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1.
The impacts of thinning, fertilization and crown position on seasonal growth of current-year shoots and foliage were studied in a 13-year-old loblolly pine (Pinus taeda L.) plantation in the sixth post-treatment year (1994). Length of new flushes, and their needle length, leaf area, and oven-dry weight were measured in the upper and lower crown from March through November. Total shoot length was the cumulative length of all flushes on a given shoot and total leaf area per shoot was the sum of leaf areas of the flushes.

By the end of June, first-flush foliage reached 70% of the November needle length (14.3 cm) and 65% of the final leaf area (15.0 cm2). Cumulative shoot length of first- and second-flush shoots achieved 95% of the annual length (30.3 cm), whereas total leaf area per shoot was 55% of the final value (75.3 dm2). Fertilization consistently stimulated fascicle needle length, dry weight, and leaf area in the upper crown. Mean leaf area of upper-crown shoots was increased by 64% six years after fertilization. A significant thinning effect was found to decrease mean leaf area per shoot in the crown. For most of the growing season, the thinned-fertilized trees produced substantially more leaf area per shoot throughout the crown than the thinned-nonfertilized trees. These thinned-fertilized trees also had greater needle length and dry weight, longer first flush shoots, and more leaf area per flush than trees in the thinned-nonfertilized plots. Needle length and leaf area of first flush shoots between April and July were linearly related to previous-month canopy air temperature (Ta). Total shoot length strongly depended on vertical light gradient (PPFD) within the canopy, whereas shoot leaf area was a function of both PPFD and Ta. Thus, trees produced larger and heavier fascicles, more and longer flush shoots, and more leaf area per shoot in the upper crown than the lower crown. We conclude that thinning, fertilization, and crown position regulate annual leaf area production of current-year shoots largely by affecting the expansion of first flush shoots and their foliage during the first half of the growing season.  相似文献   


2.
Total foliage dry mass and leaf area at the canopy hierarchical level of needle, shoot, branch and crown were measured in 48 trees harvested from a 14-year-old loblolly pine (Pinus taeda L.) plantation, six growing seasons after thinning and fertilization treatments.

In the unthinned treatment, upper crown needles were heavier and had more leaf area than lower crown needles. Branch- and crown-level leaf area of the thinned trees increased 91 and 109%, respectively, and whole-crown foliage biomass doubled. The increased crown leaf area was a result of more live branches and foliated shoots and larger branch sizes in the thinned treatment. Branch leaf area increased with increasing crown depth from the top to the mid-crown and decreased towards the base of the crown. Thinning stimulated foliage growth chiefly in the lower crown. At the same crown depth in the lower crown, branch leaf area was greater in the thinned treatment than in the unthinned treatment. Maximum leaf area per branch was located nearly 3–4 m below the top of the crown in the unthinned treatment and 4–5 m in the thinned treatment. Leaf area of the thinned-treatment trees increased 70% in the upper crown and 130% in the lower crown. Fertilization enhanced needle size and leaf area in the upper crown, but had no effect on leaf area and other variables at the shoot, branch and crown level. We conclude that the thinning-induced increase in light penetration within the canopy leads to increased branch size and crown leaf area. However, the branch and crown attributes have little response to fertilization and its interaction with thinning.  相似文献   


3.
In order to quantify the effects of thinning on biochemical photosynthesis parameters and changes in leaf nitrogen contents associated with the process of crown reclosure, the maximum rate of carboxylation (V cmax), the leaf nitrogen concentration per unit area (N a), and the photosynthetic photon flux density (PPFD) were measured at four crown heights in both thinned (1500 trees ha−1) and unthinned control (3000 trees ha−1) stands of ten-year-old Chamaecyparis obtusa (36°3′N, 140°7′E) trees during four consecutive growing seasons after thinning. Thinning increased V cmax in the lower and middle crowns in the first year after thinning, and leaves in the lower crown of the thinned stand maintained high V cmax for four years, whereas they abscised in the second year in the control stand. Significant increases in V cmax were detected even in the upper crowns of trees in the thinned stand in the second year. Thinning did not affect N a at any of the crown positions in the first year, but significantly increased N a in the middle crowns from the second year after thinning. Thus, the redistribution of nitrogen between leaves, driven by increases in light and nutrient availability due to the 50% thinning, appears to have enhanced photosynthetic rates in the thinned stand. Thinning also significantly affected the slope of the linear relationship between N a and V cmax initially after thinning, but its effect on this relationship was negligible after the second year. These quantitative results may be used to simplify the estimation of the likely effects of management practices on carbon fixation in forest canopies.  相似文献   

4.
The stand density of a forest affects the vertical distribution of foliage. Understanding the dynamics of this response is important for the study of crown structure and function, carbon-budget estimation, and forest management. We investigated the effect of tree density on the vertical distribution of foliage, branch, and stem growth, and ratio of biomass increment in aboveground tissues; by monitoring all first-order branches of five trees each from thinned and unthinned control stands of 10-year-old Chamaecyparis obtusa for four consecutive years. In the control stand, the foliage crown shifted upward with height growth but the foliage quantity of the whole crown did not increase. In addition, the vertical distribution of leaf mass shifted from lower-crown skewed to upper-crown skewed. In the thinned stand in contrast, the foliage quantity of individual crowns increased two-fold within 4 years, while the vertical distribution of leaf mass remained lower-crown skewed. The two stands had similar production rates, numbers of first-order branches per unit of tree height, and total lengths of first-order branches. However, the mortality rate of first-order branches and self-pruning within a first-order branch were significantly higher in the control stand than in the thinned stand, which resulted in a higher ratio of biomass increment in branch. Thinning induced a higher ratio of biomass increment in foliage and lower in branch. The increased foliage quantity and variation in ratio of biomass increment after thinning stimulated stem growth of residual trees. These results provide information that will be useful when considering thinning regimes and stand management.  相似文献   

5.
Several heavy wet snowfalls occurred during 2007-2009 across a broad-scale thinning and fertilization experiment to bring overstocked juvenile lodgepole pine (Pinus contorta var. latifolia) in the foothills of Alberta, Canada into an intensive management regime. We examined the bending and breakage of trees in relation to thinning and fertilization and used a multimodel information-theoretic approach to model stand and tree level predictors of snow damage. Fertilized stands suffered the greatest amount of snow damage, and this was most noteworthy when stands were also thinned; here 22% (17% broken stems) of trees were damaged compared to 8% (4% broken stems) in the thinned and unfertilized stands. At the stand level, needle weight and crown cover were reliable predictors of snow damage. At the tree level, separate models were developed for each combination of thinning and fertilization. All models used total tree volume; usually the smaller trees in the stands were more susceptible to damage but in the thinned and fertilized stands larger but slender trees with large asymmetrical crowns tended to be damaged. Also, trees with lower total stem volume were more susceptible to damage. Only in the thinned and fertilized stands were variables related to crown shape and asymmetry important predictors of snow damage. We conclude that snow damage is an important agent for self-thinning in unthinned stands and fertilization tends to exacerbate damage because of increase in foliage size. In areas with regular occurrence of heavy snow, we do not recommend fertilization at the same time as thinning, as the larger and more economically important trees in the stand are at risk.  相似文献   

6.
Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.  相似文献   

7.
Physiological parameters were measured under natural light conditions and needle orientation from towers and walkways erected in the canopy of a loblolly pine (Pinus taeda L.) plantation. Four silvicultural treatments were randomly assigned to the twelve plots in the fall of 1988. Plots were thinned to a density of 731 trees per hectare or left unthinned, at a density of 2990 trees per hectare. The plots were left unfertilized or fertilized with 744 kg/ha of diammonium triple superphosphate was applied. During the fifth growing season (1993) following thinning and fertilization, needle level physiology was not different with respect to the thinning treatment for fertilized or unfertilized plots. In contrast, upper crown levels within the fertilized and unfertilized plots had significantly higher light levels and photosynthetic rates than lower crown foliage. Light levels were greater in the thinned, fertilized plots than in the unthinned, fertilized plots. In contrast, no effect of thinning on canopy light levels was found in the unfertilized plots. Within crown variation in photosynthesis was strongly dependent on canopy light levels. A strong interaction of canopy level with thinning was apparent for net photosynthesis. Loblolly pine, being a shade intolerant species, showed only small physiological differences between needles from different parts of the crown. Because of the variability found in this study, more extensive sampling is needed to correctly describe the physiology of a forest canopy with adequate precision.  相似文献   

8.
Scaling leaf-level measurements to estimate carbon gain of entire leaf crowns or canopies requires an understanding of the distribution of photosynthetic capacity and corresponding light microenvironments within a crown. We have compared changes in the photosynthetic light response and nitrogen (N) content (per unit leaf area) of Pinus contorta Dougl. ssp. latifolia Engelm. (lodgepole pine) leaves in relation to their age and light microenvironment. The vertical gradient in integrated daily photosynthetic photon flux density (PPFD) from the upper to the lower crown of lodgepole pine was similar in magnitude to the horizontal gradient in daily PPFD along shoots from young to old leaves. The relationship between light-saturated net photosynthesis (A(max)) and daily PPFD was significant for both young and old leaves. However, old leaves had a lower A(max) than young leaves in a similar daily irradiance regime. For leaves of all ages from throughout the crown, A(max) was linearly related to the estimated daily net carbon gain that leaves could achieve in their natural PPFD environment (estimated A(day)) (r(2) = 0.84, P < 0.001, n = 39), indicating that estimated A(day) may be dominated by carbon fixed when leaves are light-saturated and operating at A(max). Comparison of the PPFD required to achieve A(max) and the PPFD available to the leaves showed that all of the measured leaves (n = 39), regardless of their position in the crown or age, were in light environments that could light-saturate photosynthesis for a similar proportion of the day. For all data pooled, foliar N was weakly correlated with daily PPFD. Analyzing each leaf age class separately showed that foliar N was significantly related to daily PPFD, A(max), and estimated A(day) for the youngest leaves but not for middle-aged or old leaves. Therefore, the general theory that foliar N is allocated within a crown according to total daily light availability was supported only for young (1-4 years old) leaves in this study.  相似文献   

9.
Biophysical constraints on leaf expansion in a tall conifer   总被引：3，自引：0，他引：3  
The physiological mechanisms responsible for reduced extension growth as trees increase in height remain elusive. We evaluated biophysical constraints on leaf expansion in old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Needle elongation rates, plastic and elastic extensibility, bulk leaf water (Psi(L)) and osmotic (Psi(pi)) potential, bulk tissue yield threshold and final needle length were characterized along a height gradient in crowns of > 50-m-tall trees during the period between bud break and full expansion (May to June). Although needle length decreased with increasing height, there was no height-related trend in leaf plastic extensibility, which was highest immediately after bud break (2.9%) and declined rapidly to a stable minimum value (0.3%) over a 3-week period during which leaf expansion was completed. There was a significant positive linear relationship between needle elongation rates and plastic extensibility. Yield thresholds were consistently lower at the upper and middle crown sampling heights. The mean yield threshold across all sampling heights was 0.12 +/- 0.03 MPa on June 8, rising to 0.34 +/- 0.03 MPa on June 15 and 0.45 +/- 0.05 MPa on June 24. Bulk leaf Psi(pi) decreased linearly with increasing height at a rate of 0.004 MPa m(-1) during the period of most rapid needle elongation, but the vertical osmotic gradient was not sufficient to fully compensate for the 0.015 MPa m(-1) vertical gradient in Psi(L), implying that bulk leaf turgor declined at a rate of about 0.011 MPa m(-1) increase in height. Although height-dependent reductions in turgor appeared to constrain leaf expansion, it is possible that the impact of reduced turgor was mitigated by delayed phenological development with increasing height, which resulted in an increase with height in the temperature during leaf expansion.  相似文献   

10.
We studied the effects of two types of selective thinning on beech stands formed by a shelterwood cut in 1910 — with lower number of crop trees and higher thinning intensity (T1) and higher number of crop trees with lower thinning intensity (T2). The stands were thinned in 1980, 1991 and 2001. Despite a lower stand density after thinning, the annual basal area increments of thinned stands in both thinning periods (1980–1991 and 1991–2002) were around 20% higher compared to those of the control (unthinned) stands. The mean annual basal area increment of dominant trees was 30–56% larger in the thinned plots compared to the control plots. Of 176 initial crop trees in the T1, 72% were chosen again during the last thinning. In the T2, 258 crop trees were chosen in the first thinning, and only 62% of these trees were chosen again during the last thinning. Only crown suppression and diameter classes of crop trees significantly influenced their basal area increment when diameter classes, crown size, crown suppression, and social status were tested. In the thinned stands, the dominant trees are more uniformly distributed if compared to the dominant trees in the control plots. Finally, the herbaceous cover and the species diversity were higher in the thinned plots.  相似文献   

11.
We studied the relief of water stress associated with fruit thinning in pear (Pyrus communis L.) trees during drought to determine what mechanisms, other than stomatal adjustment, were involved. Combinations of control irrigation (equal to crop water use less effective rainfall) and deficit irrigation (equal to 20% of control irrigation), fruit load (unthinned and thinned to 40 fruits per tree) and root pruning (pruned and unpruned) treatments were applied to pear (cv. 'Conference') trees during Stage II of fruit development. Daily patterns of midday stem water potential (Psi(stem)) and leaf conductance to water vapor (g(l)) of deficit-irrigated trees differed after fruit thinning. In response to fruit thinning, gl progressively declined with water stress until 30 days after fruit thinning and then leveled off, whereas the effects of decreased fruit load on Psi(stem) peaked 30-40 days after fruit thinning and then tended to decline. Soil water depletion was significantly correlated with fruit load during drought. Our results indicate that stomatal adjustment and the resulting soil water conservation were the factors determining the Psi(stem) response to fruit thinning. However, these factors could not explain differences in daily patterns between g(l) and Psi(stem) after fruit thinning. In all cases, effects of root pruning treatments on Psi(stem) in deficit-irrigated trees were transitory (Psi(stem) recovered from root pruning in less than 30 days), but the recovery of Psi(stem) after root pruning was faster in trees with low fruit loads. This behavior is compatible with the concept that the water balance (reflected by Psi(stem) values) was better in trees with low fruit loads compared with unthinned trees, perhaps because more carbon was available for root growth. Thus, a root growth component is hypothesized as a mechanism to explain the bimodal Psi(stem) response to fruit thinning during drought.  相似文献   

12.
Effects of needle water potential (Psi(l)) on gas exchange of Scots pine (Pinus sylvestris L.) grown for 4 years in open-top chambers with elevated temperature (ET), elevated CO(2) (EC) or a combination of elevated temperature and CO(2) (EC + ET) were examined at a high photon flux density (PPFD), saturated leaf to air water vapor pressure deficit (VPD) and optimal temperature (T). We used the Farquhar model of photosynthesis to estimate the separate effects of Psi(l) and the treatments on maximum carboxylation efficiency (V(c,max)), ribulose-1,5-bisphosphate regeneration capacity (J), rate of respiration in the light (R(d)), intercellular partial pressure of CO(2) (C(i)) and stomatal conductance (G(s)). Depression of CO(2) assimilation rate at low Psi(l) was the result of both stomatal and non-stomatal limitations on photosynthetic processes; however, stomatal limitations dominated during short-term water stress (Psi(l) < -1.2 MPa), whereas non-stomatal limitations dominated during severe water stress. Among the nonstomatal components, the decrease in J contributed more to the decline in photosynthesis than the decrease in V(c,max). Long-term elevation of CO(2) and temperature led to differences in the maximum values of the parameters, the threshold values of Psi(l) and the sensitivity of the parameters to decreasing Psi(l). The CO(2) treatment decreased the maximum values of V(c,max), J and R(d) but significantly increased the sensitivity of V(c,max), J and R(d) to decreasing Psi(l) (P < 0.05). The effects of the ET and EC + ET treatments on V(c,max), J and R(d) were opposite to the effects of the EC treatment on these parameters. The values of G(s), which were measured simultaneously with maximum net rate of assimilation (A(max)), declined in a curvilinear fashion as Psi(l) decreased. Both the EC + ET and ET treatments significantly decreased the sensitivity of G(s) to decreasing Psi(l). We conclude that, in the future, acclimation to increased atmospheric CO(2) and temperature could increase the tolerance of Scots pine to water stress.  相似文献   

13.
Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).  相似文献   

14.
Mori A  Takeda H 《Tree physiology》2004,24(6):661-670
Light-related plasticity of crown morphology and within-crown characteristics were investigated in understory sun and shade saplings of three codominant subalpine conifers, Abies mariesii M.T. Mast., Abies veitchii Lindl. and Picea jezoensis var. hondoensis (Mayr) Rehd. Compared with those of sun saplings, current-year shoots of shade saplings allocated less biomass to needles, resulting in less dense needle packing and hence less mutual needle shading. The proportion of lateral branch biomass in foliage was either similar in sun and shade saplings or greater in shade saplings, depending on the species, suggesting that, over the lifetime of a branch, greater needle longevity in shade compensates for reduced biomass investment in needles of current-year shoots of shade saplings. Saplings with slower-growing branches tended to have greater needle life spans, suggesting that plasticity of branch growth rate and plasticity of needle life span are interdependent. Both Abies species showed greater light-related plasticity of needle life span and branch growth than P. jezoensis. The greater shade tolerance of the Abies species derives from their broad flattened crowns with slow-growing branches. This type of crown development incurs substantial support costs, but the long needle life span of shade saplings of the Abies species compensates, at least in part, for their low annual investment in foliage, especially in the case of A. mariesii, which has a longer needle life span and slower-growing and stouter branches than A. veitchii. Compared with the Abies species, P. jezoensis had a less plastic crown morphology, and less variability of needle life span and branch growth in response to light, resulting in lower shade tolerance. However, compared with the flattened crown of Abies shade saplings, the conical crown of P. jezoensis saplings imposes a smaller support cost, making this species better adapted to rapid height growth than to survival in shade.  相似文献   

15.
We compared leaf gas exchange and water potential among the dominant tree species and major size classes of trees in an upland, pine-oak forest in northern Arizona. The study included old-growth Gambel oak (Quercus gambelii Nutt.), and sapling, pole, and old-growth ponderosa pines (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Old-growth oak had higher predawn leaf water potential (Psi(leaf)) than old-growth pine, indicating greater avoidance of soil water stress by oak. Old-growth oak had higher stomatal conductance (G(w)), net photosynthetic rate (P(n)), and leaf nitrogen concentration, and lower daytime Psi(leaf) than old-growth pine. Stomatal closure started at a daytime Psi(leaf) of about -1.9 MPa for pine, whereas old-growth oak showed no obvious reduction in G(w) at Psi(leaf) values greater than -2.5 MPa. In ponderosa pine, P(n) and G(w) were highly sensitive to seasonal and diurnal variations in vapor pressure deficit (VPD), with similar sensitivity for sapling, pole, and old-growth trees. In contrast, P(n) and G(w) were less sensitive to VPD in Gambel oak than in ponderosa pine, suggesting greater tolerance of oak to atmospheric water stress. Compared with sapling pine, old-growth pine had lower morning and afternoon P(n) and G(w), predawn Psi(leaf), daytime Psi(leaf), and soil-to-leaf hydraulic conductance (K(l)), and higher foliar nitrogen concentration. Pole pine values were intermediate between sapling and old-growth pine values for morning G(w) and daytime Psi(leaf), similar to sapling pine for predawn Psi(leaf), and similar to old-growth pine for morning and afternoon P(n), afternoon G(w), K(l), and foliar nitrogen concentration. For the pines, low predawn Psi(leaf), daytime Psi(leaf), and K(l) were associated with low P(n) and G(w). Our data suggest that hydraulic limitations are important in reducing P(n) in old-growth ponderosa pine in northern Arizona, and indicate greater avoidance of soil water stress and greater tolerance of atmospheric water stress by old-growth Gambel oak than by old-growth ponderosa pine.  相似文献   

16.
Water use, hydraulic properties and xylem vulnerability to cavitation were studied in the coffee (Coffea arabica L.) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown morphology and total leaf surface area. Sap flow, stomatal conductance (g(s)), crown conductance (g(c)), apparent hydraulic conductance of the soil-leaf pathway (G(t)), leaf water potential (Psi(L)) and xylem vulnerability to loss of hydraulic conductivity were assessed under well-watered conditions and during a 21-day period when irrigation was withheld. Sap flow, g(c), and G(t) were greatest in Typica both with and without irrigation, lowest in San Ramon, which was relatively unresponsive to the withholding of irrigation, and intermediate in Yellow Caturra. The cultivars had similar g(s) when well watered, but withholding water decreased g(s) more in Typica and Yellow Caturra than in San Ramon. Typica had substantially lower Psi(L) near the end of the unirrigated period than the other cultivars (-2.5 versus -1.8 MPa), consistent with the relatively high sap flow in this cultivar. Xylem vulnerability curves indicated that Typica was less susceptible to loss of hydraulic conductivity than the other cultivars, consistent with the more negative Psi(L) values of Typica in the field during the period of low soil water availability. During soil drying, water use declined linearly with relative conductivity loss predicted from vulnerability curves. However, cultivar-specific relationships between water use and predicted conductivity loss were not observed because of pronounced hysteresis during recovery of water use following soil water recharge. All cultivars shared the same functional relationship between integrated daily sap flow and G(t), but they had different operating ranges. The three cultivars also shared common functional relationships between hydraulic architecture and water use despite consistent differences in water use under irrigated and dry soil conditions. We conclude that hydraulic architectural traits, rate of water use per plant and crown architecture are important determinants of short- and long-term variations in the water balance of Coffea arabica.  相似文献   

17.
We tested the effects of thinning on allometry and needle-age distribution in natural stands of Abies sachalinensis Masters by comparing a thinned stand to an unthinned, control stand. Specifically, we attempted to clarify how allometry was altered after a thinning. We assumed that the needle-age distribution of trees in the thinned stand would show a younger composition than in the control stand, given the effect of improved light conditions on needle dynamics following a thinning. These investigations were conducted in dense Abies stands located in central Hokkaido, northern Japan, 19 years post-thinning. In the thinned stand, the ratio of individual needle mass to stem mass increased significantly, as compared to the control. A difference in the H–DBH relationship between the stands was probably related to this tendency. Mean needle age of trees differed significantly between the two stands, and the thinned stand showed a younger needle age than the control. Within each stand, dominant trees showed older mean needle age than codominant or suppressed trees. These tendencies may have been caused by differential needle dynamics affected by light conditions in the stands, and by different crown positions among the trees within a stand. In summary, trees in the thinned stand showed increased growth rates after thinning, which were caused by increased needle mass, younger composition of needles, and improved light conditions.  相似文献   

18.
It is a well-described phenomenon that plant leaves respond to changes in light intensity and duration by adjusting leaf hydraulic efficiency, and there is current consensus that up- or down-regulation of water channels (aquaporins) in the plasma membrane of the bundle sheath and mesophyll cells play a central role in the underlying mechanisms. Recently, experimental evidence has been provided also for light-mediated changes of stem hydraulic conductance (K(stem)) in field-grown laurel plants. This effect was attributed to differences in potassium ion concentration of xylem sap as a function of light conditions. In the present article, we report evidence obtained in silver birch (Betula pendula Roth), supporting the concept of light-mediated modulation of K(stem). Both canopy position (long-term effect) and current photosynthetic photon flux density (PPFD; short-term effect) had a significant impact (P < 0.001) on K(stem) measured in shoots taken from the lower (shade shoots) and upper (sun shoots) third of the crowns of ～25-year-old trees growing in a natural forest stand. The shade shoots responded more sensitively to light manipulation: K(stem) increased by 51% in shade shoots and 26% in sun shoots when PPFD increased from 70 to 330 μmol m?2 s?1. In 4-year-old trees growing in a dense experimental plantation, K(stem), specific conductivity of branch-wood (k(bw)) and potassium ion concentration ([K(+)]) in xylem sap varied in accordance with canopy position (P < 0.001). Both K(stem) and k(bw) increased considerably with light availability, increasing within the tree crowns from bottom to top; there was a strong relationship between mean values of K(stem) and [K(+)] in hydraulically sampled branches.  相似文献   

19.
Leaves, the distal section of the soil-plant-atmosphere continuum, exhibit the lowest water potentials in a plant. In contrast to angiosperm leaves, knowledge of the hydraulic architecture of conifer needles is scant. We investigated the hydraulic efficiency and safety of Pinus pinaster needles, comparing different techniques. The xylem hydraulic conductivity (k(s)) and embolism vulnerability (P(50)) of both needle and stem were measured using the cavitron technique. The conductance and vulnerability of whole needles were measured via rehydration kinetics, and Cryo-SEM and 3D X-ray microtomographic observations were used as reference tools to validate physical measurements. The needle xylem of P. pinaster had lower hydraulic efficiency (k(s)?=?2.0?×?10(-4) m(2) MPa(-1) s(-1)) and safety (P(50)?=?-?1.5 MPa) than stem xylem (k(s)?=?7.7?×?10(-4) m(2) MPa(-1) s(-1); P(50)?=?-?3.6 to?-?3.2 MPa). P(50) of whole needles (both extra-vascular and vascular pathways) was?-?0.5 MPa, suggesting that non-vascular tissues were more vulnerable than the xylem. During dehydration to?-?3.5 MPa, collapse and embolism in xylem tracheids, and gap formation in surrounding tissues were observed. However, a discrepancy in hydraulic and acoustic results appeared compared with visualizations, arguing for greater caution with these techniques when applied to needles. Our results indicate that the most distal parts of the water transport pathway are limiting for hydraulics of P. pinaster. Needle tissues exhibit a low hydraulic efficiency and low hydraulic safety, but may also act to buffer short-term water deficits, thus preventing xylem embolism.  相似文献   

20.
A 23 factorial experimental testing the effects of urea (N), superphosphate (P) and two thinning types (diagonal-line thinning and thinning from below) on the basal-are increment of 18-year-old Pinus radiata (D. Don) is described. Although the trees had received localized applications of superphosphate shortly after establishment they were considered at the age of 18 years to be phosphate-deficient because concentrations of phosphorus in the foliage were low (0.065% P). Application of superphosphate (200 kg P ha−1) increased concentrations in the foliage (0.132% P) and increased the 7-year basal-area increment by approximately 70%. No growth response was obtained when urea (476 kg N ha−1) was applied alone. When both fertilizers were applied there was a further increase in basal-area increment of 81% on line-thinned plots and 26% on plots which had been thinned from below. This interaction reversed the normal trend in which plots thinned from below, by virtue of their greater initial basal area, produced 11–16% more increment than those which had been line-thinned.

A model of tree growth using initial basal area and a competition index as independent variables was adequate to describe differences in basal-area growth between thinning types except where urea and superphosphate had both been applied. In the case of N + P-treated plots, the reciprocal of the competition index was required as an additional variable. Analysis of the development of the growth responses over time indicated that application of superphosphate had produced sustained improvement in growth rates but that the response to urea occurred only in some growing-seasons. In line-thinned plots the response to urea in addition to phosphate occurred in the 2nd, 3rd and 4th growing-seasons, but the response occurred only in the 2nd and 4th growing-seasons in plots thinned from below. It is argued that this differential response between thinning types can be attributed to differences in stand density and water availability. In order to maximise the growth response to added nitrogen it is important to reduce competition by paying particular attention to both tree spacing and residual basal area.  相似文献   


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