Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

Hydraulic resistance in Acer saccharum shoots and its influence on leaf water potential and transpiration

A new method is presented for measuring whole-shoot hydraulic conductance, K(T) (kg s(-1) MPa(-1)). The method was also used to determine other conductance values in maple (Acer saccharum Marsh.) stem segments of differing diameter including: K(h) (absolute conductance or conductance per unit pressure gradient, kg s(-1) m MPa(-1)), K(s) (specific conductance or K(h) per unit wood area, kg s(-1) m(-1) MPa(-1)), and LSC (leaf specific conductance or K(h) per unit leaf area, kg s(-1) m(-1) MPa(-1)). A regression of K(T) versus stem basal diameter, D (m), gave K(T) = 5.998 x 10(-2) D(1.402) (R(2) = 0.986 for D from 0.001 to 0.1 m) and a regression for leaf area, A(L) (m(2)), gave A(L) = 4.667 x 10(3) D(2.007) (R(2) = 0.981 for D from 0.001 to 0.3 m). More than 50% of the resistance to water flow in large shoots (0.1 m in diameter and 8 to 10 m long) was contained in branches less than 0.012 m in diameter, i.e., in the distal 1.5 m of branches. We used the regressions to predict the steady state difference in pressure potential, P, between the base of a shoot of diameter D and the average pressure potential at the apices of the shoot; the relation is given by P = 7.781 x 10(4) E D(0.605), where E is the average evaporative flux density (kg s(-1) m(-2)) in the leaves attached to the shoot. After comparing the predictions of this equation to field observations of E and leaf water potential and stomatal conductance, we concluded that the hydraulic conductance of large maple shoots is sufficiently low to prevent maximum stomatal conductance in maple leaves.     

Fall lifting and long-term freezer storage of ponderosa pine seedlings: effects on post-storage leaf water potential, stomatal conductance, and root growth potential

Post-storage water relations, stomatal conductance, and root growth potential were examined in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings from high- and low-elevation seed sources that had been lifted either in October or November and freezer stored, or in March, and then grown hydroponically in a greenhouse for 31 days. Seedlings lifted in October had poor root initiation (< 17 new roots per seedling), low predawn leaf water potentials (< -1.5 MPa), and low stomatal conductance (7.10 mmol m(-2) s(-1)) compared with seedlings lifted in November or March. There was little difference in post-storage water relations and stomatal conductance between seedlings lifted in November and those lifted in March. Throughout the 31-day test, seedlings from the high-elevation seed source produced 3-9 times more new roots, had higher predawn leaf water potentials (-0.6 to -0.7 MPa versus -1.1 to -1.6 MPa), and 1.3-5 times greater stomatal conductance than seedlings from the low-elevation seed source. For all seedlings on Day 31, the number of new roots was significantly related to predawn leaf water potential (r(2) = 0.65) and stomatal conductance (r(2) = 0.82). Similarly, the dry weight of new roots per seedling on Day 31 accounted for a significant amount of the variation in predawn leaf water potential (r(2) = 0.81) and stomatal conductance (r(2) = 0.49).     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Interactive effects of nocturnal transpiration and climate change on the root hydraulic redistribution and carbon and water budgets of southern United States pine plantations

Deep root water uptake and hydraulic redistribution (HR) have been shown to play a major role in forest ecosystems during drought, but little is known about the impact of climate change, fertilization and soil characteristics on HR and its consequences on water and carbon fluxes. Using data from three mid-rotation loblolly pine plantations, and simulations with the process-based model MuSICA, this study indicated that HR can mitigate the effects of soil drying and had important implications for carbon uptake potential and net ecosystem exchange (NEE), especially when N fertilization is considered. At the coastal site (C), characterized by deep organic soil, HR increased dry season tree transpiration (T) by up to 40%, and such an increase affected NEE through major changes in gross primary productivity (GPP). Deep-rooted trees did not necessarily translate into a large volume of HR unless soil texture allowed large water potential gradients to occur, as was the case at the sandy site (S). At the Piedmont site (P) characterized by a shallow clay-loam soil, HR was low but not negligible, representing up to 10% of T. In the absence of HR, it was predicted that at the C, S and P sites, annual GPP would have been diminished by 19, 7 and 9%, respectively. Under future climate conditions HR was predicted to be reduced by up to 25% at the C site, reducing the resilience of trees to precipitation deficits. The effect of HR on T and GPP was predicted to diminish under future conditions by 12 and 6% at the C and P sites, respectively. Under future conditions, T was predicted to stay the same at the P site, but to be marginally reduced at the C site and slightly increased at the S site. Future conditions and N fertilization would decrease T by 25% at the C site, by 15% at the P site and by 8% at the S site. At the C and S sites, GPP was estimated to increase by 18% and by >70% under future conditions, respectively, with little effect of N fertilization. At the P site, future conditions would stimulate GPP by only 12%, but future conditions plus N fertilization would increase GPP by 24%. As a consequence, in all sites, water use efficiency was predicted to improve dramatically with future conditions. Modeling the effect of reduced annual precipitation indicated that limited water availability would decrease all carbon fluxes, including NEE and respiration. Our simulations highlight the interactive effects of nutrients and elevated CO(2), and showed that the effect of N fertilization would be greater under future climate conditions.     

Stomatal behavior of four woody species in relation to leaf-specific hydraulic conductance and threshold water potential

Midday stomatal closure is mediated by the availability of water in the soil, leaf and atmosphere, but the response to these environmental and internal variables is highly species specific. We tested the hypothesis that species differences in stomatal response to humidity and soil water availability can be explained by two parameters: leaf-specific hydraulic conductance (K(L)) and a threshold leaf water potential (Psi(threshold)). We used a combination of original and published data to estimate characteristic values of K(L) and Psi(threshold) for four common tree species that have distinctly different stomatal behaviors: black cottonwood (Populus trichocarpa Torr. & Gray.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), red alder (Alnus rubra Bong.) and western hemlock (Tsuga heterophylla (Raf.) Sarg.). We used the values to parameterize a simple, nonelastic model that predicts stomatal conductance by linking hydraulic flux to transpirational flux and maintaining Psi(leaf) above Psi(threshold). The model successfully predicted fundamental features of stomatal behavior that have been reported in the literature for these species. We conclude that much of the variation among the species in stomatal response to soil and atmospheric water deficits can be explained by K(L) and Psi(threshold). The relationship between Psi(threshold) and xylem vulnerability to cavitation differed among these species.     

Model-based analysis of the spatial variability and long-term trends of soil drought at Scots pine stands in northeastern Germany

Soil water availability determines the vitality of trees and forest stands to a large degree. Over the last decades, an increasing number of drought spells has been observed in several parts of Europe. Our study aims to estimate long-term trends of soil drought at Scots pine (Pinus sylvestris L.) stands along the prevailing climatic gradient in Mecklenburg-Western Pomerania, northeastern Germany. To this end, soil water balance simulations were carried out at 24 sites over the period from 1951 to 2009 with the physically based model LWF-BROOK90. As a threshold for soil water stress, we used 40% of relative extractable water (REW). The results indicated an increased number of drought days further east, together with declining totals of precipitation. However, specific site conditions had a large influence on the occurrence of soil drought, partly overriding the climatic differences across the study area. Soil drought has distinctly increased in the recent past, both in duration and in intensity, affecting the eastern sites more than the western sites. The increased soil dryness could be attributed to higher atmospheric evaporative demand due to higher temperatures, as well as slightly lower precipitation sums during the summer months. To mitigate the negative effects of future climate change, adaptation measures should preferably be conducted in the eastern parts of northeastern Germany.     

Influence of rate of change in stomatal conductance to fluctuating irradiance on estimates of daily water use by Pinus taeda leaves

Pinus taeda L. stomata respond slowly to changes in irradiance. Because incident irradiance on a leaf varies constantly, the rate of change in stomatal conductance to fluctuating irradiance may have a large effect on plant water use. We estimated total daily water use of Pinus taeda foliage for 10 days with very different irradiance patterns, assuming that rates of stomatal opening and closing were similar. To determine how the most extreme imbalance in rates of stomatal opening and closing affects estimates of water use, we also estimated total daily water use assuming instantaneous stomatal opening and a realistic rate of stomatal closing. Total daily water use was calculated by summing estimates of transpiration based on irradiance and vapor pressure deficit measured every minute at locations atop and within a canopy. Estimates of total daily water use calculated on the basis of realistic rates of stomatal opening and closing were similar to estimates calculated assuming instantaneous stomatal change (mean difference between methods of calculation was less than 0.2%). Estimates of total daily water use assuming instantaneous stomatal opening and a realistic rate of closing differed from estimates of total daily water use based on similar rates of stomatal opening and closing. The discrepancy was greater within the canopy (mean difference 6%) than at the top of the canopy (mean difference 1%). Calculation of mean daily conductance from mean daily irradiance, without accounting for minute-by-minute variations in irradiance, resulted in overestimations of daily stomatal conductance (13% mean error) and the magnitude of the error was directly related to the variation in irradiance for that day. We conclude that, provided variation in irradiance is accounted for, rates of stomatal opening and closing have little effect on estimates of daily water use.     

Functional convergence in hydraulic architecture and water relations of tropical savanna trees: from leaf to whole plant

Functional convergence in hydraulic architecture and water relations, and potential trade-offs in resource allocation were investigated in six dominant neotropical savanna tree species from central Brazil during the peak of the dry season. Common relationships between wood density and several aspects of plant water relations and hydraulic architecture were observed. All species and individuals shared the same negative exponential relationship between sapwood saturated water content and wood density. Wood density was a good predictor of minimum (midday) leaf water potential and total daily transpiration, both of which decreased linearly with increasing wood density for all individuals and species. With respect to hydraulic architecture, specific and leaf-specific hydraulic conductivity decreased and the leaf:sapwood area ratio increased more than 5-fold as wood density increased from 0.37 to 0.71 g cm(-3) for all individuals and species. Wood density was also a good predictor of the temporal dynamics of water flow in stems, with the time of onset of sap flow in the morning and the maximum sap flow tending to occur progressively earlier in the day as wood density increased. Leaf properties associated with wood density included stomatal conductance, specific leaf area, and osmotic potential at the turgor loss point, which decreased linearly with increasing wood density. Wood density increased linearly with decreasing bulk soil water potential experienced by individual plants during the dry season, suggesting that wood density was greatest in individuals with mostly shallow roots, and therefore limited access to more abundant soil water at greater depths. Despite their taxonomic diversity and large intrapopulation differences in architectural traits, the six co-occurring species and their individuals shared similar functional relationships between all pairs of variables studied. Thus, rather than differing intrinsically in physiological responsiveness, the species and the individuals appeared to have distinct operating ranges along common physiological response curves dictated by plant architectural and structural features. The patterns of water uptake and access to soil water during the dry season appeared to be the main determinant of wood density, which constrained evolutionary options related to plant water economy and hydraulic architecture, leading to functional convergence in the neotropical savanna trees studied.     

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.     

Comparison of the measurements of leaf water potential between a hydraulic press and a pressure chamber in six Sahelian woody species

The aim of this methodological study was to quantify differences between water potential measured with a pressure chamber (PC) and with a hydraulic press (HP) in six north Sahelian dominant species of the woody strata across the range of their local environmental conditions in the Malian Gourma. Mean annual rainfall is 372 mm, falling from June to September, followed by 8–10 months of dry season. The daily course of Leaf Water Potential (LWP) was monitored in 2–6 m tall healthy individuals. Water potential measured with the two instruments were statistically comparable (R2 > 40%) except in A. senegal. However, the HP under-estimated LWP and revealed smaller ranges of water potential than the PC. In the Sahelian shrubs studied here, for the precise measurement of a water potential gradient in the soil–plant-atmosphere continuum and for inter-specific comparisons, the PC is more appropriate than the HP. However, the HP may be useful for intra-species comparison in large sampled fields, since calibrations will be checked across a wider range of dates and a large number of sites.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Analysis on the daily courses of water potential of nine woody species from Cerrado vegetation during wet season

IntroductionAS the concept of water pot6ntial (% was introduced by Saltyer (1967), there were some argumentson it among some physiologist. Some physiologistsopposed it beCause they believed Tvalue might notbe the beSt parameter for describing physiologicalprocesses. However, this concept has been accept6dand used by most plant physiologists for many years(Kramer 1988, Schulze et al. 1988, Passioura 1988,Boyer 1989).ACcording tO our experience (Prado et al. 1994,1995), we believed-t.hat th…     

Influence of stand,site and meteorological variables on the maximum leaf area index of beech,oak and Scots pine

Different multiple linear regression models of maximum leaf area index (LAI_max) based on stand characteristics, site quality, meteorological variables and their combinations were constructed and cross-validated for three economically important tree species in Flanders, Belgium: European beech (Fagus sylvatica L.), Pedunculate oak (Quercus robur L.) and Scots pine (Pinus sylvestris L.). The models were successfully tested on similar datasets of experimental sites across Europe. For each species, ten homogeneous and mature stands were selected, covering the species’ entire stand productivity range based on an a priori site index classification. LAI_max was derived from measurements of leaf area index (LAI) made by means of hemispherical digital photography over the whole growing season (mid-April till end October 2008). Species-specific models of LAI_max for beech and oak were mostly driven by management practice affecting stand characteristics and tree growth. Tree density and dominant height were main predictors for beech, while stand age and tree-ring growth were important in the oak models. Scots pine models were more affected by site quality and meteorological variables. The beech meteorological model showed very good agreement with LAI at several European sites. Scots pine’s stand model predicted well LAI across Europe. Since the species-specific models did not share common predictors, generic models of LAI_max were developed for the 30 studied sites. Dominant height was found to be the best predictor in those generic models. As expected, they showed a lower predictive performance than species-specific ones.     

两种松树种子发芽率与生活力测定试验

以湿地松和马尾松2种松类树种种子为试材,进行发芽率与生活力测定试验。结果表明：湿地松和马尾松种子均以沙床上的发芽率最高,不同发芽床上湿地松种子的发芽率差异较大,马尾松种子的发芽率差异相对较小;湿地松和马尾松种子的发芽率与生活力之间均存在极显著正相关,相关系数（R）分别为0．9931和0．9913;湿地松种子发芽率与生活力的回归方程为y=0．9125x-1．7247,马尾松种子发芽率与生活力的回归方程为y=0．9703x-3．4769。     

Daily sap flow and maximum daily trunk shrinkage measurements for diagnosing water stress in early maturing peach trees during the post-harvest period

We compared the sensitivity of two continuously recorded plant-based water stress indicators (sap flow, SF, and maximum daily trunk shrinkage, MDS) to detect changes in the water status of 4-year-old early maturing peach trees (Prunus persica (L.) Batsch cv. Flordastar grafted on GF-677 peach rootstock) during a cycle of deficit irrigation and recovery. The feasibility of obtaining SF and MDS reference equations for use in irrigation scheduling during the post-harvest period was also studied in trees irrigated in excess of crop water requirements. We found that MDS was a more sensitive and reliable detector of changes in plant water status than SF, making it a more precise tool for irrigation scheduling. Baseline relationships between SF or MDS and the climatic variables (air temperature, vapor pressure deficit (VPD) and crop reference evapotranspiration (ETo)) were established, despite some scatter in the data. Among the climatic variables, SF correlated more closely with ETo, whereas MDS correlated more closely with mean daily air temperature (T (m)). The fits of the regressions between MDS and ETo, midday air temperature and T (m) for individual periods were better than those obtained in the overall regressions, confirming that daily stem diameter variations must be considered not only in the context of plant water status but also in the context of plant carbon status.     

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.