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.     

Interactive effects of elevated CO2 and drought on nocturnal water fluxes in Eucalyptus saligna

Nocturnal water flux has been observed in trees under a variety of environmental conditions and can be a significant contributor to diel canopy water flux. Elevated atmospheric CO(2) (elevated [CO(2)]) can have an important effect on day-time plant water fluxes, but it is not known whether it also affects nocturnal water fluxes. We examined the effects of elevated [CO(2)] on nocturnal water flux of field-grown Eucalyptus saligna trees using sap flux through the tree stem expressed on a sapwood area (J(s)) and leaf area (E(t)) basis. After 19 months growth under well-watered conditions, drought was imposed by withholding water for 5 months in the summer, ending with a rain event that restored soil moisture. Reductions in J(s) and E(t) were observed during the severe drought period in the dry treatment under elevated [CO(2)], but not during moderate- and post-drought periods. Elevated [CO(2)] affected night-time sap flux density which included the stem recharge period, called 'total night flux' (19:00 to 05:00, J(s,r)), but not during the post-recharge period, which primarily consisted of canopy transpiration (23:00 to 05:00, J(s,c)). Elevated [CO(2)] wet (EW) trees exhibited higher J(s,r) than ambient [CO(2)] wet trees (AW) indicating greater water flux in elevated [CO(2)] under well-watered conditions. However, under drought conditions, elevated [CO(2)] dry (ED) trees exhibited significantly lower J(s,r) than ambient [CO(2)] dry trees (AD), indicating less water flux during stem recharge under elevated [CO(2)]. J(s,c) did not differ between ambient and elevated [CO(2)]. Vapour pressure deficit (D) was clearly the major influence on night-time sap flux. D was positively correlated with J(s,r) and had its greatest impact on J(s,r) at high D in ambient [CO(2)]. Our results suggest that elevated [CO(2)] may reduce night-time water flux in E. saligna when soil water content is low and D is high. While elevated [CO(2)] affected J(s,r), it did not affect day-time water flux in wet soil, suggesting that the responses of J(s,r) to environmental factors cannot be directly inferred from day-time patterns. Changes in J(s,r) are likely to influence pre-dawn leaf water potential, and plant responses to water stress. Nocturnal fluxes are clearly important for predicting effects of climate change on forest physiology and hydrology.     

Interspecific variation in nighttime transpiration and stomatal conductance in a mixed New England deciduous forest

Transpiration is generally assumed to be insignificant at night when stomata close in response to the lack of photosynthetically active radiation. However, there is increasing evidence that the stomata of some species remain open at night, which would allow for nighttime transpiration if there were a sufficient environmental driving force. We examined nighttime water use in co-occurring species in a mixed deciduous stand at Harvard Forest, MA, using whole-tree and leaf-level measurements. Diurnal whole-tree water use was monitored continuously with Granier-style sap flux sensors in paper birch (Betula papyrifera Marsh.), red oak (Quercus rubra L.) and red maple (Acer rubrum L.). An analysis was conducted in which nighttime water flux could be partitioned between refilling of internal water stores and transpiration. Substantial nighttime sap flux was observed in all species and much of this flux was attributed to the refilling of depleted water stores. However, in paper birch, nighttime sap flux frequently exceeded recharge estimates. Over 10% of the total daily sap flux during the growing season was due to transpiration at night in paper birch. Nighttime sap flux was over 8% of the total daily flux in red oak and 2% in red maple; however, this flux was mainly associated with recharge. On nights with elevated vapor pressure deficit, sap flux continued through the night in paper birch, whereas it reached zero during the night in red oak and red maple. Measurements of leaf-level gas exchange on a night with elevated vapor pressure deficit showed stomatal conductance dropping by only 25% in paper birch, while approaching zero in red oak and red maple. The study highlighted differences in ecophysiological controls on sap flux exerted by co-occurring species. Paper birch is a fast-growing, shade-intolerant species with an earlier successional status than red oak and red maple. Risking water loss through nighttime transpiration may provide paper birch with an ecological advantage by enabling the species to maximize photosynthesis and support rapid growth. Nighttime transpiration may also be a mechanism for delivering oxygen to respiring cells in the deep sapwood of paper birch.     

Sap-flux-scaled transpiration responses to light,vapor pressure deficit,and leaf area reduction in a flooded Taxodium distichum forest

We used 20-mm-long, Granier-type sensors to quantify the effects of tree size, azimuth and radial position in the xylem on the spatial variability in xylem sap flux in 64-year-old trees of Taxodium distichum L. Rich. growing in a flooded forest. This information was used to scale flux to the stand level to investigate variations in half-hourly and daily (24-hour) sums of sap flow, transpiration per unit of leaf area, and stand transpiration in relation to vapor pressure deficit (D) and photosynthetically active radiation (Q(o)). Measurements of xylem sap flux density (J(s)) indicated that: (1) J(s) in small diameter trees was 0.70 of that in medium and large diameter trees, but the relationship between stem diameter as a continuous variable and J(s) was not significant; (2) J(s) at 20-40 mm depth in the xylem was 0.40 of that at 0-20 mm depth; and (3) J(s) on the north side of trees was 0.64 of that in directions 120 degrees from the north. Daily transpiration was linearly related to daily daytime mean D, and reached a modest value of 1.3 mm day(-1), reflecting the low leaf area index (LAI = 2.2) of the stand. Because there was no soil water limitation, half-hourly water uptake was nearly linearly related to D at D < 0.6 kPa during both night and day, increasing to saturation during daytime at higher values of D. The positive effect of Q(o) on J(s) was significant, but relatively minor. Thus, a second-order polynomial with D explained 94% of the variation in J(s) and transpiration. An approximately 40% reduction in LAI by a hurricane resulted in decreases of about 18% in J(s) and stand transpiration, indicating partial stomatal compensation.     

Nocturnal sap flow characteristics and stem water recharge of Acacia mangium

In this paper, we studied the nocturnal stem water recharge of Acacia mangium. It is helpful to improve the precision of canopy transpiration estimation and canopy stomatal conductance, and to further understand the lag time of canopy transpiration to stem sap flow. In this study, the whole-tree sap flow in an A. mangium forest was measured by using Granier’s thermal dissipation probe for over two years in the hilly land of South China. The environmental factors, including relative humidity (RH), precipitation, vapor pressure deficit (VPD), photosynthetically active radiation (PAR), and air temperature (T _a) were recorded simultaneously. The stem water recharge of A. mangium was analyzed on both daily and monthly scales. Sap flux density was lower at night than during the day. The time range of nighttime sap flux density was longer in the dry season than in the wet season. The water recharging mainly occurred from sunset to midnight. No significant differences were observed among inter-annual nighttime water recharges. Nighttime water recharge had no significant correlation with environmental factors, but was well correlated with the diameter at breast height, tree height, and crown size. In the dry season the contribution of nighttime water recharge to total transpiration had significant correlations with daytime transpiration, total transpiration, VPD, PAR and T _a, while in the wet season it was significantly correlated with daily transpiration and total transpiration. __________ Translated from Chinese Journal of Ecology, 2007, 26(4): 476–482 [译自: 生态学杂志]     

Stomatal conductance, transpiration and sap flow of tropical montane rain forest trees in the southern Ecuadorian Andes

We investigated tree water relations in a lower tropical montane rain forest at 1950-1975 m a.s.l. in southern Ecuador. During two field campaigns, sap flow measurements (Granier-type) were carried out on 16 trees (14 species) differing in size and position within the forest stand. Stomatal conductance (g(s)) and leaf transpiration (E(l)) were measured on five canopy trees and 10 understory plants. Atmospheric coupling of stomatal transpiration was good (decoupling coefficient Omega = 0.25-0.43), but the response of g(s) and E(l) to the atmospheric environment appeared to be weak as a result of the offsetting effects of vapor pressure deficit (VPD) and photosynthetic photon flux (PPF) on g(s). In contrast, sap flow (F) followed these atmospheric parameters more precisely. Daily F depended chiefly on PPF sums, whereas on short time scales, VPD impeded transpiration when it exceeded a value of 1-1.2 kPa. This indicates an upper limit to transpiration in the investigated trees, even when soil water supply was not limiting. Mean g(s) was 165 mmol m(-2) s(-1) for the canopy trees and about 90 mmol m(-2) s(-1) for the understory species, but leaf-to-leaf as well as tree-to-tree variation was large. Considering whole-plant water use, variation in the daily course of F was more pronounced among trees differing in size and crown status than among species. Daily F increased sharply with stem diameter and tree height, and ranged between 80 and 120 kg day(-1) for dominant canopy trees, but was typically well below 10 kg day(-1) for intermediate and suppressed trees of the forest interior.     

Hydraulic responses of whole vines and individual roots of kiwifruit (Actinidia chinensis) following root severance

Whole vine (K(plant)) and individual root (K(root)) hydraulic conductances were measured in kiwifruit (Actinidia chinensis Planch. var. chinensis 'Hort16A') vines to observe hydraulic responses following partial root system excision. Heat dissipation and compensation heat pulse techniques were used to measure sap flow in trunks and individual roots, respectively. Sap flux and measurements of xylem pressure potential (Ψ) were used to calculate K(plant) and K(root) in vines with zero and ～80% of roots severed. Whole vine transpiration (E), Ψ and K(plant) were significantly reduced within 24 h of root pruning, and did not recover within 6 weeks. Sap flux in intact roots increased within 24 h of root pruning, driven by an increase in the pressure gradient between the soil and canopy and without any change in root hydraulic conductance. Photosynthesis (A) and stomatal conductance (g(s)) were reduced, without significant effects on leaf internal CO(2) concentration (c(i)). Shoot growth rates were maintained; fruit growth and dry matter content were increased following pruning. The woody roots of kiwifruit did not demonstrate a rapid dynamic response to root system damage as has been observed previously in monocot seedlings. Increased sap flux in intact roots with no change in K(root) and only a moderate decline in shoot A suggests that under normal growing conditions root hydraulic conductance greatly exceeds requirements for adequate shoot hydration.     

Nocturnal transpiration causing disequilibrium between soil and stem predawn water potential in mixed conifer forests of Idaho

Soil water potential (Psi(s)) is often estimated by measuring leaf water potential before dawn (Psi(pd)), based on the assumption that the plant water status has come into equilibrium with that of the soil. However, it has been documented for a number of plant species that stomata do not close completely at night, allowing for nocturnal transpiration and thus preventing nocturnal soil-plant water potential equilibration. The potential for nighttime transpiration necessitates testing the assumption of nocturnal equilibration before accepting Psi(pd) as a valid estimate of Psi(s). We determined the magnitude of disequilibrium between Psi(pd) and Psi(s) in four temperate conifer species across three height classes through a replicated study in northern Idaho. Based on both stomatal conductance and sap flux measurements, we confirmed that the combination of open stomata and high nocturnal atmospheric vapor pressure deficit (D) resulted in nocturnal transpiration in all four species. Nocturnal stomatal conductance (g(s-noc)) averaged about 33% of mid-morning conductance values. We used species-specific estimates of g(s-noc) and leaf specific conductance to correct Psi(pd) values for nocturnal transpiration at the time the samples were collected. Compared with the unadjusted values, corrected values reflected a significantly higher Psi(pd) (when D > 0.12 kPa). These results demonstrate that comparisons of Psi(pd) among species, canopy height classes and sites, and across growing seasons can be influenced by differential amounts of nocturnal transpiration, leading to flawed results. Consequently, it is important to account for the presence of nocturnal transpiration, either through a properly parameterized model or by making Psi(pd) measurements when D is sufficiently low that it cannot drive nocturnal transpiration. Violating these conditions will likely result in underestimation of Psi(s).     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

Environmental controls on sap flow in a northern hardwood forest

Our objective was to gain a detailed understanding of how photosynthetically active radiation (PAR), vapor pressure deficit (D) and soil water interact to control transpiration in the dominant canopy species of a mixed hardwood forest in northern Lower Michigan. An improved understanding of how these environmental factors affect whole-tree water use in unmanaged ecosystems is necessary in assessing the consequences of climate change on the terrestrial water cycle. We used continuously heated sap flow sensors to measure transpiration in mature trees of four species during two successive drought events. The measurements were scaled to the stand level for comparison with eddy covariance estimates of ecosystem water flux (Fw). Photosynthetically active radiation and D together explained 82% of the daytime hourly variation in plot-level transpiration, and low soil water content generally resulted in increased stomatal sensitivity to increasing D. There were also species-specific responses to drought. Quercus rubra L. showed low water use during both dry and wet conditions, and during periods of high D. Among the study species, Acer rubrum L. showed the greatest degree of stomatal closure in response to low soil water availability. Moderate increases in stomatal sensitivity to D during dry periods were observed in Populus grandidentata Michx. and Betula papyrifera Marsh. Sap flow scaled to the plot level and Fw demonstrated similar temporal patterns of water loss suggesting that the mechanisms controlling sap flow of an individual tree also control ecosystem evapotranspiration. However, the absolute magnitude of scaled sap flow estimates was consistently lower than Fw. We conclude that species-specific responses to PAR, D and soil water content are key elements to understanding current and future water fluxes in this ecosystem.     

Estimating sap flux densities in date palm trees using the heat dissipation method and weighing lysimeters

In a world of diminishing water reservoirs and a rising demand for food, the practice and development of water stress indicators and sensors are in rapid progress. The heat dissipation method, originally established by Granier, is herein applied and modified to enable sap flow measurements in date palm trees in the southern Arava desert of Israel. A long and tough sensor was constructed to withstand insertion into the date palm's hard exterior stem. This stem is wide and fibrous, surrounded by an even tougher external non-conducting layer of dead leaf bases. Furthermore, being a monocot species, water flow does not necessarily occur through the outer part of the palm's stem, as in most trees. Therefore, it is highly important to investigate the variations of the sap flux densities and determine the preferable location for sap flow sensing within the stem. Once installed into fully grown date palm trees stationed on weighing lysimeters, sap flow as measured by the modified sensors was compared with the actual transpiration. Sap flow was found to be well correlated with transpiration, especially when using a recent calibration equation rather than the original Granier equation. Furthermore, inducing the axial variability of the sap flux densities was found to be highly important for accurate assessments of transpiration by sap flow measurements. The sensors indicated no transpiration at night, a high increase of transpiration from 06:00 to 09:00, maximum transpiration at 12:00, followed by a moderate reduction until 08:00; when transpiration ceased. These results were reinforced by the lysimeters' output. Reduced sap flux densities were detected at the stem's mantle when compared with its center. These results were reinforced by mechanistic measurements of the stem's specific hydraulic conductivity. Variance on the vertical axis was also observed, indicating an accelerated flow towards the upper parts of the tree and raising a hypothesis concerning dehydrating mechanisms of the date palm tree. Finally, the sensors indicated reduction in flow almost immediately after irrigation of field-grown trees was withheld, at a time when no climatic or phenological conditions could have led to reduction in transpiration.     

Forest transpiration—targeted through xylem sap flux assessment versus hydrological modeling

The assessment of forest transpiration rates is crucial for determining plant-available soil water consumption and drought risk of trees. Xylem sap flux measurements have been used increasingly to quantify stand transpiration in forest ecosystems. Here, we compare this empirical approach with hydrological modeling on the basis of a stand transpiration dataset of adult beech (Fagus sylvatica), which was acquired across Bavaria, Germany, at eight forest sites. Xylem sap flux sensors were installed in five dominant trees each. Two tree to stand upscaling approaches, related to site-specific (1) sapwood area or (2) to leaf area index, were compared. The outcome was examined each in relation to process-based stand hydrological modeling, using LWF-BROOK90. Distinct relationships between tree diameter at breast height (1.30 m) and sapwood area-weighted sap flux along the radial profile became apparent across the study sites, confirming a generic allometric basis for stand-level upscaling of transpiration. The two upscaling approaches did not differ in outcome, representatively covering stand structure for comparison with modeling. Differential analysis yielded high agreement between the empirical and modeling approaches throughout most of the study period, although LWF-BROOK90 tended to overestimate sap flux measurements under low soil moisture. The two empirical approaches proved reliable for even-aged beech stands, as performance under high stand-structural heterogeneity awaits clarification. Findings advance stand-level hydrological modeling regarding coverage of stomatal behavior during temporary limitation in water availability.     

Analyses of assumptions and errors in the calculation of stomatal conductance from sap flux measurements

We analyzed assumptions and measurement errors in estimating canopy transpiration (E(L)) from sap flux (J(S)) measured with Granier-type sensors, and in calculating canopy stomatal conductance (G(S)) from E(L) and vapor pressure deficit (D). The study was performed in 12-year-old Pinus taeda L. stands with a wide range in leaf area index (L) and growth rate. No systematic differences in J(S) were found between the north and south sides of trees. However, J(S) in xylem between 20 and 40 mm from the cambium was 50 and 39% of J(S) in the outer 20-mm band of xylem in slow- and fast-growing trees, respectively. Sap flux measured in stems did not lag J(S) measured in branches, and time and frequency domain analyses of time series indicated that variability in J(S) in stems and branches is mostly explained by variation in D. Therefore, J(S) was used to estimate transpiration, after accounting for radial patterns. There was no difference between D and leaf-to-air vapor pressure gradient, and D did not have a vertical profile in stands of either low or high L suggesting a strong canopy-atmosphere coupling. Therefore, D estimated at one point in the canopy can be used to calculate G(S) in such stands. Given the uncertainties in J(S), relative humidity, and temperature measurements, to keep errors in G(S) estimates to less than 10%, estimates of G(S) should be limited to conditions in which D >/= 0.6 kPa.     

Diurnal patterns of water use in Eucalyptus victrix indicate pronounced desiccation-rehydration cycles despite unlimited water supply

Knowledge about nocturnal transpiration (E(night)) of trees is increasing and its impact on regional water and carbon balance has been recognized. Most of this knowledge has been generated in temperate or equatorial regions. Yet, little is known about E(night) and tree water use (Q) in semi-arid regions. We investigated the influence of atmospheric conditions on daytime (Q(day)) and nighttime water transport (Q(night)) of Eucalyptus victrix L.A.S. Johnson & K.D. Hill growing over shallow groundwater (not >1.5 m in depth) in semi-arid tropical Australia. We recorded Q(day) and Q(night) at different tree heights in conjunction with measurements of stomatal conductance (g(s)) and partitioned E(night) from refilling processes. Q of average-sized trees (200-400 mm diameter) was 1000-3000 l month(-1), but increased exponentially with diameter such that large trees (>500 mm diameter) used up to 8000 l month(-1). Q was remarkably stable across seasons. Water flux densities (J(s)) varied significantly at different tree heights during day and night. We show that g(s) remained significantly different from zero and E(night) was always greater than zero due to vapor pressure deficits (D) that remained >1.5 kPa at night throughout the year. Q(night) reached a maximum of 50% of Q(day) and was >0.03 mm h(-1) averaged across seasons. Refilling began during afternoon hours and continued well into the night. Q(night) eventually stabilized and closely tracked D(night). Coupling of Q(night) and D(night) was particularly strong during the wet season (R2?=?0.95). We suggest that these trees have developed the capacity to withstand a pronounced desiccation-rehydration cycle in a semi-arid environment. Such a cycle has important implications for local and regional hydrological budgets of semi-arid landscapes, as large nighttime water fluxes must be included in any accounting.     

Tree water storage and its diurnal dynamics related to sap flow and changes in stem volume in old-growth Douglas-fir trees

Diurnal and seasonal tree water storage was studied in three large Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) trees at the Wind River Canopy Crane Research site. Changes in water storage were based on measurements of sap flow and changes in stem volume and tissue water content at different heights in the stem and branches. We measured sap flow by two variants of the heat balance method (with internal heating in stems and external heating in branches), stem volume with electronic dendrometers, and tissue water content gravimetrically. Water storage was calculated from the differences in diurnal courses of sap flow at different heights and their integration. Old-growth Douglas-fir trees contained large amounts of free water: stem sapwood was the most important storage site, followed by stem phloem, branch sapwood, branch phloem and needles. There were significant time shifts (minutes to hours) between sap flow measured at different positions within the transport system (i.e., stem base to shoot tip), suggesting a highly elastic transport system. On selected fine days between late July and early October, when daily transpiration ranged from 150 to 300 liters, the quantity of stored water used daily ranged from 25 to 55 liters, i.e., about 20% of daily total sap flow. The greatest amount of this stored water came from the lower stem; however, proportionally more water was removed from the upper parts of the tree relative to their water storage capacity. In addition to lags in sap flow from one point in the hydrolic pathway to another, the withdrawal and replacement of stored water was reflected in changes in stem volume. When point-to-point lags in sap flow (minutes to hours near the top and stem base, respectively) were considered, there was a strong linear relationship between stem volume changes and transpiration. Volume changes of the whole tree were small (equivalent to 14% of the total daily use of stored water) indicating that most stored water came from the stem and from its inelastic (sapwood) tissues. Whole tree transpiration can be maintained with stored water for about a week, but it can be maintained with stored water from the upper crown alone for no more than a few hours.     

Size-mediated tree transpiration along soil drainage gradients in a boreal black spruce forest wildfire chronosequence

Boreal forests are crucial to climate change predictions because of their large land area and ability to sequester and store carbon, which is controlled by water availability. Heterogeneity of these forests is predicted to increase with climate change through more frequent wildfires, warmer, longer growing seasons and potential drainage of forested wetlands. This study aims at quantifying controls over tree transpiration with drainage condition, stand age and species in a central Canadian black spruce boreal forest. Heat dissipation sensors were installed in 2007 and data were collected through 2008 on 118 trees (69 Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce), 25 Populus tremuloides Michx. (trembling aspen), 19 Pinus banksiana Lamb. (jack pine), 3 Larix laricina (Du Roi) K. Koch (tamarack) and 2 Salix spp. (willow)) at four stand ages (18, 43, 77 and 157 years old) each containing a well- and poorly-drained stand. Transpiration estimates from sap flux were expressed per unit xylem area, J(S), per unit ground area, E(C) and per unit leaf area, E(L), using sapwood (A(S)) and leaf (A(L)) area calculated from stand- and species-specific allometry. Soil drainage differences in transpiration were variable; only the 43- and 157-year-old poorly-drained stands had?～?50% higher total stand E(C) than well-drained locations. Total stand E(C) tended to decrease with stand age after an initial increase between the 18- and 43-year-old stands. Soil drainage differences in transpiration were controlled primarily by short-term physiological drivers such as vapor pressure deficit and soil moisture whereas stand age differences were controlled by successional species shifts and changes in tree size (i.e., A(S)). Future predictions of boreal climate change must include stand age, species and soil drainage heterogeneity to avoid biased estimates of forest water loss and latent energy exchanges.     

Nighttime water use in an irrigated Eucalyptus grandis plantation

Sap flow measurements showed that a well-watered four-year-old plantation of Eucalyptus grandis (Hill ex Maiden) at Wagga Wagga, New South Wales, Australia, used 0.8 mm of water between 2100 and 0500 h on the midwinter night of July 30. Sap flow ceased for 2 to 3 h after sunset before recommencing at high rates that reached a maximum of 0.3 mm per h between 0200 and 0300 h. This pattern is inconsistent with the replenishment of tissue water reserves depleted during the day. Moreover, maximum leaf conductance at night was about 20 times maximum cuticular conductance values reported in the literature, which strongly suggests that stomata were partly open and that there was substantial water loss by way of the foliage. In an 8-month period from late winter to mid-autumn, comparable rates of nighttime water use were observed on only one other occasion. However, water use at rates of 0.1 mm per h or more occurred on 24 other nights. Almost 70% of the variation in nighttime sap velocity was explained by nighttime mean vapor pressure deficit and nighttime mean wind speed. Total recorded nighttime water use of the plantation was 48 mm, or 5% of total transpiration during the 8-month study. In view of the insensitivity of heat pulse measurements at low sap flows, this value may be an underestimate of actual nighttime transpiration.     

Estimating stand transpiration in a Eucalyptus populnea woodland with the heat pulse method: measurement errors and sampling strategies

Sap flow measurement techniques, such as the heat pulse (compensation) method, are practical means for estimating the water use of individual trees and are often the only reasonable alternative for measuring forest and woodland transpiration in complex heterogeneous terrain. The need to scale estimates of water use from a sample of individual stems to a stand (population) of known area may be satisfied by applying scalars of flux based on tree size or domain. We estimated the aggregate errors in applying the heat pulse technique to the estimation of stand transpiration in a poplar box (Eucalyptus populnea F.J. Muell.) woodland in southeastern Queensland, Australia, by a combination of precision analyses, experimental validation and Monte Carlo simulations of sampling errors. Errors in sap flux density measurements were approximately 13%. The potential error in the flux estimates for individual stems with stratified sampling of sap flux density with depth and bole quadrant based on four sensors was an additional 25%. Conducting wood area, diameter at 1.3 m, leaf area and domain based on Ecological Field Theory all proved excellent scalars of flux at the stand level. With a sample size of six trees stratified by diameter, coefficients of variation in scaling to the stand level were approximately 5% for any of these scalars. The greatest potential source of error in estimating stand transpiration by the heat pulse method was in the measurement of the fluxes of individual stems; scaling these measurements to a homogeneous stand of trees involved less uncertainty.     

Processes preventing nocturnal equilibration between leaf and soil water potential in tropical savanna woody species

The impact of nocturnal water loss and recharge of stem water storage on predawn disequilibrium between leaf (psiL) and soil (psiS) water potentials was studied in three dominant tropical savanna woody species in central Brazil (Cerrado). Sap flow continued throughout the night during the dry season and contributed from 13 to 28% of total daily transpiration. During the dry season, psiL was substantially less negative in covered transpiring leaves, throughout the day and night, than in exposed leaves. Before dawn, differences in psiL between covered and exposed leaves were about 0.4 MPa. When relationships between sap flow and psiL of exposed leaves were extrapolated to zero flow, the resulting values of psiL (a proxy of weighted mean soil water potential) in two of the species were similar to predawn values of covered leaves. Consistent with substantial nocturnal sap flow, stomatal conductance (gs) never dropped below 40 mmol m(-2) s(-1) at night, and in some cases, rose to as much as 100 mmol m(-2) s(-1) before the end of the dark period. Nocturnal gs decreased linearly with increasing air saturation deficit (D), but there were species-specific differences in the slopes of the relationships between nocturnal gs and D. Withdrawal and recharge of water from stem storage compartments were assessed by monitoring diel fluctuations of stem diameter with electronic dendrometers. Stem water storage compartments tended to recharge faster when nocturnal transpiration was reduced by covering the entire plant. Water potential of covered leaves did not stabilize in any of the plants before the end of the dark period, suggesting that, even in covered plants, water storage tissues were not fully rehydrated by dawn. Patterns of sap flow and expansion and contraction of stems reflected the dynamics of water movement during utilization and recharge of stem water storage tissues. This study showed that nighttime transpiration and recharge of internal water storage contribute to predawn disequilibrium in water potential between leaves and soil in neotropical savanna woody plants.     

Radial profiles of sap flow with increasing tree size in maritime pine

We investigated the radial variation of sap flow within sapwood below the live crown in relation to tree size in 10-, 32-, 54- and 91-year-old maritime pine stands (Pinus pinaster Ait.). Radial variations were determined with two thermal dissipation sensors; one measured sap flux in the outer 20 mm of the xylem (Jref), whereas the other was moved radially across the sapwood in 20-mm increments to measure sap flux at multiple depths (Jref). For all tree sizes, sap flow ratios (Ri = JiJref (-1)) declined with increasing sapwood depth, but the decrease was steeper in trees with large diameters. Correction factors (C) were calculated to extrapolate Jref for an estimate of whole-tree sap flux. A negative linear relationship was established between stem diameter and C, the latter ranging from 0.6 to 1.0. We found that neglecting these radial corrections in 10-, 32-, 54- and 91-year-old trees would lead to overestimation of stand transpiration by 4, 14, 26 and 47%, respectively. Therefore, it is necessary to account for the differential radial profiles of sap flow in relation to tree size when comparing tree transpiration and hydraulic properties among trees differing in size.