A new experimental setup to measure hydraulic conductivity of plant segments.

Plant hydraulic conductivity and its decline under water stress are the focal point of current plant hydraulic research. The common methods of measuring hydraulic conductivity control a pressure gradient to push water through plant samples, submitting them to conditions far away from those that are experienced in nature where flow is suction driven and determined by the leaf water demand. In this paper, we present two methods for measuring hydraulic conductivity under closer to natural conditions, an artificial plant setup and a horizontal syringe pump setup. Both approaches use suction to pull water through a plant sample while dynamically monitoring the flow rate and pressure gradients. The syringe setup presented here allows for controlling and rapidly changing flow and pressure conditions, enabling experimental assessment of rapid plant hydraulic responses to water stress. The setup also allows quantification of dynamic changes in water storage of plant samples. Our tests demonstrate that the syringe pump setup can reproduce hydraulic conductivity values measured using the current standard method based on pushing water under above-atmospheric pressure. Surprisingly, using both the traditional and our new syringe pump setup, we found a positive correlation between changes in flow rate and hydraulic conductivity. Moreover, when flow or pressure conditions were changed rapidly, we found substantial contributions to flow by dynamic and largely reversible changes in the water storage of plant samples. Although the measurements can be performed under sub-atmospheric pressures, it is not possible to subject the samples to negative pressures due to the presence of gas bubbles near the valves and pressure sensors. Regardless, this setup allows for unprecedented insights into the interplay between pressure, flow rate, hydraulic conductivity and water storage in plant segments. This work was performed using an Open Science approach with the original data and analysis to be found at https://doi.org/10.5281/zenodo.7322605.

Downwelling longwave radiation and sensible heat flux observations are critical for surface temperature and emissivity estimation from flux tower data.

Land surface temperature (LST) is a preeminent state variable that controls the energy and water exchange between the Earth's surface and the atmosphere. At the landscape-scale, LST is derived from thermal infrared radiance measured using space-borne radiometers. In contrast, plot-scale LST estimation at flux tower sites is commonly based on the inversion of upwelling longwave radiation captured by tower-mounted radiometers, whereas the role of the downwelling longwave radiation component is often ignored. We found that neglecting the reflected downwelling longwave radiation leads not only to substantial bias in plot-scale LST estimation, but also have important implications for the estimation of surface emissivity on which LST is co-dependent. The present study proposes a novel method for simultaneous estimation of LST and emissivity at the plot-scale and addresses in detail the consequences of omitting down-welling longwave radiation as frequently done in the literature. Our analysis uses ten eddy covariance sites with different land cover types and found that the LST values obtained using both upwelling and downwelling longwave radiation components are 0.5-1.5 K lower than estimates using only upwelling longwave radiation. Furthermore, the proposed method helps identify inconsistencies between plot-scale radiometric and aerodynamic measurements, likely due to footprint mismatch between measurement approaches. We also found that such inconsistencies can be removed by slight corrections to the upwelling longwave component and subsequent energy balance closure, resulting in realistic estimates of surface emissivity and consistent relationships between energy fluxes and surface-air temperature differences. The correspondence between plot-scale LST and landscape-scale LST depends on site-specific characteristics, such as canopy density, sensor locations and viewing angles. Here we also quantify the uncertainty in plot-scale LST estimates due to uncertainty in tower-based measurements using the different methods. The results of this work have significant implications for the combined use of aerodynamic and radiometric measurements to understand the interactions and feedbacks between LST and surface-atmosphere exchange processes.

Gross primary productivity and water use efficiency are increasing in a high rainfall tropical savanna.

Despite their size and contribution to the global carbon cycle, we have limited understanding of tropical savannas and their current trajectory with climate change and anthropogenic pressures. Here we examined interannual variability and externally forced long-term changes in carbon and water exchange from a high rainfall savanna site in the seasonal tropics of north Australia. We used an 18-year flux data time series (2001-2019) to detect trends and drivers of fluxes of carbon and water. Significant positive trends in gross primary productivity (GPP, 15.4 g C m2  year-2 ), ecosystem respiration (Reco , 8.0 g C m2  year-2 ), net ecosystem productivity (NEE, 7.4 g C m2  year-2 ) and ecosystem water use efficiency (WUE, 0.0077 g C kg H2 O-1  year-1 ) were computed. There was a weaker, non-significant trend in latent energy exchange (LE, 0.34 W m-2  year-1 ). Rainfall from a nearby site increased statistically over a 45-year period during the observation period. To examine the dominant drivers of changes in GPP and WUE, we used a random forest approach and a terrestrial biosphere model to conduct an attribution experiment. Radiant energy was the dominant driver of wet season fluxes, whereas soil water content dominated dry season fluxes. The model attribution suggested that [CO2 ], precipitation and Tair accounting for 90% of the modelled trend in GPP and WUE. Positive trends in fluxes were largest in the dry season implying tree components were a larger contributor than the grassy understorey. Fluxes and environmental drivers were not significant during the wet season, the period when grasses are active. The site is potentially still recovering from a cyclone 45 years ago and regrowth from this event may also be contributing to the observed trends in sequestration, highlighting the need to understand fluxes and their drivers from sub-diurnal to decadal scales.

Organizing principles for vegetation dynamics.

Plants and vegetation play a critical-but largely unpredictable-role in global environmental changes due to the multitude of contributing processes at widely different spatial and temporal scales. In this Perspective, we explore approaches to master this complexity and improve our ability to predict vegetation dynamics by explicitly taking account of principles that constrain plant and ecosystem behaviour: natural selection, self-organization and entropy maximization. These ideas are increasingly being used in vegetation models, but we argue that their full potential has yet to be realized. We demonstrate the power of natural selection-based optimality principles to predict photosynthetic and carbon allocation responses to multiple environmental drivers, as well as how individual plasticity leads to the predictable self-organization of forest canopies. We show how models of natural selection acting on a few key traits can generate realistic plant communities and how entropy maximization can identify the most probable outcomes of community dynamics in space- and time-varying environments. Finally, we present a roadmap indicating how these principles could be combined in a new generation of models with stronger theoretical foundations and an improved capacity to predict complex vegetation responses to environmental change.

Two sides to every leaf: water and CO2 transport in hypostomatous and amphistomatous leaves.

Leaves with stomata on both upper and lower surfaces, termed amphistomatous, are relatively rare compared with hypostomatous leaves with stomata only on the lower surface. Amphistomaty occurs predominantly in fast-growing herbaceous annuals and in slow-growing perennial shrubs and trees. In this paper, we present the current understanding and hypotheses on the costs and benefits of amphistomaty related to water and CO2 transport in contrasting leaf morphologies. First, there is no evidence that amphistomatous species achieve higher stomatal densities on a projected leaf area basis than hypostomatous species, but two-sided gas exchange is less limited by boundary layer effects. Second, amphistomaty may provide a specific advantage in thick leaves by shortening the pathway for CO2 transport between the atmosphere and the chloroplasts. In thin leaves of fast-growing herbaceous annuals, in which both the adaxial and abaxial pathways are already short, amphistomaty enhances leaf-atmosphere gas-exchange capacity. Third, amphistomaty may help to optimise the leaf-interior water status for CO2 transport by reducing temperature gradients and so preventing the condensation of water that could limit CO2 diffusion. Fourth, a potential cost of amphistomaty is the need for additional investments in leaf water transport tissue to balance the water loss through the adaxial surface.

Wind increases leaf water use efficiency.

A widespread perception is that, with increasing wind speed, transpiration from plant leaves increases. However, evidence suggests that increasing wind speed enhances carbon dioxide (CO2 ) uptake while reducing transpiration because of more efficient convective cooling (under high solar radiation loads). We provide theoretical and experimental evidence that leaf water use efficiency (WUE, carbon uptake per water transpired) commonly increases with increasing wind speed, thus improving plants' ability to conserve water during photosynthesis. Our leaf-scale analysis suggests that the observed global decrease in near-surface wind speeds could have reduced WUE at a magnitude similar to the increase in WUE attributed to global rise in atmospheric CO2 concentrations. However, there is indication that the effect of long-term trends in wind speed on leaf gas exchange may be compensated for by the concurrent reduction in mean leaf sizes. These unintuitive feedbacks between wind, leaf size and water use efficiency call for re-evaluation of the role of wind in plant water relations and potential re-interpretation of temporal and geographic trends in leaf sizes.

Soil Penetration by Earthworms and Plant Roots--Mechanical Energetics of Bioturbation of Compacted Soils.

We quantify mechanical processes common to soil penetration by earthworms and growing plant roots, including the energetic requirements for soil plastic displacement. The basic mechanical model considers cavity expansion into a plastic wet soil involving wedging by root tips or earthworms via cone-like penetration followed by cavity expansion due to pressurized earthworm hydroskeleton or root radial growth. The mechanical stresses and resulting soil strains determine the mechanical energy required for bioturbation under different soil hydro-mechanical conditions for a realistic range of root/earthworm geometries. Modeling results suggest that higher soil water content and reduced clay content reduce the strain energy required for soil penetration. The critical earthworm or root pressure increases with increased diameter of root or earthworm, however, results are insensitive to the cone apex (shape of the tip). The invested mechanical energy per unit length increase with increasing earthworm and plant root diameters, whereas mechanical energy per unit of displaced soil volume decreases with larger diameters. The study provides a quantitative framework for estimating energy requirements for soil penetration work done by earthworms and plant roots, and delineates intrinsic and external mechanical limits for bioturbation processes. Estimated energy requirements for earthworm biopore networks are linked to consumption of soil organic matter and suggest that earthworm populations are likely to consume a significant fraction of ecosystem net primary production to sustain their subterranean activities.

Using an optimality model to understand medium and long-term responses of vegetation water use to elevated atmospheric CO2 concentrations.

Vegetation has different adjustable properties for adaptation to its environment. Examples include stomatal conductance at short time scale (minutes), leaf area index and fine root distributions at longer time scales (days-months) and species composition and dominant growth forms at very long time scales (years-decades-centuries). As a result, the overall response of evapotranspiration to changes in environmental forcing may also change at different time scales. The vegetation optimality model simulates optimal adaptation to environmental conditions, based on the assumption that different vegetation properties are optimized to maximize the long-term net carbon profit, allowing for separation of different scales of adaptation, without the need for parametrization with observed responses. This paper discusses model simulations of vegetation responses to today's elevated atmospheric CO2 concentrations (eCO2) at different temporal scales and puts them in context with experimental evidence from free-air CO2 enrichment (FACE) experiments. Without any model tuning or calibration, the model reproduced general trends deduced from FACE experiments, but, contrary to the widespread expectation that eCO2 would generally decrease water use due to its leaf-scale effect on stomatal conductance, our results suggest that eCO2 may lead to unchanged or even increased vegetation water use in water-limited climates, accompanied by an increase in perennial vegetation cover.

Stomatal control and leaf thermal and hydraulic capacitances under rapid environmental fluctuations.

Leaves within a canopy may experience rapid and extreme fluctuations in ambient conditions. A shaded leaf, for example, may become exposed to an order of magnitude increase in solar radiation within a few seconds, due to sunflecks or canopy motions. Considering typical time scales for stomatal adjustments, (2 to 60 minutes), the gap between these two time scales raised the question whether leaves rely on their hydraulic and thermal capacitances for passive protection from hydraulic failure or over-heating until stomata have adjusted. We employed a physically based model to systematically study effects of short-term fluctuations in irradiance on leaf temperatures and transpiration rates. Considering typical amplitudes and time scales of such fluctuations, the importance of leaf heat and water capacities for avoiding damaging leaf temperatures and hydraulic failure were investigated. The results suggest that common leaf heat capacities are not sufficient to protect a non-transpiring leaf from over-heating during sunflecks of several minutes duration whereas transpirative cooling provides effective protection. A comparison of the simulated time scales for heat damage in the absence of evaporative cooling with observed stomatal response times suggested that stomata must be already open before arrival of a sunfleck to avoid over-heating to critical leaf temperatures. This is consistent with measured stomatal conductances in shaded leaves and has implications for water use efficiency of deep canopy leaves and vulnerability to heat damage during drought. Our results also suggest that typical leaf water contents could sustain several minutes of evaporative cooling during a sunfleck without increasing the xylem water supply and thus risking embolism. We thus submit that shaded leaves rely on hydraulic capacitance and evaporative cooling to avoid over-heating and hydraulic failure during exposure to typical sunflecks, whereas thermal capacitance provides limited protection for very short sunflecks (tens of seconds).

Maximum entropy production allows a simple representation of heterogeneity in semiarid ecosystems.

Feedbacks between water use, biomass and infiltration capacity in semiarid ecosystems have been shown to lead to the spontaneous formation of vegetation patterns in a simple model. The formation of patterns permits the maintenance of larger overall biomass at low rainfall rates compared with homogeneous vegetation. This results in a bias of models run at larger scales neglecting subgrid-scale variability. In the present study, we investigate the question whether subgrid-scale heterogeneity can be parameterized as the outcome of optimal partitioning between bare soil and vegetated area. We find that a two-box model reproduces the time-averaged biomass of the patterns emerging in a 100 x 100 grid model if the vegetated fraction is optimized for maximum entropy production (MEP). This suggests that the proposed optimality-based representation of subgrid-scale heterogeneity may be generally applicable to different systems and at different scales. The implications for our understanding of self-organized behaviour and its modelling are discussed.

A canopy-scale test of the optimal water-use hypothesis.

Common empirical models of stomatal conductivity often incorporate a sensitivity of stomata to the rate of leaf photosynthesis. Such a sensitivity has been predicted on theoretical terms by Cowan and Farquhar, who postulated that stomata should adjust dynamically to maximize photosynthesis for a given water loss. In this study, we implemented the Cowan and Farquhar hypothesis of optimal stomatal conductivity into a canopy gas exchange model, and predicted the diurnal and daily variability of transpiration for a savanna site in the wet-dry tropics of northern Australia. The predicted transpiration dynamics were then compared with observations at the site using the eddy covariance technique. The observations were also used to evaluate two alternative approaches: constant conductivity and a tuned empirical model. The model based on the optimal water-use hypothesis performed better than the one based on constant stomatal conductivity, and at least as well as the tuned empirical model. This suggests that the optimal water-use hypothesis is useful for modelling canopy gas exchange, and that it can reduce the need for model parameterization.

A test of the optimality approach to modelling canopy properties and CO2 uptake by natural vegetation.

Photosynthesis provides plants with their main building material, carbohydrates, and with the energy necessary to thrive and prosper in their environment. We expect, therefore, that natural vegetation would evolve optimally to maximize its net carbon profit (NCP), the difference between carbon acquired by photosynthesis and carbon spent on maintenance of the organs involved in its uptake. We modelled N(CP) for an optimal vegetation for a site in the wet-dry tropics of north Australia based on this hypothesis and on an ecophysiological gas exchange and photosynthesis model, and compared the modelled CO2 fluxes and canopy properties with observations from the site. The comparison gives insights into theoretical and real controls on gas exchange and canopy structure, and supports the optimality approach for the modelling of gas exchange of natural vegetation. The main advantage of the optimality approach we adopt is that no assumptions about the particular vegetation of a site are required, making it a very powerful tool for predicting vegetation response to long-term climate or land use change.