Temperature and leaf osmotic potential as factors in the acclimation of photosynthesis to high temperature in desert plants

Seasonal changes in the high temperature limit for photosynthesis of desert winter annuals growing under natural conditions in Death Valley, California were studied using an assay based upon chlorophyll fluorescence. All species of this group were 6 to 9°C more tolerant of high temperature at the end of the growing season (May) than at its beginning (February). Over this same time period, the mean daily maximum air temperatures increased by 12°C. Laboratory studies have demonstrated that increases in thermal tolerance could be induced by increasing growth temperature alone. For plants growing under field conditions there was also a good correlation between the thermal tolerance of leaves and the osmotic potential of leaf water, indicating that increases in the concentrations of some small molecules might also confer increased thermal tolerance. Isolated chloroplast thylakoids subjected to increasing concentrations of sorbitol could be demonstrated to have increased thermal tolerance.     

Carbon and nitrogen content of congeneric annual and perennial grass species: relationships with growth

Seven annual-perennial pairs of grass species (six congeneric and one pair taken at random) were grown under productive conditions in the laboratory in order to investigate which plant characters were responsible for the higher relative growth rate (RGR) of annuals as compared to perennials under these conditions. The nitrogen and carbon concentrations of shoot organs and of the whole plant were higher in annuals than in perennials. This was also the case for the specific absorption rate for nitrate and nitrogen productivity (on whole plant and leaf basis). The range of RGR displayed by the 14 species was large enough (0.15–0.33d⁻¹) to examine the general relationships between RGR and the various parameters measured in the present study. RGR was positively related to plant, leaf blade and sheath nitrogen concentrations, but there was no relationship between RGR and any of the carbon concentrations. RGR also strongly correlated with specific absorption rate for nitrate and with nitrogen productivity. A new factorization of this latter parameter led to the definition of the ‘leaf nitrogen productivity’ (N_LP), which is likely to depend on photosynthetic nitrogen use efficiency. RGR was shown to be strongly correlated with N_LP, but not with the second term of the factorization, namely the proportion of plant nitrogen allocated to the leaves.     

The carbon gain benefits of solar tracking in a desert annual

Abstract. A comparison between two sympatric winter desert annuals, Camissonia claviformis and Malvastrum rotundi folium showed that both gained similar amounts of carbon during a spring day, although by very different means. Camissonia has horizontally fixed leaves which have a very high photosynthetic capacity. The temperature optimum of photosynthesis for this species is near 20°C. Malvastrum has leaves with a lower photosynthetic capacity and a photosynthetic temperature optimum near 30°C. Leaves of the latter species remain normal to the sun throughout the course of the day. The tracking response and high temperature optimum for photosynthesis of Malvastrum result in a high daily carbon gain and also a high water-use efficiency.     

Relationships between photosynthetic activity and silica accumulation with ages of leaf in Sasa veitchii (Poaceae, Bambusoideae)

BACKGROUND AND AIMS: Bamboos have long-lived, evergreen leaves that continue to accumulate silica throughout their life. Silica accumulation has been suggested to suppress their photosynthetic activity. However, nitrogen content per unit leaf area (N(area)), an important determinant of maximum photosynthetic capacity per unit leaf area (P(max)), decreases as leaves age and senescence. In many species, P(max) decreases in parallel with the leaf nitrogen content. It is hypothesized that if silica accumulation affects photosynthesis, then P(max) would decrease faster than N(area), leading to a decrease in photosynthetic rate per unit leaf nitrogen (photosynthetic nitrogen use efficiency, PNUE) with increasing silica content in leaves. METHODS: The hypothesis was tested in leaves of Sasa veitchii, which have a life span of 2 years and accumulate silica up to 41 % of dry mass. Seasonal changes in P(max), stomatal conductance, N(area) and silica content were measured for leaves of different ages. KEY RESULTS: Although P(max) and PNUE were negatively related with silica content across leaves of different ages, the relationship between PNUE and silica differed depending on leaf age. In second-year leaves, PNUE was almost constant although there was a large increase in silica content, suggesting that leaf nitrogen was a primary factor determining the variation in P(max) and that silica accumulation did not affect photosynthesis. PNUE was strongly and negatively correlated with silica content in third-year leaves, suggesting that silica accumulation affected photosynthesis of older leaves. CONCLUSIONS: Silica accumulation in long-lived leaves of bamboo did not affect photosynthesis when the silica concentration of a leaf was less than 25 % of dry mass. Silica may be actively transported to epidermal cells rather than chlorenchyma cells, avoiding inhibition of CO2 diffusion from the intercellular space to chloroplasts. However, in older leaves with a larger silica content, silica was also deposited in chlorenchyma cells, which may relate to the decrease in PNUE.     

Decline of photosynthetic capacity with leaf age and position in two tropical pioneer tree species

The effect of leaf age on photosynthetic capacity, a critical parameter in the theory of optimal leaf longevity, was studied for two tropical pioneer tree species, Cecropia longipes and Urera caracasana, in a seasonally dry forest in Panama. These species continuously produce short-lived leaves (74 and 93 d, respectively) during the rainy season (May-December) on orthotropic branches. However, they differ in leaf production rate, maximum number of leaves per branch, light environment experienced by the leaves, leaf mass per unit area, and nitrogen content. Light-saturated photosynthetic rates for marked leaves of known ages (±1 wk) were measured with two contrasting schemes (repeated measurements vs. chronosequence within branch), which overall produced similar results. In both species, photosynthetic rates and nitrogen use efficiency were negatively correlated with leaf age and positively correlated with light availability. Photosynthetic rates declined faster with leaf age in Cecropia than in Urera as predicted by the theory. The rate of decline was faster for leaves on branches with faster leaf turnover rates. Nitrogen per unit leaf area decreased with leaf age only for Urera. Leaf mass per unit area increased with leaf age, either partly (in Cecropia) or entirely (in Urera) due to ash accumulation.     

氮素对不同开花期棉铃纤维比强度形成的生理基础的影响

于2005年在江苏南京(长江流域下游棉区)和徐州(黄河流域黄淮棉区)棉田设置不同氮素水平(零氮：0 kg N·hm^-2,适氮：240 kg N·hm^-2,高氮：480 kg N·hm^-2)试验,研究氮素对不同开花期棉铃(伏前桃、伏桃和秋桃)纤维比强度形成生理基础的影响.结果表明：与适氮处理相比,零氮处理显著降低了棉铃对位叶氮浓度,增加了C/N,影响程度随开花期的推迟而加大,导致伏桃、秋桃对位叶制造和运输光合产物的能力在棉铃发育中后期大幅度下降,棉纤维的相对生长速率以及纤维发育关键酶蔗糖合成酶和β-1,3-葡聚糖酶活性降低,纤维素快速累积持续期缩短,纤维比强度显著降低;高氮处理显著增加了棉铃对位叶氮浓度,降低了C/N,影响程度随开花期的推迟而降低,其降低了伏前桃、伏桃发育过程中光合产物向纤维分配的比例、棉铃发育前中期的纤维发育关键酶活性及纤维素累积速率,导致其纤维比强度亦显著降低.综合分析认为,适宜的施氮量可以协调棉花的“源库”关系,有利于促进不同开花期棉铃高纤维比强度的形成.与适氮处理相比,零氮处理的伏前桃、伏桃和秋桃纤维比强度分别降低了1.8%、5.8%和13.0%,高氮处理则分别降低了8.2%、7.4%和-2.4%.     

Photosynthesis,photoinhibition and low temperature acclimation in cold tolerant plants

Cold acclimation requires adjustment to a combination of light and low temperature, conditions which are potentially photoinhibitory. The photosynthetic response of plants to low temperature is dependent upon time of exposure and the developmental history of the leaves. Exposure of fully expanded leaves of winter cereals to short-term, low temperature shiftsinhibits whereas low temperature growthstimulates electron transport capacity and carbon assimilation. However, the photosynthetic response to low temperature is clearly species and cultivar dependent. Winter annuals and algae which actively grow and develop at low temperature and moderate irradiance acquire a resistance to irradiance 5- to 6-fold higher than their growth irradiance. Resistance to short-term photoinhibition (hours) in winter cereals is a reflection of the increased capacity to keep Q_A oxidized under high light conditions and low temperature. This is due to an increased capacity for photosynthesis. These characteristics reflect photosynthetic acclimation to low growth temperature and can be used to predict the freezing tolerance of cereals. It is proposed that the enhanced photosynthetic capacity reflects an increased flux of fixed carbon through to sucrose in source tissue as a consequence of the combined effects of increased storage of carbohydrate as fructans in the vacuole of leaf mesophyll cells and an enhanced export to the crown due to its increased sink activity. Long-term exposure (months) of cereals to low temperature photoinhibition indicates that this reduction of photochemical efficiency of PS II represents a stable, long-term down regulation of PS II to match the energy requirements for CO₂ fixation. Thus, photoinhibition in vivo should be viewed as the capacity of plants to adjust photosynthetically to the prevailing environmental conditions rather than a process which necessarily results in damage or injury to plants. Not all cold tolerant, herbaceous annuals use the same mechanism to acquire resistance to photoinhibition. In contrast to annuals and algae, overwintering evergreens become dormant during the cold hardening period and generally remain susceptible to photoinhibition. It is concluded that the photosynthetic response to low temperatures and susceptibility to photoinhibition are consequences of the overwintering strategy of the plant species.     

Effect of nitrogen nutrition on the carbohydrate repression of photosynthesis in leaves of Phaseolus vulgaris L.

When carbohydrates accumulate in leaves, photosynthesis is repressed. Limited nitrogen nutrition is thought to enhance this repressing effect. However, the interaction between carbohydrate and nitrogen limitation in leaf photosynthesis has not been examined intensively. In this study, we grew Phaseolus vulgaris L. plants at three different nitrogen levels, and examined the effects of sucrose feeding to the roots on the nitrogen content, carbohydrate content and photosynthetic properties of the primary leaves. Nitrogen content and photosynthetic rate were lower and the carbohydrate content was greater in plants grown with limited nitrogen than in well-fertilized plants. Sucrose feeding to the plants increased carbohydrate content and decreased photosynthetic rate and nitrogen content. The increase in carbohydrate content and the decreases in nitrogen content and photosynthetic rate occurred at the same time, and the negative relationship between the carbohydrate content and photosynthetic rate did not differ among nitrogen nutrition levels. These results show that carbohydrate accumulation in the leaves leads to a decrease in photosynthetic rate. At low nitrogen nutrition levels, carbohydrates accumulated markedly, which accelerated this effect. It appears that the nitrogen nutrition level influences leaf photosynthesis through changing the carbohydrate level rather than through modifying sensitivity of the leaf to the carbohydrate level.     

抗旱性不同品种的小麦叶片中光合电子传递和分配对氮素水平的响应

在大田控制条件下,以植物叶片气体交换和叶绿素荧光测定相结合的方法,研究抗旱性不同品种冬小麦拔节期叶片的光合电子传递及激发能利用分配对氮素响应的结果表明,施氮可提高抗旱性不同品种小麦叶片天线色素吸收光能的能力,虽然氮素不能改变激发能在光合碳还原（PCR）和光合碳氧化（PCO）之间的分配比例,但可提高PSⅡ总电子传递速率（JF）和Pn。低氮下不同品种小麦叶片的热耗散比例有差异,但中高氛下叶片之间无显著差异。旱地品种的鼻值随氮素水平的提高而先增加后下降,而水地品种则表现为持续升高：2个小麦品种的叶片J0值随氮素水平的提高而呈持续升高的变化趋势。氮素对叶片PSⅡ反应中心活性有影响．而不同抗旱性品种之间亦有差别,说明施氮可改善小麦叶片热耗散和光化学反应对激发能的竞争关系,从而增强光合机构的自我保护能力。     

Effects of leaf age, nitrogen nutrition and photon flux density on the organization of the photosynthetic apparatus in leaves of a vine (Ipomoea tricolor Cav.) grown horizontally to avoid mutual shading of leaves

Effects of leaf age, nitrogen nutrition and photon flux density (PFD) on the organization of the photosynthetic apparatus in leaves were investigated in a vine, Ipomoea tricolor Cav., which was grown horizontally so as to avoid mutual shading of leaves. The plants were grown hydroponically at two nitrate levels under two growth light treatments. For one group of the plants, leaves were exposed to full sunlight. For another group, respective leaves were artificially shaded in a manner that simulated changes in the light gradient with the development of an erect herbaceous canopy: old leaves were placed under progressively shadier conditions with growth of the plants (canopy-type shading). In all the treatments, chlorophyll (Chl) content gradually decreased with leaf age. Photosystem I (PSI) per Chl was constant, independent of leaf age, nitrogen nutrition and/or PFD. Photosystem II (PSII) and cytochrome / per Chl, and Chl a/b ratio were independent of leaf age and/or nitrogen nutrition but decreased with the decrease in growth PFD. Ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39, RuBPCase) per Chl steeply decreased with decrease in PFD. When leaves grown at the same PFD were compared, RuBPCase/Chl was lower in the plants grown under lower nitrogen availability and also decreased with leaf age in the plants grown without shading. These decreases were attributed to the curvilinear relationship between RuBPCase and Chl in leaves grown at full sunlight, that was independent of nitrogen availability and leaf age. From these results, it is concluded that the composition of the photosynthetic apparatus is independent of leaf age but changes depending on the light environment and total amount of photosynthetic components of the leaf.Abbreviations Chl chlorophyll - cyt f cytochrome f - PFD photon flux density - RuBPCase ribulose-1,5-bisphosphate carboxylase The author thanks Drs. K. Sonoike, Y. Kashino, K. Okada, H. Hatanaka, Y. Suzuki and A. Aoyama for technical advise. The author also thanks Drs. I. Terashima, A. Makino (Tohoku University, Sendai, Japan), Dr. J.R. Evans (Research School of Biological Sciences, Australian National University, Canberra) and Prof. A. Watanabe for valuable suggestions.     

Photosynthetic pathways and ecological distribution ofEuphorbia species in Egypt

Summary The photosynthetic pathways of 42 species of the genusEuphorbia growing wild, naturalized or cultivated in Egypt were investigated. The criteria used included the δ¹³C- and δD-values and Kranz anatomy of the leaves. There is a relationship between the photosynthetic pathway and the ecological conditions in the habitat of a particular species. All 4 CAM species are succulent shrubs, wild or cultivated. The 11 species with C₄ pathways are mainly summer annuals of tropical origin and flourish under the hot summer conditions. The 27 C₃ species are either winter annuals, perennials, perennials flourishing in winter or shrubs; the majority are Mediterranean, European or Saharo-Arabian. Summer annuals with C₃ pathways grow under the shade of the summer crops. Generally, C₃ plants grow under conditions of relatively better water resources and lower temperature than the C₄ plants. The majority of the CAM and C₄ species occur in the southern part of the country, where high temperature is a common feature of the climate.     

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.     

The effect of altered sink-source relations on photosynthetic traits and matter transport during the phase of reproductive growth in the annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis>

Annual plants transport a large portion of carbohydrates and nitrogenous compounds from leaves to seeds during the phase of reproductive growth. This study aimed to clarify how reproductive growth affects photosynthetic traits in leaves and matter transport within the plant in the annual herb Chenopodium album L. Plants were grown in pots and either reproductive tissues or axillary leaves were removed at anthesis. Matter transport was evaluated as temporal changes in dry mass (as a substitute of carbohydrates) and nitrogen content of aboveground organs: leaves, axillary leaves, stems and reproductive tissues. Photosynthetic capacity (light-saturated photosynthetic rate under ambient CO₂ concentration), nitrogen, chlorophyll and soluble protein content were followed in the 20^th leaf that was mature at the start of the experiment. Removal of reproductive tissues resulted in accumulation of dry mass in leaves and axillary leaves, and accumulation of nitrogen in stem as nitrogen resorption from leaves and axillary leaves proceeded with time. Removal of axillary leaves proportionally reduced dry mass and nitrogen allocation to reproductive tissues, thus affecting the quantity but not quality of seeds. Removal treatments did not alter the time course of photosynthetic capacity, nitrogen, chlorophyll or soluble protein content during senescence in the 20^th leaf, but changed the photosynthetic capacity per unit of leaf nitrogen according to demand from reproductive tissues. Together, the results indicate that reproductive tissues affected carbon and nitrogen economy separately. The amount of carbon was adjusted in leaves through photosynthetic capacity and carbohydrate export from them, and the amount of nitrogen was adjusted by transport from stem to reproductive tissues. The plant’s ability to independently regulate carbon and nitrogen economy should be important in natural habitats where the plant carbon-nitrogen balance can easily be disturbed by external factors.     

大气CO2浓度倍增和施氮对冬小麦光合及叶绿素荧光特性的影响

采用开顶式气室,通过土培盆栽实验研究了不同大气CO2浓度(背景空气浓度375μmol·mol-1和倍增浓度750μmol·mol-1)和氮素水平(不施氮和施氮0.25 g/kg)下两个冬小麦品种(小偃6号和小偃22)主要生育期(拔节、孕穗、扬花、灌浆期)叶片叶绿素含量和荧光动力学参数的变化.结果显示,与背景CO2浓度相比,在不施氮条件下大气CO2浓度倍增处理的小麦叶片出现明显的光合下调现象,而施氮时变化不明显;同时,CO2浓度倍增后小麦各主要生育期叶片叶绿素含量均有不同程度地下降,荧光参数初始荧光(F0)值明显提高,最大荧光(Fm)、可变荧光(Fv)、最大光能转换效率(Fv/Fm)和PSⅡ潜在活性(Fv/F0)值均显著降低.施氮可提高冬小麦各个时期叶片叶绿素含量、Fm、Fv、和Fv/F0值,降低F0值;不施氮条件下,大气CO2浓度倍增对冬小麦各主要生育时期叶绿素含量和荧光参数的影响明显,而施氮后影响微弱.研究表明,大气CO2浓度升高对冬小麦光合速率、叶绿素含量和光系统Ⅱ(PSⅡ)的光合电子传递和潜在活性具有一定抑制作用,通过施氮可以有效地缓解其负面效应.     

不同发育时期追施氮肥对冬小麦旗叶中光合酶活性的影响

小麦开花后,随着旗叶的衰老,旗叶中1,5-二磷酸核酮糖羧化酶（RuBPC）、磷酸烯醇式丙酮酸羧化酶（PEPC）和乙醇酸氧化酶（GO）活性呈下降趋势。随着追施氮肥时期的推迟,光合酶活性呈增加趋势,这意味着氮肥追施时间后移有利于提高小麦光合速率。在旗叶衰老后期,大穗型品种小麦旗叶中光合酶活性略高于多穗型品种小麦。     

Decline of photosynthetic capacity with leaf age in relation to leaf longevities for five tropical canopy tree species

The effect of leaf aging on photosynthetic capacities was examined for upper canopy leaves of five tropical tree species in a seasonally dry forest in Panama. These species varied in mean leaf longevity between 174 and 315 d, and in maximum leaf life span between 304 and 679 d. The light-saturated CO2 exchange rates of leaves produced during the primary annual leaf flush measured at 7-8 mo of age were 33-65% of the rates measured at 1-2 mo of age for species with leaf life span of < 1 yr. The negative regression slopes of photosynthetic capacity against leaf age were steeper for species with shorter maximum leaf longevity. In all species, regression slopes were less steep than the slopes predicted by assuming a linear decline toward the maximum leaf age (20-80% of the predicted decline rate). Maximum oxygen evolution rates and leaf nitrogen content declined faster with age for species with shorter leaf life spans. Statistical significance of regression slopes of oxygen evolution rates against leaf age was strongest on a leaf mass basis (r = 0.49-0.87), followed by leaf nitrogen basis (r = 0.48-0.77), and weakest on a leaf area basis (r = 0.35-0.70).     

增强UV-B辐射和氮对谷子叶光合色素及非酶促保护物质的影响

以谷子(Setaria italica(L)Beauv.)为对象,从拔节期开始持续给予低氮(1.875 mmol/L)和高氮(15 mmol/L)两种氮供应条件并从抽穗期开始进行26 d两种强度(4.29、7.12 kJ·m-2·d-1)的增强UV-B辐射处理,研究了谷子叶中光合色素含量、类黄酮含量和苯丙氨酸解氨酶(PAL)活性的变化.结果表明:与高氮供应条件相比,低氮供应条件明显降低了谷子叶中光合色素含量但提高了类胡萝卜素/叶绿素含量比值;在开花期中段和灌浆期中段,高氮供应条件下谷子叶中光合色素含量对增强UV-B辐射比低氮供应条件下的谷子更敏感.从灌浆期开始到处理结束,两种影响因子对谷子叶中类黄酮含量均有显著影响,增强UV-B辐射导致谷子叶中类黄酮含量逐渐升高,且相同增强UV-B辐射强度下低氮供应条件下的谷子叶中类黄酮含量明显高于高氮供应条件下的谷子.谷子叶中PAL活性对两种影响因子的响应较类黄酮含量更加敏感,低氮供应条件使谷子叶中PAL活性明显提高.结合上述指标的相关性分析结果可知,低氮供应条件加强了处于繁殖期主要阶段的谷子叶中类黄酮的积累,并使谷子叶中的类胡萝卜素/叶绿素含量比值明显提高,进而有助于维持谷子叶中光合色素含量在增强UV-B辐射条件下的相对稳定性,对植株抵抗UV-B辐射伤害有利.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.