Damien Landais

Elevated CO2 maintains grassland net carbon uptake under a future heat and drought extreme

Significance Ecosystems are responding to climate change and increasing atmospheric CO2 concentrations. Interactions between these factors have rarely been assessed experimentally during and after extreme climate events despite their predicted increase in intensity and frequency and their negative impact on primary productivity and soil carbon stocks. Here, we document how a grassland exposed to a forecasted 2050s climate shows a remarkable recovery of ecosystem carbon uptake after a severe drought and heat wave, this recovery being amplified under elevated CO2. Over the growing season, elevated CO2 entirely compensated for the negative impact of extreme heat and drought on net carbon uptake. This study highlights the importance of incorporating all interacting factors in the predictions of climate change impacts. Extreme climatic events (ECEs) such as droughts and heat waves are predicted to increase in intensity and frequency and impact the terrestrial carbon balance. However, we lack direct experimental evidence of how the net carbon uptake of ecosystems is affected by ECEs under future elevated atmospheric CO2 concentrations (eCO2). Taking advantage of an advanced controlled environment facility for ecosystem research (Ecotron), we simulated eCO2 and extreme cooccurring heat and drought events as projected for the 2050s and analyzed their effects on the ecosystem-level carbon and water fluxes in a C3 grassland. Our results indicate that eCO2 not only slows down the decline of ecosystem carbon uptake during the ECE but also enhances its recovery after the ECE, as mediated by increases of root growth and plant nitrogen uptake induced by the ECE. These findings indicate that, in the predicted near future climate, eCO2 could mitigate the effects of extreme droughts and heat waves on ecosystem net carbon uptake.

Processes driving nocturnal transpiration and implications for estimating land evapotranspiration

Evapotranspiration is a major component of the water cycle, yet only daytime transpiration is currently considered in Earth system and agricultural sciences. This contrasts with physiological studies where 25% or more of water losses have been reported to occur occurring overnight at leaf and plant scales. This gap probably arose from limitations in techniques to measure nocturnal water fluxes at ecosystem scales, a gap we bridge here by using lysimeters under controlled environmental conditions. The magnitude of the nocturnal water losses (12–23% of daytime water losses) in row-crop monocultures of bean (annual herb) and cotton (woody shrub) would be globally an order of magnitude higher than documented responses of global evapotranspiration to climate change (51–98 vs. 7–8 mm yr−1). Contrary to daytime responses and to conventional wisdom, nocturnal transpiration was not affected by previous radiation loads or carbon uptake, and showed a temporal pattern independent of vapour pressure deficit or temperature, because of endogenous controls on stomatal conductance via circadian regulation. Our results have important implications from large-scale ecosystem modelling to crop production: homeostatic water losses justify simple empirical predictive functions, and circadian controls show a fine-tune control that minimizes water loss while potentially increasing posterior carbon uptake.

Endogenous circadian rhythms in pigment composition induce changes in photochemical efficiency in plant canopies

There is increasing evidence that the circadian clock is a significant driver of photosynthesis that becomes apparent when environmental cues are experimentally held constant. We studied whether the composition of photosynthetic pigments is under circadian regulation, and whether pigment oscillations lead to rhythmic changes in photochemical efficiency. To address these questions, we maintained canopies of bean and cotton, after an entrainment phase, under constant (light or darkness) conditions for 30-48 h. Photosynthesis and quantum yield peaked at subjective noon, and non-photochemical quenching peaked at night. These oscillations were not associated with parallel changes in carbohydrate content or xanthophyll cycle activity. We observed robust oscillations of Chl a/b during constant light in both species, and also under constant darkness in bean, peaking when it would have been night during the entrainment (subjective nights). These oscillations could be attributed to the synthesis and/or degradation of trimeric light-harvesting complex II (reflected by the rhythmic changes in Chl a/b), with the antenna size minimal at night and maximal around subjective noon. Considering together the oscillations of pigments and photochemistry, the observed pattern of changes is counterintuitive if we assume that the plant strategy is to avoid photodamage, but consistent with a strategy where non-stressed plants maximize photosynthesis.

Circadian rhythms have significant effects on leaf-to-canopy gas exchange under field conditions

Molecular clocks drive oscillations in leaf photosynthesis, stomatal conductance and other cell and leaf level processes over ~24 h under controlled laboratory conditions. The influence of such circadian regulation over whole canopy fluxes remains uncertain and diurnal CO2 and H2O vapor flux dynamics in the field are currently interpreted as resulting almost exclusively from direct physiological responses to variations in light, temperature and other environmental factors. We tested whether circadian regulation would affect plant and canopy gas exchange at the CNRS Ecotron. Canopy and leaf level fluxes were constantly monitored under field-like environmental conditions, and also under constant environmental conditions (no variation in temperature, radiation or other environmental cues). Here we show first direct experimental evidence at canopy scales of circadian gas exchange regulation: 20-79% of the daily variation range in CO2 and H2O fluxes occurred under circadian entrainment in canopies of an annual herb (bean) and of a perennial shrub (cotton). We also observed that considering circadian regulation improved performance in commonly used stomatal conductance models. Overall, our results show that overlooked circadian controls affect diurnal patterns of CO2 and H2O fluxes in entire canopies and in field-like conditions, although this process is currently unaccounted for in models.

Dynamic niche partitioning in root water uptake facilitates efficient water use in more diverse grassland plant communities

Summary 1.Efficient extraction of soil water is essential for the productivity of plant communities. However, research on the complementary use of resources in mixed plant communities, and especially the impact of plant species richness on root water uptake, is limited. So far, these investigations have been hindered by a lack of methods allowing for the estimation of root water uptake profiles. 2.The overarching aim of our study was to determine whether diverse grassland plant communities in general exploit soil water more deeply and whether this shift occurs all the time or only during times of enhanced water demand. 3.Root water uptake was derived by analyzing the diurnal decrease of soil water content separately at each measurement depth, thus yielding root water uptake profiles for 12 experimental grasslands communities with two different levels of species richness (4 and 16 sown species). Additional measurements of leaf water potential, stomatal conductance, and root traits were used to identify differences in water relations between plant functional groups. 4.Although the vertical root distribution did not differ between diversity levels, root water uptake shifted towards deeper layers (30 cm and 60 cm) in more diverse plots during periods of high vapor pressure deficit. Our results indicate that the more diverse communities were able to adjust their root water uptake, resulting in increased water uptake per root area compared to less diverse communities (52% at 20 cm, 118% at 30 cm, and 570% at 60 cm depth) and a more even distribution of water uptake over depth. Tall herbs, which had lower leaf water potential and higher stomatal conductance in more diverse mixtures, contributed disproportionately to dynamic niche partitioning in root water uptake. 5.This study underpins the role of diversity in stabilizing ecosystem function and mitigating drought stress effects during future climate change scenarios. Furthermore, the results provide evidence that root water uptake is not solely controlled by root length density distribution in communities with high plant diversity but also by spatial shifts in water acquisition. This article is protected by copyright. All rights reserved.

Functional composition rather than species richness drive carbon gain and allocation in experimental grasslands

Numerous experiments have shown positive diversity effects on plant productivity, but little is known about related processes of carbon gain and allocation. We investigated these processes in a controlled environment (Montpellier European Ecotron) applying a continuous 13CO2 label for three weeks to 12 soil-vegetation monoliths originating from a grassland biodiversity experiment (Jena Experiment) and representing two diversity levels (4 and 16 sown species). Plant species richness did not affect community- and species-level 13C abundances neither in total biomass nor in non-structural carbohydrates (NSC). Community-level 13C excess tended to be higher in the 16-species than in the 4-species mixtures. Community-level 13C excess was positively related to canopy leaf nitrogen (N), i.e. leaf N per unit soil surface. At the species level shoot 13C abundances varied among plant functional groups and were larger in legumes and tall herbs than in grasses and small herbs and correlated positively with traits as leaf N concentrations, stomatal conductance and shoot height. The 13C abundances in NSC were larger in transport sugars (sucrose, raffinose-family oligosaccharides) than in free glucose, fructose and compounds of the storage pool (starch) suggesting that newly assimilated carbon is to a small portion allocated to storage. Our results emphasize that the functional composition of communities is key in explaining carbon assimilation in grasslands.

Does circadian regulation lead to optimal gas exchange regulation

Optimal stomatal theory is an evolutionary model proposing that leaves trade-off Carbon (C) for water to maximise C assimilation (A) and minimise transpiration (E), thereby generating a marginal water cost of carbon gain (λ) that remains constant over short temporal scales. The circadian clock is a molecular timer of metabolism that controls A and stomatal conductance (gs), amongst other processes, in a broad array of plant species. Here, we test whether circadian regulation contributes towards achieving optimal stomatal behaviour. We subjected bean (Phaseolus vulgaris) and cotton (Gossypium hirsutum) canopies to fixed, continuous environmental conditions of photosynthetically active radiation, temperature and vapour pressure deficit over 48 hours. We observed a significant and self-sustained circadian oscillation in A and in stomatal conductance (gs) which also led to a circadian oscillation in λ. The lack of constant marginal water cost indicates that circadian regulation does not directly lead to optimal stomatal behaviour. However, the temporal pattern in gas exchange, indicative of either maximizing A or of minimizing E, depending upon time of day, indicates that circadian regulation could contribute towards optimizing stomatal responses. More broadly, our results add to the emerging field of plant circadian ecology and show that molecular controls may partially explain leaf-level patterns observed in the field.