Showing papers in "Tree Physiology in 2002"

The influence of the forest canopy on nutrient cycling.

Abstract: Rates of key soil processes involved in recycling of nutrients in forests are governed by temperature and moisture conditions and by the chemical and physical nature of the litter. The forest canopy influences all of these factors and thus has a large influence on nutrient cycling. The increased availability of nutrients in soil in clearcuts illustrates how the canopy retains nutrients (especially N) on site, both by storing nutrients in foliage and through the steady input of available C in litter. The idea that faster decomposition is responsible for the flush of nitrate in clearcuts has not been supported by experimental evidence. Soil N availability increases in canopy gaps as small as 0.1 ha, so natural disturbances or partial harvesting practices that increase the complexity of the canopy by creating gaps will similarly increase the spatial variability in soil N cycling and availability within the forest. Canopy characteristics affect the amount and composition of leaf litter produced, which largely determines the amount of nutrients to be recycled and the resulting nutrient availability. Although effects of tree species on soil nutrient availability were thought to be brought about largely through differences in the decomposition rate of their foliar litter, recent studies indicate that the effect of tree species can be better predicted from the mass and nutrient content of litter produced, hence total nutrient return, than from litter decay rate. The greater canopy complexity in mixed species forests creates similar heterogeneity in nutritional characteristics of the forest floor. Site differences in slope position, parent material and soil texture lead to variation in species composition and productivity of forests, and thus in the nature and amount of litter produced. Through this positive feedback, the canopy accentuates inherent differences in site fertility.

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees

Abstract: We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt) Liebl) and beech (Fagus sylvatica L) trees during an annual cycle Radial distribution of carbohydrates was investigated according to ring age On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak Seasonal dynamics of TNC concentration were significantly (P < 005) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)) A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees The innermost sapwood rings of oak trees had low TNC concentrations Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees

Universal scaling in tree and vascular plant allometry: toward a general quantitative theory linking plant form and function from cells to ecosystems.

Abstract: A general theory of allometric scaling that predicts how the proportions of vascular plants and the characteristics of plant communities change or scale with plant size is outlined. The theory rests, in part, on the assumptions of (1) minimal energy dissipation in the transport of fluid through space-filling, fractal-like, branching vascular networks; and (2) the absence of scaling with plant size in the anatomical and physiological attributes of leaves and xylem. The theory shows how the scaling of metabolism with plant size is central to the scaling of whole-plant form and function. It is shown how allometric constraints influence plant populations and, potentially, processes in plant evolution. Rapidly accumulating evidence in support of the general allometric model is reviewed and new evidence is presented. Current work supports the notion that scaling of how plants utilize space and resources is central to the development of a general synthetic and quantitative theory of plant form, function, ecology and diversity.

Hydraulic redistribution of soil water during summer drought in two contrasting Pacific Northwest coniferous forests

Abstract: Summary The magnitude of hydraulic redistribution of soil water by roots and its impact on soil water balance were estimated by monitoring time courses of soil water status at multiple depths and root sap flow under drought conditions in a dry ponderosa pine (Pinus ponderosa Dougl. ex Laws) ecosystem and in a moist Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) ecosystem. The fate of deuterated water applied to small plots to create a strong horizontal soil water potential gradient was also monitored to assess the potential for horizontal redistribution of water and utilization of redistributed water by co-occurring shallow-rooted plants. In a 20-year-old Douglas-fir stand, approximately 28% of the water removed daily from the upper 2 m of soil was replaced by nocturnal hydraulic redistribution during late August. In an old-growth ponderosa pine stand, approximately 35% of the total daily water utilization from the upper 2 m of soil appeared to be replaced by hydraulic redistribution during July and August. By late September, hydraulic redistribution in the ponderosa pine stand was no longer apparent, even though total water use from the upper 2 m of soil was nearly identical to that observed earlier. Based on these results, hydraulic redistribution would allow 21 and 16 additional days of stored water to remain in the upper soil horizons in the ponderosa pine and Douglas-fir stands, respectively, after a 60-day drought. At both sites, localized applications of deuterated water induced strong reversal of root sap flow and caused soil water content to cease declining or even temporarily increase at locations too distant from the site of water application to have been influenced by movement of water through the soil without facilitation by roots. Xylem water deuterium values of ponderosa pine seedlings suggested utilization of redistributed water. Therefore, hydraulic redistribution may enhance seedling survival and maintain overstory transpiration during summer drought. These first approximations of the extent of hydraulic redistribution in these ecosystems suggest that it is likely to be an important process in both wet and dry forests of the Pacific Northwest.

Successional changes in live and dead wood carbon stores: implications for net ecosystem productivity

Abstract: If forests are to be used in CO2 mitigation projects, it is essential to understand and quantify the impacts of disturbance on net ecosystem productivity (NEP; i.e., the change in ecosystem carbon (C) storage with time). We examined the influence of live tree and coarse woody debris (CWD) on NEP during secondary succession based on data collected along a 500-year chronosequence on the Wind River Ranger District, Washington. We developed a simple statistical model of live and dead wood accumulation and decomposition to predict changes in the woody component of NEP, which we call NEP(w). The transition from negative to positive NEP(w), for a series of scenarios in which none to all wood was left after disturbance, occurred between 0 and 57 years after disturbance. The timing of this transition decreased as live-tree growth rates increased, and increased as CWD left after disturbance increased. Maximum and minimum NEP(w) for all scenarios were 3.9 and -14.1 Mg C ha-1 year-1, respectively. Maximum live and total wood C stores of 319 and 393 Mg C ha(-1), respectively, were reached approximately 200 years after disturbance. Decomposition rates (k) of CWD ranged between 0.013 and 0.043 year-1 for individual stands. Regenerating stands took 41 years to attain a mean live wood mass equivalent to the mean mass of CWD left behind after logging, 40 years to equal the mean CWD mass in 500-year-old forest, and more than 150 years to equal the mean total live and dead wood in an old-growth stand. At a rotation age of 80 years, regenerating stands stored approximately half the wood C of the remaining nearby old-growth forests (predominant age 500 years), indicating that conversion of old-growth forests to younger managed forests results in a significant net release of C to the atmosphere.

An investigation of hydraulic limitation and compensation in large, old Douglas-fir trees.

Abstract: The hydraulic limitation hypothesis (Ryan and Yoder 1997) proposes that leaf-specific hydraulic conductance (kl) and stomatal conductance (gs) decline as trees grow taller, resulting in decreased carbon assimilation. We tested the hydraulic limitation hypothesis by comparison of canopy-dominant Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) trees in stands that were approximately 15 m (20 years old), 32 m (40 years old) and 60 m (> 450 years old) tall in Wind River, Washington, USA. Carbon isotope discrimination (Delta) declined with tree height (18.6, 17.6 and 15.9 per thousand for stands 15, 32 and 60 m tall, respectively) indicating that gs may have declined proportionally with tree height in the spring months, when carbon used in the construction of new foliage is assimilated. Hydraulic conductance decreased by 44% as tree height increased from 15 to > 32 m, and showed a further decline of 6% with increasing height. The general nonlinear pattern of kl versus height was predicted by a model based on Darcy's Law. Stemwood growth efficiency also declined nonlinearly with height (60, 35 and 28 g C m-2 leaf area for the 15-, 32- and 60-m stands, respectively). Unlike kl and growth efficiency, gs and photosynthesis (A) during summer drought did not decrease with height. The lack of decline in cuvette-based A indicates that reduced A, at least during summer months, is not responsible for the decline in growth efficiency. The difference between the trend in gs and A and that in kl and D may indicate temporal changes (spring versus summer) in the response of gas exchange to height-related changes in kl or it may be a result of measurement inadequacies. The formal hydraulic limitation hypothesis was not supported by our mid-summer gs and A data. Future tests of the hydraulic limitation hypothesis in this forest should be conducted in the spring months, when carbon uptake is greatest. We used a model based on Darcy's Law to quantify the extent to which compensating mechanisms buffer hydraulic limitations to gas exchange. Sensitivity analyses indicated that without the observed increases in the soil-to-leaf water potential differential (DeltaPsi) and decreases in the leaf area/sapwood area ratio, kl would have been reduced by more than 70% in the 60-m trees compared with the 15-m trees, instead of the observed decrease of 44%. However, compensation may have a cost; for example, the greater DeltaPsi of the largest trees was associated with smaller tracheid diameters and increased sapwood cavitation, which may have a negative feedback on kl and gs.

Photosynthetic differences between saplings and adult trees: an integration of field results by meta-analysis.

Abstract: Ontogenetic changes in gas exchange parameters provide both insight into mechanisms underlying tree growth patterns, and data necessary to scale environmental impacts on young trees to predict responses of older trees. We present a quantitative review and meta-analysis of field measurements of gas exchange parameters in saplings and mature trees of 35 tree species (seven conifers, seven temperate deciduous trees, and 21 tropical evergreen trees). Data for saplings were obtained in both understory environments and open areas or large gaps. We also present data on ontogenetic changes in photosynthesis for Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg., species of particular interest because of their large maximal heights and long life-spans. Among tree species, there is evidence for both ontogenetic increases and ontogenetic decreases in photosynthetic capacity on a leaf area basis (A(area)). Overall, A(area) is generally higher for upper-canopy leaves of adult trees than for saplings, especially in temperate deciduous trees. However, the pattern for photosynthetic capacity on a leaf mass basis (A(mass)) is the reverse of that observed for A(area). Saplings of both conifers and broad-leaved trees, even when acclimated to low-light conditions, characteristically have a higher A(mass) than adult trees. This pattern is driven largely by an ontogenetic increase in leaf mass per unit area (LMA), as found in 100% of studies reviewed. Data for Pacific Northwest conifers, although including measurements on some of the tallest trees studied, did not differ greatly from patterns found in other tree species. We conclude that ontogenetic changes in LMA are the single most consistent difference between saplings and adult trees, and that changes in LMA and related aspects of leaf morphology may be critical to understanding both variation in gas exchange during tree growth, and stage-dependent responses of trees to environmental change.

Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris

Abstract: Foliar light-saturated net assimilation rates (A) generally decrease with increasing tree height (H) and tree age (Y), but it is unclear whether the decline in A is attributable to size- and age-related modifications in foliage morphology (needle dry mass per unit projected area; M(A)), nitrogen concentration, stomatal conductance to water vapor (G), or biochemical foliage potentials for photosynthesis (maximum carboxylase activity of Rubisco; V(cmax)). I studied the influences of H and Y on foliage structure and function in a data set consisting of 114 published studies reporting observations on more than 200 specimens of various height and age of Picea abies (L.) Karst. and Pinus sylvestris L. In this data set, foliar nitrogen concentrations were independent of H and Y, but net assimilation rates per unit needle dry mass (A(M)) decreased strongly with increasing H and Y. Although M(A) scaled positively with H and Y, net assimilation rates per unit area (A(A) = M(A) x A(M)) were strongly and negatively related to H, indicating that the structural adjustment of needles did not compensate for the decline in mass-based needle photosynthetic rates. A relevant determinant of tree height- and age-dependent modifications of A was the decrease in G. This led to lower needle intercellular CO2 concentrations and thereby to lower efficiency with which the biochemical photosynthetic apparatus functioned. However, V(cmax) per unit needle dry mass and area strongly decreased with increasing H, indicating that foliar photosynthetic potentials were lower in larger trees at a common intercellular CO2 concentration. Given the constancy of foliar nitrogen concentrations, but the large decline in apparent V(cmax) with tree size and age, I hypothesize that the decline in Vcmax results from increasing diffusive resistances between the needle intercellular air space and carboxylation sites in chloroplasts. Increased diffusive limitations may be the inevitable consequence of morphological adaptation (changes in M(A) and needle density) to greater water stress in needles of larger trees. Foliage structural and physiological variables were nonlinearly related to H and Y, possibly because of hyperbolic decreases in shoot hydraulic conductances with increasing tree height and age. Although H and Y were correlated, foliar characteristics were generally more strongly related to H than to Y, suggesting that increases in height rather than age are responsible for declines in foliar net assimilation capacities.

Heat dissipation sensors of variable length for the measurement of sap flow in trees with deep sapwood

Abstract: Robust thermal dissipation sensors of variable length (3 to 30 cm) were developed to overcome limitations to the measurement of radial profiles of sap flow in large-diameter tropical trees with deep sapwood. The effective measuring length of the custom-made sensors was reduced to 1 cm at the tip of a thermally nonconducting shaft, thereby minimizing the influence of nonuniform sap flux density profiles across the sapwood. Sap flow was measured at different depths and circumferential positions in the trunks of four trees at the Parque Natural Metropolitano canopy crane site, Panama City, Republic of Panama. Sap flow was detected to a depth of 24 cm in the trunks of a 1-m-diameter Anacardium excelsum(Bertero & Balb. ex Kunth) Skeels tree and a 0.65-m-diameter Ficus insipida Willd. tree, and to depths of 7 cm in a 0.34-m-diameter Cordia alliodora (Ruiz & Pav.) Cham. trunk, and 17 cm in a 0.47-m-diameter Schefflera morototoni (Aubl.) Maguire, Steyerm. & Frodin trunk. Sap flux density was maximal in the outermost 4 cm of sapwood and declined with increasing sapwood depth. Considerable variation in sap flux density profiles was observed both within and among the trees. In S. morototoni, radial variation in sap flux density was associated with radial variation in wood properties, particularly vessel lumen area and distribution. High variability in radial and circumferential sap flux density resulted in large errors when measurements of sap flow at a single depth, or a single radial profile, were used to estimate whole-plant water use. Diurnal water use ranged from 750 kg H 2 O day −1 for A. excelsum to 37 kg H 2 O day −1 for C. alliodora.

Radial patterns of sap flow in woody stems of dominant and understory species: scaling errors associated with positioning of sensors

Abstract: We studied sap flow in dominant coniferous (Pinus sylvestris L.) and broadleaf (Populus canescens L.) species and in understory species (Prunus serotina Ehrh. and Rhododendron ponticum L.) by the heat field deformation (HFD) method. We attempted to identify possible errors arising during flow integration and scaling from single-point measurements to whole trees. Large systematic errors of -90 to 300% were found when it was assumed that sap flow was uniform over the sapwood depth. Therefore, we recommend that the radial sap flow pattern should be determined first using sensors with multiple measuring points along a stem radius followed by single-point measurements with sensors placed at a predetermined depth. Other significant errors occurred in the scaling procedure even when the sap flow radial pattern was known. These included errors associated with uncertainties in the positioning of sensors beneath the cambium (up to 15% per 1 mm error in estimated xylem depth), and differences in environmental conditions when the radial profile applied for integration was determined over the short term (up to 47% error). High temporal variation in the point-to-area correction factor along the xylem radius used for flow integration is also problematic. Compared with midday measurements, measurements of radial variation of sap flow in the morning and evening of sunny days minimized the influence of temporal variations on the point-to-area correction factor, which was especially pronounced in trees with a highly asymmetric sap flow radial pattern because of differences in functioning of the sapwood xylem layers. Positioning a single-point sensor at a depth with maximum sap flow is advantageous because of the high sensitivity of maximum sap flow to water stress conditions and changes in micro-climate, and because of the lower random errors associated with the positioning of a single-point sensor along the xylem radius.

How the environment, canopy structure and canopy physiological functioning influence carbon, water and energy fluxes of a temperate broad-leaved deciduous forest--an assessment with the biophysical model CANOAK.

Abstract: This paper focuses on how canopy structure, its physiological functioning and the environment interact to control and drive the exchange of carbon dioxide (CO2) and water vapor between a temperate forest stand and the atmosphere. First, we present an overview of how temporal and spatial variations in canopy structure (e.g., leaf area index, species, leaf inclination angles, leaf clumping) and physiological functioning (e.g., maximal stomatal conductance, photosynthetic capacity) modulate CO2 and water vapor fluxes. Then, with the biophysical model CANOAK, we quantify the effects of leaf dimension and thickness, vertical variations in leaf area and photosynthetic capacity, leaf clumping, leaf inclination angles, photosynthetic capacity, stomatal conductance and weather on the annual sums of CO2, water vapor and sensible heat exchange. Finally, we discuss how much detail is needed in a model to predict fluxes of CO2 and water vapor with acceptable fidelity.

Age- and size-related trends in woody plant shoot development: regulatory pathways and evidence for genetic control.

Abstract: Woody plants exhibit significant and predictable patterns of change in morphology and physiology as they become older and larger. Four models of potential pathways controlling these changes are presented: a stimulus-response model in which fully developed organs respond to changes in environment (defined here as everything external to the organ); an extrinsic model in which the attributes of developing organs are determined by environmental factors; an intrinsic model in which changes are a result of programmed changes in gene expression; and an extrinsic-intrinsic model in which changes in gene expression are induced by environmental factors. We review evidence that a genetic component is involved in controlling age- and size-related changes in foliar morphology and physiology and discuss the possibility of complex interactions among model pathways.

Photosynthetic responses to water deficit in six Mediterranean sclerophyll species: possible factors explaining the declining distribution of Rhamnus ludovici-salvatoris, an endemic Balearic species.

Abstract: We sought to explain the declining distribution in the Balearic Islands of the endemic shrub Rhamnus ludovici-salvatoris R. Chodat, by comparing its photosynthetic response to drought with that of several widely distributed, competing Mediterranean species (R. alaternus L., Quercus ilex L., Pistacia lentiscus L., Q. humilis Mill. and P. terebinthus L.). All of the study species, except for the two Rhamnus species, avoided desiccation by rapidly adjusting their stomatal conductance at the onset of drought, and maintaining constant leaf relative water content. The two Rhamnus species showed desiccation-tolerant behavior; i.e., as drought progressed, their predawn leaf relative water content decreased simultaneously with stomatal closure. All four desiccation-avoiding species showed a significant positive correlation between leaf thermal dissipation (estimated by the fluorescence parameter NPQ (non-photochemical quenching)) and the de-epoxidation state of the xanthophyll cycle (DPS). The two Rhamnus species exhibited maximum DPS regardless of treatment, but only R. alaternus increased NPQ in response to drought. Rhamnus ludovici-salvatoris had a high ratio of photorespiration to photosynthesis and a low intrinsic water-use efficiency; traits that are likely to be unfavorable for plant productivity under arid conditions. It also had the lowest DPS and thermal dissipation among the six species. We conclude that the photosynthetic traits of R. ludovici-salvatoris account for its limited ability to compete with other species in the Mediterranean region.

Age- and position-related changes in hydraulic versus mechanical dysfunction of xylem: inferring the design criteria for Douglas-fir wood structure

Abstract: Summary We do not know why trees exhibit changes in wood characteristics as a function of cambial age. In part, the answer may lie in the existence of a tradeoff between hydraulic properties and mechanical support. In conifers, longitudinal tracheids represent 92% of the cells comprising the wood and are involved in both water transport and mechanical support. We used three hydraulic parameters to estimate hydraulic safety factors at several vertical and radial locations in the trunk and branches: vulnerability to cavitation; variation in xylem water potential (Ψ); and xylem relative water content. The hydraulic safety factors for 12 and 88 percent loss of conductivity (SH12 and SH88, representing the hydraulic safety factors for the air entry point and full embolism point, respectively) were determined. We also estimated the mechanical safety factor for maximum tree height and for buckling. We estimated the dimensionless hydraulic and mechanical safety factors for six seedlings (4 years old), six saplings (10 years old) and six mature trees (> 110 years old) of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco). Over the natural range of Ψ, SH12 decreased linearly from treetop to a minimum of 0.95 at the tree base. Young and mature trees had SH12 values 1.4 and 1.3 times higher, respectively, at their tips (juvenile wood) than at their bases (mature wood). Modeling analyses indicated that if trees were made entirely of mature wood, SH12 at the stem base would be only 0.7. The mechanical safety factor was 1.2 times higher for the base of the tree than for the rest of the tree. The minimum mechanical safety factor—1.6 for the critical buckling height and 2.2 for the critical buckling load—occurred at the base of the live crown. Modeling analysis indicated that if trees were made only of mature wood, these values would increase to 1.7 and 2.3, respectively. Hydraulic safety factors had values that were less than half those for mechanical safety factors, suggesting that wood structure in Douglas-fir has evolved primarily as a result of selection for hydraulic safety rather than mechanical safety. The results suggest that forest managers must consider the role of juvenile wood in tree physiology to avoid producing plantations vulnerable to drought.

Canopy and hydraulic conductance in young, mature and old Douglas-fir trees.

Abstract: We tested for reductions in water transport with increasing tree size, a key component in determining whether gas exchange and growth are hydraulically limited in tall trees. During the summers of 1998 and 1999, we measured water transport with Granier-type, constant-heat sap flow probes, vapor pressure deficit, and leaf and soil water potentials in overstory Pseudotsuga menziesii (Mirb.) Franco trees in three stands differing in size and age (15, 32 and 60 m in height and about 20, 40 and 450 years in age, respectively) in a P. menziesii-dominated forest in the Pacific Northwest, USA. A total of 24 trees were equipped with sap flow sensors--six 60-m trees, nine 32-m trees and nine 15-m trees. Based on the sap flow measurements and leaf area information estimated from leaf area-sapwood area relationships, we estimated crown-averaged stomatal conductance (GS) and leaf-specific hydraulic conductance (KL). We tested the hypothesis that GS and KL vary inversely with tree height (15 > 32 > 60 m). Analysis of variance of GS ranked as 15 = 60 > 32 m during the early summer and 15 > 60 > 32 m during late season drought. Over the growing season, mean daily GS (+/- SE) was 29.2 +/- 4.4, 24.0 +/- 6.8 and 17.7 +/- 7.2 mmol m-2 s-1 for the 15-, 60- and 32-m trees, respectively. The value of K(L) differed among tree heights only during late season drought and ranked 15 > 32 = 60 m. A hydraulic mass balance suggests that greater sapwood conductivity in 60-m trees compared with 32- and 15-m trees is a likely cause for the departure of the above rankings from those predicted by height and leaf-to-sapwood area ratio.

Response of old-growth conifers to reduction in stand density in western Oregon forests.

Abstract: Summary The positive growth response of healthy young trees to density reduction is well known. In contrast, large old trees are usually thought to be intrinsically limited in their ability to respond to increased growing space; therefore, density reduction is seldom used in stands of old-growth trees. We tested the null hypothesis that old-growth trees are incapable of responding with increased growth following density reduction. The diameter growth response of 271 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), ponderosa pine (Pinus ponderosa Dougl. ex Laws) and sugar pine (Pinus lambertiana Dougl.) trees ranging in age from 158 to 650 years was examined 20 to 50 years after density reduction. Density reduction involved either light thinning with removal of less vigorous trees, or shelterwood treatments in which overstory trees were not removed. Ratios of basal area growth after treatment to basal area growth before treatment, and several other measures of growth, all indicated that the old trees sometimes benefited and were not harmed by density reduction. Growth increased by 10% or more for 68% of the trees in treated stands, and nearly 30% of trees increased growth by over 50%. This growth response persisted for at least 20 years. During this 20-year period, only three trees in treated stands (1.5%) exhibited a rapid decrease in growth, whereas growth decreased in 64% of trees in untreated stands. The length of time before a growth response to density reduction occurred varied from 5 to 25 years, with the greatest growth response often occurring 20 to 25 years after treatment. These results have important implications both for the basic biology of aging in woody plants as well as for silvicultural practices in forests with old-growth trees.

Hydraulic redistribution of soil water by neotropical savanna trees

Abstract: The magnitude and direction of water transport by the roots of eight dominant Brazilian savanna (Cerrado) woody species were determined with a heat pulse system that allowed bidirectional measurements of sap flow. The patterns of sap flow observed during the dry season in species with dimorphic root systems were consistent with the occurrence of hydraulic redistribution of soil water, the movement of water from moist to drier regions of the soil profile via plant roots. In these species, shallow roots exhibited positive sap flow (from the soil into the plant) during the day and negative sap flow (from the plant into the soil) during the night. Sap flow in the taproots was positive throughout the 24-h period. Diel fluctuations in soil water potential, with maximum values occurring at night, provided evidence for partial rewetting of upper soil layers by water released from shallow roots. In other species, shallow roots exhibited negative sap flow during both the day and night, indicating that hydraulic redistribution was occurring continuously. A third sap flow pattern was observed at the end of the dry season after a heavy rainfall event when sap flow became negative in the taproot, and positive in the small roots, indicating movement of water from upper soil layers into shallow roots, and then into taproots and deeper soil layers. Experimental manipulations employed to evaluate the response of hydraulic redistribution to changes in plant and environmental conditions included watering the soil surface above shallow roots, decreasing transpiration by covering the plant and cutting roots where probes were inserted. Natural and manipulated patterns of sap flow in roots and stems were consistent with passive movement of water toward competing sinks in the soil and plant. Because dry shallow soil layers were often a stronger sink than the shoot, we suggest that the presence of a dimorphic root system in deciduous species may play a role in facilitating leaf expansion near the end of the dry season when the soil surrounding shallow lateral roots is still dry.

Physiological traits of two Populus x euramericana clones, Luisa Avanzo and Dorskamp, during a water stress and re-watering cycle.

Abstract: We compared responses to drought and re-watering of greenhouse-grown cuttings of Populus x euramericana (Dode) Guinier clones, Luisa Avanzo and Dorskamp. Total leaf area, leaf number, leaf area increment and stomatal conductance were evaluated periodically during a 29-day drought period and for 16 days after re-watering. Soil water content and predawn leaf water potential (psi(wp)) were measured on Days 29 and 45. On the same days, relative water content (RWC), specific leaf area (SLA), nitrogen, chlorophyll, soluble sugars, total phenols, flavanols and antioxidant activity were determined for leaves taken from the bottom to the top of each cutting. Leaves of Luisa Avanzo cuttings grew more rapidly than leaves of Dorskamp and exhibited higher SLA, but lower concentrations of nitrogen, chlorophyll and soluble sugars and lower antioxidant activity per unit area. On Day 29, after withholding water, both clones had closed their stomata, reduced rates of leaf growth, and lower psi(wp) and RWC; however, the clones differed in their responses to soil water depletion. Compared to Dorskamp, Luisa Avanzo closed its stomata earlier and maintained higher psi(wp), but lower RWC and leaf sugar concentrations. Antioxidant activity of leaf methanolic extracts decreased in response to water stress only in Luisa Avanzo. Leaf physiology and its modulation by water stress were age dependent in Luisa Avanzo.

Growth of young apple trees in relation to reserve nitrogen and carbohydrates.

Abstract: Bench-grafted Fuji/M.26 apple (Malus domestica Borkh.) trees were fertilized with a nutrient solution (fertigation) containing 0, 2.5, 5, 7.5, 10, 15 or 20 mM nitrogen (N) in a modified Hoagland's solution from June 30 to September 1. In mid-October, half of the trees in each N treatment were sprayed twice with 3% urea, 1 week apart. The remaining trees served as controls. All trees were harvested after leaf fall and stored at 2 degrees C over winter. One group of trees from each treatment was destructively sampled before bud break to determine amounts of reserve N and total nonstructural carbohydrates (TNC); the remaining trees were transplanted to N-free medium in the spring. These trees were supplied with Hoagland's solution with or without 10 mM N (from 15N-depleted NH4NO3) for 60 days, starting from bud break. With increasing N supply from fertigation, tree N concentration increased, whereas TNC concentration decreased. Foliar urea applications increased tree N concentration and decreased TNC concentration in each N fertigation treatment. There was a negative linear relationship between tree N concentration and TNC concentration. Irrespective of whether N was provided the following spring, trees with high N reserves but low carbohydrate reserves produced a larger total leaf area at the end of the regrowth period than trees with low N reserves but high carbohydrate reserves. The pooled data on reserve N used for new growth showed that, regardless of the spring N supply, there was a linear relationship between total N accumulated in the tree during the previous season and the amount of reserve N remobilized for new shoot and leaf growth. About 50% of tree N content was remobilized to support new shoot and leaf growth over the range of tree N status examined. We conclude that the initial growth of young apple trees in the spring is determined mainly by reserve N, not reserve carbohydrates. The amount of reserve N remobilized for new growth in spring was proportional to tree N status and was unaffected by current N supply.

Winter embolism, mechanisms of xylem hydraulic conductivity recovery and springtime growth patterns in walnut and peach trees

Abstract: Xylem vessels of Prunus persica Batsch (peach) and Juglans regia L. (walnut) are vulnerable to frost-induced embolism. In peach, xylem embolism increased progressively over the winter, reaching a maximum of 85% loss of hydraulic conductivity (PLC) in early March. Over winter, PLC in walnut approached 100%, but the degree of xylem embolism varied during the winter, reflecting the ability of walnut to generate positive xylem pressures in winter and spring. In contrast, positive xylem pressures were not observed in peach. Controlled freeze-thaw experiments showed that frost alone is insufficient to increase embolism in peach; evaporative conditions during thawing are also required. However, when both species were protected from frost, PLC was zero. At bud break, there was complete recovery from embolism in walnut, whereas PLC remained high in peach. Three mechanisms responsible for the restoration of branch hydraulic conductivity were identified in walnut: the development of stem pressure, the development of root pressure and the formation of a new ring of functional xylem, whereas only one mechanism was observed in peach (new functional ring). The climatic conditions necessary for the manifestation of these mechanisms were investigated.

Genomic DNA methylation-demethylation during aging and reinvigoration of Pinus radiata.

Abstract: In animals, DNA methylation is related to gene silencing during ontogenic development. Little is known about DNA methylation in plants, although occasional changes in the DNA methylation state of specific gene promoters have been reported in angiosperms during some developmental processes. We found large differences in the extent of DNA methylation between meristematic areas of juvenile and mature Pinus radiata D. Don. trees, whereas differences in the extent of DNA methylation between differentiated tissues of juvenile and mature trees were small. In meristematic areas, there was a gradual decrease in extent of DNA methylation as the degree of reinvigoration increased. The observed changes in extent of DNA methylation during aging and reinvigoration indicate that reinvigoration could be a consequence of epigenetic modifications opposite in direction to those that occur during aging.

Influence of natural temperature gradients on measurements of xylem sap flow with thermal dissipation probes. 1. Field observations and possible remedies.

Abstract: The thermal dissipation method is simple and widely used for measuring sap flow in large stems. As with several other thermal methods, natural temperature gradients are assumed to be negligible in the sapwood being measured. We studied the magnitude and variability of natural temperature gradients in sapwood of Acacia trees growing in the Sahelian zone of Senegal, analyzed their effects on sap flow measurements, and investigated possible solutions. A new measurement approach employing cyclic heating (45 minutes of heating and 15 minutes of cooling; 45/15) was also tested. Three-day measurement sequences that included 1 day without heating, a second day with continuous heating and a third day with cyclic heating were recorded during a 6.5-month period using probes installed at three azimuths in a tree trunk. Natural temperature gradients between the two probes of the sensor unit, spaced 8 to 10 cm vertically, were rarely negligible (i.e., < 0.2 degrees C): they were positive during the night and negative during the day, with an amplitude ranging from 0.3 to 3.5 degrees C depending on trunk azimuth, day and season. These temperature gradients had a direct influence on the signal from the continuously heated sensors, inducing fluctuations in the nighttime reference signal. The resulting errors in sap flow estimates can be greater than 100%. Correction protocols have been proposed in previous studies, but they were unsuitable because of the high spatial and temporal variability of the natural temperature gradients. We found that a measurement signal derived from a noncontinuous heating system could be an attractive solution because it appears to be independent of natural temperature gradients. The magnitude and variability of temperature gradients that we observed were likely exacerbated by the combination of open stand, high solar radiation and low sap flow rate. However, for all applications of the thermal dissipation method, it is wise to check regularly for natural temperature gradients by switching off the heater.

Comparison of water-use efficiency of seedlings from two sympatric oak species: genotype × environment interactions

Abstract: Seedlings of two sympatric oak species, Quercus robur L. and Quercus petraea (Matt.) Liebl., were grown in common garden conditions to test for potential interspecific differences in intrinsic water-use efficiency (WUE). Intrinsic water-use efficiency was estimated based on carbon isotope composition of shoots (delta13C) and on gas exchange measurements (ratio of net CO2 assimilation rate to stomatal conductance (A/g(sw))). In addition, genotype x environment interactions were tested by subjecting the seedlings to four irradiance treatments (8, 18, 48 and 100% of incident solar irradiance) imposed by neutral shading nets, and, in the 100% irradiance treatment, two watering regimes. In all treatments, initial growth of Q. robur was faster than that of Q. petraea. In both species, there was a tight correlation between delta13C and A/g(sw). Intrinsic water-use efficiency increased with increasing irradiance (almost doubling from 8 to 100% irradiance), and this effect paralleled the increase in A with increasing irradiance. In full sun, WUE of Q. petraea seedlings was 10-15% higher than in Q. robur seedlings, with the difference attributable to a difference between the species in g(sw). The interspecific difference in WUE was maintained during drought, despite the appreciable increase in WUE and decrease in growth imposed by drought. No interspecific differences in WUE were observed at low irradiances, suggesting a strong genotype x environment interaction for WUE. These findings confirm the existence of interspecific genetic differences in WUE, but also show that there is large intraspecific variability and plasticity in WUE. The initially greater height and biomass increments in Q. robur seedlings illustrate the ability of this species to out-compete Q. petraea in the early stages of forest regeneration. For adult trees growing in closed canopies, the high WUE of Q. petraea may contribute significantly to its survival during dry years, whereas the low WUE of Q. robur may account for the frequently observed declines in adult trees of this species following drought.

SAGE profiling and demonstration of differential gene expression along the axial developmental gradient of lignifying xylem in loblolly pine (Pinus taeda).

Abstract: Wood formation has been studied extensively at the cellular and biochemical levels, but remains poorly understood with respect to gene expression and regulation. As a first step toward identifying genes specifically involved in wood formation and characterizing their roles in determining wood quality, serial analysis of gene expression (SAGE) was used to quantify gene expression in lignifying xylem from a single, 10-year-old loblolly pine (Pinus taeda L.). Two SAGE libraries were generated based on lignifying xylem isolated from either the upper (crown) or lower (base) portions of the trunk. Over 85,000 tags representing a maximum of 27,398 expressed genes were analyzed from the crown wood library, and more than 65,000 tags, representing a maximum of 25,983 expressed genes, were analyzed in the base wood library. Combining these data sets to reflect the sum of genes expressed in lignifying xylem, 150,855 tags were cataloged, representing a maximum of 42,641 different genes. Currently, this study represents the most extensive analysis of its kind in a higher plant and provides a quantitative description of the transcriptome representing the lignifying xylem of a 10-year-old loblolly pine.

When branch autonomy fails: Milton's Law of resource availability and allocation.

Abstract: The branch autonomy principle states that the critical characteristics of a branch's carbohydrate economy (photosynthesis, respiration, growth, etc.) are largely independent of the tree to which the branch is attached, as long as light is the primary factor limiting photosynthesis and growth. However, this may not be generally true because in the spring, photosynthates are translocated from a tree stem into branches, and the amount of photosynthate available for translocation should be a function of the tree's canopy status. And the correlative inhibition principle states that a branch's priority for allocation of carbon and other resources is controlled not only by its own environment, but also by its position relative to other branches on the same tree. A study of the lower limit of branch growth and survival in trees of different sizes shows that the latter principle is more important: even though dominant trees have more resources to allocate, branches on suppressed trees are able to grow and produce new foliage at solar irradiances where branches on dominant trees die. Thus branches are sufficiently interdependent that a positive carbon budget by itself does not ensure branch survival; branch position relative to other branches on the same tree is also important. Other findings indicate that this result is quite general: regardless of the stress involved, a stressed branch on a tree where all other branches are also stressed does better than a similarly stressed branch on a tree where some branches are relatively unstressed. Although branch autonomy is an important and useful principle, it is not an absolute rule governing branch growth.

Derivation of stem taper from the pipe theory in a carbon balance framework

Abstract: A dynamic tree growth model is described. The model derives the development of stem taper and vertical distribution of branch basal area from the pipe model, assuming that reuse of active pipes is regulated by foliage dynamics in a vertically explicit crown with a foliage distribution of constant shape. Based on empirical findings, the pipe model was modified slightly to allow the foliage/sapwood ratio to vary as a function of distance from the treetop. Growth was derived from carbon balance in a stand of different size trees that may shade each other. The model was applied to old and middle-aged trees growing in dense and sparse stands of Scots pine for which stand-level measurements are available as a chronosequence, but individual trees have been measured only once. Measured trees were compared with corresponding simulated trees for stem taper and vertical distribution of branch basal area. The results indicated that the pipe model assumptions, combined with a model of tree growth, are capable of producing realistic predictions of the vertical distribution of stem and branch diameter in trees of different sizes in the stand. A comparison of the results with a simple form of the uniform stress theory showed good agreement between the two models. However, a significant difference was found between the measured relative contribution of heartwood to total stem diameter and the predicted share of disused pipes in the stem. A possible explanation for this discrepancy is that the transition from sapwood to heartwood is gradual rather than abrupt as assumed in the model. A modification of the pipe model to incorporate a gradual transition is outlined.

Response of a mature Pinus laricio plantation to a three-year restriction of water supply: structural and functional acclimation to drought.

Abstract: The response of mature forest stands to a reduction in water availability has received little attention In particular, the extent to which a short-term reduction in gas exchange can be alleviated in the long-term by acclimation processes is not well understood We studied the impact of a severe reduction in water availability on the water relations and growth of 35-year-old Pinus laricio Poiret trees in a replicated experiment Sapwood and needle increments, soil and tree water status, stand transpiration, xylem embolism and plant hydraulic architecture were monitored over a 3-year period in control and drought-treated plots Needle length was reduced in drought-treated trees by 30, 19 and 29%, and sapwood increments by 50, 27 and 24% over the 3 years Drought did not result in tree mortality or in extensive xylem embolism or foliage dieback On the contrary, a conservative water-use strategy was observed, because minimum leaf water potentials did not differ between treatments or over the season Plant hydraulic resistance was also unaffected by restricted water availability Stand transpiration was strongly reduced by drought treatment over the summer, but not during the winter, despite significant differences in leaf area between control and drought-treated trees, implying higher transpiration rates per unit leaf area in the droughted plants This suggests that water transport capacity, rather than the amount of leaf area, controlled stand transpiration, which is at variance with expectations based on experiments with seedlings and short-term experiments with mature trees

Age-related effects on leaf area/sapwood area relationships, canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge, Germany.

Abstract: Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.

Leaf area dynamics of a boreal black spruce fire chronosequence.

Abstract: Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P 40%) of total leaf area in all stands except the oldest.

Changes in foliar spectral reflectance and chlorophyll fluorescence of four temperate species following branch cutting.

Abstract: Spectral reflectance and chlorophyll fluorescence are rapid non-invasive methods that can be used to quantify plant stress. Because variation in ambient light (e.g., diurnal patterns of solar radiation) may have a confounding effect on these measurements, branches are often excised in the field and then measured under controlled conditions in the laboratory. We studied four temperate tree species (Abies balsamea (L.) Mill. (balsam fir), Betula papyrifera var. cordifolia (Regel) Fern. (paper birch), Picea rubens Sarg. (red spruce) and Sorbus americana Marsh. (mountain-ash)) to determine how quickly reflectance and fluorescence change following branch cutting. We hypothesized that conifer species, which have tough xeromorphic foliage, would exhibit changes more slowly than broadleaf species. Furthermore, we hypothesized that keeping broadleaf samples cool and moist would delay the onset of reflectance and fluorescence changes. In one set of experiments, we did not use any treatments to maintain the freshness of cut branches. During the first 12 h following cutting, changes in reflectance and fluorescence were slight for all species. Two or 3 days after branch cutting, the two conifers still showed only small changes in the ratio of variable to maximum fluorescence (F v /F m ) and most reflectance indices, whereas paper birch and mountain-ash showed larger and more rapid declines in F v /F m and most reflectance indices. We attribute these declines to loss of water. As a consequence of xeromorphic leaf structure, the conifers were better able to minimize water loss than the two broadleaf species. In another experiment, paper birch that had been kept cool and moist after cutting showed only slight changes in fluorescence and reflectance, even after 3 days, indicating that with careful handling the time interval between collection and measurement of reflectance and fluorescence of many broadleaf specimens can be extended to several days. We conclude that measurements of reflectance and fluorescence need not be made in situ to be accurate and reliable.