Impacts of tree height on leaf hydraulic architecture and stomatal control in Douglas-fir

This study investigated the mechanisms involved in the regulation of stomatal closure in Douglas-fir and evaluated the potential impact of compensatory adjustments in response to increasing tree height upon these mechanisms. In the laboratory, we measured leaf hydraulic conductance (K(leaf)) as leaf water potential (Psi(l)) declined for comparison with in situ diurnal patterns of stomatal conductance (g(s)) and Psi(l) in Douglas-fir across a height gradient, allowing us to infer linkages between diurnal changes in K(leaf) and g(s). A recently developed timed rehydration technique was used in conjunction with data from pressure-volume curves to develop hydraulic vulnerability curves for needles attached to small twigs. Laboratory-measured K(leaf) declined with increasing leaf water stress and was substantially reduced at Psi(l) values of -1.34, -1.45, -1.56 and -1.92 MPa for foliage sampled at mean heights of approximately 20, 35, 44 and 55 m, respectively. In situ g(s) measurements showed that stomatal closure was initiated at Psi(l) values of -1.21, -1.36, -1.74 and -1.86 MPa along the height gradient, which was highly correlated with Psi(l) values at loss of K(leaf). Cryogenic scanning electron microscopy (SEM) images showed that relative abundances of embolized tracheids in the central vein increased with increasing leaf water stress. Leaf embolism appeared to be coupled to changes in g(s) and might perform a vital function in stomatal regulation of plant water status and water transport in conifers. The observed trends in g(s) and K(leaf) in response to changes in Psi(l) along a height gradient suggest that the foliage at the tops of tall trees is capable of maintaining stomatal conductance at more negative Psi(l). This adaptation may allow taller trees to continue to photosynthesize during periods of greater water stress.     

Vein recovery from embolism occurs under negative pressure in leaves of sunflower (Helianthus annuus)

Leaf veins undergo cavitation at water potentials (Psi(leaf)) commonly experienced by field-growing plants. Theoretically, embolism reversal should not be possible until xylem pressures rise by several kilopascals of atmospheric pressure, but recent evidence suggests that embolized conduits can be refilled even when surrounded by others at substantial tension (novel refilling). The present study reports 'novel refilling' occurring in leaf veins of sunflower (Helianthus annuus L.) while at Psi(leaf) = -0.33 MPa. Sixty per cent loss of vein hydraulic conductance (K(vein)) was recorded at Psi(leaf) < -0.65 MPa, while stem hydraulic conductance (K(stem)) was unaffected even at Psi(leaf) = -1.1 MPa. Loss of K(vein) was accompanied by stomatal closure. Water-stressed plants (Psi(leaf) = -1.1 MPa) were rehydrated overnight to different target water potentials achieved by using PEG at different concentrations as irrigation medium. K(vein) recovered by 50% at Psi(leaf) = -0.47 MPa and vein refilling was complete at Psi(leaf) = -0.33 MPa, i.e. well below the theoretical limit for conduit refilling (-0.05 MPa as calculated for sunflower minor veins). Mercurials supplied to detached leaves had no effect on the refilling process. Upon rehydration, recovery of K(vein) was not paralleled by recovery of whole-plant hydraulic conductance or leaf conductance to water vapour (g(L)), as a likely consequence of hydraulic failure of other components of the water pathway (root system or extravascular leaf compartments) and/or root-to-leaf chemical signalling. This is the first study providing experimental evidence for 'novel refilling' in a herbaceous dicot and highlighting the importance of this process in the leaf.     

The hydraulic architecture of Juniperus communis L. ssp. communis: shrubs and trees compared

Juniperus communis ssp. communis can grow like a shrub or it can develop a tree-like habit. In this study, the hydraulic architecture of these contrasting growth forms was compared. We analysed the hydraulic efficiency (leaf-specific conductivity, k(l); specific conductivity, k(s); Huber value, HV) and the vulnerability to cavitation (the water potential corresponding to a 50% loss of conductivity, Psi(50)), as well as anatomical parameters [mean tracheid diameter, d; mean hydraulic diameter, d(h); cell wall reinforcement (t/b)(h)(2)] of shrub shoots, tree stems and tree branches. Shrub shoots were similar to tree branches (especially to lower branches) in growth form and conductivity (k(l) = 1.93 +/- 0.11 m(2) s(-1) MPa(-1) 10(-7), k(s) = 5.71 +/- 0.19 m(2) s(-1) MPa(-1) 10(-4)), but were similar to tree stems in their vulnerability to cavitation (Psi(50) = -5.81 +/- 0.08 MPa). Tree stems showed extraordinarily high k(l) and k(s) values, and HV increased from the base up. Stem xylem was more vulnerable to cavitation than branch xylem, where Psi(50) increased from lower (Psi(50) = -6.44 +/- 0.19 MPa) to upper branches (Psi(50) = -5.98 +/- 0.13 MPa). Conduit diameters were correlated with k(l) and k(s). Data indicate that differences in hydraulic architecture correspond to changes in growth form. In some aspects, the xylem hydraulics of tree-like Juniperus communis differs from that of other coniferous tree species.     

Effects of light intensity and duration on leaf hydraulic conductance and distribution of resistance in shoots of silver birch (Betula pendula)

Variation in leaf hydraulic conductance (K(L)) and distribution of resistance in response to light intensity and duration were examined in shoots of silver birch (Betula pendula Roth). K(L) was determined on detached shoots using the evaporative flux method (transpiration was measured with a porometer and water potential drop with a pressure chamber). Although K(L) depended on light duration per se, its dynamics was largely determined by leaf temperature (T(L)). Both upper-crown leaves and branches developed in well-illuminated environment exhibited higher hydraulic efficiency compared with the lower crown, accounting for vertical trends of apparent soil-to-leaf hydraulic conductance in canopy of silver birch revealed in our previous studies. K(L) varied significantly with light intensity, the highest values for both shade and sun foliage were recorded at photosynthetic photon flux density of 330 micromol m(-2) s(-1). Light responses of K(L) were associated evidently with an irradiance-mediated effect on extravascular tissues involving regulation of cell membrane aquaporins. Effects of irradiance on K(L) resulted in changes of Psi(L), bringing about considerable alteration in partitioning of the resistance between leaves and branch (leafless shoot stem): the contribution of leaves to the shoot total resistance decreased from 94% at -1.0 MPa to 75% at -0.2 MPa. Treatment with HgCl2 decreased hydraulic conductance of both leaves and branches, implying that condition of bordered pit membranes or shoot living tissues may be involved in responses of xylem conductance to Hg2+.     

Hydraulic efficiency and safety of leader shoots and twigs in Norway spruce growing at the alpine timberline

Xylem within trees varies in its hydraulic efficiency and safety. Trees at the alpine timberline were expected to exhibit a hydraulic architecture protecting the leader shoot from winter embolism. Hydraulic and related anatomical parameters were compared as well as seasonal courses of winter embolism in leader shoots and twigs of Norway spruce trees growing at 2000 m. Leader shoots had a 1.4-fold higher specific hydraulic conductivity (ks) as well as a 4.9-fold higher leaf specific conductivity (kl) than side twigs. Vulnerability to drought-induced embolism was lower in leader shoots with a 50% loss of conductivity occurring at a water potential (Psi 50) 0.7 MPa lower than in twigs. Higher ks and kl were related to 1.2-fold wider tracheid diameters in leader shoots. Lower vulnerability corresponded to smaller pit dimensions but not to wood density. High ks and kl reflect the hydraulic dominance of the leader shoot, which is important for its water supply during summer. Low vulnerability protects the leader shoot from embolism during the winter season. In field measurements at the timberline during the winter of 2001/2002, conductivity losses of up to 56% were observed only in twigs while leader shoots showed little or no embolism. Results demonstrate that leader shoot xylem is both hydraulically efficient and safe.     

Stomatal closure during leaf dehydration,correlation with other leaf physiological traits

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.     

Hydrostatic constraints on morphological exploitation of light in tall Sequoia sempervirens trees

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown. C isotope composition of leaves (delta(13)C) increased with increasing height indicating greater photosynthetic water use efficiency in the upper crown. Leaf and shoot morphology changed continuously with height. In contrast, their relationships with light availability were discontinuous: morphological characteristics did not correspond to increasing light availability above 55-85 m. Mass-based chlorophyll concentration (chl) decreased with increasing height and increasing light availability. In contrast, area-based chl remained constant or increased with increasing height. Mass-based maximum rate of net photosynthesis (P (max)) decreased with increasing height, whereas area-based P (max) reached maximum at 78.4 m and decreased with increasing height thereafter. Mass-based P (max) increased with increasing shoot mass per area (SMA), whereas area-based P (max) was not correlated with SMA in the upper crown. Our results suggest that hydrostatic limitation of morphological development constrains exploitation of light in the upper crown and contributes to reduced photosynthetic rates and, ultimately, reduced height growth at the tops of tall S. sempervirens trees.     

Effects of prolonged drought on the anatomy of sun and shade needles in young Norway spruce trees

Predicted increases in the frequency and duration of drought are expected to negatively affect tree vitality, but we know little about how water shortage will influence needle anatomy and thereby the trees’ photosynthetic and hydraulic capacity. In this study, we evaluated anatomical changes in sun and shade needles of 20‐year‐old Norway spruce trees exposed to artificial drought stress. Canopy position was found to be important for needle structure, as sun needles had significantly higher values than shade needles for all anatomical traits (i.e., cross‐sectional needle area, number of tracheids in needle, needle hydraulic conductivity, and tracheid lumen area), except proportion of xylem area per cross‐sectional needle area. In sun needles, drought reduced all trait values by 10–40%, whereas in shade needles, only tracheid maximum diameter was reduced by drought. Due to the relatively weaker response of shade needles than sun needles in drought‐stressed trees, the difference between the two needle types was reduced by 25% in the drought‐stressed trees compared to the control trees. The observed changes in needle anatomy provide new understanding of how Norway spruce adapts to drought stress and may improve predictions of how forests will respond to global climate change.     

Increases in water potential gradient reduce xylem conductivity in whole plants. Evidence from a low-pressure conductivity method

A new method using hydrostatic suctions (less than 0.02 MPa) was used to measure whole-root conductivity (K(r)) in saplings of two angiosperm pioneer trees (Eucalyptus regnans and Toona australis) and two rainforest conifers (Dacrycarpus dacrydioides and Nageia fleurii). The resultant K(r) was combined with measurements of stem and leaf hydraulic conductivity to calculate whole-plant conductivity and to predict leaf water potential (Psi(l)) during transpiration. At normal soil temperatures there was good agreement between measured and predicted Psi(l) during transpiration in all species. Changes in the soil-to-leaf water potential gradient were produced by root chilling, and in three of the four species, changes in Psi(l) corresponded to those expected by the effect of increased water viscosity on K(r). In one species, however, root chilling produced severe plant wilting and a decline in Psi(l) significantly below the predicted value. In this species Psi(l) decreased to a value close to, or below, the Psi(l) at 50% xylem cavitation. It is concluded that decreased whole-plant conductivity in T. australis resulted from a decrease in xylem conductivity due to stress-induced cavitation.     

The hydraulic conductivity of the xylem in conifer needles (Picea abies and Pinus mugo)

Main resistances of the plant water transport system are situated in leaves. In contrast to angiosperm leaves, knowledge of conifer needle hydraulics and of the partitioning of resistances within needles is poor. A new technique was developed which enabled flow-meter measurements through needles embedded in paraffin and thus quantification of the specific hydraulic conductivity (K(s)) of the needle xylem. In Picea abies, xylem K(s) of needle and axes as well as in needles of different age were compared. In Pinus mugo, resistance partitioning within needles was estimated by measurements of xylem K(s) and leaf conductance (K(leaf), measured via 'rehydration kinetics'). Mean K(s) in P. abies needles was 3.5×10(-4) m(2) s(-1) MPa(-1) with a decrease in older needles, and over all similar to K(s) of corresponding axes xylem. In needles of P. mugo, K(s) was 0.9×10(-4) m(2) s(-1) MPa(-1), and 24% of total needle resistance was situated in the xylem. The results indicate species-specific differences in the hydraulic efficiency of conifer needle xylem. The vascular section of the water transport system is a minor but relevant resistance in needles.     

Effects of the experimental blockage of the major veins on hydraulics and gas exchange of Prunus laurocerasus L. leaves

The impact of leaf vein blockage on leaf hydraulic conductance (K(L)), gas exchange (g(L)) and water potential (Psi(L)) was studied in Prunus laurocerasus L., a broad-leaved evergreen. For this purpose, leaves were measured for the three variables above, either with an intact leaf blade (controls) or with the midrib cut a third of the way up (cut a), or with the midrib cut at three different points and the first-order veins cut through near their insertion to the midrib (cut b), or with the midrib cut at 2 mm from the leaf base (cut c). All the cut surfaces were sealed with cyanoacrylate. A serial decrease of K(L) was recorded from cut a to cut c with respect to that measured for the controls, i.e. a K(L) loss of about 37% (cut a), 57% (cut b) and 87% (cut c). A positive linear relationship appeared to exist between g(L) and K(L) with a high correlation coefficient (r(2)=0.99) and a high statistical significance (P <0.01). Even under a severe drop in K(L) (as that induced by cut c), leaf water potential remained approximately constant and not statistically different from Psi(L) measured for the controls. In fact, Psi(L) ranged between -0.83 and -0.98 MPa, i.e. within the cavitation threshold of leaves in terms of the critical Psi(L) inducing a significant production of ultrasound acoustic emissions which was -0.94+/-0.09 MPa. The conclusion was that stomata were very sensitive to changes in K(L) and that stomatal closure led to the homeostatic maintenance of Psi(L) and cavitation avoidance.     

Low air humidity increases leaf-specific hydraulic conductance of Arabidopsis thaliana (L.) Heynh (Brassicaceae)

The typical isohydric plant response to low relative humidity involves stomatal closure, followed by long-term responses like adjustment of shoot-to-root ratios. Little information is available on the early responses of the root system to exposure of shoots to low humidity, nor is it clear to what extent responses of Arabidopsis thaliana conform to the isohydric model. In this study, A. thaliana plants grown hydroponically at high humidity were exposed to two constant relative humidities, 17% and 77%, while the root system remained in aerated nutrient solution. Leaf conductance (g(s)), transpiration, water potential (Psi(l)), osmotic potential, and whole plant hydraulic conductance (K) were determined for the following time intervals: 0-10, 10-20, and 20-40 min, and 0-5, 5-10, and 24-29 h. At low relative humidity, no change in g(s) was detected. Psi(l) decreased by 0.28 MPa during the first 5 h and then remained stable. During the first hour, leaf-specific K averaged 1.6 x 10(-5) kg MPa(-1) m(-2) s(-1) at high humidity. At low humidity it increased >3-fold to 5.8 x 10(-5) kg MPa(-1) m(-2) s(-1). Similar significant differences in K were observed during all time periods. Low concentration mercury amendments in the hydroponic solution (5 microM and 10 microM HgCl(2)) had no discernible influence, but pre-exposure to 50 microM HgCl(2) reduced K differences between humidity treatments. As HgCl(2) is known to be a potent inhibitor of aquaporin function, this suggests that aquaporins may have played a role in the fast hydraulic response of plants transferred to low humidity. The rapid hydraulic response and the influence of mercury raise the possibility that an alternative response to atmospheric dryness is increased K modulated by aquaporins.     

Water stress, shoot growth and storage of non-structural carbohydrates along a tree height gradient in a tall conifer

We analysed concentrations of starch, sucrose, glucose and fructose in upper branch wood, foliage and trunk sapwood of Douglas-fir trees in height classes ranging from ~2 to ~57 m. Mean concentrations of non-structural carbohydrates (NSC) for all tissues were highest in the tallest height class and lowest in the lowest height class, and height-related trends in NSC were most pronounced in branches. Throughout a 17-month sampling period, mean values of branch NSC from the 57 m trees ranged between 30 and 377% greater than the 2 m trees. Branch NSC was inversely correlated with midday shoot water potential (Ψ(l)), shoot osmotic potential at full turgor (Ψ) and shoot extension. Temporal fluctuation in branch NSC was inversely correlated with height, and positively correlated with midday Ψ(l) , Ψ and shoot extension. The positive correlation between height and storage of NSC, and the negative correlation between NSC storage and shoot extension provide evidence that size-related growth decline in trees is not strongly associated with constraints on photosynthesis. The negative correlation between height and fluctuation in NSC suggests that mobilization of photosynthate in taller trees is constrained by some factor such as reductions in turgor-driven cell expansion or constraints on phloem transport.     

Morphological and stomatal responses of Norway spruce foliage to irradiance within a canopy depending on shoot age

Morphological and stomatal responses of Norway spruce (Picea abies) foliage to light availability were studied in respect to shoot age. Needle minor diameter (D(1), anatomical width), major diameter (D(2), anatomical thickness), dry weight (M), and tissue density index (I(D)) increased, and needle flatness (Fl) and specific leaf area (SLA) decreased with foliage age, while shade foliage demonstrated higher morphological plasticity as compared to sun foliage. Needle minor diameter, dry weight, and the ratio of total to projected leaf area increased, and needle flatness and specific leaf area decreased with daily average photosynthetic photon flux density (Q(D)). The current-year foliage exhibited the highest variation with irradiance, while the morphological plasticity decreased with needle ageing. The morphological characteristics of needles were independent of irradiance if Q(D) was above 300 μmol m(-2) s(-1). D(1) was the only linear needle characteristic which significantly changed with light availability within a canopy, and thus determined needle flatness, SLA, as well as the ratio of total to projected leaf area (TLA/PLA). Needle flatness was a characteristic responding most sensitively to the photosynthetic photon flux density, R(2) was 0.68, 0.44, and 0.49 for the current-year, 1-year-old, and 2-year-old foliage, respectively. TLA/PLA ranged from 2.2 to 4.0 depending on D(1). Variation in SLA in response to light availability can be attributed to changes both in needle shape and tissue density. Stomatal responses to photosynthetic photon flux density (Q(P)) depended on foliage type (sun or shade) and age. Sun needles demonstrated higher daily maximum leaf conductances to water vapour compared to shade needles. The shade needles responded more sensitively to changes in Q(P) at dawn and sunset than the sun needles, while older needles of both foliage types exhibited faster stomatal responses. The light-saturation of leaf conductance (g(L)) was achieved by 20 μmol m(-2) s(-1) for shade foliage, and approximately by 50 μmol m(-2) s(-1) for sun foliage. As a rule, g(L) changed in response to irradiance faster in the evening, i.e. at decreasing irradiance. Stomata were not usually completely closed in the dark before sunrise and after sunset, the phenomenon being more pronounced in older shoots and sun needles. Nightly water losses from spruce foliage are attributable primarily to older shoots, and are related to age-dependent changes in stomatal responsiveness.     

Bayesian analysis of Douglas-fir hydraulic architecture at multiple scales

We used a Bayesian hierarchical model to analyze the variation in xylem anatomy, hydraulic properties, and the relationship between anatomy and properties within Douglas-fir trees. The hierarchical scales in our study included fertilization treatments (fertilized and unfertilized), trees within the treatments, and positions within the trees. We measured tracheid diameter, tracheid length, percent latewood, number of pits per cell, density, and specific conductivity (K _s) on seven positions in each of 16 fertilized and 16 unfertilized trees: the trunk at cambial age 52 (breast height), 25, and 5; a branch at cambial age 20 and 7; and a root at cambial age 42 and 22. Vulnerability to embolism was also measured on the oldest trunk, branch, and root positions. For any measurement, there was little variation between treatments, however, there was great variation among positions. Tracheid diameter, tracheid length, number of pits per cell, K _s, and vulnerability to embolism decreased vertically from the roots to the branches. Correlations were evident between some positions for tracheid diameter, percent earlywood, pits per cell, and vulnerability to embolism, mostly in the fertilized treatment. We found evidence for large-scale relationships (among all observations from all trees) between density and tracheid diameter, K _s and diameter, vulnerability and diameter, K _s and pits per cell, and vulnerability and pits per cell. At a smaller scale of within position, however, usually only the branches and roots maintained the relationship.     

Interactive effects of water stress and xylem-limited bacterial infection on the water relations of a host vine

Xylella fastidiosa, a xylem-limited bacterial pathogen that causes bacterial leaf scorch in its hosts, has a diverse and extensive host range among plant species worldwide. Previous work has shown that water stress enhances leaf scorch symptom severity and progression along the stem in Parthenocissus quinquefolia infected by X. fastidiosa. The objective here was to investigate the mechanisms underlying the interaction of water stress and infection by X. fastidiosa. Using the eastern deciduous forest vine, P. quinquefolia, infection and water availability were manipulated while measuring leaf water potentials (psi(L)), stomatal conductance (g(s)), whole shoot hydraulic conductance (K(h)), per cent xylem embolism, and xylem vessel dimensions. No significant differences in any of the physiological measurements were found between control and infected plants prior to drought. Drought treatment significantly reduced psi(L) and g(s) at all leaf positions throughout the day in late summer in both years of the study. In addition, infection significantly reduced psi(L) and g(s) in the most basal leaf positions in late summer in both years. Whole shoot hydraulic conductance was reduced by both low water and infection treatments. However, per cent embolized vessels and mean vessel diameter were affected by drought treatment only. These results imply that the major effect of infection by X. fastidiosa occurs due to reduced hydraulic conductance caused by clogging of the vessels, and not increased cavitation and embolism of xylem elements. The reduced K(h) caused by X. fastidiosa infection acts additively with the water limitation imposed by Drought stress.     

High apoplastic solute concentrations in leaves alter water relations of the halophytic shrub, Sarcobatus vermiculatus

Predawn plant water potential (Psi(w)) is used to estimate soil moisture available to plants because plants are expected to equilibrate with the root-zone Psi(w). Although this equilibrium assumption provides the basis for interpreting many physiological and ecological parameters, much work suggests predawn plant Psi(w) is often more negative than root-zone soil Psi(w). For many halophytes even when soils are well-watered and night-time shoot and root water loss eliminated, predawn disequilibrium (PDD) between leaf and soil Psi(w) can exceed 0.5 MPa. A model halophyte, Sarcobatus vermiculatus, was used to test the predictions that low predawn solute potential (Psi(s)) in the leaf apoplast is a major mechanism driving PDD and that low Psi(s) is due to high Na+ and K+ concentrations in the leaf apoplast. Measurements of leaf cell turgor (Psi(p)) and solute potential (Psi(s)) of plants grown under a range of soil salinities demonstrated that predawn symplast Psi(w) was 1.7 to 2.1 MPa more negative than predawn xylem Psi(w), indicating a significant negative apoplastic Psi(s). Measurements on isolated apoplastic fluid indicated that Na+ concentrations in the leaf apoplast ranged from 80 to 230 mM, depending on salinity, while apoplastic K+ remained around 50 mM. The water relations measurements suggest that without a low apoplastic Psi(s), predawn Psi(p) may reach pressures that could cause cell damage. It is proposed that low predawn apoplastic Psi(s) may be an efficient way to regulate Psi(p) in plants that accumulate high concentrations of osmotica or when plants are subject to fluctuating patterns of soil water availability.     

Root pressurization affects growth-induced water potentials and growth in dehydrated maize leaves

Profiles of water potential (Psi w) were measured from the soil to the tips of growing leaves of maize (Zea mays L.) when pressure (P) was applied to the soil/root system. At moderately low soil Psi w, leaf elongation was somewhat inhibited, large tensions existed in the xylem, and Psi w were slightly lower in the elongating leaf tissues than in the xylem, i.e. a growth-induced Psi w was present but small. With P, the tension was relieved, enlarging the difference in Psi w between the xylem and the elongating tissues, i.e. enlarging the growth-induced Psi w, which is critical for growth. Guttation occurred, confirming the high Psi w of the xylem, and the mature leaf tissue rehydrated. Water uptake increased and met the requirements of transpiration. Leaf elongation recovered to control rates. Under more severe conditions at lower soil Psi w, P induced only a brief elongation and the growth-induced Psi w responded only slightly. Guttation did not occur, water flow did not meet the requirements of transpiration, and the mature leaf tissues did not rehydrate. A rewatering experiment indicated that a low conductance existed in the severely dehydrated soil, which limited water delivery to the root and shoot. Therefore, the initial growth inhibition appeared to be hydraulic because the enlargement of the growth-induced Psi w by P together with rehydration of the mature leaf tissue were essential for growth recovery. In more severe conditions, P was ineffective because the soil could not supply water at the required rate, and metabolic factors began to contribute to the inhibition.     

樟子松人工固沙林衰退的主要特征

针对辽宁省章古台地区樟子松人工固沙林出现的衰退枯死现象,系统地研究了樟子松人工固沙林衰退的主要特征,结果表明,衰退的樟子松人工固沙林外貌景观呈灰绿色,针叶纤细,开花结实率低,平均单株球果数量为10．4～16．5个,成熟种子千粒重为6．96～7.39g,种子空粒、涩粒较多．生长季内2年生针叶营养元素季节变化规律相似,但N、P含量下降,K含量明显增高,表明养分循环失调;衰退林分2年生针叶叶绿素含量较高,健康林分1年生较高且增幅较大．松枯梢病的侵害是樟子松人工固沙林衰退的最明显标志．林分衰退后,树高和胸径生长量下降明显,林分胸径分布结构“左移”(径级小的株数增多),衰弱(亡)木数量增加了15.9％～27．2％;根量分别减少了22．9％～28．9％。其中吸收根减少量最大。