遮荫对库拉索芦荟细胞超微结构和芦荟素含量的影响

分别用透射电子显微镜技术、高效液相色谱法研究了生长在遮荫和自然光照条件下库拉索芦荟叶片的超微结构和芦荟素含量.电镜观察结果表明,遮荫处理6个月后,库拉索芦荟叶片细胞中叶绿体基粒数量减少,类囊体片层数目减少且排列疏松,质体中原来积累的淀粉粒逐渐减少直至消失;内质网、高尔基体等内膜系统不发达.高效液相色谱分析结果显示,生长在遮荫条件下的库拉索芦荟叶片,芦荟素含量明显低于生长在自然光照下的含量.其中,遮荫下幼叶芦荟素的含量是自然光照下的63.33%,而成熟叶芦荟素含量仅有自然光照下的23.77%.无论自然光下还是遮荫条件下生长的芦荟,芦荟素的含量与叶龄有显著的负相关性,遮荫对成熟叶的影响更大.综合两方面的实验结果认为,遮荫首先影响芦荟叶片细胞内膜系统的发育,进而限制了芦荟素的合成和运输,使芦荟叶片中芦荟素含量降低.     

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.     

Possible Mechanisms of Adaptive Leaf Senescence

Abstract: Availability of nitrogen almost always limits plant growth. Therefore, efficient use of nitrogen is essential for the plants. In upright plants, especially when they form dense plant stands, old, lower leaves are shaded by young, upper leaves. Nitrogenous compounds in such shaded leaves are degraded and re-allocated to the developing young, upper leaves. These processes raise efficiency of nitrogen use in photosynthetic production of the plant. For this to occur in the most effective way, leaves would need to sense their photosynthetic status in a plant and increase, maintain or decrease their photosynthetic capacity accordingly. Hypotheses that explain how a leaf can sense its photosynthetic status in the plant are reviewed. They include systems involving phytochrome, sugar-sensing, or cytokinin. Our experimental results with Helianthus annuus and Phaseolus vulgaris plants, which were subject to various shading treatments, are examined in the light of these hypotheses. Our experimental results favoured the sugar-sensing hypothesis: A leaf can sense demand of other plant parts for photosynthates produced by it and nitrogen abundance or deficiency by monitoring its sugar concentration. Problems that are to be challenged in the near future are also pointed out.     

Effects of Shading on the Senescence and Photosynthetic Physiology of the Early-Flowering Rice Mutant FTL10 at Noon

FTL10 is an early-flowering mutant of rice (Oryza sativa L.) with a premature senescent phenotype. Early leaf senescence can cause negative effects on rice yield. Moreover, rice leaves are damaged under high-light conditions, which promote rice senescence. Artificial shading can reduce the amount of light absorbed by rice leaves. The aim of this study was to investigate the effects of shading at noon (11:30–14:00) on the senescence and yield of FTL10. The results showed that shading improved the total antioxidant capacity of rice leaves, reduced the accumulation of reactive oxygen species (ROS) and reduced the expression of genes related to senescence. In the shaded group, the degradation rate of chlorophyll and Rubisco proteins, which are related to photosynthesis, was relatively slow, and the photosynthetic rate was relatively high. Compared with those under the natural growth conditions, the proportion of photosynthetic electron allocated to photorespiration in the shaded group rice leaves was lower, and the proportion allocated to carbon fixation was higher. The yield data showed that the single-spike weight and yield per plant of rice significantly increased after shading. Therefore, our research shows that shading at noon could delay FTL10 senescence and increase yields.

     

The Effect of Light Intensity on Certain Biological Characters of Rauvolfia

The effect of light intensity on Rauvolfia yunnanensis was studied under four different light conditions, namely: 1. 100% daylight (shadeless); 2.73% (27% shaded); 3. 48% (52% shaded) ; 4. 28% (72% shaded). The most suitable light intensity required for the growth of plant, both on the plant height and diameter, was varying from 28% daylight (at dry season) to 73% daylight (at foggy season) at different seasons. Under the half shaded condition (48% light intensity), the annual production of roots, leaves and stems was greater than under any other light conditions (100%, 73% or 28% light intensity). The total number of flowers was depressed markedly by shading, but the number of fruits yielded was higher under 73% light intensity. The number of embryoless seeds was as high as 96% under full light condition (100% light intensity), but decreased apparently under the shading of forest canopy. The size of leaves, fruits and seeds was varying quite a bit with different light intensities. Finally, we have to mention that the alkaloid contents in the leaves tended to increase with the decreasing of the light intensity.     

Shade avoidance in Trifolium repens: costs and benefits of plasticity in petiole length and leaf size

We tested whether the degree of shade-induced plasticity in petiole length and leaf area is related to the mean trait value expressed under high-light conditions, and to what extent trait values expressed under high-light and shaded conditions affect plant performance. Thirty-four Trifolium repens genotypes were used with a wide range of petiole lengths and leaf areas. Plants were subjected to a high-light environment and two shading regimes: homogeneous shading and a vertical light gradient. Absolute petiole elongation in response to both shading treatments and absolute leaf area expansion in response to homogeneous shading were independent of the trait values expressed in high light. Consequently, relative plasticity was higher for genotypes with lower high-light trait values. Plasticity was associated with enhanced plant performance in a vertical light gradient but not in homogeneously shaded conditions. We also found costs associated with the ability to express plasticity. Our results suggest that selection can act separately on trait values expressed under high-light conditions and on the degree of plasticity.     

Improving photosynthetic performance and some fruit quality traits in mango trees by shading

Excess solar radiation under hot climate can lead to decline in photosynthetic activity with detrimental effects on growth and yield. The aim of this study was to evaluate the use of a transparent plastic roof as shading for diurnal changes in photosynthetic gas exchange, chlorophyll fluorescence, fruit set and quality of mango (Mangifera indica L.) cv. ‘Nam Dok Mai’ growth in the field conditions. Fully expanded leaves were examined either shaded by the plastic roof or sunlit under natural conditions. Leaf temperature and leaf-to-air vapour pressure deficit of the shaded leaves measured on the clear day were lowered compared to those of the sunlit leaves. It resulted in increased stomatal conductance and photosynthetic rates of the shaded leaves compared to those of the sunlit leaves, especially from the morning to midday. Furthermore, the reversible decrease of the maximal quantum yield of PSII was more pronounced in the sunlit leaves than that in the shaded ones. Shading increased the total fruit number; the shaded fruits developed better external color than that of the sun-exposed fruits. Our results indicated that shading could maintain the high photosynthetic activity by reducing stomatal limitations for carbon supply and was effective in alleviating the photoinhibitory damage to PSII during bright and clear days with excessive radiation. Finally, shading could increase the number of fruits and improve mango peel color.     

Local weak light induces the improvement of photosynthesis in adjacent illuminated leaves in maize seedlings

To copy with highly heterogeneous light environment, plants can regulate photosynthesis locally and systemically, thus, maximizing the photosynthesis of individual plants. Therefore, we speculated that local weak light may induce the improvement of photosynthesis in adjacent illuminated leaves in plants. In order to test this hypothesis, maize seedlings were partially shaded, and gas exchange, chlorophyll a fluorescence and biochemical analysis were carefully assessed. It was shown that local shading exacerbated the declines in the photosynthetic rates, chlorophyll contents, electron transport and carbon assimilation‐related enzyme activities in shaded leaves as plants growth progressed. While, the decreases of these parameters in adjacent illuminated leaves of shaded plants were considerably alleviated compared to the corresponding leaves of control plants. Obviously, the photosynthesis in adjacent illuminated leaves in shaded plants was improved by local shading, and the improvement in adjacent lower leaves was larger than that in adjacent upper ones. As growth progressed, local shading induced higher abscisic acid contents in shaded leaves, but it alleviated the increase in the abscisic acid contents in adjacent leaves in shaded plants. Moreover, the difference in sugar content between shaded leaves and adjacent illuminated ones was gradually increased. Consequently, local weak light suppressed the photosynthesis in shaded leaves, while it markedly improved the photosynthesis of adjacent illuminated ones. Sugar gradient between shaded leaves and adjacent illuminated ones might play a key role in photosynthetic regulation of adjacent illuminated leaves.     

苗期遮光光质对生姜光合及生长的影响

以不同颜色塑料薄膜为遮光材料,研究了苗期遮光光质对生姜生长及光合作用的影响.结果表明:幼苗覆膜期,生姜叶片叶绿素含量以蓝膜及绿膜处理较高,白膜次之,红膜较低;叶片P_n则以绿膜处理较高,为14.9 μmol·m^-2·s^-1（第4叶）,分别较白膜、红膜及蓝膜提高5.7%、10.4%和18.3%.旺盛生长期撤膜后,P_n较幼苗期升高,但处理间的变化趋势与幼苗期相似;新生叶片叶绿素含量除红膜处理较低外,其它处理无显著差异,但下位叶片叶绿素含量则以蓝膜和红膜处理显著低于绿膜和白膜处理.蓝膜处理生姜植株茎秆增高、变细,分枝数较少;绿膜处理植株根、茎、叶及根茎鲜质量较高,白膜、红膜及蓝膜处理依次降低,收获时,其产量分别达57 000、53 709、51 487和48 712 kg·hm^-2.说明生姜苗期采用绿膜遮光,可增强叶片光合作用,促进植株生长,提高生姜产量.     

分株比例对异质光环境下美丽箬竹克隆系统光合生理的影响

分株数量或生物量比例差异会明显影响克隆系统对资源异质性环境的生态适应性, 地下茎木质化、连接稳固的竹类植物在生长过程中相连克隆分株通常会生活在异质光环境中, 但其叶片光合生理特性对异质光环境的响应及其分株比例效应则未见报道。该研究以地下茎相连的美丽箬竹(Indocalamus decorus)克隆系统为实验材料, 设置2个遮光率(分别为50% ± 5%和75% ± 5%)和3个分株比例(遮光与未遮光分株比例分别为1:3、2:2、3:1)处理, 分株数量为4株。分别测定了遮光处理后30、90、150天遮光和未遮光分株叶片光响应特征、气体交换参数、光合色素含量, 分析了异质光环境下美丽箬竹光合生理的变化规律。结果显示: 分株比例对美丽箬竹光合生理有显著影响, 且其与遮光、处理时间交互作用显著。美丽箬竹克隆系统遮光分株比例越大, 即遮光相对分株数量越多, 其表观量子效率(AQE)、光饱和点、最大净光合速率(P_{n max})、净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、水分利用效率越大, 光补偿点、暗呼吸速率越小, 其光合效率越高, 光能利用能力越强, 而与之相连的未遮光分株则相反; 随遮光分株比例的增大, 遮光分株叶片叶绿素a、叶绿素b含量呈先升高而后下降的变化趋势, 类胡萝卜素(Car)含量则持续下降, 而与之相连的未遮光分株叶片光合色素含量则呈下降趋势; 遮光率提高, 相同分株比例美丽箬竹克隆系统遮光分株叶片AQE、P_{n max}、P_n、G_s、胞间CO₂浓度(C_i)和光合色素含量总体升高, 而与之相连的未遮光分株叶片P_{n max}、C_i以及Car含量则总体下降。研究结果表明异质光环境下, 遮光分株光合效率和弱光利用能力明显增强, 而未遮光分株则相反, 克隆系统内分株间发生了明显的克隆分工, 且2:2、3:1分株比例克隆系统较1:3分株比例对异质光环境具有更好的适应能力。美丽箬竹克隆系统可通过差异性调节分株光合生理特性和光合色素含量, 提高遮光分株光能利用和光合效率来适应异质光环境, 以提高克隆系统的适合度。     

遮荫和全光下生长的棉花光合作用和叶绿素荧光特征

 遮荫条件下（遮荫下光强相当于自然光强的40％左右）棉花(Gossypium hirsutum)叶片光合速率明显降低,仅为自然光强下生长叶片的30%～40％,叶片中RuBP羧化酶活性降低,而表观量子效率(AQY)较高。不同光照条件下生长的棉花叶片对短时间持续光强的光合诱导过程有明显的差异,由弱光转到强光下,自然光强下生长的叶片的Pn、Gs、ΦPSⅡ及非光化学猝灭系数(NPQ)都能在较短的时间内达到最大值,而遮荫叶片需要的时间较长;遮荫下生长的棉花叶片的实际光化学效率,随光强的增加下降幅度较大,而自然光照下生长的叶片下降幅度较小;自然光照下生长的叶片的NPQ随光强的升高达到较高水平,而遮荫叶片在较低的光强下即达到最大值,此时NPQ较低,遮荫叶片依赖于叶黄素循环的能量耗散水平较低。遮荫叶片较低的光合速率以及过剩光能耗散能力是其转入自然强光后光抑制严重的主要原因。     

Improved growth and water use efficiency of cherry saplings under reduced light intensity

Cherry (Prunus avium L.) saplings were grown under natural sunlight (controls) or moderate shading (up to 30%, depending on the incident light intensity and the hour of the day). Reduced light intensity increased the dry mass of each of the plant components studied. Consequently, the total dry mass of shaded plants was significantly greater than that of controls at the end of the growing season. However, the diurnal trend in the level of photosynthesis (per unit of leaf area) of shaded plants was similar to the controls in August, but lower in September. As the growing season proceeded, reduced photosynthetic rates, thinner mesophyll and larger specific leaf area in the shaded plants indicated that leaf development had adapted to shaded conditions throughout the growing season. It is suggested that increased growth of shaded plants was caused by a higher initial relative growth rate and a greater whole-plant photosynthesis. Shading consistently reduced transpiration over the season, therefore improving water use efficiency of shaded leaves. Our results suggest that a moderate reduction in light intensity can be a useful method for improving growth and saving water in hot and dry environments.     

弱光胁迫对不同基因型玉米生长发育和产量的影响

以不同基因型玉米为材料,在玉米生长发育的3个主要阶段(苗期、穗期、粒期)进行分期遮光试验,研究不同时期弱光胁迫对不同基因型玉米生长发育和产量的影响。结果表明,遮光延缓了玉米叶片的出生速度,使叶片变薄;遮光可以延缓叶片的衰老,但遮光解除后则加速叶片的衰老;遮光造成植株高度增加,但恢复正常光照后,其株高却逐渐低于对照;遮光使干物质积累下降,抽雄吐丝日期推迟,尤其是吐丝日期推迟更多,并使产量降低,但不同基因型玉米不同遮光处理下降程度不同。试验的4个品种中,掖单2 2和豫玉2号受遮光影响较小,而掖单36 38和丹玉13受影响较大,即不同基因型玉米对弱光胁迫的敏感性不同     

The Effect of Shading on the Photosynthetic Rate and Longevity of Grass Leaves

The rate at which the net photosynthesis of grass leaves grownin bright light (119 W m^–2) decreased as they aged wasincreased by severe shading (to 21 W m^–2 or less). However,less severe shading (light intensities of 36 W m^–2 ormore) had no effect. The decrease in photosynthesis was unaffectedby whether the whole plant was shaded or only the leaf whosephotosynthesis was measured. In both shaded and unshaded leaves, photosynthesis measuredin bright light fell faster as the leaf aged than did photosynthesisin dim light. Both mesophyll and stomatal diffusion resistancesrose as the leaf aged but the former rose faster. The chlorophyllcontent fell only towards the end of the life of the leaves.     

遮荫对甜瓜叶片光合特性的影响

以适宜不同栽培条件的3个甜瓜品种为材料,研究了遮荫对其光合色素含量、净光合速率及比叶重等叶片特性的影响。结果表明:与正常光照相比,遮光处理能诱使甜瓜叶片叶绿素和类胡萝卜素含量显著提高,叶绿素a/b显著降低。全光照条件下,3个甜瓜品种的净光合速率日变化曲线均呈双峰型,有明显的"光合午休"现象,而在遮荫条件下则呈单峰曲线变化,且光合峰值出现的时间比全光照下推迟;遮光条件下甜瓜叶片气孔导度日变化曲线与其净光合速率日变化类似。3个品种间比叶重在全光照条件下差异不显著,但遮荫显著降低了壮龄叶片的比叶重,且遮荫强度越重,比叶重越小;品种‘黄河蜜3号’壮龄叶的比叶重降幅(31.83%)显著大于‘银帝’(27.22%)和‘玉金香’(26.01%)。可见,遮荫降低了甜瓜叶片的净光合速率和功能叶片的比叶重,植株通过增加自身叶片光合色素含量以增强对环境的适应性,缓解遮荫对其的影响,品种‘银帝’表现出较强的耐弱光性。     

Response of total night-time respiration to differences in total daily photosynthesis for leaves in a Quercus rubra L. canopy: implications for modelling canopy CO2 exchange

Measurements of photosynthesis and respiration were made on leaves in summer in a Quercus rubra L. canopy at approximately hourly intervals throughout 5 days and nights. Leaves were selected in the upper canopy in fully sunlit conditions (upper) and in the lower canopy (lower). In addition, leaves in the upper canopy were shaded (upper shaded) to decrease photosynthesis rates. The data were used to test the hypothesis that total night‐time respiration is dependent on total photosynthesis during the previous day and that the response is mediated through changes in storage in carbohydrate pools. Measurements were made on clear sunny days with similar solar irradiance and air temperature, except for the last day when temperature, especially at night, was lower than that for the previous days. Maximum rates of photosynthesis in the upper leaves (18.7 μmol m^?2 s^?1) were approximately four times higher than those in the lower leaves (4.3 μmol m^?2 s^?1) and maximum photosynthesis rates in the upper shaded leaves (8.0 μmol m^?2 s^?1) were about half those in the upper leaves. There was a strong linear relationship between total night‐time respiration and total photosynthesis during the previous day when rates of respiration were normalized to a fixed temperature of 20°C, removing the effects of temperature from this relationship. Measurements of specific leaf area, nitrogen and chlorophyll concentration and calculations of the maximum rate of carboxylation activity, V_cmax, were not significantly different between upper and upper shaded leaves 5 days after the shading treatment was started. There were small, but significant decreases in the rate of apparent maximum electron transport at saturating irradiance, J_max (P>0.05), and light use efficiency, ? (P<0.05), for upper shaded leaves compared with those for upper leaves. This suggests that the duration of shading in the experiment was sufficient to initiate changes in the electron transport, but not the carboxylation processes of photosynthesis. Support for the hypothesis was provided from analysis of soluble sugar and starch concentrations in leaves. Respiration rates in the upper shaded leaves were lower than those expected from a relationship between respiration and soluble sugar concentration for fully exposed upper and lower leaves. However, there was no similar difference in starch concentrations. This suggests that shading for the duration of several days did not affect sugar concentrations but reduced starch concentrations in leaves, leading to lower rates of respiration at night. A model was used to quantify the significance of the findings on estimated canopy CO₂ exchange for the full growing season. Introducing respiration as a function of total photosynthesis on the previous day resulted in a decrease in growing season night‐time respiration by 23% compared with the value when respiration was held constant. This highlights the need for a process‐based approach linking respiration to photosynthesis when modelling long‐term carbon exchange in forest ecosystems.     

The Effect of Natural and Artificial Shade on the Macrophytes of Lowland Streams and the Use of Shade as a Management Technique

The reduction of the biomass of three main problem species of submerged aquatic macrophytes was proportional to the light at the stream surface both under artificially shaded sections of stream and in naturally shaded areas when compared to the biomass in adjacent open and unshaded sections of stream. The effect of marginal vegetation in varying the shading effect given to streams of differing width and orientation are described. It is recommended that light should be reduced to about half that presently available in the open, by shading from marginal vegetation but it is warned that too much shade is detrimental to the fish populations of the stream and leads to accentuated local accumulations of leaves. The long term effects are considered but it is expected that partial shading will increase the diversity of submerged plant species. It is suggested that the practice of stream realignment is discontinued and that the natural tendency of streams to create their own meandering channels is allowed but within some defined and generally acceptable framework.     

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.     

Response of soybean photosynthesis and chloroplast membrane function to canopy development and mutual shading

The effect of natural shading on photosynthetic capacity and chloroplast thylakoid membrane function was examined in soybean (Glycine max. cv Young) under field conditions using a randomized complete block design. Seedlings were thinned to 15 plants per square meter at 20 days after planting. Leaves destined to function in the shaded regions of the canopy were tagged during early expansion at 40 days after planting. To investigate the response of shaded leaves to an increase in available light, plants were removed from certain plots at 29 or 37 days after tagging to reduce the population from 15 to three plants per square meter and alter the irradiance and spectral quality of light. During the transition from a sun to a shade environment, maximum photosynthesis and chloroplast electron transport of control leaves decreased by two- to threefold over a period of 40 days followed by rapid senescence and abscission. Senescence and abscission of tagged leaves were delayed by more than 4 weeks in plots where plant populations were reduced to three plants per square meter. Maximum photosynthesis and chloroplast electron transport activity were stabilized or elevated in response to increased light when plant populations were reduced from 15 to three plants per square meter. Several chloroplast thylakoid membrane components were affected by light environment. Cytochrome f and coupling factor protein decreased by 40% and 80%, respectively, as control leaves became shaded and then increased when shaded leaves acclimated to high light. The concentrations of photosystem I (PSI) and photosystem II (PSII) reaction centers were not affected by light environment or leaf age in field grown plants, resulting in a constant PSII/PSI ratio of 1.6 ± 0.3. Analysis of the chlorophyll-protein composition revealed a shift in chlorophyll from PSI to PSII as leaves became shaded and a reversal of this process when shaded leaves were provided with increased light. These results were in contrast to those of soybeans grown in a growth chamber where the PSII/PSI ratio as well as cytochrome f and coupling factor protein levels were dependent on growth irradiance. To summarize, light environment regulated both the photosynthetic characteristics and the timing of senescence in soybean leaves grown under field conditions.     

Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirritaautumnata. Branch‐wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy‐wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non‐shaded leaves. Leaf sugars were lower and protein‐bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell‐wall‐bound proanthocyanidins did not differ between shaded and non‐shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch‐shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r=0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.