Xylem Cross-sectional Area Research Articles

Xylem cell size regulation is a key adaptive response to water deficit in Eucalyptus grandis.

Future climatic scenarios forecast increasingly frequent droughts that will pose substantial consequences on tree mortality. In light of this, drought-tolerant eucalypts have been propagated; however, the severity of these conditions will invoke adaptive responses, impacting the commercially valuable wood properties. To determine what mechanisms govern the wood anatomical adaptive response, highly controlled drought experiments were conducted in Eucalyptus grandis W. Hill ex Maiden, with the tree physiology and transcriptome closely monitored. In response to water deficit, E. grandis displays an isohydric stomatal response to conserve water and enable stem growth to continue, albeit at a reduced rate. Maintaining gaseous exchange is likely a critical short-term response that drives the formation of hydraulically safer xylem. For instance, the development of significantly smaller fibers and vessels was found to increase cellular density, thereby promoting drought tolerance through improved functional redundancy, as well as implosion and cavitation resistance. The transcriptome was explored to identify the molecular mechanisms responsible for controlling xylem cell size during prolonged water deficit. Downregulation of genes associated with cell wall remodeling and the biosynthesis of cellulose, hemicellulose and pectin appeared to coincide with a reduction in cellular enlargement during drought. Furthermore, transcript levels of NAC and MYB transcription factors, vital for cell wall component biosynthesis, were reduced, while those linked to lignification increased. The upregulation of EgCAD and various peroxidases under water deficit did not correlate with an increased lignin composition. However, with the elevated cellular density, a higher lignin content per xylem cross-sectional area was observed, potentially enhancing hydraulic safety. These results support the requirement for higher density, drought-adapted wood as a long-term adaptive response in E. grandis, which is largely influenced by the isohydric stomatal response coupled with cellular expansion-related molecular processes.

Normalization criteria determine the interpretation of nitrogen effects on the root hydraulics of pine seedlings.

Plant hydraulics is key for plant survival and growth because it is linked to gas exchange and drought resistance. Although the environment influences plant hydraulics, there is no clear consensus on the effect of nitrogen (N) supply, which may be, in part, due to different hydraulic conductance normalization criteria and studied species. The objective of this study was to compare the variation of root hydraulic properties using several normalization criteria in four pine species in response to three contrasting N fertilization regimes. We studied four closely related, yet ecologically distinct species: Pinus nigra J.F. Arnold, Pinus pinaster Ait., Pinus pinea L. and Pinus halepensis Mill. Root hydraulic conductance (Kh) was measured with a high-pressure flow meter, and values were normalized by total leaf area (leaf specific conductance, Kl), xylem cross-section area (xylem specific conductance, Ks), total root area (root specific conductance, Kr) and the area of fine roots (fine root specific conductance, Kfr). Controlling for organ size differences allowed comparison of the hydraulic efficiency of roots to supply or absorb water among fertilization treatments and species. The effect of N on the root hydraulic efficiency depended on the normalization criteria. Increasing N availability reduced Kl and Ks, but increased Kh, Kr and especially Kfr. The positive effect of N on Kr and Kfr was positively related to seedling relative growth rate and was also consistent with published results at the interspecific level, whereby plant hydraulics is positively linked to photosynthesis and transpiration rate and fast growth. In contrast, normalization by leaf area and xylem cross-sectional area (Kl and Ks) reflected opposite responses to Kr and Kfr. This indicates that the normalization criteria determine the interpretation of the effect of N on plant hydraulics, which can limit species and treatment comparisons.

Costs and benefits of gas inside wood and its relationship with anatomical traits: a contrast between trees and lianas.

Gas inside wood plays an important role in plant functioning, but there has been no study examining the adaptive nature of gas inside wood across plants differing in biomechanical demands. Using a comparative approach, we measured gas volumetric content, xylem's anatomical traits and wood density of 15 tree and 16 liana species, to test whether gas content varies between these plant types strongly differing in their biomechanical demands. We asked (i) whether trees and lianas differ in gas content and (ii) how anatomical traits and wood density are related to gas content. Lianas had significantly less gas content in their branches compared with tree species. In tree species, gas content scaled positively with fiber, vessel and xylem cross-sectional area and fiber and vessel diameter, and negatively with dry-mass density. When pooling trees and lianas together, fiber cross-sectional area was the strongest predictor of gas content, with higher xylem cross-sectional area of fiber associated with higher gas content. In addition, we showed, through a simple analytical model, that gas inside wood increases the minimum branch diameter needed to prevent rupture, and this effect was stronger on trees compared with lianas. Our results support the view that gas inside wood plays an important role in the evolution of biomechanical functioning in different plant forms. Gas inside wood may also play an important role in physiological activities such as water transport, storage, photosynthesis and respiration, but it is still unknown whether these roles are or are not secondary to the mechanical support.

Increasing air humidity influences hydraulic efficiency but not functional vulnerability of xylem in hybrid aspen

Climate models predict greater increases in the frequency than in the amount of precipitation and a consequent rise in atmospheric humidity at high latitudes by the end of the century. We investigated the responses of hydraulic and relevant anatomical traits of xylem to elevated relative humidity of air on a 1-yr-old coppice of hybrid aspen (Populus×wettsteinii) growing in the experimental stand at the Free Air Humidity Manipulation site in Eastern Estonia. The hydraulic conductivity of stems was measured with a high pressure flow meter; artificial cavitation in the stem segments was induced by the air injection method. Specific conductivity of xylem decreased from 4.42 in the control to 3.94kgm−1s−1MPa−1 in the humidification treatment, while the trend was well correlated with increasing wood density. Humidified trees exhibited smaller leaf area at the same xylem cross-sectional area, resulting in 34% higher average Huber values compared to the control. Control and humidity-treated trees differed by neither native embolism level nor susceptibility to dehydration-induced cavitation. Increasing atmospheric humidity reduces the hydraulic efficiency of hybrid aspen trees expressed on a xylem area basis and causes substantial changes in resource allocation between photosynthetic and water transport tissues. This climate trend does not influence stem vulnerability to cavitation.

Weather effects of stalk necrosis in Vitis

A 5-step physiological explanation is offered for correlations found by THEILER and MULLER (1986) between aspects of the weather at flowering and the incidence of stalk necrosis in grape. Four of the steps are well supported in the literature and are discussed briefly. The other step proposes that the development of xylem in the peduncle is stimulated by floral evapotranspiration. This postulate is tested with measurements of xylem cross-sectional area in clusters taken from vineyards located in diverse regions of New Zealand for which meteorological data were available. Statistical analysis indicates a clear distinction between material from the climatic extremes and a significant (P = 0.05) correlation between calculated values of evapotranspiration during flowering and peduncular xylem area. The beginning of a physiological explanation for the seasonal effects on the incidence of stalk necrosis in grape has stimulated studies which may allow the selection of non-sensitive varieties.

Elevated air humidity affects hydraulic traits and tree size but not biomass allocation in young silver birches (Betula pendula).

As changes in air temperature, precipitation, and air humidity are expected in the coming decades, studies on the impact of these environmental shifts on plant growth and functioning are of major importance. Greatly understudied aspects of climate change include consequences of increasing air humidity on forest ecosystems, predicted for high latitudes. The main objective of this study was to find a link between hydraulic acclimation and shifts in trees’ resource allocation in silver birch (Betula pendula Roth) in response to elevated air relative humidity (RH). A second question was whether the changes in hydraulic architecture depend on tree size. Two years of application of increased RH decreased the biomass accumulation in birch saplings, but the biomass partitioning among aboveground parts (leaves, branches, and stems) remained unaffected. Increased stem Huber values (xylem cross-sectional area to leaf area ratio) observed in trees under elevated RH did not entail changes in the ratio of non-photosynthetic to photosynthetic tissues. The reduction of stem–wood density is attributable to diminished mechanical load imposed on the stem, since humidified trees had relatively shorter crowns. Growing under higher RH caused hydraulic conductance of the root system (KR) to increase, while KR (expressed per unit leaf area) decreased and leaf hydraulic conductance increased with tree size. Saplings of silver birch acclimate to increasing air humidity by adjusting plant morphology (live crown length, slenderness, specific leaf area, and fine-root traits) and wood density rather than biomass distribution among aboveground organs. The treatment had a significant effect on several hydraulic properties of the trees, while the shifts were largely associated with changes in tree size but not in biomass allocation.

Phenotypic and developmental plasticity of xylem in hybrid poplar saplings subjected to experimental drought, nitrogen fertilization, and shading.

Variation in xylem structure and function has been extensively studied across different species with a wide taxonomic, geographical, and ecological coverage. In contrast, our understanding of how xylem of a single species can adjust to different growing condition remains limited. Here phenotypic and developmental plasticity in xylem traits of hybrid poplar (Populus trichocarpa×deltoides) was studied. Clonally propagated saplings were grown under experimental drought, nitrogen fertilization, and shade for >30 d. Xylem hydraulic and anatomical traits were subsequently examined in stem segments taken from two different vertical positions along the plant’s main axis. The experimental treatments affected growth and development and induced changes in xylem phenotype. Across all treatments, the amount of leaf area supported by stem segments (AL) scaled linearly with stem native hydraulic conductivity (K native), suggesting that the area of assimilating leaves is constrained by the xylem transport capacity. In turn, K native was mainly driven by the size of xylem cross-sectional area (AX). Moreover, the structural and functional properties of xylem varied significantly. Vulnerability to cavitation, measured as the xylem pressure inducing 50% loss of conductivity (P50), ranged from –1.71MPa to –0.15MPa in saplings subjected to drought and nitrogen fertilization, respectively. Across all treatments and stem segment positions, P50 was tightly correlated with wood density. In contrast, no relationship between P50 and xylem-specific conductivity (K S) was observed. The results of this study enhance our knowledge of plant hydraulic acclimation and provide insights into common trade-offs that exist in xylem structure and function.

Size-dependent variability of leaf and shoot hydraulic conductance in silver birch

Variation in leaf and shoot hydraulic conductance was examined on detached shoots of silver birch (Betula pendula Roth), cut from the lower third (shade leaves) and upper third of the crown (sun leaves) of large trees growing in a natural temperate forest stand. Hydraulic conductances of whole shoots (K S), leaf blades (K lb), petioles (K P) and branches (i.e. leafless stem; K B) were determined by water perfusion using a high-pressure flow meter in quasi-steady state mode. The shoots were exposed to irradiance of photosynthetic photon flux density of 200–250 μmol m−2 s−1, using different light sources. K lb depended significantly (P < 0.001) on light quality, canopy position and leaf blade area (A L). K lb increased from crown base to tree top, in parallel with vertical patterns of A L. However, the analysis of data on shade and sun leaves separately revealed an opposite trend: the bigger the A L the higher K lb. Leaf anatomical study of birch saplings revealed that this trend is attributable to enhanced vascular development with increasing leaf area. Hydraulic traits (K S, K B, K lb) of sun shoots were well co-ordinated and more strongly correlated with characteristics of shoot size than those of shade shoots, reflecting their greater evaporative load and need for stricter adjustment of hydraulic capacity with shoot size. K S increased with increasing xylem cross-sectional area to leaf area ratio (Huber value; P < 0.01), suggesting a preferential investment in water-conducting tissue (sapwood) relative to transpiring tissue (leaves), and most likely contributing to the functional stability of the hydraulic system, essential for fast-growing pioneer species.

Effects of reduced irradiance on hydraulic architecture and water relations of two olive clones with different growth potentials

The hydraulic architecture and water relations of two olive genotypes, ‘Leccino Dwarf’ (LD) and ‘Leccino Minerva’ (LM) growing at two irradiance levels i.e. full sunlight irradiance (HI) and 50% sunlight irradiance (LI) were studied. The two clones showed similar plant hydraulic conductances ( K plant) and similar conductance of roots and leaves ( K root and K leaf) when growing at equal irradiance levels. However, both K plant and K root were significantly lower in LI plants than in HI ones. On the contrary, K leaf was unaffected by the light regime. One-year-old twigs of LI plants produced longer xylem conduits but lower average diameter of conduits and less conduits per unit xylem cross-sectional area compared to HI plants. As a consequence total conductive cross-sectional area of twigs was computed to be about 16% smaller in LI individuals than in HI ones. The LM genotype resulted potentially more vulnerable to cavitation than the LD one, although shading did not influence this variable. Shading influenced root biomass negatively with stronger reduction in LM genotype than in LD one. Although transpiration rates were substantially lower in shaded than in HI plants minimum diurnal leaf water potential was about −1.2 MPa for both clones regardless the irradiance regime. Our conclusion is that the hydraulic efficiency of both olive clones was adjusted to meet the evaporative demand imposed by the irradiance regime with consequently similar equal hydraulic sufficiency.

Evaluating variability of root size system and its constitutive traits in hot pepper ( Capsicum annum L.) under water stress

The importance of root size system has long been recognized as crucial to cope with drought conditions. This investigation was conducted to: (i) evaluate the variability in root size system of hot pepper at maturity; (ii) estimate the effect of root size system on yield under drought conditions; and (iii) effect of water stress on xylem vessel development and total xylem cross-sectional area in roots of hot pepper cultivars. Twelve diverse hot pepper cultivars were grown in wooden boxes with two different water treatments, normal and in 50% water application as water deficit condition. Mean primary root length (PRL) showed a significant positive correlation with final fruit yield at normal as well as stressed condition. Total dry mass of fruit was reduced by 34.7% in drought treatments (DI) compared to full watered treatment (FI). At harvest, water-stressed plants had 21% lower root dry weight mass but higher root:shoot ratio other than FI. PRL, lateral root density, total xylem area per root cross-section showed a significant positive relationship with fruit yield. Also, lateral root density was higher in cultivars with higher xylem density, particularly in tolerant cultivars. Lateral root density ( r = 0.847, P < 0.001) and total xylem cross-sectional area in root ( r = 0.926, P < 0.001) were tightly related with total biomass production. The importance of root traits contributing to withstand drought in hot pepper is discussed.

Combined application of computer tomography and light microscopy for analysis of conductive xylem area in coarse roots of European beech and Norway spruce

Axial water transport in trees is mainly deter- mined by the gradient of negative water pressure and the structure of conductive xylem elements (i.e. conduits) con- necting the fine roots with the foliage. There is still an essential lack of knowledge concerning the relationship between wood structure and hydraulic properties, especially of coarse roots. To this end, the study aimed (1) to work out a novel approach, based on the combination of computer tomography (CT) and light microscopy (LM), for determin- ing the cumulative cross-sectional lumen area of conduits involved in the water transport of coarse roots in European beech (Fagus sylvatica) and Norway spruce (Picea abies) and (2) to demonstrate its adequacy in quantifying the functional relationship between sapwood anatomy and ascending water mass flow in the xylem. The cross-sectional sapwood area of coarse roots was assessed through CT. The cumulative cross- sectional lumen area of conduits in the sapwood (i.e. the lumen area of conductive conduits) was measured by LM in combination with interactive image analysis. The new approach was developed with coarse roots of both the tree species growing in a 60-year-old mixed forest in Bavaria, Germany. The combination of the two methods unveiled spruce to possess a distinct sapwood/heartwood boundary in small-diameter roots, whereas such roots of beech reflected a gradual transition zone; only large-diameter roots displayed a distinct boundary in beech. Additionally, the cumulative lumen area of conductive conduits was found to be approxi- mately 12% of the total coarse root cross-sectional area in both the tree species. The new approach of measuring the conductive lumen area of coarse-root conduits yielded levels of specific sap flow (i.e. axial conductivity) that substantially differed from those derived from commonly applied meth- ods, which were based on sap flow per unit of total cross- sectional root area or xylem cross-sectional area of individual roots. The combination of CT and LM will facilitate func- tional comparisons of woody roots differing in diameter and of tree species of different anatomical xylem structure.

Age-related trends in red spruce foliar plasticity in relation to declining productivity

Phenotypic plasticity in needle morphology with increasing tree size and age was investigated by comparing four age classes of red spruce (Picea rubens Sarg.) ranging from juvenile (3-12 years old) to mature (over 100 years old). With increase in tree age there were significant increases in leaf mass per unit area (LMA), mesophyll and vascular bundle area as a percentage of total needle cross-sectional area, and stomatal density. Within the vascular bundle, both xylem cross-sectional area and tracheid lumen area increased significantly, whereas air space as a percentage of total cross-sectional area decreased. These morphological changes were associated with a significant decrease in photosynthetic capacity and stomatal conductance, and an increase in (13)C enrichment. Although both photosynthetic capacity and whole-tree conductance decreased significantly between age classes 3 and 12 years, they did not differ between age classes 53 and 127 years, even though needle (13)C enrichment was significantly greater in the 127-year age class. Thus there appear to be compensatory mechanisms that maintain photosynthetic capacity as trees increase in size and vascular complexity, which in red spruce and other species, may affect leaf hydraulic conductance. Although increased LMA may contribute to reduced photosynthetic capacity in red spruce, similar relationships are not seen in other conifers.

Elevated temperature and CO2 concentration effects on xylem anatomy of Scots pine

We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross-sectional area, the greatest decrease occurring in the elevated [CO(2)] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO(2)] and that the effects of temperature were confined to the earlywood.

Why are Chloris gayana leaves shorter in salt-affected plants? Analyses in the elongation zone

Reduced hydraulic conductance calculated from growth data was suggested to be the main reason for reduced leaf expansion in salt-stressed Chloris gayana (Rhodes grass). In this work, xylem vessel cross-sections and wall enzyme activities were analysed to re-examine the effects of salinity on leaf growth in this species. Maximal segmental growth rates were 20% lower and the growth zone was 23% shorter in leaves from salinized plants than in controls; however, growth rates between 0 mm and 15 mm from the ligule were similar in both types of leaves. Xylem cross-sectional areas in this region were about 65% smaller in leaves of salinized plants, suggesting that hydraulic restrictions in the leaves of salinized plants were much higher than overall growth reductions. Extractable xyloglucan endotransglucosylase activity in this zone was twice as high in leaves of salinized plants as in leaves of controls. Nevertheless, the activity of the extracted enzyme was not affected by up to 1 M NaCl added to the reaction medium. Therefore, increased xyloglucan endotransglucosylase activity under salinity may be due to a promotion of transcription of XTH (xyloglucan endotransglucosylase/hydrolases) genes and/or translation of preformed transcripts. These results suggest that, as in drought stress, increased activity of cell wall enzymes associated with wall loosening may contribute to the maintenance of growth under saline conditions despite hydraulic restrictions.

Controlled Drought Affects Morphology and Anatomy of Salvia splendens

Polyethylene glycol 8000 (PEG-8000) was applied to a soilless growing medium at the concentrations of 0, 15, 20, 30, 42, or 50 g·L-1 to impose controlled drought. Salvia (Salvia splendens F. Sellow. ex Roem &amp; Shult.) seeds were planted in the growing medium to determine if controlled drought affects morphology and anatomy of salvia. Polyethylene glycol decreased emergence percentage and delayed emergence up to 5 days. Stem elongation of salvia treated with the five lowest concentrations was reduced up to 35% (21 days after seeding), and salvia were a maximum of 53% shorter and the canopy was 20% more narrow compared to nontreated seedlings 70 days after seeding. These morphological changes were attributed to PEG-8000 mediated reduction in leaf water potential (Ψw). The growing medium Ψw ranged from -0.29 to -0.85 MPa in PEG-8000 treated plants, and plant height was positively correlated with Ψw 21 days after seeding. Stem diameter of PEG-treated seedlings was reduced up to 0.4 mm mainly due to reductions in vascular cross-sectional area. Xylem cross-sectional area decreased more than stem and phloem cross-sectional area. Polyethylene glycol 8000 reduced vessel element number, but not diameter.

13C-photosynthate accumulation in Japanese pear fruit during the period of rapid fruit growth is limited by the sink strength of fruit rather than by the transport capacity of the pedicel

In Japanese pear, the application of GA3+4 during the period of rapid fruit growth resulted in a marked increase in pedicel diameter and bigger fruit at harvest. To elucidate the relationship between pedicel capacity and fruit growth and to determine the main factor responsible for larger fruit size at harvest, fruit growth and pedicel vascularization after GA application were examined and the carbohydrate fluxes were monitored in a spur unit by non-invasive techniques using 13C tracer. Histological studies of fruit revealed that GA increased the cell size of the mesocarp but not the cell number and core size. The investigation of carbon partitioning showed that an increase in the specific rate of carbohydrate accumulation in fruit or the strength of fruit should be responsible for an increase of fruit weight in GA-treated trees. Observation of pedicel vascularization showed that an increase in pedicel cross-sectional area (CSA) by GA application mainly resulted from phloem and xylem CSA, but it is unlikely that an increase in the transport system is the direct factor for larger fruit size. Therefore, it can be concluded that larger fruit size resulting from GA application during the period of rapid fruit growth caused an increase in the cell size of the mesocarp and increased carbon partitioning to the fruit. Although GA is closely involved with pedicel vascularization, it seems that photosynthate accumulation in fruit is limited by the sink strength of fruit rather than by the transport capacity of the pedicel.

Variation in drought response of sal (Shorea robusta) seedlings.

Plant development and distribution in areas with seasonal rainfall are often related to the ability of plants to postpone desiccation or tolerate low water potentials during drought. Regeneration of Shorea robusta Gaertn. (sal), a commercially valuable, widely distributed tree of the Indian tropical belt, is unsuccessful at the base of the Himalaya. Seedling shoots die back repeatedly during the long drought that follows the monsoon rain. During the course of one year, we monitored changes in plant and soil water potentials (Psi), leaf conductance (gw), osmotic and elastic adjustment, and xylem conductance of sal seedlings of different sizes from three landforms: an alluvial plain at 540 m elevation, a slope at 510 m, and a montane site at 1370 m. Predawn plant Psi and gw were lowest in the smallest seedlings (< 20 cm tall). Across sites and seasons, seedlings > 100 cm tall had higher morning gw than seedlings in the other size classes. In all size classes, plant Psi was lowest during early summer, when leafing begins. Among sites, Psi and gw were lowest in seedlings at the montane site. Osmotic potential was lowest during leaf development and highest during the rainy season, and tissue elasticity was highest during winter. As leaf area increased during leaf development, xylem conductance per unit of xylem cross-sectional area also increased. We conclude that low Psi is unlikely to be a major cause of seedling mortality. Small seedlings, with low Psi, had low leaf conductance. Adjustments of osmotic and elastic properties appear to aid responses of seedlings to drought.

Root and stem hydraulic conductivity as determinants of growth potential in grafted trees of apple (Malus pumila Mill.).

The anatomy of the graft tissue between a rootstock and its shoot (scion) can provide a mechanistic explanation of the way dwarfing Malus rootstocks reduce shoot growth. Considerable xylem tissue disorganization may result in graft tissue having a low hydraulic conductivity (k(h)), relative to the scion stem. The graft may influence the movement of substances in the xylem such as ions, water and plant-growth-regulating hormones. Measurements were made on 3-year-old apple trees with a low-pressure flow system to determine k(h) of root and scion stem sections incorporating the graft tissue. A range of rootstocks was examined, with different abilities of dwarfing; both ungrafted and grafted with the same scion shoot cultivar. The results showed that the hydraulic conductivity (k(hroot)) of roots from dwarfing rootstocks was lower compared with semi-vigorous rootstocks, at least for the size class of root measured (1.5 mm diameter). Scion hydraulic conductivity (k(hs)) was linked to leaf area and also to the rootstock on to which it was grafted, i.e. hydraulic conductivity was greater for the scion stem on the semi-vigorous rootstock. Expressing conductivities relative to xylem cross-sectional areas (k(s)) did not remove these differences suggesting that there were anatomical changes induced by the rootstock. The calculated hydraulic conductivity of the graft tissue was found to be lower for grafted trees on dwarfing rootstocks compared to invigorating rootstocks. These observations are discussed in relation to the mechanism(s) by which rootstock influences shoot growth in grafted trees.

Radial variation in sap flow in five laurel forest tree species in Tenerife, Canary Islands.

Variations in radial patterns of xylem water content and sap flow rate were measured in five laurel forest tree species (Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng., Myrica faya Ait., Erica arborea L. and Ilex perado Ait. ssp. platyphylla (Webb & Berth.) Tutin) growing in an experimental plot at Agua García, Tenerife, Canary Islands. Measurements were performed around midday during warm and sunny days by the heat field deformation method. In all species, water content was almost constant (around 35% by volume) over the whole xylem cross-sectional area. There were no differences in wood color over the whole cross-sectional area of the stem in most species with the exception of E. arborea, whose wood became darker in the inner layers. Radial patterns of sap flow were highly variable and did not show clear relationships with tree diameter or species. Sap flow occurred over the whole xylem cross-sectional area in some species, whereas it was limited to the outer xylem layers in others. Sap flow rate was either similar along the xylem radius or exhibited a peak in the outer part of the xylem area. Low sap flow rates with little variation in radial pattern were typical for shaded suppressed trees, whereas dominant trees exhibited high sap flow rates with a peak in the radial pattern. Stem damage resulted in a significant decrease in sap flow rate in the outer xylem layers. The outer xylem is more important for whole tree water supply than the inner xylem because of its larger size. We conclude that measurement of radial flow pattern provides a reliable method of integrating sap flow from individual measuring points to the whole tree.

Xylem hydraulic characteristics of subtropical trees from contrasting habitats grown under identical environmental conditions

Five evergreen subtropical tree species growing under identical environmental conditions were investigated to establish which hydraulic properties are genotypically rigid and which show phenotypic plasticity. Maximum xylem-specific conductivity (ks) correlated well with the anatomical characteristics (conduit diameter and density) for the four angiosperms Tecomaria capensis, Trichilia dregeana, Cinnamomum camphora and Barringtonia racemosa; the anatomy of the gymnosperm Podocarpus latifolius was not assessed. Huber values (functional xylem cross-sectional area [ratio ] leaf area) varied inversely with ks among species. Maximum leaf-specific conductivity was similar in the five unrelated species. Vulnerability of xylem to cavitation differed between species, as did the relationship between transpiration and water potential. Models of these parameters and isolated midday readings confirm that these trees operate at similar maximum leaf-specific conductivity (kl) values. The data are consistent with the hypothesis that conductivity characteristics (kl, ks) are influenced by environment, whereas vulnerability to cavitation is genetically determined.