Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

落叶阔叶林内的光合作用和气孔传导率特征(英文)

本文根据Wang和BMdocchi(1989)最近提出的冠层辐射模型,进一步给出了一个模拟冠层光合作用速率和气孔传导率的模式.模式将冠层中每一层的叶面积分为向光叶、半影叶、和全遮荫叶三种,并分别计算其光合作用速率和气孔传导率。计算得到的光合速率廓线表明,在落叶阔叶林内,冠层下部的叶片常处于光照不足状态;半影效应使得透过林冠达于底部的辐射量增大,这对于林下植物的光合作用是有利的。 模式计算值与实测值之间的微弱差别应归因于纯辐射模型无法考虑湍流输送机制造成的CO_2传输和冠层底部耐荫性叶对于低光照的适应能力。     

Photosynthetic induction state of leaves in a soybean canopy in relation to light regulation of ribulose-1-5-bisphosphate carboxylase and stomatal conductance

Photosynthetic induction state, stomatal conductance and light regulation of ribulose-1,5-bisphosphate carboxylase (rubisco) were examined for leaves in a mature, closed soybean (Glycine max) canopy (leaf area index approximately 5) with the objective to determine the extent to which these factors may be limiting the capacity to respond to light transients during sunflecks. When sampled along a vertical gradient, leaves near the bottom of the canopy had lower rubisco contents and chlorophyll a/b ratios as compared with upper leaves. Leaves sampled at midcanopy showed a wide variation in photosynthetic induction state (ratio of the photosynthetic rate achieved after 1 minute exposure to high light to the steady-state assimilation rate achieved after 20 minutes exposure). Both photosynthetic induction state and the initial rubisco activity varied in parallel with stomatal conductance. By contrast there was no correlation between total rubisco activity and stomatal conductance. The results indicate that induction state, as determined by the light regulation of both rubisco activity and stomatal conductance, is an important limitation to the ability of leaves in a soybean canopy to respond to light transients that occur during sunflecks.     

Time constant for water transport in loblolly pine trees estimated from time series of evaporative demand and stem sapflow

 The use of stem sap flow data to estimate diurnal whole-tree transpiration and canopy stomatal conductance depends critically upon knowledge of the time lag between transpiration and water flux through the stem. In this study, the time constant for water movement in stems of 12-year-old Pinus taeda L. individuals was estimated from analysis of time series data of stem water flux and canopy transpiration computed from mean daytime canopy conductance, and diurnal vapor pressure deficit and solar radiation measurements. Water uptake through stems was measured using a constant-heat sapflow probe. Canopy transpiration was correlated to stem uptake using a resistance-capacitance equation that incorporates a time constant parameter. A least-squares auto-regression determined the parameters of the resistance-capacitance equation. The time constants for ten loblolly pine trees averaged 48.0 (SE = 2.0) min and the time lag for the diurnal frequency averaged 47.0 (SE = 2.0) min. A direct-cross correlation analysis between canopy transpiration and sap flow time series showed maximum correlation at an approximately 30 min lag. Residuals (model-predicted minus actual stem flow data) increased with increasing soil moisture depletion. While the time constants did not vary significantly within the range of tree sizes studied, hydraulic resistance and capacitance terms were individually dependent on stem cross-sectional area: capacitance increased and resistance decreased with stem volume. This result may indicate an inverse adjustment of resistance and capacitance to maintain a similar time constant over the range of tree sizes studied.     

麦田冠层气孔导度的分层研究

小麦灌浆期和乳熟期冠层各层叶片上、下表面的气孔导度之间呈正相关关系;冠层不同层的叶片气孔导度从早到傍晚有平行变化的趋势,数值上存在较大的差异,一般从冠层上到下递减。经分析,这主要与冠层叶片接受的光强自上而下递减有关,且这时所对应的叶片水势自冠层上到下递增的幅度大。测算结果表明,冠层气孔导度白天亦呈明显的日变化,灌浆期的值大于乳熟期的值。     

Canopy gradients in leaf intercellular CO2 mole fractions revisited: interactions between leaf irradiance and water stress need consideration

Intercellular CO₂ mole fractions (C_i) are lower in the upper canopy relative to the lower canopy leaves. This canopy gradient in C_i has been associated with enhanced rates of carbon assimilation at high light, and concomitant greater draw‐downs in C_i. However, increases in irradiance in the canopy are generally also associated with decreases in leaf water availability. Thus, stress effects on photosynthesis rates (A) and stomatal conductance (G), may provide a further explanation for the observed C_i gradients. To test the hypotheses of the sources of canopy variation in C_i, and quantitatively assess the influence of within‐canopy differences in stomatal regulation on A, the seasonal and diurnal variation in G was studied in relation to seasonal average daily integrated quantum flux density (Q_int) in tall shade‐intolerant Populus tremula L. trees. Daily time‐courses of A were simulated using the photosynthesis model of Farquhar et al. (Planta 149, 78–90, 1980). Stable carbon isotope composition of a leaf carbon fraction with rapid turnover rate was used to estimate canopy gradient in C_i during the simulations. Daily maximum G (G_max) consistently increased with increasing Q_int. However, canopy differences in G_max decreased as soil water availability became limiting during the season. In water‐stressed leaves, there were strong mid‐day decreases in G that were poorly associated with vapour pressure deficits between the leaf and atmosphere, and the magnitude of the mid‐day decreases in G occasionally interacted with long‐term leaf light environment. Simulations indicated that the percentage of carbon lost due to mid‐day stomatal closure was of the order of 5–10%, and seasonal water stress increased this percentage up to 20%. The percentage of carbon lost due to stomatal closure increased with increasing Q_int. Canopy differences in light environment resulted in a gradient of daily average C_i of approximately 20 µmol mol^?1. The canopy variation in seasonal and diurnal reductions in G led to a C_i gradient of approximately 100 µmol mol^?1, and the actual canopy C_i gradient was of the same magnitude according to leaf carbon isotope composition. This study demonstrates that stress effects influence C_i more strongly than within‐canopy light gradients, and also that leaves acclimated to different irradiance and water stress conditions may regulate water use largely independent of foliar photosynthetic potentials.     

桃树冠层蒸腾动态的数学模拟

将气孔导度公式、Penman—Monteith公式和土壤水分限制模型相结合,可以模拟出不同环境因子对植物蒸腾进程的影响。通过对盆栽桃树（Prunus persica var．nectadna Maxim．）数值模拟发现：影响桃树蒸腾速率的主要气象因子是太阳辐射、大气温度和湿度。植物通过气孔导度的改变来响应气象因子的变化,蒸腾的日变化主要是由气象因子的日变化引起的。土壤的水分状况也对气孔导度有显著的影响,进而影响植物的蒸腾大小。通过数值模拟还发现植物的蒸腾量并不总是随叶面积的增大而增大,对于桃树而言叶面积指数为4左右时日蒸腾量达到最大值。通过对气孔导度和蒸腾速率的模拟值和实测值进行检验发现,两者基本吻合,说明利用数学模拟的方法可以求出不同环境条件和不同叶面积桃树冠层的蒸腾速率。     

Adjustments in hydraulic architecture of Pinus palustris maintain similar stomatal conductance in xeric and mesic habitats

We investigated relationships between whole-tree hydraulic architecture and stomatal conductance in Pinus palustris Mill. (longleaf pine) across habitats that differed in soil properties and habitat structure. Trees occupying a xeric habitat (characterized by sandy, well-drained soils, higher nitrogen availability and lower overstory tree density) were shorter in stature and had lower sapwood-to-leaf area ratio (A(S):A(L)) than trees in a mesic habitat. The soil-leaf water potential gradient (psiS - psiL) and leaf-specific hydraulic conductance (kL) were similar between sites, as was tissue-specific hydraulic conductivity (Ks) of roots. Leaf and canopy stomatal conductance (gs and Gs, respectively) were also similar between sites, and they tended to be somewhat higher at the xeric site during morning hours when vapour pressure deficit (D) was low. A hydraulic model incorporating tree height, A(S):A(L) and psiS-psiL accurately described the observed variation in individual tree G(Sref) (G(S) at D = 1 kPa) across sites and indicated that tree height was an important determinant of G(Sref) across sites. This, combined with a 42% higher root-to-leaf area ratio (A(R):A(L)) at the xeric site, suggests that xeric site trees are hydraulically well equipped to realize equal--and sometimes higher potential for conductance compared with trees on mesic sites. However, a slightly more sensitive stomatal closure response to increasing D observed in xeric site trees suggests that this potential for higher conductance may only be reached when D is low and when the capacity of the hydraulic system to supply water to foliage is not greatly challenged.     

水稻冠层光合有效辐射的时空分布特征

以2个不同株型水稻品种为材料,设置高、中、低3个施氮水平,利用SunScan冠层分析仪于灌浆期系统测定了不同施氮水平下不同株型水稻品种植株形态和冠层内光合有效辐射（PAR）的时空分布状况.结果表明：施氮量对水稻株高、穗弯曲度和茎叶夹角有明显影响;群体叶面积的垂直分布呈中部>上部>下部的分布特征,最大分层叶面积指数（LAI）出现在0.60相对高度处.冠层内平均PAR透光率从顶部向下递减,且在冠层上中部递减迅速,下部递减缓慢;平均PAR透光率随施氮量的增加而递减;平均PAR透光率日变化表现为早晚较低,中午较高;平均PAR透光率随向下累积LAI的增加呈指数递减,群体消光系数K的日变化表现为早晚较高,中午较低,灌浆期的K值介于0.35～0.50.水稻冠层内PAR的三维空间分布表现为冠层上中部水平面上PAR透光率高,光斑面积大;下部水平面上PAR透光率低,光斑少;同一冠层高度水平面上的PAR光强呈不均匀分布.株型紧凑的水稻品种,冠层透光率高,透光率日变化大,群体消光系数小.     

Vertical gradients in photosynthetic physiology diverge at the latitudinal range extremes of white spruce

Light availability drives vertical canopy gradients in photosynthetic functioning and carbon (C) balance, yet patterns of variability in these gradients remain unclear. We measured light availability, photosynthetic CO₂ and light response curves, foliar C, nitrogen (N) and pigment concentrations, and the photochemical reflectance index (PRI) on upper and lower canopy needles of white spruce trees (Picea glauca) at the species' northern and southern range extremes. We combined our photosynthetic data with previously published respiratory data to compare and contrast canopy C balance between latitudinal extremes. We found steep canopy gradients in irradiance, photosynthesis and leaf traits at the southern range limit, but a lack of variation across canopy positions at the northern range limit. Thus, unlike many tree species from tropical to mid-latitude forests, high latitude trees may not require vertical gradients of metabolic activity to optimize photosynthetic C gain. Consequently, accounting for self-shading is less critical for predicting gross primary productivity at northern relative to southern latitudes. Northern trees also had a significantly smaller net positive leaf C balance than southern trees suggesting that, regardless of canopy position, low photosynthetic rates coupled with high respiratory costs may ultimately constrain the northern range limit of this widely distributed boreal species.     

Effects of Density and Extinction Coefficient on Size Variability in Plant Populations

The effects of density and extinction coefficient on size variability,as measured by the coefficient of variation of plant weightin even-aged monocultures, were investigated theoretically usinga diffusion model of growth and size distribution and a canopyphotosynthesis model over the range of densities at which self-thinning(size-dependent mortality) does not occur. Size inequality (thecoefficient of variation of plant weight) increases with increasingdensity or leaf area index at each growth stage. Plants witherect leaves are prone to lower size inequality than plantswith horizontal leaves. These results agree well with existingobservations on even-aged plant monocultures and suggest thatcompetition between plants is mainly one-sided (competitionfor light). One sided competition affects size variability througha G(t, x) function (mean growth of plants of size x at timet per unit time). Two-sided competition (including competitionfor nutrients) affects size variability through a D(t, x) function(variance of growth of plants of size x at time t per unit time).In this case, size inequality decreases with increasing density.The importance of studying size variability is emphasized. Helianthus annus L., size variability, size inequality, coefficient of variation, competition, density effect, extinction coefficient, diffusion model, canopy photosynthesis model     

不同光强对吴茱萸生长动态和光合日变化的影响

报道了不同生长光强(透光率分别为15%、30%、50%、100%)对吴茱萸的生长和光合日变化的影响。结果表明:生长环境光强对吴茱萸苗期株高、地径和冠面积的影响较大,100%RI下的株高、地径和冠面积明显小于各遮荫处理,其中30%RI下吴茱萸的株高、地径和冠面积最大,这表明苗期强光不利于吴茱萸的生长。随着光强的减弱,吴茱萸增大冠面积与株高有利于截获更多光能。10月下旬,100%RI下吴茱萸净光合速率(Pn)日变化曲线呈"双峰型",11:30～14:30,吴茱萸的Pn、胞间二氧化碳浓度与气孔导度均下降,表明此时的光合午休现象是由气孔部分关闭造成的。各遮荫处理下,吴茱萸的Pn日变化均呈"单峰型",并无第二峰的出现,可能与遮荫条件下,下午光强较弱,环境温度较高,空气相对湿度较低有关。     

Partitioning controls on Amazon forest photosynthesis between environmental and biotic factors at hourly to interannual timescales

Gross ecosystem productivity (GEP) in tropical forests varies both with the environment and with biotic changes in photosynthetic infrastructure, but our understanding of the relative effects of these factors across timescales is limited. Here, we used a statistical model to partition the variability of seven years of eddy covariance‐derived GEP in a central Amazon evergreen forest into two main causes: variation in environmental drivers (solar radiation, diffuse light fraction, and vapor pressure deficit) that interact with model parameters that govern photosynthesis and biotic variation in canopy photosynthetic light‐use efficiency associated with changes in the parameters themselves. Our fitted model was able to explain most of the variability in GEP at hourly (R² = 0.77) to interannual (R² = 0.80) timescales. At hourly timescales, we found that 75% of observed GEP variability could be attributed to environmental variability. When aggregating GEP to the longer timescales (daily, monthly, and yearly), however, environmental variation explained progressively less GEP variability: At monthly timescales, it explained only 3%, much less than biotic variation in canopy photosynthetic light‐use efficiency, which accounted for 63%. These results challenge modeling approaches that assume GEP is primarily controlled by the environment at both short and long timescales. Our approach distinguishing biotic from environmental variability can help to resolve debates about environmental limitations to tropical forest photosynthesis. For example, we found that biotically regulated canopy photosynthetic light‐use efficiency (associated with leaf phenology) increased with sunlight during dry seasons (consistent with light but not water limitation of canopy development) but that realized GEP was nonetheless lower relative to its potential efficiency during dry than wet seasons (consistent with water limitation of photosynthesis in given assemblages of leaves). This work highlights the importance of accounting for differential regulation of GEP at different timescales and of identifying the underlying feedbacks and adaptive mechanisms.     

长白落叶松冠层光合作用的空间异质性

以2014年黑龙江省帽儿山林场14年生人工长白落叶松为研究对象,对比分析了各项光合指标、环境因子及光合生理参数在冠层内的空间差异性,并探讨了净光合速率(P_n)与其他指标的关系.结果表明: 在树冠垂直方向,上层P_n、气孔导度(g_s)和蒸腾速率(T_r)显著高于中层和下层,胞间CO₂浓度(C_i)表现为下层>中层>上层;光合有效辐射(PAR)从上层外部到下层内部呈显著降低趋势,水汽压差(VPD)和叶片温度(T_l)表现为上层显著高于中层和下层,相对湿度(RH)则无显著差异;最大净光合速率(P_{n max})、暗呼吸速率(R_d)、光补偿点(LCP)和光饱和点(LSP)均表现为上层>中层>下层,下层比上层分别降低32.7%、55.8%、80.2%和51.6%,表观量子效率(AQY)表现为下层>中层>上层,下层分别是中层和上层的1.2和1.3倍.水平方向,光合指标和环境因子的差异性主要体现在树冠上层,P_n、g_s、T_r、PAR和VPD表现为树冠外部显著高于树冠内部,而C_i和RH差异不显著;P_{n max}、R_d、LCP和LSP表现为外部>内部,内部比外部分别降低0.4%、37.7%、42.0%和16.4%,而AQY在内部比外部高0.7%.C_i是限制P_n的主要生理因子,PAR是影响P_n的主要环境因子,尤其在弱光区域PAR对P_n的影响十分明显.因此,在模拟和预估树木冠层光合作用时,考虑空间异质性有一定的必要性.     

Microclimate, Canopy Structure and Photosynthesis in Canopies of Three Contrasting Temperate Forage Grasses: II. Microclimate and Canopy Structure

Profiles of shortwave radiation, net radiation and temperaturewere measured in swards of three grasses of contrasting structureLolium perenne cv. S24, L. perenne cv. Reveille and Festucaarundinacea cv. S170. Measurements were also made of the reflectionof shortwave radiation, leaf water potential and stomatal resistance.Differences in canopy structure influenced the absorption andreflection of radiation by the varieties. The absorption ofnet radiation and its influence on air temperature inside thecanopy was shown to vary with canopy structure. Calculationsshowed that diurnal changes in the reflection and transmissionof light (400–700 nm) would have little effect on canopyphotosynthesis. No clear relationship between leaf extensionrate, temperature and leaf water potential could be established,although decreases in water potential did appear to reduce thepotential response of leaf extension rate to temperature.     

黄土高原油松冠层气孔导度和蒸腾变化特征与模拟

油松是黄土高原重要的造林树种,模拟其冠层气孔导度和蒸腾对区域水量平衡计算和人工林可持续经营具有重要意义。基于2015—2018年TDP(Thermal dissipation probes)方法所测得液流数据,分析了黄土高原地区油松冠层平均气孔导度(g_c)与冠层蒸腾(Tr)的变化特征与影响因素,并采用Penman-Monteith公式和Jarvis型气孔导度模型模拟了其g_c和Tr的变化过程,结果表明：(1)该地区油松g_c和Tr日内变化均呈现单峰型,日均蒸腾耗水量为(1.25±0.57) mm/d,生长季(4—10月)总蒸腾耗水量均值为195.47 mm。(2)g_c的日内变化受太阳辐射(Rad)驱动(偏相关系数为0.65),当Rad高于300 W/m²时,驱动作用减弱;g_c的日内变化受水汽压亏缺(VPD)控制(偏相关系数为-0.41),随VPD的增加而降低;g_c的日际变化受土壤水分限制(偏相关系数为0.46),当根区相对有效含水率(RE...     

Stomatal responses of pearl millet (Pennisetum americanum [L.] Leeke) to leaf water status and environmental factors in the field

Abstract. Factors affecting stomatal conductance (g₁) of pearl millet ( Pennisetum americanum [L.] Leeke), cultivar BJ 104, were examined in the field in India during the dry season.
Diurnal changes in g₁ were evaluated for upper expanded leaves at flowering on two occasions using plants subjected to varying degrees of water stress. Except for the most severely stressed treatment, diurnal changes in g₁ closely matched changes in irradiance ( I ), the promotive effect of which largely overcame opposing influences on g₁ of increasing atmospheric vapour pressure deficit, and decreasing leaf water and turgor potentials (Ψ, Ψ_p).
Two main effects of water stress on g₁ were evident: (i) a decrease in the amplitude of the mid-day peak in g₁, and (ii) a decrease in the time over which high g₁ was maintained, resulting in early (mid-day) closure and hysteresis in the relationship between g₁ and I .
Leaf conductance was greatest for upper leaves and decreased down the canopy. At equivalent depths in the canopy g₁ was higher in flowering than in photoperiodically-retarded plants of the same age. The magnitude of water stress-induced stomatal closure increased down the plant, and was more marked in retarded than in flowering plants.
Within individual stress treatments Ψ of upper leaves decreased linearly as transpiration flux increased. It is concluded that stomatal behaviour of upper leaves of pearl millet at flowering largely operates to maximize assimilation rather than to minimize water loss.     

A Comparison of Leaf Anatomy in Field-grown Gossypium hirsutum and G. barbadense

Gossypium hirsutum L. (upland cotton) and G. barbadense L. (Pimacotton) are two of the most important fibre producing cottonspecies in cultivation. When grown side-by-side in the field,G.hirsutum has higher photosynthetic and transpiration rates (Luet al., 1997. Australian Journal of Plant Physiology24: 693–700).The present study was undertaken to determine if the differencesin physiology can be explained by leaf and canopy morphologyand anatomy. Scanning electron microscopy was used to comparethe leaf anatomy of field-grown upland (‘Delta’and ‘Pine Land 50’) and Pima (‘S6’)cotton. Compared to G. hirsutum, mature leaves of G. barbadenseare larger and thinner, with a thinner palisade layer. G. barbadenseleaves show significant cupping or curling which allows fora more even absorption of insolation over the course of theday and much more light penetration into the canopy. AlthoughG. barbadense leaves have a 70–78% higher stomatal densityon both the abaxial and the adaxial surfaces, its stomates areonly one third the size of those of G. hirsutum. This resultsin G. barbadense having only about 60% of the stomatal surfacearea per leaf surface area compared to G. hirsutum. These resultsare indicative of the anatomical and physiological differencesthat may limit the yield potential of G. barbadense in certaingrowing environments. Copyright 2000 Annals of Botany Company Cotton, leaf anatomy, leaf development, photosynthesis, Gossypium hirsutum, Gossypium barbadense, stomatal density     

Critical Water Potential for Stomatal Closure in Sitka Spruce

Steady state rates of net photosynthesis and stomatal conductance at high water potentials were measured under controlled conditions in a leaf chamber on Sitka spruce [Picea sitchensis (Bong.) Carr.] shoots detached from the forest canopy or on seedlings. The water supply to the seedlings was terminated by excision and the shoot water potential (or critical water potential) and osmotic potential at the onset of stomatal closure measured. The turgor potential was calculated. The initial osmotic potential before insertion of the shoot into the chamber was also measured. Shoot water potential and osmotic potential at stomatal closure, and initial osmotic potential were significantly higher (less negative) in foliage from the lowest level in the canopy compared with foliage in the upper canopy, and higher in shoots of seedlings transferred to low light than in those at high light. Critical water potential also varied with season, being higher in July than in October and November. In all except one instance, turgor potential at the onset of stomatal closure was negative, possibly because of dilution of the cell sap by the extracellular water during the estimate of osmotic potential. Over all the experiments variation in critical water potential was correlated with variation in critical osmotic potential and, to a lesser extent, the initial osmotic potential. However, turgor potential at the critical potential varied from +0.6 to -4.6 bar. This suggests that difference in turgor between the guard cells and subsidiary cells, which controls stomatal aperture, is only loosely coupled with the bulk leaf turgor and hence that bulk leaf turgor is not a good index of the turbor relations of the guard cells.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. I. Leaf model parametrization

In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m^?2 s^?1) than in sun leaves (11–5μmol m^?2 s^?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).