干旱胁迫对真桦光合特性及渗透调节物质含量的影响

以真桦盆栽幼苗为试材,研究干旱胁迫对真桦叶片光合作用和渗透调节物质含量的影响。结果表明:干旱胁迫使真桦幼苗净光合速率、蒸腾速率和气孔导度显著降低,降低幅度随胁迫程度的加剧而加大。胁迫初期,轻度和中度干旱胁迫组真桦幼苗净光合速率降低的同时气孔导度和胞间CO2浓度同时降低,光合速率降低的主要原因是气孔因素;胁迫中期和后期,各干旱处理组光合速率降低的同时胞间CO2浓度均显著升高,非气孔限制是净光合速率降低的主要原因。真桦在干旱胁迫过程中,可溶性蛋白变化不显著,可溶性糖含量增加,为主要的渗透调节物质。     

Effects of potassium supply on limitations of photosynthesis by mesophyll diffusion conductance in Carya cathayensis

Potassium (K) influences the photosynthesis process in a number of ways; however, the mechanisms underlying the photosynthetic response to differences in K supply are not well understood. Concurrent measurements of gas exchange and chlorophyll fluorescence were made to investigate the effect of K nutrition on photosynthetic efficiency and mesophyll conductance (g(m)) in hickory seedlings (Carya cathayensis Sarg.) in a greenhouse. The results show that leaf K concentrations < 0.7-0.8% appeared to limit the leaf net CO2 assimilation rate (A), and that the relative limitation of photosynthesis due to g(m) and stomatal conductance (g(s)) decreased with increasing supplies of K. However, a sensitivity analysis indicated that A was most sensitive to the maximum carboxylation rate of Rubisco (V(c,max)) and the maximum rate of electron transport (J(max)). These results indicate that the photosynthetic rate is primarily limited by the biochemical processes of photosynthesis (V(c,max) and J(max)), rather than by g(m) and g(s) in K-deficient plants. Additionally, g(m) was closely correlated with g(s) and the leaf dry mass per unit area (M(A)) in hickory seedlings, which indicates that decreased g(m) and g(s) may be a consequence of leaf anatomical adaptation.     

Seasonal photosynthetic responses to light and temperature in white spruce (Picea glauca) seedlings planted under an aspen (Populus tremuloides) canopy and in the open

Photosynthetic light and temperature response curves were measured seasonally in seedlings of white spruce (Picea glauca (Moench.) Voss) grown for two years in the understory of aspen (Populus tremuloides Michx.) or in the open in central Alberta. Light-saturated rate of net photosynthesis, the optimum temperature for net photosynthesis, transpiration rate, photochemical efficiency, and stomatal and mesophyll conductances increased from spring to summer and declined thereafter, whereas dark respiration rate and compensation and saturation points were highest in spring. Depression of photosynthetic parameters was greater in open-grown seedlings than in understory seedlings during the periods in spring and autumn when night frosts were common. Net photosynthetic rates were similar in understory and open-grown seedlings in summer, but they were significantly lower in open-grown seedlings in spring and autumn. Significantly lower transpiration rates and stomatal conductances in open-grown seedlings than in understory seedlings were also observed at 15 and 25 degrees C in the autumn. Shoot and needle growth were less in open-grown seedlings than in understory seedlings. In summer, when irradiances were low in the aspen understory, understory white spruce seedlings maintained a positive carbon balance by decreasing their compensation and saturation points and increasing their photochemical efficiency compared to spring and autumn.     

Gas exchange during a soil drying cycle in seedlings of four black walnut (Juglans nigra L.) families

Photosynthetic and stomatal responses to a soil drying cycle were examined in half-sib seedlings of four walnut (Juglans nigra L.) families. Well-watered seedlings of an Iowa seed source had significantly higher rates of net photosynthesis than seedlings from New York or Michigan sources. This superior photosynthetic potential was associated with both greater stomatal conductance and mesophyll capacity for CO(2) fixation. In a drying soil, net photosynthesis and leaf conductance to water vapor of all families declined substantially, even under mild water stress. These responses were more strongly related to soil water status, as estimated by predawn leaf water potential, than to leaf water potential at the time of gas exchange measurement. There were no differences among families in the pattern of gas exchange response to developing water stress; however, families differed in capacity for recovery of gas exchange from water stress following rehydration. Sensitivity of photosynthesis of black walnut seedlings to water stress may be associated with poor growth and survival of this species in xeric habitats.     

多重复干旱循环对北美短叶松和黑云杉苗木气体交换及水分利用效率的影响

本文研究了多重复干旱循环对１年生北美短叶松（ＰｉｎｕｓｂａｎｋｓｉａｎａＬａｍｂ．）和黑云杉（Ｐｉｃｅａｍａｒｉａｎａ［Ｍｉｌ］Ｂ．Ｓ．Ｄ．）苗木的气体交换速率及水分利用效率的影响。结果表面，多重干旱循环对它们的气体交换（Ｃｓ，Ｐｎ，Ｔｒ）有显著影响（Ｐ＜０．５），而对其水分利用效率（ＷＵＥ）影响不大（Ｐ＞０．１）。尽管北美短叶松的气孔对轻度干旱胁迫不如黑云杉敏感，但是它对中度及严重干旱胁迫的敏感程度却高于黑云杉。在轻度及中度干旱胁迫下，北美短叶松的光合作用主要受非气孔因素的影响，而黑云杉则主要受气孔因素的影响。解除干旱胁迫后，黑云杉的气孔敏感性、光合能力及水分利用效率的恢复都要比北美短叶松更快．我们认为，延迟脱水是北美短叶松的主要耐旱机理，而忍耐脱水则是黑云杉重要的耐旱途径。轻度的干旱胁迫锻炼可以帮助北美短叶松在更严重的干旱胁迫下保持固有而较强的耐旱能力。然而，通过多重复干旱循环锻炼后黑云杉在改善耐旱能力的强度方面则大于北美短叶松     

Responses of black spruce (Picea mariana) and tamarack (Larix laricina) to flooding and ethylene

Black spruce (Picea mariana (Mill.) BSP) and tamarack (Larix laricina (Du Roi) K. Koch) are the predominant tree species in the boreal peatlands of Alberta, Canada, where low nutrient availability, low soil temperature and a high water table limit their growth. Effects of flooding for 28 days on morphological and physiological responses were investigated in greenhouse-grown black spruce and tamarack seedlings in a growth chamber. Flooding reduced root hydraulic conductance, net assimilation rate and stomatal conductance, and increased water-use efficiency (WUE) and needle electrolyte leakage in both species. Although flooded black spruce seedlings maintained higher net assimilation rates and stomatal conductance than flooded tamarack seedlings, flooded tamarack seedlings were able to maintain higher root hydraulic conductance than flooded black spruce seedlings. Needles of flooded black spruce developed tip necrosis and electrolyte leakage after 14 days of flooding, and these symptoms were subsequently more prominent than in needles of flooded tamarack seedlings. Flooded tamarack seedlings exhibited no visible injury symptoms and developed hypertrophied lenticels at their stem base. Application of exogenous ethylene resulted in a significant reduction in net assimilation, stomatal conductance and root respiration, whereas root hydraulic conductivity increased in both species. Thus, although flooded black spruce seedlings maintained a higher stomatal conductance and net assimilation rate than tamarack seedlings, black spruce did not cope with the deleterious effects of prolonged soil flooding and exogenous ethylene as well as tamarack.     

土壤干旱胁迫对巨龙竹实生苗光合特性和抗氧化酶活性的影响

试验选取盆栽巨龙竹(Dendrocalamus sinicus)实生苗，通过控制土壤水分含量设置轻度干旱、中度干旱和重度干旱3个胁迫水平研究不同干旱胁迫处理对巨龙竹实生苗的光合参数和抗氧化酶活性的影响。结果表明：随干旱程度的增加，土壤含水量显著下降，巨龙竹实生苗叶片的叶绿素含量(SPAD值)、净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)均呈下降趋势，而胞间CO₂浓度(Ci)在中度和重度干旱时呈上升趋势；在3种干旱强度下，SOD活性呈现先上升后下降再上升的变化趋势，APX活性和丙二醛含量均高于对照，CAT均低于对照。由研究结果可见，干旱会引起巨龙竹实生苗光合参数下降，光合速率下降的原因表现为随着干旱强度的增加由气孔限制原因转为非气孔限制原因，并且SOD和APX是巨龙竹在干旱胁迫下起主要作用的2种酶。巨龙竹实生苗在轻度干旱环境中表现出良好的适应性，但在中度和重度干旱环境中其适应性较差。     

Effects of soil drought stress on photosynthetic gas exchange traits and chlorophyll fluorescence in Forsythia suspensa

To clarify the changes in plant photosynthesis and mechanisms underlying those responses to gradually increasing soil drought stress and reveal quantitative relationships between photosynthesis and soil moisture,soil water conditions were controlled in greenhouse pot experiments using 2-year-old seedlings of Forsythia suspensa(Thunb.) Vahl. Photosynthetic gas exchange and chlorophyll fluorescence variables were measured and analyzed under 13 gradients of soil water content. Net photosynthetic rate(PN), stomatal conductance(gs), and water-use efficiency(WUE) in the seedlings exhibited a clear threshold response to the relative soil water content(RSWC). The highest PNand WUEoccurred at RSWCof51.84 and 64.10%, respectively. Both PNand WUEwere higher than the average levels at 39.79% B RSWCB 73.04%. When RSWCdecreased from 51.84 to 37.52%,PN, gs, and the intercellular CO2 concentration(Ci)markedly decreased with increasing drought stress; the corresponding stomatal limitation(Ls) substantially increased, and nonphotochemical quenching(NPQ) also tended to increase, indicating that within this range of soil water content, excessive excitation energy was dispersed from photosystem II(PSII) in the form of heat, and the reduction in PNwas primarily due to stomatal limitation.While RSWCdecreased below 37.52%, there were significant decreases in the maximal quantum yield of PSII photochemistry(Fv/Fm) and the effective quantum yield of PSII photochemistry(UPSII), photochemical quenching(qP), and NPQ; in contrast, minimal fluorescence yield of the dark-adapted state(F0) increased markedly. Thus,the major limiting factor for the PNreduction changed to a nonstomatal limitation due to PSII damage. Therefore, an RSWCof 37.52% is the maximum allowable water deficit for the normal growth of seedlings of F. suspensa, and a water content lower than this level should be avoided in field soil water management. Water contents should be maintained in the range of 39.79% B RSWCB 73.04% to ensure normal function of the photosynthetic apparatus and high levels of photosynthesis and efficiency in F.suspensa.     

淹水胁迫对南五味子和海南红豆幼苗光合特性的影响

对南五味子和海南红豆的1年生实生苗在不同淹水时间下的叶片的光合特性进行了研究。结果表明:随着淹水时间的持续,南五味子幼苗的净光合速率(Pn)小幅波动后升高,而气孔导度(Gs)、蒸腾速率(Tr)和胞间CO2浓度(Ci)先降后升,至淹水20 d时各指标均高于对照;海南红豆幼苗的Pn、Gs、Ci、Tr先降后升,至淹水20 d没有恢复到对照水平;2种幼苗的气孔限制值(Ls)均为先升高后下降;南五味子和海南红豆都有一定的抗涝性,前者具有更强的耐淹水能力。     

Relationships between gas exchange adaptation of Sitka x interior spruce genotypes and ribosomal DNA markers

Adaptive physiological changes were investigated in seven populations of Sitka (Picea sitchensis (Bong.) Carr.) x interior spruce (P. glauca (Moench) Voss x P. engelmannii Parry ex Engelm.) spanning the Nass-Skeena transition zone in British Columbia, Canada. Each population was represented by an Si rDNA index that was calculated from the relative optical densities on a gel autoradiogram of five ribosomal DNA bands characteristic of Sitka spruce and interior spruce. This index estimates the proportion of the genome contributed by interior spruce. Physiological adaptations were assessed by gas exchange parameters measured under both well-watered and drought conditions. Under well-watered conditions, Sitka spruce populations had higher maximal photosynthesis at saturating light and ambient CO(2), higher quantum yield at the light compensation point, and higher dark respiration than interior spruce populations. Sitka spruce populations also reached maximal photosynthesis at lower photosynthetically active radiation and higher CO(2) concentrations, and had higher stomatal densities that resulted in lower stomatal limitations to photosynthesis than interior spruce populations. In contrast, interior spruce populations exhibited greater drought tolerance than Sitka spruce populations. Their gas exchange rates declined at a slower rate in response to drought. They maintained higher gas exchange rates in response to moderate to severe drought (predawn plant water potentials = -1.5 MPa), and their photosynthetic rates recovered faster when they were rewatered after exposure to drought. Comparison of the seven populations indicated that physiological parameters were significantly related to the Si rDNA index. An increase in Si rDNA index was associated with proportional changes in physiological measurements, suggesting that genetic interchange among species with contrasting ecological adaptations can enhance the environmental adaptation of natural populations.     

Growth,morphology and photosynthetic activity in flooded <Emphasis Type="Italic">Alnus japonica</Emphasis> seedlings

This study was conducted on Alnus japonica seedlings subjected to flooding for 2, 4, and 6 weeks to examine responses in growth, morphology, and photosynthesis to different periods of flooding. Seedlings subjected to flooding for 2 and 4 weeks were drained after flooding then watered daily. Increases in biomass of leaves, roots, and whole plants were less for 6-week-flooded seedlings. Rate of photosynthesis and stomatal conductance of flooded seedlings decreased within 2 weeks. For 2-week-flooded seedlings recovery from reduced stomatal conductance and recovery of photosynthetic activity occurred after drainage. For the 6-week-flooded seedlings stomatal conductance recovered by the end of the experiment. Adventitious root formation by the 4 and 6-week-flooded seedlings was observed from the third week of flooding. These results suggest that recovery of reduced function in leaves may progress with development of adventitious roots during the period of flooding.     

短期干旱和复水对麻栎幼苗光合及叶绿素荧光的影响

麻栎是鲁中山区森林植被恢复的重要阔叶树种之一,本文以1年生麻标幼苗为试验对象,研究短期干旱和复水对其光合及叶绿素荧光等生理生态特征的影响。结果表明：在干旱胁迫初期,干旱组的各项指标均优于对照组,说明过高的土壤含水量并不利于麻栎幼苗的生长;随着土壤干旱程度的进一步加剧,幼苗的气孔导度减少,导致光合和蒸腾作用的下降,水分利用效率升高;胁迫末期各项光合指标均降到最低。从叶绿素荧光参数可以看出,干旱胁迫并未使幼苗的光合机构发生伤害,幼苗在适宜的水分条件时产生光合能力上调,当水分匮缺时,以叶黄素循环为主的非光化学淬灭耗散掉过剩的光能达到保护光合机构的目的。复水后麻栎幼苗的各项生理生态指标恢复正常,但气孔导度的增大存在一定的滞后效应,导致了水分利用效率的反冲,同时叶黄素库仍然维持较高水平,麻栎幼苗通过物理和化学的双重保护来抵御干旱胁迫和进行胁迫后的恢复。     

Quantifying stomatal and non-stomatal limitations to carbon assimilation resulting from leaf aging and drought in mature deciduous tree species

Gas exchange techniques were used to investigate light-saturated carbon assimilation and its stomatal and non-stomatal limitations over two seasons in mature trees of five species in a closed deciduous forest. Stomatal and non-stomatal contributions to decreases in assimilation resulting from leaf age and drought were quantified relative to the maximum rates obtained early in the season at optimal soil water contents. Although carbon assimilation, stomatal conductance and photosynthetic capacity (V(cmax)) decreased with leaf age, decreases in V(cmax) accounted for about 75% of the leaf-age related reduction in light-saturated assimilation rates, with a secondary role for stomatal conductance (around 25%). However, when considered independently from leaf age, the drought response was dominated by stomatal limitations, accounting for about 75% of the total limitation. Some of the analytical difficulties associated with computing limitation partitioning are discussed, including path dependence, patchy stomatal closure and diffusion in the mesophyll. Although these considerations may introduce errors in our estimates, our analysis establishes some reasonable boundaries on relative limitations and shows differences between drought and non-drought years. Estimating seasonal limitations under natural conditions, as shown in this study, provides a useful basis for comparing limitation processes between years and species.     

Physiological changes in red spruce seedlings during a simulated winter thaw

We evaluated net photosynthesis, respiration, leaf conductance, xylem pressure potential (XPP) and cold hardiness in red spruce (Picea rubens Sarg.) seedlings exposed to either a continuous thaw (CT) or a daytime thaw with freezing nights (FN) for 8 days during mid-winter. Physiological differences between CT and FN seedlings were evident for all measured parameters. However, the temporal expression of treatment differences varied among parameters. When compared to FN seedlings, CT seedlings had higher rates of respiration following 24 h of treatment, and a higher net photosynthetic rate, leaf conductance and XPP after 48 h of treatment. The CT seedlings were significantly less cold tolerant than the FN seedlings following 4 days of thaw, whereas FN seedlings did not deharden over the 8 days of treatment. Examination of temporal trends among thaw-associated changes in physiology suggested that, although greater carbon exchange occurred as stomatal conductance increased, the transition from negative to positive net photosynthesis was not the result of increases in conductance, but may have been associated with thaw-induced increases in XPP. Because thaw-associated changes in gas exchange and cold hardiness were offset in time, we conclude that, if changes in these processes are physiologically linked, the linkage is indirect.     

Diurnal Course of Gas Exchange and Water Use Efficiency of Platycladus orientalis in the Semiarid Region of the Loess Plateau

The daily gas exchange, stomatal conductance, and water use efficiency of Platycladus orientalis in the test field of Fangshan county of Shanxi Province in the semiarid region of the Loess Plateau were measured with portable Li-6200 gas analysis system in natural conditions. The results showed that the diurnal course of net photosynthetic rate displayed a two-peak pattern, that of stomatal conductance displayed a hollow pattern, and that of transpiration rate displayed a signal-peak pattern. Water use efficiency culminated in the early morning. On the basis of two criteria of stomatal limitation of photosynthesis suggested by Farquhar and Sharkey, the predominant limiting factor of photosynthesis was the stomatal conductance of stomatal limitation in the morning (10:00–12:00). However, the midday depression of photosynthesis at noon (12:00–14:00) and the decrease in photosynthesis in the afternoon (16:00–18:00) were the results of nonstomatal limitation, such as low carboxylation capacity of the leaf mesophyll.     

Drought resistance of Ailanthus altissima: root hydraulics and water relations

Drought resistance of Ailanthus altissima (Mill.) Swingle is a major factor underlying the impressively wide expansion of this species in Europe and North America. We studied the specific mechanism used by A. altissima to withstand drought by subjecting potted seedlings to four irrigation regimes. At the end of the 13-week treatment period, soil water potential was -0.05 MPa for well-watered control seedlings (W) and -0.4, -0.8 and -1.7 MPa for drought-stressed seedlings (S) in irrigation regimes S1, S2 and S3, respectively. Root and shoot biomass production did not differ significantly among the four groups. A progressively marked stomatal closure was observed in drought-stressed seedlings, leading to homeostasis of leaf water potential, which was maintained well above the turgor loss point. Root and shoot hydraulics were measured with a high-pressure flow meter. When scaled by leaf surface area, shoot hydraulic conductance did not differ among the treated seedlings, whereas root hydraulic conductance decreased by about 20% in S1 and S2 seedlings and by about 70% in S3 seedlings, with respect to the well-watered control value. Similar differences were observed when root hydraulic conductance was scaled by root surface area, suggesting that roots had become less permeable to water. Anatomical observations of root cross sections revealed that S3 seedlings had shrunken cortical cells and a multilayer endodermal-like tissue that probably impaired soil-to-root stele water transport. We conclude that A. altissima seedlings are able to withstand drought by employing a highly effective water-saving mechanism that involves reduced water loss by leaves and reduced root hydraulic conductance. This water-saving mechanism helps explain how A. altissima successfully competes with native vegetation.     

Acidic mist reduces foliar membrane-associated calcium and impairs stomatal responsiveness in red spruce

Acidic deposition can leach essential pools of calcium (Ca) directly from plant foliage. Because of the central role of Ca in environmental signal transduction, disruptions of labile foliar Ca pools could impair physiological responses to a variety of environmental stimuli and stressors. We investigated the possibility that acidic mist-induced depletion of membrane-associated Ca (mCa), which is one form of labile Ca, may alter stomatal responsiveness to water stress, a process known to include Ca in signal transduction cascades. Red spruce (Picea rubens Sarg.) seedlings were exposed to either pH 3.0 or pH 5.0 mist treatments for one growing season. Foliar nutrition was assessed following treatments, and declines in stomatal conductance and net photosynthesis were measured on current-year shoots following stem excision. Seedlings exposed to pH 3.0 acidic mist treatments had reduced mCa relative to the pH 5.0 treated seedlings. Seedlings subjected to the pH 3.0 acidic mist treatment exhibited impaired stomatal functions, including a smaller maximum aperture, slower closure and an increased lag time between stomatal closure and photosynthetic decline following experimental water stress. Delayed stomatal closure could undermine desiccation avoidance mechanisms. Previous work has demonstrated that acidic mist treatments deplete mCa in red spruce and impair cold tolerance, with similar effects in other species. The results we present provide further evidence that acidic mist-induced mCa depletion may cause disruption of a broad range of plant stress responses.     

Effects of foliar nitrogen concentration on photosynthesis and water use efficiency in Douglas-fir

Leaf-level physiological processes were studied in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to determine whether apparent increases in stand-level water use efficiency (WUE) observed in response to nitrogen (N) fertilization were attributable to foliar N effects on carbon fixation rates or on stomatal control of water loss. Photosynthesis and transpiration were measured at different light intensities and ambient CO(2) molar fractions and comparisons were made between current-year shoots with average foliar N concentrations of 1.58% (High-N) and 1.25% (Low-N). Photosynthetic rates and foliar N concentrations were positively correlated. In response to light, photosynthesis and stomatal conductance were closely coupled and a similar coupling was observed in response to different ambient CO(2) concentrations. Partitioning the photosynthetic responses into mesophyll and stomatal components indicated that foliar N altered mesophyll conductance but not stomatal control of water loss. High-N shoots had significantly greater rates of photosynthesis and transpiration than Low-N shoots and, as a result, instantaneous WUE did not differ significantly between High-N and Low-N shoots.     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

Age-related trends in red spruce foliar plasticity in relation to declining productivity

Phenotypic plasticity in needle morphology with increasing tree size and age was investigated by comparing four age classes of red spruce (Picea rubens Sarg.) ranging from juvenile (3-12 years old) to mature (over 100 years old). With increase in tree age there were significant increases in leaf mass per unit area (LMA), mesophyll and vascular bundle area as a percentage of total needle cross-sectional area, and stomatal density. Within the vascular bundle, both xylem cross-sectional area and tracheid lumen area increased significantly, whereas air space as a percentage of total cross-sectional area decreased. These morphological changes were associated with a significant decrease in photosynthetic capacity and stomatal conductance, and an increase in (13)C enrichment. Although both photosynthetic capacity and whole-tree conductance decreased significantly between age classes 3 and 12 years, they did not differ between age classes 53 and 127 years, even though needle (13)C enrichment was significantly greater in the 127-year age class. Thus there appear to be compensatory mechanisms that maintain photosynthetic capacity as trees increase in size and vascular complexity, which in red spruce and other species, may affect leaf hydraulic conductance. Although increased LMA may contribute to reduced photosynthetic capacity in red spruce, similar relationships are not seen in other conifers.