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Aerosols can contribute to plant nutrition via foliar uptake. The conditions for this are best at night because the humidity is high and hygroscopic, saline deposits can deliquesce as a result. Still, stomata tend to be closed at night to avoid unproductive water loss. However, if needed, nutrients are on the leaf surface, and plants could benefit from nocturnal stomatal opening because it further increases humidity in the leaf boundary layer and allows for stomatal nutrient uptake. We tested this hypothesis on P-deficient soil by comparing the influence of ambient aerosols and additional foliar P application on nocturnal transpiration. We measured various related leaf parameters, such as the foliar water loss, minimum leaf conductance (gmin), turgor loss point, carbon isotope ratio, contact angle, specific leaf area (SLA), tissue element concentration, and stomatal and cuticular characteristics. For untreated leaves grown in filtered, aerosol-free air (FA), nocturnal transpiration consistently decreased overnight, which was not observed for leaves grown in unfiltered ambient air (AA). Foliar application of a soluble P salt increased nocturnal transpiration for AA and FA leaves. Crusts on stomatal rims were shown by scanning electron microscopy, supporting the idea of stomatal uptake of deliquescent salts. Turgor loss point and leaf moisture content indicated a higher accumulation of solutes, due to foliar uptake by AA plants than FA plants. The hypothesis that deliquescent leaf surface salts may play a role in triggering nocturnal transpiration was supported by the results. Still, further experiments are required to characterize this phenomenon better.

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BACKGROUND: Stomatal pores in many species are separated from the atmosphere by different anatomical obstacles produced by leaf epidermal cells, especially by sunken stomatal crypts, stomatal antechambers and/or hairs (trichomes). The evolutionary driving forces leading to sunken or 'hidden' stomata whose antechambers are filled with hairs or waxy plugs are not fully understood. The available hypothetical explanations are based mainly on mathematical modelling of water and CO2 diffusion through superficial vs. sunken stomata, and studies of comparative autecology. A better understanding of this phenomenon may result from examining the interactions between the leaf cuticle and stomata and from functional comparisons of sunken vs. superficially positioned stomata, especially when transpiration is low, for example at night or during severe drought. SCOPE: I review recent ideas as to why stomata are hidden and test experimentally whether hidden stomata may behave differently from those not covered by epidermal structures and so are coupled more closely to the atmosphere. I also quantify the contribution of stomatal vs. cuticular transpiration at night using four species with sunken stomata and three species with superficial stomata. CONCLUSIONS: Partitioning of leaf conductance in darkness (gtw) into stomatal and cuticular contributions revealed that stomatal conductance dominated gtw across all seven investigated species with antechambers with different degrees of prominence. Hidden stomata contributed, on average, less to gtw (approx. 70 %) than superficial stomata (approx. 80 %) and reduced their contribution dramatically with increasing gtw. In contrast, species with superficial stomata kept their proportion in gtw invariant across a broad range of gtw. Mechanisms behind the specific behaviour of hidden stomata and the multipurpose origin of sunken stomata are discussed.

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Nocturnal transpiration is widely observed across species and biomes, and may significantly impact global water, carbon, and energy budgets. However, it remains elusive why plants lose water at night and how to model it at large scales. We hypothesized that plants optimize nighttime leaf diffusive conductance (gwn ) to balance potential daytime photosynthetic benefits and nocturnal transpiration benefits. We quantified nighttime benefits from respiratory reductions due to evaporative leaf cooling. We described nighttime costs in terms of a reduced carbon gain during the day because of water use at night. We measured nighttime stomatal responses and tested our model with water birch (Betula occidentalis) saplings grown in a glasshouse. The gwn of water birch decreased with drier soil, higher atmospheric CO2 , wetter air, lower leaf temperature, and lower leaf respiration rate. Our model predicted all these responses correctly, except for the response of gwn to air humidity. Our results also suggested that the slow decrease in gwn after sunset could be associated with decreasing leaf respiration. The optimality-based nocturnal transpiration model smoothly integrates with daytime stomatal optimization approaches, and thus has the potential to quantitatively predict nocturnal transpiration across space and time.

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Non-negligible nighttime transpiration rates (TRN) have been identified in grasses such as wheat and barley. Evidence from the last 30 years indicate that in drought-prone environments with high evaporative demand, TRN could amount to 8-55% of daytime TR, leading several investigators to hypothesize that reducing TRN might represent a viable water-saving strategy that minimizes seemingly 'wasteful' water loss that is not traded for CO2 fixation. More recently however, evidence suggests that actual increases in TRN during pre-dawn hours, which are presumably controlled by the circadian clock, mediate drought tolerance - not through water conservation - but by enabling maximized gas exchange early in the morning before midday depression sets in. Finally, new findings point to a previously undocumented role for leaf sheaths as substantial contributors (up to 45%) of canopy TRN, although the extent of their involvement in these two strategies remains unknown. In this paper, we synthesize and reconcile key results from experimental and simulation-based modeling efforts conducted at scales ranging from the leaf tissue to the field plot on wheat and barley to show that both strategies could in fact concomitantly enable yield gains under limited water supply. We propose a simple framework highlighting the role played by TRN dynamics in drought tolerance and provide a synthesis of potential research directions, with an emphasis on the need for further examining the role played by the circadian clock and leaf sheath gas exchange.

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MAIN CONCLUSION: Nocturnal transpiration, through its circadian control, plays a role in modulating daytime transpiration response to increasing evaporative demand, to potentially enable drought tolerance in wheat. Limiting plant transpiration rate (TR) in response to increasing vapor pressure deficit (VPD) has been suggested to enable drought tolerance through water conservation. However, there is very little information on the extent of diversity of TR response curves to "true" VPD (i.e., independent from temperature). Furthermore, new evidence indicate that water-saving could operate by modulating nocturnal TR (TRN), and that this response might be coupled to daytime gas exchange. Based on 3 years of experimental data on a diverse group of 77 genotypes from 25 countries and 5 continents, a first goal of this study was to characterize the functional diversity in daytime TR responses to VPD and TRN in wheat. A second objective was to test the hypothesis that these traits could be coupled through the circadian clock. Using a new gravimetric phenotyping platform that allowed for independent temperature and VPD control, we identified three and fourfold variation in daytime and nighttime responses, respectively. In addition, TRN was found to be positively correlated with slopes of daytime TR responses to VPD, and we identified pre-dawn variation in TRN that likely mediated this relationship. Furthermore, pre-dawn increase in TRN positively correlated with the year of release among drought-tolerant Australian cultivars and with the VPD threshold at which they initiated water-saving. Overall, the study indicates a substantial diversity in TR responses to VPD that could be leveraged to enhance fitness under water-limited environments, and that TRN and its circadian control may play an important role in the expression of water-saving.

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During the night, plant water loss can occur either through the roots, as hydraulic redistribution (HR), or through the leaves via the stoma, as nocturnal transpiration (En), which was methodologically difficult to separate from stem refilling (Re). While HR and En have been reported across a range of species, ecosystem, and climate zone, there is little understanding on the interactions between En and/or Re and HR. As water movement at night occurs via gradients of water potential, it is expected that during periods of high atmospheric vapor pressure deficit (VPD), water loss via En will override water loss via HR. To test this hypothesis, sap flow in stems and roots of Populus euphratica Oliv. trees, growing in a riparian zone in a hyperarid climate, was measured once in a year. Nocturnal stem sap flow was separated into En and Re using the "forecasted refilling" method. Substantial nocturnal sap flow (38% of 24-hr flux on average) was observed and positively correlated with VPD; however, the strength of the correlation was lower (R2 = .55) than diurnal sap flow (Ed) (R2 = .72), suggesting that nocturnal stem sap flow was attributed to both water loss through the canopy and replenishment of water in stem tissues. Partitioning of nocturnal sap flow shows that Re constituted approximately 80%, and En ~20%, of nocturnal sap flow. The amount of root sap flow attributed to redistribution was negatively related to Ed (R2 = .69) and the amount of acropetally sap flow in stems, Re (R2 = .41) and En (R2 = .14). It was suggested that the magnitude of HR is more strongly depressed by Re that was recharge to the water loss via Ed than by En. It was consistent with whole-tree water balance theory, that the nighttime upward sap flow to xylem, stem refilling and transpiration, may depress hydraulic redistribution of roots.

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Hydraulic lift, the passive movement of water through plant roots from wet to dry soil, is an important ecohydrological process in a wide range of water-limited ecosystems. This phenomenon may also alter plant functioning, growth, and survival in mesic grasslands, where soil moisture is spatially and temporally variable. Here, we monitored diurnal changes in the isotopic signature of soil and plant xylem water to assess (1) whether hydraulic lift occurs in woody and herbaceous tallgrass prairie species (Rhus glabra, Amorpha canescens, Vernonia baldwinii, and Andropogon gerardii), (2) if nocturnal transpiration or grazing by large ungulates limits hydraulic lift, and (3) if a dominant grass, A. gerardii, utilizes water lifted by other tallgrass prairie species. Broadly, the results shown here suggest that hydraulic lift does not appear to be widespread or common in this system, but isolated instances suggest that this process does occur within tallgrass prairie. The isolated instance of hydraulic lift did not vary by grazing treatment, nor did they result in facilitation for neighboring grasses. We suggest that the topographic complexity of this tallgrass prairie and the high rates of nocturnal transpiration observed in this study likely limit the frequency and occurrence of hydraulic lift. These results suggest that hydraulic lift can be a patchy process, particularly in heterogeneous landscapes.

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In addition to buffering plants from water stress during severe droughts, plant water storage (PWS) alters many features of the spatio-temporal dynamics of water movement in the soil-plant system. How PWS impacts water dynamics and drought resilience is explored using a multi-layer porous media model. The model numerically resolves soil-plant hydrodynamics by coupling them to leaf-level gas exchange and soil-root interfacial layers. Novel features of the model are the considerations of a coordinated relationship between stomatal aperture variation and whole-system hydraulics and of the effects of PWS and nocturnal transpiration (Fe,night) on hydraulic redistribution (HR) in the soil. The model results suggest that daytime PWS usage and Fe,night generate a residual water potential gradient (Δψp,night) along the plant vascular system overnight. This Δψp,night represents a non-negligible competing sink strength that diminishes the significance of HR. Considering the co-occurrence of PWS usage and HR during a single extended dry-down, a wide range of plant attributes and environmental/soil conditions selected to enhance or suppress plant drought resilience is discussed. When compared with HR, model calculations suggest that increased root water influx into plant conducting-tissues overnight maintains a more favorable water status at the leaf, thereby delaying the onset of drought stress.

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Circadian resonance, whereby a plant's endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C3 plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole-plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian-regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.

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Nighttime transpiration is a substantial portion of ecosystem water budgets, but few studies compare water use of closely related co-occurring species in a phylogenetic context. Nighttime transpiration can range up to 69% of daytime rates and vary between species, ecosystem, and functional type. We examined leaf-level daytime and nighttime gas exchange of five species of the genus Rubus co-occurring in the Pacific Northwest of western North America in a greenhouse common garden. Contrary to expectations, nighttime transpiration was not correlated to daytime water use. Nighttime transpiration showed pronounced phylogenetic signals, but the proportion of variation explained by different phylogenetic groupings varied across datasets. Leaf osmotic water potential, water potential at turgor loss point, stomatal size, and specific leaf area were correlated with phylogeny but did not readily explain variation in nighttime transpiration. Patterns in interspecific variation as well as a disconnect between rates of daytime and nighttime transpiration suggest that variation in nighttime water use may be at least partly driven by genetic factors independent of those that control daytime water use. Future work with co-occurring congeneric systems is needed to establish the generality of these results and may help determine the mechanism driving interspecific variation in nighttime water use.

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Nocturnal sap flow (Qn) has been found to occur across many taxa, seasons and biomes. There is no general understanding as to how much Qn occurs and whether it is a significant contribution to total daily sap flow (Q). A synthesis of the literature and unpublished data was made to determine how significant is Qn, as a proportion of Q (%Qn), across seasons, biomes, phylogenetic groups and different thermometric sap flow methods. A total of 98 species were analysed to find that %Qn, on average, was 12.03% with the highest average dataset of 69.00%. There was significantly less %Qn in winter than in other temperate seasons, and significantly less %Qn in the wet season than in the dry season. The equatorial and tropical biomes had significantly higher %Qn than the warm temperate and nemoral biomes. The heat ratio method (HRM) and the thermal dissipation (TDP) method had significantly higher %Qn than the heat balance method. Additional analysis between HRM and TDP found HRM to have significantly higher %Qn in winter, wet season and various biomes. In all but one out of 246 cases Qn occurred, demonstrating that Qn is significant and needs to be carefully considered in sap flow and related studies.

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We measured sap flux (S) and environmental variables in four monospecific stands of alpine ash (Eucalyptus delegatensis R. Baker, AA) and snowgum (E. pauciflora Sieb. ex Spreng., SG) in Australia's Victorian Alps. Nocturnal S was 11.8 ± 0.8% of diel totals. We separated transpiration (E) and refilling components of S using a novel modeling approach based on refilling time constants. The nocturnal fraction of diel water loss (f(n)) averaged 8.6 ± 0.6% for AA and 9.8 ± 1.7% for SG; f(n) differed among sites but not species. Evaporative demand (D) was the strongest driver of nocturnal E (E(n)). The ratio E(n)/D (G(n)) was positively correlated to soil moisture in most cases, whereas correlations between wind speed and G(n) varied widely in sign and strength. Our results suggest (1) the large, mature trees at our subalpine sites have greater f(n) than the few Australian native tree species that have been studied at lower elevations, (2) AA and SG exhibit similar f(n) despite very different size and life history, and (3) f(n) may differ substantially among sites, so future work should be replicated across differing sites. Our novel approach to quantifying f(n) can be applied to S measurements obtained by any method.