Ecological applications of differences in the hydraulic efficiency of palms and broad-leaved trees

Key message

In tropical forests, co-occurring woody monocot and dicot species adapted different water use strategies highly depending on their investment in the hydraulic conduit properties.

Abstract

We studied the hydraulic efficiency of palms and broad-leaved tropical tree species from a moist tropical lowland forest in the Central Brazilian Amazon. Therefore, we harvested 34 trees and 10 açai palms and measured vessel size and frequency at diameter at breast height and additionally at the base of the crown shaft for the palms. Further, we assessed the active xylem area to estimate the hydraulic conductivity through Hagen Poiseuille’s adapted theoretical equation. Mean vessel diameter in dicot trees was 127.62 ± 49.22 μm with an average 9.09 ± 6.50 vessels per mm². Mean conduits sizes at the base (h = 0.10 m) of palm trees were larger with 288.20 ± 32.96 μm and less frequent with 1.40 ± 0.46 vessels per mm². Hydraulic conductivity was on average 3.31 ± 4.59 kg m^?1 s^?1 MPa^?1 for dicot trees. Mean hydraulic conductivity in açai palms was 20.45 ± 10.6 kg m^?1 s^?1 MPa^?1 at the base, and increased to 124.73 ± 55.2 kg m^?1 s^?1 MPa^?1 at the crown base. Hydraulic conductivity at the base of the crown was higher than in the base of the trunk due to the high density of vessels in a small cross-section in this height. Furthermore, we found a species-independent relationship between vessel diameter and frequency. We conclude that the differences found in the hydraulic efficiency give some evidence that palms have a lower occurrence of embolism and cavitation than trees, which is due to stiffer and stronger conduit pathways and efficient drought-avoiding strategies. The differences in hydraulic architecture between palms and trees imply different water use patterns thus varying niche differentiation, but this does not consequently need to be an excluding factor for coexistence in the same environment.

     

Xylem Hydraulic Characteristics, Water Relations and Wood Anatomy of the Resurrection PlantMyrothamnus flabellifoliusWelw.

Myrothamnus flabellifoliusWelw. is a desiccation-tolerant (‘resurrection’)plant with a woody stem. Xylem vessels are narrow (14 µmmean diameter) and perforation plates are reticulate. This leadsto specific and leaf specific hydraulic conductivities thatare amongst the lowest recorded for angiosperms (k_s0.87 kg m^-1MPa^-1s^-1;k_l3.28x10^-5kg m^-1MPa^-1s^-1, stem diameter 3 mm). Hydraulic conductivitiesdecrease with increasing pressure gradient. Transpiration ratesin well watered plants were moderate to low, generating xylemwater potentials of -1 to -2 MPa. Acoustic emissions indicatedextensive cavitation events that were initiated at xylem waterpotentials of -2 to -3 MPa. The desiccation-tolerant natureof the tissue permits this species to survive this interruptionof the water supply. On rewatering the roots pressures thatwere developed were low (2.4 kPa). However capillary forceswere demonstrated to be adequate to account for the refillingof xylem vessels and re-establishment of hydraulic continuityeven when water was under a tension of -8 kPa. During dehydrationand rehydration cycles stems showed considerable shrinking andswelling. Unusual knob-like structures of unknown chemical compositionwere observed on the outer surface of xylem vessels. These maybe related to the ability of the stem to withstand the mechanicalstresses associated with this shrinkage and swelling.Copyright1998 Annals of Botany Company cavitation, desiccation, hydraulic conductivity, refilling, resurrection plant, root pressure, xylem anatomy,Myrothamnus flabellifolius     

Embolism vulnerability of an evergreen tree

Leaf bearing stems ofCuratella americana L. were very susceptible to induced cavitation: embolisms began at a pressure of 0.5 MPa (15 %) and at 2.0 MPa most of the conductivity was lost (85 %). Nevertheless, in nature similar leaf specific conductivities, of about 90 x 10^-5 kg m^-2 s^-1 MPa^-1 during both wet and dry seasons indicated absence of drought induced embolisms. Leaf water potentials were neither very negative or considerably different between seasons but stomatal conductance decreased from 236 mmol m^-2 s^-1 measured during wet period to 100 mmol m^-2 s^-1 during drought season. Therefore, it was concluded thatCuratella had an accurate homeostatic balance of leaf water status to keep up xylem integrity. Acknowledgements: Financial support was provided by Decanato de Investigaciones - USB (S1-CB-811). Dr John Sperry (University of Utah) allowed me to learn how to use and to build the equipment used in this study. Mr. M. Edreida and Mr. T. Perez helped me in the field and in the laboratory, respectively. Dr D. HenrIquez corrected the English grammar.     

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest

Background and Aims

The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.

Methods

A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.

Key Results

It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D_h) and higher mass-based photosynthetic rate (A_m); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A_m.

Conclusions

The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

Contribution of vines to the evapotranspiration of a secondary forest in eastern Amazonia

The contribution of vines to the evapotranspiration (ET) of a secondary forest in eastern Amazonia was estimated based on field measurements of vine and tree transpiration, and seasonal changes in soil water content to 12 meters depth. Transpiration of vines and trees was measured with sapflow gauges placed around stems or branches. Total ET of the secondary forest was estimated as the sum of rainfall and reductions in soil moisture measured using Time Domain Reflectometry sensors installed in the walls of soil shafts. Our results suggest that vines transpire more than trees with stems of similar diameter, and with similar leaf crown exposure to sunlight. Trees experienced a smaller reduction in transpiration from the wet to the dry season than did vines. During the dry season, vines represented 8% (0.4 mm d^–1) of total secondary forest ET (5.4 mm d^–1), but they represented only 5.5% (0.5 m² ha^–1) of total secondary forest basal area (9.6 m² ha^–1). Considering that transpiration corresponds to 66–90% of forest ET, vines may contribute 9–12% to the transpiration of the forest. Hence, vine cutting, which is a commonly recommended management practice to favor the growth of tropical timber trees, may result in a proportionally larger reduction in evapotranspiration than in forest basal area.     

Physiological acclimation to drought stress in Solidago canadensis

Solidago canadensis is an invasive species from North America that is spreading across Europe, Australia and temperate Asia. We hypothesized that the species' wide ecological amplitude is also based on its potential in hydraulic acclimation, and analyzed hydraulic and anatomical properties along a transect with decreasing soil humidity. Stem hydraulic conductivity, vulnerability to drought‐induced embolism, stomatal closure during dehydration and xylem‐anatomical parameters were quantified at three sites. At the humid site, specific hydraulic conductivity of stems (1.0 ± 0.2 kg m^–1 MPa^–1 s^–1) was about twofold higher, and leaf‐specific conductivity about 1.5 times higher (3.1 ± 0.5 kg m^–1 MPa^–1 s^–1) than at the dry site. Water potential (Ψ) at 50% loss of conductivity was ?3.7 ± 0.1 MPa at the dry site and ?3.1 ± 0.2 MPa at the humid site (September). Vulnerability to drought‐induced embolism decreased along the transect and over the vegetation period. At drier sites, stomata started closing at lower Ψ while complete stomatal closure was reached at less negative Ψ (12% of maximum stomatal conductance: –2.5 ± 0.0 and ?3.0 ± 0.2 MPa at the dry and humid site). The safety margin between stomatal closure and 50% loss of conductivity was 1.2 and 0.2 MPa at the dry and humid sites. The observed variability indicated an efficient acclimation in hydraulic conductivity and safety: plants at dry sites exhibited lower specific hydraulic conductivity, higher embolism resistance and broader safety margins, signifying a trade‐off between the hydraulic safety and efficiency. The observed intraspecific plasticity in hydraulic and anatomical traits may help to explain the invasive potential of this species.     

鼎湖山南亚热带天然针阔叶混交林臭氧吸收特征

针阔叶混交林是我国南亚热带针叶林向地带性常绿阔叶林演替的中间林分类型,为我国南亚地区主要森林类型,发挥着重要的生态系统服务功能。基于树干液流技术和对臭氧浓度的连续监测,评价该森林类型的臭氧吸收特征和能力有着重要的环境生态学意义。对鼎湖山天然针阔叶混交林优势种马尾松(Pinus manssoniana)、锥栗(Castanopsis chinensis)、木荷(Schima superba)和华润楠(Machilus chinensis)在自然环境条件下的臭氧吸收能力进行了分析研究。结果表明:在日尺度上,4个优势树种的冠层气孔对臭氧导度(GO_3)和臭氧吸收通量(FO_3)均呈单峰型曲线,其最大值的时间在干季(10月至竖年3月)比湿季(4月至9月)滞后;季节尺度上,臭氧浓度在湿季达到最大值48.94 n L/L,湿季GO_3、FO_3和年臭氧吸收累积量(accumulative stomatal O_3flux,AFst)均显著高于干季(P 0.01),华润楠的臭氧吸收能力最强,在干季和湿季可分别达1.11 nmol m~(-2)s~(-1)和1.71nmol m~(-2)s~(-1)。随着水汽压亏缺(VPD)增大,优势种GO_3降低。光合有效辐射(PAR)超过1500 umol m~(-2)s~(-1)时,优势树种GO_3和FO_3呈下降趋势。针阔叶混交林的年臭氧吸收累积量超过了保护森林树木所采用的临界阈值,可认为鼎湖山针阔叶混交林受臭氧危害的潜在风险较高。     

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.     

The hydraulic architecture of balsam fir (Abies balsamea)

Leaf-specific conductivities (LSCs – hydraulic conductivity per dry weight of supplied leaves). Huber values (transverse sapwood area per dry weight of supplied leaves), specific conductivity (hydraulic conductivity per transverse sapwood area) and tracheid diameters were measured throughout the trunk and crown of 20-year-old trees of Abies balsamca (L.) Mill. Measured specific conductivity was proportional to the radius to the fourth power of tracheids. LSCs, which indicate the relative water availability to different plant parts, are much higher in the trunk than in first order branches, and lowest in second order branches. The structural basis for this "hydraulic hierarchy" lies both in Huber values and in tracheid diameters. For similar diameter stem segments, there was no statistically significant difference for trunks versus branches in specific conductivity. However, in old parts of the tree, trunks are wider than supported branches and producer wider tracheids resulting in greater specific conductivities than in branches. In vigorous trees with strong apical control, Huber values were 12.0 times greater in the trunk than in similar diameter branch segments. In slow-growing trees with weak apical control, Huber values were 2.2 times greater in the trunk versus similar branch segments.     

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements

Summary Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day^–1 tree^–1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m² of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s^–1 that is equivalent to 112 mmol m^–2 s^–1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m^–2 s^–1 at the top and 150 mmol m^–2 s^–1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was –27.76±0.27 (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.This paper is dedicated to Prof. Dr. O.L. Lange on the occasion of his 65th birthday     

广西弄岗五桠果叶木姜子群落木质藤本空间格局及其与树种间的关系

木质藤本是生物多样性的重要组成,木质藤本通过影响支持木进而影响群落的结构和功能,但在生物多样性丰富的北热带喀斯特森林中,木质藤本与支持木的关系鲜为人知。以喀斯特季节性雨林的五桠果叶木姜子（Litsea dilleniifolia）群落为研究对象,对木质藤本的密度、分布格局及其与主要树种的关系进行调查研究,分析木质藤本对树木的影响。结果显示：（1）五桠果叶木姜子群落内木质藤本平均密度为0.0913株/m²,木质藤本在0-20m空间尺度整体表现为聚集分布,且随着尺度增大,聚集强度逐渐减弱;不同径级木质藤本在不同尺度上的分布格局不同。（2）木质藤本对不同径级、不同种类、不同聚集强度的支持木选择表现以下体征：随着支持木径级增加,木质藤本攀附的比例和每木藤本数有增加趋势,且木质藤本胸径与支持木胸径呈极显著正相关;附藤率较高的支持木有紫葳科（Bignoniaceae）种类和东京桐（Deutzianthus tonkinensis）,单木附藤数量多的是南方紫金牛（Ardisia thyrsiflora）;物种的聚集强度与附藤率、附藤数量呈负相关。（3）木质藤本的密度与支持木死亡率关系不显著,而物种的附藤率与死亡率呈极显著负相关。以上结果表明,木质藤本密度在原生性喀斯特季节性雨林中并不高,且木质藤本对支持木具有选择性,但其对五桠果叶木姜子群落的死亡率并未产生显著影响。该研究可为喀斯特原生性季节性雨林的物种共存、极小植物种群保育提供理论依据,也可为石漠化区域的植被修复提供科学参考。     

Induction of rehydration and bud break by irrigation or rain in decidous trees of a tropical dry forest in Costa Rica

Summary Clusters of 2–4 bare, deciduous hardwood trees and woody vines in a dry upland forest in Costa Rica were surrounded by scaffolding and rehydration was induced during the dry season by irrigation of 9–50 m² plots with 200 mm water. The resulting changes in water status preceding bud break were monitored. Following irrigation, stem water potentials increased from < –4 MPa to about –1.5 MPa within 24 h and to > –0.3 MPa within 48 h. Rehydration of stem tissues by lateral transport, measured as an increase in electric conductivity, continued for 4–8 days. Terminal flower buds in Tabebuia ochracea began to expand 48 h after irrigation and trees were in full bloom 4 days later. In all experimental species, lateral vegetative buds began to expand 5–7 days after irrigation and leaves were fully expanded 2 weeks later. After the first rains of the rainy season (100 mm in 48 hr) all trees in the dry forest rehydrated and leaves emerged in synchrony slightly faster than after irrigation. In response to rain or irrigation drought-stressed tropical hardwood trees thus rehydrated at rates similar to those of desert succulents and their development resumed much faster than that of deciduous cold-temperate trees in spring.     

Abundance of climbing plants in a southern temperate rain forest: host tree characteristics or light availability?

Question: In a southern temperate rain forest, we addressed three questions: (1) Does the abundance of climbing plants increase with light availability? (2) Do host tree species differ in their susceptibility to vine infestation? (3) How does the relationship between host tree trunk diameter and relative abundance of vines vary with their climbing mechanism? Location: Two sites in the temperate evergreen rain forest of southern Chile: Puyehue (40°39′S, 72°09′W; 350 m a.s.l.) and Pastahue (42°22′S, 73°49′W; 285 m a.s.l.). Methods: We sampled vines in 60 25‐m² plots, with 20 plots in each of three light environments: mature forest, forest edges and canopy gaps. In each plot, for every tree ≥1.50‐m tall of any diameter we counted and identified all climbing plant individuals at a height of 1.30 m. We also counted, measured (trunk diameter at 1.30 m) and identified all these trees, and determined prevalence of vine infestation for each tree species. Results: Light availability in forest plots did not affect vine abundance when the number and size of host trees was taken into account. Overall, vine abundance increased with host tree trunk diameter. Tree species did not differ in the prevalence of vine infestation. The relative abundance of stem twiners and adhesive climbers decreased and increased with trunk diameter, respectively. The densities of stem twiners and adhesive climbers were negatively correlated across the forest. Conclusion: We provide further evidence that the pattern of vine abundance is independent of light availability in southern temperate rain forests, in contrast to results commonly reported for tropical rain forests. We also show that support suitability across the forest varies with the mechanism by which vines climb, probably due in part to biomechanical constraints and in part to vine interspecific competition, a virtually unexplored ecological factor.     

Through-flow of water in leaves of a submerged plant is influenced by the apical opening

Submerged aquatic higher plants maintain acropetal water transport to the young leaves in active growth to satisfy their demand for nutrients and hormones derived from the roots. We here present the first measurements of hydraulic properties for a submerged plant, the monocotyledon Sparganium emersum Rehman. The hydraulic conductance per unit length, K_h, was measured in leaf segments without the leaf tip and shown to be greater in old, fully developed leaves (1.5 · 10⁻¹⁰ · m⁴ · MPa⁻¹ · s⁻¹) than in young leaves (1.0 · 10⁻¹⁰ · m⁴ · MPa⁻¹ · s⁻¹). In leaves with intact leaf tips, however, K_h was significantly greater in the youngest leaves, which suggests that the leaf tip with the hydathode influences resistance and thus flow. Microscopy confirmed that the hydathodal area, which is an apical opening, undergoes structural changes with leaf age; a matrix of microorganisms develops in the older leaves and probably restricts water flow by clogging the hydathodes. The leaf specific conductivity expressing transport capacity relative to the leaf area supplied, of S. emersum (0.1 · 10⁻⁸ to 9 · 10⁻⁸ · m² MPa⁻¹· s⁻¹) was within the same range as for various species of terrestrial ferns, vines and trees. This finding does not support the traditional concept of functionally reduced vascular transport in Received: 15 July 1996 / Accepted: 30 November 1996     

Lateral hydraulic conductivity of early metaxylem vessels in Zea mays L. roots

The hydraulic conductivity of the lateral walls of early metaxylem vessels (Lp_x in m · s^–1 · MPa^–1) was measured in young, excised roots of maize using a root pressure probe. Values for this parameter were determined by comparing the root hydraulic conductivities before and after steam-ringing a short zone on each root. Killing of living tissue virtually canceled its hydraulic resistance. There were no suberin lamellae present in the endodermis of the roots used. The value of Lp_x ranged between 3 · 10^–7 and 35 · 10^–7 m · s^–1 · MPa^–1 and was larger than the hydraulic conductivity of the untreated root (Lp_r = 0.7 · 10^–7 to 4.0 · 10^–7 m · s^–1 · MPa^–1) by factor of 3 to 13. Assuming that all flow through the vessel walls was through the pit membranes, which occupied 14% of the total wall area, an upper limit of the hydraulic conductivity of this structure could be given(Lp_pm=21 · 10^–7 to 250 · 10^–7 m · s^–1 · MPa^–1). The specific hydraulic conductivity (Lp_cw) of the wall material of the pit membranes (again an upper limit) ranged from 0.3 · 10^–12 to 3.8 · 10^–12 m² · s^–1 · MPa^–1 and was lower than estimates given in the literature for plant cell walls. From the data, we conclude that the majority of the radial resistance to water movement in the root is contributed by living tissue. However, although the lateral walls of the vessels do not limit the rate of water flow in the intact system, they constitute 8–31% of the total resistance, a value which should not be ignored in a detailed analysis of water flow through roots.Abbreviatations and Symbols k_wr (T ₁ ^2/W ) rate constant (half-time) of water exchange across root (s^–1 or s, respectively) - Lp_cw specific hydraulic conductivity of wall material (m² · s^–1 · MPa^–1) - Lp_pm hydraulic conductivity of pit membranes (m · s ^–1 · MPa^–1) - Lp_r hydraulic conductivity of root (m · s^–1 · MPa^–1) - Lp_x lateralhydraulic conductivity of walls of root xylem (m · s ^–1 · MPa^–1) This research was supported by a grant from the Bilateral Exchange Program funded jointly by the Natural Sciences and Engineering Research Council of Canada and the Deutsche Forschungsgemeinschaft to C.A.P., and by a grant from the Deutsche Forschungsgemeinschaft, Sonderforschungsbereich 137, to E.S. The expert technical help of Mr. Burkhard Stumpf and the work of Ms. Martina Murrmann and Ms. Hilde Zimmermann in digitizing chart-recorder strips is gratefully acknowledged.     

Do vine species in neotropical forests see the forest or the trees?

Questions: Is the occurrence of vine species in neotropical rain forests primarily determined by properties of the forest (environmental factors), by properties of the trees (tree species or tree size) or are vines randomly distributed? Location: Maya Biosphere Reserve, Guatemala. Methods: In five 1‐ha plots that span variation from unlogged forest to forest impacted by recurrent human disturbance we recorded the presence of all climbing vine species on every tree. The presence of all free standing vine species and 11 environmental variables were recorded in 100‐m² subplots. The relationship of host tree diameter and host tree identity on single tree vine species richness was investigated by GLM modelling. Partial redundancy analyses were used to partition the variation in vine species composition on two sources: environmental factors and tree species identity. Results: Single tree vine richness increased with increasing host tree DBH and differed significantly among host species. For climbing vines, the ratio of variation in subplot presence explained by tree species and by environmental variables was ca. 4:1 (in the most disturbed logged plots slightly lower), for free standing vines this ratio varied from 1:2 in the most disturbed logged plots to 9:1 in reserve plots, while a ratio of ca. 1:1 was found for all plots analysed together. Conclusion: Different tree species have different probabilities of being infested by vines. Vines see both the forest and the trees; the environment is more important in earlier developmental stages, properties of individual trees become more important from the time vines start to climb.     

Fluxes of carbon, water and energy over Brazilian cerrado: an analysis using eddy covariance and stable isotopes

We present the energy and mass balance of cerrado sensu stricto (a Brazilian form of savanna), in which a mixture of shrubs, trees and grasses forms a vegetation with a leaf area index of 1·0 in the wet season and 0·4 in the dry season. In the wet season the available energy was equally dissipated between sensible heat and evaporation, but in the dry season at high irradiance the sensible heat greatly exceeded evaporation. Ecosystem surface conductance g_s in the wet season rose abruptly to 0·3 mol m^?2 s^?1 and fell gradually as the day progressed. Much of the total variation in g_s was associated with variation in the leaf-to-air vapour pressure deficit of water and the solar irradiance. In the dry season the maximal g_s values were only 0·1 mol m^?2 s^?1. Maximal net ecosystem fluxes of CO₂ in the wet and dry season were –10 and –15 μmol CO₂ m^?2 s^?1, respectively (sign convention: negative denotes fluxes from atmosphere to vegetation). The canopy was well coupled to the atmosphere, and there was rarely a significant build-up of respiratory CO₂ during the night. For observations in the wet season, the vegetation was a carbon dioxide sink, of maximal strength 0·15 mol m^?2 d^?1. However, it was a source of carbon dioxide for a brief period at the height of the dry season. Leaf carbon isotopic composition showed all the grasses except for one species to be C₄, and all the palms and woody plants to be C₃. The CO₂ coming from the soil had an isotopic composition that suggested 40% of it was of C₄ origin.     

The effects of cambial age and position within the stem on specific conductivity in Douglas-fir (Pseudotsuga menziesii) sapwood

Specific conductivity (k_s, m²s^-1MPa^-1) describes the permeability of xylem and is determined by all aspects of xylem anatomy that create resistance to the flow of water. Here we test the hypothesis that k_s is a function of radial and vertical position within the stem, rather than solely a function of cambial age (ring number from the pith), by measuring k_s on samples excised from 35-year-old Douglas-fir [Pseudotsuga menziesii var. menziesii (Mirb.) Franco] trees at six heights and two or three radial positions. Sapwood k_s decreased from the cambium to the heartwood boundary, and the difference between outer and inner sapwood increased with height in the tree. Beneath the live crown, inner sapwood had 80-90% the k_s of outer sapwood, but only 55% just 10 m higher in the stem (about 10 nodes down from the tree top). Outer sapwood k_s peaked near the base of the crown and declined toward both the base and top of the stem. These patterns can be explained by two superimposed effects: the effect of cambial age on the dimensions of tracheids as they are produced, and the effect of xylem aging, which may include accumulation of emboli and aspiration of bordered pits. Tracheid lumen diameter and earlywood and latewood density and width, all factors known to vary with cambial age, were measured on different trees of the same age and from the same stand. Lumen diameter increased with cambial age, whereas the proportion of latewood and growth ring density increased after an initial decrease in the first 5 years. Our results suggest that the effect of cambial age on xylem anatomy is not sufficient to explain variation in k_s. Instead, physical position (both vertical and radial) in the stem and cambial age must be considered as determinants of conductivity.     

Stem hydraulic capacitance decreases with drought stress: implications for modelling tree hydraulics in the Mediterranean oak Quercus ilex

Hydraulic modelling is a primary tool to predict plant performance in future drier scenarios. However, as most tree models are validated under non‐stress conditions, they may fail when water becomes limiting. To simulate tree hydraulic functioning under moist and dry conditions, the current version of a water flow and storage mechanistic model was further developed by implementing equations that describe variation in xylem hydraulic resistance (R_X) and stem hydraulic capacitance (C_S) with predawn water potential (Ψ_PD). The model was applied in a Mediterranean forest experiencing intense summer drought, where six Quercus ilex trees were instrumented to monitor stem diameter variations and sap flow, concurrently with measurements of predawn and midday leaf water potential. Best model performance was observed when C_S was allowed to decrease with decreasing Ψ_PD. Hydraulic capacitance decreased from 62 to 25 kg m^?3 MPa^?1 across the growing season. In parallel, tree transpiration decreased to a greater extent than the capacitive water release and the contribution of stored water to transpiration increased from 2.0 to 5.1%. Our results demonstrate the importance of stored water and seasonality in C_S for tree hydraulic functioning, and they suggest that C_S should be considered to predict the drought response of trees with models.     

Climate-related trends in sapwood biophysical properties in two conifers: avoidance of hydraulic dysfunction through coordinated adjustments in xylem efficiency, safety and capacitance

In the Pacific north‐west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas‐fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas‐fir. Radial conductivity was higher in west‐side populations of both species, but axial conductivity was higher in the east‐side populations and in Douglas‐fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west‐side populations. Sapwood capacitance was similar between species, but was about twice as great in east‐side populations (580 kg m^?3 MPa^?1) as in west‐side populations (274 kg m^?3 MPa^?1). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity.