A requirement for sucrose in xylem sap flow from dormant maple trees

The response of excised stem segments of several tree species to freezing and thawing cycles was studied. All species studied (Thuja occidentalis, Fagus grandifolia, and Betula papyrifera) except maple (Acer spp.) exuded sap while freezing and absorbed on thawing. Maple stems absorbed sap while freezing and exuded sap during the thaw only when sucrose was present in the vessel solution. Increased concentration of sucrose in the vessel sap led to increased exudation. In the absence of sucrose, maple stems absorbed sap on thawing. The presence of sucrose enhanced sap absorption during freezing cycles in maples. In general, large sugars, disaccharides and larger, could substitute for sucrose in the maple exudation response while sugar hexoses could not. The results are discussed in relation to the O'Malley-Milburn model (1983 Can J Bot 61: 3100-3106) of sap flow in maples.     

Maple sap uptake, exudation, and pressure changes correlated with freezing exotherms and thawing endotherms

Sap flow rates and sap pressure changes were measured in dormant sugar maple trees (Acer saccharum Marsh.). In the forest, sap flow rates and pressure changes were measured from tap holes drilled into tree trunks in mature trees and sap flow rates were measured from the base of excised branches. Excised branches were also brought into the laboratory where air temperature could be carefully controlled in a refrigerated box and sap flow rates and sap pressures were measured from the cut base of the branches.

Under both forest and laboratory conditions, sap uptake occurred as the wood temperature declined but much more rapid sap uptake correlated with the onset of the freezing exotherm. When sap pressures were measured under conditions of negligible volume displacement, the sap pressure rapidly fell to −60 to −80 kilopascals at the start of the freezing exotherm. The volume of water uptake and the rate of uptake depended on the rate of freezing. A slow freezing rate correlated with a large volume of water uptake, a fast freezing rate induced a smaller volume of water uptake. The volume of water uptake ranged from 0.02 to 0.055 grams water per gram dry weight of sapwood. The volume of water exuded after thawing was usually less than the volume of uptake so that after several freezing and thawing cycles the sapwood water content increased from 0.7 to 0.8 grams water per gram dry weight.

These results are discussed in terms of a physical model of the mechanism of maple sap uptake and exudation first proposed by P. E. R. O'Malley. The proposed mechanism of sap uptake is by vapor distillation in air filled wood fiber lumina during the freezing of minor branches. Gravity and pressurized air bubbles (compressed during freezing) cause sap flow from the canopy down the tree after the thaw.

     

Scots pine root distribution derived from radial sap flow patterns in stems of large leaning trees

This study characterizes whole tree root system distribution in a non-destructive way based on its functional parameters, particularly the sap flow patterns in stems. This approach particularly considers sap flow variation across stems, both radial and circumferential patterns of flow that are usually used for a better integration of sap flow density at the whole tree level. We focused at: (1) Showing examples of sap flow variation across stems at a defined situation (high midday values at the period of non-limiting water supply; (2) Analyzing radial flow patterns in terms of root distribution; (3) Validating these results at the stand level (mean data of series of individual trees) using results of classical biometric methods used at the same site; and (4) Applying the results for evaluation of root distribution around leaning trees. Sap flow rate was measured by the heat deformation method on a set of 14 trees at an experimental pine forest stand in Brasschaat (Belgium) during the growing season of 2000. Sap flow variation across stems was measured at a total of 700 points. Amounts of water supplied by superficial (horizontally oriented) and sinker (vertically oriented) roots were estimated from sap flow patterns. The vertical distribution of absorbing roots as derived from the analysis of sap flow patterns in stem sapwood was very similar to the distribution determined by the classical biometric analysis of fine roots. Trees leaning to the East had stem radii at the stump level and crown radii enhanced in the leaning direction. Sinker roots showed higher absorption activities in the leaning direction, but superficial roots were more absorbing in the opposite direction. The application of the above-described method allows for a better evaluation of the whole-tree behavior and facilitates the evaluation of tree and stand properties in traditional forest stands, which are not equipped for detailed scientific research. This may also facilitate practical applications in landscape-level studies.     

Measurement of sap flow in plant stems

Transpiration rates for whole plants, individual branches ortillers can be determined by techniques which measure the rateat which sap ascends stems. All of these methods use heat asa tracer for sap movement, but they are fundamentally differentin their operating principles. Two methods commonly employed,the stem heat balance and trunk sector heat balance methods,use the heat balance principle; the stem is heated electricallyand the heat balance is solved for the amount of heat takenup by the moving sap stream, which is then used to calculatethe mass flow of sap in the stem. In the heat-pulse method,rather than using continuous heating, short pulses of heat areapplied and the mass flow of sap is determined from the velocityof the heat pulses moving along the stem. In addition, ratesof sap flow can be determined empirically, using the thermaldissipation technique, from the temperature of sapwood neara continuously-powered heater implanted in the stem. Users mustunderstand the theory underlying each of these methods, so thatthey can select the method most appropriate to their applicationand take precautions against potential sources of error. Whenattempting to estimate transpiration by stands of vegetationfrom measurements of sap flow in individual plants, users mustalso select an appropriate sampling strategy and scaling method. Key words: Sap flow, transpiration, stem heat balance, heat pulse velocity, review     

Gas-exchange and sap flow measurements of Salix viminalis trees in short-rotation forest

Simultaneous field measurements of transpiration and sap flow were performed on short-rotation Salix viminalis trees ranging in diameter from 1.5 to 3.5 cm (2-year-old shoots on 8-year-old stumps). Transpiration was measured using an open-top ventilated chamber enclosing the whole foliage of a tree. Sap flow was measured using a tree-trunk heat balance (THB) technique with a constant temperature difference and variable heat input. Both the instantaneous and daily values of water flux measured by the two absolute techniques agreed well with a difference of up to about 5%. In July, the hourly transpiration reached a maximum of about 0.2 kg m^–2 (leaf area) or 0.45 kg tree^–1, whereas maximum daily integrals reached 4 kg tree^–1. The response of sap flow rate to abrupt flux change when inducing emboli by cutting-off the stem was very rapid: the registered signal dropped by 85% within 10 min for a specimen with a projected leaf area of 2 m². For S. viminalis trees, transpiration was linearly correlated with stem cross-sectional area and with leaf area.     

Validating sap flow measurement in field-grown sunflower and corn

Sap flow measurement has been widely used recently in studyingplant response to the environment, but results have not alwaysbeen satisfactory. The possibility that the non-uniform distributionof the conducting elements in the stem cross-sectional areaintroduces error in the measurements was examined. The heatpulse method, combined with anatomical observations of the stemat the point of sap flow measurement, was applied in corn andsunflower, to study the interaction between anatomical structureand sap flow distribution. The probability of the temperaturesensor being in contact with conducting elements was stronglyaffected by its insertion depth into the stem; the measuredheat pulse velocity for a given transpirational flux was correlatedwith the number of conducting elements in contact with the sensor.Consequently, the calibration coefficient is affected by theposition of the thermocouple junction in relation to the conductingelements. This dependence of apparent heat pulse velocity ondensity of conducting elements suggests that thermal equilibrationin species with large stems may not be achieved within the limitedtime of heat dissipation. The relationship between sap flowand leaf area of single plants was used to test the consistencyand the accuracy of the sap flow measurements. The ratio ofmeasured to potential transpiration was used to validate theuse under field conditions, of the calibration coefficient determinedin potted plants. Key words: Heat pulse, transpiration, LAI, potential transpiration     

干旱区枸杞树干液流变化特征及其影响因素

树干液流作为植物蒸散作用的水分来源,是植物水分消耗的直观量化监测指标,利用包裹式树干液流监测技术获取干旱区枸杞全生育期树干液流实时数据,分析了不同时间尺度树干液流变化特征及各气象要素对树干液流的影响,为明晰枸杞耗水规律及其影响因素提供了重要的佐证。结果表明:枸杞的树干液流量昼夜差异较大,白天液流量是夜间的10倍左右;晴天液流速率、日累积量及变化幅度均大于阴雨天气,晴天液流速率变化曲线且呈宽峰型,在06:30左右启动较阴天提前30min;夏季树干液流启动时间为6:00比秋季提前1h左右,夏季的峰值123g/h。盛果期液流速率最大10.32g/h,营养生长期最小1.35 g/h;6—8月旺盛生长季,平均日耗水1388.3g/d,5—11月全生育期日均耗水1102.7g/d;树干液流速率与太阳辐射、空气温度均呈极显著正相关关系,与相对湿度呈负相关关系;枸杞树干液流(F)与太阳辐射(S)、温度(T)、相对湿度(H)及饱和水汽压(VPD)符合方程F=41.5+0.167S-0.563H+1.36T-9.67VPD(R~2=0.6547)。     

荒漠防护林典型树种液流特征及其对环境因子的响应

利用基于热补偿理论的SF300分体液流仪对干旱荒漠区人工防护林典型树种(俄罗斯杨、胡杨、榆树、沙枣)树干液流全天候监测,自动气象站同步记录相关环境因子变化。研究表明:①4种防护林树种茎干液流日变化除沙枣树外均存在明显昼夜节律,液流速度在同属种间差异较小,在不同属种间差异显著,俄罗斯杨的日平均液流速度可以达到沙枣的13.8倍,耗水量排序为俄罗斯杨胡杨榆树沙枣树。②水分充足条件下,增加实验地灌溉量使4种树木蒸腾受到抑制,液流流速降低,水分利用效率降低。③液流流速因所处树干径向位点不同而存在差异,俄罗斯杨、榆树、沙枣液流速度表现出由形成层到髓心的递减趋势,胡杨树干径向位点液流没有表现一定规律。④树干液流流速与环境因子进行相关分析,通过逐步回归分析建立了4个典型树种茎干液流速度与环境因子关系估算模型,分析认为4种树木的环境敏感性排序为俄罗斯杨榆树胡杨沙枣。     

胡杨液流对地下水埋深变化的响应

以塔里木河流域荒漠河岸林主要建群种胡杨(Populus euphratica)为研究对象, 结合中下游不同断面地下水埋深和胡杨液流变化的监测数据, 分析了胡杨茎干液流与地下水埋深变化的关系, 探讨荒漠环境下天然胡杨生长的合理生态水位。研究表明, 胡杨液流通量密度随地下水埋深即干旱胁迫程度的加大而减小, 两者呈极显著负相关, 相关系数达-0.887; 胡杨液流通量在地下水埋深位于4.5-5 m时出现异常变化, 表明此时胡杨的正常生长受到胁迫, 胡杨通过自身调节降低蒸腾耗水以适应环境; 土壤盐分不是影响塔里木河中下游各断面胡杨液流变化的主要因子; 对植物样地调查结果分析显示, 胡杨盖度、密度和频度均在地下水埋深在4-6 m梯度下开始表现为降低趋势。综合分析认为维系塔里木河中下游天然胡杨正常生长的生态水位为地下水埋深4.5 m以内。     

An Improved Thermoelectric Probe for Measurements of Apparent Sap Flow Velocity in Intact Plant Stems

Gray, H. R-, Erickson, P. I. and Stone, J. F. 1985. An improvedthermoelectric probe for measurement of apparent sap flow velocityin intact plant stems.—J. exp. Bot. 36: 1320–1324. An improved thermoelectric technique for non-intrusive, non-destructive,and direct sensing of the apparent velocity of sap flow in intactplant stems is presented. The thermoelectric probe was madeof extruded Teflon (DuPont) rod. One of two heating elements,positioned 0.5 cm and 1.0 cm below the temperature sensor, wasused to generate the heat pulse. Heating element selection wasbased on daily environmental conditions. A thermistor touchingthe stem sensed the heat pulse. A second thermistor mountedin dose proximity was used to cancel ambient temperature fluctuations. The improved thermoelectric probe was tested using irrigatedpeanuts (Arachis hypogeae L.). After canopy closure was completein both treatments, apparent sap flow velocity(V_a) was measuredon 17 dat 1200 s intervals between 0830 h and 1630 h apparentsolar time. Successful V_a, measurements were accomplished on94% of the attempts. In order to verify performance on otherspecies with different types of stems, non-replicated V_a, determinationswere also made on cotton (Gossypiwn spp.), prickly sida (Sidasptnosa L.), goosegrass (Eleusine indica (L.) Gaertn.), Palmerpigweed (Amaranthus Palmerii S. Wats.), and cottonwood (Populusdeltoides Marsh.). Success of the probe was due in part to the design of the probewhich included remotely selectable spacing between heater andthermistor and adequate insulation and shielding of the probeand sensed portion of the stem. In addition, acclimatizationof the probe to the operating environment was essential. Key words: Sap velocity, instrumentation     

Heat balance sap flow gauge for small diameter stems

Applying heat balance sap flow gauges to plant stems <10 mm in diameter has been difficult because a miniature design is needed that can be attached to a range of stem geometries. This report presents a modified gauge design for use on small plant stems of irregular geometry and shows results from Glycine max with stem diameters of 3–4 mm. The gauge was evaluated on container-grown plants by comparing gauge measurements of flow to gravimetric estimates of transpiration. Experiments were conducted in the laboratory and greenhouse, using artificial and natural lighting, respectively. Laboratory comparisons of gauge versus gravimetric water loss measurements indicated that the instrument was accurate to within ±5% when soil water was not limiting. Similar results were obtained from greenhouse tests except when soil water availability was low and cumulative gauge estimates became 30–45% less than gravimetric measurements. Differences may have reflected reduced plant water uptake or errors in sap flow estimates associated with low flow rates. Gauge accuracy was not improved by including the rate change in heat storage (S) in the flow calculations because S was always less then 3% of the total heat balance. Relationships between system temperature and sap flow rate suggested a diagnostic test for determining optimum power input. A time constant of 15 s indicated potential application in many agronomic and physiological studies.     

Changes in stem water content influence sap flux density measurements with thermal dissipation probes

Key message

Stem WC may decline during the day. Zero-flow dT _m increases when WC decreases. Use of nighttime dT _m in the calculation of sap flux density during the day might introduce errors.

Abstract

There is increasing evidence of diel variation in water content of stems of living trees as a result of changes in internal water reserves. The interplay between dynamic water storage and sap flow is of current interest, but the accuracy of measurement of both variables has come into question. Fluctuations in stem water content may induce inaccuracy in thermal-based measurements of sap flux density because wood thermal properties are dependent on water content. The most widely used thermal method for measuring sap flux density is the thermal dissipation probe (TDP) with continuous heating, which measures the influence of moving sap on the temperature difference between a heated needle and a reference needle vertically separated in the flow stream. The objective of our study was to investigate how diel fluctuations in water content could influence TDP measurements of sap flux density. We analysed the influence of water content on the zero-flow maximum temperature difference, dT _m, which is used as the reference for calculating sap flux density, and present results of a dehydration experiment on cut branch segments of American sycamore (Platanus occidentalis L.). We demonstrate both theoretically and experimentally that dT _m increases when stem water content declines. Because dT _m is measured at night when water content is high, this phenomenon could result in underestimations of sap flux density during the day when water content is lower. We conclude that diel dynamics in water content should be considered when TDP is used to measure sap flow.     

Sapflow+: a four-needle heat-pulse sap flow sensor enabling nonempirical sap flux density and water content measurements

? To our knowledge, to date, no nonempirical method exists to measure reverse, low or high sap flux density. Moreover, existing sap flow methods require destructive wood core measurements to determine sapwood water content, necessary to convert heat velocity to sap flux density, not only damaging the tree, but also neglecting seasonal variability in sapwood water content. ? Here, we present a nonempirical heat-pulse-based method and coupled sensor which measure temperature changes around a linear heater in both axial and tangential directions after application of a heat pulse. By fitting the correct heat conduction-convection equation to the measured temperature profiles, the heat velocity and water content of the sapwood can be determined. ? An identifiability analysis and validation tests on artificial and real stem segments of European beech (Fagus sylvatica L.) confirm the applicability of the method, leading to accurate determinations of heat velocity, water content and hence sap flux density. ? The proposed method enables sap flux density measurements to be made across the entire natural occurring sap flux density range of woody plants. Moreover, the water content during low flows can be determined accurately, enabling a correct conversion from heat velocity to sap flux density without destructive core measurements.     

Effects of xylem sap flow on carbon dioxide efflux from stems of birch (Betula pendula Roth)

The effect of xylem sap flow in stems of mature Betula pendula Roth on radial CO₂ efflux was studied from April to October 2001. Temperature-controlled respiration cuvettes allowed measurements of CO₂ efflux without interference from temperature gradients between stem surface and sapwood. Variations of sap flow in different stem sectors, and in a given sector at different heights were analysed. Daytime reduction of CO₂ efflux caused by sap flow was expressed as the difference between gross and apparent CO₂ release. Gross CO₂ release was calculated from Arrhenius-equations derived from night-time data records of the same day, which were free from interference by sap flow. In mid-July, daytime reductions of CO₂ efflux reached 1.8–3.9 μmol CO₂ m⁻² g⁻¹ xylem sap transpired. Assuming tree-specific maximum transpiration rates of 30 kg H₂O d⁻¹ this is up to 40% of gross CO₂ release. In relation to photosynthetic CO₂ fixation the endogenous supply of dissolved CO₂ to the leaves acccounted for 0.5–3.7%. This study indicates a negative correlation between sap flow velocity and radial CO₂ efflux from B. pendula stems. Periods of unbalanced CO₂ partial pressures between aqueous and gaseous pathways during increase and decrease of sap flow seem to affect gaseous CO₂ release through lenticels. It is concluded that CO₂ efflux rates are not simply equivalent to respiration rates because of the interference of aqueous CO₂ transport by xylem sap flow in the wood-body of trees.     

马占相思树干液流与光合有效辐射和水汽压亏缺间的时滞效应

应用Granier热消散探针测定华南丘陵马占相思的树干液流,将液流与对应的光合有效辐射和水汽压亏缺数据列分别进行逐行错位分析和时间序列分析,探讨树干液流与蒸腾驱动因子之间的时滞效应,并对结果进行互相验证.结果表明:马占相思树木蒸腾主要驱动因子是光合有效辐射和水汽压亏缺,树干液流的变化更多地依赖光合有效辐射的变化,而且干季的依赖性比湿季更强;无论是干季还是湿季,树干液流都滞后于光合有效辐射,提前于水汽压亏缺;时滞效应季节差异显著;不同径级马占相思的时滞效应差异不显著;树高、胸径、冠幅并不能解释树干液流与光合有效辐射、水汽压亏缺之间的时滞效应;干季树干液流与水汽压亏缺之间的时滞效应与夜间水分补充量显著相关,湿季则相反.     

Xylem sap flow as a major pathway for oxygen supply to the sapwood of birch (Betula pubescens Ehr.)

The role of xylem sap flow as an aqueous pathway for oxygen supply to the wood parenchyma of Betula pubescens saplings was investigated. Using micro‐optode sensors the oxygen status of the sapwood was quantified in relation to mass flow of xylem sap. Sap flow was gradually reduced by an increasing oxygen depletion in the root space. The effect of sap flow on radial O₂ transport between stem and atmosphere was assessed by a stoichiometrical approach between respiratory CO₂ production and O₂ consumption. Restriction of sap flow set in 36.5 h after the onset of O₂ depletion, and was complete after 71 h. Interruption of sap flow drastically increased the O₂ deficit in the sapwood to 70%. Sap flow contributed about 60% to the total oxygen supply to the sapwood. Diurnal O₂ flow rates varied between 3 and 6.3 nmol O₂ m^?2 leaf area (LA) s^?1 during night‐ and daytime, respectively. Maximum O₂ flow rates of 20 nmol O₂ m^?2 LA s^?1 were reached at highest sap flow rates of 5.7 mmol H₂O m^?2 LA s^?1. Sap flow not only affected the oxygen status of the sapwood but also had an effect on radial O₂ transport between stem and atmosphere.     

Evaluation of the heat pulse velocity technique for measurement of sap flow in rainforest and eucalypt forest species of south-eastern Australia

Sap flow in the stems of two cut saplings each of Eucalyptus maculata (a canopy eucalypt forest tree), Doryphora sassafras and Ceratopetalum apetalum (both canopy rainforest trees of south-eastern coastal Australia) was measured by the heat pulse velocity technique and compared with water uptake from a potometer. Scanning electron micrographs of wounding caused by implantation of temperature sensor and heater probes into the sapwood showed that wounding was similar in rainforest and eucalypt species and was elliptical in shape. A circular wound has been implicitly assumed in previous studies. Accurate measurements of sapling water use were obtained using the smaller transverse wound dimension rather than the larger longitudinal dimension because maximum disruption of sap flow through the xylem vessels occurred in the transverse plane. Accurate measurements of sap flux were obtained above a minimum threshold sap velocity. These velocities were 15·7,10·9 and 9·4 cm h^?1 for E. maculata, C. apetalum and D. sassafras, respectively. Below the threshold sap velocity, however, sap flow could not be accurately calculated from measurements of heat pulse velocity. The minimum threshold sap velocity appeared to be determined by probe construction and xylem anatomy. Despite the elliptical wounding and inaccurate measurement of sap flow below the threshold sap velocity, total sap flow over the experimental period for two saplings of each species was within 7% of water use measured by the potometer.     

Phloem exudation from castor bean: Induction by massage

Summary Exudation can be induced or greatly enhanced in Ricinus by massaging the stem periodically some time before tapping the sap by making an incision. Nevertheless temporary massage during exudation stops flow completely indicating that flow is prevented normally by a pressure sensitive sealing mechanism which becomes desensitised by regular massage.     

Daily sap flow rate as an indicator of drought avoidance mechanisms in five Mediterranean perennial species in semi-arid southeastern Spain

Daily sap flow rate was determined in five Mediterranean species (Pinus halepensis, Quercus coccifera, Pistacia lentiscus, Erica multiflora, and Stipa tenacissima) under two slope aspects (north- and south-facing) in a semi-arid area (Alicante, SE Spain). Sap flow velocity was measured in January, May, August and October of two consecutive years (1998 and 1999) using the stem heat balance (SHB) method. Our results have demonstrated the effects of global radiation (R _g), vapour pressure deficit (VPD) on the sap flow velocity per unit of leaf area. Mean daily sap flow rates (Q _md) showed values between 0.001 and 0.202 g H₂O cm⁻² leaf area day⁻¹. Q _md values were higher on the south-facing slope than on the north-facing slope. In most species, the Q _md was higher in 1998 than in 1999 due to the higher soil water content, temperature and VPD in 1998. In all five species, a decrease in predawn leaf water potential was accompanied by a decrease in mean daily sap flow rates; nevertheless, the responses of the five species to water deficit conditions were different. In this context, we have linked the drought avoidance mechanisms of the different species through the combined use of daily sap flow rate and predawn leaf water potential under different water deficit conditions. We conclude that Pinus halepensis, Pistacia lentiscus and Erica multiflora show water-savers mechanisms to cope with drought, while Quercus coccifera and Stipa tenacissima show water-spenders mechanisms.     

Effect of soil temperature on stem sap flow,shoot gas exchange and water potential of Picea engelmannii (Parry) during snowmelt

Summary The effect of cold soils on stem sap flow, shoot gas exchange and water potential of Picea engelmannii (Parry) was investigated during the snowmelt period in the Medicine Bow Mountains, Wyoming, USA. Shoot net photosynthetic rates were higher in young trees (1.5–1.8 m in height) growing in cold soils (<3.5° C) associated with snowpack, than trees in warm soils until about 1500 h. Higher shoot photosynthetic rates of trees in cold soils continued after snow was removed and could not be completely explained by higher visible irradiance over highly reflective snow. Following soil warming higher photosynthetic rates were evident in these trees for five days. High nutrient availability associated with snowmelt may improve shoot nutrient status leading to higher gas-exchange rates during snowmelt. Shoot conductance to water vapor was higher in trees in cold soil until midday, when declining shoot conductance led to lower intercellular CO₂ concentrations. Midday through afternoon shoot water potentials of trees in cold soils were similar or higher than those of trees in warm soils and the lower afternoon shoot conductances in cold soils were not the result of lower bulk shoot water potentials. Decline in net photosynthesis of trees in cold soils at 1500 h paralleled increases in intercellular CO₂ concentrations, implying a nonstomatal limitation of photosynthesis. This scenario occurred consistently in mid-afternoon following higher morning and midday photosynthesis in cold soils, suggesting a carbohydrate feedback inhibition of photosynthesis. Diurnal patterns in stem sap flow of all trees (cold and warm soils) reflected patterns of shoot conductance, although changes in stem sap flow lagged 1–3 h behind shoot conductance apparently due to stem water storage. Total daily stem sap flow was similar in trees in cold and warm soils, although diel patterns differed. The morning surge and night-time drop in sap flow commenced 1–2 h earlier in trees in cold soils. Overnight stem sap flow was lower in trees in cold soils, possibly due to higher resistance to root water uptake in cold soils, which may explain lower predawn shoot water potentials. However, midday shoot water potentials of trees in cold soils equalled or exceeded those of trees in warm soils. Higher resistance to root water uptake in P. engelmannii in cold soils was apparently overshadowed by transpirational forces and significant shoot water deficits did not develop.