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1.
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.  相似文献   

2.
Functional relationships between wood density and measures of xylem hydraulic safety and efficiency are ambiguous, especially in wet tropical forests. In this meta-analysis, we move beyond wood density per se and identify relationships between xylem allocated to fibers, parenchyma, and vessels and measures of hydraulic safety and efficiency. We analyzed published data of xylem traits, hydraulic properties and measures of drought resistance from neotropical tree species retrieved from 346 sources. We found that xylem volume allocation to fiber walls increases embolism resistance, but at the expense of specific conductivity and sapwood capacitance. Xylem volume investment in fiber lumen increases capacitance, while investment in axial parenchyma is associated with higher specific conductivity. Dominant tree taxa from wet forests prioritize xylem allocation to axial parenchyma at the expense of fiber walls, resulting in a low embolism resistance for a given wood density and a high vulnerability to drought-induced mortality. We conclude that strong trade-offs between xylem allocation to fiber walls, fiber lumen, and axial parenchyma drive drought resistance in neotropical trees. Moreover, the benefits of xylem allocation to axial parenchyma in wet tropical trees might not outweigh the consequential low embolism resistance under more frequent and severe droughts in a changing climate.  相似文献   

3.
Defoliation by herbivores can reduce carbon assimilation, change plant water relations, and even shift the biotic structure of plant communities. In this study, we took advantage of a long-term deer exclosure experiment to examine the consequences of persistent deer herbivory on plant water relations and the xylem structure–function relationships in Ceanothus rigidus, a maritime chaparral shrub in coastal California. Browsed plants had thicker stems with many intertwined short distal twigs, and significantly higher sapwood-to-leaf area ratios than their non-browsed counterparts. Leaf area-specific hydraulic conductivity was similar in both browsed and non-browsed plants, but xylem area-specific conductivity was significantly lower in the browsed plants. Vessel diameters were equivalent in both plant groups, but the number of vessels on a transverse area basis was nearly 40 % lower in the browsed plants, accounting for their lower transport efficiency. Mid-day in situ water potentials and losses of hydraulic conductivity due to embolism were similar in both groups of plants but stomatal conductance was higher in the browsed shrubs in the early part of the growing season. We discuss our findings in the context of whole-plant ecophysiology, and explore the consequences of herbivory on hormonal signals, wood anatomy, and xylem function.  相似文献   

4.
Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (Ks) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of Kh and leaf specific conductivity (KL) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.  相似文献   

5.
Mangrove trees dominate coastal vegetation in tropical regions, but are completely replaced by herbaceous salt marshes at latitudes above 32 degrees N and 40 degrees S. Because water deficit can increase damage caused by freezing, we hypothesized that mangroves, which experience large deficits as a result of saline substrates, would suffer freeze-induced xylem failure. Vulnerability to freeze-induced xylem embolism was examined in the most poleward mangrove species in North America, in an area where freezing is rare but severe, and in Australia, in an area where freezing is frequent but mild. Percentage loss in hydraulic conductivity was measured following manipulations of xylem tension; xylem sap ion concentration was determined using X-ray microanalysis. Species with wider vessels suffered 60-100% loss of hydraulic conductivity after freezing and thawing under tension, while species with narrower vessels lost as little as 13-40% of conductivity. These results indicate that freeze-induced embolism may play a role in setting the latitudinal limits of distribution in mangroves, either through massive embolism following freezing, or through constraints on water transport as a result of vessel size.  相似文献   

6.
The exotic temperate liana (woody vine) Celastrus orbiculatus has become a weed in Michigan, occurring in many of the same habitats as the native liana Vitis riparia. However, C. orbiculatus frequently develops into extensive monospecific infestations, while V. riparia does not. Freezing-induced embolism may be responsible for limiting liana distribution. Root pressure has been observed in numerous tropical lianas and temperate species of Vitis and has been implicated as vital to the recovery of xylem function in wide vessels following winter freezes. For both of these co-occurring lianas we investigated root pressure and water conductance as possible explanatory factors for their differential spread. According to our hypothesis, C. orbiculatus should have produced greater or more frequent root pressures than V. riparia. However, the reverse proved true, indicating that root pressure is not a prerequisite for weedy proliferation of C. orbiculatus. Additionally, the seasonal patterns of specific conductivity of stem xylem indicate that each species responds differently to environmental constraints. Vitis riparia establishes conductivity early in the growing season, before the leaves emerge, using root pressure to reverse embolism, but loses conductivity with the first freeze in early autumn. Celastrus orbiculatus is slow to establish conductivity, depending on new wood production, but leafs out sooner than V. riparia and maintains green leaves after the first freeze. Vulnerability curves of xylem to cavitation caused by water stress for the two species indicate that they respond similarly to dehydration. These results indicate that root pressures are not responsible for the invasive success of C. orbiculatus and suggest that other factors must be key to its prolific invasion.  相似文献   

7.
We present a practical for university-level students aimed at measuring and comparing xylem hydraulic properties of co-existing plant species. After sampling branches of several woody species in the field, their main hydraulic properties were measured using a simple set-up. Hydraulic conductivity (Kh ) was calculated as the ratio between water flow through a plant segment and the pressure gradient driving the flow. The percent reduction in conductivity due to xylem embolism (i.e. air-filled conduits) was estimated by comparing Kh before and after flushing the measure segments to remove all native embolism. Raw hydraulic conductivity was standardised by cross-sectional wood area or supported leaf area to obtain more meaningful measures of conducting capacity. The results showed differences among study species, particularly between conifers and angiosperms. These differences are briefly discussed in terms of wood anatomy and the general biology of the species. Overall the practical provides a good opportunity for students to appreciate the main aspects of xylem water transport and the constraints it imposes on plant water relations.  相似文献   

8.
Background and Aims When the orientation of the stems of conifers departs from the vertical as a result of environmental influences, conifers form compression wood that results in restoration of verticality. It is well known that intercellular spaces are formed between tracheids in compression wood, but the function of these spaces remains to be clarified. In the present study, we evaluated the impact of these spaces in artificially induced compression wood in Chamaecyparis obtusa seedlings.Methods We monitored the presence or absence of liquid in the intercellular spaces of differentiating xylem by cryo-scanning electron microscopy. In addition, we analysed the relationship between intercellular spaces and the hydraulic properties of the compression wood.Key Results Initially, we detected small intercellular spaces with liquid in regions in which the profiles of tracheids were not rounded in transverse surfaces, indicating that the intercellular spaces had originally contained no gases. In the regions where tracheids had formed secondary walls, we found that some intercellular spaces had lost their liquid. Cavitation of intercellular spaces would affect hydraulic conductivity as a consequence of the induction of cavitation in neighbouring tracheids.Conclusions Our observations suggest that cavitation of intercellular spaces is the critical event that affects not only the functions of intercellular spaces but also the hydraulic properties of compression wood.  相似文献   

9.
The occurrence of root pressure, the vulnerability of xylemvessels to drought-induced cavitation, and the seasonal changesin hydraulic conductivity due to embolism were studied in theculms of Rhipidocladum racemiflorum (Steud.) McClure, a tropicalvine-like bamboo from central Panama. Positive hydrostatic potentialsup to 120 kPa occurred only during the wet season when the transpirationrate of the plant was low, i.e. at night or during rain events.Although the xylem vessels were large and efficient for conductingwater, they were highly resistant to cavitation. Xylem waterpotentials lower than –4.5 MPa were required to induce50% loss of hydraulic conductivity in culms. The minimum waterpotential reached –3.75 MPa at the end of the 1993 dryseason, so loss of hydraulic conductivity due to embolism remained<10%. The species is adapted to drier habitats both by wayof a low vulnerability to xylem cavitation and by root pressuresin the wet season that could refill vessels that became embolizedduring a severe dry season. Key words: Rhipidocladum racemiflorum, root pressure, cavitation, embolism, water relations  相似文献   

10.

Background and Aims

The hydraulic architecture and water relations of fruits and leaves of Capsicum frutescens were measured before and during the fruiting phase in order to estimate the eventual impact of xylem cavitation and embolism on the hydraulic isolation of fruits and leaves before maturation/abscission.

Methods

Measurements were performed at three different growth stages: (1) actively growing plants with some flowers before anthesis (GS1), (2) plants with about 50 % fully expanded leaves and immature fruits (GS2) and (3) plants with mature fruits and senescing basal leaves (GS3). Leaf conductance to water vapour as well as leaf and fruit water potential were measured. Hydraulic measurements were made using both the high-pressure flow meter (HPFM) and the vacuum chamber (VC) technique.

Key Results

The hydraulic architecture of hot pepper plants during the fruiting phase was clearly addressed to favour water supply to growing fruits. Hydraulic measurements revealed that leaves of GS1 plants as well as leaves and fruit peduncles of GS2 plants were free from significant xylem embolism. Substantial increases in leaf petiole and fruit peduncle resistivity were recorded in GS3 plants irrespective of the hydraulic technique used. The higher fraction of resistivity measured using the VC technique compared with the HPFM technique was apparently due to conduit embolism.

Conclusions

The present study is the first to look at the hydraulics of leaves and fruits during growth and maturation through direct, simultaneous measurements of water status and xylem efficiency of both plant regions at different hours of the day.  相似文献   

11.
Kocacinar F  Sage RF 《Oecologia》2004,139(2):214-223
Xylem structure and function is proposed to reflect an evolutionary balance between demands for efficient movement of water to the leaf canopy and resistance to cavitation during high xylem tension. Water use efficiency (WUE) affects this balance by altering the water cost of photosynthesis. Therefore species of greater WUE, such as C4 plants, should have altered xylem properties. To evaluate this hypothesis, we assessed the hydraulic and anatomical properties of 19 C3 and C4 woody species from arid regions of the American west and central Asia. Specific conductivity of stem xylem (Ks ) was 16%–98% lower in the C4 than C3 shrubs from the American west. In the Asian species, the C3 Nitraria schoberi had similar and Halimodendron halodendron higher Ks values compared with three C4 species. Leaf specific conductivity (KL ; hydraulic conductivity per leaf area) was 60%–98% lower in the C4 than C3 species, demonstrating that the presence of the C4 pathway alters the relationship between leaf area and the ability of the xylem to transport water. C4 species produced similar or smaller vessels than the C3 shrubs except in Calligonum, and most C4 shrubs exhibited higher wood densities than the C3 species. Together, smaller conduit size and higher wood density indicate that in most cases, the C4 shrubs exploited higher WUE by altering xylem structure to enhance safety from cavitation. In a minority of cases, the C4 shrubs maintained similar xylem properties but enhanced the canopy area per branch. By establishing a link between C4 photosynthesis and xylem structure, this study indicates that other phenomena that affect WUE, such as atmospheric CO2 variation, may also affect the evolution of wood structure and function.  相似文献   

12.
 以同处于干旱区的塔里木河下游(铁干里克)和黑河下游(乌兰图格)断面为研究区, 比较了荒漠河岸林主要建群种胡杨(Populus euphratica)、柽柳(Tamarix spp.)、疏叶骆驼刺(Alhagi sparsifolia)和花花柴(Karelinia caspia)在长期遭受不同干旱胁迫下的根、枝条木质部导水力和栓塞化程度的变化特征, 并分析了木质部导水对干旱胁迫的响应及适应策略。结果表明: 1) 黑河下游荒漠河岸林植物的导水能力显著高于塔里木河下游, 其中柽柳、胡杨、疏叶骆驼刺和花花柴根木质部的初始比导率(Ks0)分别高11.97、6.74、7.10和3.73倍, 枝条的Ks0分别高9.48、3.65、2.07和1.88倍, 地下水埋深导致的干旱胁迫程度不同是诱发荒漠植物导水能力差异的根本原因; 2)柽柳耐干旱能力最强, 适应范围较宽, 而花花柴、疏叶骆驼刺的耐旱性相对较弱, 适生范围较窄, 这可能与植物的根系分布有关; 3)干旱胁迫较轻时, 枝条木质部是荒漠河岸林植物水分传输的主要阻力部位, 干旱胁迫严重时, 根木质部是限制植株水流的最大阻碍部位; 4)荒漠河岸林植物主要通过调节枝条木质部的水流阻力来适应干旱胁迫, 且其适应策略与干旱胁迫程度有关, 干旱胁迫轻时, 植物通过限制枝条木质部水流来协调整株植物的均匀生长; 干旱胁迫严重时, 植物通过牺牲劣势枝条、增强优势枝条水流来提高植株整体生存的机会。  相似文献   

13.

Background and Aims

Despite the importance of vessels in angiosperm roots for plant water transport, there is little research on the microanatomy of woody plant roots. Vessels in roots can be interconnected networks or nearly solitary, with few vessel–vessel connections. Species with few connections are common in arid habitats, presumably to isolate embolisms. In this study, measurements were made of root vessel pit sizes, vessel air-seeding pressures, pit membrane thicknesses and the degree of vessel interconnectedness in deep (approx. 20 m) and shallow (<10 cm) roots of two co-occurring species, Sideroxylon lanuginosum and Quercus fusiformis.

Methods

Scanning electron microscopy was used to image pit dimensions and to measure the distance between connected vessels. The number of connected vessels in larger samples was determined by using high-resolution computed tomography and three-dimensional (3-D) image analysis. Individual vessel air-seeding pressures were measured using a microcapillary method. The thickness of pit membranes was measured using transmission electron microscopy.

Key Results

Vessel pit size varied across both species and rooting depths. Deep Q. fusiformis roots had the largest pits overall (>500 µm) and more large pits than either shallow Q. fusiformis roots or S. lanuginosum roots. Vessel air-seeding pressures were approximately four times greater in Q. fusiformis than in S. lanuginosum and 1·3–1·9 times greater in shallow roots than in deep roots. Sideroxylon lanuginosum had 34–44 % of its vessels interconnected, whereas Q. fusiformis only had 1–6 % of its vessels connected. Vessel air-seeding pressures were unrelated to pit membrane thickness but showed a positive relationship with vessel interconnectedness.

Conclusions

These data support the hypothesis that species with more vessel–vessel integration are often less resistant to embolism than species with isolated vessels. This study also highlights the usefulness of tomography for vessel network analysis and the important role of 3-D xylem organization in plant hydraulic function.  相似文献   

14.
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.  相似文献   

15.
Catastrophic hydraulic failure will likely be an important mechanism contributing to large‐scale tree dieback caused by increased frequency and intensity of droughts under global climate change. To compare the susceptibility of 22 temperate deciduous tree and shrub species to hydraulic failure during a record drought in the southeastern USA, we quantified leaf desiccation, native embolism, wood density, stomatal conductance and predawn and midday leaf water potential at four sites with varying drought intensities. At the two driest sites, there was widespread leaf wilting and desiccation, and most species exhibited predawn leaf water potentials of ≤3 MPa and >60% loss of xylem conductivity in branches. Although species with high wood density were more resistant to cavitation, they had higher levels of native embolism and greater canopy dieback than species with low wood density. This unexpected result can be explained by the failure of species with dense wood to avert a decline in water potential to dangerous levels during the drought. Leaf water potential was negatively correlated with wood density, and the relationship was strongest under conditions of severe water deficit. Species with low wood density avoided catastrophic embolism by relying on an avoidance strategy that involves partial drought deciduousness, higher sensitivity of stomata to leaf water potential and perhaps greater rooting depth. These species therefore maintained water potential at levels that ensured a greater margin of safety against embolism. These differences among species may mediate rapid shifts in species composition of temperate forests if droughts intensify due to climate change.  相似文献   

16.
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.  相似文献   

17.
Background and Aims The efficiency and safety functions of xylem hydraulics are strongly dependent on the pits that connect the xylem vessels. However, little is known about their biochemical composition and thus about their hydraulic properties. In this study, the distribution of the epitopes of different wall components (cellulose, hemicelluloses, pectins and lignins) was analysed in intervessel pits of hybrid poplar (Populus tremula × alba).Methods Immunogold labelling with transmission electron microscopy was carried out with a set of antibodies raised against different epitopes for each wall polysaccharide type and for lignins. Analyses were performed on both immature and mature vessels. The effect of sap ionic strength on xylem conductance was also tested.Key Results In mature vessels, the pit membrane (PM) was composed of crystalline cellulose and lignins. None of the hemicellulose epitopes were found in the PM. Pectin epitopes in mature vessels were highly concentrated in the annulus, a restricted area of the PM, whereas they were initially found in the whole PM in immature vessels. The pit border also showed a specific labelling pattern, with higher cellulose labelling compared with the secondary wall of the vessel. Ion-mediated variation of 24 % was found for hydraulic conductance.Conclusions Cellulose microfibrils, lignins and annulus-restricted pectins have different physicochemical properties (rigidity, hydrophobicity, porosity) that have different effects on the hydraulic functions of the PM, and these influence both the hydraulic efficiency and vulnerability to cavitation of the pits, including ion-mediated control of hydraulic conductance. Impregnation of the cellulose microfibrils of the PM with lignins, which have low wettability, may result in lower cavitation pressure for a given pore size and thus help to explain the vulnerability of this species to cavitation.  相似文献   

18.
极端干旱环境下的胡杨木质部水力特征   总被引:2,自引:0,他引:2  
胡杨作为我国西北干旱区重要的乔木树种,研究其木质部水力特征对了解此树种适应极端干旱环境的生物学背景具有较重要的意义。本研究以塔里木河下游的胡杨成株和2年生胡杨幼苗为研究材料,对其木质部最大导水能力(ks(max))和自然栓塞程度(PLC)等木质部水力特征及其水力特征有关的木质部导管(或管饱)数量特征进行研究。结果表明,成株胡杨多年生枝条和侧根(2≤d<5 mm)木质部自然栓塞程度均较高,PLC均值高于50%,其中多年生枝条栓塞程度具有一定的日变化规律,清晨的PLC均值(58%)小于正午的(67%);河道边上成株胡杨侧根的均ks(max)和PLC均值都小于距河道200 m处的。随着土壤干旱程度的加剧,幼苗胡杨侧根的自然栓塞程度随之增加,而叶片气孔导度随之降低,土壤含水率与侧根自然栓塞程度,叶片气孔导度之间分别存在显著负相关关系(R =-0.9、R =-0.811)。在统一直径范围内(2≤d<5 mm),成株胡杨侧根均导管直径(dmean)和水力直径均大于(d95%、dh)胡杨幼苗,而导管密度胡杨幼苗高于成株胡杨;胡杨侧根木质部最大导水能力与均导管直径、水力直径之间具有显著正相关关系(R>0.9).  相似文献   

19.
Salix psammophila and Caragana korshinskii are two common shrubs in the southern Mu Us Desert, China. Their hydraulic strategies for adapting to this harsh, dry desert environment are not yet clear. This study examined the hydraulic transport efficiency, vulnerability to cavitation, and daily embolism refilling in the leaves and stems of these two shrubs during the dry season. In order to gain insight into water use strategies of whole plants, other related traits were also considered, including daily changes in stomatal conductance, leaf mass per area, leaf pressure–volume parameters, wood density and the Huber value. The leaves and stems of S. psammophila had greater hydraulic efficiency, but were more vulnerable to drought-induced hydraulic dysfunction than C. korshinskii. The difference between leaf and stem water potential at 50 % loss of conductivity was 0.12 MPa for S. psammophila and 0.81 MPa for C. korshinskii. Midday stomatal conductance decreased by 74 % compared to that at 8:30 in S. psammophila, whereas no change occurred in C. korshinskii. Daily embolism and refilling occurred in the stems of S. psammophila and leaves of C. korshinskii. These results suggest that a stricter stomatal regulation, daily embolism repair in stems, and a higher stem water capacitance could be partially compensating for the greater susceptibility to xylem embolism in S. psammophila, whereas higher leaf elastic modulus, greater embolism resistance in stems, larger difference between leaf and stem hydraulic safety, and drought-induced leaf shedding in C. korshinskii were largely responsible for its more extensive distribution in arid and desert steppes.  相似文献   

20.
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