Giulia Vico

Hydraulic limits on maximum plant transpiration and the emergence of the safety–efficiency trade‐off

Soil and plant hydraulics constrain ecosystem productivity by setting physical limits to water transport and hence carbon uptake by leaves. While more negative xylem water potentials provide a larger driving force for water transport, they also cause cavitation that limits hydraulic conductivity. An optimum balance between driving force and cavitation occurs at intermediate water potentials, thus defining the maximum transpiration rate the xylem can sustain (denoted as E(max)). The presence of this maximum raises the question as to whether plants regulate transpiration through stomata to function near E(max). To address this question, we calculated E(max) across plant functional types and climates using a hydraulic model and a global database of plant hydraulic traits. The predicted E(max) compared well with measured peak transpiration across plant sizes and growth conditions (R = 0.86, P < 0.001) and was relatively conserved among plant types (for a given plant size), while increasing across climates following the atmospheric evaporative demand. The fact that E(max) was roughly conserved across plant types and scales with the product of xylem saturated conductivity and water potential at 50% cavitation was used here to explain the safety-efficiency trade-off in plant xylem. Stomatal conductance allows maximum transpiration rates despite partial cavitation in the xylem thereby suggesting coordination between stomatal regulation and xylem hydraulic characteristics.

Effects of stomatal delays on the economics of leaf gas exchange under intermittent light regimes

• Understory plants are subjected to highly intermittent light availability and their leaf gas exchanges are mediated by delayed responses of stomata and leaf biochemistry to light fluctuations. In this article, the patterns in stomatal delays across biomes and plant functional types were studied and their effects on leaf carbon gains and water losses were quantified. • A database of more than 60 published datasets on stomatal responses to light fluctuations was assembled. To interpret these experimental observations, a leaf gas exchange model was developed and coupled to a novel formulation of stomatal movement energetics. The model was used to test whether stomatal delays optimize light capture for photosynthesis, whilst limiting transpiration and carbon costs for stomatal movement. • The data analysis showed that stomatal opening and closing delays occurred over a limited range of values and were strongly correlated. Plant functional type and climate were the most important drivers of stomatal delays, with faster responses in graminoids and species from dry climates. • Although perfectly tracking stomata would maximize photosynthesis and minimize transpiration at the expense of large opening costs, the observed combinations of opening and closure times appeared to be consistent with a near-optimal balance of carbon gain, water loss and movement costs.

Modelling C3 and C4 photosynthesis under water-stressed conditions

Despite the observed impact of water stress on photosynthesis, some of the most used models of CO2 assimilation in C3 and C4 functional types do not directly account for it. We discuss an extension of these models, which explicitly includes the metabolic and diffusive limitations due to water stress on photosynthesis. Functional relationships describing the photosynthetic processes and CO2 diffusion inside leaves are modified to account for leaf water status on the basis of experimental results available in the literature. Extensive comparison with data shows that the model is suitable to describe the reduction in CO2 assimilation rate with decreasing leaf water potentials in various species. A simultaneous analysis of photosynthesis, transpiration and soil moisture dynamics is then carried out to explore the actual impact of drought on different photosynthesis processes and on the overall plant activity. The model well reproduces measured CO2 assimilation rate as a function of soil moisture and could be useful to formulate hypotheses for detailed experiments as well as to simulate in detail transpiration and photosynthesis dynamics under water stress.

Increasing water use efficiency along the C3 to C4 evolutionary pathway: a stomatal optimization perspective

Summary Using a stomatal optimization model, an abrupt change is foundnd in the relationship between carbon and water fluxes along the evolutionary gradient from C3 to C4 photosynthetic types.

Optimal plant water‐use strategies under stochastic rainfall

Plant hydraulic traits have been conjectured to be coordinated, thereby providing plants with a balanced hydraulic system that protects them from cavitation while allowing an efficient transport of water necessary for photosynthesis. In particular, observations suggest correlations between the water potentials at which xylem cavitation impairs water movement and the one at stomatal closure, and between maximum xylem and stomatal conductances, begging the question as to whether such coordination emerges as an optimal water-use strategy under unpredictable rainfall. Here mean transpiration is used as a proxy for long-term plant fitness and its variations as a function of the water potentials at 50% loss of stem conductivity due to cavitation and at 90% stomatal closure are explored. It is shown that coordination between these hydraulic traits is necessary to maximize , with rainfall patterns altering the optimal range of trait values. In contrast, coordination between ecosystem-level conductances appears not necessary to maximize . The optimal trait ranges are wider under drier than under mesic conditions, suggesting that in semiarid systems different water use strategies may be equally successful. Comparison with observations across species from a range of ecosystems confirms model predictions, indicating that the coordinated functioning of plant organs might indeed emerge from an optimal response to rainfall variability.

On the complementary relationship between marginal nitrogen and water-use efficiencies among Pinus taeda leaves grown under ambient and CO2-enriched environments

BACKGROUND AND AIMS Water and nitrogen (N) are two limiting resources for biomass production of terrestrial vegetation. Water losses in transpiration (E) can be decreased by reducing leaf stomatal conductance (g(s)) at the expense of lowering CO(2) uptake (A), resulting in increased water-use efficiency. However, with more N available, higher allocation of N to photosynthetic proteins improves A so that N-use efficiency is reduced when g(s) declines. Hence, a trade-off is expected between these two resource-use efficiencies. In this study it is hypothesized that when foliar concentration (N) varies on time scales much longer than g(s), an explicit complementary relationship between the marginal water- and N-use efficiency emerges. Furthermore, a shift in this relationship is anticipated with increasing atmospheric CO(2) concentration (c(a)). METHODS Optimization theory is employed to quantify interactions between resource-use efficiencies under elevated c(a) and soil N amendments. The analyses are based on marginal water- and N-use efficiencies, λ = (∂A/∂g(s))/(∂E/∂g(s)) and η = ∂A/∂N, respectively. The relationship between the two efficiencies and related variation in intercellular CO(2) concentration (c(i)) were examined using A/c(i) curves and foliar N measured on Pinus taeda needles collected at various canopy locations at the Duke Forest Free Air CO(2) Enrichment experiment (North Carolina, USA). KEY RESULTS Optimality theory allowed the definition of a novel, explicit relationship between two intrinsic leaf-scale properties where η is complementary to the square-root of λ. The data support the model predictions that elevated c(a) increased η and λ, and at given c(a) and needle age-class, the two quantities varied among needles in an approximately complementary manner. CONCLUSIONS The derived analytical expressions can be employed in scaling-up carbon, water and N fluxes from leaf to ecosystem, but also to derive transpiration estimates from those of η, and assist in predicting how increasing c(a) influences ecosystem water use.

Trade-offs between seed output and life span - a quantitative comparison of traits between annual and perennial congeneric species.

Perennial plants allocate more resources belowground, thus sustaining important ecosystem services. Hence, shifting from annual to perennial crops has been advocated towards a more sustainable agriculture. Nevertheless, wild perennial species have lower seed production than selected annuals, raising the questions of whether there is a fundamental trade-off between reproductive effort and life span, and whether such trade-off can be overcome through selection. In order to address these questions and to isolate life span from phylogenetic and environmental factors, we conducted a meta-analysis encompassing c. 3000 congeneric annual/perennial pairs from 28 genera. This meta-analysis is complemented with a minimalist model of long-term productivity in perennial species. Perennials allocate more resources belowground and less to seeds than congeneric annuals, independently of selection history. However, existing perennial wheat and rice could achieve yields similar to annuals if they survived three years and each year doubled their biomass, as other perennial grasses do. Selected perennial crops maintain the large belowground allocation of wild perennials, and thus can provide desired regulatory ecosystem services. To match the seed yield of annuals, biomass production of perennial grains must be increased to amounts attained by some perennial grasses - if this goal can be met, perennial crops can provide a more sustainable alternative to annuals.

Ecohydrological modeling in agroecosystems: Examples and challenges

Human societies are increasingly altering the water and biogeochemical cycles to both improve ecosystem productivity and reduce risks associated with the unpredictable variability of climatic drivers. These alterations, however, often cause large negative environmental consequences, raising the question as to how societies can ensure a sustainable use of natural resources for the future. Here we discuss how ecohydrological modeling may address these broad questions with special attention to agroecosystems. The challenges related to modeling the two-way interaction between society and environment are illustrated by means of a dynamical model in which soil and water quality supports the growth of human society but is also degraded by excessive pressure, leading to critical transitions and sustained societal growth-collapse cycles. We then focus on the coupled dynamics of soil water and solutes (nutrients or contaminants), emphasizing the modeling challenges, presented by the strong nonlinearities in the soil and plant system and the unpredictable hydroclimatic forcing, that need to be overcome to quantitatively analyze problems of soil water sustainability in both natural and agricultural ecosystems. We discuss applications of this framework to problems of irrigation, soil salinization, and fertilization and emphasize how optimal solutions for large-scale, long-term planning of soil and water resources in agroecosystems under uncertainty could be provided by methods from stochastic control, informed by physically and mathematically sound descriptions of ecohydrological and biogeochemical interactions.

Coupled carbon and water fluxes in CAM photosynthesis: modeling quantification of water use efficiency and productivity.

Background and AimsDue to their high water use efficiency, Crassulacean acid metabolism (CAM) plants are of environmental and economic importance in the arid and semiarid regions of the world. Moreover, many CAM plants (e.g., Agave tequilana) have attractive qualities for biofuel production such as a relatively low lignin content and high amount of soluble carbohydrates. However, the current estimates of CAM productivity are based on empirical stress indices that create large uncertainties. As a first step towards a more accurate quantification of CAM productivity, this paper introduces a new model that couples both soil and atmosphere conditions to CAM photosynthesis.MethodsThe new CAM model is based upon well established C3 photosynthesis models coupled to a nonlinear circadian rhythm oscillator for the control of the photosynthesis carbon fluxes. The leaf-level dynamics are coupled to a simple, yet realistic description of the soil-plant-atmosphere continuum, including a plant water capacitance module.ResultsThe resulting model reproduces the four phases of CAM photosynthes is and the evolution of their dynamics during a soil moisture drydown, as a function of soil type, plant features, and climatic conditions.ConclusionThe results help quantify the impact of soil water availability on CAM carbon assimilation and transpiration flux.

Probabilistic description of topographic slope and aspect

[1] Local topographic features such as slope and aspect play a crucial role in a number of morphological, ecological, and hydrological processes. We propose a simple yet realistic probabilistic description of local slope and aspect as a function of properties of the field of elevation changes. We consider different classes of models of elevation changes and obtain the theoretical distribution of slope and aspect. We relate the features of the obtained distributions to large-scale landscape structures, such as regional trends and anisotropy. We find that the theoretical distribution of slope is strongly impacted by the parameters used to represent the distribution of elevation changes, while large-scale features play a secondary role. Conversely, the distribution of aspect is also controlled by regional trends and anisotopy, even when they are weak. The proposed statistical description of slope and aspect is applied to assess the effects of topographic features on direct solar radiation mean and standard deviation. The main control on direct solar radiation is exerted by the partial derivative variance. We consider four different landscapes across the continental United States and compare the proposed theoretical description of slope and aspect distributions to the observed histograms.