Interspecific variation in nighttime transpiration and stomatal conductance in a mixed New England deciduous forest

Transpiration is generally assumed to be insignificant at night when stomata close in response to the lack of photosynthetically active radiation. However, there is increasing evidence that the stomata of some species remain open at night, which would allow for nighttime transpiration if there were a sufficient environmental driving force. We examined nighttime water use in co-occurring species in a mixed deciduous stand at Harvard Forest, MA, using whole-tree and leaf-level measurements. Diurnal whole-tree water use was monitored continuously with Granier-style sap flux sensors in paper birch (Betula papyrifera Marsh.), red oak (Quercus rubra L.) and red maple (Acer rubrum L.). An analysis was conducted in which nighttime water flux could be partitioned between refilling of internal water stores and transpiration. Substantial nighttime sap flux was observed in all species and much of this flux was attributed to the refilling of depleted water stores. However, in paper birch, nighttime sap flux frequently exceeded recharge estimates. Over 10% of the total daily sap flux during the growing season was due to transpiration at night in paper birch. Nighttime sap flux was over 8% of the total daily flux in red oak and 2% in red maple; however, this flux was mainly associated with recharge. On nights with elevated vapor pressure deficit, sap flux continued through the night in paper birch, whereas it reached zero during the night in red oak and red maple. Measurements of leaf-level gas exchange on a night with elevated vapor pressure deficit showed stomatal conductance dropping by only 25% in paper birch, while approaching zero in red oak and red maple. The study highlighted differences in ecophysiological controls on sap flux exerted by co-occurring species. Paper birch is a fast-growing, shade-intolerant species with an earlier successional status than red oak and red maple. Risking water loss through nighttime transpiration may provide paper birch with an ecological advantage by enabling the species to maximize photosynthesis and support rapid growth. Nighttime transpiration may also be a mechanism for delivering oxygen to respiring cells in the deep sapwood of paper birch.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.     

Environmental controls on sap flow in a northern hardwood forest

Our objective was to gain a detailed understanding of how photosynthetically active radiation (PAR), vapor pressure deficit (D) and soil water interact to control transpiration in the dominant canopy species of a mixed hardwood forest in northern Lower Michigan. An improved understanding of how these environmental factors affect whole-tree water use in unmanaged ecosystems is necessary in assessing the consequences of climate change on the terrestrial water cycle. We used continuously heated sap flow sensors to measure transpiration in mature trees of four species during two successive drought events. The measurements were scaled to the stand level for comparison with eddy covariance estimates of ecosystem water flux (Fw). Photosynthetically active radiation and D together explained 82% of the daytime hourly variation in plot-level transpiration, and low soil water content generally resulted in increased stomatal sensitivity to increasing D. There were also species-specific responses to drought. Quercus rubra L. showed low water use during both dry and wet conditions, and during periods of high D. Among the study species, Acer rubrum L. showed the greatest degree of stomatal closure in response to low soil water availability. Moderate increases in stomatal sensitivity to D during dry periods were observed in Populus grandidentata Michx. and Betula papyrifera Marsh. Sap flow scaled to the plot level and Fw demonstrated similar temporal patterns of water loss suggesting that the mechanisms controlling sap flow of an individual tree also control ecosystem evapotranspiration. However, the absolute magnitude of scaled sap flow estimates was consistently lower than Fw. We conclude that species-specific responses to PAR, D and soil water content are key elements to understanding current and future water fluxes in this ecosystem.     

Linking leaf and tree water use with an individual-tree model

We tested the ability of a model to scale gas exchange from leaf level to whole-tree level by: (1) measuring leaf gas exchange in the canopy of 10 trees in a tall Eucalyptus delegatensis RT Baker forest in NSW, Australia; (2) monitoring sap flow of the same 10 trees during the measurement week; and (3) using an individual-tree-based model (MAESTRA) to link the two sets of measurements. Photosynthesis and stomatal conductance components of the model were parameterized with the leaf gas exchange data, and canopy structure was parameterized with crown heights, dimensions and leaf areas of each of the measurement trees and up to 45 neighboring trees. Transpiration of the measurement trees was predicted by the model and compared with sap flow data. Leaf gas exchange parameters were similar for all 10 trees, with the exception of two smaller trees that had relatively low stomatal conductances. We hypothesize that these trees may have experienced water stress as a result of competition from large neighboring trees. The model performed well, and in most cases, was able to replicate the time course of tree transpiration. Maximum rates of transpiration were higher than measured rates for some trees and lower than measured rates for others, which may have been a result of inaccuracy in estimating tree leaf area. There was a small lag (about 15-30 minutes) between sap flow and modeled transpiration for some trees in the morning, likely associated with use of water stored in stems. The model also captured patterns of variation in sap flow among trees. Overall, the study confirms the ability of models to estimate forest canopy transpiration from leaf-level measurements.     

Long-term species and structural changes after cleaning young even-aged northern hardwoods in New Hampshire,USA

The effects of four cleaning practices (including a control) on species composition and structural characteristics were studied over a 31-year period following treatment of an even-aged 25-year-old northern-hardwood stand that originated after complete clearcutting in 1933–1935. The treatments consisted of: a heavy and a light crop tree cleaning; a drastic species-cleaning treatment that removed nearly all pin cherry (Prunus pensylvanica L.f.), aspen (Populus tremuloides Michx. and grandidentata Michx.), striped maple (Acer pensylvanicum L.), and red maple (Acer rubrum L.) sprout clumps followed by a crop-tree cleaning; and an uncut control. There were no significant differences among treatments in species and structural characteristics in the 56-year-old stand at the end of the study period, except for the presence of a moderate aspen component in the light cleaning and the control. Although previous research shows that cleaning treatments in young northern hardwoods may have silvicultural and economic benefits, the impact of such treatments on long-term stand development is relatively minor.     

Seasonal and interannual variability of canopy transpiration of a hedgerow in southern England

Transpiration from a hawthorn (Crataegus monogyna L.) dominated hedgerow in southern England was measured continuously over two growing seasons by the sap flow technique. Accompanying measurements of structural parameters, microclimate and leaf stomatal and boundary layer conductances were used to establish the driving factors of hedgerow transpiration. Observed transpiration rates, reaching peak values of around 8 mm day(-1) and a seasonal mean of about 3.5 mm day(-1), were higher than those reported for most other temperate deciduous woodlands, except short-rotation coppice and wet woodlands. The high rates were caused by the structural and physiological characteristics of hawthorn leaves, which exhibited much higher stomatal and boundary-layer conductances than those of the second-most abundant woody species in the hedgerow, field maple (Acer campestre L.). Only in the hot summer of 2003 did stomatal conductance, and thus transpiration, decrease substantially. The hedgerow canopy was always closely coupled to the atmosphere. Hedgerow transpiration equaled potential evaporation (calculated by the Priestley-Taylor formula) in 2003 and exceeded it in 2004, which meant that a substantial fraction of the energy (21% in 2003 and more than 37% in 2004) came from advection. Hedgerow canopy conductance (g(c)), as inferred from the sap flow data by inverting the Penman-Monteith equation, responded to solar radiation (R(G)) and vapor pressure deficit (D). Although the response to R(G) showed no systematic temporal variation, the response to D, described as g(c)(D) = g(cref) - mln(D), changed seasonally. The reference g(c) depended on leaf area index and the ratio of -m/g(cref) on long-term mean daytime D. A model is proposed based on these observations that predicts canopy conductance for the hawthorn hedge from standard weather data.     

彩叶植物在北方园林中的适应性评价

随着人类对居住环境要求的不断提高,彩叶植物以其色彩艳丽,观赏期长,营造的园林景观稳定性强,色相变化明显等无可比拟的优越性在园林行业中占有相当大的优势。选取紫叶矮樱、紫叶李、紫叶碧桃、美人梅等九种彩叶植物对其生态适应性和景观效果进行了分析评价。     

Development of canopy stratification during early succession in northern hardwoods

Canopy development on a 6-year-old strip cut was analyzed by measuring the heights to terminal buds and bud scale scars of the tallest individuals of each species present on 50 plots of radius 6 m. Phenology of height growth was monitored during the following growing season. Pin cherry (Prunus pensylvania L.), an intolerant short-lived tree, had the fastest growth rate and was on the average the tallest species from the second to the sixth year of regrowth. Although advance regeneration of sugar maple (Acer saccharum Marsh.) and beech (Fagus grandifoloa Ehrh.) were the tallest trees during the first growing season, their slower growth rate insured that they would not keep up with the pin cherry. Trembling aspen (Populus tremuloides Michx.), striped maple (Acer pensylvanicum L.) and yellow birch (Betula alleghaniensis Britt.) occupied an intermediate position in the canopy by the end of the sixth growing season, and showed relatively greater annual height increment than beech or sugar maple. Height growth phenology differed slightly for each species. Beech, ash (Fraxinus americana L.) and sugar maple commenced growth early, grew rapidly and set buds all by 1 August (beech by 15 June). Yellow birch, pin cherry and trembling aspen started growing as early as the others, grew more slowly at first but then grew for a longer period of time. Striped maple seemed to be somewhat intermediate. Growth phenology and growth rate are related to the tolerance and growth form type (e.g. determinate or indeterminate) of the species. The most tolerant species tend to be determinate in growth form, have slower growth rates and complete height growth earlier. The intolerant species tend to be indeterminate, have a faster growth rate and continue to grow for a longer period. These may be mechanisms by which many species can grow together and avoid adverse effects such as suppression.     

10种彩叶树种光合特性研究

采用Li-6400便携式光合测定仪,对福建沙县引种的银杏、半枫荷、枫香、黄山栾、蓝果树、北美红枫、红叶李、无患子、海棠和碧桃等10种彩叶树种的净光合速率和蒸腾速率进行测定.结果 表明:(1)银杏、蓝果树、红叶李和碧桃净光合速率的日变化呈单峰曲线;而半枫荷、枫香、黄山栾、北美红枫、无患子和海棠则呈双峰曲线,观察到"午休"...     

The Effect of Landfill Leachate Spraying on Foliar Nutrient Concentrations and Leaf Transpiration in a Northern Hardwood Forest, Canada

The effect of municipal solid waste (MSW) leachate spray irrigationon a mature northern hardwood forest was investigated. Canopyfoliar samples and stem increment cores were collected fromtwo indicative species, sugar maple (Acer saccharum Marsh.)and American beech (Fagus grandifolia Ehrh.), within each ofa heavily sprayed, lightly sprayed and control area. Foliarconcentrations of N and P were significantly higher in bothmaple and beech foliage within the sprayed areas when comparedto an unsprayed area (control). Levels of Mg and K were markedlyhigher in maple but not beech foliar samples within the heavilysprayed areas when compared to foliage sampled within the unsprayedcontrol. While no significant trends were observed within themaple foliage, both Fe and B levels increased significantlyin beech foliar samples obtained from within the heavily sprayedarea in comparison to foliage samples from the control. Directporometric measurements of the transpiration rate and diffusive(stomatal) resistance of canopy and understory plant leavesrevealed a significant increase in diffusive resistance anda decline in transpiration rate with leachate spraying. Afterfour years of spraying a significant effect of leachate applicationon radial stem growth of both maple and beech trees has notbeen observed.     

Osmotic potential of several hardwood species as affected by manipulation of throughfall precipitation in an upland oak forest during a dry year

Components of dehydration tolerance, including osmotic potential at full turgor (Psi(pio)) and osmotic adjustment (lowering of Psi(pio)), of several deciduous species were investigated in a mature, upland oak forest in eastern Tennessee. Beginning July 1993, the trees were subjected to one of three throughfall precipitation treatments: ambient, ambient minus 33% (dry treatment), and ambient plus 33% (wet treatment). During the dry 1995 growing season, leaf water potentials of all species declined to between -2.5 and -3.1 MPa in the dry treatment. There was considerable variation in Psi(pio) among species (-1.0 to -2.0 MPa). Based on Psi(pio) values, American beech (Fagus grandifolia Ehrh.), dogwood (Cornus florida L.), and sugar maple (Acer saccharum Marsh.) were least dehydration tolerant, red maple (A. rubrum L.) was intermediate in tolerance, and white oak (Quercus alba L.) and chestnut oak (Quercus prinus L.) were most tolerant. During severe drought, overstory chestnut oak and understory dogwood, red maple and chestnut oak displayed osmotic adjustment (-0.12 to -0.20 MPa) in the dry treatment relative to the wet treatment. (No osmotic adjustment was evident in understory red maple and chestnut oak during the previous wet year.) Osmotic potential at full turgor was generally correlated with leaf water potential, with both declining over the growing season, especially in species that displayed osmotic adjustment. However, osmotic adjustment was not restricted to species considered dehydration tolerant; for example, dogwood typically maintained high Psi(pio) and displayed osmotic adjustment to drought, but had the highest mortality rates of the species studied. Understory saplings tended to have higher Psi(pio) than overstory trees when water availability was high, but Psi(pio) of understory trees declined to values observed for overstory trees during severe drought. We conclude that Psi(pio) varies among deciduous hardwood species and is dependent on canopy position and soil water potential in the rooting zone.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of young plants of four temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid (ABA) concentration ([ABA]) and stomatal aperture were compared in drought-sensitive black walnut (Juglans nigra L.) and black willow (Salix nigra Marsh.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.). Strong correlations among reduction in predawn water potential, increase in xylem sap [ABA] and stomatal closure were observed in all species. Stomatal response was more highly correlated with xylem [ABA] than with ABA flux. Xylem sap pH and ion concentrations appeared not to play a major role in the stomatal response of these species. Stomata were more sensitive to relative changes in [ABA] in drought-sensitive black walnut and black willow than in sugar maple and white oak. In the early stages of drought, increased [ABA] in the xylem sap of black walnut and black willow was probably of root origin and provided a signal to the shoot of the water status of the roots. In sugar maple and white oak, leaf water potential declined with the onset of stomatal closure, so that stomatal closure also may have occurred in response to the change in leaf water potential.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.     

茶条槭山梨山桃苗木对土壤水分胁迫的适应性

在温室内用盆栽法栽植了1年生茶条槭、山梨和山桃苗木,并采用4种土壤水分处理,土壤相对含水量分别为75.0%、61.1%、46.4%和35.4%。结果表明:随土壤含水量降低,3树种苗木净光合速率,蒸腾速率和气孔导度均下降,土壤含水量最低时,茶条槭苗木各生理指标降低幅度最大,山梨和山桃次之。茶条槭和山梨水分利用效率随土壤含水量的减少呈降低趋势,山桃水分利用效率呈升高趋势。水分胁迫下,3树种苗木根分配生物量显著增加。茶条槭在水分胁迫下,根冠比增加幅度最大,山梨次之,山桃最小。此外,茶条槭叶形态可塑性强,随土壤含水量降低,茶条槭单叶叶面积和总叶面积显著减小,叶厚度增加。山梨和山桃在水分胁迫下单叶叶面积及总叶面积无显著变化。综合3树种苗木在水分胁迫下的生理和形态指标及生物量分配的变化,茶条槭对水分胁迫的适应能力强于山桃和山梨。表3参31。     

Structural differences and functional similarities between two sugar maple (Acer saccharum) stands

The spatially inexplicit or functional multilayer models used to predict canopy transpiration or photosynthesis are based on the assumption that closed stands show less functional variability than structural variability, because foliage tends to arrange itself in space to optimize the capture of light. To validate this assumption, we compared the structural and functional properties, and the measured and modeled transpiration fluxes of two sugar maple (Acer saccharum Marsh.) stands of comparable leaf mass but differing in height and diameter distributions. One stand was characterized by a well-developed single-layer canopy, whereas the other stand had a multilayered canopy and a stem diameter distribution of the classical inverse-J shape. Stand differences in height and diameter distribution, and canopy gap fraction, were highly significant. There were minor but significant differences in leaf mass and leaf mass per unit leaf area (LMA) distributions. We found no differences in tree-level relationships between basal area and either transpiration flux or sapwood area. We compared measurements of stand transpiration with transpiration estimates obtained from a multilayer gas exchange model, in which only the nonspatial inputs, leaf area index and LMA frequency distribution described stand structure. For both stands, modeled values of daily transpiration closely followed measured values (r(2) = 0.94). These results support use of the nonspatially explicit approach to estimating canopy gas exchange, especially if the intent is to scale-up to larger portions of the landscape.     

Transpiration and forest structure in relation to soil waterlogging in a Hawaiian montane cloud forest

Transpiration, leaf characteristics and forest structure in Metrosideros polymorpha Gaud. stands growing in East Maui, Hawaii were investigated to assess physiological limitations associated with flooding as a mechanism of reduced canopy leaf area in waterlogged sites. Whole-tree sap flow, stomatal conductance, microclimate, soil oxidation-reduction potential, stand basal area and leaf area index (LAI) were measured on moderately sloped, drained sites with closed canopies (90%) and on level, waterlogged sites with open canopies (50-60%). The LAI was measured with a new technique based on enlarged photographs of individual tree crowns and allometric relationships. Sap flow was scaled to the stand level by multiplying basal area-normalized sap flow by stand basal area. Level sites had lower soil redox potentials, lower mean stand basal area, lower LAI, and a higher degree of soil avoidance by roots than sloped sites. Foliar nutrients and leaf mass per area (LMA) in M. polymorpha were similar between level and sloped sites. Stomatal conductance was similar for M. polymorpha saplings on both sites, but decreased with increasing tree height (r(2) = 0.72; P < 0.001). Stand transpiration estimates ranged from 79 to 89% of potential evapotranspiration (PET) for sloped sites and from 28 to 51% of PET for level sites. Stand transpiration estimates were strongly correlated with LAI (r(2) = 0.96; P < 0.001). Whole-tree transpiration was lower at level sites with waterlogged soils, but was similar or higher for trees on level sites when normalized by leaf area. Trees on level sites had a smaller leaf area per stem diameter than trees on sloped sites, suggesting that soil oxygen deficiency may reduce leaf area. However, transpiration per unit leaf area did not vary substantially, so leaf-level physiological behavior was conserved, regardless of differences in tree leaf area.     

Growth and mortality in underplanted tree seedlings in response to variations in canopy closure of Norway spruce stands

Six broadleaved tree species and Picea abies (L.) Karst. wereplanted under spruce plantations of varying densities, in Sweden.Treatments included control (994 stems ha^–1), dense (538stems ha^–1), sparse (294 stems ha^–1) and gap (0stems ha^–1) overstory treatments. There was an increasein height and diameter growth from control to sparse overstorytreatment of all underplanted tree species except for ash (Fraxinusexcelsior L.) and Norway maple (Acer platanoides L.). Site conditionsmay have hampered the growth of these species, as well as wildcherry (Prunus avium L.). Both oak (Quercus robur L.) and sprucehad greater growth in the gap treatment, relative to the othertreatments. Insecticide application did not influence seedlinggrowth or survival. The performance of beech (Fagus sylvaticaL.), lime (Tilia cordata Mill.), spruce and oak was consistentwith shade tolerance ranking. Beech and lime had a very highsurvival rate, even under the densest canopy. The growth andmortality of ash, maple and wild cherry differed significantlyfrom what was expected. This experiment demonstrated significantvariation in interspecific growth and mortality between sevenplanted tree species in relation to canopy density. Correctsite and species selection is crucial when underplanting inshelterwood systems.     

Modelling the effect of climate on maple syrup production in Québec,Canada

Due to the exceptional sweetness of its sap, sugar maple (Acer saccharum Marsh.) is economically exploited at a commercial scale for maple syrup production in northeastern North America. Approximately 80% of world production is realised in the province of Quebec, Canada, where it is economically important for rural communities. Despite important financial investments in industrial infrastructure over recent decades, the maple syrup yield (ml of sap/tap/year) has followed a general declining trend over the last 15 years, presumably because of unfavourable climatic conditions. In this study, the relationship between climate and maple syrup yield by tap for the whole province was investigated.     

用热扩散式茎流计测定园林树木蒸腾耗水量

采用热扩散式边材液流探针和环境自动监测系统对北京3种园林树木的树干液流及主要环境因子进行了一个生长季的同步观测.结果表明,3树种树干液流的日变化呈明显的单峰曲线,晴天的液流速率大于多云天和阴天,紫叶李和悬铃木的日耗水量明显大于元宝枫;对不同天气3树种每h的液流速率与相应的环境因子进行逐步回归分析.结果显示,影响3树种液流速率的主要环境因子是空气温度、空气相对湿度、辐射强度和5 cm土层温度,在不同天气里起主导作用的因子不同,环境因子与树干液流之间的数量关系能较好地预测树木的蒸腾耗水量.     

Modeling intra-crown and intra-canopy interactions in red maple: assessment of light transfer on carbon dioxide and water vapor exchange

Daily and seasonal net photosynthesis (Anet), transpiration (E), absorbed photosynthetically active radiation (Qa) and light-use efficiency (epsilonc) in a red maple container nursery were simulated with MAESTRA, a three-dimensional canopy model. Effects of canopy heterogeneity were simulated by imposing changes in crown spacing. The light transfer sub-model, a distribution model of incident, direct, diffuse and scattered radiation within MAESTRA, was validated against field measurements of light interception on an intra-crown scale. In the container nursery, we found that a fiber-optic-based method of integrating photosynthetically active radiation (Q) was more suitable for crown-layer light transfer measurements and adjustments than either orthogonal line or individual quantum sensor measurements. The model underestimated intercepted Q by 9.3, 18 and 11.1% for crown layers 1, 2 and 3, respectively; however, there were linear relationships between model estimates and observations made with each of the three measurement methods. We used the validated and parameterized light transfer model to assess intra-crown and intra-canopy light transfer on a layer, crown and canopy basis, and investigated effects of tree size ratio and tree spacing interactions on Anet, E, Qa and epsilonc in the container nursery. Heterogeneous crown and canopy photosynthesis were predicted to exceed values for a uniform canopy under space-limiting conditions. Tree size ratio had large effects on Anet, E, Qa and epsilonc when light to lower-canopy layers was limited by inadequate space between crowns. Increasing Qa at lower-crown layers had the largest impact on whole-crown and whole-canopy Anet, E, Qa and epsilonc. Increases in canopy productivity led to increased water use. Simulations of heterogeneous stands with adequate soil water indicated that light absorption is maximized under space-limiting conditions as a canopy crown moves toward heterogeneity. Nursery and plantation productivity per unit land area was optimized by tactical placement of trees of several sizes, but this was accompanied by increased canopy water use.