Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of droughtinduced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function

Abstract. The global hydrological cycle is predicted to become more intense in future climates, with both larger precipitation events and longer times between events in some regions. Redistribution of precipitation may occur both within and across seasons, and the resulting wide fluctuations in soil water content (SWC) may dramatically affect plants. Though these responses remain poorly understood, recent research in this emerging field suggests the effects of redistributed precipitation may differ from predictions based on previous drought studies. We review available studies on both extreme precipitation (redistribution within seasons) and seasonal changes in precipitation (redistribution across seasons) on grasslands and forests.Extreme precipitation differentially affected above-ground net primary productivity (ANPP), depending on whether extreme precipitation led to increased or decreased SWC, which differed based on the current precipitation and aridity index of the site. Specifically, studies to date reported that extreme precipitation decreased ANPP in mesic sites, but, conversely, increased ANPP in xeric sites, suggesting that plant-available water is a key factor driving responses to extreme precipitation. Similarly, the effects of seasonal changes in precipitation on ANPP, phenology, and leaf and fruit development varied with the effect on SWC. Reductions in spring or summer generally had negative effects on plants, associated with reduced SWC, while subsequent reductions in autumn or winter had little effect on SWC or plants. Similarly, increased summer precipitation had a more dramatic impact on plants than winter increases in precipitation.The patterns of response suggest xeric biomes may respond positively to extreme precipitation, while comparatively mesic biomes may be more likely to be negatively affected. Moreover, seasonal changes in precipitation during warm or dry seasons may have larger effects than changes during cool or wet seasons. Accordingly, responses to redistributed precipitation will involve a complex interplay between plant-available water, plant functional type and resultant influences on plant phenology, growth and water relations. These results highlight the need for experiments across a range of soil types and plant functional types, critical for predicting future vegetation responses to future climates.

Total daily water use is a key factor influencing the growth of many terrestrial plants, and reflects both day-time and nocturnal water fluxes. However, while nocturnal sap flow (En) and stomatal conductance (gs,n) have been reported across a range of species, ecosystems and microclimatic conditions, the regulation of these fluxes remains poorly understood. Here, we present a framework describing the role of abiotic and biotic factors in regulating En and gs,n highlighting recent developments in this field. Across ecosystems, En and gs,n generally increased with increasing soil water content and vapor pressure deficit, but the interactive effects of these factors and the potential roles of wind speed and other abiotic factors remain unclear. On average, gs,n and En are higher in broad-leaved compared with needle-leaved plants, in C3 compared with C4 plants, and in tropical compared with temperate species. We discuss the impacts of leaf age, elevated [CO2] and refilling of capacitance on night-time water loss, and how nocturnal gs,n may be included in vegetation models. Younger leaves may have higher gs,n than older leaves. Embolism refilling and recharge of capacitance may affect sap flow such that total plant water loss at night may be less than estimated solely from En measurements. Our estimates of gs,n for typical plant functional types, based on the published literature, suggest that nocturnal water loss may be a significant fraction (10-25%) of total daily water loss. Counter-intuitively, elevated [CO2] may increase nocturnal water loss. Assumptions in process-based ecophysiological models and dynamic global vegetation models that gs is zero when solar radiation is zero are likely to be incorrect. Consequently, failure to adequately consider nocturnal water loss may lead to substantial under-estimation of total plant water use and inaccurate estimation of ecosystem level water balance.

To investigate if Eucalyptus species have responded to industrial-age climate change, and how they may respond to a future climate, we measured growth and physiology of fast-(E. saligna) and slow-growing (E. sideroxylon) seedlings exposed to preindustrial (290), current (400) or projected (650 lL L À1 ) CO 2 concentration ([CO 2 ]) and to current or projected (current 1 4 1C) temperature. To evaluate maximum potential treatment responses, plants were grown with nonlimiting soil moisture. We found that: (1) E. sideroxylon responded more strongly to elevated [CO 2 ] than to elevated temperature, while E. saligna responded similarly to elevated [CO 2 ] and elevated temperature; (2) the transition from preindustrial to current [CO 2 ] did not enhance eucalypt plant growth under ambient temperature, despite enhancing photosynthesis; (3) the transition from current to future [CO 2 ] stimulated both photosynthesis and growth of eucalypts, independent of temperature; and (4) warming enhanced eucalypt growth, independent of future [CO 2 ], despite not affecting photosynthesis. These results suggest large potential carbon sequestration by eucalypts in a future world, and highlight the need to evaluate how future water availability may affect such responses.

Summary• The response of nocturnal stomatal conductance (g s,n ) to rising atmospheric CO 2 concentration ([CO 2 ]) is currently unknown, and may differ from responses of daytime stomatal conductance (g s,d ). Because night-time water fluxes can have a significant impact on landscape water budgets, an understanding of the effects of [CO 2 ] and temperature on g s,n is crucial for predicting water fluxes under future climates.• Here, we examined the effects of [CO 2 ] (280, 400 and 640 lmol mol )1 ), temperature (ambient and ambient + 4°C) and drought on g s,n, and g s,d in Eucalyptus sideroxylon saplings.• g s,n was substantially higher than zero, averaging 34% of g s,d . Before the onset of drought, g s,n increased by 85% when [CO 2 ] increased from 280 to 640 lmol mol )1 , averaged across both temperature treatments. g s,n declined with drought, but an increase in [CO 2 ] slowed this decline. Consequently, the soil water potential at which g s,n was zero (W 0 ) was significantly more negative in elevated [CO 2 ] and temperature treatments. g s,d showed inconsistent responses to [CO 2 ] and temperature.• g s,n may be higher in future climates, potentially increasing nocturnal water loss and susceptibility to drought, but cannot be predicted easily from g s,d . Therefore, predictive models using stomatal conductance must account for both g s,n and g s,d when estimating ecosystem water fluxes.

We tested the hypothesis that mycorrhizal infection benefits wild plants to a lesser extent than cultivated plants. This hypothesis stems from two observations: (1) mycorrhizal infection improves plant growth primarily by increasing nutrient uptake, and (2) wild plants often possess special adaptations to soil infertility which are less pronounced in modern cultivated plants. In the first experiment, wild (Avena fatua L.) and cultivated (A. sativa L.) oats were grown hydroponically at four different phosphorus levels. Wild oat was less responsive (in shoot dry weight) to increasing phosphorus availability than cultivated oat. In addition, the root: shoot ratio was much more plastic in wild oat (varying from 0.90 in the low phosphorus solution to 0.25 in the high phosphorus solution) than in cultivated oat (varying from 0.44 to 0.17). In the second experiment, mycorrhizal and non-mycorrhizal wild and cultivated oats were grown in a phosphorus-deficient soil. Mycorrhizal infection generally improved the vegetative growth of both wild and cultivated oats. However, infection significantly increased plant lifespan, number of panicles per plant, shoot phosphorus concentration, shoot phosphorus content, duration of flowering, and the mean weight of individual seeds in cultivated oat, while it had a significantly reduced effect, no effect, or a negative effect on these characters for wild oat. Poor positive responsiveness of wild oat in these characters was thus associated with what might be considered to be inherent adaptations to nutrient deficiency: high root: shoot ratio and inherently low growth rate. Infection also increased seed phosphorus content and reproductive allocation.

To identify environmental and biological drivers of nocturnal vapour exchange, we quantified intra-annual, intra- and inter-specific variation in nocturnal water transport among ecologically diverse Eucalyptus species. We measured sap flux (J(s)) and leaf physiology (leaf surface conductance (g(s)), transpiration (E) and water potential (Psi(l))) in three to five trees of eight species. Over 1 year, nocturnal J(s) (J(s,n)) contributed 5-7% of total J(s) in the eight species. The principal environmental driver of J(s,n) was the product of atmospheric vapour pressure deficit (D) and wind speed (U). Selected observations suggest that trees with higher proportions of young foliage may exhibit greater J(s,n) and nocturnal g(s) (g(s,n)). Compared with other tree taxa, nocturnal water use in Eucalyptus was relatively low and more variable within than between species, suggesting that (i) Eucalyptus as a group exerts strong nocturnal stomatal control over water loss and (ii) prediction of nocturnal flux in Eucalyptus may depend on simultaneous knowledge of intra-specific tree traits and nocturnal atmospheric conditions.

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