盐碱胁迫下枸杞和柽柳的水力学特性和碳代谢

盐碱胁迫是全球范围内重要的非生物胁迫形式之一,但目前对植物水力学特性和碳代谢应对盐碱胁迫响应的研究还不多。本研究以耐盐碱植物枸杞（Lycium chinense Miller）和柽柳（Tamarix chinensis Lour.）为对象,测定不同盐碱程度下两种植物的枝条水势和导水损失率（PLC）、叶片光合作用和气孔导度、不同部位的非结构性碳（NSC）浓度以及植株生长情况。结果显示,重度盐碱胁迫显著降低了两种植物凌晨和正午水势、光合速率和枝条PLC,重度胁迫下柽柳的光合速率、气孔导度和枝条PLC下降程度均大于枸杞,重度盐碱胁迫下枸杞不同部位的NSC浓度均显著降低,但柽柳的地上部分NSC浓度显著增加,根部NSC浓度显著减低。两种植物有不同的应对盐碱胁迫策略,枸杞有较强的气孔调节能力,对水力结构的维持有利,但会限制碳摄取,柽柳气孔调节能力弱,水力结构易受影响,但对碳平衡维持有利。     

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。     

最优气孔行为理论和气孔导度模拟

气孔调节功能是陆地生态系统碳-水耦合过程中最重要的环节。与即时的气孔导度测量相比, 气孔导度斜率能有效地反映气孔导度对CO₂浓度、饱和水汽压亏缺和光合作用的敏感性, 包含了环境因子对光合作用和临界水分利用效率等的综合影响, 为研究全球变化下陆地生态系统碳-水耦合关系提供了一个简明且综合的框架。气孔导度模型从经验模型、半经验模型发展到机理模型, 经过很多学者的改进, 但是模型参数的生物学意义和变化规律还不明确。鉴于气孔导度斜率方面研究的重要性和研究的不足, 为了加强对气孔导度调节规律的认识, 并减少气孔导度模拟的不确定性, 该文主要综述了长期以来国内外关于最优气孔行为理论和气孔导度模拟方面的研究成果, 其中包括广泛使用的气孔导度模型及参数意义, 讨论影响气孔导度斜率的主要因素以及气孔导度机理模型的应用, 并对最优气孔行为理论和气孔导度模拟方面的研究做了简单展望。     

春玉米持续干旱过程中常用气孔导度模型的比较研究

气候变化背景下,干旱频发导致的土壤水分变化将影响气孔导度模型的适用性,进而影响生态系统碳-氮-水循环模拟的准确性。基于春玉米持续干旱田间模拟试验资料,比较了常用气孔导度模型在干旱条件下的模拟效果,评价了土壤水分响应函数对气孔导度模型效果的影响,并探讨了气孔导度模型的适用土壤水分范围。结果表明,在持续干旱过程中,模型模拟效果表现为BBL模型最优,其次是USO模型和BWB模型,Jarvis模型最差;引入土壤水分响应函数,提高了BWB模型和USO模型的模拟效果,而降低了Jarvis模型和BBL模型模拟效果,模型模拟效果表现为USO修正模型最优,其次是BBL修正模型和BWB修正模型,Jarvis修正模型最差。在持续干旱过程中,Jarvis模型和BWB模型的剩余气孔导度较大,而BBL模型和USO模型的剩余气孔导度相对较小,表明BBL模型和USO模型在干旱条件下具有一定的稳定性。基于95%置信区间判断表明:Jarvis模型、BBL模型和USO模型在土壤相对湿度范围为33%—83%条件下适用,而BWB模型的适用土壤相对湿度范围为33%—76%,引入水分响应函数后可在试验条件下适用。研究结果可为干旱条件下选取合适的气孔导度模型以准确模拟陆地生态系统碳循环和水循环提供依据,并为改善农业水资源的有效使用和评估提供支撑。     

叶片水力性状研究进展

叶片水力性状表征了叶片为适应外在环境而形成的水分传输方面的生存策略。叶片水力性状会限制整个植株的水分传输,并影响植物的气体交换及其对干旱的响应,因此关于叶片水力性状的研究已成为植物水分关系领域的研究热点之一。本文概括了叶片水力性状的基本指标(包括叶片整体水力导度(Kleaf)、叶片木质部水力导度(Kxylem)、叶片木质部外水力导度(Kout-xylem)等)和叶片水力导度的5种主要测量方法;总结了叶脉网络结构和环境因素对叶片水力性状的影响、叶片水力性状与叶片功能指标(气孔导度、叶片水势、叶片最大光合速率)的匹配与权衡关系,以及叶片水力性状与植物抗旱性关系的最新研究进展;对今后叶片水力性状的研究提出了两点建议:1)将叶片水力性状与气体交换和叶解剖结构等相结合,构建叶片碳-水耦合模型,揭示叶片应对外界环境变化而采取的生态策略,以及植物的水-碳投资机理;2)开展植株各部分(根-茎-叶)间水分传输的交互作用研究,筛选出水力系统高效安全的物种。     

干旱胁迫对植物气孔特性影响研究进展

综述干旱胁迫对植物叶片气孔特性的影响,从气孔大小、气孔密度、气孔导度等方面概括植物气孔对干旱胁迫的响应,分析植物气孔特性与其抗旱性的关系,并对气孔特性的研究方向进行展望。     

Changes in hydraulic traits of nine vascular plants from different evolutionary lineages

维管植物从蕨类植物、裸子植物到被子植物的演化过程中, 生理结构和功能都被认为更趋向于适应干旱的环境。但是关于3类植物是否提升了水分传导和水分利用效率的研究目前多基于叶脉和气孔的形态结构推断, 而对水分传导速率的直接测量则集中于被子植物内部。因此, 同时测定3类植物叶片和枝条尺度的水力性状, 可以研究维管植物水力性状的演化趋势和协同演化关系。该研究测定了中国科学院华南植物园内的蕨类植物(芒萁(Dicranopteris pedata)、华南毛蕨(Cyclosorus parasiticus)、乌毛蕨(Blechnum orientale)), 裸子植物(罗汉松(Podocarpus macrophyllus)、竹柏(Podocarpus nagi)、落羽杉(Taxodium distichum))和被子植物(海南木莲(Manglietia fordiana var. hainanensis)、东京油楠(Sindora tonkinensis)、羊蹄甲(Bauhinia purpurea))共9种植物的水力结构与功能的相关性状。发现9种植物从原始到更演化物种的枝条比导水率(K_S)、叶片比导水率(K_L)和叶片水力导度(K_leaf)依次升高; 叶片蒸腾速率(E)和潜在水分利用效率(WUE_i)升高趋势不显著; 气孔面积指数(SPI)和木材密度(WD)在3个类群间无明显变化趋势。水力功能性状在枝条(K_S)和叶片(K_L、K_leaf和E)尺度显著相关, 但和两个结构性状并无显著相关性。对显著相关的性状进行系统发育独立差分析发现相关性依然存在, 表明枝条导水和叶片导水、叶片失水的协同性独立于系统发育而存在。该文以9种维管植物为例, 发现演化过程中水分传导相关的功能性状有显著提升, 且导水和失水的性状存在着协同演化关系。上述维管植物水力性状演化趋势的结论需要对更多物种、更精细的水力结构性状的进行测定验证。     

九种维管植物水力性状的演化趋势

维管植物从蕨类植物、裸子植物到被子植物的演化过程中, 生理结构和功能都被认为更趋向于适应干旱的环境。但是关于3类植物是否提升了水分传导和水分利用效率的研究目前多基于叶脉和气孔的形态结构推断, 而对水分传导速率的直接测量则集中于被子植物内部。因此, 同时测定3类植物叶片和枝条尺度的水力性状, 可以研究维管植物水力性状的演化趋势和协同演化关系。该研究测定了中国科学院华南植物园内的蕨类植物(芒萁(Dicranopteris pedata)、华南毛蕨(Cyclosorus parasiticus)、乌毛蕨(Blechnum orientale)), 裸子植物(罗汉松(Podocarpus macrophyllus)、竹柏(Podocarpus nagi)、落羽杉(Taxodium distichum))和被子植物(海南木莲(Manglietia fordiana var. hainanensis)、东京油楠(Sindora tonkinensis)、羊蹄甲(Bauhinia purpurea))共9种植物的水力结构与功能的相关性状。发现9种植物从原始到更演化物种的枝条比导水率(K_S)、叶片比导水率(K_L)和叶片水力导度(K_leaf)依次升高; 叶片蒸腾速率(E)和潜在水分利用效率(WUE_i)升高趋势不显著; 气孔面积指数(SPI)和木材密度(WD)在3个类群间无明显变化趋势。水力功能性状在枝条(K_S)和叶片(K_L、K_leaf和E)尺度显著相关, 但和两个结构性状并无显著相关性。对显著相关的性状进行系统发育独立差分析发现相关性依然存在, 表明枝条导水和叶片导水、叶片失水的协同性独立于系统发育而存在。该文以9种维管植物为例, 发现演化过程中水分传导相关的功能性状有显著提升, 且导水和失水的性状存在着协同演化关系。上述维管植物水力性状演化趋势的结论需要对更多物种、更精细的水力结构性状的进行测定验证。     

半干旱地区3种植物叶片水平的抗旱耐旱特性分析——两个气孔导度模型的应用和比较

在对半干旱区3种植物进行生理生态特性测定的基础上,应用两种气孔导度模型进行参数的非线性拟合,BBL模型平均可以解释77.6%的结果,Gao模型平均可以解释59.3%的结果。但Gao模型作为一个机理性的模型,其参数具有明确的物理意义。模型的行为和敏感性分析结果说明,用BBL计算的气孔导度一般大于Gao模型。BBL模型对于干旱胁迫下的土壤水分亏缺没有响应, 因而不适合用作干旱半干旱区的植物生理生态学分析和生态系统模拟。而Gao模型可以描述在各种水分条件下植物气孔导度的响应。Gao模型的结果说明,与油松 (Pinus tabulaeformis) 和中间锦鸡儿 (Caragana intermedia) 比较,小叶杨 (Populus simonii) 具有最小的抗旱和耐旱能力,油松具有最好的叶片水平的耐旱和抗旱特性,但其气孔导度对土壤水分的不敏感意味着在干旱条件下维持光合作用的同时,也可能会导致过多的水分损失。中间锦鸡儿具有很强的耐旱性,且其气孔导度对土壤水分的变化敏感,二者相结合,中间锦鸡儿可以在土壤水分条件较好的情况下,维持较大的气孔导度以满足光合作用的需要,但在土壤水分胁迫严重的时候能迅速降低气孔导度以保持土壤水分。     

植物气孔气态失水与SPAC系统液态供水的相互调节作用研究进展

讨论了植物气孔气态失水与SPAC系统液态供水相互作用研究领域的一些重要现象和行为.当植物水力信号和化学信号共同作用促进气孔对叶水势的调节时,植物对叶水势的调节表现为等水行为.气孔对环境湿度变化响应的反馈机制可用来解释土壤干旱条件下气孔和光合的午休现象,以及气孔导度和水流导度之间的相关关系;而气孔对环境湿度变化响应的前馈机制,则可用来解释气孔导度对大气 叶片间水汽饱和差的滞后反应.植物最大限度地利用木质部传输水分的策略,要求气孔快速响应以避免木质部过度气穴化和短时间内将气穴逆转的相应机制.     

Stomatal characteristics of ferns and angiosperms and their responses to changing light intensity at different habitats

气孔是植物与大气环境进行气体交换的重要通道, 在调控植物碳水平衡方面发挥着重要作用。为探讨生境和植物类型对气孔形态特征的影响以及气孔对光强变化的响应格局在不同植物间和不同生境条件下的变异, 选取开阔生境和林下生境的5种蕨类植物和4种被子植物, 测定了它们的气孔形态特征和气孔导度对光强变化的响应。此外, 还收集了8篇文献中开阔和林下生境的45种蕨类植物和70种被子植物的气孔密度和气孔长度数据, 以增大样本量从而更好地探讨不同生境条件下蕨类和被子植物气孔密度及长度的变异格局, 并通过分析生境和植物类型对气孔形态特征的影响来推测生境和植物类型对气孔响应行为的可能影响。实验结果表明, 与林下植物相比, 开阔环境下的植物气孔密度更大, 气孔长度更小, 气孔对光强降低的响应更敏感; 但植物类型对气孔形态特征的影响以及对气孔响应光强的敏感程度的影响均不显著。对文献数据的分析表明, 生境和植物类型对气孔形态特征均有显著影响。考虑到气孔响应快慢与气孔形态特征密切相关, 与蕨类植物相比, 被子植物小而密的气孔可能为其更快地响应环境变化提供了基础。研究表明生境和植物类型对气孔响应行为均有显著影响。     

不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应

气孔是植物与大气环境进行气体交换的重要通道, 在调控植物碳水平衡方面发挥着重要作用。为探讨生境和植物类型对气孔形态特征的影响以及气孔对光强变化的响应格局在不同植物间和不同生境条件下的变异, 选取开阔生境和林下生境的5种蕨类植物和4种被子植物, 测定了它们的气孔形态特征和气孔导度对光强变化的响应。此外, 还收集了8篇文献中开阔和林下生境的45种蕨类植物和70种被子植物的气孔密度和气孔长度数据, 以增大样本量从而更好地探讨不同生境条件下蕨类和被子植物气孔密度及长度的变异格局, 并通过分析生境和植物类型对气孔形态特征的影响来推测生境和植物类型对气孔响应行为的可能影响。实验结果表明, 与林下植物相比, 开阔环境下的植物气孔密度更大, 气孔长度更小, 气孔对光强降低的响应更敏感; 但植物类型对气孔形态特征的影响以及对气孔响应光强的敏感程度的影响均不显著。对文献数据的分析表明, 生境和植物类型对气孔形态特征均有显著影响。考虑到气孔响应快慢与气孔形态特征密切相关, 与蕨类植物相比, 被子植物小而密的气孔可能为其更快地响应环境变化提供了基础。研究表明生境和植物类型对气孔响应行为均有显著影响。     

Adjustments of water use efficiency by stomatal regulation during drought and recovery in the drought-adapted Vitis hybrid Richter-110 (V. berlandieri x V. rupestris)

The hybrid Richter-110 (Vitis berlandieri x Vitis rupestris) (R-110) has the reputation of being a genotype strongly adapted to drought. A study was performed with plants of R-110 subjected to water withholding followed by re-watering. The goal was to analyze how stomatal conductance (g(s)) is regulated with respect to different physiological variables under water stress and recovery, as well as how water stress affects adjustments of water use efficiency (WUE) at the leaf level. Water stress induced a substantial stomatal closure and an increase in WUE, which persisted many days after re-watering. The g(s) during water stress was mainly related to the content of ABA in the xylem and partly related to plant hydraulic conductivity but not to leaf water potential. By contrast, low g(s) during re-watering did not correlate with ABA contents and was only related to a sustained decreased hydraulic conductivity. In addition to a complex physiological regulation of stomatal closure, g(s) and rate of transpiration (E) were strongly affected by leaf-to-air vapor pressure deficit (VPD) in a way dependent of the treatment. Interestingly, E increased with increasing VPD in control plants, but decreased with increasing VPD in severely stressed plants. All together, the fine stomatal regulation in R-110 resulted in very high WUE at the leaf level. This genotype is revealed to be very interesting for further studies on the physiological mechanisms leading to regulation of stomatal responsiveness and WUE in response to drought.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.     

干旱对陆地生态系统生产力的影响

该文综述了干旱对陆地生态系统生产力的影响,分析了其影响机制,并总结了植被对干旱的响应与适应及其机理机制。干旱通过抑制光合作用来降低陆地生态系统总初级生产力,干旱还可以降低生态系统的自养呼吸和异养呼吸。同时干旱还可以通过影响其它干扰形式来间接影响陆地生态系统生产力,如增加火干扰的发生频率和强度,增加植物的死亡率,增加病虫害的发生等。在生态系统水平上干旱可以降低碳固定,减弱碳汇功能,甚至把生态系统从碳汇改变成碳源。目前生态系统水平上的干旱影响研究主要通过两种方法实现,一种是模型模拟,另一种就是大型模拟实验。作为陆地生态系统生产力的实现者,在干旱胁迫条件下,植物也会采取积极的适应策略以减弱干旱对生态系统生产力的影响,其适应策略主要分以下3种:在一些周期性发生干旱的地区,植物会调整生长期以避开干旱或通过休眠来减弱干旱所造成的伤害;还有一些植物会通过调节体内的代谢过程,改变一些生理特性来抵御干旱;而长期生活在干旱条件下的植物则通过进化来改变了自身的生理生化代谢过程,形成耐旱机制。目前,植物对干旱响应的分子学机制,以及生态系统水平上对干旱的响应和适应仍然是薄弱的领域,也必然成为未来研究的重点。     

Wide vessels sustain marginal transpiration flux and do not optimize inefficient gas exchange activity under impaired hydraulic control and salinity

Plants optimize water use and carbon assimilation via transient regulation of stomata resistance and by limiting hydraulic conductivity in a long-term response of xylem anatomy. We postulated that without effective hydraulic regulation plants would permanently restrain water loss and photosynthetic productivity under salt stress conditions. We compared wild-type tomatoes to a transgenic type (TT) with impaired stomatal control. Gas exchange activity, biomass, starch content, leaf area and root traits, mineral composition and main stems xylem anatomy and hydraulic conductivity were analyzed in plants exposed to salinities of 1 and 4 dS m⁻¹ over 60 days. As the xylem cannot easily readjust to different environmental conditions, shifts in its anatomy and the permanent effect on plant hydraulic conductivity kept transpiration at lower levels under unstressed conditions and maintained it under salt-stress, while sustaining higher but inefficient assimilation rates, leading to starch accumulation and decreased plant biomass, leaf and root area and root length. Narrow conduits in unstressed TT plants were related to permanent restrain of hydraulic conductivity and plant transpiration. Under salinity, TT plants followed the atmospheric water demand, sustained similar transpiration rate from unstressed to salt-stressed conditions and possibly maintained hydraulic integrity, due to likely impaired hydraulic regulation, wider conduits and higher hydraulic conductivity. The accumulation of salts and starch in the TT plants was a strong evidence of salinity tolerance via osmotic regulation, also thought to help to maintain the assimilation rates and transpiration flux under salinity, although it was not translated into higher growth.     

Rapid hydraulic recovery in Eucalyptus pauciflora after drought: linkages between stem hydraulics and leaf gas exchange

In woody plants, photosynthetic capacity is closely linked to rates at which the plant hydraulic system can supply water to the leaf surface. Drought‐induced embolism can cause sharp declines in xylem hydraulic conductivity that coincide with stomatal closure and reduced photosynthesis. Recovery of photosynthetic capacity after drought is dependent on restored xylem function, although few data exist to elucidate this coordination. We examined the dynamics of leaf gas exchange and xylem function in Eucalyptus pauciflora seedlings exposed to a cycle of severe water stress and recovery after re‐watering. Stomatal closure and leaf turgor loss occurred at water potentials that delayed the extensive spread of embolism through the stem xylem. Stem hydraulic conductance recovered to control levels within 6 h after re‐watering despite a severe drought treatment, suggesting an active mechanism embolism repair. However, stomatal conductance did not recover after 10 d of re‐watering, effecting tighter control of transpiration post drought. The dynamics of recovery suggest that a combination of hydraulic and non‐hydraulic factors influenced stomatal behaviour post drought.     

Hydraulic properties of fern sporophytes: Consequences for ecological and evolutionary diversification

? Premise of the study: Ferns are an important component of both tropical and temperate forests; yet, our understanding of the water relations of their sporophyte generation is limited. Indeed, to date there has been no large scale survey that attempts to clarify how ferns fit into current ideas of plant water relations. This study examines several tropical ferns with the goal of understanding how these characters vary between species from various habitats and across life forms ? Methods: We measured stipe hydraulic conductivity, water potential, and vulnerability to cavitation along with photosynthetic variables and leaf allometry of 21 species from 14 genera to identify physiological trait assemblages across taxa. ? Key results: Epiphytic ferns have significantly lower hydraulic conductivity and a vascular system more resistant to cavitation (i.e., higher P(50) values). They reached lower mid-day water potentials and produced leaves with reduced stipe lengths and reduced laminar area relative to terrestrial species. Xylem specific hydraulic conductivity (K(S)) was correlated with the mean hydraulic diameter of tracheids in terrestrial species, but not in epiphytes. There was no evidence of safety-efficiency trade-offs in any group. ? Conclusions: When compared across life forms, our data shed light on physiological mechanisms that may have allowed for terrestrial ferns to move into the epiphytic habit. When compared across a diverse assemblage of terrestrial plants, we find that resistance to water flow in fern stipes is significantly higher than that recorded from the stems of seed plants.     

Stomatal dynamics are limited by leaf hydraulics in ferns and conifers: results from simultaneous measurements of liquid and vapour fluxes in leaves

Stomatal responsiveness to vapour pressure deficit (VPD) results in continuous regulation of daytime gas‐exchange directly influencing leaf water status and carbon gain. Current models can reasonably predict steady‐state stomatal conductance (g_s) to changes in VPD but the g_s dynamics between steady‐states are poorly known. Here, we used a diverse sample of conifers and ferns to show that leaf hydraulic architecture, in particular leaf capacitance, has a major role in determining the g_s response time to perturbations in VPD. By using simultaneous measurements of liquid and vapour fluxes into and out of leaves, the in situ fluctuations in leaf water balance were calculated and appeared to be closely tracked by changes in g_s thus supporting a passive model of stomatal control. Indeed, good agreement was found between observed and predicted g_s when using a hydropassive model based on hydraulic traits. We contend that a simple passive hydraulic control of stomata in response to changes in leaf water status provides for efficient stomatal responses to VPD in ferns and conifers, leading to closure rates as fast or faster than those seen in most angiosperms.