Seasonal changes in photoprotective mechanisms of leaves from shaded and unshaded field-grown coffee (<Emphasis Type="Italic">Coffea arabica</Emphasis> L.) trees

Coffee is native to shady environments, but often grows and yields better without shade. Thus, it may be reasoned that coffee leaves should display enough plasticity to acclimate themselves to contrasting light environments. However, little is known about mechanisms associated with such plasticity in coffee. This work aimed, therefore, to explore differences in leaf photoprotective mechanisms. Plants were grown in the field and received either 48 or 100% natural light. Evaluations were made using outer leaves from the sun-facing sides of the coffee hedgerow in Viçosa (Brazil) in August and October, when growth and photosynthetic rates are expected to be minimal and maximal, respectively, and in December, when temporary depressions in those variables are common. Regardless of light treatments, coffee leaves showed: (1) very low photosynthetic rates (generally below 2.5 μmol m^?2 s^?1), (2) chronic photoinhibition in August (dry, cool season) that was accompanied by strong loss of pigment concentration, and (3) discrete, dynamic photoinhibition in October and December (rainy, warm season). Compared with shaded leaves, sunlit leaves generally exhibited lower pigment concentration, lower quantum yield of electron transport, steeper inclinations and similar electron transport rate. Total ascorbate pool tended to be larger in sunlit than in shaded leaves (but with similar redox state), whereas activities of key antioxidant enzymes, as well as malondialdehyde accumulation and electrolyte leakage, were similar between those leaf types. As a whole, the photosynthetic apparatus of the coffee tree showed a low phenotypic plasticity to varying irradiance.     

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.     

In High-Light-Acclimated Coffee Plants the Metabolic Machinery Is Adjusted to Avoid Oxidative Stress Rather than to Benefit from Extra Light Enhancement in Photosynthetic Yield

Coffee (Coffea arabica L.) has been traditionally considered as shade-demanding, although it performs well without shade and even out-yields shaded coffee. Here we investigated how coffee plants adjust their metabolic machinery to varying light supply and whether these adjustments are supported by a reprogramming of the primary and secondary metabolism. We demonstrate that coffee plants are able to adjust its metabolic machinery to high light conditions through marked increases in its antioxidant capacity associated with enhanced consumption of reducing equivalents. Photorespiration and alternative pathways are suggested to be key players in reductant-consumption under high light conditions. We also demonstrate that both primary and secondary metabolism undergo extensive reprogramming under high light supply, including depression of the levels of intermediates of the tricarboxylic acid cycle that were accompanied by an up-regulation of a range of amino acids, sugars and sugar alcohols, polyamines and flavonoids such as kaempferol and quercetin derivatives. When taken together, the entire dataset is consistent with these metabolic alterations being primarily associated with oxidative stress avoidance rather than representing adjustments in order to facilitate the plants from utilizing the additional light to improve their photosynthetic performance.     

杂种杨无性系的光系统Ⅱ放氧活性、光合色素及叶绿体超微结构对光胁迫的响应

 用氧电极仪、红外CO2气体分析仪及叶绿素荧光仪,结合透射电镜技术对几个杂种杨无性系在光胁迫下的光系统Ⅱ活性、光合色素及叶绿体超微结构进行了测定。随着预处理光强的增加,各无性系叶片的净光合速率和光系统Ⅱ放氧活性明显下降。二倍体无性系B11的光系统Ⅱ最大光化学效率（Fv/Fm）及实际光化学效率［(Fm′-F)/Fm′］高于两个三倍体无性系B346和B342。强光下三倍体无性系B346的非光化学淬灭远高于B342和二倍体无性系B11。叶绿素含量和光合速率没有明显的线性关系,叶绿素a/b含量在自然光胁迫下存在季节变化,受强光胁迫2个三倍体无性系的叶绿素a/b增大。预处理光强PFD超过3 000 μmol photons·m-2·s-1,各无性系的基粒类囊体片层结构遭到破坏,但二倍体无性系的类囊体片层结构受破坏程度较三倍体无性系轻。叶绿体蛋白合成抑制剂（硫酸链霉素,SM）可加剧叶绿体超微结构的破坏。     

Benzo[1,2‐b:4,5‐b′]Dithiophene–6,7‐Difluoroquinoxaline Small Molecule Donors with >8% BHJ Solar Cell Efficiency

Solution‐processable small molecule (SM) donors are promising alternatives to their polymer counterparts in bulk‐heterojunction (BHJ) solar cells. While SM donors with favorable spectral absorption, self‐assembly patterns, optimum thin‐film morphologies, and high carrier mobilities in optimized donor–acceptor blends are required to further BHJ device efficiencies, material structure governs each one of those attributes. As a result, the rational design of SM donors with gradually improved BHJ solar cell efficiencies must concurrently address: (i) bandgap tuning and optimization of spectral absorption (inherent to the SM main chain) and (ii) pendant‐group substitution promoting structural order and mediating morphological effects. In this paper, the rational pendant‐group substitution in benzo[1,2‐b:4,5‐b′]dithiophene–6,7‐difluoroquinoxaline SMs is shown to be an effective approach to narrowing the optical gap (E_opt) of the SM donors ( SM1 and SM2 ), without altering their propensity to order and form favorable thin‐film BHJ morphologies with PC₇₁BM. Systematic device examinations show that power conversion efficiencies >8% and open‐circuit voltages (V_OC) nearing 1 V can be achieved with the narrow‐gap SM donor analog ( SM2 , E_opt = 1.6 eV) and that charge transport in optimized BHJ solar cells proceeds with minimal, nearly trap‐free recombination. Detailed device simulations, light intensity dependence, and transient photocurrent analyses emphasize how carrier recombination impacts BHJ device performance upon optimization of active layer thickness and morphology.     

The functional divergence of biomass partitioning,carbon gain and water use in Coffea canephora in response to the water supply: Implications for breeding aimed at improving drought tolerance

Robusta coffee (Coffea canephora) is widely cultivated in regions where water availability is the major environmental constraint affecting crop production. The functional divergence associated with biomass partitioning, carbon gain and water use in response to water supply was examined in 10 one-year-old clones of robusta coffee with varying degrees of drought tolerance. The plants were grown outdoors in 24 L pots and either irrigated or subjected to a four-month water deficit. Under conditions of ample irrigation, clones with superior water use ability (i.e., a higher water potential, transpiration rate, apparent hydraulic conductance and biomass partitioning into roots and a lower wood density) displayed enhanced carbon gains. In contrast, under drought conditions, clones that postponed dehydration via more conservative water use rates showed lower relative decreases in stomatal conductance, photosynthetic rates and biomass accumulation. Isotopic signatures (δ¹³C) might be useful for identifying clones with improved performance under drought conditions. Our results suggest that combining useful morphological and physiological traits facilitates the successful assessment of coffee clonal performance in response to drought at the seedling stage. This strategy may be valuable when exploring a large number of genotypes in coffee-breeding programs because it reduces the time and resource costs that would otherwise be wasted on potentially undesirable genotypes.     

Further Studies on Differentiation of Photosynthetic Properties in Sun and Shade Ecotypes of Solidago virgaurea

Measurements of the fraction of the incident light absorbed by diverse Solidago leaves revealed that differences in light harvesting capacity cannot explain the differences in efficiency of utilization of weak light in photosynthesis that have previously been shown to exist between sun and shade ecotypes when these have been grown in strong light and between identical clones of shade ecotypes when grown at different light intensities. Photosynthesis measurements at low and normal oxygen concentrations, provided no evidence that a different degree of inhibition of photo-synthetic CO₂ uptake by atmospheric oxygen is responsible for the observed differences in photosynthetic efficiency, at low or high light intensities. These results support the conclusion that the markedly less efficient use of weak light by shaded habitat clones grown in strong as compared with weak light is caused primarily by damage to the photosystems, or to a site close to them. Measurements of Emerson enhancement and of light-induced absorbance changes provide some evidence that photoreaction II is more affected than I. Enzyme extracts prepared from clones native to an exposed habitat were found to contain considerably higher activities of carboxydismutase (ribulosc-l,5-diphos-phate carboxylase) than from clones native to a shaded habitat when the plants were previously grown at a moderately high light intensity. Exposed habitat clones apparently have a genetically determined, higher capacity to produce the carboxyla-tion enzyme than shaded habitat clones. The high degree of correlation found when the light-saturated rate of CO₂ uptake in vivo of a number of individual Solidago leaves is plotted against the carboxydismutase activities found in the extracts of these same leaves suggests that low carboxydismutase activity is one of the intrinsic properties responsible for the low capacity for light-saturated photosynthesis of clones from shaded habitats. It is concluded from this and other investigations that differentiation between plants from habitats with contrasting light intensities, whether unrelated species or ecotypos of the same species, probably involves the capacity of several component steps of the photosynthetic process.     

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.     

Photosynthetic Adaptation to Light Intensity in Plants Native to Shaded and Exposed Habitats

Photosynthetic adaptation to light intensity has been studied in clones of populations from shaded and exposed habitats of Rumex acetosa and Geum rivale. Clones of the shade species Lamium galeobdolon and the sun species Plantago lanceolata were also included for comparison. The plants were grown under controlled conditions at a high and a low light intensity. The capacity of photosynthetic carbon dioxide uptake at low as well as at saturating light intensities was determined on single attached leaves. As was previously demonstrated in Solidago virgaurea, clones of populations native to shaded and to exposed environments show differences in the photosynthetic response to light intensity during growth. The data provide evidence that populations of the same species native to habitats with contrasting light intensities differ in their photosynthetic properties in an adaptive manner Ln a similar mode as sun and shade species.     

Increased light‐use efficiency sustains net primary productivity of shaded coffee plants in agroforestry system

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light‐use efficiency, thereby reducing the difference in net primary productivity between shaded and non‐shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light‐use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light‐use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.     

Complementary ecosystem services provided by pest predators and pollinators increase quantity and quality of coffee yields

Wild animals substantially support crop production by providing ecosystem services, such as pollination and natural pest control. However, the strengths of synergies between ecosystem services and their dependencies on land-use management are largely unknown. Here, we took an experimental approach to test the impact of land-use intensification on both individual and combined pollination and pest control services in coffee production systems at Mount Kilimanjaro. We established a full-factorial pollinator and vertebrate exclosure experiment along a land-use gradient from traditional homegardens (agroforestry systems), shaded coffee plantations to sun coffee plantations (total sample size = 180 coffee bushes). The exclusion of vertebrates led to a reduction in fruit set of ca 9%. Pollinators did not affect fruit set, but significantly increased fruit weight of coffee by an average of 7.4%. We found no significant decline of these ecosystem services along the land-use gradient. Pest control and pollination service were thus complementary, contributing to coffee production by affecting the quantity and quality of a major tropical cash crop across different coffee production systems at Mount Kilimanjaro.     

Testing ant predation on the coffee berry borer in shaded and sun coffee plantations in Colombia

Soil‐dwelling ants, many of which are generalist predators, are more diverse in shaded than in sun coffee plantations without trees. We compared ant predation on the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Curculionidae: Scolytinae) in three shaded and three sun coffee plantations in Apía, Colombia, in both the wet and the dry seasons. We found that H. hampei adults exposed to ants for 5 days suffered higher removal in shaded plantations and in the wet season. In the laboratory, we observed that ants killed 74–99% of H. hampei adults over the course of 5 days. Ants appear to be important predators of H. hampei, particularly in shaded coffee plantations and in the wet season.     

Limitations to photosynthesis in coffee leaves from different canopy positions.

Limitations to photosynthesis were explored in leaves from four canopy positions of field-grown, unshaded coffee (Coffea arabica L.), a tropical tree species classified as shade-obligatory. Overall, compared to shade (lower) leaves, sun (upper) leaves had higher net carbon assimilation rate (A) (4.5 against 2.0mumolm(-2)s(-1) at most) associated with higher electron transport rate (due to a greater irradiance availability) but unrelated to stomatal and mesophyll conductances, which were similar regardless of leaf position. Neither physiological variable directly involved with photosynthetic carbon gain nor those involved with light capture were able to adjust themselves to match the capacity of the photosynthetic machinery to the light supply. We concluded that: (i) there was no major difference in photosynthetic capacity between sun and shade leaves; (ii) the intrinsic low A in coffee was greatly associated with remarkable low diffusive limitations rather than with biochemical or photochemical constraints; and (iii) morphological (e.g., variations in specific leaf area and leaf inclination) or anatomical plasticity should be of greater acclimative value than physiological plasticity as a mean of coffee leaves to respond to changing irradiance.     

Spatial autocorrelation in two Iris pumila populations estimated on morphological data from natural clones and their samples grown in two different habitats

Morphological data from two Iris pumila populations (measured on native clones, on their replants into the same habitat, and on their transplants into alternative habitat) were combined with native clones spatial position and spatial autocorrelations (SA) were calculated. Naturally growing I. pumila clones revealed significant SA that were positive on small distances and negative on medium ones in both open Hillock and shaded Woodland populations. No significant SA were detected when calculated with original clone positions, but with morphometric data from replants into the experimental plot in the same habitat. Some significant SA were, however, detected when morphometric data from transplants to alternative habitat were used. Detected SA on I. pumila clones were primarily a consequence of spatial structuring of environmental factors but also, in a lesser degree, a result of genetic spatial arrangements (most probably due to patterns of gene flow). The text was submitted by the authors in English.     

Resorption protection. Anthocyanins facilitate nutrient recovery in autumn by shielding leaves from potentially damaging light levels

The resorption protection hypothesis, which states that anthocyanins protect foliar nutrient resorption during senescence by shielding photosynthetic tissues from excess light, was tested using wild-type (WT) and anthocyanin-deficient mutants of three deciduous woody species, Cornus sericea, Vaccinium elliottii (Chapmn.), and Viburnum sargentii (Koehne). WT Betula papyrifera (Marsh) was included to compare the senescence performance of a species that does not produce anthocyanins in autumn. Plants were subjected to three environmental regimes during senescence: an outdoor treatment; a 5-d high-stress (high light and low temperature) treatment followed by transfer to a low-stress environment and a low-stress treatment that served as control. In the outdoor treatment, the appearance of anthocyanins in senescing leaves of WT plants was concomitant with the development of photo-inhibition in mutant plants of all three anthocyanin-producing species. In the high-stress environment, WT plants maintained higher photochemical efficiencies than mutants and were able to recover when transferred to the low-stress environment, whereas mutant leaves dropped while still green and displayed signs of irreversible photooxidative damage. Nitrogen resorption efficiencies and proficiencies of all mutants in both stressful treatments were significantly lower than the WT counterparts. B. papyrifera displayed photochemical efficiencies and nitrogen resorption performance comparable with the highest of the anthocyanin-producing species in all three senescing environments, indicating a photoprotective strategy divergent from the other species studied. These results strongly support the resorption protection hypothesis of anthocyanins in senescing leaves.     

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。     

Spatial autocorrelation in two Iris pumila populations estimated on morphological data from natural clones and their samples grown in two different habitats

Morphological data from two Iris pumila populations (measured on native clones, on their replants into the same habitat, and on their transplants into alternative habitat) were combined with native clones spatial position and spatial autocorrelations (SA) were calculated. Naturally growing I. pumila clones revealed significant SA that were positive on small distances and negative on medium ones in both open Hillock and shaded Woodland populations. No significant SA were detected when calculated with original clone positions, but with morphometric data from replants into the experimental plot in the same habitat. Some significant SA were, however, detected when morphometric data from transplants to alternative habitat were used. Detected SA on I. pumila clones were primarilly a consequence of spatial structuring of environmental factors but also, in lesser degree, a result of genetic spatial arrangements (most probably due to patterns of gene flow).     

Do Clonal Plants Show Greater Division of Labour Morphologically and Physiologically at Higher Patch Contrasts?

Background

When growing in reciprocal patches in terms of availability of different resources, connected ramets of clonal plants will specialize to acquire and exchange locally abundant resources more efficiently. This has been termed division of labour. We asked whether division of labour can occur physiologically as well as morphologically and will increase with patch contrasts.

Methodology/Principal Findings

We subjected connected and disconnected ramet pairs of Potentilla anserina to Control, Low, Medium and High patch contrast by manipulating light and nutrient levels for ramets in each pair. Little net benefit of inter-ramet connection in terms of biomass was detected. Shoot-root ratio did not differ significantly between paired ramets regardless of connection under Control, Low and Medium. Under High, however, disconnected shaded ramets with ample nutrients showed significantly larger shoot-root ratios (2.8∼6.5 fold) than fully-lit but nutrient-deficient ramets, and than their counterparts under any other treatment; conversely, fully-lit but nutrient-deficient ramets, when connected to shaded ramets with ample nutrients, had significantly larger shoot-root ratios (2.0∼4.9 fold) than the latter and than their counterparts under any other treatment. Only under High patch contrast, fully-lit ramets, if connected to shaded ones, had 8.9% higher chlorophyll content than the latter, and 22.4% higher chlorophyll content than their isolated counterparts; the similar pattern held for photosynthetic capacity under all heterogeneous treatments.

Conclusions/Significance

Division of labour in clonal plants can be realized by ramet specialization in morphology and in physiology. However, modest ramet specialization especially in morphology among patch contrasts may suggest that division of labour will occur when the connected ramets grow in reciprocal patches between which the contrast exceeds a threshold. Probably, this threshold patch contrast is the outcome of the clone-wide cost-benefit tradeoff and is significant for risk-avoidance, especially in the disturbance-prone environments.     

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.     

Could shading reduce the negative impacts of drought on coffee? A morphophysiological analysis

Based on indirect evidence, it was previously suggested that shading could attenuate the negative impacts of drought on coffee (Coffea arabica), a tropical crop species native to shady environments. A variety (47) of morphological and physiological traits were examined in plants grown in 30-l pots in either full sunlight or 85% shade for 8 months, after which a 4-month water shortage was implemented. Overall, the traits showed weak or negligible responses to the light × water interaction, explaining less than 10% of the total data variation. Only slight variations in biomass allocation were observed in the combined shade and drought treatment. Differences in relative growth rates were mainly associated with physiological and not with morphological adjustments. In high light, drought constrained the photosynthetic rate through stomatal limitations with no sign of apparent photoinhibition; in low light, such constraints were apparently linked to biochemical factors. Sun-grown plants displayed osmotic adjustments, decreased tissue elasticities and improved long-term water use efficiencies, especially under drought. Regardless of the water availability, higher concentrations of lipids, total phenols, total soluble sugars and lignin were found in high light compared to shade conditions, in contrast to the effects on cellulose and hemicellulose concentrations. Proline concentrations increased in water-deprived plants, particularly those grown under full sun. Phenotypic plasticity was much higher in response to the light than to the water supply. Overall, shading did not alleviate the negative impacts of drought on the coffee tree.