Coordination and trade‐offs between leaf and stem hydraulic traits and stomatal regulation along a spectrum of isohydry to anisohydry

The degree of plant iso/anisohydry, a widely used framework for classifying species‐specific hydraulic strategies, integrates multiple components of the whole‐plant hydraulic pathway. However, little is known about how it associates with coordination of functional and structural traits within and across different organs. We examined stem and leaf hydraulic capacitance and conductivity/conductance, stem xylem anatomical features, stomatal regulation of daily minimum leaf and stem water potential (Ψ), and the kinetics of stomatal responses to vapour pressure deficit (VPD) in six diverse woody species differing markedly in their degree of iso/anisohydry. At the stem level, more anisohydric species had higher wood density and lower native capacitance and conductivity. Like stems, leaves of more anisohydric species had lower hydraulic conductance; however, unlike stems, their leaves had higher native capacitance at their daily minimum values of leaf Ψ. Moreover, rates of VPD‐induced stomatal closure were related to intrinsic rather than native leaf capacitance and were not associated with species' degree of iso/anisohydry. Our results suggest a trade‐off between hydraulic storage and efficiency in the leaf, but a coordination between hydraulic storage and efficiency in the stem along a spectrum of plant iso/anisohydry.     

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.     

三株耐铅锌菌的分离、鉴定及其吸附能力

以铅锌矿渣盆栽试验中长势较好的耐性植物夹竹桃(Nerium indicum)的根际土壤为材料,进行耐铅锌优势菌株的分离鉴定,探讨影响铅锌吸附的因素及其吸附机理。结果表明:(1)从土样中分离筛选出3株耐铅锌菌株(B1、B4、B14),3株菌均能在Pb2+、Zn2+浓度为600 mg·L-1的牛肉膏蛋白胨培养基上生长,经形态和分子生物学鉴定分别为蜡样芽孢杆菌(Bacillus cereus)或炭疽杆菌(Bacillus anthracis)、解硫胺素硫胺素芽孢杆菌(Aneurinibacillus aneurinilyticus)和藤黄微球菌(Micrococcus luteus)。(2)对影响菌株吸附铅、锌的p H、吸附时间、初始菌量3个因素进行分析,发现菌株B1在p H为5.0、吸附时间为50 min、初始菌量为0.06 g时,对Pb2+、Zn2+的去除率分别可达84.22%和70.66%。菌株B4在p H为6.0、吸附时间为50 min、初始菌量为0.18 g时,对Pb2+、Zn2+的去除率分别可达72.63%和54.17%。菌株B14在p H为4.0、吸附时间为60 min、初始菌量为0.10 g时对Pb2+、Zn2+的吸附率分别为77.56%和50.63%。(3)扫描电镜观察和红外光谱分析显示:3株菌对Pb2+、Zn2+的吸附主要是细胞表面的吸附,还存在一定的内部吸收;羟基(O-H)、胺基(N-H)、烷基、酰胺基(CONH-)是吸附、络合或螯合金属离子或原子的主要活性基团,重金属与菌株表面的活性基团结合反应是其吸附Pb2+、Zn2+的主要作用机制。     

Stem water storage capacity and efficiency of water transport: their functional significance in a Hawaiian dry forest

We investigated the contribution of internal water storage and efficiency of water transport to the maintenance of water balance in six evergreen tree species in a Hawaiian dry forest. Wood‐saturated water content, a surrogate for relative water storage capacity, ranged from 70 to 105%, and was inversely related to its morphological correlate, wood density, which ranged between 0·51 and 0·65 g cm^?3. Leaf‐specific conductivity (k_L) measured in stem segments from terminal branches ranged from 3 to 18 mmol m^?1 s^?1 MPa^?1, and whole‐plant hydraulic efficiency calculated as stomatal conductance (g) divided by the difference between predawn and midday leaf water potential (Ψ_L), ranged from 70 to 150 mmol m^?2 s^?1 MPa^?1. Hydraulic efficiency was positively correlated with k_L (r² = 0·86). Minimum annual Ψ_L ranged from ? 1·5 to ? 4·1 MPa among the six species. Seasonal and diurnal variation in Ψ_L were associated with differences among species in wood‐saturated water content, wood density and k_L. The species with higher wood‐saturated water content were more efficient in terms of long‐distance water transport, exhibited smaller diurnal variation in Ψ_L and higher maximum photosynthetic rates. Smaller diurnal variation in Ψ_L in species with higher wood‐saturated water content, k_L and hydraulic efficiency was not associated with stomatal restriction of transpiration when soil water deficit was moderate, but avoidance of low minimum seasonal Ψ_L in these species was associated with a substantial seasonal decline in g. Low seasonal minimum Ψ_L in species with low k_L, hydraulic efficiency, and wood‐saturated water content was associated with higher leaf solute content and corresponding lower leaf turgor loss point. Despite the species‐specific differences in leaf water relations characteristics, all six evergreen tree species shared a common functional relationship defined primarily by k_L and stem water storage capacity.     

Wood anatomical correlates with theoretical conductivity and wood density across China: evolutionary evidence of the functional differentiation of axial and radial parenchyma

Background and Aims

In recent years considerable effort has focused on linking wood anatomy and key ecological traits. Studies analysing large databases have described how these ecological traits vary as a function of wood anatomical traits related to conduction and support, but have not considered how these functions interact with cells involved in storage of water and carbohydrates (i.e. parenchyma cells).

Methods

We analyzed, in a phylogenetic context, the functional relationship between cell types performing each of the three xylem functions (conduction, support and storage) and wood density and theoretical conductivity using a sample of approx. 800 tree species from China.

Key Results

Axial parenchyma and rays had distinct evolutionary correlation patterns. An evolutionary link was found between high conduction capacity and larger amounts of axial parenchyma that is probably related to water storage capacity and embolism repair, while larger amounts of ray tissue have evolved with increased mechanical support and reduced hydraulic capacity. In a phylogenetic principal component analysis this association of axial parenchyma with increased conduction capacity and rays with wood density represented orthogonal axes of variation. In multivariate space, however, the proportion of rays might be positively associated with conductance and negatively with wood density, indicating flexibility in these axes in species with wide rays.

Conclusions

The findings suggest that parenchyma types may differ in function. The functional axes represented by different cell types were conserved across lineages, suggesting a significant role in the ecological strategies of the angiosperms.     

A new method for vulnerability analysis of small xylem areas reveals that compression wood of Norway spruce has lower hydraulic safety than opposite wood

Compression wood is formed at the underside of conifer twigs to keep branches at their equilibrium position. It differs from opposite wood anatomically and subsequently in its mechanical and hydraulic properties. The specific hydraulic conductivity (k_s) and vulnerability to drought‐induced embolism (loss of conductivity versus water potential ψ) in twigs of Norway spruce [Picea abies (L.) Karst.] were studied via cryo‐scanning electron microscope observations, dye experiments and a newly developed ‘Micro‐Sperry’ apparatus. This new technique enabled conductivity measurements in small xylem areas by insertion of syringe cannulas into twig samples. The hydraulic properties were related to anatomical parameters (tracheid diameter, wall thickness). Compression wood exhibited 79% lower k_s than opposite wood corresponding to smaller tracheid diameters. Vulnerability was higher in compression wood despite its narrower tracheids and thicker cell walls. The P₅₀ (ψ at 50% loss of conductivity) was ?3.6 MPa in opposite but only ?3.2 MPa in compression wood. Low hydraulic efficiency and low hydraulic safety indicate that compression wood has primarily a mechanical function.     

Tapering of xylem conduits and hydraulic limitations in sycamore (Acer pseudoplatanus) trees

Vertical conduit tapering is proposed as an effective mechanism to almost eliminate the increase in hydraulic resistance with increased height. Despite this potential role, very little is known about its changes during ontogeny. Here, conduit tapering and stem morphology of young/small and old/tall individuals of Acer pseudoplatanus in the field, as well as 3-yr-old grafted trees from both age classes, were analysed. The distribution of hydraulic resistance along stems was also determined in a subsample of trees. Substantial conduit tapering was found in small trees (field-grown and grafted from both age classes), whereas values were lower in tall trees, indicating that tapering was a size-related, not an age-related process. Apical conduit diameters were larger in tall trees and were inversely correlated with the degree of tapering. Hydraulic resistance increased less than linearly with distance from the apex. Its scaling against distance was indistinguishable from that predicted from anatomical measurements. Conduit tapering was an effective but partial mechanism of compensation for the increase in hydraulic resistance with tree height. Size-related changes in tapering and in apical conduit diameters may be explained by the combined need to reduce the build-up of hydraulic resistance while minimizing the carbon costs of building vessel walls.     

How does a gymnosperm branch (Pseudotsuga menziesii) assume the hydraulic status of a main stem when it takes over as leader?

In most gymnosperms, resistance to the flow of water per unit path length through the main stem is less than that of lateral branches. Using branches, leaders, and branches that have replaced missing leaders ('branch-leaders'), we tested the hypothesis that branch-leaders are at a hydraulic disadvantage. Reduced xylem transport efficiency in branch-leaders relative to leaders could be expected both because of an initial disparity in hydraulic capacity, and because of the relatively impermeable compression wood formed in branch-leaders during shoot reorientation. By subsampling branch-leaders, we also tested the hypothesis that opposite wood (formed directly opposite compression wood) is more permeable than normal wood, and could, therefore, compensate for the presence of compression wood at the whole shoot level. Fifteen months after leader removal, branch-leaders were intermediate between branches and leaders in their ability to supply foliage with water, suggesting a transition towards leader status that was not yet complete. Increased hydraulic capacity in branch-leaders was the result of increased xylem cross-sectional area per unit foliage, rather than an increase in permeability. Among subsampled wood types from basal branch-leader segments, opposite wood was significantly less permeable than normal wood, suggesting that it does not compensate for the presence of compression wood.     

The earliest wood and its hydraulic properties documented in c. 407‐million‐year‐old fossils using synchrotron microtomography

We document xylem structure and hydraulic properties in the earliest woody plant A rmoricaphyton chateaupannense gen. nov. & sp. nov. based on c. 407‐million‐year‐old fossils from the Armorican Massif, western France. The plant was small, and the woody axes were narrow and permineralized in pyrite (FeS₂). We used standard palaeobotanical methods and employed propagation phase contrast X‐ray synchrotron microtomography (PPC‐SRμCT) to create three‐dimensional images of the wood and to evaluate its properties. The xylem comprised tracheids and rays, which developed from a cambium. Tracheids possessed an early extinct type of scalariform bordered pitting known as P‐type. Our observations indicate that wood evolved initially in plants of small stature that were members of Euphyllophytina, a clade that includes living seed plants, horsetails and ferns. Hydraulic properties were calculated from measurements taken from the PPC‐SRμCT images. The specific hydraulic conductivity of the xylem area was calculated as 8.7 kg m^?1 s^?1 and the mean cell thickness‐to‐span ratio (t/b)² of tracheids was 0.0372. The results show that the wood was suited to high conductive performance with low mechanical resistance to hydraulic tension. We argue that axis rigidity in the earliest woody plants initially evolved through the development of low‐density woods. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 423–437.     

Mechanical reinforcement of tracheids compromises the hydraulic efficiency of conifer xylem

Wood structure and function of juvenile wood from 18 conifer species from four conifer families (Araucariaceae, Cupressaceae, Pinaceae and Podocarpaceae) were examined for a trade-off between wood reinforcement and hydraulic efficiency. Wood density and tracheid 'thickness-to-span' ratio were used as anatomical proxies for mechanical properties. The thickness:span represented the ratio of tracheid double wall thickness to lumen diameter. Hydraulic resistivity (R) of tracheids on a cross-sectional area basis (RCA) increased over 50-fold with increasing density and thickness:span, implying a strength versus efficiency conflict. The conflict arose because density and thickness:span were increased by narrowing tracheid diameter rather than by thickening walls, which may be developmentally difficult. In the Pinaceae and Cupressaceae species, density and thickness:span correlated strongly with protection from drought-induced embolism, suggesting that mechanical strength was required in part to withstand tracheid collapse by negative sap pressure. These species showed a corresponding trade-off between increasing RCA and embolism protection. In contrast, species of Podocarpaceae and Araucariaceae were overbuilt for their embolism protection and were hydraulically inefficient, having greater density, thickness:span and RCA, none of which were correlated with vulnerability to embolism.     

Long-term acclimatization of hydraulic properties, xylem conduit size, wall strength and cavitation resistance in Phaseolus vulgaris in response to different environmental effects

Phaseolus vulgaris grown under various environmental conditions was used to assess long-term acclimatization of xylem structural characteristics and hydraulic properties. Conduit diameter tended to be reduced and 'wood' density (of 'woody' stems) increased under low moisture ('dry'), increased soil porosity ('porous soil') and low phosphorus ('low P') treatments. Dry and low P had the largest percentage of small vessels. Dry, low light ('shade') and porous soil treatments decreased P50 (50% loss in conductivity) by 0.15-0.25 MPa (greater cavitation resistance) compared with 'controls'. By contrast, low P increased P50 by 0.30 MPa (less cavitation resistance) compared with porous soil (the control for low P). Changes in cavitation resistance were independent of conduit diameter. By contrast, changes in cavitation resistance were correlated with wood density for the control, dry and porous soil treatments, but did not appear to be a function of wood density for the shade and low P treatments. In a separate experiment comparing control and porous soil plants, stem hydraulic conductivity (kh), specific conductivity (ks), leaf specific conductivity (LSC), total pot water loss, plant biomass and leaf area were all greater for control plants compared to porous soil plants. Porous soil plants, however, demonstrated higher midday stomatal conductance to water vapour (gs), apparently because they experienced proportionally less midday xylem cavitation.     

Testing the divergent adaptation of two congeneric tree species on a rainfall gradient using eco‐physio‐morphological traits

In tropical Africa, evidence of widely distributed genera transcending biomes or habitat boundaries has been reported. The evolutionary processes that allowed these lineages to disperse and adapt into new environments are far from being resolved. To better understand these processes, we propose an integrated approach, based on the eco‐physio‐morphological traits of two sister species with adjacent distributions along a rainfall gradient. We used wood anatomical traits, plant hydraulics (vulnerability to cavitation, wood volumetric water content, and hydraulic capacitance), and growth data from the natural habitat, in a common garden, to compare species with known phylogeny, very similar morphologically, but occupying contrasting habitats: Erythrophleum ivorense (wet forest) and Erythrophleum suaveolens (moist forest and forest gallery). We identified some slight differences in wood anatomical traits between the two species associated with strong differences in hydraulics, growth, and overall species distribution. The moist forest species, E. suaveolens, had narrower vessels and intervessel pits, and higher vessel cell‐wall reinforcement than E. ivorense. These traits allow a high resistance to cavitation and a continuous internal water supply of the xylem during water shortage, allowing a higher fitness during drought periods, but limiting growth. Our results confirm a trade‐off between drought tolerance and growth, controlled by subtle adaptations in wood traits, as a key mechanism leading to the niche partitioning between the two Erythrophleum species. The generality of this trade‐off and its importance in the diversification of the African tree flora remains to be tested. Our integrated eco‐physio‐morpho approach could be the way forward.     

Radial Gradients in Wood Specific Gravity,Water and Gas Content in Trees of a Mexican Tropical Rain Forest

Cell walls, water, and gas that have mechanical and physiological functions in wood, and wood specific gravity (WSG) is related to demographic traits. To understand variation in wood structure and function, we analyzed radial changes in WSG, in the gas and the water fractions from trees growing in four different habitats in a southern Mexican rain forest. Mean WSG was 0.55 ± 0.16, slightly lower than reported for other tropical forests. In 27 species, WSG decreased and in two species, it increased from pith to bark with a strong (r² = 0.65) negative correlation between WSG in the center of the tree and the radial WSG gradient. Habitat had some effect on mean WSG and trees growing on karst had significantly higher WSG than the same species growing on alluvial soil. The cell wall, water, and gas fractions accounted for 35 percent (range: 16–50), 42 percent (28–65), and 23 percent (2–56), respectively, of wood volume, with a negative correlation between the gas and the cell wall and between the gas and the water fractions, but not between the cell wall and water fractions. Radially increasing WSG is advantageous for pioneer trees with fast initial growth. We found that the water displacement method may result in biased WSG estimates. To increase the accuracy of WSG data, we suggest to measure sample volume geometrically using a constant diameter (that of the borer tip), to include radial variation in WSG, and to consider for possible site effects on species‐specific WSG.     

木材上的微生物类群对木材的分解及其演替规律

论述了木材上的微生物类群以及各类群在木材生物分解过程中的演替规律 ,并解释了活立木心材能够发生腐朽的原因。结果表明 :能够生长在木材上的微生物类群有木材腐朽菌、木材软腐菌、木材变色菌、污染性霉菌、细菌、放线菌等多种。这些微生物类群共同合作完成对复杂的木质有机物质的生物分解。它们按一定次序进行作用 ,在木材生物分解的不同时期显示出明显的菌种协调与演替规律。一般情况是细菌、一些半知菌、接合菌和子囊菌等先驱微生物首先侵入 ,然后草本对策的木材腐朽菌开始出现 ,最后由竞争对策或忍耐对策的木腐菌取代草本对策的木腐菌 ,这时木材的分解过程就进入稳定的发展阶段 ,最后使木材分解或腐朽。木材腐朽最终是腐殖化阶段 ,这时微生物群落被土壤习居菌如毛霉、青霉、木霉、镰刀菌及细菌与放线菌等所取代。     

Studying global change through investigation of the plastic responses of xylem anatomy in tree rings

Variability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, sub-annual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that low-cost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.