Seasonal acclimation of photosystem II in Pinus sylvestris. I. Estimating the rate constants of sustained thermal energy dissipation and photochemistry

Acclimation of the partitioning of absorbed light energy in Photosystem II (PSII) between photochemical and non-photochemical processes includes short-term adjustments that are rapidly reversed in the dark and seasonal acclimation processes that are unaffected by dark acclimation. Thus, by using dark-acclimated leaves to study the seasonal acclimation of PSII, the confounding effect of short-term adjustments is eliminated. The maximum quantum yield of photochemistry, estimated by chlorophyll fluorescence analysis as F(v)/F(m), where F(v) = (F(m) - F(o)), and F(m) and F(o) are maximum and minimum chlorophyll fluorescence, respectively, has been widely used to follow the seasonal acclimation of PSII, because it is measured in dark-acclimated leaves. Seasonal changes in F(v)/F(m) can be caused by adjustments in either the photochemical capacity in PSII, or the capacity of thermal dissipation in PSII, or both. However, there is a lack of chlorophyll fluorescence parameters that can distinguish between these processes. In this study, we introduce two new parameters: the rate constants of sustained thermal energy dissipation (k(NPQ)) and of photochemistry (k(P)). We estimated k(NPQ) and k(P) from dark-acclimated F(o) and F(m) measured during spring recovery of photosynthesis in Scots pine (Pinus sylvestris L.) trees. We suggest that k(NPQ) and k(P) be used to study the mechanisms underlying the observed seasonal acclimation in PSII, because these parameters provide quantitative data that complement and extend F(v)/F(m) measurements.     

Leaf physiological versus morphological acclimation to high-light exposure at different stages of foliar development in oak

We investigated light acclimation in seedlings of the temperate oak Quercus petraea (Matt.) Liebl. and the co-occurring sub-Mediterranean oak Quercus pyrenaica Willd. Seedlings were raised in a greenhouse for 1 year in either 70 (HL) or 5.3% (LL) of ambient irradiance of full sunlight, and, in the following year, subsets of the LL-grown seedlings were transferred to HL either before leaf flushing (LL-HLBF plants) or after full leaf expansion (LL-HLAF plants). Gas exchange, chlorophyll a fluorescence, nitrogen fractions in photosynthetic components and leaf anatomy were examined in leaves of all seedlings 5 months after plants were moved from LL to HL. Differences between species in the acclimation of LL-grown plants to HL were minor. For LL-grown plants in HL, area-based photosynthetic capacity, maximum rate of carboxylation, maximum rate of electron transport and the effective photochemical quantum yield of photosystem II were comparable to those for plants grown solely in HL. A rapid change in nitrogen distribution among photosynthetic components was observed in LL-HLAF plants, which had the highest photosynthetic nitrogen-use efficiency. Increases in mesophyll thickness and dry mass per unit area governed leaf acclimation in LL-HLBF plants, which tended to have less nitrogen in photosynthetic components and a lower assimilation potential per unit of leaf mass or nitrogen than LL-HLAF plants. The data indicate that the phenological state of seedlings modified the acclimatory response of leaf attributes to increased irradiance. Morphological adaptation of leaves of LL-HLBF plants enhanced photosynthetic capacity per unit leaf area, but not per unit leaf dry mass, whereas substantial redistribution of nitrogen among photosynthetic components in leaves of LL-HLAF plants enhanced both mass- and area-based photosynthetic capacity.     

Photosynthetic acclimation to light changes in tropical monsoon forest woody species differing in adult stature

We studied morphological and physiological leaf and whole-plant features of seedlings of six late-successional woody species common in the Xishuangbanna lowland rain forest in southwest China. Study species differed in adult stature and shade tolerance and included the shrubs Lasianthus attenuatus Jack and Lasianthus hookeri C.B. Clarke ex Hook. f.; the sub-canopy species Barringtonia macrostachya (Jack) Kurz and Linociera insignis C.B. Clarke; the canopy tree Pometia tomentosa (Blume) Teijsm. & Binn.; and the emergent species Shorea chinensis (Wang Hsie) H. Zhu. After 1 year of growth in low light (4.5% full sun), seedlings were transferred to high light (24.5% full sun) to investigate acclimation responses of existing leaves to forest gap opening and to determine whether seedling capacity for acclimation is a limiting factor in its natural regeneration. Leaves of the shrub species are shade-adapted, as indicated by their low photosynthetic capacity, efficiency in using sunflecks, low stomatal density, low Chl a/b ratio and high spongy/palisade mesophyll ratio. The shrub species utilized sunflecks efficiently because they had a short photosynthetic induction time and low induction loss. In all species, transfer of seedlings to high light resulted in a substantial initial reduction in the dark-adapted quantum yield of photosystem II (variable chlorophyll fluorescence/maximum chlorophyll fluorescence; Fv/Fm) at midday. Predawn Fv/Fm of the taller species did not change greatly, but predawn Fv/Fm of the shrub species decreased significantly without complete recovery within 25 days of transfer to high light, indicating chronic photoinhibition and damage to the previously shade-adapted leaves. Maximum net photosynthetic rate and dark respiration of the four taller species increased considerably after transfer to high light, but not in the shrub species. Similar trends were observed for the number of newly formed leaves and relative height growth rate. We conclude that the shrubs L. hookeri and L. attenuatus have limited potential for developmental and physiological acclimation to high light, which explains their absence from forest gaps. Compared with the shrub species, the taller tree species, which are more likely to experience high light during their life span, showed a greater potential for light acclimation. Physiological differences among the four tree species were not consistent with differences in adult stature.     

Increase of nitrogen to promote growth of poplar seedlings and enhance photosynthesis under NaCl stress

The solution culture method was used to study the effect of increasing nitrogen on the growth and pho-tosynthesis of poplar seedlings under 100 mmol L-1 NaCl stress. I Increase in nitrogen reduced stomatal limitation of leaves under NaCl stress, improved utilization of CO2 by mesophyll cells, enhanced photosynthetic carbon assimi-lation capacity, significantly alleviated saline damage of NaCl, and promoted the accumulation of aboveground and root biomass. I Increased nitrogen enhanced photochemical efficiency (ΦPSⅡ) and electron transport rates, relieved the reduction of maximum photochemical efficiency (Fv/Fm) under NaCl, and reduced the degree of photoinhibition caused by NaCl stress. Increased nitrogen applications reduced the proportion of energy dissipating in the form of ineffective heat energy and hence a greater proportion of light energy absorbed by leaves was allocated to photo-chemical reactions. Under treatment with increased nitro-gen, the synergistic effect of heat dissipation and the xanthophyll cycle in the leaves effectively protected pho-tosynthetic PSⅡ and enhanced light energy utilization of leaves under NaCl stress. The increased nitrogen promoted photosynthetic electron supply and transport ability under NaCl stress evident in enhanced functioning of the oxygen-evolving complex on the electron donor side of PS Ⅱ. It increased the ability of the receptor pool to accept electrons on the PSII electron acceptor side and improved the sta-bility of thylakoid membranes under NaCl stress. Therefore, increasing nitrogen applications under NaCl stress can promote poplar growth by improving the effi-ciency of light energy utilization.     

Light acclimation potential and carry-over effects vary among three evergreen tree species with contrasting patterns of leaf emergence and maturation

We compared light acclimation potential among three evergreen broadleaved species with contrasting patterns of shoot elongation, leaf emergence and leaf maturation. Understory saplings were transferred to a high-light environment before bud break, grown for 13 months, and then transferred back to the understory to observe subsequent carry-over effects. Acclimation potential was highest and sapling mortality was lowest for Cinnamomum japonicum Sieb. ex Nakai. Indeterminate growth and successive leaf emergence allowed this species to acclimate to both high and low light by adjusting leaf production as well as leaf properties. Sapling mortality occurred after both transfers for Camellia japonica L., which also has indeterminate growth and successive leaf emergence. In this species, carry-over effects were observed at the individual level, but leaf-level acclimation potential was high. Acclimation potential was lowest and sapling mortality occurred soon after the transfer to high light for Quercus glauca Thunb. ex Murray. Determinate growth and flush-type leaf emergence resulted in significant carry-over effects in this species. Indeterminate growth and successive leaf emergence increase whole-plant acclimation potential by extending the period of growth and architectural development during the growing season. Similarly, we inferred that delayed leaf maturation, observed in many evergreen species, increases the acclimation potential of current-year leaves by extending the period of leaf development. In evergreen species, the acclimation potential of preexisting leaves determines the role that leaf turnover plays in whole-plant light acclimation, resulting in diverse strategies for light acclimation among species, as observed in this study.     

Changes during early development in photosynthetic light acclimation capacity explain the shade to sun transition in Nothofagus nitida

Nothofagus nitida (Phil.) Krasser, an emergent tree of the Chilean evergreen forest, regenerates under the canopy. Nonetheless, it is common to find older saplings in clear areas. We hypothesized that this transition from shade to sun during the early developmental stages is made possible by an ontogenetic increase in the light acclimation capacity of photosynthesis. To test our hypothesis, we studied photosynthetic performance and photoprotection in N. nitida saplings at different developmental stages corresponding with three different height classes (short: 16.2 cm; medium-height: 48.0 cm; and tall: 73.7 cm) grown under contrasting light conditions (photosynthetic photon flux (PPF) of 20, 300 or 600 micromol m(-2) s(-1)) until newly expanded leaves had developed. Light-saturated CO(2) assimilation rate and light compensation and saturation points increased with increasing PPF. Medium-height and tall saplings acclimated to high light had higher electron transport rates and higher proportions of open Photosystem II reaction centers than shorter plants acclimated to high light. Short saplings showed higher thermal dissipation and contents of xanthophylls than taller saplings. Only medium-height and tall saplings acclimated to high light recovered after photoinhibition. State transitions were higher in short plants growing in low light, and decreased with plant height and growth irradiance. Thus, during development, N. nitida changes the balance of light energy utilization and photoprotective mechanisms, supporting a phenotypic transition from shade to sun during its early ontogeny.     

Acclimation of shade-developed leaves on saplings exposed to late-season canopy gaps

We hypothesized that photoinhibition of shade-developed leaves of deciduous hardwood saplings would limit their ability to acclimate photosynthetically to increased irradiance, and we predicted that shade-tolerant sugar maple (Acer saccharum Marsh.) would be more susceptible to photoinhibition than intermediately shade-tolerant red oak (Quercus rubra L.). After four weeks in a canopy gap, photosynthetic rates of shade-developed leaves of both species had increased in response to the increase in irradiance, although final acclimation was more complete in red oak. However, photoinhibition occurred in both species, as indicated by short-term reductions in maximum rates of net photosynthesis and the quantum yield of oxygen evolution, and longer-term reductions in the efficiency of excitation energy capture by open photosystem II (PSII) reaction centers (dark-adapted F(v)/F(m)) and the quantum yield of PSII in the light (phi(PSII)). The magnitude and duration of this decrease were greater in sugar maple than in red oak, suggesting greater susceptibility to photoinhibition in sugar maple. Photoinhibition may have resulted from photodamage, but it may also have involved sustained rates of photoprotective energy dissipation (especially in red oak). Photosynthetic acclimation also appeared to be linked to an ability to increase leaf nitrogen content. Limited photosynthetic acclimation in shade-developed sugar maple leaves may reflect a trade-off between shade-tolerance and rapid acclimation to a canopy gap.     

Interactive effects of leaf age and self-shading on leaf structure,photosynthetic capacity and chlorophyll fluorescence in the rain forest tree,Dryobalanops aromatica

In the tropical canopy tree, Dryobalanops aromatica Gaertn. f., upper-canopy leaves (UL) develop under sunlit conditions but are subjected to self-shading within the crown as they age. In contrast, lower-canopy leaves (LL) are exposed to uniform dim light conditions throughout their life span. By comparing leaf morphology and physiology of UL and LL, variations in leaf characteristics were related to leaf age and self-shading. Mass-based chlorophyll (chl) concentration and the chlorophyll/nitrogen (chl/N) ratio were lower and the chl a/b ratio was higher in UL than in LL. In UL, the chl/N ratio gradually increased and the chl a/b ratio gradually decreased with leaf aging, whereas these ratios remained unchanged with leaf age in LL. The effective quantum yield of photosystem II (PSII) (DeltaF/F(m)') at a given irradiance remained unchanged with leaf age in LL, whereas DeltaF/F(m)' changed with leaf age in UL. These data indicate N reallocation within the leaves from carbon fixation components to light harvesting components and a dynamic regulation of photochemical processes of PSII in response to increased self-shading of UL. Despite the difference in light environment with leaf age between UL and LL, maximum photosynthetic rates and nitrogen-use efficiency decreased with leaf aging in both UL and LL. Constancy in the chl/N ratio with leaf age in LL indicated that the decrease in photosynthetic capacity was caused by effects other than shading, such as leaf aging. We conclude that N reallocation and acclimation of PSII to self-shading occurred even in mature leaves, whereas the change in photosynthetic capacity with leaf age was more conservative.     

Seasonal acclimation of photosystem II in Pinus sylvestris. II. Using the rate constants of sustained thermal energy dissipation and photochemistry to study the effect of the light environment

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.     

Seasonality of photosynthetic parameters in a multi-specific and vertically complex forest ecosystem in the Sierra Nevada of California

Understanding seasonal variations of photosynthetic parameters is critical for accurate modeling of carbon dioxide (CO2) uptake by ecosystems. Maximum carboxylation velocity (Vcmax), maximum rate of electron transport (Jmax), leaf respiration in the light (R(day)), light-saturated assimilation (Amax) and maximum quantum yield (Phi) were calculated from leaf gas exchange measurements made monthly throughout the year on leaves of three co-occuring evergreen species in a Pinus ponderosa Dougl. ex P. Laws. & C. Laws. forest with shrubs in the understory (Arctostaphylos manzanita Parry and Ceanothus cordulatus Kellogg.). The seasonality and relationships of the photosynthetic parameters with environmental and physiological variables differed among the species. The nitrogen-fixing species, C. cordulatus had the highest values of the parameters and the largest seasonal variation, whereas A. manzanita exhibited the lowest seasonality and weaker correlations with environmental variables. In general, variations in Vcmax were highly correlated with light, leaf mass per area and leaf nitrogen content on an area basis. Temporal scaling of the parameters with each other seemed possible for C. cordulatus and P. ponderosa. However, lags between these variables and Vcmax likely reflect the influences of other factors. The acclimation relationships found along vertical light gradients within canopies in other studies cannot be applied to seasonal variations. The Jmax to Vcmax ratio varied seasonally for P. ponderosa and A. manzanita, being lower at high light, high air temperature and low soil water content.     

Tradeoff between shade adaptation and mitigation of photoinhibition in leaves of Quercus mongolica and Acer mono acclimated to deep shade

We investigated susceptibility to photoinhibition in leaves acclimated to different light regimes in intermediately shade-tolerant Japanese oak (Quercus mongolica Fisch. ex Turcz. var. crispula (Blume) Ohashi) and shade-tolerant Japanese maple (Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara), to elucidate adaptability to gap formation in leaves differing in shade acclimation. We hypothesized that there is a tradeoff between shade adaptation and capacity to mitigate photoinhibition associated with leaf morphology. We simultaneously measured chlorophyll fluorescence and gas exchange in seedlings that had been grown in full sunlight (open), 10% of full sun (moderate shade) and 5% of full sun (deep shade). Shade-tolerant A. mono adapted to deep shade through changes in leaf morphology, lowering its leaf mass per area (LMA), but Q. mongolica showed little change in LMA between moderate and deep shade. Photochemical quenching (qP) did not differ between species in full sunlight and moderate shade; however, in deep shade, qP of Q. mongolica was higher than that of A. mono, suggesting that Q. mongolica grown in deep shade is less susceptible to photoinhibition at gap formation. This is consistent with the finding that chronic photoinhibition 3 days after the transfer to full sunlight, indicated by the decrease in maximum photochemical efficiency, Fv/Fm, at predawn, was less in deep-shade-grown Q. mongolica than in deep shade-grown A. mono. In deep shade, the electron transport rate (ETR) of Q. mongolica was higher than that of A. mono, whereas thermal energy dissipation through photosystem II antennae, indicated by non-photochemical quenching, was lower in Q. mongolica than in A. mono. In deep shade, the greater ETR capacity in Q. mongolica in association with higher LMA and higher leaf N content could contribute to maintaining high qP and mitigating photoinhibition. These results indicate that, by maintaining a high electron transport capacity even in deep shade, the gap-dependent and intermediate-shade-tolerant Q. mongolica trades improved shade adaptation for higher growth potential when a gap event occurs.     

一品红组培苗移栽期叶片生理与解剖变化

一品红(Euphorbia pulcherrima),别名象牙红、老来娇、圣诞花、猩猩木,为大戟科大戟属的重要花卉,世界各地广为栽培,为圣诞节、元旦、春节的优良装饰花卉。目前我国市场的矮化一品红主要由进口种苗培育而成,国内利用组织培养技术培育矮化一品红种苗已有不少成功的报道(钟士传等,1999;蒋小满等,2002),但一品红试管苗移栽时叶片较易萎蔫、成活率较低,为其种苗生产带来了困难。本试验研究一品红试管苗在移栽驯化过程中叶片蒸腾、光合特性及解剖结构的变化,探索提高其移栽成活率的方法,以便为矮化一品红种苗产业化提供技术支持。1材料与方法1·…     

Acclimation of tropical tree species to hurricane disturbance: ontogenetic differences

We investigated acclimation responses of seedlings and saplings of the pioneer species Cecropia schreberiana Miq. and three non-pioneer species, Dacryodes excelsa Vahl, Prestoea acuminata (Willdenow) H.E. Moore var. montana (Graham) Henderson and Galeano, and Sloanea berteriana Choisy ex DC, following a hurricane disturbance in a lower montane wet forest in Puerto Rico. Measurements were made, shortly after passage of the hurricane, on leaves expanded before the hurricane (pre-hurricane leaves) and, at a later time, on recently matured leaves that developed after the hurricane (post-hurricane leaves) from both seedlings and saplings at sites that were severely damaged by the hurricane (disturbed sites) and at sites with little disturbance (undisturbed sites). Pre-hurricane leaves of the non-pioneer species had relatively low light-saturated photosynthetic rates (A(max)) and stomatal conductance (g(s)); neither A(max) nor g(s) responded greatly to the increase in irradiance that resulted from the disturbance, and there were few significant differences between seedlings and saplings. Pre-hurricane leaves of plants at undisturbed sites had low dark respiration rates per unit area (R(d)) and light compensation points (LCP), whereas pre-hurricane leaves of plants at disturbed sites had significantly higher R(d) and LCP. Post-hurricane leaves of plants at disturbed sites had significantly higher A(max) and R(d) than plants at undisturbed sites. Compared with seedlings, saplings had higher A(max) and R(d) and showed greater acclimation to the increase in irradiance that followed the disturbance. Post-hurricane leaves of the non-pioneer species had significantly lower A(max) and were less responsive to changes in irradiance than the pioneer species C. schreberiana. Variation in A(max) across light environments and stages was strongly related to differences in leaf mass per unit area (LMA), especially in the non-pioneer species. As indicated by V(cmax) or J(max) per unit nitrogen, light acclimation of A(max) was determined by leaf morphology (LMA) for the non-pioneer species and by both leaf morphology and leaf biochemistry for C. schreberiana. Ontogenetic changes in A(max) were attributable to changes in leaf morphology. The ontogenetic component of variation in A(max) across light environments and stages differed among species, ranging from 36 to 59% for the non-pioneer species (D. excelsa, 59.3%; P. acuminata var. montana, 44.7%; and S. berteriana, 36.3%) compared with only 17% in the pioneer species C. schreberiana.     

Photosynthetic light acclimation in peach leaves: importance of changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning

Photosynthetic light acclimation of leaves can result from (i) changes in mass-based leaf nitrogen concentration, Nm, (ii) changes in leaf mass:area ratio, Ma, and (iii) partitioning of total leaf nitrogen among different pools of the photosynthetic machinery. We studied variations in Nm and Ma within the crowns of two peach (Prunus persica L. Batsch) trees grown in an orchard in Portugal, and one peach tree grown in an orchard in France. Each crown was digitized and a 3-D radiation transfer model was used to quantify the intra-crown variations in time-integrated leaf irradiance, . Nitrogen concentration, leaf mass:area ratio, chlorophyll concentration, and photosynthetic capacity were also measured on leaves sampled on five additional peach trees in the orchard in Portugal. The data were used to compute the coefficients of leaf nitrogen partitioning among carboxylation, bioenergetics, and light harvesting pools. Leaf mass:area ratio and area-based leaf nitrogen concentration, Na, were nonlinearly related to , and photosynthetic capacity was linearly related to Na. Photosynthetic light acclimation resulted mainly from changes in Ma and leaf nitrogen partitioning, and to a lesser extent from changes in Nm. This behavior contrasts with photosynthetic light acclimation observed in other tree species like walnut (Juglans regia L.) in which acclimation results primarily from changes in Ma.     

A biochemical model of photosynthesis for mango leaves: evidence for the effect of fruit on photosynthetic capacity of nearby leaves

Variations in leaf nitrogen concentration per unit mass (Nm) and per unit area (Na), mass-to-area ratio (Ma), total nonstructural carbohydrates (Ta), and photosynthetic capacity (maximum carboxylation rate, electron transport capacity, rate of phosphate release in triose phosphate utilization and dark respiration rate) were studied within the digitized crowns of two 3-year-old mango trees (Mangifera indica L.) on La Réunion Island. Additional measurements of Nm, Na, Ma, Ta and photosynthetic capacities were performed on young, fully expanded leaves of 11-year-old mango trees. Leaves of similar gap fractions were taken far from and close to developing fruits. Unlike Nm, both Na and Ta were linearly correlated to gap fraction. Similar relationships were found for all leaves whatever their age and origin, except for Ta, for which we found a significant tree effect. Photosynthetic capacity was nonlinearly correlated to Na, and a unique relationship was obtained for all types of leaves. Photosynthetic acclimation to light was mainly driven by changes in Ma, but allocation of total leaf N between the different photosynthetic functions also played a substantial role in acclimation to the lowest irradiances. Leaves close to developing fruits exhibited a higher photosynthetic capacity than other leaves, but similar Ta. Our data suggest that Ta does not control photosynthetic capacity in mango leaves. We used the data to parameterize a biochemically based model of photosynthesis and an empirical stomatal conductance model, allowing accurate predictions of net photosynthesis of leaves in field-grown mango trees.     

Tree size and light availability increase photochemical instead of non-photochemical capacities of Nothofagus nitida trees growing in an evergreen temperate rain forest

Nothofagus nitida (Phil.) Krasser (Nothofagaceae) regenerates under the canopy in microsites protected from high light. Nonetheless, it is common to find older saplings in clear areas and adults as emergent trees of the Chilean evergreen forest. We hypothesized that this shade to sun transition in N. nitida is supported by an increase in photochemical and non-photochemical energy dissipation capacities of both photosystems in parallel with the increase in plant size and light availability. To dissect the relative contribution of light environment and plant developmental stage to these physiological responses, the photosynthetic performance of both photosystems was studied from the morpho-anatomical to the biochemical level in current-year leaves of N. nitida plants of different heights (ranging from 0.1 to 7 m) growing under contrasting light environments (integrated quantum flux (IQF) 5-40 mol m(-2). Tree height (TH) and light environment (IQF) independently increased the saturated electron transport rates of both photosystems, as well as leaf and palisade thickness, but non-photochemical energy flux, photoinhibition susceptibility, state transition capacity, and the contents of D1 and PsbS proteins were not affected by IQF and TH. Spongy mesophyll thickness and palisade cell diameter decreased with IQF and TH. A(max), light compensation and saturation points, Rubisco and nitrogen content (area basis) only increased with light environment (IQF), whereas dark respiration (R(d)) decreased slightly and relative chlorophyll content was higher in taller trees. Overall, the independent effects of more illuminated environment and tree height mainly increased the photochemical instead of the non-photochemical energy flux. Regardless of the photochemical increase with TH, carbon assimilation only significantly improved with higher IQF. Therefore it seems that mainly acclimation to the light environment supports the phenotypic transition of N. nitida from shade to sun.     

Nitrogen allocation and the fate of absorbed light in 21-year-old Pinus radiata

We investigated effects of nitrogen (N) fertilizer and canopy position on the allocation of N to Rubisco and chlorophyll as well as the distribution of absorbed light among thermal energy dissipation, photochemistry, net CO2 assimilation and alternative electron sinks such as the Mehler reaction and photorespiration. The relative reduction state of the primary quinone receptor of photosystem II (QA) was used as a surrogate for photosystem II (PSII) vulnerability to photoinactivation. Measurements were made on needles from the lower, mid and upper canopy of 21-year-old Pinus radiata D. Don trees grown with (N+) and without (N0) added N fertilizer. Rubisco was 45 to 60% higher in needles of N+ trees than in needles of N0 trees at all canopy positions. Chlorophyll was approximately 80% higher in lower- and mid-canopy needles of N+ trees than of N0 trees, but only approximately 20% higher in upper-canopy needles. Physiological differences between N+ and N0 trees were found only in the lower- and mid- canopy positions. Needles of N+ trees dissipated up to 30% less light energy as heat than needles of N0 trees and had correspondingly more reduced QA. Net CO2 assimilation and the proportions of electrons used by alternative electron sinks such as the Mehler reaction and photorespiration were unaffected by N treatment regardless of canopy position. We conclude that the application of N fertilizer mainly affected the biochemistry and light-use physiology in lower- and mid-canopy needles by increasing the amount of chlorophyll and hence the amount of light harvested. This, however, did not improve photochemistry or safe dissipation, but increased PSII vulnerability to photoinactivation, an effect with likely significant consequences during sunflecks or sudden gap formation.     

Acclimation of Pacific yew (Taxus brevifolia) foliage to sun and shade

The success in clinical trials of the anti-cancer drug, Taxol(R), obtained from the bark of Pacific yew (Taxus brevifolia Nutt.), has raised interest in cultivation and regeneration of this little-known species. Pacific yew is shade-tolerant and it is not known whether the foliage can tolerate the high solar irradiances found on an open forest regeneration site or a nursery. Acclimation of Pacific yew to sun and shade was studied by comparing foliar physiology and morphology of male and female trees growing in full sun or shade. Interspecific foliar acclimation to sun was studied by comparing sun-grown English yew (Taxus baccata L.) with Pacific yew. No sex-specific acclimation was found in foliar physiology or morphology in either species. Sun-grown foliage of Pacific yew and English yew differed with respect to light harvesting, transpiration, stomatal conductance, leaf structure, stomatal distribution and foliar N concentrations and contents. Chlorophyll a fluorescence measurements indicated that shade-grown foliage of Pacific yew had larger and more efficient light harvesting systems than sun-grown foliage. Rates of CO(2) uptake and transpiration were similar in sun- and shade-grown foliage indicating acclimation of photosynthesis to the growth irradiance. Specific leaf area was significantly higher in shade-grown foliage of Pacific yew than in sun-grown foliage and was diagnostic of the light environment in which the foliage grew. Foliar N concentrations were not significantly different between sun- and shade-grown leaves of Pacific yew but sun-grown foliage had a higher N content. Physiological and morphological adjustments of Pacific yew foliage conferred tolerance to both high light and shade, enabling the trees to survive in a variety of light environments and indicating that Pacific yew is suited to nursery cultivation and regeneration of open sites.     

Effects of severe dehydration on leaf photosynthesis in Quercus petraea (Matt.) Liebl.: photosystem II efficiency, photochemical and nonphotochemical fluorescence quenching and electrolyte leakage

Leaf disks of oak (Quercus petraea (Matt.) Liebl.) trees were subjected to rapid dehydration in air in the dark. Optimal photochemical efficiency of PS II (F(v)/F(M)), photochemical (q(P)) and nonphotochemical (q(NP)) quenchings of chlorophyll a fluorescence, and relative conductivity (C(r)) of leaf disk diffusate were measured in leaf disks with different water deficits (D). No effect of dehydration was detected before D reached values above 0.30. When D increased from 0.30 to 0.50, q(NP) increased without any change in q(P), which may indicate that thermal deexcitation of PS II increased, allowing reduced photochemical activity and maintenance of a large pool of oxidized primary acceptors (QA), although carbon reduction was impaired. Large changes in electron transport chain activity, leading to decreases in both q(P) and q(NP), appeared only in leaf disks subjected to severe water deficits (D > 0.60) and were correlated with a modification of membrane structure. However, stability of F(v)/F(M) indicated that the functional integrity of PS II was not altered until D reached values above 0.75. We conclude that the photosynthetic apparatus of Q. petraea is rather insensitive to leaf dehydration per se during drought under natural conditions.     

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.