Effects of duration of a simulated winter thaw on dieback and xylem conductivity of Betula papyrifera

Stems or roots + stems of potted, 2-year-old paper birch (Betula papyrifera L.) were subjected to simulated winter thaws of various durations in climate-controlled chambers. The simulated thaws induced dieback of shoots of the treated plants. Although the stem thaw treatment did not significantly increase dieback, there were significant (P < 0.05) correlations between growing degree days above 4 degrees C and both shoot dieback and percent reduction in conductive xylem. All trees that received > 60 growing degree days (GDD) > 4 degrees C died back to some extent. Plants in the root + stem thaw treatment that received more than 60 GDD > 4 degrees C showed a significant (P < 0.05) increase in dieback and a significant (P < 0.05) loss of conducting xylem after a period of growth and recovery in the greenhouse, especially in the xylem of 1-year-old stems. Furthermore, higher correlations between GDD > 4 degrees C during a thaw and both the extent of dieback and the loss in conductive xylem were found in trees subjected to the root + stem thaw treatments than in trees exposed only to the stem thaw treatments (P < 0.05). The root + stem thaw treatments also resulted in highly significant relationships (P < 0.05-0.001) between loss in conductive xylem and dieback. The occurrence of dieback in response to winter thaws, and its close correlation with irreversible losses of xylem conductivity due to embolisms, coupled with an inability to refill the xylem because of root damage, support the view that these processes may be key factors in initiating birch decline.     

Chlorophyll fluorescence and CO(2) assimilation of black spruce seedlings following frost in different temperature and light conditions

Effects of artificial frosts on light-saturated photosynthesis (A(max)) and ground, maximal and variable fluorescence variables (F(o), F(m), and F(v) and F(v)/F(m)) were monitored on 1-year-old foliage of black spruce seedlings (Picea mariana (Mill.) BSP) grown at high (25 degrees C), moderate (15 degrees C) and low (5 degrees C) temperatures and moderate (240 &mgr;mol m(-2) s(-1)) and low (80 &mgr;mol m(-2) s(-1)) irradiances. Photoinhibition of 1-year-old foliage was greater in seedlings grown in moderate light than in seedlings grown in low light. Photoinhibition increased with decreasing growth chamber temperature at both irradiances. Most changes in F(v)/F(m) were caused by changes in F(v). Exposure to -4 degrees C decreased both F(v)/F(m) and A(max) compared with control values. The effect of the -4 degrees C frost treatment was greater in seedlings grown in low light than in seedlings grown in moderate light, probably because seedlings grown in moderate light were already partially photoinhibited before the frost treatment. Following -4 degrees C treatment, neither F(v)/F(m) nor A(max) recovered in seedlings grown in low light. Light-saturated photosynthesis decreased with decreasing growth chamber temperature. Light-saturated photosynthesis was more sensitive to the -3 and -4 degrees C frost treatments in seedlings grown at 25 degrees C than in seedlings grown at 15 and 5 degrees C. The A(max) of seedlings grown at 15 degrees C was sensitive only to the -4 degrees C frost treatment, whereas A(max) of seedlings grown at 5 degrees C was not sensitive to any of the frost treatments. Recovery of A(max) following frost took longer in seedlings grown at high temperatures than in seedlings grown at low temperatures. For seedlings grown at the same temperature but under different irradiances, both A(max) and F(v)/F(m) reflected damage to the photosynthetic system following a moderate frost. However, for seedlings grown at the same irradiance but different temperatures, A(max) provided a more sensitive indicator of frost damage to the photosynthetic system than F(v)/F(m) ratio.     

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 simulated thaw on xylem cavitation,residual embolism,spring dieback and shoot growth in yellow birch

Yellow birch seedlings (Betula alleghaniensis Britt.) that had lost more than 90% of their stem hydraulic conductivity during ambient winter temperatures were exposed to 0 and 20 days of a simulated winter thaw followed by a 48-h freezing treatment at 0, -5, -10, -20 and -30 degrees C. After measuring freezing injury to shoots and roots, the seedlings were placed in a greenhouse where recovery of xylem conductivity and new growth were measured. Shoot xylem cavitation was measured as percent loss of hydraulic conductivity. Shoot freezing injury was assessed by electrolyte leakage (EL) and root freezing injury was assessed by EL and triphenyl tetrazolium chloride reduction. Seedlings pretreated with thaw had higher stem water contents and suffered more freezing damage to roots and shoots (at -20 and -30 degrees C, respectively) than unthawed seedlings. After 3 weeks in a greenhouse, seedlings from the 0, -5 and -10 degrees C freezing treatments showed complete recovery of xylem conductivity, with substantially increased stem water contents. Poor recovery of hydraulic conductivity was observed only in seedlings that were subjected to freezing treatments at -20 and -30 degrees C, regardless of thaw treatment. Of these embolized seedlings, however, only those not previously thawed showed recovery of hydraulic conductivity or regained stem water content after 9 weeks in the greenhouse. Shoot dieback, bud burst and length of new shoots were significantly related to the extent of stem xylem cavitation and freezing injury. We conclude that (1) the simulated winter thaw predisposed yellow birch seedlings to freezing damage in shoots and roots by dehardening tissues and increasing their water content; (2) root freezing damage in turn affected the seedlings' ability to refill embolized stem xylem, resulting in considerable residual xylem embolism after spring refilling; (3) further recovery of stem xylem conductivity was attributable to growth of new vessels; (4) and the permanent residual embolism, together with root and shoot freezing injury, caused increased dieback, bud mortality and reduced growth of new shoots.     

Assimilation and stomatal conductance responses of red spruce to midwinter frosts and the constituent ions of acid mist

Red spruce (Picea rubens Sarg.) seedlings growing outside in open-top chambers were sprayed twice weekly with artificial mists at either pH 2.5 or 5.6, for five months during the 1988 growing season. The mists contained one of the following: water, pH 5.6 (control); (NH(4))(2)SO(4), pH 5.6; NH(4)NO(3), pH 5.6; HNO(3), pH 2.5; H(2)SO(4), pH 2.5; or (NH(4))(2)SO(4) + NH(4)NO(3), pH 2.5. During January 1989, the light responses of assimilation and stomatal conductance were assessed in the laboratory following a 4-day equilibration at 12 degrees C. The aerial portions of the intact trees were then subjected to a mild (-10 degrees C) frost for three hours during the night and the rate of recovery of light-saturated assimilation (A(max)) was determined the following day using the same branches as were used for the assimilation studies before the frost treatment. The same trees were then subjected to a second frost of -18 degrees C for three hours during the following night and the recovery of A(max) of the same branches was measured the next day. All of the acid mist treatments increased A(max) and apparent quantum yield relative to the control treatment when measured before the frost treatments. Frosts of -10 and -18 degrees C resulted in a significant decline in A(max) of seedlings in all treatments except the control. Stomatal conductance increased with increasing irradiance in seedlings in the acid mist treatments that did not contain SO(4) (2-) ion. Stomatal conductance of seedlings in acid mist treatments containing SO(4) (2-) ion was insensitive to changes in irradiance over the range 50-1500 micro mol m(-2) s(-1). It is concluded that acid precipitation increased the sensitivity of the assimilation response to midwinter frosts that follow a brief warm period. The SO(4) (2-) ion appears to be significant in causing increased sensitivity to frost and in causing stomatal insensitivity to light flux density.     

Soil temperature,gas exchange and nitrogen status of 5-year-old Norway spruce seedlings

Five-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment chambers. Plants were acclimated to a soil temperature of 16 degrees C during the first and third growing seasons, but were allocated at random to soil temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season. Low soil temperature during the second growing season depressed stomatal conductance and photosynthetic rate (A) per unit of projected leaf area, although intercellular CO2 concentrations did not differ significantly between treatments. At all soil temperatures, total chlorophyll concentration first decreased and then increased, although the rate of increase and the final concentration increased with soil temperature, which may explain the effect of soil temperature on A. Neither chlorophyll a/b ratio nor leaf nitrogen concentration was significantly affected by soil temperature. Treatment differences disappeared during the third simulated growing season when plants were again acclimated to a soil temperature of 16 degrees C.     

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.     

Frost hardiness of Picea rubens growing in spruce decline regions of the Appalachians

It has been proposed that pollutants predispose Picea rubens Sarg. growing in the high Appalachians to frost damage. The pattern of autumn hardening of P. rubens growing at Whiteface Mountain, NY, and Newfound Gap, NC, was monitored by detaching shoots at 1-3 weekly intervals, air freighting them to Scotland, and freeze-testing them. The temperatures that produced freezing injury from August 1986 to January 1987 were compared with minimum air temperatures recorded in those months at nearby meteorological stations over 22 previous years. There was only weak evidence that the onset or degree of frost hardening was inadequate to protect the trees from direct freezing injury (as opposed to winter desiccation). Historically, minimum air temperatures occasionally fell below the lethal temperature for a 10% kill (LT(10)), but they rarely fell below the LT(50). The trees hardened rapidly in the autumn (max. 2.2 degrees C day(-1)) to between -30 degrees C and -40 degrees C by January (LT(50)), including trees showing visible decline on Clingman's Dome, TN. Individual trees differed in hardiness by up to 10 degrees C. It is concluded that any pollutant-induced susceptibility to freezing injury is insufficient, on its own, to account for forest decline in the Appalachians.     

Planting stress in newly planted jack pine and white spruce. 2. Changes in tissue water potential components

Water relations of bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) planted in a greenhouse and on a boreal cut-over site were examined during the first growing season. In field-planted trees, maximum stomatal conductances (g(wv)) were initially low (< 0.10 cm s(-1)). Base and minimum xylem pressure potentials (Psi(x(base)) and Psi(x(min))) were less than -1.5 and -1.7 MPa for jack pine and -2.0 and -2.6 MPa for white spruce, respectively. During the growing season, maximum g(wv) increased in both species to around 0.2 cm s(-1). Base and minimum xylem pressure potentials also increased in both species to around -0.5 and -1.0 MPa in jack pine and -1.0 and -1.5 MPa in white spruce, respectively. Minimum xylem pressure potentials in white spruce fell below the turgor loss point during the first half of the growing season. Osmotic potential at the turgor loss point Psi(pi(TLP)) decreased after field planting to around -2.7 and -2.3 MPa in jack pine and white spruce, respectively. In the greenhouse, minimum values of Psi(pi(TLP)) were -2.2 and -2.3 MPa in jack pine and white spruce, respectively. Maximum bulk modulus of elasticity was greater in white spruce and underwent greater seasonal change than in jack pine. Relative water content (RWC) at turgor loss ranged between 71 and 74% in jack pine and 80 and 87% in white spruce. Available turgor (T(avail)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and xylem pressure potential at the turgor loss point, was similar in jack pine and white spruce just after field planting. For the rest of the growing season, however, T(avail) in jack pine was two to three times that in white spruce. Diurnal turgor (T(diurnal)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and Psi(x(min)), as a percent of T(avail) was higher in field-planted white spruce than jack pine until the end of the season. Dynamics of tissue water potential components are discussed in relation to plantation establishment.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Bole girdling affects metabolic properties and root, trunk and branch hydraulics of young ponderosa pine trees

Effects of trunk girdling on seasonal patterns of xylem water status, water transport and woody tissue metabolic properties were investigated in ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) trees. At the onset of summer, there was a sharp decrease in stomatal conductance (g(s)) in girdled trees followed by a full recovery after the first major rainfall in September. Eliminating the root as a carbohydrate sink by girdling induced a rapid reversible reduction in g(s). Respiratory potential (a laboratory measure of tissue-level respiration) increased above the girdle (branches and upper trunk) and decreased below the girdle (lower trunk and roots) relative to control trees during the growing season, but the effect was reversed after the first major rainfall. The increase in branch respiratory potential induced by girdling suggests that the decrease in g(s) was caused by the accumulation of carbohydrates above the girdle, which is consistent with an observed increase in leaf mass per area in the girdled trees. Trunk girdling did not affect native xylem embolism or xylem conductivity. Both treated and control trunks experienced loss of xylem conductivity ranging from 10% in spring to 30% in summer. Girdling reduced xylem growth and sapwood to leaf area ratio, which in turn reduced branch leaf specific conductivity (LSC). The girdling-induced reductions in g(s) and transpiration were associated with a decrease in leaf hydraulic conductance. Two years after girdling, when root-to-shoot phloem continuity had been restored, girdled trees had a reduced density of new wood, which increased xylem conductivity and whole-tree LSC, but also vulnerability to embolism.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Net CO(2) assimilation of cacao seedlings following dark chilling

Overnight exposure of cacao (Theobroma cacao L.) seedlings to chilling temperatures between 4.7 and 15.8 degrees C reduced net CO(2) assimilation rate (A) and stomatal conductance to water vapor (g(s)), with temperatures below 10 degrees C causing severe inhibition. Net CO(2) assimilation rates of chilled seedlings recovered to those of nonchilled plants within 7 days. No differences in daytime intercellular CO(2) concentration (c(i)) with overnight temperature were observed on the first day after the chilling treatment, which indicates that the reduction in photosynthesis was not caused by the reduction in stomatal conductance. However, c(i) of chilled plants was much less than that of nonchilled plants on the second day after treatment, which suggests that chilling caused a change in stomatal response to CO(2) concentration. Even 7 days after treatment, when A had recovered to control values, g(s) of chilled leaves was only approximately 70% that of controls. Chilling did not inhibit A through an effect on leaf water potential, which was higher in chilled plants than in unchilled plants.     

Planting stress in newly planted jack pine and white spruce. 1. Factors influencing water uptake

Bareroot jack pine (Pinus banksiana Lamb.) seedlings (2 + 0) and bareroot white spruce (Picea glauca (Moench) Voss) transplants (1 1/2 + 1 1/2) were taken from cold storage and planted on a clearcut forest site in northeastern Ontario on several dates between May 6 and June 5 during which period soil temperature at 15 cm depth increased from 0 to 18 degrees C. Additional cold-stored trees were transferred to a greenhouse where they were grown in pots for 0, 7 or 28 days and then placed with their roots in aerated water maintained at one of a range of constant temperatures between 0 and 22 degrees C. In both species, daytime xylem pressure potentials (Psi(x)) and needle conductances (g(wv)) decreased with decreasing soil or water temperature. At all root temperatures, g(wv) was lower, and Psi(x) higher, in jack pine than in white spruce. After 28 days in the greenhouse, g(wv) of jack pine seedlings, and Psi(x) of white spruce, was higher than in plants just removed from cold storage. In both species, water-flow resistance through the soil-plant-atmosphere continuum (RSPAC) increased as root temperature decreased. At all root temperatures, RSPAC was higher in plants just removed from cold storage than in plants grown in the greenhouse for 28 days, during which time many new unsuberized roots were formed. At root temperatures above 10 degrees C, RSPAC of both species was higher in trees newly planted in mineral soil than in trees with roots in aerated water; presumably because the roots of planted trees had limited hydraulic contact with the soil. On the day following removal from cold storage, relative plant water flow resistance increased, in both species, more rapidly with declining root temperature than could be accounted for by the change with temperature in the viscosity of water, thus indicating an effect of temperature on root permeability. The same effect was evident in jack pine seedlings, but not white spruce transplants, that had been grown for 28 days in the greenhouse after removal from cold storage.     

Impacts of seasonal air and soil temperatures on photosynthesis in Scots pine trees

Seasonal courses of light-saturated rate of net photosynthesis (A360) and stomatal conductance (gs) were examined in detached 1-year-old needles of Scots pine (Pinus sylvestris L.) from early April to mid-November. To evaluate the effects of soil frost and low soil temperatures on gas exchange, the extent and duration of soil frost, as well as the onset of soil warming, were manipulated in the field. During spring, early summer and autumn, the patterns of A360 and gs in needles from the control and warm-soil plots were generally strongly related to daily mean air temperatures and the frequency of severe frost. The warm-soil treatment had little effect on gas exchange, although mean soil temperature in the warm-soil plot was 3.8 degrees C higher than in the control plot during spring and summer, indicating that A360 and gs in needles from control trees were not limited by low soil temperature alone. In contrast, prolonged exposure to soil temperatures slightly above 0 degrees C severely restricted recovery of A360 and especially gs in needles from the cold-soil treatment during spring and early summer; however, full recovery of both A360 and gs occurred in late summer. We conclude that inhibition of A360 by low soil temperatures is related to both stomatal closure and effects on the biochemistry of photosynthesis, the relative importance of which appeared to vary during spring and early summer. During the autumn, soil temperatures as low as 8 degrees C did not affect either A360 or gs.     

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.)

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).     

Boundary layer conductance,leaf temperature and transpiration of Abies amabilis branches

We used three methods to measure boundary layer conductance to heat transfer (g(bH)) and water vapor transfer (g(bV)) in foliated branches of Abies amabilis Dougl. ex J. Forbes, a subalpine forest tree that produces clumped shoot morphology on sun-formed branches. Boundary layer conductances estimated in the field from energy balance measurements increased linearly from approximately 10 mm s(-1) at low wind speeds (< 0.1 m s(-1)) to over 150 mm s(-1) at wind speeds of 2.0 m s(-1). Boundary layer conductances measured on shoot models in a wind tunnel were consistently higher than field measurements. The difference between wind tunnel values and field measurements was attributable to variation in path length between the two experimental environments. Boundary layer conductance estimated by subtracting stomatal resistance (r(sV)) measured with a porometer from the total branch vapor phase resistance were unusually small. Sensitivity analysis demonstrated that this method is not suitable for coniferous foliage or when stomatal conductance (g(sV)) is small compared with g(bV). Analysis of the relative magnitudes of g(sV) and g(bV) revealed that, under most conditions, A. amabilis branches are well coupled (i.e., g(sV) is the dominant controller of transpiration). The boundary layer conductance to heat transfer is small enough that leaf temperature can become substantially higher than air temperature when radiation is high and wind speed is low. Over a two-month period, the maximum difference between leaf and air temperatures exceeded 6 degrees C. Leaf temperature exceeded air temperature by more than 2 degrees C on 10% of the daylight hours during this period. Consideration of both the photosynthetic temperature response of A. amabilis foliage as well as the summer air temperature conditions in its habitat suggests that these elevated leaf temperatures do not have a significant impact on carbon gain during the growing season.     

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.     

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 μmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.     

Short-term effects of potassium fertilization on the hydraulic conductance of Laurus nobilis L

This study reports experimental evidence on the effect of short-term potassium fertilization on potassium uptake, tissue concentration and hydraulic conductance of pot-grown laurel plants. Potassium uptake and loading into the xylem of laurel seedlings increased within 24 h after fertilization. Potassium was not accumulated in roots and leaves, but the [K(+)] of xylem sap was 80% higher in fertilized plants (+K) than in potassium-starved plants (-K), as a likely result of recirculation between xylem and phloem. Increased xylem sap [K(+)] resulted in a 45% increase in transpiration rate, a 30% increase in plant hydraulic conductance (K(plant)) and a 120% increase in leaf-specific conductivity of the shoot (k(shoot)). We suggest that this increase was due to ion-mediated up-regulation of xylem hydraulics, possibly caused by the interaction of potassium ions with the pectic matrix of intervessel pits. The enhancement of hydraulic conductance following short-term potassium fertilization is a phenomenon that can be of advantage to plants for maintaining cell turgor, stomatal aperture and gas exchange rates under moderate drought stress. Our data provide additional support for the important role of potassium nutrition in agriculture and forestry.