Diversity of hydraulic traits in nine Cordia species growing in tropical forests with contrasting precipitation

Inter- and intraspecific variation in hydraulic traits was investigated in nine Cordia (Boraginaceae) species growing in three tropical rainforests differing in mean annual precipitation (MAP). Interspecific variation was examined for the different Cordia species found at each site, and intraspecific variation was studied in populations of the widespread species Cordia alliodora across the three sites. Strong intra- and interspecific variation were observed in vulnerability to drought-induced embolism. Species growing at drier sites were more resistant to embolism than those growing at moister sites; the same pattern was observed for populations of C. alliodora. By contrast, traits related to hydraulic capacity, including stem xylem vessel diameter, sapwood specific conductivity (K(s)) and leaf specific conductivity (K(L)), varied strongly but independently of MAP. For C. alliodora, xylem anatomy, K(s), K(L) and Huber value varied little across sites, with K(s) and K(L) being consistently high relative to other Cordia species. A constitutively high hydraulic capacity coupled with plastic or genotypic adjustment in vulnerability to embolism and leaf water relations would contribute to the ability of C. alliodora to establish and compete across a wide precipitation gradient.     

Xylem hydraulic characteristics of subtropical trees from contrasting habitats grown under identical environmental conditions

Five evergreen subtropical tree species growing under identical environmental conditions were investigated to establish which hydraulic properties are genotypically rigid and which show phenotypic plasticity. Maximum xylem-specific conductivity ( k _s) correlated well with the anatomical characteristics (conduit diameter and density) for the four angiosperms Tecomaria capensis , Trichilia dregeana , Cinnamomum camphora and Barringtonia racemosa ; the anatomy of the gymnosperm Podocarpus latifolius was not assessed. Huber values (functional xylem cross-sectional area : leaf area) varied inversely with k _s among species. Maximum leaf-specific conductivity was similar in the five unrelated species. Vulnerability of xylem to cavitation differed between species, as did the relationship between transpiration and water potential. Models of these parameters and isolated midday readings confirm that these trees operate at similar maximum leaf-specific conductivity ( k _l) values. The data are consistent with the hypothesis that conductivity characteristics ( k _l, k _s) are influenced by environment, whereas vulnerability to cavitation is genetically determined.     

木本植物木质部栓塞脆弱性研究新进展

木质部空穴化和栓塞是木本植物在干旱等条件下遭受水分胁迫时产生的木质部输水功能障碍, 在全球气候变化的大背景下, 栓塞脆弱性对干旱响应的研究已成为热点和重要内容。近年来有关木质部栓塞脆弱性与植物输水结构和耐旱性的关系已有大量研究并取得一定成果, 但是, 不同学者在不同地区对不同材料的研究结果存在很大不同。该文就近年来这一研究领域取得的成果及争议问题进行了概括和总结, 主要涉及木质部栓塞脆弱性(P₅₀)及脆弱曲线的建立方法、木质部栓塞脆弱性与木质部结构(导管直径、导管长度、纹孔膜、木质部密度、纤维及纤维管胞)间的关系和木质部栓塞脆弱性与耐旱性的关系, 并对未来工作进行展望, 提出在未来的工作中应对同一树种使用Cochard Cavitron离心机法、Sperry离心机技术与传统方法建立的脆弱曲线进行比较验证、计算P₅₀值、分析植物个体器官水平差异(根、茎、叶)、测定树种生理生态指标, 探索植物栓塞脆弱性与输水结构和耐旱性的关系, 从而评估不同类型植物在未来气候变化下的耐旱能力。     

A review of new research progress on the vulnerability of xylem embolism of woody plants

Xylem cavitation/embolism is the blockage of xylem conduits when woody plants suffer from water stress under drought and other environmental conditions, the study of embolism has become a hot and key topic under global climate change. Recent researches on the relationship between the vulnerability of xylem embolism and hydraulic architecture/drought tolerance have made some progress, however, scholars reached different conclusions based on results from different regions or different materials. This paper reviews the current achievements and controversial viewpoints, which includes indicator of xylem embolism vulnerability (P₅₀), method of vulnerability curve establishment, the relationship between embolism vulnerability and hydraulic architecture (vessel diameter, vessel length, pit area, wood density, fiber and fiber tracheid) and the relationship between embolism vulnerability and drought tolerance of woody plants. Future studies should use Cochard Cavitron centrifuge and Sperry centrifuge coupled with traditional methods to establish vulnerability curves, calculate P₅₀, analyze the difference among different organisms (root, stem, leaf), and measure physiological and ecological indexes. Future studies should be aimed to explore the relationship between the vulnerability of xylem embolism and hydraulic architecture/drought tolerance and to assess drought tolerance ability of different species under future climate change.     

The importance of xylem constraints in the distribution of conifer species

Vulnerability of stem xylem to cavitation was measured in 10 species of conifers using high pressure air to induce xylem embolism. Mean values of air pressure required to induce a 50% loss in hydraulic conductivity (φ₅₀) varied enormously between species, ranging from a maximum of 14.2±0.6 MPa (corresponding to a xylem water potential of −14.2 MPa) in the semi-arid species Actinostrobus acuminatus to a minimum of 2.3±0.2 MPa in the rainforest species Dacrycarpus dacrydioides . Mean φ₅₀ was significantly correlated with the mean rainfall of the driest quarter within the distribution of each species. The value of φ₅₀ was also compared with leaf drought tolerance data for these species in order to determine whether xylem dysfunction during drought dictated drought response at the leaf level. Previous data describing the maximum depletion of internal CO₂ concentration (c_i) in the leaves of these species during artificial drought was strongly correlated with φ₅₀ suggesting a primary role of xylem in effecting leaf drought response. The possibility of a trade-off between xylem conductivity and xylem vulnerability was tested in a sub-sample of four species, but no evidence of an inverse relationship between φ₅₀ and either stem-area specific (K_a) or leaf-area specific conductivity (K₁) was found.     

Limitation of plant water use by rhizosphere and xylem conductance: results from a model

Hydraulic conductivity ( K ) in the soil and xylem declines as water potential ( Ψ ) declines. This results in a maximum rate of steady-state transpiration ( E _crit) and corresponding minimum leaf Ψ ( Ψ _crit) at which K has approached zero somewhere in the soil–leaf continuum. Exceeding these limits causes water transport to cease. A model determined whether the point of hydraulic failure (where K = 0) occurred in the rhizosphere or xylem components of the continuum. Below a threshold of root:leaf area ( A _R: A _L), the loss of rhizosphere K limited E _crit and Ψ _crit. Above the threshold, loss of xylem K from cavitation was limiting. The A _R: A _L threshold ranged from > 40 for coarse soils and/or cavitation-resistant xylem to < 0·20 in fine soils and/or cavitation-susceptible xylem. Comparison of model results with drought experiments in sunflower and water birch indicated that stomatal regulation of E reflected the species' hydraulic potential for extracting soil water, and that the more sensitive stomatal response of water birch to drought was necessary to avoid hydraulic failure. The results suggest that plants should be xylem-limited and near their A _R: A _L threshold. Corollary predictions are (1) within a soil type the A _R: A _L should increase with increasing cavitation resistance and drought tolerance, and (2) across soil types from fine to coarse the A _R: A _L should increase and maximum cavitation resistance should decrease.     

Transport constraints on water use by the Great Basin shrub, Artemisia tridentata

As soil and plant water status decline, decreases in hydraulic conductance can limit a plant's ability to maintain gas exchange. We investigated hydraulic limitations for Artemisia tridentata during summer drought. Water use was quantified by measurements of soil and plant water potential ( Ψ ), transpiration and leaf area. Hydraulic transport capacity was quantified by vulnerability to water stress-induced cavitation for root and stem xylem, and moisture release characteristics for soil. These data were used to predict the maximum possible steady-state transpiration rate ( E _crit) and minimum leaf xylem pressure ( Ψ _crit). Transpiration and leaf area declined by ~ 80 and 50%, respectively, as soil Ψ decreased to –2·6 MPa during drought. Leaf-specific hydraulic conductance also decreased by 70%, with most of the decline predicted in the rhizosphere and root system. Root conductance was projected to be the most limiting, decreasing to zero to cause hydraulic failure if E _crit was exceeded. The basis for this prediction was that roots were more vulnerable to xylem cavitation than stems (99% cavitation at –4·0 versus –7·8 MPa, respectively). The decline in water use during drought was necessary to maintain E and Ψ within the limits defined by E _crit and Ψ _crit.     

Leaf xylem embolism, detected acoustically and by cryo-SEM, corresponds to decreases in leaf hydraulic conductance in four evergreen species

Hydraulic conductance of leaves ( K _leaf) typically decreases with increasing water stress. However, the extent to which the decrease in K _leaf is due to xylem cavitation, conduit deformation or changes in the extra-xylary pathway is unclear. We measured K _leaf concurrently with ultrasonic acoustic emission (UAE) in dehydrating leaves of two vessel-bearing and two tracheid-bearing species to determine whether declining K _leaf was associated with an accumulation of cavitation events. In addition, images of leaf internal structure were captured using cryo-scanning electron microscopy, which allowed detection of empty versus full and also deformed conduits. Overall, K _leaf decreased as leaf water potentials ( Ψ _L) became more negative. Values of K _leaf corresponding to bulk leaf turgor loss points ranged from 13 to 45% of their maximum. Additionally, Ψ _L corresponding to a 50% loss in conductivity and 50% accumulated UAE ranged from −1.5 to −2.4 MPa and from −1.1 to −2.8 MPa, respectively, across species. Decreases in K _leaf were closely associated with accumulated UAE and the percentage of empty conduits. The mean amplitude of UAEs was tightly correlated with mean conduit diameter ( R ² = 0.94, P  = 0.018). These results suggest that water stress-induced decreases in K _leaf in these species are directly related to xylem embolism.     

Evidence for xylem embolism as a primary factor in dehydration-induced declines in leaf hydraulic conductance

Hydraulic conductance of leaves (K(leaf)) typically decreases with increasing water stress and recent studies have proposed different mechanisms responsible for decreasing K(leaf) . We measured K(leaf) concurrently with ultrasonic acoustic emissions (UAEs) in dehydrating leaves of several species to determine whether declining K(leaf) was associated with xylem embolism. In addition, we performed experiments in which the surface tension of water in the leaf xylem was reduced by using a surfactant solution. Finally, we compared the hydraulic vulnerability of entire leaves with the leaf lamina in three species. Leaf hydraulic vulnerability based on rehydration kinetics and UAE was very similar, except in Quercus garryana. However, water potentials corresponding to the initial decline in K(leaf) and the onset of UAE in Q. garryana were similar. In all species tested, reducing the surface tension of water caused K(leaf) to decline at less negative water potentials compared with leaves supplied with water. Microscopy revealed that as the fraction of embolized xylem increased, K(leaf) declined sharply in Q. garryana. Measurements on leaf discs revealed that reductions in lamina hydraulic conductance with dehydration were not as great as those observed in intact leaves, suggesting that embolism was the primary mechanism for reductions in K(leaf) during dehydration.     

Leaf hydraulics and drought stress: response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential ( Ψ _leaf), leaf hydraulic conductance ( K _leaf), and midday transpiration ( E ) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of K _leaf to declining Ψ _leaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and K _leaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of K _leaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Ψ _leaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.     

The effects of acclimation to sunlight on the xylem vulnerability to embolism in Fagus sylvatica L.

We assessed the effects of irradiance received during growth on the vulnerability of Fagus sylvatica L. xylem vessels to water-stress-induced embolism. The measurements were conducted on (1) potted saplings acclimated for 2 years under 100% and 12% incident global radiation and (2) branches collected from sun-exposed and shaded sides of adult trees. Both experiments yielded similar results. Light-acclimated shoots were less vulnerable to embolism. Xylem water potential levels producing 50% loss of hydraulic conductivity were lower in sun-exposed branches and seedlings than in shade-grown ones (–3·0 versus –2·3 MPa on average). The differences in vulnerability were not correlated with differences in xylem hydraulic conductivity nor vessel diameter. Resistance to cavitation was correlated with transpiration rates, midday xylem and leaf water potentials in adult trees. We concluded that vulnerability to cavitation in Fagus sylvatica may acclimate to contrasting ambient light conditions.     

沙柳和柠条茎水力学特性对模拟降雨改变的响应

利用模拟降雨控制试验(对照、降雨增加45%和减少50%),研究了黄土高原水蚀风蚀交错带典型灌木沙柳和柠条茎水力学特性对模拟降雨改变的响应,以揭示两种灌木对未来降雨改变的适应性.结果表明: 沙柳茎比导水率(K_s)、比叶导水率(K_l)和Huber值对增水有显著响应,而对干旱无显著响应;柠条黎明前和正午叶水势、水分传输效率(K_s及K_l)对干旱有显著响应,但对增水无响应.两种灌木不同处理间抵抗栓塞能力无显著差异,沙柳不同处理间正午原位栓塞程度亦无显著差异,而柠条干旱处理正午原位栓塞程度显著增大.沙柳增水处理导管直径和导管面积占边材面积的比例显著增加,干旱导致沙柳导管密度显著增大,水力直径变小;柠条增水处理的木质部结构无明显改变,干旱导致其导管密度和木材密度显著增加.说明增水提升了沙柳的水力功能,而长期干旱显著降低了柠条水力功能,预测在未来气候旱化条件下,柠条的水力适应性可能不如沙柳.     

Xylem traits mediate a trade-off between resistance to freeze-thaw-induced embolism and photosynthetic capacity in overwintering evergreens

Hydraulic traits were studied in temperate, woody evergreens in a high-elevation heath community to test for trade-offs between the delivery of water to canopies at rates sufficient to sustain photosynthesis and protection against disruption to vascular transport caused by freeze-thaw-induced embolism. Freeze-thaw-induced loss in hydraulic conductivity was studied in relation to xylem anatomy, leaf- and sapwood-specific hydraulic conductivity and gas exchange characteristics of leaves. We found evidence that a trade-off between xylem transport capacity and safety from freeze-thaw-induced embolism affects photosynthetic activity in overwintering evergreens. The mean hydraulically weighted xylem vessel diameter and sapwood-specific conductivity correlated with susceptibility to freeze-thaw-induced embolism. There was also a strong correlation of hydraulic supply and demand across species; interspecific differences in stomatal conductance and CO(2) assimilation rates were correlated linearly with sapwood- and leaf-specific hydraulic conductivity. Xylem vessel anatomy mediated an apparent trade-off between resistance to freeze-thaw-induced embolism and hydraulic and photosynthetic capacity during the winter. These results point to a new role for xylem functional traits in determining the degree to which species can maintain photosynthetic carbon gain despite freezing events and cold winter temperatures.     

Regulatory and molecular properties of citrate synthases from methylotrophs

Abstract Gram-negative methylotrophs contain a high- M _r'large' citrate synthase. Gram-positive methylotrophs, on the other hand, contain a 'small' citrate synthase. These differences in M _r coincided partly with differences in NADH sensitivity. Citrate synthases from obligate Gram-negative and Gram-positive facultative methylotrophs were insensitive to feedback inhibition by NADH; only the enzymes from Gram-negative facultative methylotrophs were inhibited by NADH.     

Hydraulic properties of Pinus halepensis,Pinus pinea and Tetraclinis articulata in a dune ecosystem of Eastern Spain

The hydraulic properties of Pinus pinea, Pinus halepensis and Tetraclinis articulata were studied in a coastal dune area from Eastern Spain. The measured variables include vulnerability to xylem embolism (vulnerability curves), hydraulic conductivity and carbon isotopic discrimination in leaves. Leaf water potentials were also monitored in the three studied populations during an extremely dry period. Our results showed that roots had always wider vessels and higher hydraulic conductivity than branches. Roots were also more vulnerable to xylem embolism and operated closer to their hydraulic limit (i.e., with narrower safety margins). Although it was not quantified, extensive root mortality was observed in the two pines during the study period, in agreement with the high values of xylem embolism (> 75%) predicted from vulnerability curves and the water potentials measured in the field. T. articulata was much more resistant to embolism than P. pinea and P. halepensis. Since T. articulata experienced also lower water potentials, safety margins from hydraulic failure were only slightly wider in this species than in the pines. Combining species and tissues, high resistance to xylem embolism was associated with low hydraulic conductivity and with high wood density. Both relationships imply a cost of having a resistant xylem. The study outlined very different water-use strategies for T. articulata and the pines. Whereas T. articulata had a conservative strategy that relied on the low vulnerability of its conducting system to drought-induced xylem embolism, the two pines showed regulatory mechanisms at different levels (i.e., embolism, root demography) that constrained the absorption of water when it became scarce. This revised version was published online in August 2006 with corrections to the Cover Date.     

Ecological relevance of minimum seasonal water potentials

The minimum seasonal water potential a plant experiences, ψ_min, provides an important measure of plant water status, as it reflects the maximum water deficit that leaves and xylem must tolerate to maintain physiological activity. ψ_min also acts as a selective force on xylem structure which, in turn, generates correlations between ψ_min and numerous hydraulic traits. This review focuses on the ecological relevance of ψ_min as a reflection of overall plant hydraulic strategy. The focus is on plant functional strategies with respect to soil drought, but we conclude with preliminary findings on the role of atmospheric drought.     

Vulnerability of xylem to embolism in a mangrove vs an inland species of Rhizophoraceae

Vulnerability of xylem conduits to cavitation and embolism was compared in two species of Rhizophoraceae, the mangrove Rhizophora mangle L. and the tropical moist-forest Cassipourea elliptica (Sw.) Poir. Cavitation (water column breakage preceeding embolism) was monitored by ultrasonic detection; embolism was quantified by its reduction of xylem hydraulic conductivity. Acoustic data were not predictive of loss in hydraulic conductivity, probably because signals from cavitating vessels were swamped by more numerous ones from cavitating fibers. Rhizophora mangle was the less vulnerable to embolism of the two species, losing 80% of its hydraulic conductivity between – 6.0 and – 7.0 MPa. Cassipourea elliptica lost conductivity in linear proportion to decreasing xylem pressure from – 0.5 to – 7.0 MPa. Species vulnerability correlated closely with physiological demands of habitat; the mangrove Rhizophora mangle had field xylem pressures between – 2.5 and – 4.0 MPa. whereas the minimum for Cassipourea elliptica was – 1.6 MPa. Differences in vulnerability between species could be accounted for by differences in the measured air permeability of intervessel pit membranes. According to this explanation, embolism occurs when air enters a water-filled vessel from a neighboring air-filled one via pores in shared pit membranes.     

六种木本植物木质部栓塞化生理生态效应的研究

对６个树种１年生枝木质部栓塞及水势进行了１４个月的连续观测。测定结果表明,木质部栓塞化直接引起木质部导水率下降,从而对树木生长发育节律造成影响。对导水率的影响可通过脆弱曲线看出,６种落叶阔叶树的２类脆弱曲线模型建立：ｙ＝ａｅｂ／ｘ和ｙ＝ｘ＾２／（ａ＋ｂｘ＾２）。通过模型求解,探讨和确定了参数ａ、ｂ的生理生态学意义,并据此比较不同树种木质部栓塞脆弱性的大小,结果为沙棘＞刺槐≥白榆＞加杨＞榛木＞元宝枫     

长白山阔叶树种木质部环孔和散孔结构特征的分化导致其水力学性状的显著差异

木质部的解剖结构特征对树木水分传输功能有重要的影响,阔叶树种木质部环孔和散孔结构特征的分化,很可能导致两个功能类群在水力学结构上存在显著差异,但是有关两个功能类群间细致的水力学性状的对比研究还较少,二者整枝水平的导水率及纹孔水平的细致结构差异尚未见报道.本试验以长白山阔叶红松林常见的3个环孔材和4个散孔材乔木树种为研究对象,对比了两个功能类群树种的整枝导水率(k_shoot)、枝条木质部栓塞脆弱性(p_50)等重要水力学相关生理功能特征,并分析了两个功能类群间的木质部组织水平和纹孔水平上的解剖结构特征差异.结果表明:与茎段导水率差异一致,环孔材树种的整枝导水率也显著高于散孔材,但枝条木质部气穴化抵抗力显著弱于散孔材,二者的差异反映了整枝水平上木质部导水效率和安全性之间的权衡关系,与两个功能类群的水力学生理特征存在显著差异一致,二者在最大导管长度、导管直径、纹孔开口面积、纹孔开口比例等光学和扫描电镜观测解剖结构特征上都存在显著差异;木质部解剖特征(组织水平、纹孔水平)和k_shoot、p_50等生理特征间,以及木质部不同解剖特征之间存在显著的相关,且两个功能类群遵循相同的规律,反映了木质部结构对水分传输功能的重要影响,而导水率和气穴化抵抗力对木质部对立的结构要求,体现了树木水分传输系统构建的生物物理局限性.     

Changes in axial hydraulic conductivity along elongating leaf blades in relation to xylem maturation in tall fescue

Xylem maturation in elongating leaf blades of tall fescue ( Festuca arundinacea ) was studied using staining and microcasting. Three distinctive regions were identified in the blade: (1) a basal region, in which elongation was occurring and protoxylem (PX) vessels were functioning throughout; (2) a maturation region, in which elongation had stopped and narrow (NMX) and large (LMX) metaxylem vessels were beginning to function; (3) a distal, mature region in which most of the longitudinal water movements occurred in the LMX. The axial hydraulic conductivity ( K _h) was measured in leaf sections from all these regions and compared with the theoretical axial hydraulic conductivity ( K _t) computed from the diameter of individual inner vessels. K _t was proportional to K _h throughout the leaf, but K _t was about three times K _h. The changes in K _h and K _t along the leaf reflected the different stages of xylem maturation. In the basal 60 mm region, K _h was about 0.30±0.07 mmol s⁻¹ mm MPa⁻¹. Beyond that region, K _h rapidly increased with metaxylem element maturation to a maximum value of 5.0±0.3 mmol s⁻¹ mm MPa⁻¹, 105 mm from the leaf base. It then decreased to 3.5±0.2 mmol s⁻¹ mm MPa⁻¹ near the leaf tip. The basal expanding region was observed to restrict longitudinal water movement. There was a close relationship between the water deposition rate in the elongation zone and the sum of the perimeters of PX vessels. The implications of this longitudinal vasculature on the partitioning of water between growth and transpiration is discussed.