Girdling decreases photosynthetic electron fluxes and induces sustained photoprotection in mango leaves

Girdling has been suggested as a way to improve earliness and intensity of flowering in mango (Mangifer indica L.). However, the accumulation of carbohydrates associated with girdling may result in a decrease in photosynthesis. We assessed the long-term effect of branch girdling during the prefloral period on leaf net photosynthesis (A(n)) of 3-year old mango trees, cv. Cogshall, growing on La Réunion island. Leaf gas exchange and chlorophyll fluorescence parameters were measured monthly from March to August 2004 on recently matured leaves on girdled and non-girdled branches. Within 28 days after girdling, A(n) was reduced by 77% and remained at about 2 micromol CO(2) m(-2 ) s(-1) until the beginning of flowering. The decrease in photosynthetic electron transport rate (J) and sustained photoprotection (reflected by the decrease in predawn maximal efficiency of photosystem II) effectively protected leaves on girdled branches from photodamage, as shown by the vigorous recovery of A(n) and J observed immediately after the appearance of inflorescences. These increases in A(n) and J were unaccompanied by a decrease in leaf carbohydrate concentration during the first month following the onset of flowering, indicating that there are carbohydrate-dependent and carbohydrate-independent mechanisms of sink regulation of photosynthesis. It is concluded that girdling does not necessarily lead to irreversible damage, even in the presence of a fourfold increase in leaf starch concentration and in the absence of any sink activity. However, the decrease in leaf nitrogen concentration indicates that there may exist long-term negative effects of branch girdling on photosynthetic capacity. A modified version of the biochemical model of A(n) is presented that takes account of the effect of leaf starch concentration on J.     

Responses of foliar delta13C, gas exchange and leaf morphology to reduced hydraulic conductivity in Pinus monticola branches

We tested the hypothesis that branch hydraulic conductivity partly controls foliar stable carbon isotope ratio (delta13C) by its influence on stomatal conductance in Pinus monticola Dougl. Notching and phloem-girdling treatments were applied to reduce branch conductivity over the course of a growing season. Notching and phloem girdling reduced leaf-specific conductivity (LSC) by about 30 and 90%, respectively. The 90% reduction in LSC increased foliar delta13C by about 1 per thousand (P < 0.0001, n = 65), whereas the 30% reduction in LSC had no effect on foliar delta13C (P = 0.90, n = 65). Variation in the delta13C of dark respiration was similar to that of whole-tissues when compared among treatments. These isotopic measurements, in addition to instantaneous gas exchange measurements, suggested only minor adjustments in the ratio of intercellular to atmospheric CO2 partial pressures (ci/ca) in response to experimentally reduced hydraulic conductivity. A strong correlation was observed between stomatal conductance (gs) and photosynthetic demand over a tenfold range in gs. Although ci/ca and delta13C appeared to be relatively homeostatic, current-year leaf area varied linearly as a function of branch hydraulic conductivity (r2 = 0.69, P < 0.0001, n = 18). These results suggest that, for Pinus monticola, adjustment of leaf area is a more important response to reduced branch conductivity than adjustment of ci/ca.     

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.     

Stem girdling manipulates leaf sugar concentrations and anthocyanin expression in sugar maple trees during autumn

To better understand the effects of sugar accumulation on red color development of foliage during autumn, we compared carbohydrate concentration, anthocyanin expression and xylem pressure potential of foliage on girdled versus non-girled (control) branches of 12 mature, open-grown sugar maple (Acer saccharum Marsh.) trees. Half of the study trees were known to exhibit mostly yellow foliar coloration and half historically displayed red coloration. Leaves from both girdled and control branches were harvested at peak color expression (i.e., little or no chlorophyll present). Disruption of phloem export by girdling increased foliar sucrose, glucose and fructose concentrations regardless of historical tree color patterns. Branch girdling also increased foliar anthocyanin expression from 50.4 to 66.7% in historically red trees and from 11.7 to 54.2% in historically yellow trees, the latter representing about a fivefold increase compared with control branches. Correlation analyses indicated a strong and consistent relationship between foliar red coloration and sugar concentrations, particularly glucose and fructose, in both girdled and control branches. Measures of xylem pressure potentials confirmed that girdling was a phloem-specific treatment and had no effect on water transport to distal leaves. Results indicate that stem girdling increased foliar sugar concentrations and enhanced anthocyanin expression during autumn in sugar maple foliage. Native environmental stresses (e.g., low autumn temperatures) that reduce phloem transport may promote similar physiological outcomes.     

Patterns of hydraulic architecture and water relations of two tropical canopy trees with contrasting leaf phenologies: Ochroma pyramidale and Pseudobombax septenatum

Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees

We tested the hypotheses that hydraulic conductance is lower in old (about 250 years old and 30 m tall) compared to young (about 40 years old and 10 m tall) Pinus ponderosa Dougl. ex Laws. trees and that lower hydraulic conductance of old trees limits their photosynthesis. Hydraulic conductance at the end of summer 1995, calculated from leaf water potential and leaf gas exchange measurements on one-year-old needles, was 44% lower in old trees compared to young trees growing in a mixed age-class stand on the east slope of the Oregon Cascades. Whole-tree sapflow per unit leaf area averaged 53% lower in old trees compared to young trees and mean hydraulic conductance calculated from sapflow and water potential data was 63% lower in old trees than in young trees. For the entire summer, stomatal conductance (g(s)) and assimilation (A) declined more steeply with air saturation deficit (D) in old trees than in young trees. For both old and young trees, mean g(s) and A were approximately 32 and 21% lower, respectively, at typical midday D values (2.5-3.0 kPa). We hypothesized that if hydraulic conductance limits g(s) and A, then increasing or decreasing the leaf specific conductance of a branch will result in proportional changes in the responses of g(s) and A with D. Removal of 50% of the foliage from a set of experimental branches on old trees caused g(s) and A to decline less steeply with D in early summer, but values were not significantly different from control values in late summer. Cutting transverse notches in branches on young trees had no effect on the responses of g(s) and A with D. Leaf nitrogen content and photosynthetic capacity were similar suggesting that differences in g(s) and A between old and young trees were not caused by differences in photosynthetic capacity.     

Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in contrasting climates

We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.     

Gender-specific patterns of aboveground allocation, canopy conductance and water use in a dominant riparian tree species: Acer negundo

Acer negundo Sarg. (box elder) is a dioecious tree species that dominates riparian systems at mid elevations throughout the southwest and Intermountain West of the United States. Previous studies have shown that female A. negundo trees occur at higher frequencies along stream margins, whereas males occur at higher frequencies in drier microsites. To better understand the adaptive significance of sex ratio biases and their impact on the ecohydrology of riparian ecosystems, we examined whole-plant water relations and hydraulic properties of mature male and female A. negundo trees occurring within 1 m of a perennial stream channel. We hypothesized that (1) females would have significantly greater canopy water fluxes than males (particularly during periods of seed production: May-June), and (2) xylem in females is more hydraulically efficient but more vulnerable to cavitation than xylem in males. Mean sap flux density (J(s)) during the early growing season (May and June) was 43% higher in female trees than in male trees (n = 6 and 7 trees respectively, P < 0.0001). Mean J(s) in July and August remained 17% higher in females than in males (P = 0.0009). Mean canopy stomatal conductance per unit leaf area (g(s,leaf)) in May and June was on average 140% higher in females than in males (P < 0.0001). Mean g(s,leaf) in July and August remained 69% higher in female trees than in male trees (P < 0.0001). Canopy stomatal conductance scaled to basal area was 90 and 31% higher in females relative to males during May-June and July-August, respectively (P < 0.0001 during both periods). Conversely, there were no apparent differences in either branch hydraulic conductance or branch xylem cavitation vulnerability between genders. These results improve our capacity to describe the adaptive forces that shape the spatial distribution of male and female trees in dioecious species, and their consequences for ecohydrological processes in riparian ecosystems.     

Girdling effects on fruit set and quantum yield efficiency of PSII in two Citrus cultivars

Girdling effects on fruitlet abscission, leaf chlorophyll, chlorophyll a fluorescence and carbohydrate concentration in various flowering and vegetative shoots were studied during natural fruit drop in two Citrus cultivars. Irrespective of shoot type, girdling delayed fruitlet abscission, but only fruitlets borne on leafy shoots had increased final fruit set. Chlorophyll a fluorescence analysis revealed differences in quantum yield efficiency of photosystem II of light adapted leaves (Phi(PSII)) among shoot types and in response to girdling. In young leaves of vegetative shoots, girdling decreased Phi(PSII), whereas Phi(PSII) increased from Day 30 after girdling in young leaves of leafy flowering shoots; however, Phi(PSII) did not change in mature leaves during fruit set in either control or girdled trees. Girdling altered leaf carbohydrate concentrations and the photosynthetic performance of the various shoot types. Our results indicate that, in Citrus, several carbohydrate-based regulatory mechanisms of photosynthesis coexist during carbohydrate accumulation brought about by girdling. It is concluded that the delay in fruitlet abscission and the increase in Phi(PSII )observed in girdled leafy flowering shoots are the mechanisms underlying the enhancement of fruit set after girdling.     

Coordinating leaf functional traits with branch hydraulic conductivity: resource substitution and implications for carbon gain

We studied relationships among branch hydraulic conductivity, xylem embolism, stomatal conductance (gs), foliar nitrogen (N) concentration and specific leaf area (SLA) of seven tree species growing at four temperate woodland sites spanning a 464-1350 mm rainfall gradient. Specifically, we examined the question: are gs and foliar N concentration coordinated with branch hydraulic conductivity and, if so, what are the implications for carbon assimilation? Area-based, light-saturated photosynthetic rate (Aa) was uniquely and positively correlated with gs and foliar N concentration. Multiple regression analyses showed that, when variability in SLA was controlled for, the (positive) partial slope for each predictor remained significant. In contrast, there was a negative correlation between gs and foliar N concentration such that, for any given Aa, leaves with a high gs allocated less N to foliage than leaves with a low gs. Foliar N concentration was negatively correlated with branch hydraulic conductivity, whereas gs was positively correlated with branch hydraulic conductivity. These relationships were also significant when variability in leaf area to sapwood area ratio, gs and SLA were controlled for in a multiple regression, suggesting that the relationships were unique and independent of other confounding factors. Trees with low water transport capacity were able to support a high Aa by increasing investment in foliar N. Resource substitution occurred such that there was a trade-off between gs and foliar N in relation to branch hydraulic conductivity. High Aa could be sustained through either a high branch hydraulic conductivity and hence high gs and a low allocation to foliar N, or the effect of a low branch hydraulic conductivity and hence low gs could be offset by a high allocation to foliar N. The results are discussed in relation to mechanisms for minimizing the negative effects of limited water availability on carbon gain.     

Hydraulic conductivity in roots of ponderosa pine infected with black-stain (Leptographium wageneri) or annosus (Heterobasidion annosum) root disease

Roots from healthy and diseased mature ponderosa pine, Pinus ponderosa Laws., trees were excavated from a site near Burns, Oregon. The diseased trees were infected with black-stain root disease, Leptographium wageneri Kendrick, or annosus root disease, Heterobasidion annosum (Fr.) Bref., or both. Axial hydraulic conductivity of the roots was measured under a positive head pressure of 5 kPa, and the conducting area was stained with safranin dye to determine specific conductivity (k(s)). In diseased roots, only 8-12% of the cross-sectional xylem area conducted water. Resin-soaked xylem completely restricted water transport and accounted for 13-16% of the loss in conducting area. In roots with black-stain root disease, 17% of the loss in conducting area was associated with unstained xylem, possibly resulting from occlusions or embolisms. Based on the entire cross-sectional area of infected roots, the k(s) of roots infected with black-stain root disease was 4.6% of that for healthy roots, whereas the k(s) of roots infected with annosus root disease was 2.6% of that for healthy roots. Although these low values were partly the result of the presence of a large number of diseased roots (72%) with no conducting xylem, the k(s) of functional xylem of diseased roots was only 33% of that for healthy roots. The low k(s) values of functional xylem in diseased roots may be caused by fungus induced occlusions preceding cavitation and embolism of tracheids. The k(s) of disease-free roots from diseased trees was only 70% of that for healthy roots from healthy trees. The disease-free roots had the same mean tracheid diameter and tissue density as the healthy roots, suggesting that the lower k(s) in disease-free roots of diseased trees may also have been caused by partial xylary occlusions.     

Systematic variation in xylem hydraulic capacity within the crown of white ash (Fraxinus americana)

A 7-m tall white ash tree (Fraxinus americana Marsh.) was dissected, and hydraulic parameters of the xylem were determined by inducing a steady-state flow of water through the stem segments and monitoring volume and velocity flow rates. Leaf-specific conductivity (LSC) was highest in the main stem and lowest in some of the leaf-bearing lateral shoots. The LSC was higher in the main stem than in branches and higher in primary than in secondary branch axes. Terminal leaf-bearing shoots were larger and had a significantly greater mean LSC than subjacent lateral shoots. A significant reduction in LSC was associated with the transition between 1- and 2-year-old growth. In branches of the same age, there was a close correspondence among LSC, branch position and branch size. The average LSC of leaf-bearing shoots from south-facing branches was 43% greater than that of shoots from north-facing branches. Within-crown variation in LSC was associated with variation in velocity flow rate (V). By contrast, the ratio of potentially functional xylem area to supported leaf area (A(pf)/A(l)) was relatively stable throughout the crown. Stratification of stems by Strahler order accounted for approximately 70% of the total variation in LSC. These results suggest that (1) there exists a systematic pattern of variation in LSC distribution within the crown of white ash, (2) within-crown variability in LSC is primarily the result of variation in mean vessel diameter, and (3) there is a physiological linkage between LSC and crown morphology that is maintained through a positive feedback mechanism during branch ontogeny.     

How timing of stem girdling affects needle xylem structure in Scots pine

While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source–sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source–sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area.     

Formation and seasonal occurrence of xylem embolism in Alnus cordata

We investigated the vulnerability of xylem to embolism and the seasonal occurrence of xylem embolism in Italian alder (Alnus cordata Loisel.) by acoustic and hydraulic methods. Wood anatomy was also studied. More than eighty percent of the vessels were less than 50 mm long and no vessels were longer than 120 mm. Mean vessel diameter was 48 micro m. Ultrasound acoustic emissions from root and branch segments dehydrating in air followed a similar pattern: in both tissues, emission peaks were recorded when the relative water content of the xylem was around 0.2. In branches dehydrating in air, xylem embolism increased linearly as water potential decreased. In trees in the field, more than 80 percent of hydraulic conductivity was lost in the tree crowns during winter. Recovery from winter embolism occurred mostly before bud burst. In summer, xylem embolism was low (< 30%) and acoustic emissions from roots, stem and branches of trees in the field were also low.     

Viability of thinning sessile oak stands by girdling

The effects of girdling were analysed in a sessile oak (Quercuspetraea Matts. Leibl.) forest with an 80-year-old coppice withstandard structure in northern Spain. The study evaluated theviability of girdling as an alternative to felling trees duringcrown thinning. Eighty-four direct competitors of selected futurecrop trees were girdled by double notching and peeling. Threeyears after girdling, 100 per cent of the treated trees haddied. Approximately half of the girdled trees did not resprout.The occurrence of basal sprouting was very low, and more than35 per cent of the sprouts had very low vigour. No correlationwas found between sprouting and the diameter of the girdledtrees or the distance to their nearest neighbour. A negativerelationship between the diameter of the girdled tree and thestage of decline was recognized. Girdling is an economic alternativeto non-commercial thinning in oak stands and can be seen asan effective structural enrichment treatment.     

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.     

Stomatal and leaf growth responses to partial drying of root tips in willow

Root tips of intact willow (Salix dasyclados Wimm., Clone 81-090) plants were partially dried by exposure to ambient greenhouse air and then kept in water-vapor-saturated air for up to 3 days. The drying treatment increased abscisic acid (ABA) concentrations in both the root tips subjected to drying and in the xylem sap, while it reduced leaf stomatal conductance and leaf extension rate. Despite the decrease in stomatal conductance, leaf water potentials were unaffected by the root drying treatment, indicating that the treatment reduced hydraulic conductivity between roots and foliage. After roots subjected to drying were returned to a nutrient solution or excised, ABA concentrations in the remaining roots and in the xylem sap, stomatal conductance of mature leaves and extension rate of unfolding leaves all returned to values observed in control plants. The 4-fold increase in xylem sap ABA concentration following the root drying treatment was not solely the result of reduced sap flow, and thus may be considered a potential cause, not merely a consequence, of the observed reduction in stomatal conductance.     

Response of Japanese red pine to inoculation with a blue stain fungus,Ceratocystis piceae

To confirm the pathogenicity of a blue stain fungus,Ceratocystis piceae (Münch) Bakshi to the Japanese red pine (Pinus densiflora Sieb. et Zucc.), the responses of healthy young pine trees and stressed trees which were girdled by the half-circumferential girdling technique were investigated by the fungal inoculation test. Although neither of the pine trees inoculated withC. piceae in the non-girdled treatment nor the controls died, mortality of the trees girdled and inoculated withC. piceae was 28.6%. In the pine trees inoculated withC. piceae, the mean area of the necrotic lesion of the sapwood was significantly larger than that of the controls, and the mean of the water pressure potential of the xylem decreased, regardless of the girdling treatment. TheC. piceae was reisolated from the wood pieces near the inoculation points on the inoculated trees, but not from the controls. These results suggest that under strongly stressed conditions, the Japanese red pine trees might have been killed by heavy infestations ofC. piceae carried by bark beetles. A part of this paper was presented at the 103rd Annual Meeting of the Japanese Forestry Society (1992).     

Variation in seedling hydraulic architecture as a function of species and environment

Northern red oak (Quercus rubra L.) and yellow-poplar (Liriodendron tulipifera L.) were grown for two years in full sunlight (unshaded) or 20% of full sunlight (shaded) under either well-watered or drought conditions. There was a close association between evaporative flux (in situ) and leaf-specific conductivity (LSC) in stem segments of both species. Shaded, drought-stressed seedlings of both species had significantly reduced leaf area, evaporative flux, volume flow rate in xylem, flow velocity, potentially functional xylem area, and LSC than unshaded, well-watered seedlings. Reductions in LSC associated with drought or shade were similar for both species; and within a treatment, both species had similar LSC. Species differed in the manner of LSC adjustment to drought and shade. Reductions in leaf area associated with drought or shade were accompanied primarily by reductions in potentially functional xylem area for L. tulipifera, and by reductions in flow velocity for Q. rubra. These results suggest (1) the existence of a homeostatic balance between evaporative flux and LSC, (2) that species with widely different growth patterns and xylem anatomies may develop similar LSC within the same environment, and (3) a possible hydraulic basis for differences in habitat between ring- and diffuse-porous species.