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Abscisic acid is a key inducer of hydrogen peroxide production in leaves of maize plants exposed to water stress.

The histochemical and cytochemical localization of water stress-induced H(2)O(2) production in the leaves of ABA-deficient vp5 mutant and wild-type maize (Zea mays L.) plants were examined, using 3,3-diaminobenzidine and CeCl(3) staining, respectively, and the roles of endogenous ABA in the production of H(2)O(2) induced by water stress were assessed. Water stress induced by polyethylene glycol resulted in the accumulation of H(2)O(2) in mesophyll cells, bundle-sheath cells and vascular bundles of wild-type maize leaves, and the accumulation was substantially blocked in the mutant maize leaves exposed to water stress. Pre-treatments with several apoplastic H(2)O(2) manipulators abolished the majority of H(2)O(2) accumulation induced by water stress in the wild-type leaves. The subcellular localization of H(2)O(2) production was demonstrated in the cell walls, xylem vessels, chloroplasts, mitochondria and peroxisomes in the leaves of wild-type maize plants exposed to water stress, and the accumulation of H(2)O(2) induced by water stress in the cell walls and xylem vessels, but not in the chloroplasts, mitochondria and peroxisomes, was arrested in the leaves of the ABA mutant or the ABA biosynthesis inhibitor (tungstate)-pre-treated maize plants. Pre-treatments with the apoplastic H(2)O(2) manipulators also blocked the apoplastic but not the intracellular H(2)O(2) accumulation induced by water stress in the leaves of wild-type plants. These data indicate that under water stress, the apoplast is the major source of H(2)O(2) production and ABA is a key inducer of apoplastic H(2)O(2) production. These data also suggest that H(2)O(2) generated in the apoplast could not diffuse freely into subcellular compartments.

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Radial velocity profiles of water flow in trunks of Norway spruce and oak and the response of spruce to severing.

Trunk-tissue heat balance, volumetric and staining methods were used to study xylem water flow rates and pathways in mature Norway spruce (Picea abies (L.) Karst.) and pedunculate oak (Quercus robur L.) trees. The radial profile of flow velocity was confirmed to be symmetrical in spruce, i.e., maximum flow velocity was in the center of the conducting xylem and tailed with low amplitude (about 30 cm h(-1)) in the direction of the cambium and heartwood. Variability around the trunk was high. In contrast, in oak, the radial profile of flow velocity was highly asymmetrical, reaching a peak of about 45 m h(-1) in the youngest growth ring and tailing centripetally for about 10 rings, but variability around the trunk was less, under non-limiting soil water conditions, than in spruce. In spruce, the flow rate increased abruptly within seconds when the tree was severed while immersed in water, and then decreased gradually, showing significant root resistance. We conclude that water flow through an absorbing cut surface differs from the flow higher in a tree trunk because of the presence of hydraulic capacitances in the conductive pathways. The staining technique always yielded higher estimates of flow velocity than the non-destructive tree-trunk heat balance method.

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Crown architecture of stand-grown sugar maple (Acer saccharum Marsh.) in the Adirondack Mountains.

Leaf and crown morphology of shade-tolerant sugar maple (Acer saccharum Marsh.) were examined to test the hypotheses (1) that leaf area exhibits significant plasticity both within and between crown classes and individual tree crowns and (2) that leaf area is accurately predicted from estimates of crown volume. A total of 18 trees, ranging from 3.3 to 43.4 cm dbh, were felled and dissected into upper, middle, lower, and below-crown layers, for measurements of leaf, bark, and xylem dimensions. For dominant trees only, bark thickness and xylem radii were higher within the crown than below the crown. Cumulative leaf area index increased with decreasing stratum height at similar rates in all trees, except for two trees that were located in the understory. Area leaf weight declined with decreasing stratum height within the crown of all except four overstory trees. These four trees showed an increase with decreasing stratum height, i.e., leaves were heavier per unit area in the lower crown stratum and below the crown than they were at mid-crown. Within-tree leaf area density was usually higher in the upper crown of overstory trees and in the lower crown of understory trees. Total crown volume was the best predictor of whole-tree leaf area, but it was only slightly better than dbh.

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Vascular tissue anatomy of Norway spruce needles and twigs in relation to magnesium deficiency.

Vascular tissues of needles and twigs of Norway spruce with low foliar magnesium concentrations were examined by light microscopy. Observations were made on samples from (1) apparently healthy trees, (2) trees exhibiting progressive symptoms of decline, including the yellowing and fall of needles and the death of twigs and branches in the subapical canopy, and (3) trees recovering from symptoms of decline after fertilization with magnesium sulfate. Abnormalities in cambium and phloem anatomy, which were apparent in 2-year-old needles of declining trees, were seen only in needles at least 4 years old in healthy trees, suggesting that needles of declining trees are susceptible to precocious aging. Abnormalities in xylem anatomy were observed in needles 3 years or more in age in declining trees, but were never seen in needles of healthy trees. Over time, anatomical abnormalities in declining trees were seen in progressively younger twigs, which may explain the acropetal development of decline symptoms. After fertilization with magnesium sulfate, normal phloem and xylem were observed in the newly formed tissues.

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Influence of climate and fertilization on net photosynthesis of mature slash pine.

Net photosynthesis was measured under field conditions in 23-year-old slash pine (Pinus elliottii Engelm. var. elliottii) trees to determine how it was affected by fertilization and climate. There was only a small decrease in rates of net photosynthesis from late summer through winter demonstrating that appreciable carbon gain occurs throughout the year in slash pine. Although fertilization substantially increased leaf area and aboveground biomass, it only slightly increased the rate of net photosynthesis. Simultaneous measurements of gas exchange in fertilized and unfertilized (control) plots allowed the detection of a small, but statistically significant difference in average net photosynthesis of 0.14 micro mol m(-2) s(-1). Irradiance, and to a lesser extent air temperature, were the environmental factors that exerted the most control on net photosynthesis. The highest rates of net photosynthesis occurred between air temperatures of 25 and 35 degrees C. Because air temperatures were within this range for 46% of all daylight hours during the year, air temperature was not often a significant limitation. Soil and atmospheric water deficits had less effect on photosynthesis than irradiance or air temperature. Although the depth to the water table changed during the year from 10 to 160 cm, predawn and midday xylem pressure potentials only changed slightly throughout the year. Predawn values ranged from -0.63 to -0.88 MPa in the control plot and from -0.51 to -0.87 MPa in the fertilized plot and were not correlated with water table depth. There was no correlation between xylem pressure potentials and net photosynthesis, presumably because water uptake was adequate. Although vapor pressure deficits reached 3.5 kPa during the summer, they had little effect on net photosynthesis. Over a vapor pressure deficit range from 1.0 to 3.0 kPa, net photosynthesis only decreased 21%. No differences in responses to these environmental factors could be attributed to fertilization.

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Influence of water relations on Quercus cerris-Hypoxylon mediterraneum interaction: a model of drought-induced susceptibility to a weakness parasite.

The association between water stress and susceptibility of Quercus cerris to the fungal parasite Hypoxylon mediterraneum was studied in field-grown trees and greenhouse-grown seedlings. Susceptibility of Q. cerris to the fungus, expressed as extension of wood discolorations, increased in field-grown trees during drought and increased in greenhouse-grown seedlings in response to limited water supply. In both seedlings and trees, spread of the fungus in the vascular system was higher when leaf water potentials were low than when leaf water potentials were high, and was significantly correlated to loss of hydraulic conductivity of xylem. The presence of the fungus in the xylem suggests that H. mediterraneum could use embolized vessels to spread in the host.

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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.

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Canopy dynamics and the morphological development of Abies balsamea: effects of foliage age on specific leaf area and secondary vascular development.

Data were collected from two branches from each whorl of nine open-grown Abies balsamea (L.) Miller trees to test the hypothesis that specific leaf area (SLA, m(2) projected fresh leaf area kg(-1) oven-dry foliage) is constant among five foliage age classes (current-year, 1-year-old, 2-year-old, 3-year-old and 4-year-old-plus). Between-tree variation in SLA was greater than within-tree variation. Differences in SLA among the foliage age classes were small, but statistically significant, showing a trend of decreasing SLA with increasing foliage age. Using data from two previous biomass studies, we found that three different methods of calculating SLA of individual trees produced the same projected leaf area estimates. To test the hypothesis that foliage mass increases with foliage age as a result of secondary xylem or phloem development, we examined the secondary vascular development of foliage collected from five age classes and three crown sections in an open-grown A. balsamea. The number of rows of xylem cells was not constant among foliage age classes, but the differences were small and showed no consistent pattern of change with foliage age. Total number of rows of phloem cells increased, number of living rows of phloem cells decreased, and the number of rows of nonliving crushed phloem cells increased with foliage age.

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Carbon isotope variation in Douglas-fir foliage: improving the delta(13)C-climate relationship.

The natural abundance of stable carbon isotopes in the annual rings of forest trees is used as a tracer of environmental changes such as climate and atmospheric pollution. Although tree-ring delta(13)C varies by about 2 per thousand from year to year, variability within the foliage can be as high as 6 per thousand. Recent studies have shown that branch length affects stomatal response, which influences the integrated foliar delta(13)C signal. To improve the ability of delta(13)C to predict climate differences, we examined the relationship between branch length and foliar delta(13)C in Pseudotsuga menziesii (Mirb.) Franco from four sites across a steep climate gradient in Oregon. The transect spanned the boundary between the ranges of the coastal variety, P. menziesii var. menziesii (three sites), and the Rocky Mountain variety, P. menziesii var. glauca (one site). At the most maritime site, branch length explained 76% of within-site variation of 5 per thousand, whereas at the harshest site, branch length accounted for only 15% of this variation. We considered the possibility that cavitation in the water-conducting xylem obscures the branch length effect in the harsher climates. Cavitation, as measured by dye perfusion, was most extensive at sites where the branch length effect in the coastal variety was weakest. Trees at the site with the most substantial cavitation displayed seasonal xylem refilling. Branch length standardization significantly improved the relationship between delta(13)C and climate. With standardization to constant length, delta(13)C values were significantly related to the degree that climatic variables, as modeled with a forest growth simulation model, constrain transpiration (R(2) = 0.69, P < 0.0001). Without standardization, the R(2) was 0.27. We conclude that sampling standard length branches or tree rings from trees of similar shape and size is desirable when seeking correlations between isotopic composition and climate.

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Transport and metabolism of gibberellins in relation to flower bud differentiation in Norway spruce (Picea abies).

A mixture of tritiated and deuterated gibberellin A(4) (GA(4)) was injected into the xylem of Norway spruce (Picea abies (L.) Karst) propagules, below an elongating shoot, or applied directly on the needles of an elongating shoot. The distribution of [(2)H(2)]GA(4) and [(3)H]GA(4) in the needles, stems and buds was determined after 4, 12 and 24 h. After 4 h, most of the xylem-injected GA(4) was found in the needles, whereas after 24 h, most of the GA(4) was found in the stem, with a small portion in the lateral buds. Of the GA(4) applied to the needles, 51% of the radioactivity recovered after 24 h was found in the stem and 2% in the lateral buds. Mixtures of tritiated and deuterated GA(4) and GA(9) were injected into elongating shoots of one abundant-flowering family and one limited-flowering family, grown either under conditions inductive for flowering (hot and dry, HD) or under noninductive conditions for flowering (cool and wet, CW). Shoots of both CW- and HD-treated propagules converted [(2)H(2)]GA(9) to [(2)H(2)]GA(51), [(2)H(2)]GA(4), [(2)H(2)]GA(34) and [(2)H(2)]GA(1), whereas [(2)H(2)]GA(4) was converted to [(2)H(2)]GA(34), [(2)H(2)]GA(1) and [(2)H(2)]GA(8). In shoots of both CW-treated clones, the main metabolite of [(3)H]GA(9) was in the GA(51) region. The HD-treated propagules converted more [(3)H]GA(9) to putative GA(4) than the CW-treated propagules. The main metabolite of [(3)H]GA(4) was in the GA(34) region. Radioactive metabolites were also found in the GA(1) and GA(8) regions.

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Sap flow in Bornean heath and dipterocarp forest trees during wet and dry periods.

Daytime xylem sap flow was measured by the compensation heat pulse velocity technique in dipterocarp forest trees growing on sandy loam (haplic acrisol) and in heath forest trees growing on bleached sand (albic arenosol) in Brunei during wet and dry periods. In dipterocarp forest trees, daily sap flow was stable during the dry period but declined on rainy days presumably as a result of reduced transpiration. At the heath site, daily sap flow of the small trees was stable during the dry period, whereas it declined in the large trees as the dry season progressed. In some trees at both sites, the hydroactive area of xylem was less during the dry period than during the wet period. Daily sap flow was linearly related to projected crown area, and a common regression fitted trees of different species from both forest types. The dry/wet period ratio of sap flow (about 0.7) was statistically equivalent for heath and dipterocarp forest trees, whereas it had been expected that trees on the freely draining arenosol would suffer greater water stress during drought. For well-watered trees >/= 0.2 m dbh (diameter at 1.3 m above ground) in both the dipterocarp and heath forests, stand water use was estimated to be 40 mm month(-1).

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Hydraulic properties of Douglas-fir (Pseudotsuga menziesii) branches and branch halves with reference to compression wood.

Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) branch segments were used to test the hypothesis that compression wood reduces xylem transport efficiency. Whole 3-year-old segments were first measured for specific conductivity (k(s), m(2) s(-1) MPa(-1)), then split lengthwise into upper and lower halves, the latter containing all or most of the compression wood in the segment. Halves were then remeasured for k(s) using a new technique that prevents leakage of permeating fluid during measurements. Lower branch halves had significantly lower k(s) than upper halves (6.4 +/- 0.3 versus 9.3 +/- 0.3 m(2) s(-1) MPa(-1) x 10(-4), respectively; n = 36), and despite their larger size, significantly lower hydraulic conductivity (k(h), m(4) s(-1) MPa(-1)) than upper halves. Lower branch halves had higher specific gravity (0.51 +/- 0.01 versus 0.45 +/- 0.01; n = 36), lower water content (123 +/- 2% versus 155 +/- 3%; n = 36), and larger proportions of volume occupied by both cell wall and air than upper halves. Lower halves had more tracheids per annual ring than upper halves (73 +/- 3 versus 63 +/- 2 per radial transect, respectively; n = 36), but tracheids were shorter and had narrower lumens than those of upper branch halves. Differences in hydraulic properties between upper and lower halves suggest that compression wood does reduce xylem transport efficiency. In contrast, the amount of compression wood in each sample did not explain any variation in whole unsplit sample hydraulic properties.

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Effect of nitrogen on the seasonal course of growth and maintenance respiration in stems of Norway spruce trees.

To determine effects of stem nitrogen concentration ([N]) on the seasonal course of respiration, rates of stem respiration of ten control and ten irrigated-fertilized (IL), 30-year-old Norway spruce trees (Picea abies (L.) Karst.), growing in northern Sweden, were measured on seven occasions from June 1993 to April 1994. To explore sources of seasonal variation and mechanisms of fertilization effects on respiration, we separated total respiration into growth and maintenance respiration for both xylem and phloem bark. Stem respiration increased in response to the IL treatment and was positively correlated with growth rate, volume of living cells and stem nitrogen content. However, no significant effect of IL treatment or [N] in the living cells was found for respiration per unit volume of live cells. Total stem respiration during the growing season (June to September) was estimated to be 16.7 and 29.7 mol CO(2) m(-2) for control and IL-treated trees, respectively. Respiration during the growing season accounted for approximately 64% of total annual respiration. Depending on the method, estimated growth respiration varied between 40 and 60% of total respiration during the growing season. Between 75 and 80% of the live cell volume in the stems was in the phloem, and phloem maintenance accounted for about 70% of maintenance respiration. Because most of the living cells were found in the phloem, and the living xylem cells were concentrated in the outer growth rings, we concluded that the best base for expressing rates of stem growth and maintenance respiration in young Norway spruce trees is stem surface area.

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Ethylene evolution, radial growth and carbohydrate concentrations in Abies balsamea shoots ringed with Ethrel.

The terminal (1-year-old) shoot of quiescent, 2-year-old Abies balsamea (L.) Mill. seedlings was either left untreated or ringed with 0, 1 or 10 mg Ethrel g(-1) lanolin. After 5 weeks of culture under environmental conditions favorable for growth, the shoots were harvested to measure ethylene evolution and carbohydrate concentrations by gas chromatography, and tracheid number and bark radial width by microscopy. In untreated shoots, the basal rate of ethylene evolution followed the order: cambial region > cortex + periderm = xylem + pith = needles. Wound-induced ethylene production was not detected until at least 4 h after excision, but was evident in all fractions 24 h after excision; the increase in wound-induced ethylene evolution followed the order: cambial region > cortex + periderm > xylem + pith > needles. Compared with untreated controls, the application of plain lanolin, which involved the removal of needles and periderm, increased bark radial width and wound-induced ethylene production by the cambial region and the cortex + periderm, but decreased cambial region concentrations of fructose, glucose and starch at the application point. At the application point, Ethrel concomitantly increased ethylene evolution from the cambial region and the cortex + periderm, tracheid number, bark radial width, and the cambial region concentrations of fructose, glucose, sucrose and starch. No effects of Ethrel treatment were detected above or below the application point, with the exception that the 10 mg g(-1) Ethrel treatment stimulated ethylene evolution and decreased starch concentration of the cambial region. The results indicate that: (1) the cambial region is the major source of endogenous ethylene in the 1-year-old shoot; (2) the magnitude of the difference in ethylene evolution between particular shoot fractions is different before and after the start of wound-induced ethylene production; (3) the Ethrel-induced increase in tracheid number and bark radial width at the application point is positively related to ethylene evolution from the cambial region and the cortex + periderm, respectively; and (4) ethylene derived from Ethrel applied laterally to a woody stem can mobilize carbohydrates to the application point.

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Transport, storage and mobilization of nitrogen by trees and shrubs in the wet/dry tropics of northern Australia.

Xylem sap from woody species in the wet/dry tropics of northern Australia was analyzed for N compounds. At the peak of the dry season, arginine was the main N compound in sap of most species of woodlands and deciduous monsoon forest. In the wet season, a marked change occurred with amides becoming the main sap N constituents of most species. Species from an evergreen monsoon forest, with a permanent water source, transported amides in the dry season. In the dry season, nitrate accounted for 7 and 12% of total xylem sap N in species of deciduous and evergreen monsoon forests, respectively. In the wet season, the proportion of N present as nitrate increased to 22% in deciduous monsoon forest species. These results suggest that N is taken up and assimilated mainly in the wet season and that this newly assimilated N is mostly transported as amide-N (woodland species, monsoon forest species) and nitrate (monsoon forest species). Arginine is the form in which stored N is remobilized and transported by woodland and deciduous monsoon forest species in the dry season. Several proteins, which may represent bark storage proteins, were detected in inner bark tissue from a range of trees in the dry season, indicating that, although N uptake appears to be limited in the dry season, the many tree and shrub species that produce flowers, fruit or leaves in the dry season use stored N to support growth. Nitrogen characteristics of the studied species are discussed in relation to the tropical environment.

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Seasonal changes in photosynthesis of trees in the flooded forest of the Mapire River.

We studied the flood tolerance of five tree species growing in the flooded forest adjacent to the Mapire river, in SW Venezuela. Mean photosynthetic rate and leaf conductance were 11 &mgr;mol m(-2) s(-1) and 700 mmol m(-2) s(-1), respectively. Xylem water potential ranged from -0.08 to -1.15 MPa. Based on leaf gas exchange as a criterion of tolerance to flooding, two response patterns were identified: (1) decreasing photosynthetic rate with increasing flooding and leaf conductance (Psidium ovatifolium Berg. ex Desc., Campsiandra laurifolia Benth., Symmeria paniculata Benth. and Acosmium nitens (Vog.) Benth); and (2) independence of photosynthesis and leaf conductance from flooding (Eschweilera tenuifolia (Berg.) Miers.). In the first response pattern, declining photosynthetic rate with flooding may be interpreted as a sign of reduced flood tolerance, whereas the second response pattern may indicate increased flood tolerance. An increase in xylem water potential with depth of water column was found for all species (with the possible exception of P. ovatifolium), indicating that flooding does not cause water stress in these trees. Submerged leaves that had been under water for between four days and four months generally had photosynthetic rates and leaf conductances similar to those of aerial leaves, indicating maintenance of photosynthetic capacity under water. Daily positive oscillations in glucan content in submerged leaves of P. ovatifolium and C. laurifolia suggest that submerged leaves do not represent a sink for photosynthates produced by aerial leaves.

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Thinning, fertilization, and crown position interact to control physiological responses of loblolly pine.

To examine physiological responses to thinning, fertilization, and crown position, we measured net photosynthesis (P(n)), transpiration (E), vapor pressure difference (VPD), stomatal conductance (g(s)), and xylem pressure potential (Psi(1)) between 0930 and 1130 h under ambient conditions in the upper and lower crowns of a 13-year-old loblolly pine (Pinus taeda L.) plantation six years (1994) after the treatments were applied. Photosynthetic photon flux density (PPFD) and air temperature (T(a)) within the canopy were also recorded. Needle P(n) of thinned trees was significantly enhanced by 22-54% in the lower crown, because canopy PPFD increased by 28-52%. Lower crown foliage of thinned plots also had higher E and g(s) than foliage of unthinned plots, but thinning had no effect on needle Psi(1) and predawn xylem pressure potential (0430-0530 h; Psi(pd)). Tree water status did not limit P(n), E and g(s) during the late-morning measurements. Fertilization significantly decreased within-canopy PPFD and T(a). Needle Psi(1) was increased in fertilized stands, whereas P(n), E and g(s) were not significantly altered. Upper crown foliage had significantly greater PPFD, P(n), VPD, g(s), E, and more negative Psi(1) than lower crown foliage. In both crown positions, needle P(n) was closely related to g(s), PPFD and T(a) (R(2) = 0.77 for the upper crown and 0.82 for the lower crown). We conclude that (1) silvicultural manipulation causes microclimate changes within the crowns of large trees, and (2) needle physiology adjusts to the within-crown environmental conditions.

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Analyses of assumptions and errors in the calculation of stomatal conductance from sap flux measurements.

We analyzed assumptions and measurement errors in estimating canopy transpiration (E(L)) from sap flux (J(S)) measured with Granier-type sensors, and in calculating canopy stomatal conductance (G(S)) from E(L) and vapor pressure deficit (D). The study was performed in 12-year-old Pinus taeda L. stands with a wide range in leaf area index (L) and growth rate. No systematic differences in J(S) were found between the north and south sides of trees. However, J(S) in xylem between 20 and 40 mm from the cambium was 50 and 39% of J(S) in the outer 20-mm band of xylem in slow- and fast-growing trees, respectively. Sap flux measured in stems did not lag J(S) measured in branches, and time and frequency domain analyses of time series indicated that variability in J(S) in stems and branches is mostly explained by variation in D. Therefore, J(S) was used to estimate transpiration, after accounting for radial patterns. There was no difference between D and leaf-to-air vapor pressure gradient, and D did not have a vertical profile in stands of either low or high L suggesting a strong canopy-atmosphere coupling. Therefore, D estimated at one point in the canopy can be used to calculate G(S) in such stands. Given the uncertainties in J(S), relative humidity, and temperature measurements, to keep errors in G(S) estimates to less than 10%, estimates of G(S) should be limited to conditions in which D >/= 0.6 kPa.

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