Plasticity in hydraulic architecture of Scots pine across Eurasia.

Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G(s)) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G(s) to D (m/G (s,ref), where m is the slope and G(s,ref) is reference G(s) at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G (s,ref), and its absolute sensitivity to D(m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.

Quantifying ozone uptake at the canopy level of spruce, pine and larch trees at the alpine timberline: an approach based on sap flow measurement.

Micro-climatic and ambient ozone data were combined with measurements of sap flow through tree trunks in order to estimate whole-tree ozone uptake of adult Norway spruce (Picea abies), cembran pine (Pinus cembra), and European larch (Larix decidua) trees. Sap flow was monitored by means of the heat balance approach in two trees of each species during the growing season of 1998. In trees making up the stand canopy, the ozone uptake by evergreen foliages was significantly higher than by deciduous ones, when scaled to the ground area. However, if expressed per unit of whole-tree foliage area, ozone flux through the stomata into the needle mesophyll was 1.09, 1.18 and 1.40 nmol m(-2) s(-1) in Picea abies, Pinus cembra and Larix decidua, respectively. These fluxes are consistent with findings from measurements of needle gas exchange, published from the same species at the study site. It is concluded that the sap flow-based approach offers an inexpensive, spatially and temporally integrating way for estimating ozone uptake at the whole-tree and stand level, intrinsicly covering the effect of boundary layers on ozone flux.

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

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.

The relationship between tree height and leaf area: sapwood area ratio.

The leaf area to sapwood area ratio (A l:A s) of trees has been hypothesized to decrease as trees become older and taller. Theory suggests that A l:A s must decrease to maintain leaf-specific hydraulic sufficiency as path length, gravity, and tortuosity constrain whole-plant hydraulic conductance. We tested the hypothesis that A l:A s declines with tree height. Whole-tree A l:A s was measured on 15 individuals of Douglas-fir (Pseudotsuga menziesii var. menziesii) ranging in height from 13 to 62 m (aged 20-450 years). A l:A s declined substantially as height increased (P=0.02). Our test of the hypothesis that A l:A s declines with tree height was extended using a combination of original and published data on nine species across a range of maximum heights and climates. Meta-analysis of 13 whole-tree studies revealed a consistent and significant reduction in A l:A s with increasing height (P<0.05). However, two species (Picea abies and Abies balsamea) exhibited an increase in A l:A s with height, although the reason for this is not clear. The slope of the relationship between A l:A s and tree height (ΔA l:A s/Δh) was unrelated to mean annual precipitation. Maximum potential height was positively correlated with ΔA l:A s/Δh. The decrease in A l:A s with increasing tree size that we observed in the majority of species may be a homeostatic mechanism that partially compensates for decreased hydraulic conductance as trees grow in height.

Reduced uptake of oxidized low density lipoproteins in monocyte-derived macrophages from CD36-deficient subjects.

To clarify the physiological roles of CD36 as an oxidized low density lipoprotein (OxLDL) receptor, we analyzed the monocyte-derived macrophages from normal and two CD36-deficient subjects, since we identified the molecular abnormalities (Kashiwagi, H., Y. Tomiyama, Y. Kosugi, M. Shiraga, R. H. Lipsky, Y. Kanayama, Y. Kurata, and Y. Matsuzawa 1994. Blood. 83:3545-3552; and Kashiwagi, H., Y. Tomiyama, S. Honda, S. Kosugi, M. Shiraga, N. Nagao, S. Sekiguchi, Y. Kanayama, Y. Kurata, and Y. Matsuzawa. 1995. J. Clin. Invest. 95:1040-1046). Scatchard analysis of 125I-OxLDL binding showed a linear plot and the maximum binding was lower by approximately 40% in the macrophages from subjects with CD36 deficiency than those from normal controls. Competition studies showed that the uptake of 125I-OxLDL was suppressed by OKM5, an antibody against CD36, by 53% in normal control macrophages, but not in the CD36-deficient macrophages. After incubation with OxLDL for 24 h, cholesteryl ester mass accumulation was reduced by approximately 40% in the macrophages from CD36-deficient subjects than those from normal controls. These results suggest that CD36 is one of the physiological receptors for OxLDL. Since specific binding of OxLDL was only reduced by approximately 40% in spite of the complete deficiency of CD36, several other receptors also may have some role in OxLDL uptake. Further studies will be needed to assess the quantitative role of CD36 in foam cell formation in vivo.

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements.

Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day-1 tree-1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m2 of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s-1 that is equivalent to 112 mmol m-2 s-1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m-2 s-1 at the top and 150 mmol m-2 s-1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was -27.76±0.27‰ (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.

Photosynthetic capacity, chloroplast pigments, and mineral content of the previous year's spruce needles with and without the new flush: analysis of the forest-decline phenomenon of needle bleaching.

Spruce (Picea abies) damage in the Fichtelgebirge (FRG) occurs as needle bleaching and a depression of CO2 assimilation. Such injury may primarily result from the direct, above-ground effects of air pollution or indirect, below-ground changes in mineral uptake.Typically, the new flush of spruce needles is green and exhibits high photosynthetic capacity. Mies and Zöttl concluded that the older foliage is damaged when nutrients are withdrawn to supply the current year's needles. By removing the terminal buds of single branches in the spring, we produced an experimental set of the previous year's needles with greater mineral reserves than the control needles. During the course of the growing period, the performance of the experimental needles, which lacked competition from the new flush, was compared to that of the control needles of the same age-class on intact branches with the new flush.Throughout the experiment, chloroplast pigments of a healthy control tree were not affected by the elimination of the new flush. However, the chlorophyll and carotenoid content as well as the photosynthetic capacity of the previous year's needles on those branches of a heavily damaged tree where the new flush had been eliminated increased substantially. This increase was associated with an increase in minerals, which seemed to be deficient in the control needles with the new flush. Thus, in contrast to needles of the same age-class on intact branches with undisturbed new growth in the same atmospheric environment, the experimental needles escaped bleaching and a decrease in photosynthesis. It would seem that the bleaching and the loss in photosynthetic capacity typical of trees damaged by forest decline indirectly result from nutrient deficiencies through soil environment changes and/or root damage than directly from atmospheric pollutants.