A simple method to estimate photosynthetic radiation use efficiency of canopies

BACKGROUND AND AIMS: Photosynthetic radiation use efficiency (PhRUE) over the course of a day has been shown to be constant for leaves throughout a general canopy where nitrogen content (and thus photosynthetic properties) of leaves is distributed in relation to the light gradient. It has been suggested that this daily PhRUE can be calculated simply from the photosynthetic properties of a leaf at the top of the canopy and from the PAR incident on the canopy, which can be obtained from weather-station data. The objective of this study was to investigate whether this simple method allows estimation of PhRUE of different crops and with different daily incident PAR, and also during the growing season. METHODS: The PhRUE calculated with this simple method was compared with that calculated with a more detailed model, for different days in May, June and July in California, on almond (Prunus dulcis) and walnut (Juglans regia) trees. Daily net photosynthesis of 50 individual leaves was calculated as the daylight integral of the instantaneous photosynthesis. The latter was estimated for each leaf from its photosynthetic response to PAR and from the PAR incident on the leaf during the day. KEY RESULTS: Daily photosynthesis of individual leaves of both species was linearly related to the daily PAR incident on the leaves (which implies constant PhRUE throughout the canopy), but the slope (i.e. the PhRUE) differed between the species, over the growing season due to changes in photosynthetic properties of the leaves, and with differences in daily incident PAR. When PhRUE was estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident radiation above the canopy, obtained from weather-station data, the values were within 5 % of those calculated with the more detailed model, except in five out of 34 cases. CONCLUSIONS: The simple method of estimating PhRUE is valuable as it simplifies calculation of canopy photosynthesis to a multiplication between the PAR intercepted by the canopy, which can be obtained with remote sensing, and the PhRUE calculated from incident PAR, obtained from standard weather-station data, and from the photosynthetic properties of leaves at the top of the canopy. The latter properties are the sole crop parameters needed. While being simple, this method describes the differences in PhRUE related to crop, season, nutrient status and daily incident PAR.     

Effects of kaolin application on light absorption and distribution, radiation use efficiency and photosynthesis of almond and walnut canopies

BACKGROUND AND AIMS: Kaolin applied as a suspension to plant canopies forms a film on leaves that increases reflection and reduces absorption of light. Photosynthesis of individual leaves is decreased while the photosynthesis of the whole canopy remains unaffected or even increases. This may result from a better distribution of light within the canopy following kaolin application, but this explanation has not been tested. The objective of this work was to study the effects of kaolin application on light distribution and absorption within tree canopies and, ultimately, on canopy photosynthesis and radiation use efficiency. METHODS: Photosynthetically active radiation (PAR) incident on individual leaves within the canopy of almond (Prunus dulcis) and walnut (Juglans regia) trees was measured before and after kaolin application in order to study PAR distribution within the canopy. The PAR incident on, and reflected and transmitted by, the canopy was measured on the same day for kaolin-sprayed and control trees in order to calculate canopy PAR absorption. These data were then used to model canopy photosynthesis and radiation use efficiency by a simple method proposed in previous work, based on the photosynthetic response to incident PAR of a top-canopy leaf. KEY RESULTS: Kaolin increased incident PAR on surfaces of inner-canopy leaves, although there was an estimated 20 % loss in PAR reaching the photosynthetic apparatus, due to increased reflection. Assuming a 20 % loss of PAR, modelled photosynthesis and photosynthetic radiation use efficiency (PRUE) of kaolin-coated leaves decreased by only 6.3 %. This was due to (1) more beneficial PAR distribution within the kaolin-sprayed canopy, and (2) with decreasing PAR, leaf photosynthesis decreases less than proportionally, due to the curvature of the photosynthesis response-curve to PAR. The relatively small loss in canopy PRUE (per unit of incident PAR), coupled with the increased incident PAR on the leaf surface on inner-canopy leaves, resulted in an estimated increase in modelled photosynthesis of the canopy (+9 % in both walnut and almond). The small loss in PRUE (per unit of incident PAR) resulted in an increase in radiation use efficiency per unit of absorbed PAR, which more than compensated for the minor (7 %) reduction in canopy PAR absorption. CONCLUSIONS: The results explain the apparently contradictory findings in the literature of positive or no effects of kaolin applications on canopy photosynthesis and yield, despite the decrease in photosynthesis by individual leaves when measured at the same PAR.     

Leaf area index, canopy structure and photosynthesis of red clover (Trifolium pratense L.)

Abstract. The influence of leaf age, total leaf area and its dispersion in space on canopy photosynthesis were studied using microswards of red clover ( Trifolium pratense L.) which were established in the greenhouse. Two varieties, Renova (flowering) and Molstad (non-flowering), were sown in separate plastic boxes at densities of 225, 400 and 625 plants per m².
Vertical distribution of photosynthetically active radiation (PAR), leaf area, leaf age and ¹⁴CO₂-fixation were determined periodically. Net photosynthesis and dark respiration of canopies were measured. Maximum photosynthetic capacity of individual leaves was measured on plants taken from the intact canopy or from plants where shading of the growing leaves had been prevented.
Net photosynthetic rate of canopies increased linearly with leaf area index (LAI) up to an LAI of 3.5 and then declined at higher LAI, independent of variety and sowing density. Below the optimum LAI, net photosynthesis depended mainly on interception of PAR. Decrease in canopy photosynthesis above the optimum LAI was due to a higher proportion of old leaves with decreased photosynthetic capacity, and not to an increase in respiring plant parts. It is concluded that LAI and position of leaf age categories in the canopy are more important than vertical distribution of leaf area in determining canopy photosynthesis of red clover.     

Canopy Photosynthesis and Crop Growth Rate of Eight Temperate Forage Grasses

The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q₁₀ = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.     

杉木人工林单叶至冠层光合作用的扩展与模拟研究

根据野外条件下杉木（Ｃｕｎｎｉｎｇｈａｍｉａ ｌａｎｃｅｏｌａｔａ（Ｌａｍｂ．）Ｈｏｏｋ．）针叶光合作用的测定结果。考虑到光合作用对光的非线性响应特性,及其与叶所处的实际冠层环境变量和冠层的空气动力学特性的相关,以筒化的林冠辐射传输模型为基础,结合不同部位和年龄针叶的光响应曲线,实现了叶室测量结果向冠层环境的调整,并进行了冠层光合作用模拟的初步研究。经过调整后的冠层光合作用平均比未经过调整的值高１     

刺槐树冠光合作用的空间异质性

林木冠层是森林与外界环境相互作用最直接的部分,冠层光合作用是研究森林生产力的基础。为了深入了解冠层内部光合作用的差异性,以陕西省永寿县马莲滩流域阳坡和阴坡立地的刺槐林为研究对象,对比分析了光合速率(An)、蒸腾速率(E)、水分利用效率(WUE)、气孔导度(gs)、羧化效率(Vc)、水汽压亏缺(VPD)、气孔限制值(ls)、光合有效辐射(PAR)、空气温度(Ta)在树冠不同层次、不同方位,以及不同坡向之间的差异性。结果表明,刺槐树冠不同层次的光合作用差异性显著,大部分光合生理生态指标表现为:上中下。对于阳坡刺槐,VPD、Ta、gs、E是影响不同层次An的主要因子;对于阴坡刺槐,VPD、E、PAR是影响不同层次An的主要因子。光合作用在刺槐树冠的不同方位没有显著差异,大多数光合指标变化很小,E、ls、PAR、Ta是影响不同方位An的主要因子。对于刺槐冠层内部的任何层次或方位,阴坡刺槐具有更高的日均An、E、Vc、VPD、ls,而阳坡刺槐具有更高的日均WUE、gs、PAR、Ta。阳坡刺槐树冠中层西方和阴坡中层东方的日总光合速率值,可以分别代表阳坡和阴坡刺槐整个冠层的日总光合速率。研究认为,在冠层水平模拟和估计森林生产力时,必须考虑冠层光合作用的空间异质性,对于从单木到林分的尺度推演和模型拟合具有重要的意义。     

不同辐射条件下苹果叶片净光合速率模拟

以富士苹果(Malus domestica Borkh.cv.‘Fuji’)为试材,将C3植物光合生化模型、气孔导度半机理模型、叶片最大光合速率和相对光合有效辐射(RPAR)之间的经验公式相耦合,能够模拟出不同RPAR(或树冠不同部位)下叶片净光合速率(Pn)对小气候因子和叶水势(Ψl)的响应,及Pn日变化。模拟表明,不同RPAR下Pn变化主要依赖于光合有效辐射(PAR)大小,并对CO2浓度有很高敏感性。不同RPAR下叶片Pn最适温度约为20—30℃,并随PAR或CO2浓度的升高而升高。相对湿度(RH)和Ψl对不同RPAR下叶片Pn影响不大,Pn只随RH和Ψl的减小而略有降低。数值模拟表明,当RPAR减小时Pn随之迅速减小,从冠层3 m到1 m处,叶片RPAR从57.18%减少到16.22%,而最大Pn从16.65μmol.m-.2s-1减小到4.24μmol.m-.2s-1。在平均气象条件下,树冠顶部单位面积叶片每天固定CO2为420 mmol.m-.2d-1,而下层叶片只有40 mmol.m-.2d-1。当苹果树冠内叶片接受RPAR低于12%时,全天净光合总量为0,这类叶片可称为无效叶,其所在树冠空间为无效光区。果树整形修剪的主要目的就是尽量减少无效枝叶,利用该模型可确定出这类枝叶在树冠中的位置。     

Heterogeneous reflected light influences asymmetry in leaf anatomy and gas exchange

Soil and vegetative groundcovers reflect light heterogeneously in habitats lacking a continuous overhead canopy, however the effects of reflected light on vegetation in these habitats has received little attention. We test the hypothesis that reflected light flux affects leaf optical properties, anatomy, and photosynthesis of Ipomoea pes-caprae (Convolvulacae), a common sand dune vine with functionally symmetric leaves, by comparing leaves growing over patches of high and low reflected light flux. Patches of high reflected light were found directly over sand and reflected 26.0 ± 0.9% (mean ± 1 SE) of incident photosynthetically active radiation (PAR) while patches of low reflected light were found over vegetation and reflected 6.1 ± 0.7% of incident PAR. Using a novel in situ method to simultaneously illuminate and measure gas exchange of one leaf surface at a time, we show that abaxial surface photosynthetic maxima and palisade parenchyma in sand patches were, respectively, 2.6 times greater and 20% thicker than those found over vegetation patches. Our results suggest that reflected light strongly influences leaf anatomy and gas exchange of I. pes-caprae, demonstrating that reflected light can be an important component of the light environment for vegetation of habitats characterized by high-albedo substrates.     

Effect of light interception by non-photosynthetic organs on canopy photosynthetic production

A canopy photosynthesis model was derived on the assumption that the light diminution within a canopy is caused by both leaves and non-photosynthetic organs. The light diminution by leaves and that by non-photosynthetic organs were taken into account separately in the Lambert-Beer equation of light extinction. The light flux density on the leaf surface at each depth was evaluated from the leaf's share of light. The light flux density on the leaf surface thus obtained was incorporated into the Monsi-Saeki model of canopy photosynthesis. The proposed model was applied for estimating gross canopy photosynthesis in a 19-year-oldLarix leptolepis plantation where 38% of the light diminution was due to non-photosynthetic organs. The daily canopy photosynthesis on one summer day calculated using the present model was about 22% less than that calculated by the conventional Monsi-Saeki model, in which light interception by non-photosynthetic organs is neglected. The degree of such reduction in canopy photosynthesis through shading by non-photosynthetic organs was assessed in relation to parameters affecting light extinction, leaf photosynthetic characteristics, and light regime above the canopy.     

叶龄及树冠不同部位光强对黄花梨光合速率的影响

笔者以黄花梨为试材,研究了叶龄及树冠不同部位光强对其光合速率的影响。结果表明:黄花梨叶片在展叶后约11天即有少量光合产物输出,25天时,单叶净光合速率接近最大值;密植梨树高光合叶幕厚度约125cm左右。     

Model simulations of spatial distributions and daily totals of photosynthesis in Eucalyptus grandis canopies

Summary A simulation model for radiation absorption and photosynthesis was used to test the hypothesis that observed nonuniform distributions of nitrogen concentrations in young Eucalyptus grandis trees result in greater amounts of daily assimilation than in hypothetical trees with uniform N distributions. Simulations were performed for trees aged 6, 9, 12 and 16 months which had been grown in plantations under a factorial combination of two levels of fertilization and irrigation. Observed leaf N distribution patterns yielded daily assimilation rates which were only marginally greater (<5%) than for hypothetical trees with uniform distributions. Patterns of assimilation distribution in individual tree crowns closely resembled those for absorbed radiation, rather than for N. These conclusions were unaffected by three choices of alternative leaf area density distributions. The simulation model was also used to calculate hourly and daily rates of canopy assimilation to investigate the relative importance of radiation absorption and total canopy nitrogen on assimilation. Simulated hourly rates of carbon assimilation were often lightsaturated, whereas daily carbon gain was directly proportional to radiation absorbed by the tree crown and to total mass of N in the leaves. Leaf nitrogen concentrations determined photosynthetic capacity, whereas total leaf area determined the amount of radiation absorbed and thus the degree to which capacity was realized. Observed total leaf area and total crown N were closely correlated. The model predicted that nitrogen use efficiences (NUE, mol CO₂ mol^–1 N) were 60% higher for unfertilized than for fertilized trees at low levels of absorbed photosynthetically active radiation (PAR). Nitrogen use efficiency was dependent on fertilizer treatment and on the amount of absorbed PAR; NUE declined with increasing absorbed PAR, but decreased more rapidly for unfertilized than for fertilized trees. Annual primary productivity was linearly related to both radiation absorbed and to mass of N in the canopy.     

The diurnal time course of net photosynthesis of soybean leaves: Analysis with a physiologically based steady-state photosynthesis model

Summary A physiologically based steady-state model of whole leaf photosynthesis (WHOLEPHOT) is used to analyze observed net photosynthesis daily time courses of soybean, Glycine max (L.) Merr., leaves. Observations during two time periods of the 1978 growing season are analyzed and compared. After adjustment of the model for soybean, net photosynthesis rates are calculated with the model in response to measured incident light intensity, leaf temperature, air carbon dioxide concentration, and leaf diffusion resistance. The steady-state calculations closely approximate observed net photosynthesis. Results of the comparison reveal a decrease in photosynthetic capacity in leaves sampled during the second time period, which is associated with decreasing ability of leaves to respond to light intensity and internal air space carbon dioxide concentration, increasing mesophyll resistance, and increasing stomatal resistance.     

Canopy photosynthesis of six major arable crops is enhanced under diffuse light due to canopy architecture

Diffuse radiation generally increases photosynthetic rates if total radiation is kept constant. Different hypotheses have been proposed to explain this enhancement of photosynthesis, but conclusive results over a wide range of diffuse conditions or about the effect of canopy architecture are lacking. Here, we show the response of canopy photosynthesis to different fractions of diffuse light conditions for five major arable crops (pea, potato, wheat, barley, rapeseed) and cover crops characterized by different canopy architecture. We used 13 years of flux and microclimate measurements over a field with a typical 4 year crop rotation scheme in Switzerland. We investigated the effect of diffuse light on photosynthesis over a gradient of diffuse light fractions ranging from 100% diffuse (overcast sky) to 11% diffuse light (clear‐sky conditions). Gross primary productivity (GPP) increased with diffuse fraction and thus was greater under diffuse than direct light conditions if the absolute photon flux density per unit surface area was kept constant. Mean leaf tilt angle (MTA) and canopy height were found to be the best predictors of the diffuse versus direct radiation effect on photosynthesis. Climatic factors, such as the drought index and growing degree days (GDD), had a significant influence on initial quantum yield under direct but not diffuse light conditions, which depended primarily on MTA. The maximum photosynthetic rate at 2,000 µmol m^?2 s^?1 photosynthetically active radiation under direct conditions strongly depended on GDD, MTA, leaf area index (LAI) and the interaction between MTA and LAI, while under diffuse conditions, this parameter depended mostly on MTA and only to a minor extent on canopy height and their interaction. The strongest photosynthesis enhancement under diffuse light was found for wheat, barley and rapeseed, whereas the lowest was for pea. Thus, we suggest that measuring canopy architecture and diffuse radiation will greatly improve GPP estimates of global cropping systems.     

美国黑核桃实生苗生态生理过程对环境因素响应的数值模拟(Ⅲ):植株冠层光合作用数理模型

以1-2年生北加州黑核桃为试材,建立了具有较高分辨能力的植株群体结构、光分布模型和冠层光合作用模型．将植株冠层按叶面积指数划分为若干层次。上下层之间水平面上太阳辐照度按Monsi＆Saeki所提出的指数递减规律分布．冠层内太阳散射光的消光系数由冠层结构决定,而直射光的消光系数则决定于冠层结构与太阳在天空的位置．在同一层次。将叶片的叶倾角划分为6个等级。将叶片的水平位置划分为8个方位．设同一层次中水平面上的太阳辐照度相同。某一方位角和叶倾角的叶面的直接辐射由太阳视运动方程决定．以此为基础,分别计算“光斑区”和“遮荫区”内叶片的光合速率,并通过数值积分计算整个冠层的光合速率及光合日总量．用田间实测资料验证了冠层内太阳辐射分布模型和冠层光合作用模型．敏感性试验分析表明。模型对环境因子和生物学因素有良好的响应．     

The Contribution of Leaves from Different Levels within a Tomato Crop to Canopy Net Photosynthesis: An Experimental Examination of Two Canopy Models

The rates of net photosynthesis per unit ground area by a closedcanopy of tomato plants were measured over a range of naturallight flux densities. The canopy, of leaf area index 8.6, wasdivided into three horizontal layers of equal depth. On successivedays the canopy was progressively defoliated in layers fromthe ground upwards, allowing the photosynthetic contributionfrom individual leaf layers to be determined. The uppermostlayer, 23% of the total leaf area, assimilated 66% of the netCO₂ fixed by the canopy and accounted for a similar percentageof the total leaf respiration. Net photosynthesis versus light response curves for individualleaves from different positions within the canopy were alsoobtained. Leaf conductances to CO₂ transfer and the dark respirationrates of leaves from the uppermost leaf layer were approximatelyten times those from the lowest layer. The canopy data were analysed using a simple model which assumedthat the canopy was composed of leaves with identical photosyntheticand respiratory characteristics. The model fitted the data andallowed the characteristics of an ‘idealized’ leafto be estimated. The estimated values of the leaf light utilizationefficiency, ,and the leaf conductance CO₂ transfer, , were similarto values directly determined for individual leaves in the uppermostleaf layer and the estimated rate of leaf dark respiration,R_d, corresponded to measured rates for leaves much lower inthe canopy. The simple model may be used to examine gross effectsof crop environment on the leaf photosynthetic characteristicof an ‘idealized’ leaf, but cannot be used to predictaccurately canopy net photosynthesis from the photosyntheticand respiratory characteristics of any single real leaf. A moredetailed model, developed to allow explicitly for the observedvariation in and R_d within the canopy is appropriate for thispurpose.     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

Instantaneous canopy photosynthesis: analytical expressions for sun and shade leaves based on exponential light decay down the canopy and an acclimated non-rectangular hyperbola for leaf photosynthesis

Analytical expressions for the contributions of sun and shade leaves to instantaneous canopy photosynthesis are derived. The analysis is based on four assumptions. First, that the canopy is closed in the sense that it is horizontally uniform. Secondly, that there is an exponential profile of light down the canopy with the same decay constant for light from different parts of the sky. Thirdly, that the leaf photosynthetic response to incident irradiance can be described by a three-parameter non-rectangular hyperbola (NRH). And lastly, that light acclimation at the leaf level occurs in only one parameter of the NRH, that describing the light-saturated photosynthetic rate, which is assumed to be proportional to the local averaged leaf irradiance. These assumptions have been extensively researched empirically and theoretically and their limitations are quite well understood. They have been widely used when appropriate. Combining these four assumptions permits the derivation of algebraic expressions for instantaneous canopy photosynthesis which are computationally efficient because they avoid the necessity for numerical integration down the canopy. These are valuable for modelling plant and crop ecosystems, for which canopy photosynthesis is the primary driver. Ignoring the sun/shade dichotomy can result in overestimates of canopy photosynthesis of up to 20 %, but using a rectangular hyperbola instead of a non-rectangular hyperbola to estimate canopy photosynthesis taking account of sun and shade leaves can lead to a similarly sized underestimate.     

Coping with branch excision when measuring leaf net photosynthetic rates in a lowland tropical forest

Measuring leaf gas exchange from canopy leaves is fundamental for our understanding of photosynthesis and for a realistic representation of carbon uptake in vegetation models. Since canopy leaves are often difficult to reach, especially in tropical forests with emergent trees up to 60 m at remote places, canopy access techniques such as canopy cranes or towers have facilitated photosynthetic measurements. These structures are expensive and therefore not very common. As an alternative, branches are often cut to enable leaf gas exchange measurements. The effect of branch excision on leaf gas exchange rates should be minimized and quantified to evaluate possible bias. We compared light-saturated leaf net photosynthetic rates measured on excised and intact branches. We selected branches positioned at three canopy positions, estimated relative to the top of the canopy: upper sunlit foliage, middle canopy foliage, and lower canopy foliage. We studied the variation of the effects of branch excision and transport among branches at these different heights in the canopy. After excision and transport, light-saturated leaf net photosynthetic rates were close to zero for most leaves due to stomatal closure. However, when the branch had acclimated to its new environmental conditions—which took on average 20 min—light-saturated leaf net photosynthetic rates did not significantly differ between the excised and intact branches. We therefore conclude that branch excision does not affect the measurement of light-saturated leaf net photosynthesis, provided that the branch is recut under water and is allowed sufficient time to acclimate to its new environmental conditions.     

Phosphoglycerate dehydrogenase from soybean nodules : partial purification and some kinetic properties

The relationship between single leaf photosynthesis and conductance was examined in cotton (Gossypium hirsutum L.) across a range of environmental conditions. The purpose of this research was to separate and define the degree of stomatal and nonstomatal limitations in the photosynthetic process of field-grown cotton.

Photosynthetic rates were related to leaf conductance of upper canopy leaves in a curvilinear manner. Increases in leaf conductance of CO₂ in excess of 0.3 to 0.4 mole per square meter per second did not result in significant increases in gross or net photosynthetic rates. No tight coupling between environmental influences on photosynthetic rates and those affecting conductance levels was evident, since photosynthesis per unit leaf conductance did not remain constant. Slowly developing water stress caused greater reductions in photosynthesis than in leaf conductance, indicating nonstomatal limitations of photosynthesis.

Increases in external CO₂ concentration to levels above ambient did not produce proportional increases in photosynthesis even though substomatal or intercellular CO₂ concentration increased. The lack of a linear increase in photosynthetic rate in response to increases in leaf conductance and in response to increases in external CO₂ concentration demonstrated that nonstomatal factors are major photosynthetic rate determinants of cotton under field conditions.

     

Acclimation of photosynthesis to lightflecks in tomato leaves: interaction with progressive shading in a growing canopy

Plants in natural environments are often exposed to fluctuations in light intensity, and leaf‐level acclimation to light may be affected by those fluctuations. Concurrently, leaves acclimated to a given light climate can become progressively shaded as new leaves emerge and grow above them. Acclimation to shade alters characteristics such as photosynthetic capacity. To investigate the interaction of fluctuating light and progressive shading, we exposed three‐week old tomato (Solanum lycopersicum ) plants to either lightflecks or constant light intensities. Lightflecks of 20 s length and 1000 μmol m^?2 s^?1 peak intensity were applied every 5 min for 16 h per day, for 3 weeks. Lightfleck and constant light treatments received identical daily light sums (15.2 mol m^?2 day^?1). Photosynthesis was monitored in leaves 2 and 4 (counting from the bottom) during canopy development throughout the experiment. Several dynamic and steady‐state characteristics of photosynthesis became enhanced by fluctuating light when leaves were partially shaded by the upper canopy, but much less so when they were fully exposed to lightflecks. This was the case for CO₂‐saturated photosynthesis rates in leaves 2 and 4 growing under lightflecks 14 days into the treatment period. Also, leaf 2 of plants in the lightfleck treatment showed significantly faster rates of photosynthetic induction when exposed to a stepwise change in light intensity on day 15. As the plants grew larger and these leaves became increasingly shaded, acclimation of leaf‐level photosynthesis to lightflecks disappeared. These results highlight continuous acclimation of leaf photosynthesis to changing light conditions inside developing canopies.