树木高生长限制的几个假说

树木生长到一定年龄后高生长停滞,对这一现象的解释存在很多争议.成熟假说认为树木顶端分生组织分裂活性下降导致树木高生长减慢.营养限制假说认为土壤中营养元素(特别是氮素)在植物活体或枯落物中积累使土壤中可利用的养分含量降低,细根生物量增加和叶片光合能力下降导致了地上部分生长的减缓.呼吸假说认为边材呼吸消耗随个体发育的增加使投入到高生长的碳减少.水力限制假说认为水分运输阻力随高度增加而增加导致了叶片总光合碳同化下降,分配到高生长的生物量减少.树木发展假说认为植物用多种调节机制克服随个体发育增加的水力阻力,包括叶片结构和生理特征的变化,叶/边材面积比降低,边材渗透性和树干储水能力的增加等.水力限制假说得到了较多的关注,对不同高度树木的叶比导率、光合特征和树干生长量等测定结果支持这一假说.但对这一假说也存在很多的争议,主要表现在:水力阻力是否确实随高度的增加而增加,水力阻力的分布,补偿机制的作用和生物量分配转变等.本文综述了树木高生长限制的4个假说以及争论的焦点,并总结了目前研究的热点问题和今后的研究方向.     

树木储存水对水力限制的补偿研究进展

水力限制假说认为水分传输阻力与水势梯度协同调节气孔气体交换、影响CO₂的吸收进而限制树木的高生长,比较合理地解释了不同生境树木极限高度和高生长的差异.但该理论并未考虑到水力结构和其他生物学特征在树木向上生长的同时会进行适应性调整,以减弱逐渐增加的水分传输阻力.树木储存水对缓解木质部的水力限制、控制叶片水势的波动具有重要的生物学意义,也可能是气孔调节和水分状况变化格型的重要决定因素,其对水力限制可起到部分补偿作用.本文对储存水影响树木水分利用过程中的水力限制进行了综述,探讨了储存水补偿水力限制的可能性机理以及相对应的研究方法,并对未来的研究方向进行了展望.     

云南哀牢山6种常绿阔叶树木质部解剖特征的轴向和径向变化

West、Brown和Enquist提出的树木水分传导的分形网络模型(简称WBE模型)认为,树木连续分枝之间的导管或管胞直径按照一定的比率均匀变细,其总的水力阻力与水分传导的路径长度无关,从而使不同部位叶片获得基本相当的水分供应。该模型对树木高生长的水力限制假说提出了置疑。为了验证WBE模型中树木导管或管胞均匀变细的假说,该文研究了云南哀牢山中湿性常绿阔叶林中6种常绿阔叶树, 腾冲栲(Castanopsis wattii)、景东石砾(Lithocarpus chintungensis)、木果石砾(L. xylocarpus)、长尾青冈(Cyclobalanopsis stewardiana)、滇木荷(Schima noronhae)和舟柄茶(Hartia sinensis)木质部解剖特征随树高和年龄的变化。对这6个树种共14株样木进行了不同高度树干圆盘和边材生长轮取样,样木的高度为15~25 m,按照常规木材解剖的处理和分析方法,在显微镜下测定木材切片的导管直径和密度等特征。结果表明:在14株样木中,有4株树木导管直径随树木高度增加呈线性减小, 1株没有明显变化,其它9株树木导管直径在树冠以下的树干部分变化幅度较小或没有明显变化,而从树冠基部往上直到树木顶端导管直径显著减小。同一植株随着高度的增加,导管密度增加并且在树冠内增加更显著。有三分之一的树木导管占边材面积的比例随树高增加没有明显变化,其余树木导管占边材面积比在树冠以上有所减小。多数树木理论比导率在树冠以下没有明显变化而在树冠基部往上显著降低。在从髓芯开始往外的20~40个年轮范围内导管直径增加显著,但大部分植株导管直径在40个年轮后趋于稳定。不同高度圆盘导管直径随形成层发育时间的变化呈相似的趋势,并且相同发育年龄的导管直径没有明显差异。该文的研究结果说明,导管直径的轴向和径向变化一定程度上补偿了水分运输阻力随树木个体增大而增加的缺陷,但是6种常绿阔叶树树干的导管基本不按一定比率均匀变细,不支持WBE模型。     

人工林生产力年龄效应及衰退机理研究进展

研究人工林生长规律具有重要的经济和生态意义。同龄林林分郁闭后,地上部净初级生产量随着林龄增加而降低的现象近几十年引起了林业工作者的兴趣和注意并成为研究热点。多数研究试图通过光合生理、林分营养、生物量分配和林分结构等随林龄的变化规律来解释林分生产力衰退机理。研究认为,林分郁闭后水分传输阻力的增加减少了树木的光合能力;林地养分的减少使得根系生物量分配增加,导致林分叶面积减少,树木光合能力下降;对资源的竞争使得树木优势度发生变化,资源利用率降低。光合能力、林分叶面积和资源利用率的降低以及根系生物量分配的增加是林分生产力衰退的关键,而林分呼吸和林木衰老的作用不大。今后深入研究树体水分运输及其补偿机制、逆境下根系的生长过程及适应机制,并跟踪研究林分生长规律,更有助于揭示人工林生产力衰退的实质。     

4种高大树木的叶片性状及WUE随树高的变化

为了解西双版纳北热带雨林高大树木树顶叶片性状对通道阻力增长引起的水力限制增强及高光和季节性干旱等气候条件的响应,对该区乔木浆果乌桕(Sapium baccatum Roxb)、思茅木姜子(Litsea pierrei var. szemois liou)、小叶藤黄(Garcinia cowa Roxb)及共生木质藤本黑风藤(Fissisfigma polyanthum (Hook. f. et Thoms.)Merr.)的叶片形态解剖结构、光合色素、水分利用效率(WUE)等随冠层高度的变化及种间差异进行了研究。结果表明:小叶藤黄和思茅木姜子的叶片(310.14、319.73 μm)和角质层(6.06、5.13 μm)都较厚、细胞较大(21.48、27.09 μm),光合色素含量则较低;黑风藤栅栏组织所占的比例最大、光合色素含量也最高,但叶片薄、WUE最低;浆果乌桕的WUE最高。随着冠层高度的增加,4种树木的叶厚、栅栏组织及角质层厚度、LMA、P/S和TPM/LT均增加、细胞变小,其中黑风藤的变幅最大。4树种的叶绿素和类胡萝卜素含量均随冠层的增高而减少,δ¹³C和WUE则随树冠增高而增大(黑风藤的变幅小于3种乔木);Δ则相反。上述结果表明4种树木冠层上部叶片偏向旱生结构和水分利用效率增加,暗示树顶叶片可能受到了水分胁迫,从而在结构上偏向于减少水分散失、功能上提高对水分的利用效率以适应水分亏缺;同时,随冠层增加光合色素含量减少,暗示其光合碳同化能力也降低。上述结果支持了水力限制假说中由于通道阻力增大引起树顶水力限制增强,大树可能会通过减少光和碳的获得而减慢树高生长的假设。     

长白山不同海拔岳桦非结构碳水化合物含量的变化

 通常认为, 随着林木不断接近其海拔分布极限, 光合作用产量不断下降, 导致碳水化合物供应不足(碳供应限制), 或者低温限制了碳投资(生长限制)。植物组织内非结构性碳水化合物(Nonstructural carbohydrates, NSC)的含量反映了植物碳供应与碳吸收的平衡。为了检验“碳供应限制”和“生长抑制”假说, 我们对长白山海拔1 700~ 2 050 m的自然生境下生长的岳桦(Betula ermanii)的叶片和枝条组织的NSC含量进行了比较。结果表明: 岳桦叶片的NSC含量随海拔升高变化不显著, 枝条的NSC含量随海拔升高显著增加; 叶片和枝条中淀粉含量与可溶性总糖含量的比值均随海拔的升高而减小; 林线附近的岳桦林木不存在碳水化合物供应不足的问题, 这在一定程度上表明生长限制导致长白山岳桦林线的形成。     

陕北沙地高龄小叶杨光合速率下降的水力限制

黄土高原地区“小老树”现象多出现在成年树,幼树相对较少.为探讨树龄影响“小老树”形成的机制,以该地区“小老树”发生面积最大的树种小叶杨为例,研究了3个不同树龄(低龄:13～15 a;中龄:31～34 a;高龄:49～54 a)树木的生长、光合、水力学特性.结果表明: 随树龄增加,小叶杨枯稍长度显著增加,叶片光合速率、气孔导度和蒸腾速率显著下降,整株水力导度也显著下降,不同测定时间的光合速率、气孔导度与整株水力导度呈显著正相关,表明树龄增加引起的光合速率下降可能与整株水力导度下降有关.与中、低龄相比,高龄小叶杨枝干和叶片抵抗空穴化的能力(P₅₀)更强,但通过脆弱性曲线估算的不同树龄正午时的枝干栓塞程度和叶片水力导度无显著差异,表明高龄小叶杨土-根系统水流阻力的增加可能是导致其整株水力导度降低的重要原因.     

氮对黄花蒿生长、光合特性和青蒿素含量的影响

对不同氮处理黄花蒿生长、生物量分配、青蒿素含量和光合特性进行测定,结果表明:(1)供氮量在0～0.4g.kg-1之间,黄花蒿叶片单位重量的氮含量、最大净光合速率、光饱和点和表观量子效率均随供氮量的增大而增加,之后开始下降。在较大范围内,环境氮含量越高,黄花蒿的光合能力越强,能够利用的光强也更高;(2)黄花蒿根生物量分数和根冠比均随供氮量的减少而显著增大,低氮时分配更多的生物量到养分吸收器官,有利于减少氮素对生长的限制,供氮量在0.1～0.6g.kg-1之间,黄花蒿叶生物量分数随供氮量的增加而增大,高氮时更多的生物量投入到碳同化器官,提高了植株的竞争能力;(3)无论以最大净光合速率、地径、叶片生物量还是以总生物量来衡量,均以0.4g.kg-1氮处理的植株生长得最好,0.2g.kg-1氮处理的青蒿素含量最高,生产中推荐使用0.2g.kg-1剂量的氮更经济。     

中金杨幼苗叶功能性状与碳氮分配对氮添加的响应

氮沉降对森林生态系统产生了重要影响,但树木个体对氮沉降的生理生态响应过程和机制有待进一步明确。为探究杨树幼苗功能叶经济性状、植株生长以及生物量和碳氮分配对氮添加的响应,本文以欧美杂交杨中金杨2号(Populus×euramericana‘Zhongjin2’)幼苗为材料,研究了连续2年氮添加梯度处理(0、3、6和9 g NH₄NO₃·a^-1·plant^-1)对其叶片光合速率、叶经济性状、生长以及器官间碳氮分配的影响。结果表明：氮添加第一年,随氮添加浓度增加,中金杨叶片变大变薄,叶肉组织厚度显著降低,单位叶面积的净光合速率和气孔导度则随氮添加浓度增加而先升后降,而蒸腾速率、瞬时水分利用效率和光合氮利用效率不变;单株生物量和碳氮元素积累量显著增加,茎器官的碳投资显著增加而根的碳投资显著降低;氮添加第二年,功能叶光合固碳速率随氮添加量增加更快达到饱和,且高剂量氮添加处理表现出氮毒害现象;仅低量氮添加处理的单株生物量积累显著增加,并且进一步增加了对茎器官的碳投资,而中高剂量氮添加的根系生长明显受到抑制。氮添...     

树高对马占相思整树水分利用的效应

利用Granier热消散探针,于2004年观测了华南丘陵坡地常见绿化先锋树种马占相思(22年生)的树干液流,同时监测林冠上方的光合有效辐射、气温、相对湿度和0～30 cm的土壤体积含水量.结合树木的形态特征、液流密度和简化的Whitehead & Jarvis公式,分别计算了整树蒸腾、冠层气孔导度和叶面积/边材面积比值,分析了树高对整树蒸腾、冠层气孔导度和叶面积/边材面积比值的影响.结果表明：土壤水分充足时,马占相思整树蒸腾随树高呈二次多项式增加(P<0.01),冠层气孔导度日变化均呈“单峰”格型;在所有光合有效辐射范围内,高树的参比冠层气孔导度和冠层气孔导度对水汽压亏缺的敏感性均高于矮树;叶面积/边材面积比值为（1.837±0.048） m²·cm^-2,并与树高呈幂函数关系.随着树木高度的增加,马占相思没有发生明显的水力限制和补偿.     

The hydraulic limitation hypothesis revisited

We proposed the hydraulic limitation hypothesis (HLH) as a mechanism to explain universal patterns in tree height, and tree and stand biomass growth: height growth slows down as trees grow taller, maximum height is lower for trees of the same species on resource-poor sites and annual wood production declines after canopy closure for even-aged forests. Our review of 51 studies that measured one or more of the components necessary for testing the hypothesis showed that taller trees differ physiologically from shorter, younger trees. Stomatal conductance to water vapour (g(s)), photosynthesis (A) and leaf-specific hydraulic conductance (K L) are often, but not always, lower in taller trees. Additionally, leaf mass per area is often greater in taller trees, and leaf area:sapwood area ratio changes with tree height. We conclude that hydraulic limitation of gas exchange with increasing tree size is common, but not universal. Where hydraulic limitations to A do occur, no evidence supports the original expectation that hydraulic limitation of carbon assimilation is sufficient to explain observed declines in wood production. Any limit to height or height growth does not appear to be related to the so-called age-related decline in wood production of forests after canopy closure. Future work on this problem should explicitly link leaf or canopy gas exchange with tree and stand growth, and consider a more fundamental assumption: whether tree biomass growth is limited by carbon availability.     

Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns

The hydraulic limitation hypothesis of Ryan & Yoder (1997, Bioscience 47, 235-242) suggests that water supply to leaves becomes increasingly difficult with increasing tree height. Within the bounds of this hypothesis, we conjectured that the vertical hydrostatic gradient which gravity generates on the water column in tall trees would cause a progressive increase in xylem 'safety' (increased resistance to embolism and implosion) and a concomitant decrease in xylem 'efficiency' (decreased hydraulic conductivity). We based this idea on the historically recognized concept of a safety-efficiency trade-off in xylem function, and tested it by measuring xylem conductivity and vulnerability to embolism of Sequoia sempervirens branches collected at a range of heights. Measurements of resistance of branch xylem to embolism did indeed show an increase in 'safety' with height. However, the expected decrease in xylem 'efficiency' was not observed. Instead, sapwood-specific hydraulic conductivities (Ks) of branches increased slightly, while leaf-specific hydraulic conductivities increased dramatically, with height. The latter could be largely explained by strong vertical gradients in specific leaf area. The increase in Ks with height corresponded to a decrease in xylem wall fraction (a measure of wall thickness), an increase in percentage of earlywood and slight increases in conduit diameter. These changes are probably adaptive responses to the increased transport requirements of leaves growing in the upper canopy where evaporative demand is greater. The lack of a safety-efficiency tradeoff may be explained by opposing height trends in the pit aperture and conduit diameter of tracheids and the major and semi-independent roles these play in determining xylem safety and efficiency, respectively.     

Comparative hydraulic and anatomic properties in palm trees (Washingtonia robusta) of varying heights: implications for hydraulic limitation to increased height growth

As trees grow taller, the energetic cost of moving water to the leaves becomes higher and could begin to limit carbon gain and subsequent growth. The hydraulic limitation hypothesis states that as trees grow taller, the path length and therefore frictional resistance of water flow increases, leading to stomatal closure, reduced photosynthesis and decreased height growth in tall trees. Although this hypothesis is supported by the physical laws governing water movement in trees, its validation has been complicated by the complex structure of most tree species. Therefore, this study tested the hydraulic limitation hypothesis in Washingtonia robusta (H. Wendl.), a palm that, while growing to tall heights, is still structurally simple enough to act as a model organism for testing. There were no discernable relationships between tree height and stomatal conductance, stomatal densities, guard cell lengths, leaf dry mass per unit area (LMA) or sap flux, suggesting that these key aspects of hydraulic limitation are not reduced in taller palms. Taller palms did, however, have higher maximum daily photosynthetic assimilation rates, lower minimum leaf water potentials that occurred earlier in the day and fewer, smaller leaves than did shorter palms. Leaf epidermal cells were also smaller in taller palms compared with shorter ones. These findings are consistent with hydraulic compensation in that tall palms may be overcoming the increased path length resistance through smaller, more efficient leaves and lower leaf water potentials than shorter palms.     

Linking physiological processes with mangrove forest structure: phosphorus deficiency limits canopy development, hydraulic conductivity and photosynthetic carbon gain in dwarf Rhizophora mangle

Spatial gradients in mangrove tree height in barrier islands of Belize are associated with nutrient deficiency and sustained flooding in the absence of a salinity gradient. While nutrient deficiency is likely to affect many parameters, here we show that addition of phosphorus (P) to dwarf mangroves stimulated increases in diameters of xylem vessels, area of conductive xylem tissue and leaf area index (LAI) of the canopy. These changes in structure were consistent with related changes in function, as addition of P also increased hydraulic conductivity ( K _s), stomatal conductance and photosynthetic assimilation rates to the same levels measured in taller trees fringing the seaward margin of the mangrove. Increased xylem vessel size and corresponding enhancements in stem hydraulic conductivity in P fertilized dwarf trees came at the cost of enhanced midday loss of hydraulic conductivity and was associated with decreased assimilation rates in the afternoon. Analysis of trait plasticity identifies hydraulic properties of trees as more plastic than those of leaf structural and physiological characteristics, implying that hydraulic properties are key in controlling growth in mangroves. Alleviation of P deficiency, which released trees from hydraulic limitations, reduced the structural and functional distinctions between dwarf and taller fringing tree forms of Rhizophora mangle .     

Transgenic poplars with reduced lignin show impaired xylem conductivity, growth efficiency and survival

We studied xylem anatomy and hydraulic architecture in 14 transgenic insertion events and a control line of hybrid poplar (Populus spp.) that varied in lignin content. Transgenic events had different levels of down-regulation of two genes encoding 4-coumarate:coenzyme A ligase (4CL). Two-year-old trees were characterized after growing either as free-standing trees in the field or as supported by stakes in a greenhouse. In free-standing trees, a 20 to 40% reduction in lignin content was associated with increased xylem vulnerability to embolism, shoot dieback and mortality. In staked trees, the decreased biomechanical demands on the xylem was associated with increases in the leaf area to sapwood area ratio and wood specific conductivity (k(s)), and with decreased leaf-specific conductivity (k(l)). These shifts in hydraulic architecture suggest that the bending stresses perceived during growth can affect traits important for xylem water transport. Severe 4CL-downregulation resulted in the patchy formation of discoloured, brown wood with irregular vessels in which water transport was strongly impeded. These severely 4CL-downregulated trees had significantly lower growth efficiency (biomass/leaf area). These results underscore the necessity of adequate lignification for mechanical support of the stem, water transport, tree growth and survival.     

Effects of defoliation caused by the leaf miner<Emphasis Type="Italic"> Cameraria ohridella</Emphasis> on wood production and efficiency in<Emphasis Type="Italic"> Aesculus hippocastanum</Emphasis> growing in north-eastern Italy

The leaf miner Cameraria ohridella causes premature defoliation of Aesculus hippocastanum trees. Repeated defoliation has been reported to cause decrease in radial growth of trees and a progressive decline due to reduced production and allocation of photosynthates. Our study represents an attempt to estimate the impact of C. ohridella on annual wood increments and the hydraulic properties of the wood as well as on the dry mass of seeds. Twenty-two adult horse chestnut trees were selected, four of which had been chemically treated to prevent attack (controls). All other trees were heavily infested. The ground cover (GC) of each tree, measured from monthly hemispherical photographs, revealed that infested trees were completely defoliated in September and the slope of the GC-to-measurement dates relationship (named GC decrease index) was positively related to the number of mines per leaf. Anatomical observations showed that infested trees produced more wood per year than controls through more false rings with wider xylem conduits and, hence, with higher conductive area and theoretical flow. In fact, the theoretical flow was positively related to the defoliation intensity. In contrast, the allocation of photosynthates to seeds was strongly reduced in infested trees with respect to controls (up to 50% less). The hypothesis was advanced that horse chestnut trees reacted to C. ohridella attacks by increasing the hydraulic efficiency of the wood, thus ameliorating the water and nutrient supply to leaves between the spring and mid-summer and, therefore, compensating, at least partly, the reduced leaf lifespan.     

Genetic determinism of anatomical and hydraulic traits within an apple progeny

The apple tree is known to have an isohydric behaviour, maintaining rather constant leaf water potential in soil with low water status and/or under high evaporative demand. However, little is known on the xylem water transport from roots to leaves from the two perspectives of efficiency and safety, and on its genetic variability. We analysed 16 traits related to hydraulic efficiency and safety, and anatomical traits in apple stems, and the relationships between them. Most variables were found heritable, and we investigated the determinism underlying their genetic control through a quantitative trait loci (QTL) analysis on 90 genotypes from the same progeny. Principal component analysis (PCA) revealed that all traits related to efficiency, whether hydraulic conductivity, vessel number and area or wood area, were included in the first PC, whereas the second PC included the safety variables, thus confirming the absence of trade-off between these two sets of traits. Our results demonstrated that clustered variables were characterized by common genomic regions. Together with previous results on the same progeny, our study substantiated that hydraulic efficiency traits co-localized with traits identified for tree growth and fruit production.