Water relations in silver birch during springtime: How is sap pressurised?

• Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known.
• We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period.
• The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations.
• Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed.

     

Diurnal patterns in Scots pine stem oleoresin pressure in a boreal forest

Coniferous tree stems contain large amounts of oleoresin under positive pressure in the resin ducts. Studies in North‐American pines indicated that the stem oleoresin exudation pressure (OEP) correlates negatively with transpiration rate and soil water content. However, it is not known how the OEP changes affect the emissions of volatile vapours from the trees. We measured the OEP, xylem diameter changes indicating changes in xylem water potential and monoterpene emissions under field conditions in mature Scots pine (Pinus sylvestris L.) trees in southern Finland. Contrary to earlier reports, the diurnal OEP changes were positively correlated with temperature and transpiration rate. OEP was lowest at the top part of the stem, where water potentials were also more negative, and often closely linked to ambient temperature and stem monoterpene emissions. However, occasionally OEP was affected by sudden changes in vapour pressure deficit (VPD), indicating the importance of xylem water potential on OEP as well. We conclude that the oleoresin storage pools in tree stems are in a dynamic relationship with ambient temperature and xylem water potential, and that the canopy monoterpene emission rates may therefore be also regulated by whole tree processes and not only by the conditions prevailing in the upper canopy.     

Wood diameter indicates diurnal and long-term patterns of xylem water potential in Norway spruce

The stem diameter of adult Norway spruce trees was measured to see whether changes in xylem water potential lead to detectable radial deformation of the wood. The dendrometers used in these experiments measured only the dimensional changes of the woody cylinder (sap- and heartwood). Wood diameter was measured close to the ground and just below the living crown. After correction for thermal expansion of dendrometers and wood, diurnal variation of wood diameter ranged between 50 and 180 µm. Psychrometric measurements showed that xylem water potential varied in parallel to wood diameter. Diameter changes were always more pronounced at the higher stem position and exhibited a clear diurnal pattern. During the day, wood diameter decreased with increasing vapor pressure deficit and transpiration rate and with decreasing twig water potential. At night, the wood re-expanded but did not always reach the dimension of the previous day. Pre-dawn wood diameter decreased during periods of soil drought, a process which rapidly stopped and reversed after rain events. On several days, oscillation in wood diameter was observed during the mid-day hours. The oscillation had a period of approximately 50 min and showed a phase shift between different stem heights. All observed patterns of wood shrinkage and expansion were consistent with the hypothesis that xylem water tension leads to an elastic contraction of xylem conduits. The results demonstrate that xylem diameter is more suitable than whole-stem diameter for monitoring changes in xylem water potential.     

鼎湖山针阔叶混交林优势种树干液流特征及其与环境因子的关系

运用Granier热扩散式探针法,于2010年干湿季对鼎湖山自然保护区针阔混交林4种优势树种马尾松、锥栗、木荷和广东润楠的树干液流密度进行连续监测,并同步观测气温、相对湿度和光合有效辐射等环境因子的变化,研究其树干液流特征及其对环境因子的响应.结果表明:在干湿季,4种优势树种的树干液流速率日变化均呈“昼高夜低”的典型单峰曲线,阔叶树锥栗、木荷和广东润楠的平均液流速率和峰值以及日液流量均显著大于针叶树马尾松;马尾松、锥栗、木荷和广东润楠的最大树干液流密度分别为29.48、38.54、51.67、58.32g H2O·m-2·s-1.优势树种树干液流速率的变化与环境因子的昼夜变化存在时滞;液流速率变化与光合有效辐射、水汽压亏缺和气温等环境因子的变化呈显著正相关,其中湿季以光合有效辐射为主导因子,干季以气温为主导因子.     

Diurnal changes in branch diameter as indicator of water status of Hinoki cypress Chamaecyparis obtusa

We investigated use of strain gauges for monitoring the water status of trees by measuring changes in the diameter of the largest spreading branch of a 27-year-old Chamaecyparis obtusa tree. The change in xylem diameter in the branch is more closely related than the change in phloem diameter to the change in leaf water potential. Since the diurnal changes in diameter match the diurnal changes in water balance (sap flow velocity - transpiration), measuring the change in xylem diameter using a strain gauge is useful in evaluating the water status of C. obtusa.     

鼎湖山针阔叶混交林优势种树干液流特征及其与环境因子的关系

运用Granier 热扩散式探针法,于2010年干湿季对鼎湖山自然保护区针阔混交林4种优势树种马尾松、锥栗、木荷和广东润楠的树干液流密度进行连续监测,并同步观测气温、相对湿度和光合有效辐射等环境因子的变化,研究其树干液流特征及其对环境因子的响应.结果表明： 在干湿季,4种优势树种的树干液流速率日变化均呈“昼高夜低”的典型单峰曲线,阔叶树锥栗、木荷和广东润楠的平均液流速率和峰值以及日液流量均显著大于针叶树马尾松;马尾松、锥栗、木荷和广东润楠的最大树干液流密度分别为29.48、38.54、51.67、58.32 g H₂O·m-2·s-1.优势树种树干液流速率的变化与环境因子的昼夜变化存在时滞;液流速率变化与光合有效辐射、水汽压亏缺和气温等环境因子的变化呈显著正相关,其中湿季以光合有效辐射为主导因子,干季以气温为主导因子.     

黄土丘陵区辽东栎和刺槐树干液流时滞效应与蒸腾特征的关联性

运用Granier热扩散探针法,于2016年7-9月对半干旱黄土丘陵区天然次生林树种辽东栎和人工林树种刺槐的树干液流进行连续测定,并同步监测气象因子和土壤含水量,用错位相关法分析液流通量密度与空气水汽压亏缺日变化的时滞长度,研究2个树种不同径级个体在不同土壤水分条件下液流通量密度与蒸腾驱动因子之间的时滞效应.结果表明:辽东栎和刺槐液流通量密度的日变化节律与气象因子显著相关,空气水汽压亏缺峰值的出现较辽东栎树干液流通量密度滞后118.2 min,较刺槐树干液流通量密度滞后39.5 min;而光合有效辐射的峰值通常滞后于辽东栎12.4 min,提前于刺槐68.5 min.液流通量密度和空气水汽压亏缺的时滞长度与树种和土壤含水量显著相关,辽东栎、刺槐在土壤含水量较高时段的时滞长度分别大于土壤含水量较低时段32.2和68.2 min.时滞长度与径级的相关性整体上未达到显著水平,但在土壤含水量较低时段小径级刺槐的时滞长度大于大径级21.4 min,差异达到了显著水平.两树种液流通量密度与空气水汽压亏缺之间的时滞效应反映了对蒸腾驱动因子的敏感性,较好的土壤水分条件有利于液流通量密度提早达到峰值,较低土壤水分会导致树干液流对气象环境因子响应的敏感性降低;刺槐树干液流受土壤水分的影响更显著.     

Crown conductance in dwarf, medium, and tall pitch pines in the Long Island Pine Plains

The New York Pine Plains are a unique ecosystem with normal statured and a dwarfed variety of pitch pines (Pinus rigida Mill.). Growing interspersed with the dwarf pines are trees of intermediate height and features. Several hypotheses have been put forward as to why some of the trees are dwarfed, but none have been substantiated. In this study, we tested whether dwarf or medium trees are hydraulically limited compared to normally growing trees. Granier style sap flux sensors were installed in three to six trees of each tree type and sap flux was measured in early August 2004. Sap flux measurements were scaled to crown stomatal conductance using leaf area to sapwood area ratios for each tree. Contrary to expectation, dwarf and medium stature trees had very low leaf area to sapwood area ratios, but high crown stomatal conductances compared to normal trees. Analyses of leaf area, ring widths, and crown stomatal conductance indicate that differences between normal, and dwarf and medium pines are not due to hydraulic limitation, but that stunted growth may be due to other causes.     

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。     

Laurel forests in Tenerife, Canary Islands

The efficiency of the conductive system in about 40-year-old Laurus azorica trees growing in a laurel forest was evaluated by comparing main stems and leaves (petioles) on the basis of theoretical sap flow values (1) calculated from vessel anatomy (taking vessels as ideal capillaries), (2) derived from measured dye velocity and (3) data taken from direct sap flow measurements. It was found that actual sap flow rate per wood area increases in stems from the pith towards the cambium. The outermost part of the stem is the most important part of the tree for conducting water. Maximum actually measured transpiration (sap flow rate) for the stand was practically identical to the theoretical rate calculated based on petiole anatomy, but it was about 45 times lower than that calculated based on stem anatomy. This illustrates the safety features of stem wood, which due to its high vessel density, is capable of transporting all the water required even when only a small area of its vessels is working. In the petioles, xylem is more efficiently used, but almost all vessels must work in order to supply water to leaves and any disturbance may cause leaf loss.     

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.

     

大连4种城市绿化乔木树种夜间液流活动特征

夜间液流有助于树木物质运输及其体内水分的补充(water recharge), 它不仅对植物的生长发育具有重要的生理生态学意义, 而且对大尺度植物蒸腾耗水的估算可能产生重要影响。2008年6月1日至8月31日, 以热扩散探针(thermal dissipation probe, TDP)技术对大连市劳动公园内的雪松(Cedrus deodara)、大叶榉(Zelkova schneideriana)、丝棉木(Euonymus bungeanus)和水杉(Metasequoia glyptostroboides) 4种乔木树种的不同径阶样木树干边材液流进行了测定, 并结合同步土壤水分与小气候观测结果分析了树木夜间(18:00至次日5:00)液流特征。实验结果表明, 树木普遍存在可感夜间液流, 夜间液流总量占观测期液流总量的比例在样木个体间呈现显著差异, 其变化范围为0.44%-75.96%; 观测期雨天夜间液流波动活跃, 显著高于晴天, 其单日夜间液流总量可持平, 甚至高于日间液流。相关分析表明: 水汽压亏缺(vapor pressure deficit, VPD)和风速的变化与夜间蒸腾显著相关, 它们能够较好地解释液流变化(R² > 0.6); 树木夜间液流主要用于夜间蒸腾和自身水分补充, 夜间液流现象主要发生在前半夜, 后半夜液流平稳且极接近0, 夜间液流量与相应的日间流量(R² = 0.356, p = 0.00)及胸径(R²_Spearman > 0.80)显著相关, 说明植物本身的结构和生理特点也是影响树木夜间液流的重要因子。单株样木夜间液流占全天总蒸腾量的比例低于14.4%, 如不考虑夜间液流的影响, 根据日间液流通过尺度扩展推算的森林生态系统年蒸腾量可能偏低。     

测定位点对计算梨树树干液流的影响

采用热脉冲法,研究了不同测定位点对计算梨树树干液流速率和液流量的影响.结果表明,不同时期内,各方向木质部液质比、木质比径向变幅分别为0.01～0.03和0～0.02,而同一深度木质部液质比和木质比季节变幅分别为0.02～0.09和0.02～0.08. 用同一月份不同深度木质部液质比和木质比参数计算特定深度液流速率差异不显著;而用不同时期测定的同一深度木质部参数计算特定月份相应深度液流速率差异显著或极显著.内层2个、4个测点平均低估液流量是外层相应测定位点的1.5和4.9倍,距形成层0～0.6四个位点的液流量基本可以代表整树耗水量.     

Adjustment Capacity of Maritime Pine Cambial Activity in Drought-Prone Environments

Intra-annual density fluctuations (IADFs) are anatomical features formed in response to changes in the environmental conditions within the growing season. These anatomical features are commonly observed in Mediterranean pines, being more frequent in younger and wider tree rings. However, the process behind IADF formation is still unknown. Weekly monitoring of cambial activity and wood formation would fill this void. Although studies describing cambial activity and wood formation have become frequent, this knowledge is still fragmentary in the Mediterranean region. Here we present data from the monitoring of cambial activity and wood formation in two diameter classes of maritime pine (Pinus pinaster Ait.), over two years, in order to test: (i) whether the differences in stem diameter in an even-aged stand were due to timings and/or rates of xylogenesis; (ii) if IADFs were more common in large trees; and (iii) if their formation is triggered by cambial resumption after the summer drought. Larger trees showed higher rates of cell production and longer growing seasons, due to an earlier start and later end of xylogenesis. When a drier winter occurs, larger trees were more affected, probably limiting xylogenesis in the summer months. In both diameter classes a latewood IADF was formed in 2012 in response to late-September precipitation, confirming that the timing of the precipitation event after the summer drought is crucial in determining the resumption of cambial activity and whether or not an IADF is formed. It was the first time that the formation of a latewood IADF was monitored at a weekly time scale in maritime pine. The capacity of maritime pine to adjust cambial activity to the current environmental conditions represents a valuable strategy under the future climate change conditions.     

Time constant for water transport in loblolly pine trees estimated from time series of evaporative demand and stem sapflow

 The use of stem sap flow data to estimate diurnal whole-tree transpiration and canopy stomatal conductance depends critically upon knowledge of the time lag between transpiration and water flux through the stem. In this study, the time constant for water movement in stems of 12-year-old Pinus taeda L. individuals was estimated from analysis of time series data of stem water flux and canopy transpiration computed from mean daytime canopy conductance, and diurnal vapor pressure deficit and solar radiation measurements. Water uptake through stems was measured using a constant-heat sapflow probe. Canopy transpiration was correlated to stem uptake using a resistance-capacitance equation that incorporates a time constant parameter. A least-squares auto-regression determined the parameters of the resistance-capacitance equation. The time constants for ten loblolly pine trees averaged 48.0 (SE = 2.0) min and the time lag for the diurnal frequency averaged 47.0 (SE = 2.0) min. A direct-cross correlation analysis between canopy transpiration and sap flow time series showed maximum correlation at an approximately 30 min lag. Residuals (model-predicted minus actual stem flow data) increased with increasing soil moisture depletion. While the time constants did not vary significantly within the range of tree sizes studied, hydraulic resistance and capacitance terms were individually dependent on stem cross-sectional area: capacitance increased and resistance decreased with stem volume. This result may indicate an inverse adjustment of resistance and capacitance to maintain a similar time constant over the range of tree sizes studied.     

Tree Height-Related Hydraulic Strategy to Cope with Freeze-Thaw Stress in Six Common Urban Tree Species in North China

Urban trees are sensitive to extreme weather events under climate change. Freeze-thaw induced hydraulic failure could induce urban tree dieback and nullify the services they provide. Plant height is a simple but significant trait for plant ecological strategies. Understanding how urban trees with different heights adapt to freeze-thaw stress is increasingly important under climate change. We investigated the relationship between tree height and stem hydraulic functional traits of six common urban tree species in North China to explore tree height-related hydraulic strategies to cope with freeze-thaw stress. Results showed that tall trees had wider vessels, higher hydraulic conductivity, more winter embolism, but lower vessel and wood densities. Positive relationships were found between tree height and vessel diameter, hydraulic conductivity, and freeze-thaw induced embolism, and negative relationships were found between tree height and vessel and wood densities, which implied that short trees employ more conservative ecological strategies than tall trees. Tall and short tree species were well separated by multiple stem hydraulic functional traits; this is consistent with the fact that tall and short trees occupy different niches and indicates that different hydraulic strategies for freeze-thaw stress exist between them. Tall trees might face more pressure to survive under extreme cold weather caused by climate change in the future. Therefore, more attention should be paid to tall urban tree management in North China to cope with extreme cold weather.     

Comparison of sap flow, cavitation and water status of Quercus petraea and Quercus cerris trees with special reference to computer tomography

Concurrent measurements of sap velocity (heat pulse) and ultrasound acoustic emission were performed on the trunks of mature Turkey oak (Quercus cerris) and sessile oak (Quercus petraea) trees. Plant water status was assessed by measuring leaf water potential, leaf conductance and transpiration. Wood density was estimated non-destructively on the trunk section of the plants by mobile computer tomography, which measures the attenuation of a collimated beam of radiation traversing the trunk in several directions, as the device rotates around the tree. Absorption is proportional to the density of the wood. As wood density is strictly correlated to water content, this non-invasive method allows the water content in the trunk section to be evaluated as well as mapped. Leaf water potential declined each morning until a minimum was reached at midday and recovered in the afternoon, lagging behind changes in transpiration rate. Good correspondence was found between the patterns of sap velocity and cavitation rate. A close correlation was demonstrated between wood density, water content and sap velocity. Sap now was always higher in Turkey oak than in sessile oak. Trunk signatures by computer tomography appeared to differentiate the two oak species, with the Turkey oak stem clearly more hydrated than the sessile oak; water storage reservoirs could play an important role in tree survival during extended periods of low soil water availability and in the relative distribution of tree species, especially in the context of global climate change. Late-wood conducting elements of oak species seem to play a significant role in water transport. The mobile computer tomograph was confirmed as a peerless tool for investigating stem water relations. Diurnal variations in the measured parameters under natural drought conditions and the differences between the two oak species are discussed.     

黄土丘陵区辽东栎树干液流特征对边材面积和土壤水分的响应

运用Granier热扩散探针法对半干旱黄土丘陵区不同胸径辽东栎进行树干液流测定,并对太阳辐射、空气温湿度、降水量、土壤水分等环境因子进行同步观测,分析不同土壤水分条件下不同胸径辽东栎的树干液流变化特征及其对环境因子的响应.结果表明:辽东栎液流日变化特征总体上与太阳辐射和空气水汽压亏缺呈相同趋势,但液流峰值出现时间早于两个气象环境因子的峰值时间.同一树木个体在土壤水分条件较高时期的树干液流通量高于土壤水分较低时期.在相同土壤水分条件下,大径级样本液流通量显著高于小径级样本.采用指数饱和曲线函数对液流通量与太阳辐射和空气水汽压亏缺以及两因子的综合指标进行拟合,效果良好,可以反映液流通量对气象环境因子的响应规律.不同胸径辽东栎在不同土壤水分条件下的拟合曲线特征和拟合参数差异表明,在土壤水分较高时段,液流通量可快速上升至饱和值;在土壤水分较低时段,液流通量上升缓慢.小径级样本对土壤水分变化的反应更加剧烈.单位空气水汽压亏缺的日液流通量值(日液流通量与空气水汽压亏缺的比值)在两种土壤水分条件下的比值与边材面积呈线性相关,且小径级样本的斜率高于大径级样本,说明小径级样本对土壤水分的变化较为敏感,在土壤含水量较低时段,大径级样本较厚的导水组织对土壤水分供应不足起到了缓冲作用.     

Modeling xylem and phloem water flows in trees according to cohesion theory and Münch hypothesis

Water and solute flows in the coupled system of xylem and phloem were modeled together with predictions for xylem and whole stem diameter changes. With the model we could produce water circulation between xylem and phloem as presented by the Münch hypothesis. Viscosity was modeled as an explicit function of solute concentration and this was found to vary the resistance of the phloem sap flow by many orders of magnitude in the possible physiological range of sap concentrations. Also, the sensitivity of the predicted phloem translocation to changes in the boundary conditions and parameters such as sugar loading, transpiration, and hydraulic conductivity were studied. The system was found to be quite sensitive to the sugar-loading rate, as too high sugar concentration, (approximately 7 MPa) would cause phloem translocation to be irreversibly hindered and soon totally blocked due to accumulation of sugar at the top of the phloem and the consequent rise in the viscosity of the phloem sap. Too low sugar loading rate, on the other hand, would not induce a sufficient axial water pressure gradient. The model also revealed the existence of Münch “counter flow”, i.e., xylem water flow in the absence of transpiration resulting from water circulation between the xylem and phloem. Modeled diameter changes of the stem were found to be compatible with actual stem diameter measurements from earlier studies. The diurnal diameter variation of the whole stem was approximately 0.1 mm of which the xylem constituted approximately one-third.     

Traits of Masson Pine Affecting Attack of Pine Wood Nematode

Masson pine characteristics were analyzed in five sample plots in Zhejiang Province, China. Bursaphelenchus xylophilus （Steiner et Buhrer） Nickle （pine wood nematode, PWN） carried by Monochamus alternatus predominately attacked Masson pines in the lower diameter classes. Among the 10 tree characteristics examined, mean crown width, percentage of bole with crown, 5-year cumulative diameter growth, and resin amount showed significant variation between successfully attacked and unattacked trees. The attacked trees had a lower percentage of the bole covered with tree crown, lower crown width, lower radial growth in the last 5 years, and produced less induced resinosis than unattacked trees. Results allowed for effective ranking of the pine forest based on individual tree resistance to PWN. This index of resistance should be considered throughout the development of an ＂Evaluation Criterion and Indicator System＂. The preceding ranking can be used to evaluate the resistance and resiliency of the pine forest ecosystem to PWN＇s invasion, which is similar to Pest Risk Analysis （PRA）.