Temporal and spatial pattern of embolism induced by pressure collar techniques in twigs of Picea abies

The pressure collar technique enables the induction of embolism in plant xylem. This artificial cavitation is based on air seeding processes which occur when specific pressure gradients between the air and water phase of the xylem are exceeded. Standard pressure collars and a new point injection technique, which builds up a local potential gradient, were used to study the time and spatial pattern of this process. On twigs of Norway spruce (Picea abies), the cross-sectional and axial pattern, and the time-course of embolism formation were analysed via conductivity and ultrasonic measurements as well as staining experiments. Furthermore, the release of air from the twig surface was studied by immersing twig sections in water. In cross-sections, embolized areas induced by the point injection technique were smaller compared with the standard collar and restricted to a circle sector. Embolism propagated from the pressure collar towards the nearest distal and proximal nodes but not further. A release of air was also observed predominantly at the internode attached to the pressure collar. Embolism rates increased within minutes and reached approximately 80% loss of conductivity after 10 min treatment with the standard collar. The size of air entry points and embolism rates correlated significantly. Embolism formation in wood therefore depends not only on vulnerability thresholds but also on the extent of air-water interfaces within the xylem and on the time of exposure to pressure gradients. These aspects and the propagation of pressure within samples are crucial for pressure collar experiments. In addition, wood architecture influences the extent and pattern of embolism caused by air seeding processes.     

Uptake of water and solutes through twigs of Picea abies (L.) Karst

Summary Uptake of water and magnesium chloride solution was investigated through the outer surface of twigs of Picea abies (L.) Karst. Water uptake was determined by using pressure/volume (P/V) curves of the twigs as a basis for calculation to avoid problems of superficial extraneous water. When water was sprayed on bark and needles of 3- to 7-year-old twigs at a xylem water potential of -1.00 MPa, they absorbed as much as 80 mm³ water in 200 min/g twig dry weight as the twig water potential recovered to -0.15 MPa. With fluorescent dyes, pathways for absorption of water and solutes through the twig bark were found, particularly through the radially orientated ray tissue. In addition to uptake by mass flow, magnesium could also diffuse along a concentration gradient from the twig surface into the xylem. In the field, the magnitude of these uptake processes would depend on the concentration of elements deposited by atmospheric precipitation, the concentration gradient between the plant surface and the xylem sap, the xylem water potential and the intensity and duration of each precipitation event.     

Water storage in the wood and xylem cavitation in 1-year-old twigs of Populus deltoides Bartr.

The possible role of water expelled from cavitated xylem conduits in the rehydration of water-stressed leaves has been studied in one-year-old twigs of populus deltoides Bartr. Twigs were dehydrated in air. At desired values of leaf water potential (Ψ_l) (between near full turgor and -1.62 MPa), twigs were placed in black plastic bags for 1–2h. Leaf water content was measured every 3–5 min before bagging and every 10 min in the dark. Hydraulic conductivity and xylem cavitation were measured both in the open and in the dark. Cavitation was monitored as ultrasound acoustic emissions (AE). A critical Ψ_l value of -0.96 MPa was found, at which AE increased significantly while the leaf water deficit decreased by gain of water. Since the twigs were no longer attached to roots, it was concluded that water expelled from cavitated xylem conduits was transported to the leaves, thus contributing to their rehydration. Xylem cavitation is discussed in terms of a ‘leaf water deficit buffer mechanism’, under not very severe water stress conditions.     

A New Laboratory Method Used for Investigating the Uptake, Translocation and Metabolism of Bark Banded Morphactin by Trees

Cut twigs of Monterey pine(Pinus radiata D.Don) were banded with a ¹⁴C labelled morphactin formulation and the lower half of the twig placed in a Scholander type pressure bomb. Water was pumped through the twig from a reservoir inside the bomb and exudate collected from the protruding end of the twig. Using this method it was possible to determine that the slowest step in the translocation of morphactin was the passage across the outer layer of bark. Subsequent passage through the twig was not affected by girdling the bark suggesting that morphactin was translocated upwards via the xylem. The ¹⁴C compound in the exudate from live twigs differed from morphactin and was tentatively identified as the carboxylic acid derivative of the morphactin ester. The cut twig system facilitated studies on the uptake, metabolism and translocation of morphactin and should prove useful for studying the uptake and fate of other chemicals considered for bark application to trees.     

Ultrasound acoustic emissions from dehydrating leaves of deciduous and evergreen trees

While drying, detached leaves produced ultrasound acoustic emissions (UAE) comparable to emissions from stem and twig wood. Experiments on Ilex aquifolium L. showed that the main source of these signals was cavitation in the veins, to which conduits and fibres probably both contributed. Regions of the leaf blade with abundant mesophyll and only small veins emitted few signals. More signals were counted on the adaxial side of the midrib than on the abaxial one and on the proximal third than on the distal one, in accordance with the anatomical structure. Sound attenuation was pronounced. Eight species were compared with respect to cavitation behaviour, field water relations and pressure–volume curves, and the data showed differences in cumulative number of events and resistance of leaves to cavitation. Data were generally in good agreement with anatomical structure and habitat preferences. The number of signals per conduit counted on cross-sections was in some leaves much higher than unity, which suggests short xylem elements or an acoustic activity of cells other than conduits. There was no correlation between cavitation threshold or cumulative number of signals and the degree of sclerophylly; unexpectedly, there was a correlation between the cumulative number of signals at a water potential of -1.3 MPa and the bulk modulus of elasticity.     

武夷山不同海拔黄山松枝叶大小关系

对武夷山自然保护区不同海拔黄山松叶片面积、叶片数量、小枝长度及小枝直径等性状进行测定,分析不同海拔黄山松枝-叶大小间的权衡关系.结果表明: 随海拔升高,黄山松小枝的叶片数量、小枝长度、小枝直径、出叶强度及茎截面积逐渐增大,单叶面积呈逐渐减小趋势;不同海拔黄山松小枝出叶强度与单叶面积均呈显著负相关,不同海拔黄山松小枝茎截面积与总叶面积呈显著正相关;不同海拔黄山松小枝长度、小枝直径与出叶强度呈显著负相关,与单叶面积、叶片数量及总叶面积均呈显著正相关.为提高竞争优势或是资源利用效率,低海拔黄山松倾向于在短枝上着生量少但单叶面积大的针叶,而高海拔黄山松趋向于在长枝上着生量大但单叶面积小的针叶,这体现出不同海拔梯度黄山松小枝的资源利用策略及枝叶间生物量分配的权衡机制.     

Vulnerability to cavitation in Olea europaea current‐year shoots: further evidence of an open‐vessel artifact associated with centrifuge and air‐injection techniques

Different methods have been devised to analyze vulnerability to cavitation of plants. Although a good agreement between them is usually found, some discrepancies have been reported when measuring samples from long‐vesseled species. The aim of this study was to evaluate possible artifacts derived from different methods and sample sizes. Current‐year shoot segments of mature olive trees (Olea europaea), a long‐vesseled species, were used to generate vulnerability curves (VCs) by bench dehydration, pressure collar and both static‐ and flow‐centrifuge methods. For the latter, two different rotors were used to test possible effects of the rotor design on the curves. Indeed, high‐resolution computed tomography (HRCT) images were used to evaluate the functional status of xylem at different water potentials. Measurements of native embolism were used to validate the methods used. The pressure collar and the two centrifugal methods showed greater vulnerability to cavitation than the dehydration method. The shift in vulnerability thresholds in centrifuge methods was more pronounced in shorter samples, supporting the open‐vessel artifact hypothesis as a higher proportion of vessels were open in short samples. The two different rotor designs used for the flow‐centrifuge method revealed similar vulnerability to cavitation. Only the bench dehydration or HRCT methods produced VCs that agreed with native levels of embolism and water potential values measured in the field.     

生长环境与株龄对凤丹当年生果枝的生物量分配的影响

生物量分配研究是了解作物产量形成机制的基础。凤丹是以杨山牡丹（Paeonia ostii）为原种形成的栽培类群,也是新型木本油料作物油用牡丹的主栽品种。该文通过比较凤丹主要产区6龄种群、以及安徽铜陵和上海地区的4、6、8龄种群的当年生果枝大小与生物量分配,探讨环境和株龄对小枝生长与繁殖的影响。结果表明,凤丹植株的果枝数量随株龄增大而上升,不同株龄植株的单个果枝大小和果枝内生物量分配没有明显差异,果枝水平上没有株龄效应,但整株水平上株龄效应明显;果枝大小及生物量各指标之间存在明显相关性,但相关性在种群间变化较大;不同种群的果枝大小及生物量分配差异明显,果枝生物量分配明显随纬度、降水和光辐射强度变化而变化;异速生长方程模拟显示,大部分种群的果枝繁殖与营养生物量或总生物量之间为等速生长关系（斜率=1）,但种群间截距变化较大。这些结果表明栽培环境对凤丹果枝的生长与繁殖有显著效应。     

The leaf size – twig size spectrum and its relationship to other important spectra of variation among species

There is a spectrum from species with narrow, frequently branched twigs carrying small leaves and other appendages, to species with thick twigs carrying large leaves and appendages. Here we investigate the allometry of this spectrum and its relationship to two other important spectra of ecological variation between species, the seed mass-seed output spectrum and the specific leaf area-leaf lifespan spectrum. Our main dataset covered 33 woody dicotyledonous species in sclerophyll fire-prone vegetation on low nutrient soil at 1,200 mm annual rainfall near Sydney, Australia. These were phylogenetically selected to contribute 32 evolutionary divergences. Two smaller datasets, from 390 mm annual rainfall, were also examined to assess generality of cross-species patterns. There was two to three orders of magnitude variation in twig cross-sectional area, individual leaf size and total leaf area supported on a twig across the study species. As expected, species with thicker twigs had larger leaves and branched less often than species with thin twigs. Total leaf area supported on a twig was mainly driven by leaf size rather than by the number of leaves. Total leaf area was strongly correlated with twig cross-section area, both across present-day species and across evolutionary divergences. The common log-log slope of 1.45 was significantly steeper than 1. Thus on average, species with tenfold larger leaves supported about threefold more leaf area per twig cross-section, which must have considerable implications for other aspects of water relations. Species at the low rainfall site on loamy sand supported about half as much leaf area, at a given twig cross-section, as species at the low rainfall site on light clay, or at the high rainfall site. Within sites, leaf and twig size were positively correlated with seed mass, and negatively correlated with specific leaf area. Identifying and understanding leading spectra of ecological variation among species is an important challenge for plant ecology. The seed mass-seed output and specific leaf area-leaf lifespan spectra are each underpinned by a single, comprehensible trade-off and their consequences are fairly well understood. The leaf-size-twig-size spectrum has obvious consequences for the texture of canopies, but we are only just beginning to understand the costs and benefits of large versus small leaf and twig size.     

Systemic Migration and Establishment of Xanthomonas campestris pv. pruni in Plum Leaves and Twigs

The systemic migration of Xanthomonas campestris pv. pruni (Xcp) through vascular bundles of leaves and twigs of plum was investigated. A rifampicin-resistant strain of Xcp was inoculated into leaves located midway from the tip of new green twigs of Golden King plum trees in a glasshouse. High numbers of the pathogen were recovered 4 and 8 weeks after inoculation from sections of uninoculated and symptomless veins, petioles, and twig tissue. Symptoms of bacterial spot developedwithin 8 weeks on main and secondary veins of uninoculated leaves located as far as 13 cm from the inoculated leaf on the same twig. Weekly isolation indicated the constant presence of Xcp in apparently unaffected shields of twig tissue obtained from a naturally infected Golden King orchard. Xcp apparently enters plum twigs through veins of infected leaves and migrates systemically through twigs to leaves.     

浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.     

Biogeographic Patterns of Structural Traits and C:N:P Stoichiometry of Tree Twigs in China’s Forests

There have been a number of studies on biogeographic patterns of plant leaf functional traits; however, the variations in traits of other plant organs such as twigs are rarely investigated. In this study, we sampled current-year twigs of 335 tree species from 12 forest sites across a latitudinal span of 32 degrees in China, and measured twig specific density (TSD), twig dry matter content (TDMC), and carbon (C), nitrogen (N) and phosphorous (P) contents, to explore the latitudinal and environmental patterns of these twig traits. The overall mean of TSD and TDMC was 0.37 g cm⁻³ and 41%, respectively; mean twig C, N and P was 472 mg g⁻¹, 9.8 mg g⁻¹ and 1.15 mg g⁻¹, respectively, and mean N:P mass ratio was 10.6. TSD was positively correlated with TDMC which was positively associated with twig C but negatively with twig N and P. There were no significant differences in TSD between conifer, deciduous-broadleaf and evergreen-broadleaf plants, but evergreen-broadleaf plants had the lowest and conifers the highest TDMC. Conifer twigs were lowest in C, N, P and N:P, whereas deciduous-plant twigs were highest in N and P and evergreen-plant twigs were highest in C and N:P. As latitude increased or temperature/precipitation dropped, TDMC and P increased, but N:P ratio decreased. Our results also showed that the patterns of twig P and N:P stoichiometry were consistent with those reported for leaves, but no significant trends in twig N were observed along the gradient of latitude, climate and soils. This study provides the first large-scale patterns of the twig traits and will improve our understanding of the biogeochemistry of carbon and other key nutrients in forest ecosystems.     

Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park,Zhejiang Province,China

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.     

Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient: allometric scaling analysis

We studied the effects of twig size and altitude on biomass allocation within plant twigs (terminal branches of current-year shoots), to determine whether species with large twigs/leaves or living at low altitude allocate a higher proportion of biomass to laminas than their counterparts with small twigs/leaves or living at high altitude. Stem mass, lamina mass and area, and petiole mass were measured for terminal branches of current-year shoots in 80 subtropical evergreen broad-leaved species belonging to 38 genera from 24 families along an altitudinal gradient of Mt. Emei, Southwest China. The scaling relationships between the biomass allocations of within-twig components were determined using model type II regression method. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to either leaves or laminas was independent of twig mass. Petiole mass disproportionally increased with both lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. These cross-species correlations were consistent with those among evolutionary divergences. In addition, species at low altitude tended to have a greater leaf and lamina mass but a smaller stem mass at a given twig mass than at middle and high altitudes. This is possibly due to the high requirement in physical support and the low efficiency of eco-physiological transport for the species living at high altitude. In general, within-twig biomass allocation pattern was not significantly affected by twig size but was greatly modulated by altitude.     

浙江天童木本植物小枝的“大小-数量”权衡

枝条大小和数量关系反映了植物适应环境胁迫的构型和生物量分配策略。该研究以浙江天童木本植物为对象, 通过对小枝大小(横截面积)与数量(稠密度)关系的研究发现: 1)小枝稠密度与枝截面积显著负相关(斜率为-1.32, CI = -1.48- -1.17; p < 0.05); 2)在相同曝光度, 在II级曝光环境(植株40%-80%暴露在直射光中)中常绿植物比落叶植物单位小枝截面积的小枝稠密度高, 而在I级(植株<40%暴露在直射光中)和III级(>80%暴露在直射光中)曝光环境中, 小枝稠密度在两种生活型间无显著差异; 3)在不同曝光度下, 常绿植物单位枝条在I和II级比III级曝光水平具有更高的小枝稠密度; 但落叶植物单位枝条的小枝稠密度在3个曝光水平相同; 4)相同枝条大小下, 4 m以下灌木比4 m以上的亚乔木和乔木具有更高的小枝稠密度。总之, 天童地区木本植物的小枝“大小-数量”关系符合Corner法则(描述枝叶“大小-数量”关系的法则), 且在不同生活型间存在差异, 常绿植物相对于落叶植物, 灌木相对于乔木具有较高的小枝稠密度, 从而有利于它们适应光资源的限制。     

发育阶段和海拔对岷江源区陇蜀杜鹃小枝功能性状及生物量分配的影响

本研究分析了岷江源区卡卡山典型高山灌丛植物陇蜀杜鹃不同发育阶段(花芽期和开花期)及不同海拔(3600 m低海拔和3800 m高海拔)下功能小枝和叶片性状的差异,以及空间异质性对花芽期和开花期小枝功能性状相关性及其权衡作用的影响。结果表明: 低海拔同一生长期陇蜀杜鹃小枝长度显著大于高海拔,高海拔开花期小枝的花数量和花质量显著高于低海拔。同一海拔花芽期的小枝质量、叶片数量、总叶质量、总叶面积和总叶柄质量均显著大于开花期,单叶质量和单叶柄质量均显著小于开花期。与花芽期相比,开花期叶生物量占比减少了13%,而小枝质量占比显著增加。小枝的生物量权衡表明,花芽期性状倾向于小枝质量,开花期倾向于总叶质量;花芽期的叶偏好分配于单叶质量,低海拔和高海拔开花期叶片分别偏好分配到单叶柄质量和单叶质量。低海拔开花期小枝质量与总叶面积、总叶质量均为异速生长,单叶柄质量和单叶面积为异速生长。海拔和发育阶段共同影响了陇蜀杜鹃功能小枝和叶片的性状。     

Change of Organic Matter in the Decomposing Oak Twigs in the Temperate Forest Ecosystems

Litter bag method was used in this study on the twig decomposition of an oak ( Quercus liaotungensis Koidz. ) which is dominant in the warm temperate deciduous forests. A consecutive five-year investigation was carded out to measure the changes of organic components in the twig litter. The decomposition of oak twigs based on rates of the mass loss during the first five years was simulated using the Olson exponential equation. The simulated data fit well with the observed values. Oak twigs were predicted to reach 95% decomposition within 21 years. During the first five years, the concentration of protein in the remaining litter increased from 3.5 % to 5.5 %, while the concentration of semicellulose decreased from 16.0% to 8.0%. However, there was no obvious change in the concentrations of lignin and cellulose. The loss of lignin, crude-cellulose, cellulose and semicellulose could be well simulated using the Olson exponential equation. However, this was not so effective for protein.     

Xylem Cavitation in Nodes and Internodes of Whole Chorisia insignis H.B. et K. Plants Subjected to Water Stress: Relations Between Xylem Conduit Size and Cavitation

Measurements of cavitation occurring in xylem conduits of differentstem parts in whole Chorisia insignis H.B. et. K. plants subjectedto water stress are reported. Pre-stressed plants were shownto undergo cavitation over 10 times greater than watered ones.The most vulnerable parts of plants were one-year-old twigswhere cavitation reached a peak of over 50 acoustic emissions(AE) min^–1 while in two-year-old twigs AE min^–1were about one half this value. Stem zones were found wherecavitation was typically very low even during water stress:these were one-year-old nodes and junctions where branches meet.Measurements of the inside diameters of xylem conduits and distributionof conduit ends in stem parts where AE were detected, showedthat nodes have a significantly larger percentage of narrowxylem conduits than internodes. Similar ‘constricted zones’were found injunctions with respect to two-year-old twigs. Hereabout 50 per cent of the xylem conduits were as narrow as 20to 50 µm in diameter. The distribution of xylem conduitends show about 3 per cent of them ending in the nodes and 1per cent in the internodes of one-year-old twigs. About 11.6per cent of xylem conduits end in the junctions and about ahalf in two-year-old internodes. Our data would give furtherexperimental evidence to the functional concept of ‘plantsegmentation’ into zones (internodes) more efficient inwater conduction, i.e. with wider xylem conduits but more vulnerableto cavitation and others (nodes and junctions) with oppositecharacteristics. Chorisia insignis, acoustic emissions, water stress, nodes, internodes, xylem conduit size, vessel ends     

暖温带落叶阔叶林辽东栎枝条分解过程中有机物质的变化

应用网袋法对暖温带地区广泛分布的乔木树种辽东栎（ＱｕｅｒｃｕｓｌｉａｏｔｕｎｇｅｎｓｉｓＫｏｉｄｚ．）枝条分解进行了连续５年时间的观察,并测定了凋落物在分解过程中有机质（纤维素、木质素、蛋白质）的变化。用Ｏｌｓｏｎ的指数方程对分解枝条重量的减少进行了模拟,并预测暖温带地区辽东栎枝条完全（９５％）分解需要约２１年的时间。在各有机成分含量的变化上,表现出明显不同的分异。蛋白质的含量随着分解过程的不断进行,从３．５％增加至５．５％,半纤维素则从１６．０％下降至８．０％,而粗纤维和木质素的含量出现了上下波动的情形。经过拟合,凋落物的丢失速率符合Ｏｌｓｏｎ的指数降解曲线,含量较大的粗纤维和木质素的减少也较好地符合Ｏｌｓｏｎ的指数降解曲线,半纤维素也可以用Ｏｌｓｏｎ指数方程拟合,但蛋白质的拟合效果较差。