Chronic cork oak decline and water status: new insights

Chronic decline and Sudden death are two syndromes of cork oak (Quercus suber) dieback. Mortality is associated with water stress, but underlying physiological mechanisms are poorly understood. Here, we investigated the physiological performance of declining trees during the summer drought. Leaf water potential, gas-exchange, fluorescence of photosystem II and leaf and root starch concentration were compared in healthy (asymptomatic) and declining trees. Low annual cork increment in declining trees indicated tree decline for several years. All trees showed similar water status in spring. In summer, declining trees showed lower predawn leaf water potential (?2.0 vs. ?0.8 MPa), but unexpectedly higher midday leaf water potential than healthy trees (?2.8 vs. ?3.3 MPa). The higher midday water potential was linked to by means of strongly reduced stomatal conductance and, consequently, transpiration. This study is pioneer showing that declining trees had high midday water potential. A tendency for lower sap flow driving force (the difference between predawn and midday water potential) in declining trees was also associated with reduced photosynthesis, suggesting that chronic dieback may be associated with low carbon uptake. However, starch in roots and leaves was very low and not correlated to the health status of trees. Declining trees showed lower water-use efficiency and non-photochemical quenching in summer, indicating less resistance to drought. Contrarily to chronic decline, one tree that underwent sudden death presented predawn leaf water potential below the cavitation threshold.     

Xylem sap flow and drought stress of Fraxinus excelsior saplings

We report on diurnal and seasonal variations in sap flow rate and stem water potential of Fraxinus excelsior L. saplings growing at the edge of a Fraxino-Aceretum forest in western Germany. Because of shallow soil, the trees were subjected to drought in summer. When soil water availability was not limiting, sap flow rate was related to changes in solar radiation and vapor pressure deficit. Maximum transpiration rates per leaf area were 3.5-7.4 mmol m-2 s-1, and maximum daily totals were 1.7-3.3 kg m-2 day-1. Under drought conditions, stem water potential dropped to midday minima of -2.6 to -3.5 MPa and sap flow rate was strongly related to this parameter. After the drought period, reduced apparent (whole-plant) hydraulic conductance was observed, which was attributed to a continued reduction in stomatal conductance after the drought stress had ceased. A model was developed that linked sap flow rate directly to climatic variables and stem water potential. Good correlation between measured and simulated sap flow rates allowed the model to be used for data interpretation.     

Transpiration of silver Fir (Abies alba mill.) during and after drought in relation to soil properties in a Mediterranean mountain area

Context

Silver fir is declining and dying at its southern margin on the Mediterranean area, where climate is expected to be warmer and drier. At a regional scale, silver fir seems to be vulnerable to drought, and at a forest stand scale, tree death seems to be distributed according to soil water availability.

Aim

To understand the vulnerability of silver fir to drought, factors involved in the regulation of transpiration were assessed with respect to soil properties in order to document the spatial distribution of death rates.

Methods

Soil properties were characterized by electric resistivity measurements. Sap flow density and predawn needle water potential were recorded on sampled trees during several years, and crown specific transpiration was estimated. In addition, the vulnerability of coarse roots and branches to cavitation was quantified.

Results and conclusion

Trees growing on soils with a large water storage capacity were the most vulnerable to drought induced soil water deficits. Transpiration was down-regulated as soon as predawn water potential decreased. The vulnerability to cavitation was low, which protected the trees from run-away xylem embolism. Severe soil water deficits led to a rapid decrease of transpiration, which was still visible the following year. The drop-off in transpiration was mainly due to inner sap flow that almost ceased after the drought on all monitored trees. Our results suggest that root dynamic and the ability of roots to take up water were modified by soil water deficit over several years. Such a regulation scheme needs to be better documented and included in models to address silver fir forest responses to drought.     

Impact of soil drought on sap flow and water status of evergreen trees in a tropical monsoon forest in northern Thailand

Hill evergreen forest is the dominant vegetation type in northern Thailand. In this region, there is higher atmospheric evaporative demand and lower soil moisture during the 5- to 7-month dry season than in the rainy season under influences from Asian monsoons. In an earlier study we revealed that canopy-scale transpiration is actively maintained even during the latter part of the dry season in hill evergreen forest. However, the impact of soil drought on tree water use was not investigated. To clarify the ecohydrological processes at this site, we used individual tree-scale measurements during a 2-year period to base our examination of whether limited water use in individual trees is caused by soil drought in the latter part of the dry season. Sap flow and water potential measurements were conducted in four evergreen trees, two large emergent trees 29.8 and 25.4 m high, and two smaller understory trees 4.8 and 1.4 m high.The amount of rainfall preceding the late dry season of 2004 was significantly less than that preceding the late dry season of 2003. Although a distinct decrease in sap-flow velocities in individual trees due to soil water stress was not found in the late dry season of 2003, it did become comparatively apparent in the late dry season of 2004; ranging from 10 to 40% for a given atmospheric evaporative demand. Furthermore, the reductions in sap-flow velocities and predawn stem-water potential were most significant in the smallest tree. The recovery of sap-flow velocities and water potential in the smallest tree after irrigation confirmed that the reductions in sap-flow velocity and predawn stem-water potential in the smallest tree were caused by soil drought. These results suggest that shallower roots could be reason for the significant decrease in water use in the smallest trees. The deeper roots of larger trees could be the reason for the reduced impact of soil drought on water use in larger trees, and canopy-scale transpiration might be maintained by larger trees, even in an unusually severe drought. These possibilities provide a new insight for management of evergreen forests under Asian monsoon influences.     

Responses of deciduous forest trees to severe drought in Central Europe

In 2003, Central Europe experienced the warmest summer on record combined with unusually low precipitation. We studied plant water relations and phenology in a 100-year- old mixed deciduous forest on a slope (no ground water table) near Basel using the Swiss Canopy Crane (SCC). The drought lasted from early June to mid September. We studied five deciduous tree species; half of the individuals were exposed to elevated CO(2) concentration ([CO(2)]) (530 ppm) using a free-air, atmospheric CO(2)-enrichment system. In late July, after the first eight weeks of drought, mean predawn leaf water potential about 30 m above ground was -0.9 MPa across all trees, dropping to a mean of -1.5 MPa in mid-August when the top 1 m of the soil profile had no plant accessible moisture. Mean stomatal conductance and rates of maximum net photosynthesis decreased considerably in mid-August across all species. However, daily peak values of sap flow remained surprisingly constant over the whole period in Quercus petraea (Matt.) Liebl., and decreased to only about half of the early summer maxima in Fagus sylvatica L. and Carpinus betulus L. (stomatal down- regulation of flux). Although we detected no differences in most parameters between CO(2)-treated and control trees, predawn leaf water potential tended to be less negative in trees exposed to elevated [CO(2)]. Leaf longevity was greater in 2003 compared with the previous years, but the seasonal increase in stem basal area reached only about 75% of that in previous years. Our data suggest that the investigated tree species, particularly Q. petraea, did not experience severe water stress. However, an increased frequency of such exceptionally dry summers may have a more serious impact than a single event and would give Q. petraea a competitive advantage in the long run.     

Rapid response of large, drought-stressed beech trees to irrigation

Large, declining beech (Fagus sylvatica L.) trees (diameter at breast height = 50 cm), growing on heavy clay soils in the highlands near Zurich, Switzerland, were amply irrigated in late summer. During irrigation, the xylem sap flow rate, Q(wt), was measured by the stem-tissue heat balance method with internal heating and sensing. Only a gradual and slight increase in Q(wt) in response to irrigation was observed in the control trees, whereas Q(wt) in the declining trees, whose transpiration rates were only 2-20% those of the control trees, increased 2-5 times within minutes. This suggests, that severe local drought was the major factor limiting tree growth at the site. The extent of the response permits estimation of the supply-limited (soil water) and demand-limited (tree structure) components of stress. Drought caused a decline in Q(wt) in the trees with short crowns and limited root systems that had originally been growing in dense canopies and had become suddenly exposed to full illumination as a result of a severe wind storm and thinning. Trees with deep, narrow, dense crowns, growing in more open places and adapted over a long period to high irradiance remained healthy during drought. Prolonged, periodic water shortage reduced the amount of foliage up to 90% but during drought stimulated the growth of fine roots in the surface and upper soil layers. The stem conductive systems of the declining trees were still partially functional.     

Sap flow rates of Minquartia guianensis in central Amazonia during the prolonged dry season of 2015–2016

Minquartia guianensis Aubl.is a slow-growing species with several uses.In the juvenile state,it is well-adapted to low light conditions of the forest understory.However,it is still unknown how climate variability affects transpiration of this species,particularly under drought stress.In this study,we aimed to assess the effect of climatic variability on sap flow rates(SFR).SFR and radial growth were measured in six trees(14-50 cm diameter) in 2015 and 2016.Climate(precipitation,irradiance,relative humidity and temperature) and soil water content(SWC) data were also collected.SFR tended to increase in the dry season,with a negative relationship between SFR and SWC and precipitation(p 0.001),while there was a positive association between radial growth and monthly precipitation(p=0.004).Irradiance and temperature were the environmental factors more closely correlated with SFR during daytime(p0.001),whereas relative humidity and vapor pressure deficit were the most important factors at night(p0.001).Although negative SFR were sometimes recorded at night,the mean nocturnal sap flow was positive and across trees the nighttime sap flow accounted for 12.5%of the total daily sap flow.Increased transpiration during the dry season suggests that the root system of Minquartia was able to extract water from deep soil layers.These results widen our understanding of the ecophysiology of Amazonian trees under drought and provide further insight into the potential effect of the forecasted decline in precipitation in the Amazon region.     

Canopy and hydraulic conductance in young, mature and old Douglas-fir trees

We tested for reductions in water transport with increasing tree size, a key component in determining whether gas exchange and growth are hydraulically limited in tall trees. During the summers of 1998 and 1999, we measured water transport with Granier-type, constant-heat sap flow probes, vapor pressure deficit, and leaf and soil water potentials in overstory Pseudotsuga menziesii (Mirb.) Franco trees in three stands differing in size and age (15, 32 and 60 m in height and about 20, 40 and 450 years in age, respectively) in a P. menziesii-dominated forest in the Pacific Northwest, USA. A total of 24 trees were equipped with sap flow sensors--six 60-m trees, nine 32-m trees and nine 15-m trees. Based on the sap flow measurements and leaf area information estimated from leaf area-sapwood area relationships, we estimated crown-averaged stomatal conductance (GS) and leaf-specific hydraulic conductance (KL). We tested the hypothesis that GS and KL vary inversely with tree height (15 > 32 > 60 m). Analysis of variance of GS ranked as 15 = 60 > 32 m during the early summer and 15 > 60 > 32 m during late season drought. Over the growing season, mean daily GS (+/- SE) was 29.2 +/- 4.4, 24.0 +/- 6.8 and 17.7 +/- 7.2 mmol m-2 s-1 for the 15-, 60- and 32-m trees, respectively. The value of K(L) differed among tree heights only during late season drought and ranked 15 > 32 = 60 m. A hydraulic mass balance suggests that greater sapwood conductivity in 60-m trees compared with 32- and 15-m trees is a likely cause for the departure of the above rankings from those predicted by height and leaf-to-sapwood area ratio.     

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.     

Stored water use and transpiration in Scots pine: a modeling analysis with ANAFORE

We estimated daily use of stored water by Scots pine (Pinus sylvestris L.) trees growing in a temperate climate with the ANAFORE model (ANAlysis of FORest Ecosystems) and compared the simulation results with sap flow measurements. The original model was expanded with a dynamic water flow and storage model that simulates sap flow dynamics in an individual tree. ANAFORE was able to accurately simulate diurnal patterns of measured sap flow under microclimatic conditions that differ from those of the calibration period. Strong relationships were found between stored water use and several tree characteristics (diameter at breast height, sapwood area, leaf area), but not with tree height. Relative to transpiration, stored water use varied over time (between < 1% and 44% of daily transpiration). On days when transpiration was high, trees were more dependent on stored water, indicating that the contribution of internal water to transpiration is not a constant in the water budget of trees.     

Comparative water use by the riparian trees Melaleuca argentea and Corymbia bella in the wet-dry tropics of northern Australia

We examined sources of water and daily and seasonal water use patterns in two riparian tree species occupying contrasting niches within riparian zones throughout the wet-dry tropics of northern Australia: Corymbia bella Hill and Johnson is found along the top of the levee banks and Melaleuca argentea W. Fitzg. is restricted to riversides. Patterns of tree water use (sap flow) and leaf water potential were examined in four trees of each species at three locations along the Daly River in the Northern Territory. Predawn leaf water potential was higher than -0.5 MPa throughout the dry season in both species, but was lower at the end of the dry season than at the beginning of the dry season. Contrary to expectations, predawn leaf water potential was lower in M. argentea trees along the river than in C. bella trees along the levees. In contrast, midday leaf water potential was lower in the C. bella trees than in M. argentea trees. There were no seasonal differences in tree water use in either species. Daily water use was lower in M. argentea trees than in C. bella trees. Whole-tree hydraulic conductance, estimated from the slope of the relationship between leaf water potential and sap flow, did not differ between species. Xylem deuterium concentrations indicated that M. argentea trees along the riverbank were principally reliant on river water or shallow groundwater, whereas C. bella trees along the levee were reliant solely on soil water reserves. This study demonstrated strong gradients of tree water use within tropical riparian communities, with implications for estimating riparian water use requirements and for the management of groundwater resources.     

An alternative method to estimate zero flow temperature differences for Granier's thermal dissipation technique

Calibration of the Granier thermal dissipation technique for measuring stem sap flow in trees requires determination of the temperature difference (DeltaT) between a heated and an unheated probe when sap flow is zero (DeltaT(max)). Classically, DeltaT(max) has been estimated from the maximum predawn DeltaT, assuming that sap flow is negligible at nighttime. However, because sap flow may continue during the night, the maximum predawn DeltaT value may underestimate the true DeltaT(max). No alternative method has yet been proposed to estimate DeltaT(max) when sap flow is non-zero at night. A sensitivity analysis is presented showing that errors in DeltaT(max) may amplify through sap flux density computations in Granier's approach, such that small amounts of undetected nighttime sap flow may lead to large diurnal sap flux density errors, hence the need for a correct estimate of DeltaT(max). By rearranging Granier's original formula, an optimization method to compute DeltaT(max) from simultaneous measurements of diurnal DeltaT and micrometeorological variables, without assuming that sap flow is negligible at night, is presented. Some illustrative examples are shown for sap flow measurements carried out on individuals of Erica arborea L., which has needle-like leaves, and Myrica faya Ait., a broadleaf species. We show that, although DeltaT(max) values obtained by the proposed method may be similar in some instances to the DeltaT(max) predicted at night, in general the values differ. The procedure presented has the potential of being applied not only to Granier's method, but to other heat-based sap flow systems that require a zero flow calibration, such as the Cermák et al. (1973) heat balance method and the T-max heat pulse system of Green et al. (2003).     

Investigation of change in tangential strain on the inner bark of the stem and root ofBetula platyphylla var.japonica andAcer mono during sap season

Although it is well known that sap exudation during early spring in temperate deciduous trees occurs in response to daytime warming and nighttime cooling, the mechanisms of the process are not yet fully understood. Previous theories suggested that changes in stress in the wood caused by daytime heating and nighttime cooling might be linked with sap flow. Consequently, a study of itaya-kaede maple (Aver mono) and shirakamba birch (Betula platyphylla var.japonica) looked at tangential strains. One-hour intervals for 3 years of the tangential strains on the inner bark of stem and root were measured in itaya-kaede maple and shirakamba birch during the sap exudation season. The measurements indicated different mechanisms of sap exudation in these two trees. During the sap exudation season in late March, when the temperature fluctuated around 0°C, the tangential strain in the root of itaya-kaede maple showed expansion in the daytime and contraction at night. Conversely, in early April the tangential strain in the root of shirakamba birch exhibited contraction in the daytime and expansion at night. The changes in tangential strains in itaya-kaede maple were attributed to conditioning, a known concept used to explain the uptake mechanism of soil water in maple and its exudation during early spring. However, because the change in tangential strain in the roots of shirakamba birch was similar to that found during the rampant season, sap exudation was not attributed to conditioning but to the plentiful supply of water from the roots. The implications of these mechanisms are that different sap harvesting techniques may be appropriate for different tree species.     

Stable annual pattern of water use by Acacia tortilis in Sahelian Africa

Water use by mature trees of Acacia tortilis (Forsk.) Hayne ssp. raddiana (Savi) Brenan var. raddiana growing in the northern Sahel was continuously recorded over 4 years. Water use was estimated from xylem sap flow measured by transient heat dissipation. Concurrently, cambial growth, canopy phenology, leaf water potential, climatic conditions and soil water availability (SWA) were monitored. In addition to the variation attributable to interannual variation in rainfall, SWA was increased by irrigation during one wet season. The wet season lasted from July to September, and annual rainfall ranged between 146 and 367 mm. The annual amount and pattern of tree water use were stable from year-to-year despite interannual and seasonal variations in SWA in the upper soil layers. Acacia tortilis transpired readily throughout the year, except for one month during the dry season when defoliation was at a maximum. Maximum water use of about 23 l (dm sapwood area)(-2) day(-1) was recorded at the end of the wet season. While trees retained foliage in the dry season, the decline in water use was modest at around 30%. Variation in predawn leaf water potential indicated that the trees were subject to soil water constraint. The rapid depletion of water in the uppermost soil layers after the wet season implies that there was extensive use of water from deep soil layers. The deep soil profile revealed (1) the existence of living roots at 25 m and (2) that the availability of soil water was low (-1.6 MPa) down to the water table at a depth of 31 m. However, transpiration was recorded at a predawn leaf water potential of -2.0 MPa, indicating that the trees used water from both intermediary soil layers and the water table. During the full canopy stage, mean values of whole-tree hydraulic conductance were similar in the wet and dry seasons. We propose that the stability of water use at the seasonal and annual scales resulted from a combination of features, including an extensive rooting habit related to deep water availability and an effective regulation of canopy conductance. Despite a limited effect on tree water use, irrigation during the wet season sharply increased predawn leaf water potential and cambial growth of trunks and branches.     

Water-use strategies in two co-occurring Mediterranean evergreen oaks: surviving the summer drought

In the Mediterranean evergreen oak woodlands of southern Portugal, the main tree species are Quercus ilex ssp. rotundifolia Lam. (holm oak) and Quercus suber L. (cork oak). We studied a savannah-type woodland where these species coexist, with the aim of better understanding the mechanisms of tree adaptation to seasonal drought. In both species, seasonal variations in transpiration and predawn leaf water potential showed a maximum in spring followed by a decline through the rainless summer and a recovery with autumn rainfall. Although the observed decrease in predawn leaf water potential in summer indicates soil water depletion, trees maintained transpiration rates above 0.7 mm day(-1) during the summer drought. By that time, more than 70% of the transpired water was being taken from groundwater sources. The daily fluctuations in soil water content suggest that some root uptake of groundwater was mediated through the upper soil layers by hydraulic lift. During the dry season, Q. ilex maintained higher predawn leaf water potentials, canopy conductances and transpiration rates than Q. suber. The higher water status of Q. ilex was likely associated with their deeper root systems compared with Q. suber. Whole-tree hydraulic conductance and minimum midday leaf water potential were lower in Q. ilex, indicating that Q. ilex was more tolerant to drought than Q. suber. Overall, Q. ilex seemed to have more effective drought avoidance and drought tolerance mechanisms than Q. suber.     

Effects of drought stress and high density stem inoculations with Leptographium wingfieldii on hydraulic properties of young Scots pine trees

We examined drought-induced changes in susceptibility of potted Scots pine (Pinus sylvestris L.) trees to a bark-beetle associated fungus (Leptographium wingfieldii Morelet, from the bark beetle Tomicus piniperda L.). Five-year-old field-grown trees were transplanted to 50-l pots and grown for 1 year before the treatments were applied. Trees in the drought treatment were subjected to several successive, 3-week-long drought cycles, with predawn water potential dropping below -2 MPa at peak drought intensity. The experimental drought cycles were more severe than the natural drought episodes usually recorded in Scots pine stands. Trees were then mass-inoculated with L. wingfieldii at a density close to the critical threshold density of inoculations (400 m(-2)) above which tree resistance is overcome. Inoculation of well-watered trees resulted in induced reaction zones around the inoculation points and very limited damage (resinosis) in the sapwood. Drought alone had no long-lasting consequences on tree water relations, except for a decrease in hydraulic conductance in the youngest segments of the main stem. However, the combination of mass-inoculation and drought stress after inoculation resulted in a dramatic loss of stem hydraulic conductivity that was paralleled by conspicuous damage to the sapwood (resinosis, drying and blue staining). There was a close correlation between amount of visible sapwood damage and loss of hydraulic conductivity. The intensity of induced reactions in the phloem was unaffected by drought stress. We conclude that tree defence against L. wingfieldii is decreased during severe drought stress, resulting in changes in the spread and action of the fungus in the sapwood but not in the phloem.     

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.     

Sap flow characteristics of three afforestation species during the wet and dry seasons in a dry-hot valley in Southwest China

Assessing and using tree species (exotic or native) with superior tolerance to environmental stresses (such as drought and high temperature) play an important role in afforestation practices. In the present study, stem sap flow characteristics and responses to ambient meteorological factors of three tree species, Albizzia kalkora (native), Azadirachta indica (exotic), and Acacia auriculaeformis (exotic), in a dry-hot valley (Yuanmou, Yunnan Province, China) were investigated using thermal dissipation probes. The diurnal dynamics of sap flow in three studied species displayed an obvious circadian rhythm during the wet and dry seasons, with the exception of A. indica during the dry season. The sap flow velocity (SFV) in A. kalkora and A. auriculaeformis was significantly positively correlated with photosynthetically active radiation (PAR), air temperature, vapour pressure deficit (VPD) and wind speed, but negatively correlated with atmospheric relative humidity over the two seasons. The cross-correlation analysis also revealed that the SFV of the three species was significantly correlated with PAR and VPD (P < 0.001). Additionally, stem sap flow lagged behind PAR but ahead of VPD, and the diurnal sap flow was more dependent on PAR than on VPD. However, we found that the dominant climatic factor influencing the stem sap flow differed between daytime and nighttime. PAR was more influential than other meteorological factors during the daytime, while VPD or other factors were more influential overnight. When the nighttime refilling ability of the three tree species was compared, our results suggest that A. indica has higher drought resistance and better for afforestation of the studied region.     

A comparison of heat pulse and deuterium tracing techniques for estimating sap flow in Eucalyptus grandis trees

Sap flow rates were measured simultaneously by the heat pulse and deuterium tracing techniques in nine Eucalyptus grandis W. Hill ex Maiden. trees at two sites (1) to compare results from the two techniques and (2) to assess the impact of the assumptions underlying the deuterium tracing method on the calculation of sap flow for a range of tree sizes. The trees ranged in height from 4 to 14 m with leaf areas of 5 to 35 m(2). In all trees, sap flow estimated by the deuterium tracing technique was higher than sap flow estimated by the heat pulse method, with differences of 11 to 43% in eight of the trees and 113% in one tree. The largest difference was attributed to errors in the heat pulse method, as indicated by aberrant relationships between sap flow measured by the heat pulse method and tree size characteristics (i.e., diameter, sap wood area, leaf area) for that tree compared with the other experimental trees. Drilling holes in the trees to allow injection of deuterium had no significant effect on sap flow, even when 32 holes were drilled. Sap flow measured by the heat pulse method was only lower after drilling than before drilling in three trees, and the difference only persisted for about 1 h. Deuterium concentrations of water collected from the tree canopies had not returned to background values 17 days after injection. Twenty-one days after injection, sapwood and heartwood samples taken from trunks near the injection sites contained considerable concentrations of deuterium, indicating that some of the deuterium injected into the trees was still present. An experiment performed on two trees showed that deuterium was stored in the heartwood and sapwood throughout the trees, and its distribution within the trees four days after injection was similar whether it was injected into only the sapwood (where it should mix with sap and be transported from the tree most readily) or into both the sapwood and heartwood, indicating that there was considerable movement of deuterium between the heartwood and sapwood. Deuterium storage was accounted for by an approximate means in the sap flow calculations, and may have resulted in an error of about 10% in sap flow estimated by this method. We conclude that the heat pulse and deuterium tracing techniques can be used simultaneously to increase the number of sap flow measurements obtained from a forest, thereby increasing the precision of forest water use estimates. Their combination would be most effective in stands with a wide range of tree sizes and sap flow rates, where the relative differences in sap flux estimates between the methods is small compared with differences in sap flow between trees.     

Estimating water use by sugar maple trees: considerations when using heat-pulse methods in trees with deep functional sapwood

Accurate estimates of sapwood properties (including radial depth of functional xylem and wood water content) are critical when using the heat pulse velocity (HPV) technique to estimate tree water use. Errors in estimating the volumetric water content (V(h)) of the sapwood, especially in tree species with a large proportion of sapwood, can cause significant errors in the calculations ofsap velocity and sap flow through tree boles. Scaling to the whole-stand level greatly inflates these errors. We determined the effects of season, tree size and radial wood depth on V(h) of wood cores removed from Acer saccharum Marsh. trees throughout 3 years in upstate New York. We also determined the effects of variation in V(h) on sap velocity and sap flow calculations based on HPV data collected from sap flow gauges inserted at four depths. In addition, we compared two modifications of Hatton's weighted average technique, the zero-step and zero-average methods, for determining sap velocity and sap flow at depths beyond those penetrated by the sap flow gauges. Parameter V(h) varied significantly with time of year (DOY), tree size (S), and radial wood depth (RD), and there were significant DOY x S and DOY x RD interactions. Use of a mean whole-tree V(h) value resulted in differences ranging from -6 to +47% for both sap velocity and sap flow for individual sapwood annuli compared with use of the V(h) value determined at the specific depth where a probe was placed. Whole-tree sap flow was 7% higher when calculated on the basis of the individual V(h) value compared with the mean whole-tree V(h) value. Calculated total sap flow for a tree with a DBH of 48.8 cm was 13 and 19% less using the zero-step and the zero-average velocity techniques, respectively, than the value obtained with Hatton's weighted average technique. Smaller differences among the three methods were observed for a tree with a DBH of 24.4 cm. We conclude that, for Acer saccharum: (1) mean V(h) changes significantly during the year and can range from nearly 50% during winter and early spring, to 20% during the growing season;(2) large trees have a significantly greater V(h) than small trees; (3) overall, V(h) decreases and then increases significantly with radial wood depth, suggesting that radial water movement and storage are highly dynamic; and (4) V(h) estimates can vary greatly and influence subsequent water use calculations depending on whether an average or an individual V(h) value for a wood core is used. For large diameter trees in which sapwood comprises a large fraction of total stem cross-sectional area (where sap flow gauges cannot be inserted across the entire cross-sectional area), the zero-average modification of Hatton's weighted average method reduces the potential for large errors in whole-tree and landscape water balance estimates based on the HPV method.