Phenotypic plasticity in response to light in the coffee tree

Phenotypic plasticity to light availability was examined at the leaf level in field-grown coffee trees (Coffea arabica). This species has been traditionally considered as shade-demanding, although it performs well without shade and even out-yields shaded coffee. Specifically, we focused our attention on the morpho-anatomical plasticity, the balance between light capture and excess light energy dissipation, as well as on physiological traits associated with carbon gain. A wide natural light gradient, i.e., a diurnal intercepted photon irradiance differing by a factor of 25 between the deepest shade leaves and the more exposed leaves in the canopy, was explored. Responses of most traits to light were non-linear, revealing the classic leaf sun vs. leaf shade dichotomy (e.g., compared with sun leaves, shade leaves had a lower stomatal density, a thinner palisade mesophyll, a higher specific leaf area, an improved light capture, a lower respiration rate, a lower light compensating point and a limited capacity for photoprotection). The light-saturated rates of net photosynthesis were higher in sunlit than in shade leaves, although sun leaves were not efficient enough to use the extra light supply. However, sun leaves showed well-developed photoprotection mechanisms in comparison to shade leaves, which proved sufficient for avoiding photoinhibition. Specifically, a higher non-photochemical quenching coefficient was found in parallel to increases in: (i) zeaxanthin pools, (ii) de-epoxidation state of the xanthophyll cycle, and (iii) activities of some antioxidant enzymes. Intracanopy plasticity depended on the suite of traits considered, and was high for some physiological traits associated with photoprotection and maintenance of a positive carbon balance under low light, but low for most morpho-anatomical features. Our data largely explain the successful cultivation of the coffee tree in both exposed and shade environments, although with a poor resource-use efficiency in high light.     

Temporal variation of δ<Superscript>13</Superscript>C of larch leaves from a montane boreal forest in Mongolia

This paper reports the temporal variation (2002–2004) in foliar δ¹³C values, which are indicative of long-term integrated photosynthetic and water use characteristics, of Siberian larch (Larix sibirica Ledeb.) trees in a montane forest at Mongonmorit, NE Mongolia. At the stand, the δ¹³C value for understory shaded leaves was more negative by 2‰ on average than that for sunlit leaves sampled concurrently from open and sun-exposed environments in a forest gap. The δ¹³C value of both sunlit and shaded leaves showed pronounced intra- but relatively small inter-seasonal variations. The δ¹³C value was more positive for juvenile than mature leaves. We conjecture that juvenile leaves may derive carbon reserves in woody tissues (e.g., stems). Regardless of leaf habitats, the δ¹³C value was also affected by insect herbivores occurred in mid summer of 2003, being more negative in newly emerging leaves from the twigs after defoliation than in non-defoliated mature leaves. This pattern seems to contrast with that for the juvenile leaves in the early growing season. We surmise that the newly emerging leaves used stored organic carbon that was depleted due to fractionation during remobilization and translocation for leaf regrowth. There was also intra- and inter-seasonal variation in the foliar N concentrations and C:N ratios. A good positive (negative) correlation between the foliar δ¹³C values and N concentrations (C:N ratios) was also observed for both sunlit and shaded leaves, suggesting that the relationship between water and nitrogen use is a crucial factor affecting the plant carbon–water relationship in this mid latitude forest with a cold semiarid climate. Our isotopic data demonstrate that the larches in NE Mongolia exhibits relatively higher water use efficiency with a distinct within-season variability.     

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.     

Relationship between flavonoids and photoprotection in shade-developed Erigeron breviscapus transferred to sunlight

Flavonoids are thought to participate in protection of the photosynthetic apparatus against photoinhibition under excessive light. Flavone glycoside, scutellarin, is a main active ingredient extracted from Erigeron breviscapus, the plant used in Chinese medicine. Shade-developed leaves of E. breviscapus were transferred from shade to full sunlight to quantify a relationship between the concentration of leaf scutellarin and tolerance to high radiation stress or the recovery from photoinhibition. The maximal quantum yield of PSII photochemistry showed a diurnal fluctuation in both shaded and sunlit leaves throughout the day. It indicated dynamic photoinhibition in the leaves of Erigeron, i.e., higher photoinhibition at solar noon and lower one in the morning and late afternoon. The sun-developed leaves reached the higher scutellarin content and values of nonphotochemical quenching coefficient with a lower degree of photoinhibition than the shade-developed leaves. When the shade-developed leaves were transferred to full sunlight, the content of scutellarin was declining continuously for 10 d and then was increasing for 15 d. After 50 d, all leaves became the sun-developed leaves with their scutellarin contents of about 138.5 ± 5.2 mg g^?1(dry mass, DM) which was significantly higher than that of the shade-developed leaves [107.8 ± 9.8 mg g^?1(DM)]. During acclimatization, the degree of photoinhibition was negatively correlated with the scutellarin content. Our results demonstrated a synchronous fluctuation between the flavonoid content and degree of protection against photoinhibition.     

The study of a determinate growth orchid highlights the role of new leaf production in photosynthetic light acclimation

Plants usually respond to the changes of growth irradiance by a combination of the physiological modifications in their preexisting leaves and the production of new leaves. However, those with a determinate growth habit produce certain number of leaves in a growing season and cannot produce new leaves when light condition changes. We used an epiphytic orchid with only one leaf produced every growing season to examine whether and how determinate growth species adapt to changing environments after their preexisting leaves mature. Leaf photosynthesis and anatomy of Pleione aurita were investigated at full expansion and at 40 days after the fully expanded leaves were transferred from high to low light or from low to high light. Leaves show large physiological and morphological plasticity to light gradients at full expansion and the transferred leaves exhibited multiple physiological modifications, including reallocation of nitrogen between light harvesting and carbon fixation, and enhancement of thermal dissipation in their new environments, to optimize carbon assimilation and avoid photoinhibition. Irrespective of the various changes either to shade or sun, the sole preexisting leaf could not fully acclimate to new light environments due to the mesophyll thickness constraint. This leads to the consequence that only plants exposed to high light throughout the experiment had a positive annual biomass gain. Our results highlighted the importance of new leaf production in the carbon accumulation during photosynthetic light acclimation, and contribute new insights of epiphytes physiological responses to their highly dynamic arboreal habitat.     

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.     

Why is it better to produce coffee seedlings in full sunlight than in the shade? A morphophysiological approach

The coffee plant is native to shaded environments and its seedlings are often produced in shaded nurseries. However, some nursery managers, in an effort to improve the acclimation of seedlings to field conditions after transplantation, produce seedlings in full sun exposure. In this study, the morphological and physiological parameters of arabica coffee (Coffea arabica) seedlings produced in full sun (T1) and in shade (T2) were examined. The biomass accumulation and relative growth rate of T1 and T2 seedlings were similar. The T1 seedlings had less biomass allocation to shoots, a lower leaf mass ratio and a lower leaf area ratio; however, they had a greater net assimilation rate (rate of increase in plant mass per unit leaf area), which was associated with a greater net photosynthetic rate. There were no alterations in the concentrations of total chlorophylls or in the chlorophyll a/b ratio when comparing T1 and T2 seedlings. No indications of photoinhibition or photooxidative damage were observed in the T1 plants, which were shown to have a more robust antioxidant system than the T2 plants. Seedlings transferred from shade to full sun (T3) were not capable of utilising the incident extra light to fix CO₂. These seedlings showed a remarkable nocturnal retention of zeaxanthin and a significantly increased deepoxidation state of the xanthophyll cycle, even at predawn, but the activity of antioxidant enzymes was lower than in the T1 and T2 plants. Despite the acclimation capacity of T3 seedlings to the new light environment, they exhibited chronic photoinhibition and considerable photooxidative damage throughout the seven days following the transfer to full sun exposure. We further discuss the practical implications of producing coffee seedlings in full sunlight and under shade.     

Strong leaf morphological, anatomical, and physiological responses of a subtropical woody bamboo (Sinarundinaria nitida) to contrasting light environments

Dwarf bamboos are an important understory component of the lowland and montane forests in the subtropical regions of Asia and South America, yet little is known about their physiology and phenotypic plasticity in response to changing light environments. To understand how bamboo species adapt to different light intensities, we examined leaf morphological, anatomical, and physiological differentiation of Sinarundinaria nitida (Mitford) Nakai, a subtropical woody dwarf bamboo, growing in open and shaded natural habitats in the Ailao Mountains, SW China. Compared with leaves in open areas, leaves in shaded areas had higher values in leaf size, specific leaf area, leaf nitrogen, and chlorophyll concentrations per unit area but lower values in leaf thickness, vein density, stomatal density, leaf carbon concentration, and total soluble sugar concentration. However, stomatal size and leaf phosphorus concentration per unit mass remained relatively constant regardless of light regime. Leaves in the open habitat exhibited a higher light-saturated net photosynthetic rate, dark respiration rate, non-photochemical quenching, and electron transport rate than those in the shaded habitat. The results of this study revealed that the bamboo species exhibited a high plasticity of its leaf structural and functional traits in response to different irradiances. The combination of high plasticity in leaf morphological, anatomical, and physiological traits allows this bamboo species to grow in heterogeneous habitats.     

Physiological and biochemical abilities of robusta coffee leaves for acclimation to cope with temporal changes in light availability

The effects of varying intensities of light on plants depend on when they occur, even if the total amount of light received is kept constant. We designed an experiment using two clones of robusta coffee (Coffea canephora) intercropped with shelter trees in such a way that allowed us to compare coffee bushes shaded in the morning (SM) with those shaded in the afternoon (SA), and then confronting both with bushes receiving full sunlight over the course of the day (FS). The SM bushes displayed better gas‐exchange performance than their SA and FS counterparts, in which the capacity for CO₂ fixation was mainly constrained by stomatal (SA bushes) and biochemical (FS bushes) factors. Physiological traits associated with light capture were more responsive to temporal fluctuations of light rather than to the amount of light received, although this behavior could be a clone‐specific response. The activity of key antioxidant enzymes differed minimally when comparing the SM and SA clones, but was much larger in FS clones. No signs of photoinhibition or cell damage were found regardless of the light treatments. Acclimations to varying light supplies had no apparent additional cost for constructing and maintaining the leaves regardless of the light supply. Both the SM and SA individuals displayed higher return in terms of revenue streams (e.g. higher mass‐based light‐saturated photosynthetic rates, photosynthetic nitrogen use efficiencies and long‐term water use efficiencies) than their FS counterparts. In conclusion, shading may improve the physiological performance of coffee bushes growing in harsh, tropical environments.     

Functional plasticity of photosynthetic apparatus and its resistance to photoinhibition in <Emphasis Type="Italic">Plantago media</Emphasis>

Morphological and functional characteristics of Plantago media L. leaves were compared for plants growing at different light regimes on limestone outcrops in Southern Timan (62°45′N, 55°49′E). The plants grown in open areas under exposure to full sunlight had small leaves with low pigment content and high specific leaf weight; these leaves exhibited high photosynthetic capacity and elevated water use efficiency at high irradiance. The maximum photochemical activity of photosystem II (F _v/F _m) in leaves of sun plants remained at the level of about 0.8 throughout the day. The photosynthetic apparatus of sun plants was resistant to excess photosynthetically active radiation, mostly due to non-photochemical quenching of chlorophyll fluorescence (qN). This quenching was promoted by elevated deepoxiation of violaxanthin cycle pigments. Accumulation of zeaxanthin, a photoprotective pigment in sun plant leaves was observed already in the morning hours. The plant leaves grown in the shade of dense herbage were significantly larger than the sun leaves, with pigment content 1.5–2.0 times greater than in sun leaves; these leaves had low qN values and did not need extensive deepoxidation of violaxanthin cycle pigments. The data reveal the morphophysiological plasticity of plantain plants in relation to lighting regime. Environmental conditions can facilitate the formation of the ecotype with photosynthetic apparatus resistant to photoinhibition. Owing to this adjustment, hoary plantain plants are capable of surviving in ecotopes with high insolation.     

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.     

Seasonal changes in photosynthesis and nitrogen allocation in leaves of different ages in evergreen understory shrub <Emphasis Type="Italic">Daphniphyllum humile</Emphasis>

Seasonal changes in photosynthetic capacity, leaf nitrogen (N) content, leaf chlorophyll (Chl) content and leaf N allocation patterns in leaves of different ages in the evergreen understory shrub, Daphniphyllum humile Maxim, growing at a forest border and an understory site were studied. In current-year leaves at the understory site, the N and Rubisco contents increased from spring to autumn although their light-saturated photosynthetic rate at 22°C (P _max²²) remained stable, indicating that their mesophyll conductance rates declined as they completed their development and/or that they invested increasing amounts of their resources in photosynthetic enzymes during this period. In contrast, seasonal changes in P _max²² in current-year leaves at the forest border site were correlated with changes in Rubisco content. In 1-year old leaves at the understory site, P _max²² and contents of Chl, leaf N, and Rubisco remained stable from spring to autumn, while these parameters decreased in 1-year-old forest border leaves, indicating that N may have been remobilized from shaded 1-year-old leaves to sunlit current-year leaves. When leaves senesced at the forest border site the Rubisco content decreased more rapidly than that of light-harvesting proteins such as LHCII, suggesting that N remobilization from Rubisco may be more efficient, possibly because Rubisco has greater N costs and is soluble, whereas the light-harvesting proteins are membrane components.     

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.     

Preferential Feeding and Occupation of Sunlit Leaves Favors Defense Response and Development in the Flea Beetle,Altica brevicollis coryletorum – A Pest of Corylus avellana

The monophagous beetle, Altica brevicollis coryletorum, is a major leaf pest of Corylus avellana (common hazel). In contrast to majority of the other studied species of shrubs, sunlit leaves are grazed to a much greater extent than shaded leaves. Since the observation of a link between leaf irradiance level and A. brevicollis feeding is unique, we hypothesized that feeding preference of this beetle species is related to the speed needed to escape threats i.e. faster jumping. We also hypothesized that sunlit leaves are more nutritious and easier to consume than the leaves of shaded shrubs. Results indicated that beetle mass was greater in beetles occupying sunlit leaves, which is consistent with our second hypothesis. The study also confirmed under laboratory conditions, that larvae, pupae and beetles that were fed full-light (100% of full light) leaves were significantly heavier than those fed with shaded leaves (15% of full light). In the high irradiance conditions (higher temperature) duration of larval development is also reduced. Further results indicated that neither the concentration of soluble phenols, leaf toughness, or the number of trichomes could explain the insect’s preference for sunlit leaves. Notably, measurements of jump length of beetles of this species, both in the field and under laboratory conditions, indicated that the defense pattern related to jumping was associated with light conditions. The jump length of beetles in the sun was significantly higher than in the shade. Additionally, in laboratory tests, beetle defense (jumping) was more strongly affected by temperature (15, 25, or 35°C for 24h) than by leaf type. The effect of sunlit, higher nutrient leaves (greater level of non-structural carbohydrates) on defense (jumping) appears to be indirect, having a positive effect on insect mass in all developmental stages.     

Diurnal and Seasonal Changes in Photosynthetic Characteristics in Different Olive (Olea europaea L.) Cultivars

Diurnal and seasonal changes in photosynthetic characteristics, leaf area dry mass (ADM), and reducing sugar and total chlorophyll (Chl) contents of leaves of Frantoio, Leccino, and Maurino olive cultivars were investigated in Central Italy. Leaf net photosynthetic rate (P _N) per unit leaf area changed during the growing season and during the day, but the cultivar did not significantly influence the changes. In both young and one-year-old leaves the highest P _N values were observed in October, while the lowest values were recorded in August and December; during the day the highest P _N values were generally found in the morning. The pattern of photosynthetic response to photosynthetic photon flux density (PPFD) of leaves was similar in the three genotypes. Sub-stomatal CO₂ concentration (C _I) tended to increase when P _N decreased. The increase in C _I was accompanied by a stomatal conductance to water vapor (g _S) decrease. In general, P _N and dark respiration rate (R _D) were correlated. Transpiration rate (E), with no differences between the cultivars, increased from April to July, decreased greatly in August, then increased in October and finally decreased again in December. Leaf water content increased from April to June, remained high until mid July, decreased significantly in August, remaining constant until December with no differences associated with the cultivar. In both young and one-year-old leaves, the leaf water content per unit leaf area was slightly greater in Frantoio than in the other two cultivars. The one-year-old leaves had a higher Chl content than the young ones. The cultivar did not substantially influence the leaf reducing sugar content which decreased from April to August, when it reached the lowest level, then increased rapidly until October. During the day the reducing sugar content did not change significantly. The leaf ADM was slightly higher in Frantoio than in the other cultivars and one-year-old leaves had higher values than the young ones. Leaf ADM decreased from April to June and then tended to increase until December. During the day there were no substantial variations.     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

<Emphasis Type="Italic">Lantana camara</Emphasis> L.: a weed with great light-acclimation capacity

Plant invasions may be limited by low radiation levels in ecosystems such as forests. Lantana camara has been classified among the world’s 10 worst weeds since it is invading many different habitats all around the planet. Morphological and physiological responses to different light fluxes were analyzed. L. camara was able to acclimate to moderately shaded environments, showing a high phenotypic plasticity. Morphological acclimation to low light fluxes was typified by increasing leaf size, leaf biomass, leaf area index and plant height and by reduced stomatal density and leaf thickness. Plants in full sunlight produced many more inflorescences than in shaded conditions. Physiological acclimation to low radiation levels was shown to be higher stomatal conductance, higher net photosynthetic rates and higher efficiency of photosystem II (PSII). L. camara behaves as a facultative shade-tolerant plant, being able to grow in moderately sheltered environments, however its invasion could be limited in very shady habitats. Control efforts in patchy environments should be mainly directed against individuals in open areas since that is where the production of seeds would be higher and the progress of the invasion would be faster.     

Photoinhibition of holly (Ilex aquifolium) in the field during the winter

The distribution of holly ( Ilex aquifolium ) and its habitat preferences indicate a sensitivity to low temperature, particularly when exposed to high light. Experiments were conducted to determine whether photoinhibition of photosynthesis occurs in holly leaves in the field in United Kingdom during the winter. Photosynthetic efficiency was assessed in holly leaves that were exposed to or shaded from direct sunlight using measurements of photosynthetic oxygen evolution and chlorophyll fluorescence. Field measurements were conducted over 3 weeks during January and February. Correlation of the measurements of photosynthetic efficiency with weather conditions indicated that holly was suffering photoinhibition, particularly in leaves exposed to direct sunlight. Controlled environment studies demonstrated that exposure of leaves to low temperature and high light resulted in reductions in photosynthetic efficiency; however, leaves recovered rapidly when exposed to a higher temperature and reduced light level.     

Photosynthetic responses to short-term and long-term light variation in <Emphasis Type="Italic">Pinus sylvestris</Emphasis> and <Emphasis Type="Italic">Salix dasyclados</Emphasis>

Pinus sylvestris and Salix dasyclados, which differ in leaf longevity, were compared with respect to four aspects of photosynthetic light use and response: high light acclimation, photoinhibition resistance and recovery, lightfleck exposure and use and chloroplast acclimation across leaves. The first two aspects were examined using seedlings under controlled conditions and the other two were tested using trees in the field. When exposed to high light, shade leaves of Pinus acclimated completely, achieving the same photosynthetic capacities as sun leaves, whereas shade leaves of Salix did not reach sun leaf capacities although the absolute magnitude of their acclimation was larger. Shade leaves of Pinus were also more resistant to photoinhibition than those of Salix. Much of the direct light supplied within the canopy was in the form of rapid fluctuations, lightflecks, for Pinus and Salix alike. They exploited short lightflecks with similar efficiency. The greater proportion of diffuse light in the canopy for Pinus than Salix seems to lead to a lesser degree of differential intra-leaf acclimation of chloroplasts, in turn leading to lower efficiency of photosynthesis under unilateral light as reflected by a lower convexity, rate of bending, of the light–response curve. The differences in light use and responses are discussed in relation to possible differences in characteristics of the long and short-lived leaf.     

Paraheliotropic leaf movement in Siratro as a protective mechanism against drought-induced damage to primary photosynthetic reactions: damage by excessive light and heat

Damage to primary photosynthetic reactions by drought, excess light and heat in leaves of Macroptilium atropurpureum Dc. cv. Siratro was assessed by measurements of chlorophyll fluorescence emission kinetics at 77 K (-196°C). Paraheliotropic leaf movement protected waterstressed Siratro leaves from damage by excess light (photoinhibition), by heat, and by the interactive effects of excess light and high leaf temperatures. When the leaves were restrained to a horizontal position, photoinhibition occurred and the degree of photoinhibitory damage increased with the time of exposure to high levels of solar radiation. Severe inhibition was followed by leaf death, but leaves gradually recovered from moderate damage. This drought-induced photoinhibitory damage seemed more closely related to low leaf water potential than to low leaf conductance. Exposure to leaf temperatures above 42°C caused damage to the photosynthetic system even in the dark and leaves died at 48°C. Between 42 and 48°C the degree of heat damage increased with the time of exposure, but recovery from moderate heat damage occurred over several days. The threshold temperature for direct heat damage increased with the growth temperature regime, but was unaffected by water-stress history or by current leaf water status. No direct heat damage occurred below 42°C, but in water-stressed plants photoinhibition increased with increasing leaf temperature in the range 31–42°C and with increasing photon flux density up to full sunglight values. Thus, water stress evidently predisposes the photosynthetic system to photoinhibition and high leaf temperature exacerbates this photoinhibitory damage. It seems probable that, under the climatic conditions where Siratro occurs in nature, but in the absence of paraheliotropic leaf movement, photoinhibitory damage would occur more frequently during drought than would direct heat damage.Abbreviations and symbols PFD photon flux area density - PSI, PSII photosyntem I, II - F _M, F _O, F _V maximum, instantaneous, variable fluorescence emission - PLM paraheliotropic leaf movement; all data of parameter of variation are mean ± standard error