disappeared when the soils had been sterilized by gamma-irradiation. Mixing plants themselves did not result in overyielding except when grown in the soil of one of the species (Leucanthemum vulgare), where growth of one species disproportionally increased in mixture compared to monoculture. * Soil nutrient availability could not explain differences in growth between the non-sterilized soils. Therefore, our results suggest that plant species–specific soil biota rather than the plants have contributed to the plant community overyielding. * Species biomass ranking in mixtures highly differed between non-sterilized soils of different histories of soil conditioning, whilst the ranking was more consistent in sterilized soil. Sterilized soils of different origin differed significantly in nutrient availability. These results suggest that shifts in competitive hierarchies depend on plant species–specific interactions influenced by soil biota and cannot be induced by mineral nitrogen. * Synthesis. Our results show that overyielding in four plant species mixtures can be due to species-specific interactions between plants and their specific soil biota. Neither mixing the plant species alone nor the differential responses of species to mineral nitrogen influenced community productivity, but mixing soil biota did. [KEYWORDS: Anthoxanthum odoratum biodiversity experiment biodiversity–productivity relationship competitive hierarchy determinants of plant community diversity and structure grasslands Leucanthemum vulgare micro-organisms pathogens plant–soil feedback]

Towards a multidimensional root trait framework: a tree root review

Contents 1159 I. 1159 II. 1161 III. 1164 IV. 1166 1167 References 1167 SUMMARY: The search for a root economics spectrum (RES) has been sparked by recent interest in trait-based plant ecology. By analogy with the one-dimensional leaf economics spectrum (LES), fine-root traits are hypothesised to match leaf traits which are coordinated along one axis from resource acquisitive to conservative traits. However, our literature review and meta-level analysis reveal no consistent evidence of an RES mirroring an LES. Instead the RES appears to be multidimensional. We discuss three fundamental differences contributing to the discrepancy between these spectra. First, root traits are simultaneously constrained by various environmental drivers not necessarily related to resource uptake. Second, above- and belowground traits cannot be considered analogues, because they function differently and might not be related to resource uptake in a similar manner. Third, mycorrhizal interactions may offset selection for an RES. Understanding and explaining the belowground mechanisms and trade-offs that drive variation in root traits, resource acquisition and plant performance across species, thus requires a fundamentally different approach than applied aboveground. We therefore call for studies that can functionally incorporate the root traits involved in resource uptake, the complex soil environment and the various soil resource uptake mechanisms - particularly the mycorrhizal pathway - in a multidimensional root trait framework.