木本植物水力系统对干旱胁迫的响应机制

干旱导致树木死亡对生态系统功能和碳平衡有重大影响。植物水分运输系统失调是引发树木死亡的主要机制。然而, 树木对干旱胁迫响应的多维性和复杂性, 使人们对植物水分运输系统在极端干旱条件下的响应以及植物死亡机理的认识还不清楚。该文首先评述衡量植物抗旱性的指标, 着重介绍可以综合评价植物干旱抗性特征的新参数——气孔安全阈值(SSM)。SSM越高, 表明气孔和水力性状之间的协调性越强, 木质部栓塞的可能性越低, 水力策略越保守。然后, 阐述木本植物应对干旱胁迫的一般响应过程。之后, 分别综述植物不同器官(叶、茎和根)对干旱胁迫的响应机制。植物达到死亡临界阈值的概率和时间, 取决于相关生理和形态学特征的相互作用。最后, 介绍木本植物水力恢复机制, 并提出3个亟待开展的研究问题: (1)改进叶片水分运输(木质部和木质部外水力导度)的测量方法, 量化4种不同途径的叶肉水分运输的相对贡献; (2)量化叶片表皮通透性变化, 以便更好地理解植物水分利用策略; (3)深入研究树木水碳耦合机制, 将个体结构和生理特征与群落/景观格局和过程相关联, 以便更好地评估和监测干旱诱导树木死亡的风险。     

7年穿透雨减少对锐齿槲栎光合固碳及生物量碳储量的影响

气候变化背景下不断加剧的干旱事件对树木的生长及碳积累产生显著影响。然而,树木光合固碳能力及生物量碳储量对相对长期干旱的连续响应机制的研究仍然有限。选择70年生的天然锐齿槲栎(Quercus aliena var.acuteserrata)林,探究长期模拟穿透雨减少对锐齿槲栎光合固碳潜力和生物量碳储量的影响。研究结果表明,连续7年的穿透雨减少处理显著降低了锐齿槲栎的光合固碳能力,其叶片净光合速率(A)、最大羧化速率(V_cmax)、最大电子传递速率(J_max)、最大光化学效率(F_v/F_m)均明显降低,且穿透雨减少处理增强了A与气孔导度(g_s)、J_max、F_v/F_m之间的相关性。在适应长期干旱过程中,锐齿槲栎通过增加比叶面积(SLA)、叶片栅栏组织与海绵组织的比值、气孔密度等叶片形态及结构特性变化,降低冠层叶面积(LAI)指数和蒸腾水分散失及提高水分利用效率(WUE)缓解和适应干旱胁迫的不利影响。但是,长期穿透雨减少仍...     

Plant water-regulation strategies: Isohydric versus anisohydric behavior

Water is a vital resource for plant survival, growth and distribution, and it is of significance to explore mechanisms of plant water-relations regulation and responses to drought in ecophysiology and global change ecology. Plants adapt to different climates and soil water regimes and develop divergent water-regulation strategies involving a suite of related traits, of which two typical types are isohydric and anisohydric behaviors. It is critical to distinguish water-regulation strategies of plants and reveal the underlying mechanisms for plant breeding and vegetation restoration especially in xeric regions; and it is also important for developing more accurate vegetation dynamic models and predicting vegetation distribution under climate change scenarios. In this review, we first recalled the definitions of isohydric and anisohydric regulations and three quantitative classification methods that were established based on the relationships (1) between stomatal conductance and leaf water potential, (2) between stomatal conductance and vapor pressure deficit, (3) between predawn and midday leaf water potentials. We then compared the two water-regulation strategies in terms of hydraulics and carbon-economics traits. We synthesized the mechanisms of plant water-regulation and found that the interaction between hydraulic and chemical signals was the dominant factor controlling plant water-regulation behavior. Last, we proposed three promising aspects in this field: (1) to explore reliable and universal methods for classifying plant water-regulation strategies based on extensive investigation of the traits related with plant water-relations in various regions; (2) to explore relationships between plant water-regulation strategies and traits of hydraulics, morphology, structure, and function in order to provide reliable parameters for improving vegetation dynamic models; and (3) to deeply understand the processes of plant water-regulation at different spatial and temporal scales, and reveal mechanisms of plants’ responses and adaption to environmental stresses (especially drought).     

气候过渡带锐齿栎林土壤呼吸对降雨改变的响应

作为大气与陆地生态系统之间的第二大碳通量,土壤呼吸是评价陆地生态系统碳循环及碳汇能力的不确定性来源之一。降雨格局改变及其导致的土壤水分变化是调节土壤呼吸的重要驱动。气候过渡带的水热状况受全球降雨格局改变的影响更为明显,揭示该区域森林土壤呼吸对降雨改变的响应规律有助于改善碳循环模型的预测精度。然而,气候过渡区的土壤碳排放过程如何响应降雨格局改变尚不清楚。通过在亚热带-暖温带的过渡区(宝天曼)开展降雨改变实验,以阐明锐齿栎林土壤呼吸及其温度敏感性对降雨增加(50%)和减少(50%)的响应规律。结果表明,降雨增加显著提高土壤湿度(+8.92%)而不影响土壤温度。与对照相比,降雨增加导致土壤呼吸显著提高80.5%,其土壤呼吸的温度敏感性(4.07)显著高于对照样地(2.66)。增雨处理下的土壤呼吸与土壤湿度呈负相关。降雨减少则显著降低土壤湿度(-10.25%),并对土壤呼吸有促进趋势,然而,对土壤呼吸的温度敏感性(2.64)无显著影响。减雨处理下的土壤呼吸强度与土壤湿度呈正相关。这意味着在我国亚热带—暖温带过渡区,降雨增加或减少均对土壤呼吸有不同程度的刺激作用,进而很可能减弱该区域森林生态系统土壤的固碳潜力。     

白刺叶片气孔特征对人工模拟降雨的响应

以荒漠生态系统典型植物白刺(Nitraria tangutorum Bobr)叶片为研究对象,利用数码图像显微镜处理系统,研究了不同人工模拟增雨处理下的白刺叶片气孔密度及形态特征的变化情况。结果表明,荒漠植物固有特征决定了白刺叶片下表皮气孔密度大于上表皮,上表皮、下表皮气孔密度对增雨响应差异不显著(P0.05)。增雨处理上表皮、下表皮气孔密度与对照差异显著(P0.05)。相同增雨季节,50%处理下叶片气孔密度高于100%处理;不同增雨季节,气孔密度对生长季后期增雨响应更明显。白刺叶表皮气孔分布遵循"一细胞间隔(one cell spacing rule)"法则。增雨后叶片上表皮和下表皮气孔长度、宽度均有不同程度的增加,气孔形态特征对100%处理的响应较50%处理更为明显,且生长季后期增雨对叶片气孔形态特征的影响更大。     

水分和盐分胁迫对番茄植株生长和木质部水力特性的影响

木质部是植株体内水分传输的主要通路,其水力特性的变化会影响植株的水分关系和果实的水分积累。目前关于番茄植株木质部解剖结构和水力特性对水分和盐分胁迫的响应及其与植株生长和果实含水量之间的关系尚不明确。本研究通过日光温室番茄盆栽试验,设置3个处理:对照,土壤含水量(θ)为75%～95%田间持水量(FC),初始电导率(EC)为0.398 dS·m^-1;水分胁迫,开花前θ为75%～95% FC,开花后至成熟期θ为45%～65% FC,EC为0.398 dS·m^-1;盐分胁迫,θ为75%～95% FC,EC为1.680 dS·m^-1,研究了樱桃型番茄(红宝石)和中果型番茄(北番501)植株在水分和盐分胁迫下的植株生长、果实含水量以及木质部水力特性的变化。结果表明: 与对照相比,水分和盐分胁迫下茎秆横截面积和木质部导管直径分别减小了22.0%～40.7%和10.0%～18.3%,茎秆比导水率和桁架柄比导水率分别降低了8.8%～41.1%和12.9%～28.4%,抑制了植株生长,减少了地上部鲜重、果实大小、果实鲜重和含水量,且与樱桃型番茄相比,中果型番茄的降幅更大。此外,果实含水量分别与茎秆和桁架柄比导水率呈显著正相关。综上,番茄植株在水分和盐分胁迫下木质部水力特性指标减小,生长被抑制,果实鲜重显著降低,最终导致产量降低。其中,中果型番茄相较于樱桃型番茄对水分和盐分胁迫更敏感。     

昆虫对降雨和干旱的响应与适应

水分因子在昆虫的生长发育及其整个生活史中起着至关重要的作用。降雨作为改变环境水分的方式之一,其机械冲刷作用对昆虫具有直接的致死效应,并影响昆虫的生长发育、繁殖及其产卵和取食行为等;干旱作为降雨减少导致的极端环境形式,不仅对昆虫生理产生直接影响,而且还会通过影响寄主植物而间接作用于昆虫;同时,干旱还会改变同一寄主植物上昆虫之间的种间关系,导致群落多样性和稳定性的变化以及种群演替的发生。本文综述了气候变化背景下降雨和干旱对昆虫生长发育和繁殖的影响,并介绍了迁飞型昆虫、群居型昆虫、土壤害虫等对降雨和干旱的行为反应;此外,还详细介绍了昆虫对降雨和干旱的适应对策(包括对水分因子的行为适应对策、滞育和迁飞对策等),并建议利用环境水分(尤其是土壤水分)调控措施(如人工降雨和灌溉等)来防治农业害虫。     

模拟降雨减少对刺槐人工林土壤养分和微生物群落的影响

降雨是黄土高原地区土壤水分的最主要来源。为探明降雨减少对黄土高原半湿润区土壤养分和土壤微生物群落的影响,以刺槐人工林为研究对象,通过搭设透光遮雨板将部分穿透雨导流至样地外,实施了4年减少总降雨输入约47%的处理,测定了在不同降雨条件下林地的土壤养分含量、土壤微生物多样性及群落结构特征,分析了土壤养分和土壤微生物对降雨减少的响应。结果表明：减雨处理4年后(1)处理样地土壤有机碳、全氮、全磷含量均低于对照样地,全氮含量差异达到显著水平(P<0.05)。(2)对照样地生长季和非生长季土壤微生物多样性差异不显著,减雨处理样地非生长季的细菌均匀度显著低于处理样地生长季和对照样地非生长季(P<0.05);降雨减少对细菌和真菌的群落结构有显著影响(P<0.05),降雨减少显著降低了酸杆菌门(Acidobacteria)的相对丰度,提高了放线菌门(Actinobacteria)、绿弯菌门(Chloroflexi)、子囊菌门(Ascomycota)和被孢霉门(Mortierellomycota)的相对丰度(P<0.05)。(3)土壤有机碳、全氮含量与部分细菌相对丰度显著相关,土壤...     

伊犁河谷退化野果林中新疆野苹果茎的水力结构

为揭示伊犁河谷退化野果林中新疆野苹果植株个体的死亡机理,对比研究3种长势(Ⅰ级,20%枯枝率;Ⅱ级,40%~60%枯枝率;Ⅲ级,>80%枯枝率)新疆野苹果茎的水力结构特征的差异以及水分运输有效性和安全性及其影响因素。结果表明: 随野果林退化程度的增加,3种长势的新疆野苹果茎的边材比导水率和叶比导水率呈下降趋势,但差异不显著;傍晚时的栓塞程度、栓塞脆弱性呈显著增加的趋势,其中Ⅰ、Ⅱ和Ⅲ级长势个体的导水率损失50%时木质部水势分别为-1.87、-1.35和-0.53 MPa;黎明前和正午叶水势、水力安全边际均为Ⅰ级>Ⅱ级>Ⅲ级;与水力学相关的木质部解剖结构以及枝叶性状在3种长势个体之间存在显著差异。相关性分析表明,新疆野苹果茎木质部水分运输的有效性和安全性之间存在弱的权衡关系。新疆野苹果茎的水力结构变化伴随着新疆野果林的衰退过程。野果林的退化会加剧新疆野苹果木质部导管的栓塞化程度,降低其抵抗空穴化栓塞的能力,新疆野苹果面临水力失衡的威胁更大。     

锐齿栎幼苗生长、光合作用和非结构性碳对遮阴和模拟光斑的响应

近年发现宝天曼锐齿栎(Quercus aliena var.acuteserrata)成熟林难以更新,通过对锐齿栎幼苗不同遮阴处理下生长、非结构性碳(NSC)和光合作用进行研究,为理解宝天曼锐齿栎树种的更新困难和加强锐齿栎林的科学抚育提供理论依据。本研究通过搭建遮阴棚对盆栽锐齿栎幼苗设置遮阴处理,并用中午开放遮阴网的方法模拟光斑,设置了对照(自然光照)、遮阴80%中午开放1 h(T80-1)、一直遮阴80%(T80)、遮阴98%中午开放1 h(T98-1)、一直遮阴98%(T98)5处理,测定了遮阴后锐齿栎幼苗的干物质量、比叶面积、NSC浓度、存活率、光合光响应曲线参数、光合日变化、荧光参数等指标。结果表明:遮阴处理的幼苗干物质量和NSC浓度都显著低于对照,模拟光斑处理使幼苗NSC浓度显著高于一直遮阴处理的幼苗;遮阴严重的T98处理使幼苗出现了较高的死亡率;在高光强下,T80、T98-1、T98的净光合速率(P_n)因非气孔限制显著低于对照;且T98的光化学反应能力最低;遮阴后锐齿栎幼苗表现出一定的遮阴适应性,如光饱和点(LCP)和暗呼吸速率(R_d)降低,T98的P_n高于T98-1。本研究证明,在严重遮阴环境下锐齿栎幼苗易因难以维持碳的收支平衡而死亡,但光斑缓解了遮阴对幼苗碳储备的不利影响,对塑造幼苗在高光照条件下的光合能力也有显著作用。     

Water deficits and hydraulic limits to leaf water supply

Many aspects of plant water use -- particularly in response to soil drought -- may have as their basis the alteration of hydraulic conductance from soil to canopy. The regulation of plant water potential (Psi) by stomatal control and leaf area adjustment may be necessary to maximize water uptake on the one hand, while avoiding loss of hydraulic contact with the soil water on the other. Modelling the changes in hydraulic conductance with pressure gradients in the continuum allows the prediction of water use as a function of soil environment and plant architectural and xylem traits. Large differences in water use between species can be attributed in part to differences in their 'hydraulic equipment' that is presumably optimized for drawing water from a particular temporal and spatial niche in the soil environment. A number of studies have identified hydraulic limits as the cause of partial or complete foliar dieback in response to drought. The interactions between root:shoot ratio, rooting depth, xylem properties, and soil properties in influencing the limits to canopy water supply can be used to predict which combinations should optimize water use in a given circumstance. The hydraulic approach can improve our understanding of the coupling of canopy processes to soil environment, and the adaptive significance of stomatal behaviour.     

Size-dependent mortality in a Neotropical savanna tree: the role of height-related adjustments in hydraulic architecture and carbon allocation

Size-related changes in hydraulic architecture, carbon allocation and gas exchange of Sclerolobium paniculatum (Leguminosae), a dominant tree species in Neotropical savannas of central Brazil (Cerrado), were investigated to assess their potential role in the dieback of tall individuals. Trees greater than ∼6-m-tall exhibited more branch damage, larger numbers of dead individuals, higher wood density, greater leaf mass per area, lower leaf area to sapwood area ratio (LA/SA), lower stomatal conductance and lower net CO₂ assimilation than small trees. Stem-specific hydraulic conductivity decreased, while leaf-specific hydraulic conductivity remained nearly constant, with increasing tree size because of lower LA/SA in larger trees. Leaves were substantially more vulnerable to embolism than stems. Large trees had lower maximum leaf hydraulic conductance ( K _leaf) than small trees and all tree sizes exhibited lower K _leaf at midday than at dawn. These size-related adjustments in hydraulic architecture and carbon allocation apparently incurred a large physiological cost: large trees received a lower return in carbon gain from their investment in stem and leaf biomass compared with small trees. Additionally, large trees may experience more severe water deficits in dry years due to lower capacity for buffering the effects of hydraulic path-length and soil water deficits.     

Co‐occurring woody species have diverse hydraulic strategies and mortality rates during an extreme drought

From 2011 to 2013, Texas experienced its worst drought in recorded history. This event provided a unique natural experiment to assess species‐specific responses to extreme drought and mortality of four co‐occurring woody species: Quercus fusiformis, Diospyros texana, Prosopis glandulosa, and Juniperus ashei. We examined hypothesized mechanisms that could promote these species' diverse mortality patterns using postdrought measurements on surviving trees coupled to retrospective process modelling. The species exhibited a wide range of gas exchange responses, hydraulic strategies, and mortality rates. Multiple proposed indices of mortality mechanisms were inconsistent with the observed mortality patterns across species, including measures of the degree of iso/anisohydry, photosynthesis, carbohydrate depletion, and hydraulic safety margins. Large losses of spring and summer whole‐tree conductance (driven by belowground losses of conductance) and shallower rooting depths were associated with species that exhibited greater mortality. Based on this retrospective analysis, we suggest that species more vulnerable to drought were more likely to have succumbed to hydraulic failure belowground.     

模拟穿透雨减少对锐齿栎(Quercus aliena var. acuteserrata)树干液流密度的影响

降水格局变化是全球气候变化的重要特征之一,未来气候变化下,较为频繁和严峻的干旱将威胁地球中纬度部分地区的森林,但森林植被如何响应季节性干旱胁迫及其机制尚不清楚。北亚热带-暖温带过渡区分布着以锐齿栎(Quercus aliena var.acuteserrata)为优势树种的落叶阔叶林,研究其水分蒸腾代谢过程对干旱的响应是评估气候变化对过渡区天然落叶阔叶林生态系统水碳影响的关键科学问题。在典型的锐齿栎天然林中通过开展模拟穿透雨减少大型野外实验,采用Granier热扩散式探针技术监测锐齿栎树干液流密度的动态变化,研究了不同径级锐齿栎树干液流密度对模拟干旱的响应规律。结果表明:(1)穿透雨减少对树干液流密度的影响呈现季节变异。在7月份,林内穿透雨减少显著降低了锐齿栎的树干液流密度,但生长季后期的10月份林内穿透雨减少反而使锐齿栎树干液流密度显著升高。(2)不同径级的锐齿栎树干液流密度在生长季内对干旱有不同的响应,特别是小径级的树干液流密度与其他径级有较多的不同。小径级的锐齿栎树干液流密度在5、7月份表现为减雨样地显著小于对照样地,在9、10月份则表现为减雨样地显著大于对照样地。中径级的锐齿栎树干液流密度在5、10月份表现为减雨样地显著大于对照样地,在7月份则表现为减雨样地极显著小于对照样地。大径级的锐齿栎树干液流密度在6、7月份表现为减雨样地显著小于对照样地,在10月份则表现为减雨样地显著大于对照样地。     

栓皮栎根系解剖结构、水力特性及碳、氮含量研究

为阐明栓皮栎根系随径级的变化规律,探究其细根的合理划分标准。以1年生栓皮栎幼苗为研究对象,将其根系分为1、1～2、2～3、3～4 mm四个径级,分别制作石蜡切片观察解剖结构,比较木质部水力特性,测定碳氮含量及其比值,并采用主成分法对根系进行分类。结果表明:(1)随着径级增加,栓皮栎根系周皮、韧皮部和形成层组织厚度增加而占径比降低,木质部直径及其占径比均增加。(2)直径2 mm以上的栓皮栎根系木质部平均最大和最小导管直径、根比导水率和栓塞脆弱性指数增加显著;而导管密度显著下降,导管面积与木质部面积之比变化不显著。(3)直径2 mm以上栓皮栎根系碳含量表现出显著增加,随着径级增加,根系氮含量下降、碳氮比升高。(4)主成分分析表明,13项根系结构和元素含量指标降维后,前2个主分量方差贡献率达62%,PCA双序轴显示栓皮栎根系可划分为2 mm以下的吸收根群和2 mm以上的运输根群。综上认为,以2 mm作为栓皮栎细根划分的标准兼顾了形态和功能的特点,更具有准确性。     

胡杨木质部水分传导对盐胁迫的响应与适应北大核心CSCD

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象,系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明:(1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶,盐胁迫下根系生长和根尖数显著受到抑制,根木质部易于发生栓塞,导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定,轻度(0.05 mol·L–1 Na Cl)和中度(0.15 mol·L–1 Na Cl)盐胁迫下,胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率,维持植物正常生长;重度(0.30 mol·L–1 Na Cl)盐胁迫下,胡杨茎木质部导管输水有效性和安全性均明显降低,木质部导水率显著下降,并伴随叶片气孔导度的显著降低,从而严重抑制了胡杨的光合和生长。     

Limits to water transport in Juniperus osteosperma and Pinus edulis: implications for drought tolerance and regulation of transpiration

1. An air-injection method was used to study loss of water transport capacity caused by xylem cavitation in roots and branches of Pinus edulis (Colorado Pinyon) and Juniperus osteosperma (Utah Juniper). These two species characterize the Pinyon–Juniper communities of the high deserts of the western United States. Juniperus osteosperma can grow in drier sites than P. edulis and is considered the more drought tolerant.
2. Juniperus osteosperma was more resistant to xylem cavitation than P. edulis in both branches and roots. Within a species, branches were more resistant to cavitation than roots for P. edulis but no difference was seen between the two organs for J. osteosperma . There was also no difference between juveniles and adults in J. osteosperma ; this comparison was not made for P. edulis .
3. Tracheid diameter was positively correlated with xylem cavitation pressure across roots and stems of both species. This relation suggests a trade-off between xylem conductance and resistance to xylem cavitation in these species.
4. During summer drought, P. edulis maintained higher predawn xylem pressures and showed much greater stomatal restriction of transpiration, consistent with its greater vulnerability to cavitation, than J. osteosperma .
5. These results suggest that the relative drought tolerance of P. edulis and J. osteosperma results in part from difference in their vulnerability to xylem cavitation.     

Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.