Nighttime Water Loss Research Articles

Nighttime transpiration represents a negligible part of water loss and does not increase the risk of water stress in grapevine.

Nighttime transpiration has been previously reported as a significant source of water loss in many species; however, there is a need to determine if this trait plays a key role in the response to drought. This study aimed to determine the magnitude, regulation and relative contribution to whole plant water‐use, of nighttime stomatal conductance (g night) and transpiration (E night) in grapevine (Vitis vinifera L.). Our results showed that nighttime water loss was relatively low compared to daytime transpiration, and that decreases in soil and plant water potentials were mainly explained by daytime stomatal conductance (g day) and transpiration (E day). Contrary to E day, E night did not respond to VPD and possible effects of an innate circadian regulation were observed. Plants with higher g night also exhibited higher daytime transpiration and carbon assimilation at midday, and total leaf area, suggesting that increased g night may be linked with daytime behaviors that promote productivity. Modeling simulations indicated that g night was not a significant factor in reaching critical hydraulic thresholds under scenarios of either extreme drought, or time to 20% of soil relative water content. Overall, this study suggests that g night is not significant in exacerbating the risk of water stress and hydraulic failure in grapevine.

Biophysical controls on nocturnal sap flow in plantation forests in a semi-arid region of northern China

Recent studies have recognized the importance of nocturnal sap flow (Qn) in affecting forest carbon and water budgets and responses to climate change at stand, regional, and global scales. However, biophysical controls on Qn are not fully understood, and their implications for land surface and vegetation models are unclear. We measured growing season sap flow of two widely distributed afforestation species, Pinus tabuliformis and Acer truncatum, in a middle-aged and a young monoculture forest stand, respectively, in a semi-arid mountainous area of northern China. We found a convergence in Qn between the two species and in the proportion of Qn to the total sap flow (12.2–15.0%) across species and ages. Total growing season Qn was higher for middle-aged stands than for young stands because of larger diameters at breast height of older stands. Nighttime vapor pressure deficit (VPDn) influence on Qn of young stands was soil moisture dependent. Nighttime wind speed indirectly controlled Qn through enhancing VPDn in young stands and directly promoted Qn in middle-aged stands with relatively low tree densities. For each species, both increased and decreased soil water content were able to promote Qn in stands with relatively dry soils, which might be due to enhanced nighttime water recharge for capacitance refilling and for avoiding hydraulic failure under prolonged water stress, respectively. Total effects of these three environmental factors explained less than 55% of the Qn variations. This study highlights uncertain physiological influences of VPDn on nighttime stomatal water loss, the nighttime water loss induced by wind, region-specific patterns of nighttime water recharge, and the importance of biotic controls on Qn. Our findings help to improve the existing VPD-based method for partitioning nighttime transpiration and water recharge at tree and stand levels, and suggest the importance of incorporating nocturnal sap flow into large-scale models.

Tolerance of ozone and drought in common waterhemp (Amaranthus tuberculatus)

ABSTRACTBiological invasions in established cropping systems impact biodiversity and agricultural sustainability. Under conditions of multi-year drought and elevated tropospheric ozone (O3), several Amaranthus species have recently become problematic weeds in California. We examined responses of common waterhemp (A. tuberculatus) to 12-h mean O3 exposures of 4, 59, or 114 nmol O3 mol air−1, with two levels of irrigation, 33% (deficit irrigation, DI) or 100% (well-watered, WW) of field capacity. Daytime stomatal conductance (gs) was reduced by O3 but not by DI. Nocturnal gs was uncommonly large but unaffected by either treatment. Plant productivity was not affected by O3 or DI. Leaf mass per unit area and root mass per unit leaf area were reduced by O3 but not by DI. Tolerance of this species to O3 and deficit irrigation may increase its competitiveness with crops. High night-time water loss may reduce water availability for competing crops. The competitiveness of common waterhemp may increase, as climate change creates conditions that are adverse to major crops.

Partitioning between evaporation and transpiration from Agrostis stolonifera L. during light and dark periods

Pressures on water availability for irrigation of turfgrasses continue in many parts of the United States as climate and weather patterns shift and populations increase. It is essential to understand underlying factors controlling water loss to more precisely predict irrigation requirements and develop new strategies for improving effective use of water. In this study, we investigate two key components of potential water loss from a bentgrass (Agrostis stolonifera L.) system that have not previously been examined in detail: 1) water loss in darkness, and 2) water loss through evaporation directly from the soil. The experiments were conducted in controlled environment chambers with intact cores from the field. An automated gravimetric system and soil moisture probes allowed precise measurements of water loss over ranges of vapor pressure deficits (VPD). The gravimetric and soil probe results indicated that substantial evapotranspiration occurred in darkness, at rates 40 to 60% of that in the light across VPDs. Simulations using field weather data from dry and humid environments indicated nighttime water loss rates would be expected to be 30 to 40% of that in the light. Using cores treated with a fast-acting, desiccating herbicide that eliminated transpiration but kept core resistances intact, evaporation directly from the soil surface was estimated to account for 40% of total water loss in the light and 60 to 70% in the dark. The results, collectively, indicated that water loss in darkness must be separately accounted for to accurately estimate daily evapotranspiration totals and irrigation requirements. Furthermore, because of the very high potential for evaporative water loss in the light and dark, efforts to improve water use efficiencies in the turfgrass system should include strategies that regulate both transpiration by the plant and evaporation from the soil surface.

The role of nighttime water balance on Olea europaea plants subjected to contrasting water regimes

The climate change scenarios besides foreseeing a severe drought imposition also emphasize the temperature rising in the Mediterranean region, with special prominence at nighttime. Despite the high olive tree tolerance to severe environmental conditions, stomatal nighttime water loss can change plant water relations, and the related consequences and opportunities, especially under water scarcity, must be clarified. A set of 3-year-old potted olive trees were subjected to three cycles of drought, imposed by withholding irrigation, while another group were continuously irrigated. At the end of the latter and more severe drought cycle, daytime gas exchange parameters, water status and membrane integrity was negatively affected by drought imposition. Moreover, the nighttime transpiration rate was far above cuticular water loss, suggesting sustained stomatal aperture during nighttime, leading to substantial water losses, which was higher under drought in the first hours of darkness. The higher nighttime stomatal conductance of droughted plants were related with higher starch concentration in their leaves, a thicker trichome layer and a lower intercellular CO2 concentration, in a closely association with an inferior nighttime respiration. Still, whole-plant transpiration on droughted plants were much lower than leaf transpiration-based estimates, which is interpreted as compensation by water inputs due to dew deposition on leaves. Although unexpected, the increased of stomatal conductance in the first hours of the night, until a certain level of water deficit intensity, could be linked with potential benefits to the plants.

Tolerance of Elevated Ozone and Water Stress in a California Population of Palmer Amaranth (Amaranthus palmeri)

Palmer amaranth is a highly invasive weed species causing huge economic losses in agricultural cropping systems under a broad range of environmental conditions. Sensitivity of this species to ozone (O3) air pollution and to soil water deficit, relative to native species or competing crops, may affect its competitiveness and invasive potential. In recent years, both high tropospheric O3and soil water deficiency have become common in the San Joaquin Valley of California. Responses to these environmental parameters may help predict the invasiveness of this species and have implications for landscape hydrology. We assessed the impact of O3and soil water deficit on Palmer amaranth. Five- to seven-leaf–stage potted plants were placed in continuous stirred tank reactor chambers and maintained for 30 to 35 d under 12-h mean daylight O3exposures (0700–1900 hours) of 4, 59, or 114 ppb O3. Within the chambers the plants were either well-watered (WW) or exposed to regulated deficit irrigation (RDI) and grown for about 7 wk. Dry weights of the leaves, stems, roots, and leaf area were determined. Day- and nighttime stomatal conductances (gs) were measured at 1.5-h intervals. Nocturnalgswas about 16 to 29% of daytimegs; this suggests that the species could have substantial nighttime water loss, uncoupled from carbon gain in the weed, and could affect water availability for crops and reduce irrigation efficiency. Nocturnalgswas lower in the RDI than in the WW, but daytimegswas not affected by O3or irrigation regime. Neither O3nor irrigation regime affected root or shoot parameters. As O3and drought are two key stressors in the San Joaquin Valley, to which potential competing species have been found to be sensitive, Palmer amaranth may proliferate and become more invasive in the future, potentially altering landscape hydrology and reducing irrigation efficiency.

Environmental controls in the water use patterns of a tropical cloud forest tree species, Drimys brasiliensis (Winteraceae).

Trees from tropical montane cloud forest (TMCF) display very dynamic patterns of water use. They are capable of downwards water transport towards the soil during leaf-wetting events, likely a consequence of foliar water uptake (FWU), as well as high rates of night-time transpiration (Enight) during drier nights. These two processes might represent important sources of water losses and gains to the plant, but little is known about the environmental factors controlling these water fluxes. We evaluated how contrasting atmospheric and soil water conditions control diurnal, nocturnal and seasonal dynamics of sap flow in Drimys brasiliensis (Miers), a common Neotropical cloud forest species. We monitored the seasonal variation of soil water content, micrometeorological conditions and sap flow of D. brasiliensis trees in the field during wet and dry seasons. We also conducted a greenhouse experiment exposing D. brasiliensis saplings under contrasting soil water conditions to deuterium-labelled fog water. We found that during the night D. brasiliensis possesses heightened stomatal sensitivity to soil drought and vapour pressure deficit, which reduces night-time water loss. Leaf-wetting events had a strong suppressive effect on tree transpiration (E). Foliar water uptake increased in magnitude with drier soil and during longer leaf-wetting events. The difference between diurnal and nocturnal stomatal behaviour in D. brasiliensis could be attributed to an optimization of carbon gain when leaves are dry, as well as minimization of nocturnal water loss. The leaf-wetting events on the other hand seem important to D. brasiliensis water balance, especially during soil droughts, both by suppressing tree transpiration (E) and as a small additional water supply through FWU. Our results suggest that decreases in leaf-wetting events in TMCF might increase D. brasiliensis water loss and decrease its water gains, which could compromise its ecophysiological performance and survival during dry periods.

Consequences of nocturnal water loss: a synthesis of regulating factors and implications for capacitance, embolism and use in models.

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.

Rates of nocturnal transpiration in two evergreen temperate woodland species with differing water-use strategies

Nocturnal fluxes may be a significant factor in the annual water budget of forested ecosystems. Here, we assessed sap flow in two co-occurring evergreen species (Eucalyptus parramattensis and Angophora bakeri) in a temperate woodland for 2 years in order to quantify the magnitude of seasonal nocturnal sap flow (E(n)) under different environmental conditions. The two species showed different diurnal water relations, demonstrated by different diurnal curves of stomatal conductance, sap flow and leaf water potential. The relative influence of several microclimatic variables, including wind speed (U), vapour pressure deficit (D), the product of U and D (UD) and soil moisture content, were quantified. D exerted the strongest influence on E(n) (r² = 0.59-0.86), soil moisture content influenced E(n) when D was constant, but U and UD did not generally influence E(n). In both species, cuticular conductance (G(c)) was a small proportion of total leaf conductance (G(s)) and was not a major pathway for E(n). We found that E(n) was primarily a function of transpiration from the canopy rather than refilling of stem storage, with canopy transpiration accounting for 50-70% of nocturnal flows. Mean E(n) was 6-8% of the 24-h flux across seasons (spring, summer and winter), but was up to 19% of the 24-h flux on some days in both species. Despite different daytime strategies in water use of the two species, both species demonstrated low night-time water loss, suggesting similar controls on water loss at night. In order to account for the impact of E(n) on pre-dawn leaf water potential arising from the influence of disequilibria between root zone and leaf water potential, we also developed a simple model to more accurately predict soil water potential (ψ(s)).

Soil nitrogen limitation does not impact nighttime water loss in Populus

Nighttime transpirational water loss from C(3) trees occurs without carbon gain and is both common and substantial. However, the magnitude of this water loss varies and a better understanding of the environmental factors driving this variation is needed. We investigated the response of nighttime conductance (g(night)) and transpiration (E(night)) to soil nitrogen limitation. We used instantaneous gas exchange measurements in greenhouse studies of Populus angustifolia James (narrowleaf cottonwood) and Populus balsamifera L. spp. trichocarpa (Torr. & A. Gray ex Hook.) Brayshaw (black cottonwood). g(night) for sufficiently watered plants ranged from 0.045 to 0.308 mol m(-2) s(-1) for P. balsamifera and 0.037 to 0.188 mol m(-2) s(-1) for P. angustifolia, which was much larger than minimum leaf conductance (g(min); up to 0.005 mol m(-2) s(-1) in the dark). Long-term nitrogen limitation sufficient to substantially reduce biomass did not affect g(night) or E(night) when potentially confounding water stress effects were eliminated. We conclude that nighttime water loss from two Populus species is large and although it is under stomatal control is not regulated at night in response to soil nitrogen availability.

Environmental stress and genetics influence night-time leaf conductance in the C4 grass Distichlis spicata.

Growing awareness of night-time leaf conductance (gnight) in many species, as well as genetic variation in gnight within several species, has raised questions about how genetic variation and environmental stress interact to influence the magnitude of gnight. The objective of this study was to investigate how genotype salt tolerance and salinity stress affect gnight for saltgrass [Distichlis spicata (L.) Greene]. Across genotypes and treatments, night-time water loss rates were 5-20% of daytime rates. Despite growth declining 37-87% in the high salinity treatments (300 mm and 600 mm NaCl), neither treatment had any effect on gnight in four of the six genotypes compared with the control treatment (7 mm NaCl). Daytime leaf conductance (gday) also was not affected by salinity treatment in three of the six genotypes. There was no evidence that more salt tolerant genotypes (assessed as ability to maintain growth with increasing salinity) had a greater capacity to maintain gnight or gday at high salinity. In addition, gnight as a percentage of gday was unaffected by treatment in the three most salt tolerant genotypes. Although gnight in the 7 mm treatment was always highest or not different compared with the 300 mm and 600 mm treatments, gday was generally highest in the 300 mm treatment, indicating separate regulation of gnight and gday in response to an environmental stress. Thus, it is clear that genetics and environment both influence the magnitude of gnight for this species. Combined effects of genetic and environmental factors are likely to impact our interpretation of variation of gnight in natural populations.

Genetic and Physiological Analysis of an Irradiated Bloomless Mutant (Epicuticular Wax Mutant) of Sorghum

An irradiation‐induced bloomless mutant of sorghum [Sorghum bicolor (L.) Moench.], KFS2021, which visually exhibits an absence of white fluffy epicuticular wax in leaf and sheath, was characterized using a combination of genetic and physiological approaches. Study of the phenotypic segregation for the bloomless trait in F2 and F2:3 populations from a cross between KFS2021 and BTx623 (a cultivar with bloom showing profuse deposition of white epicuticular wax) suggests that bloomless is controlled by a single nuclear recessive gene. The bloomless parent (KFS2021) and F2 individuals had lower frequency of guttation, leakier epidermal layer (based on percentage of chlorophyll leaching), and higher rate of seedling water loss than the BTx623 and F2 bloom individuals. Bloomless F2 individuals showed 3‐ to 6‐fold higher nighttime transpiration rates relative to F2 bloom individuals based on nighttime conductance. Correlation analysis showed significant negative associations between leaf epicuticular wax load with epidermal permeability and nighttime conductance, which indicate the important role of epicuticular wax in these traits. These results suggest that epicuticular wax may enhance water use efficiency of sorghum by regulating nighttime water loss.

What the towers don't see at night: nocturnal sap flow in trees and shrubs at two AmeriFlux sites in California

At the leaf scale, it is a long-held assumption that stomata close at night in the absence of light, causing transpiration to decrease to zero. Energy balance models and evapotranspiration equations often rely on net radiation as an upper bound, and some models reduce evapotranspiration to zero at night when there is no solar radiation. Emerging research is showing, however, that transpiration can occur throughout the night in a variety of vegetation types and biomes. At the ecosystem scale, eddy covariance measurements have provided extensive data on latent heat flux for a multitude of ecosystem types globally. Nighttime eddy covariance measurements, however, are generally unreliable because of low turbulence. If significant nighttime water loss occurs, eddy flux towers may be missing key information on latent heat flux. We installed and measured rates of sap flow by the heat ratio method (Burgess et al. 2001) at two AmeriFlux (part of FLUXNET) sites in California. The heat ratio method allows measurement and quantification of low rates of sap flow, including negative rates (i.e., hydraulic lift). We measured sap flow in five Pinus ponderosa Dougl. ex Laws. trees and three Arctostaphylos manzanita Parry and two Ceanothus cordulatus A. Kellog shrubs in the Sierra Nevada Mountains, and in five Quercus douglasii Hook and Arn. trees at an oak savanna in the Central Valley of California. Nocturnal sap flow was observed in all species, and significant nighttime water loss was observed in both species of trees. Vapor pressure deficit and air temperature were both well correlated with nighttime transpiration; the influence of wind speed on nighttime transpiration was insignificant at both sites. We distinguished between storage-tissue refilling and water loss based on data from Year 2005, and calculated the percentage by which nighttime transpiration was underestimated by eddy covariance measurements at both sites.

Significant transpirational water loss occurs throughout the night in field-grown tomato.

Incomplete stomatal closure at night can result in substantial water loss at times when photosynthetic carbon gain is not occurring in C3 and C4 plant species. To investigate the magnitude of nighttime water loss for a crop species in the field, measurements of nighttime water loss by tomato (Lycopersicon esculentum Mill. cv. Heinz 8892) were made by three methods: a field-scale lysimeter and two leaf-level instruments, an automated viscous flow porometer and a portable photosynthesis system. The portable photosynthesis system indicated nighttime transpiration of 10% of maximal daytime transpiration and the viscous flow porometer demonstrated partially open stomata. Integrated crop water loss during the dark, non-photosynthetic hours measured on the lysimeter was 3-10.8% of total daily water loss. In the glasshouse, a survey of closely related wild and cultivated tomato species showed that under ambient conditions nighttime transpiration varied within and among species and was 8-33% of maximal daytime transpiration. Implications of such a substantial fraction of total daily crop water use occurring during the night are significant in agronomic, environmental, and economic terms. Further, variation within and among species in nighttime water loss has implications for breeding to improve crop water use efficiency.

Helianthus nighttime conductance and transpiration respond to soil water but not nutrient availability.

We investigated the response of Helianthus species nighttime conductance (g(night)) and transpiration (E(night)) to soil nutrient and water limitations in nine greenhouse studies. The studies primarily used wild Helianthus annuus, but also included a commercial and early domesticate of H. annuus and three additional wild species (Helianthus petiolaris Nutt., Helianthus deserticola Heiser, and Helianthus anomalus Blake). Well-watered plants of all species showed substantial g(night) (0.023-0.225 mol m(-2) s(-1)) and E(night) (0.29-2.46 mmol m(-2) s(-1)) measured as instantaneous gas exchange. Based on the potential for transpiration to increase mass flow of mobile nutrients to roots, we hypothesized that g(night) and E(night) would increase under limiting soil nutrients but found no evidence of responses in all six studies testing this. Based on known daytime responses to water limitation, we hypothesized that g(night) and E(night) would decrease when soil water availability was limited, and results from all four studies testing this supported our hypothesis. We also established that stomatal conductance at night was on average 5 times greater than cuticular conductance. Additionally, g(night) and E(night) varied nocturnally and across plant reproductive stages while remaining relatively constant as leaves aged. Our results further the ability to predict conditions under which nighttime water loss will be biologically significant and demonstrate that for Helianthus, g(night) can be regulated.

Night-time conductance in C3 and C4 species: do plants lose water at night?

Significant night-time stomatal conductance and transpiration were found for 11 out of 17 species with a range of life histories (herbaceous annual, perennial grass, shrub, tree), photosynthetic pathways (C(3), C(4)), and habitats in the western United States. Across species and habitats, higher night-time conductance and transpiration were associated with higher daytime values. The prevalence, mechanisms and ecological implications of substantial night-time water loss deserve further investigation.

Thermal Energy Exchange Model and Water Loss of a Barrel Cactus, Ferocactus acanthodes

The influences of various diurnal stomatal opening patterns, spines, and ribs on the stem surface temperature and water economy of a CAM succulent, the barrel cactus Ferocactus acanthodes, were examined using an energy budget model. To incorporate energy exchanges by shortwave and longwave irradiation, latent heat, conduction, and convection as well as the heat storage in the massive stem, the plant was subdivided into over 100 internal and external regions in the model. This enabled the average surface temperature to be predicted within 1 C of the measured temperature for both winter and summer days.Reducing the stem water vapor conductance from the values observed in the field to zero caused the average daily stem surface temperature to increase only 0.7 C for a winter day and 0.3 C for a summer day. Thus, latent heat loss does not substantially reduce stem temperature. Although the surface temperatures averaged 18 C warmer for the summer day than for the winter day for a plant 41 cm tall, the temperature dependence of stomatal opening caused the simulated nighttime water loss rates to be about the same for the 2 days.Spines moderated the amplitude of the diurnal temperature changes of the stem surface, since the daily variation was 17 C for the winter day and 25 C for the summer day with spines compared with 23 C and 41 C, respectively, in their simulated absence. Ribs reduced the daytime temperature rise by providing 54% more area for convective heat loss than for a smooth circumscribing surface. In a simulation where both spines and ribs were eliminated, the daytime average surface temperature rose by 5 C.