Subtle variation in shade avoidance responses may have profound consequences for plant competitiveness

Background and Aims: Although phenotypic plasticity has been shown to be beneficial for plant competitiveness for light, there is limited knowledge on how variation in these plastic responses plays a role in determining competitiveness. Methods: A combination of detailed plant experiments and functional–structural plant (FSP) modelling was used that captures the complex dynamic feedback between the changing plant phenotype and the within-canopy light environment in time and 3-D space. Leaf angle increase (hyponasty) and changes in petiole elongation rates in response to changes in the ratio between red and far-red light, two important shade avoidance responses in Arabidopsis thaliana growing in dense population stands, were chosen as a case study for plant plasticity. Measuring and implementing these responses into an FSP model allowed simulation of plant phenotype as an emergent property of the underlying growth and response mechanisms. Key Results: Both the experimental and model results showed that substantial differences in competitiveness may arise between genotypes with only marginally different hyponasty or petiole elongation responses, due to the amplification of plant growth differences by small changes in plant phenotype. In addition, this study illustrated that strong competitive responses do not necessarily have to result in a tragedy of the commons; success in competition at the expense of community performance. Conclusions: Together, these findings indicate that selection pressure could probably have played a role in fine-tuning the sensitive shade avoidance responses found in plants. The model approach presented here provides a novel tool to analyse further how natural selection could have acted on the evolution of plastic responses

Species richness, functional traits and climate interactively affect tree survival in a large forest biodiversity experiment

1. Tree survival affects forest biodiversity, structure and functioning. However, little is known about feedback effects of biodiversity on survival and its dependence on functional traits and interannual climatic variability. 2. With an individual-based dataset from a large subtropical forest biodiversity experiment, we evaluated how species richness, functional traits and time-dependent covariates affected annual tree survival rates from age 3–12 (years) after planting 39 species across a diversity gradient from 1 to 2, 4, 8 and 16 tree species. 3. We found that overall survival rates marginally increased with diversity at the plot level, with large variation among plots within diversity levels. Significant variation among species in survival responses to diversity and changes in these responses with age were related to species functional traits and climatic conditions. Generally, survival rates of conservative species (evergreen, late-successional species with thick leaves and high carbon to nitrogen ratio but low specific leaf area, leaf phosphorus and hydraulic conductivity) increased with diversity, age and yearly precipitation, whereas acquisitive species showed opposite responses. 4. Synthesis. Our results indicate that interactions between diversity, species functional traits and yearly climatic conditions can balance survival among species in diverse forests. Planting mixtures of species that differ in functional traits in afforestation projects may lead to a positive feedback loop where biodiversit

Species richness stabilizes productivity via asynchrony and drought-tolerance diversity in a large-scale tree biodiversity experiment

Extreme climatic events threaten forests and their climate mitigation potential globally. Understanding the drivers promoting ecosystem stability is therefore considered crucial for mitigating adverse climate change effects on forests. Here, we use structural equation models to explain how tree species richness, asynchronous species dynamics, species-level population stability, and drought-tolerance traits relate to the stability of forest productivity along an experimentally manipulated species richness gradient ranging from 1 to 24 tree species. Tree species richness improved community stability by increasing asynchrony. That is, at higher species richness, interannual variation in productivity among tree species buffered the community against stress-related productivity declines. This effect was positively related to variation in stomatal control and resistance-acquisition strategies among species, but not to the community-weighted means of these trait syndromes. The identified mechanisms by which tree species richness stabilizes forest productivity emphasize the importance of diverse, mixed-species forests to adapt to climate change

Tree species and genetic diversity increase productivity via functional diversity and trophic feedbacks

Addressing global biodiversity loss requires an expanded focus on multiple dimensions of biodiversity. While most studies have focused on the consequences of plant interspecific diversity, our mechanistic understanding of how genetic diversity within plant species affects plant productivity remains limited. Here, we use a tree species × genetic diversity experiment to disentangle the effects of species diversity and genetic diversity on tree productivity, and how they are related to tree functional diversity and trophic feedbacks. We found that tree species diversity increased tree productivity via increased tree functional diversity, reduced soil fungal diversity, and marginally reduced herbivory. The effects of tree genetic diversity on productivity via functional diversity and soil fungal diversity were negative in monocultures but positive in the mixture of the four tree species tested. Given the complexity of interactions between species and genetic diversity, tree functional diversity and trophic feedbacks on productivity, we suggest that both tree species and genetic diversity should be considered in afforestation

Species richness stabilizes productivity via asynchrony and drought-tolerance diversity in a large-scale tree biodiversity experiment

Extreme climatic events threaten forests and their climate mitigation potential globally. Understanding the drivers promoting ecosystem stability is therefore considered crucial for mitigating adverse climate change effects on forests. Here, we use structural equation models to explain how tree species richness, asynchronous species dynamics, species-level population stability, and drought-tolerance traits relate to the stability of forest productivity along an experimentally manipulated species richness gradient ranging from 1 to 24 tree species. Tree species richness improved community stability by increasing asynchrony. That is, at higher species richness, interannual variation in productivity among tree species buffered the community against stress-related productivity declines. This effect was positively related to variation in stomatal control and resistance-acquisition strategies among species, but not to the community-weighted means of these trait syndromes. The identified mechanisms by which tree species richness stabilizes forest productivity emphasize the importance of diverse, mixed-species forests to adapt to climate change

Functional-structural plant models to boost understanding of complementarity in light capture and use in mixed-species forests

Positive relationships between tree species richness and productivity have been observed in both natural and experimental forests. Complementarity in resource capture and use has been put forward as one important mechanism for the positive richness-productivity relationship. However, inference that complementarity drives this relationship is often based on statistical modelling or the use of functional diversity indices and these methods do not consider resource capture or use among individuals from various species within a community. Here I introduce functional-structural plant models as tool to study species complementarity in light capture and use in mixed-species forests. These models consider the interplay between structure, physiology and the environment in 3D and scale from organ-specific characteristics to tree and community performance. Functional-structural plant models that represent mixed-species forests have therefore the potential to disentangle the effects of structural and functional differences among species on community light capture and use using the 3D setting of the forest. Knowing how community light capture and use in mixed-species forests depends on structural and functional differences among species will shed light on the potential of species complementarity in light capture and use to drive species richness-productivity relationships. (C) 2020 Gesellschaft fur okologie. Published by Elsevier GmbH. All rights reserved

How virtual shade sheds light on plant plasticity

Phenotypic plasticity is the ability of a genotype to express multiple phenotypes in accordance with different environments. Although variation in plasticity has been observed, there is limited knowledge on how this variation results from natural selection. This thesis analyses how variation in the level of plasticity influences light competition between plants and how this variation could result from selection, driven by light competition, in various environments. As an exemplary case of phenotypic plasticity, this thesis focusses on phenotypic responses of the annual rosette plant Arabidopsis thaliana (Brassicaceae) in response to the proximity of neighbour plants, as signalled through the red : far—red (R:FR) ratio, which are responses associated with the shade avoidance syndrome (SAS). Plant experiments were conducted to measure variation in these plastic responses and a functional-structural plant (FSP) model was created that simulates plant structures in 3D and includes these organ-level plastic responses while simulating explicitly a heterogeneous light environment. Simulating individual plants that explicitly compete for light, while their phenotype changes through plasticity, gave insights in the role of the level of phenotypic plasticity and site of signal perception on plant competitiveness. In addition, an analysis on how natural selection in different environments acts on the level of plasticity was performed by combining FSP simulations and evolutionary game theoretical (EGT) principles.</p

Reducing shade avoidance can improve Arabidopsis canopy performance against competitors

Plants that grow in high density communities activate shade avoidance responses to consolidate light capture by individuals. Although this is an evolutionary successful strategy, it may not enhance performance of the community as a whole. Resources are invested in shade responses at the expense of other organs and light penetration through the canopy is increased, allowing invading competitors to grow better. Here we investigate if suppression of shade avoidance responses would enhance group performance of a monoculture community that is invaded by a competitor. Using different Arabidopsis genotypes, we show that suppression of shade-induced upward leaf movement in the pif7 mutant increases the pif7 communal performance against invaders as compared to a wild-type canopy. The invaders were more severely suppressed and the community grew larger as compared to wild type. Using computational modelling, we show that leaf angle variations indeed strongly affect light penetration and growth of competitors that invade the canopy. Our data thus show that modifying specific shade avoidance aspects can improve plant community performance. These insights may help to suppress weeds in crop stands

Genetic richness affects trait variation but not community productivity in a tree diversity experiment

Biodiversity–ecosystem functioning experiments found that productivity generally increases with species richness, but less is known about effects of within‐species genetic richness and potential interactions between the two. While functional differences between species can explain species richness effects, empirical evidence regarding functional differences between genotypes within species and potential consequences for productivity is largely lacking. We therefore measured within‐ and among‐species variation in functional traits and growth and determined stand‐level tree biomass in a large forest experiment factorially manipulating species and genetic richness in subtropical China. Within‐species variation across genetic seed families, in addition to variation across species, explained a substantial amount of trait variation. Furthermore, trait responses to species and genetic richness varied significantly within and between species. Multivariate trait variation was larger among individuals from species mixtures than those from species monocultures, but similar among individuals from genetically diverse vs genetically uniform monocultures. Correspondingly, species but not genetic richness had a positive effect on stand‐level tree biomass. We argue that identifying functional diversity within and among species in forest communities is necessary to separate effects of species and genetic diversity on tree growth and community productivity

Reducing shade avoidance can improve Arabidopsis canopy performance against competitors

Plants that grow in high density communities activate shade avoidance responses to consolidate light capture by individuals. Although this is an evolutionary successful strategy, it may not enhance performance of the community as a whole. Resources are invested in shade responses at the expense of other organs and light penetration through the canopy is increased, allowing invading competitors to grow better. Here we investigate if suppression of shade avoidance responses would enhance group performance of a monoculture community that is invaded by a competitor. Using different Arabidopsis genotypes, we show that suppression of shade-induced upward leaf movement in the pif7 mutant increases the pif7 communal performance against invaders as compared to a wild-type canopy. The invaders were more severely suppressed and the community grew larger as compared to wild type. Using computational modelling, we show that leaf angle variations indeed strongly affect light penetration and growth of competitors that invade the canopy. Our data thus show that modifying specific shade avoidance aspects can improve plant community performance. These insights may help to suppress weeds in crop stands