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.     

Leaf display and photosynthesis of tree seedlings in a cool-temperate deciduous broadleaf forest understorey

To examine a possible convergence in leaf photosynthetic characteristics and leaf display responses to light environment in seedlings of three canopy and two shrub tree species in understorey of cool-temperate deciduous broadleaf forest, relationships between light environment, leaf orientation and leaf light-photosynthetic response were measured. Light capture of the seedlings (17-24 individuals with 2-12 leaves for each species) was assessed with a three dimensional geometric modeling program Y-plant. Leaf photosynthetic characteristics of the five species were found to have acclimated to the understorey light environment, i.e., low light compensation point and high apparent quantum yield. In addition, light-saturated photosynthetic rates were higher in seedlings inhabiting microsites with higher light availability. Efficiencies of light capture and carbon gain of the leaf display were evaluated by simulating the directionalities of light capture and daily photosynthesis for each seedling using hemispherical canopy photography. The results showed that most of the seedlings orientated their leaves in a way to increase the daily photosynthesis during the direct light periods (sunflecks) rather than maximize daily photosynthesis by diffuse light. Simulations also showed that daily photosynthesis would increase only 10% of that on actual leaf display when the leaves orientated to maximize the diffuse light interception. Simulations in which leaf orientations were varied showed that when the leaf display fully maximized direct light interception, the time that leaves were exposed to excessive photon flux density of >800 mumol photons m(-2) s(-1) were doubled. The understorey seedlings studied responded to the given light environments in a way to maximize the efficiency of acquisition and use of light during their short (approximately 3 month) seasonal growth period.     

Bundle sheath leakiness and light limitation during C4 leaf and canopy CO2 uptake

Perennial species with the C(4) pathway hold promise for biomass-based energy sources. We have explored the extent that CO(2) uptake of such species may be limited by light in a temperate climate. One energetic cost of the C(4) pathway is the leakiness () of bundle sheath tissues, whereby a variable proportion of the CO(2), concentrated in bundle sheath cells, retrodiffuses back to the mesophyll. In this study, we scale from leaf to canopy level of a Miscanthus crop (Miscanthus x giganteus hybrid) under field conditions and model the likely limitations to CO(2) fixation. At the leaf level, measurements of photosynthesis coupled to online carbon isotope discrimination showed that leaves within a 3.3-m canopy (leaf area index = 8.3) show a progressive increase in both carbon isotope discrimination and as light decreases. A similar increase was observed at the ecosystem scale when we used eddy covariance net ecosystem CO(2) fluxes, together with isotopic profiles, to partition photosynthetic and respiratory isotopic flux densities (isofluxes) and derive canopy carbon isotope discrimination as an integrated proxy for at the canopy level. Modeled values of canopy CO(2) fixation using leaf-level measurements of suggest that around 32% of potential photosynthetic carbon gain is lost due to light limitation, whereas using determined independently from isofluxes at the canopy level the reduction in canopy CO(2) uptake is estimated at 14%. Based on these results, we identify as an important limitation to CO(2) uptake of crops with the C(4) pathway.     

Sexes show contrasting patterns of leaf and crown carbon gain in a dioecious rainforest shrub

The sexes of dioecious species may differ in a range of vegetative and reproductive traits as well as in physiological traits. In Siparuna grandiflora, a Neotropical dioecious shrub, we examined differences in leaf-level photosynthesis of different classes of leaf age and, using simulation models, explored whether differences in leaf-level carbon gain led to sex differences in whole-plant daily carbon gain. Male plants had higher photosynthetic capacity at the leaf level. As leaves of both sexes aged their photosynthetic capacity and specific leaf area declined as expected. Simulations of daily carbon gain using the architecturally explicit model Y-Plant and a non-architectural model incorporating a wide range of realistic light environments revealed that the difference in leaf-level photosynthetic capacity did not translate into greater crown-level carbon gain for males. Rather, differences in patterns of allocation to leaf area allow females to achieve higher crown-level carbon gain. The results demonstrate that sex differences at the leaf level do not necessarily predict patterns at the whole-plant level.     

Altered physiological function,not structure,drives increased radiation-use efficiency of soybean grown at elevated CO<Subscript>2</Subscript>

Previous studies of elevated carbon dioxide concentration ([CO₂]) on crop canopies have found that radiation-use efficiency is increased more than radiation-interception efficiency. It is assumed that increased radiation-use efficiency is due to changes in leaf-level physiology; however, canopy structure can affect radiation-use efficiency if leaves are displayed in a manner that optimizes their physiological capacity, even though the canopy intercepts the same amount of light. In order to determine the contributions of physiology and canopy structure to radiation-use and radiation-interception efficiency, this study relates leaf-level physiology and leaf display to photosynthetic rate of the outer canopy. We used a new imaging approach that delivers three-dimensional maps of the outer canopy during the growing season. The 3D data were used to model leaf orientation and mean photosynthetic electron transport of the outer canopy to show that leaf orientation changes did not contribute to increased radiation-use; i.e. leaves of the outer canopy showed similar diurnal leaf movements and leaf orientation in both treatments. Elevated [CO₂] resulted in an increased maximum electron transport rate (ETR_max) of light reactions of photosynthesis. Modeling of canopy light interception showed that stimulated leaf-level electron transport at elevated [CO₂], and not alterations in leaf orientation, was associated with stimulated radiation-use efficiency and biomass production in elevated [CO₂]. This study provides proof of concept of methodology to quantify structure–function relationships in combination, allowing a quantitative estimate of the contribution of both effects to canopy energy conversion under elevated [CO₂].     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

Does Long-Term Elevation of CO2 Concentration Increase Photosynthesis in Forest Floor Vegetation? (Indiana Strawberry in a Maryland Forest)

As the partial pressure of CO2 (pCO2) in the atmosphere rises, photorespiratory loss of carbon in C3 photosynthesis will diminish and the net efficiency of light-limited photosynthetic carbon uptake should rise. We tested this expectation for Indiana strawberry (Duchesnea indica) growing on a Maryland forest floor. Open-top chambers were used to elevate the pCO2 of a forest floor habitat to 67 Pa and were paired with control chambers providing an ambient pCO2 of 38 Pa. After 3.5 years, D. indica leaves grown and measured in the elevated pCO2 showed a significantly greater maximum quantum efficiency of net photosynthesis (by 22%) and a lower light compensation point (by 42%) than leaves grown and measured in the control chambers. The quantum efficiency to minimize photorespiration, measured in 1% O2, was the same for controls and plants grown at elevated pCO2. This showed that the maximum efficiency of light-energy transduction into assimilated carbon was not altered by acclimation and that the increase in light-limited photosynthesis at elevated pCO2 was simply a function of the decrease in photorespiration. Acclimation did decrease the ribulose-1,5-bisphosphate carboxylase/oxygenase and light-harvesting chlorophyll protein content of the leaf by more than 30%. These changes were associated with a decreased capacity for light-saturated, but not light-limited, photosynthesis. Even so, leaves of D. indica grown and measured at elevated pCO2 showed greater light-saturated photosynthetic rates than leaves grown and measured at the current atmospheric pCO2. In situ measurements under natural forest floor lighting showed large increases in leaf photosynthesis at elevated pCO2, relative to controls, in both summer and fall. The increase in efficiency of light-limited photosynthesis with elevated pCO2 allowed positive net photosynthetic carbon uptake on days and at locations on the forest floor that light fluxes were insufficient for positive net photosynthesis in the current atmospheric pCO2.     

Nonstomatal inhibition of photosynthesis in sunflower at low leaf water potentials and high light intensities

The inhibition of photosynthesis at low leaf water potentials was studied in soil-grown sunflower to determine the degree to which photosynthesis under high light was affected by stomatal and nonstomatal factors. Below leaf water potentials of −11 to −12 bars, rates of photosynthesis at high light intensities were insensitive to external concentrations of CO₂ between 200 and 400 microliters per liter. Photosynthesis also was largely insensitive to leaf temperature between 10 and 30 C. Changes in CO₂ concentration and temperature had negligible effect on leaf diffusive resistance. The lack of CO₂ and temperature response for both photosynthesis and leaf diffuse resistance indicates that rates of photosynthesis were not limited by either CO₂ diffusion or a photosynthetic enzyme. It was concluded that photosynthesis under high light was probably limited by reduced photochemical activity of the leaves at water potentials below −11 to −12 bars.     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

Physiological and morphological acclimation of shade-grown tree seedlings to late-season canopy gap formation

Because acclimation to canopy gaps may involve coordination of new leaf production with morphological or physiological changes in existing, shade-developed leaves, we examined both new leaf production and photosynthesis of existing leaves on shade-grown seedlings after exposure to a late-season canopy gap. Midway through the summer, we transferred potted, shade-grown seedlings of four co-occurring temperate deciduous tree species representing a range of shade-tolerance categories and leaf production strategies to gaps. Shade-tolerant Acer saccharum was the least responsive to gap conditions. It produced few new, high-light acclimated leaves and increases in photosynthetic rates of shade-developed leaves appeared stomatally limited. Intermediately shade-tolerant Fraxinus americana and Quercus rubra responded most, by producing new leaves and increasing photosynthetic rates of existing shade-developed leaves to levels not significantly different from gap-grown controls within four weeks of gap exposure. Shade-intolerant Liriodendron tulipifera was intermediate in response. In these species, the degree of shoot-level morphological acclimation (new leaf production) and leaf-level physiological acclimation (photosynthetic increases in existing leaves) appear coupled. Mechanisms of acclimation also appear related to intrinsic patterns of nitrogen use and mobilization, the ability to adjust stomatal conductance, and shade tolerance.     

Photosynthesis of Eucalyptus globulus with Mycosphaerella leaf disease

Mycosphaerella leaf disease (MLD) is a major cause of foliage damage in Eucalyptus globulus plantations. Our study is the first to describe the physiological effects of MLD on E. globulus leaves. It involved measurements on both field and potted plants. Changes in photosynthetic parameters in response to MLD were quantified in a study using gas exchange techniques. There was a negative linear relationship between light-saturated photosynthesis (A(max)) and leaf-level damage from MLD. Reductions in A(max) were proportionally greater than might be expected from the reduction in green leaf area as a result of the disease, indicating that asymptomatic tissue also was affected by MLD. The reductions in A(max) were not related to increases in stomatal resistance, but were a result of reduced activity of ribulose bisphosphate carboxylase (Rubisco) and changes in the capacity for ribulose bisphosphate (RuBP) regeneration. Changes in mesophyll resistance to CO2 were also implicated. The effect of MLD was similar at different sites and irrespective of tree-level infection, suggesting a general leaf-level response of E. globulus to MLD.     

Thermal acclimation of photosynthesis: on the importance of adjusting our definitions and accounting for thermal acclimation of respiration

While interest in photosynthetic thermal acclimation has been stimulated by climate warming, comparing results across studies requires consistent terminology. We identify five types of photosynthetic adjustments in warming experiments: photosynthesis as measured at the high growth temperature, the growth temperature, and the thermal optimum; the photosynthetic thermal optimum; and leaf-level photosynthetic capacity. Adjustments of any one of these variables need not mean a concurrent adjustment in others, which may resolve apparently contradictory results in papers using different indicators of photosynthetic acclimation. We argue that photosynthetic thermal acclimation (i.e., that benefits a plant in its new growth environment) should include adjustments of both the photosynthetic thermal optimum (T _opt) and photosynthetic rates at the growth temperature (A _growth), a combination termed constructive adjustment. However, many species show reduced photosynthesis when grown at elevated temperatures, despite adjustment of some photosynthetic variables, a phenomenon we term detractive adjustment. An analysis of 70 studies on 103 species shows that adjustment of T _opt and A _growth are more common than adjustment of other photosynthetic variables, but only half of the data demonstrate constructive adjustment. No systematic differences in these patterns were found between different plant functional groups. We also discuss the importance of thermal acclimation of respiration for net photosynthesis measurements, as respiratory temperature acclimation can generate apparent acclimation of photosynthetic processes, even if photosynthesis is unaltered. We show that while dark respiration is often used to estimate light respiration, the ratio of light to dark respiration shifts in a non-predictable manner with a change in leaf temperature.     

Effects of rising temperatures and [CO2] on the physiology of tropical forest trees

Using a mixture of observations and climate model outputs and a simple parametrization of leaf-level photosynthesis incorporating known temperature sensitivities, we find no evidence for tropical forests currently existing "dangerously close" to their optimum temperature range. Our model suggests that although reductions in photosynthetic rate at leaf temperatures (TL) above 30 degrees C may occur, these are almost entirely accountable for in terms of reductions in stomatal conductance in response to higher leaf-to-air vapour pressure deficits D. This is as opposed to direct effects of TL on photosynthetic metabolism. We also find that increases in photosynthetic rates associated with increases in ambient [CO2] over forthcoming decades should more than offset any decline in photosynthetic productivity due to higher D or TL or increased autotrophic respiration rates as a consequence of higher tissue temperatures. We also find little direct evidence that tropical forests should not be able to respond to increases in [CO2] and argue that the magnitude and pattern of increases in forest dynamics across Amazonia observed over the last few decades are consistent with a [CO2]-induced stimulation of tree growth.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

An Explanation for the Difference in Photosynthetic Capabilities of Healthy and Beet Yellows Virus-infected Sugar Beets (Beta vulgaris L.)

Sugar beets (Beta vulgaris L.) infected with the Beet Yellows Virus exhibit lower rates of net photosynthesis at light saturation than do healthy plants. These Pn reductions were correlated with increases in leaf resistance to water vapor loss. Theoretical analyses demonstrated that, although the leaf resistance to water vapor loss increases could account for a major part of the net photosynthesis decreases, some other aspect of leaf functioning also was debilitated by infection. Both the levels and the activities of ribulose-1, 5-diP carboxylase were less on a leaf area basis in extracts from infected leaves than from healthy ones. Soluble carbohydrates accumulate in Beet Yellows Virus-infected leaves, but inhibiting translocation in several ways provided no evidence in support of the hypothesis that the accumulation of photosynthates in leaves has a direct, short term, feed-back effect upon the photosynthetic rate.     

甜瓜幼苗叶片光合变化特性

为探讨甜瓜光响应变化特性与环境因子的关系,选择光响应曲线适宜测定的时段,以甜瓜幼苗为试材,将1 d分为3个时段:10:00-12:00、12:00-15:00和15:00-17:00,每个叶位叶片测定1 d,并采用直角双曲线修正模型拟合光响应曲线,研究不同时段下甜瓜叶片光响应曲线、光响应参数的变化趋势和不同叶位叶片光响应参数特性。结果表明:当环境中光合有效辐射增强,叶面温度(Tl)升高,空气相对湿度(RH)降低;当环境中光合有效辐射减弱,Tl降低,RH升高。10:00-12:00光响应曲线和12:00-15:00的第1-4叶光响应曲线呈双曲线,在强光下趋向饱和状况,12:00-15:00的第5叶光合速率和15:00-17:00光合速率在强光下出现明显的光抑制现象。1 d的不同时段均表现为10:00-12:00最大净光合速率(P_max)和光饱和点(LSP)最高,12:00-17:00降低;12:00-15:00光补偿点(LCP)和暗呼吸速率(R_d)较高,其它两个时段较低,10:00-17:00光补偿点量子效率(φ_c)、气孔导度(G_s)和胞间CO₂浓度(C_i)总体呈降低趋势,气孔限制值(L_s)升高。10:00-15:00相同时段测得的不同叶位叶片光响应参数,以第4-5叶光合性能较好,15:00-17:00以第3叶P_max最高,第5叶次之;10:00-17:00 G_s和C_i以第5叶较低,第1叶较高,L_s以第5叶较高,第1叶较低。RH为影响P_max的主要决策因子,测定时段、叶面饱和蒸汽压亏缺(Vpdl)和Tl为主要限制因子。10:00-12:00适宜进行光响应曲线测定,气孔限制为不同时段光合作用不同的主要因素,非气孔限制为影响不同叶位叶片光合作用的主要因素。     

Influence of light intensity during growth on photosynthesis and activity of several key photosynthetic enzymes in a C4 plant (Zea mays)

Maize ( Zea mays L. Hybrid Sweet Corn, Royal Crest), a C₄ plant, was grown under different light regimes, after which the rate of photosynthesis and activities of several photosynthetic enzymes (per unit leaf chlorophyll) were measured at different light intensities. Plants were grown outdoors under direct sunlight or 23% of direct sunlight, and in growth chambers at photosynthetic photon flux densities of about 20% and 8% of direct sunlight. The plants grown under direct sunlight had a higher light compensation point than plants grown under lower light. At a light intensity about 25% of direct sunlight, plants from all growth regimes had a similar rate of photosynthesis. Under saturating levels of light the plants grown under direct sunlight had a substantially higher rate of photosynthesis than plants grown under the lower light regimes. The higher photosynthetic capacity in the plants grown under direct sunlight was accompanied by an increased activity of several photosynthetic enzymes and in the amount of the soluble protein in the leaf. Among five photosynthetic enzymes examined, RuBP carboxylase (EC 4.1.1.39) and pyruvate, Pi dikinase (EC 2.7.9.1) were generally just sufficient to account for rates of photosynthesis under saturating light; thus, these may be rate limiting enzymes in C₄ photosynthesis. Pyruvate, Pi dikinase and NADP-malate dehydrogenase (EC 1.1.1.82) were the only enzymes examined which were light activated and increased in activity with increasing light intensity. In the low light grown plants the activity of pyruvate, Pi dikinase closely paralleled the photosynthetic rate measured under different light levels. With the plants grown under direct sunlight, as light intensity was increased the activation of pyruvate, Pi dikinase and NADP⁺-malate dehydrogenase proceeded more rapidly than photosynthesis.     

The Control of the Light Acclimation of Photosynthesis in Glycine max: Dependence on Import as Modified by Intraplant Shading

The dependence of photosynthetic capacity on imported and locally-assimilatedsupplies of carbon during leaf development under different irradianceswas investigated in Glycine max. The potential export of carbonto the developing, mainstem trifoliate leaf (source-potential)was restricted non-destructively by shading all lower, sourceleaves (source-shading), while local photosynthesis was modifiedconcurrently by exposing the young leaf to different light levelsduring development. When source-shading was applied below the2nd mainstem trifoliate leaf at the bud stage of development,photosynthetic capacity was unaffected in leaves which had developedunder moderate and low irradiances (500 and 250 µmol PARm ^–2 s^–1 respectively), but was reduced significantlyin leaves developed under a high irradiance (900 µmolPAR m ^–2 s^–1). If source-shading was applied beneaththe 2nd leaf at unfolding, the reduction of photosynthetic capacityunder the high irradiance was relatively minor. The photosyntheticcapacity attained by the 2nd leaf during development under differentirradiances was influenced by the previous light environmentof the whole plant. In contrast to the 2nd leaf, the photosyntheticcapacities of the 1st and 4th mainstem leaves were relativelyunaffected by source-shading, even under the highest light regime.While photosynthetic capacity showed a widespread insensitivityto the light level of the lower region of the canopy, source-shadingreduced final leaf size irrespective of node position or localirradiance during leaf development. These effects were not relatedto differences in daily photosynthesis by the expanding leaf,and are discussed in terms of the source/sink balance of thedeveloping leaf. Key words: Glycine max, source-shading, photosynthetic capacity     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.