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.     

Light-use properties in two sun-adapted shrubs with contrasting canopy structures

We investigated the impact of high solar irradiance and elevated temperature on carbon gain by two, co-occurring, sun-adapted, dwarf shrub species, Planchonella obovata var. dubia (Koidz.) Hatusima and Hibiscus glaber Matsumura, growing on sun-exposed ridges in the Bonin Islands, in the subtropical Pacific Ocean. Planchonella had steeply inclined, longer lived, sclerophyllous leaves, whereas Hibiscus has thinner, more horizontally oriented, and shorter lived leaves. We tested the hypothesis that leaf physiological tolerance to high light is lower in Planchonella than in Hibiscus. Under relatively high irradiances (photosynthetic photon flux density, PPFD, > 500 micromol m(-2) s(-1)), net photosynthetic rate (P(n)) was about 8.0 and 0.4 micromol m(-2) s(-1) in mature and young leaves of Planchonella, and about 12.4 and 10.3 micromol m(-2) s(-1) in mature and young leaves of Hibiscus, respectively. Both P(n) and photosystem II (PSII) quantum yield at a given PPFD were lower in Planchonella than in Hibiscus, whereas non-photochemical quenching (NPQ) at a given PPFD was higher in Planchonella. When leaf discs were exposed to high light (1900 micromol m(-2) s(-1) PPFD) at 37, 40 or 43 degrees C for 3 h, the recovery of PSII quantum yield (F(v)/F(m)) in the following 60-min dark period was slower in Planchonella than in Hibiscus, indicating that the ability of PSII to tolerate high light and high temperature was less in Planchonella than in Hibiscus. We postulate that there is a linkage between leaf display and leaf photochemical ability in sun-adapted shrub species.     

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.     

Seasonal variation in photosystem II efficiency and photochemical reflectance index of evergreen trees and perennial grasses growing at low and high elevations in subtropical Taiwan

Three pines species, three evergreen broadleaf trees, one C(3) and two C(4) perennial grasses of subtropical Taiwan were studied to elucidate the correlation between photosystem II (PSII) efficiency and photochemical reflectance index (PRI = (R(531) - R(570))/(R(531) + R(570))). Measurements were made at two sites differing in altitude (800 and 2600 m) over several growing seasons. At high elevation, potential PSII efficiency, measured by chlorophyll fluorescence (the ratio of variable to maximal fluorescence; F(v)/F(m)) at predawn, decreased with decreasing air temperature and varied greatly among species. At the lowest air temperature (-3 degrees C) studied, variation in F(v)/F(m) among species ranged from 0.33 to 0.72. In contrast, at low elevation where air temperature was moderate, seasonal variation in F(v)/F(m) was small in all of the study species. When species, elevation and season data were pooled, despite the high variation in F(v)/F(m) among species, a good correlation between F(v)/F(m) and PRI was observed. When compared at the same value of PRI, F(v)/F(m) of evergreen trees was higher than that of perennial grasses; however, when the minimum temperature on the measurement day was below 0 degrees C, F(v)/F(m) was underestimated relative to PRI. We conclude that PRI could be used as a remote indicator of photosynthetic function when air temperature is above 0 degrees C.     

Variation in stand mortality related to successional composition

Forest ecologists have long recognized that there are large differences in the vital rates of early- and late-successional species: early-successional species typically grow faster than late-successional species, while late-successional species typically have lower rates of mortality than early-successional species. Numerous studies have shown that the differences in mortality are particularly evident when comparing per-capita rates in stands of early- and late-successional species. However, fewer studies have examined whether such differences in per-capita mortality are manifest as a difference in biomass turnover: do stands comprised late-successional species have lower rates of basal area mortality (mortality per unit basal area) than stands comprised of early-successional species? In this paper, the relationship between stand mortality and successional composition is examined using forest inventory data from the state of Michigan (USA). Mortality was quantified as the annual percentage of basal area lost to mortality, and successional composition was quantified using a continuous variable that reflects the successional status of each of the component tree species. Analysis by multivariate regression revealed that the difference in mortality between early- and late-successional stands is as great as 50%, and that the significance of this result is robust to collinearity between stand composition, stand age, and stand structure. This result suggests that successional composition could be used to better forecast changes in timber supply, habitat supply, and ecosystem function.     

Interspecific variability of stem photosynthesis among tree species

The photosynthetic characteristics of current-year stems of six deciduous tree species, two evergreen tree species and ginkgo (Ginkgo biloba L.) were compared. Gas exchange, chlorophyll concentration, nitrogen concentration and maximum quantum yield of PSII were measured in stems in summer and winter. A light-induced decrease in stem CO(2) efflux was observed in all species. The apparent gross photosynthetic rate in saturating light ranged from 0.72 micromol m(-2 )s(-1) (ginkgo, in winter) to 3.73 micromol m(-2) s(-1) (Alnus glutinosa (L.) Gaertn., in summer). Despite this variability, a unique correlation (slope = 0.75), based on our results and those reported in the literature, was found between gross photosynthetic rate and dark respiration rate. Mass-based gross photosynthetic rate decreased with stem mass per area and correlated to chlorophyll concentration and nitrogen concentration, both in summer and winter. The radial distribution of stem chlorophyll differed among species, but all species except ginkgo had chlorophyll as deep as the pith. In summer, the maximum quantum yield of stem PSII (estimated from the ratio of variable to maximal fluorescence; F (v)/F (m)) of all species was near the optimal value found for leaves. By contrast, the values were highly variable in winter, suggesting large differences in sensitivity to low-temperature photoinhibition. The winter values of Fv/Fm were only 31-60% of summer values for the deciduous species, whereas the evergreen conifer species maintained high F (v)/F (m) in winter. The results highlight the interspecific variability of gross photosynthesis in the stem and its correlation with structural traits like those found for leaves. The structural correlations suggest that the selection of photosynthetic traits has operated under similar constraints in stems and leaves.     

Impact of snow cover on photoinhibition and winter dessication in evergreen Rhododendron ferrugineum leaves during subalpine winter

Effects of winter snow cover on photoinhibition and possible interactions with winter desiccation were investigated in situ in an evergreen subalpine woody species, Rhododendron ferrugineum L., at the alpine timberline (1950 m a.s.l.). Timing and duration of complete snow cover markedly influenced potential efficiency of photosystem II (PSII; F(v) /F(m)). Lack of snow cover led to severe but mostly reversible photoinhibition with F(v)/F(m) values as low as 0.05. Complete snow cover immediately stopped further reductions in PSII efficiency. Snow cover promoted recovery from photoinhibition, but only if, in addition to shading by snow, plants were exposed to nonfreezing temperatures close to 0 degrees C. The F(v)/F(m) ratio was closely related to minimum leaf temperatures because both photoinhibition and recovery from photoinhibition were strongly influenced by temperature. The period without major reductions in PSII efficiency lasted for only two months, reflecting the extremely short growing period in the subalpine environment. Compared with complete snow cover, incomplete snow cover led to significantly higher water losses as well as lower dehydration tolerance, because both osmotic adjustment and changes in turgor maintenance capacity were significantly reduced. Interactions between photoinhibition and winter desiccation were masked by the direct effects of freezing temperatures. However, both photoinhibition and winter desiccation are closely linked and occur together under field conditions in this evergreen subalpine woody species.     

Effects of soil drought stress on photosynthetic gas exchange traits and chlorophyll fluorescence in Forsythia suspensa

To clarify the changes in plant photosynthesis and mechanisms underlying those responses to gradually increasing soil drought stress and reveal quantitative relationships between photosynthesis and soil moisture,soil water conditions were controlled in greenhouse pot experiments using 2-year-old seedlings of Forsythia suspensa(Thunb.) Vahl. Photosynthetic gas exchange and chlorophyll fluorescence variables were measured and analyzed under 13 gradients of soil water content. Net photosynthetic rate(PN), stomatal conductance(gs), and water-use efficiency(WUE) in the seedlings exhibited a clear threshold response to the relative soil water content(RSWC). The highest PNand WUEoccurred at RSWCof51.84 and 64.10%, respectively. Both PNand WUEwere higher than the average levels at 39.79% B RSWCB 73.04%. When RSWCdecreased from 51.84 to 37.52%,PN, gs, and the intercellular CO2 concentration(Ci)markedly decreased with increasing drought stress; the corresponding stomatal limitation(Ls) substantially increased, and nonphotochemical quenching(NPQ) also tended to increase, indicating that within this range of soil water content, excessive excitation energy was dispersed from photosystem II(PSII) in the form of heat, and the reduction in PNwas primarily due to stomatal limitation.While RSWCdecreased below 37.52%, there were significant decreases in the maximal quantum yield of PSII photochemistry(Fv/Fm) and the effective quantum yield of PSII photochemistry(UPSII), photochemical quenching(qP), and NPQ; in contrast, minimal fluorescence yield of the dark-adapted state(F0) increased markedly. Thus,the major limiting factor for the PNreduction changed to a nonstomatal limitation due to PSII damage. Therefore, an RSWCof 37.52% is the maximum allowable water deficit for the normal growth of seedlings of F. suspensa, and a water content lower than this level should be avoided in field soil water management. Water contents should be maintained in the range of 39.79% B RSWCB 73.04% to ensure normal function of the photosynthetic apparatus and high levels of photosynthesis and efficiency in F.suspensa.     

Drought inhibits photosynthetic capacity more in females than in males of Populus cathayana

We investigated sex-related photosynthetic responses to drought in the dioecious species, Populus cathayana Rehd. Plants were subjected to two watering regimes (100% and 30% of field capacity) in a semi-controlled environment. Drought significantly decreased leaf area (LA), total number of leaves (TNL), specific leaf area (SLA), relative water content, net photosynthetic rate (P(n)), transpiration (E), stomatal conductance (g(s)), intercellular CO(2) concentration (C(i)), light saturation point (L(SP)), apparent quantum yield (Phi), carboxylation efficiency (CE), light-saturated photosynthetic rate (P(max)), maximum efficiency of PSII (F(v)/F(m)) and maximum effective quantum yield of PSII (Yield), and increased the total chlorophyll concentration (TC), CO(2) compensation point (Gamma), non-photochemical quenching coefficient, peroxidase (POD) activity and carbon isotope composition (delta(13)C). Moreover, differences between males and females were detected in many of these responses. In the drought treatment, males exhibited significantly higher LA, TNL, TC, concentration of carotenoids (Caro), P(n), E, g(s), C(i), L(SP), Phi, CE, P(max), F(v)/F(m), photochemical quenching coefficient, POD activity and delta(13)C, but a lower SLA, chlorophyll a/b ratio, carotenoids/total chlorophyll ratio and Gamma than females. However, Caro, L(SP), Gamma, Phi, CE and POD activity were apparently associated with sex-related resource demands, because significant differences in these traits were detected between the sexes under well-watered conditions. Our results indicate that drought stress limits photosynthetic capacity more in females than in males.     

Constraints on light interception efficiency due to shoot architecture in broad-leaved Nothofagus species

Shoot architecture may significantly alter mean quantum flux on foliage and thus, photosynthetic productivity. There is currently only limited information about plastic alterations in shoot structure caused by within-canopy variation in mean integrated irradiance (Q(int)) in broad-leaved trees. We studied leaf and shoot structure, and nitrogen and carbon content in late-successional, widely distributed, temperate, broad-leaved Nothofagus taxa to determine the architectural controls on light harvesting and photosynthetic performance. Nothofagus fusca (Hook. f.) Oersted has larger leaves and less densely leaved shoots than the N. solandri varieties. Nothofagus solandri var. solandri (Hook. f.) Oersted is characterized by rounder leaves that potentially have a larger overlap than the ovate-triangular leaves of N. solandri var. cliffortioides (Hook. f.) Poole. Leaf dry mass (M(A)) and nitrogen content (N(A)) per unit area increased with increasing Q(int) in all species, demonstrating enhanced investment of photosynthetic biomass in high light. Although M(A) differed between species at a common irradiance, there was a uniform relationship between N(A) and Q(int) across species. Leaf carbon content per dry mass and leaf dry mass to fresh mass ratio also scaled positively with irradiance, suggesting greater structural investments in high light. In all species, shoots became more horizontal and flatter at lower Q(int), implying an enhanced use efficiency of direct irradiance in natural leaf positions. In contrast, irradiance effects on leaf aggregation varied among species. Across the data, leaf overlap or leaf area density was often greater at lower irradiances, possibly as a result of limited carbon availability for shoot axis extension growth. In N. fusca, leaves of which were more aggregated in high light, the shoot silhouette to total leaf area ratio (S(S)) declined strongly with increasing irradiance, demonstrating a lower light harvesting efficiency at high Q(int). This effect was only moderate in N. solandri var. cliffortioides and S(S) was independent of Q(int) in N. solandri var. solandri. Although the efficiency of light interception at high irradiances was lowest in N. fusca, this species had the greatest nitrogen content per unit shoot silhouette area (2N(A)/S(S)), indicating superior shoot-level photosynthetic potential. These data collectively demonstrate that shoot architecture significantly affects light interception and photosynthesis in broad-leaved trees, and that structural carbon limitations may constrain leaf light harvesting efficiency at low irradiance.     

Water stress responses of two Mediterranean tree species influenced by native soil microorganisms and inoculation with a plant growth promoting rhizobacterium

Soil microorganisms, such as plant growth-promoting rhizobacteria (PGPR), play crucial roles in plant growth, but their influence on plant water relations remains poorly explored. We studied the effects of native soil microorganisms and inoculation with the PGPR strain Aur6 of Pseudomonas fluorescens on water stress responses of seedlings of the drought-avoiding Pinus halepensis Mill. and the drought-tolerant Quercus coccifera L. Plant growth, nutrient concentrations and physiology (maximum photochemical efficiency of photosystem II (PSII; F(v)/F(m)), electron transport rate (ETR), stomatal conductance (g(s)) and predawn shoot water potential (Psi(PD))) were measured in well-watered plants, and in plants under moderate or severe water stress. Inoculation with PGPR and native soil microorganisms improved tree growth, and their interactions had either additive or synergistic effects. Both F(v)/F(m) and ETR were significantly affected by PGPR and native soil microorganisms. Marked differences in g(s) and Psi(PD) were found between species, confirming that they differ in mechanisms of response to water stress. A complex tree species x treatment interactive response to drought was observed. In P. halepensis, F(v)/F(m) and ETR were enhanced by PGPR and native soil microorganisms under well-watered conditions, but the effects of PGPR on Psi(PD) and g(s) were negative during a period of water stress. In Q. coccifera, F(v)/F(m) and ETR were unaffected or even reduced by inoculation under well-watered conditions, whereas Psi(PD) and g(s) were increased by PGPR during a period of water stress. Our results indicate that microbial associates of roots can significantly influence the response of tree seedlings to drought, but the magnitude and sign of this effect seems to depend on the water-use strategy of the species.     

Aluminum-induced decrease in CO2 assimilation in citrus seedlings is unaccompanied by decreased activities of key enzymes involved in CO2 assimilation

'Cleopatra' tangerine (Citrus reshni Hort. ex Tanaka) seedlings were irrigated daily for 8 weeks with 1/4 strength Hoagland's nutrient solution containing 0 (control) or 2 mM aluminum (Al). Leaves from Al-treated plants had decreased CO2 assimilation and stomatal conductance, but increased intercellular CO2 concentrations compared with control leaves. On a leaf area basis, 2 mM Al increased activities of key enzymes in the Calvin cycle, including ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), NADP-glyceraldehyde-3-phosphate dehydrogenase (GAPDH), phosphoribulokinase (PRK), stromal fructose-1,6-bisphosphatase (FBPase), and a key enzyme in starch synthesis, ADP-glucose pyrophosphorylase (AGPase), compared with control leaves. Aluminum had no effect on cytosolic FBPase activity, but it decreased sucrose phosphate synthase (SPS) activity. Aluminum had no effect on area-based concentrations of carbohydrates, glucose-6-phosphate (G6P) and fructose 6-phosphate (F6P) or the G6P:F6P ratio, but it decreased the area-based concentration of 3-phosphoglycerate (PGA). Photochemical quenching coefficient (qP) and electron transport rate through PSII were greatly reduced by Al. Non-photochemical quenching coefficient (NPQ) was less affected by Al than qP and electron transport rate through PSII. We conclude that the reduced rate of CO2 assimilation in Al-treated leaves was probably caused by a combination of factors such as reduced electron transport rate through PSII, increased closure of PSII reaction centers and increased photorespiration.     

Functional traits and plasticity in response to light in seedlings of four Iberian forest tree species

We investigated the differential roles of physiological and morphological features on seedling survivorship along an experimental irradiance gradient in four dominant species of cool temperate-Mediterranean forests (Quercus robur L., Quercus pyrenaica Willd., Pinus sylvestris L. and Pinus pinaster Ait.). The lowest photochemical efficiency (F(v)/F(m) in dark-adapted leaves) was reached in deep shade (1% of full sunlight) in all species except Q. robur, which had the lowest photochemical efficiency in both deep shade and 100% of full sunlight. Species differed significantly in their survival in 1% of full sunlight but exhibited similar survivorship in 6, 20 and 100% of full sunlight. Shade-tolerant oaks had lower leaf area ratios, shoot to root ratios, foliage allocation ratios and higher rates of allocation to structural biomass (stem plus thick roots) than shade-intolerant pines. Overall phenotypic plasticity for each species, estimated as the difference between the minimum and the maximum mean values of the ecophysiological variables studied at the various irradiances divided by the maximum mean value of those variables, was inversely correlated with shade tolerance. Observed morphology, allocation and plasticity conformed to a conservative resource-use strategy, although observed differences in specific leaf area, which was higher in shade-tolerant species, supported a carbon gain maximization strategy. Lack of a congruent suite of traits underlying shade tolerance in the studied species provides evidence of adaptation to multiple selective forces. Although the study was based on only four species, the importance of ecophysiological variables as determinants of interspecific differences in survival in limiting light was demonstrated.     

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.     

Post-breeding bird responses to canopy tree retention, stand size, and edge in regenerating Appalachian hardwood stands

Avian use of even-aged timber harvests is likely affected by stand attributes such as size, amount of edge, and retained basal area, all characteristics that can easily be manipulated in timber harvesting plans. However, few studies have examined their effects during the post-breeding period. We studied the impacts of clearcut, low-leave two-age, and high-leave two-age harvesting on post-breeding birds using transect sampling and mist-netting in north-central West Virginia. In our approach, we studied the effects of these harvest types as well as stand size and edge on species characteristic of both early-successional and mature forest habitats. In 2005-2006, 13 stands ranging from 4 to 10 years post-harvest and 4-21 ha in size were sampled from late June through mid-August. Capture rates and relative abundance were similar among treatments for generalist birds. Early-successional birds had the lowest capture rates and fewer species (∼30% lower), and late-successional birds reached their highest abundance and species totals (double the other treatments) in high-leave two-age stands. Area sensitivity was evident for all breeding habitat groups. Both generalist and late-successional bird captures were negatively related to stand size, but these groups showed no clear edge effects. Mean relative abundance decreased to nearly zero for the latter group in the largest stands. In contrast, early-successional species tended to use stand interiors more often and responded positively to stand size. Capture rates for this group tripled as stand size increased from 4 to 21 ha. Few birds in the forest periphery responded to harvest edge types despite within-stand edge effects evident for several species. To create suitable habitat for early-successional birds, large, non-linear openings with a low retained basal area are ideal, while smaller harvests and increased residual tree retention would provide habitat for more late-successional birds post-breeding. Although our study has identified habitat use patterns for different species in timber harvests, understanding habitat-specific bird survival is needed to help determine the quality of silvicultural harvests for post-breeding birds.     

Response of leaf photosynthetic characteristics of Syringa oblata and Syringa reticulata var.mandshurica to chilling stress

Syringa species not only have good ornamental properties but also play an important role in the landscaping and environmental purification of cities.To investigate the chilling stress resistance of Syringa oblata Lindl.and Syringa reticulata var.mandshurica and provide theoretical grounds for the practical cultivation of Syringa species,in vitro leaves were used to study photosynthetic gas exchange parameters and chlorophyll fluorescence parameters.After nine hours of chilling,decreasing rates of net photosynthesis,stomatal conductance,and transpiration in S.reticulata var.mandshurica leaves were significantly greater than that of the S.oblata,while intercellular CO2 concentrations in S.oblata leaves were higher than those in S.reticulata var.mandshurica.The quantum yield of PSII reaction center(APSII)declined in S.reticulata and light capture efficiency(Fv 0/Fm 0)was stable.However,reduction percentages of Fv 0/Fm 0,APSII,and Fv/Fm in S.oblata were significant higher than those of S.reticulata var.mandshurica.After nine hours of chilling,the relative variable fluorescence of VJ and VI of S.oblata increased and the increasing rate of VJ was greater than VI.In contrast,the change of VJ and VI in S.reticulata var.mandshurica leaves was relatively small.This suggests that chilling primarily damaged the electron transport process of QA to QB at the receptor site of the PSII reaction center.Photosynthetic capacity of S.oblata was more sensitive to chilling stress compared to S.reticulate var.mandshurica,which the limitations were mainly due to non-stomatal factors such as the decrease in electron transport efficiency,activity in the PSII reaction center,and the destruction of the photodamage defense system.     

Coordination of crown structure, leaf plasticity and carbon gain within the crowns of three winter-deciduous mature trees

We examined the vertical profiles of leaf characteristics within the crowns of two late-successional (Fagus crenata Blume and Fagus japonica Maxim.) and one early-successional tree species (Betula grossa Sieb. et Zucc.) in a Japanese forest. We also assessed the contributions of the leaves in each crown layer to whole-crown instantaneous carbon gain at midday. Carbon gain was estimated from the relationship between electron transport and photosynthetic rates. We hypothesized that more irradiance can penetrate into the middle of the crown if the upper crown layers have steep leaf inclination angles. We found that such a crown has a high whole-crown carbon gain, even if leaf traits do not change greatly with decreasing crown height. Leaf area indices (LAIs) of the two Fagus trees (5.26-5.52) were higher than the LAI of the B. grossa tree (4.50) and the leaves of the F. crenata tree were more concentrated in the top crown layers than were leaves of the other trees. Whole-crown carbon gain per unit ground area (micromol m(-2) ground s(-1)) at midday on fine days in summer was 16.3 for F. crenata, 11.0 for F. japonica, and 20.4 for B. grossa. In all study trees, leaf dry mass (LMA) and leaf nitrogen content (N) per unit area decreased with decreasing height in the crown, but leaf N per unit mass increased. Variations (plasticity) between the uppermost and lowermost crown layers in LMA, leaf N, the ratio of chlorophyll to N and the ratio of chlorophyll a to b were smaller for F. japonica and B. grossa than for F. crenata. The light extinction coefficients in the crowns were lower for the F. japonica and B. grossa trees than for the F. crenata tree. The leaf carbon isotope ratio (delta(13)C) was higher for F. japonica and B. grossa than for F. crenata, especially in the mid-crown. These results suggest that, in crowns with low leaf plasticity but steep leaf inclination angles, such as those of F. japonica and B. grossa trees, irradiance can penetrate into the middle of the crowns, thereby enhancing whole-crown carbon gain.     

Seasonal and yearly variations in light use and nitrogen use by seedlings of four deciduous broad-leaved tree species invading larch plantations

Several deciduous broad-leaved tree species, differing in leaf phenology, invade larch (Larix kaempferii (Lamb.) Carrière) plantations in Japan. The understory light environment of larch forests changes drastically between the leafy and leafless periods. To determine how the invading seedlings exploit the changing light environment, and if phenological differences reflect the light- and nitrogen-use traits of the seedlings, we measured leaf phenology, seasonal changes in light-saturated photosynthetic rate (P(sat)), leaf nitrogen (N) content (N(area)), chlorophyll/nitrogen ratio (Chl/N), specific leaf area (SLA) and N remobilization rate (NRMR) over 3 years. The mid-successional or gap-phase species, Magnolia hypoleuca Siebold & Zucc., had a short leafy period and high P(sat) and NRMR. In contrast, two late-successional tree species, Prunus ssiori Friedr. Schmidt, which undergoes leaf flush before larch, and Carpinus cordata Blume, which maintains green leaves until frost, both had low P(sat) and NRMR but exploited the opportunity for growth during the period when the larch canopy trees were leafless. Quercus mongolica Fisch. ex Ledeb. var. crispula (Blume) Ohashi, a mid-late-successional species that underwent leaf flush at the same time as the overstory larch, had values of photosynthetic parameters between those of the gap-phase and late-successional species. Among species, M. hypoleuca and Q. mongolica had higher photosynthetic rates and photosynthetic N-use efficiencies. In all species, the relationship between N(area) and P(sat) showed species-specific yearly fluctuations; however, there was no yearly fluctuation in the relationship between N(area) and P(sat) at CO2 saturation. Yearly fluctuations in the N(area)-P(sat) relationship appeared to be induced by changes in SLA and N-use characteristics, which in turn are affected by climatic variations.     

Acclimation of leaves to contrasting irradiance in juvenile trees differing in shade tolerance

Leaves developing in different irradiances undergo structural and functional acclimation, although the extent of trait plasticity is species specific. We tested the hypothesis that irradiance-induced plasticity of photosynthetic and anatomical traits is lower in highly shade-tolerant species than in moderately shade-tolerant species. Seedlings of two evergreen conifers, shade-tolerant Abies alba Mill. and moderately shade-tolerant Picea abies Karst., and two deciduous angiosperm species, highly shade-tolerant Fagus sylvatica L. and moderately shade-tolerant Acer pseudoplatanus L., were grown in deep shade (LL, 5% of full irradiance) or in full solar irradiance (HL) during 2003 and 2004. Steady state responses of quantum yield of PSII (Phi(PSII)), apparent electron transport rate (ETR), nonphotochemical quenching (NPQ) and photochemical quenching (qP) were generally modified by the light environment, with slower declines in Phi(PSII) and qP and greater maximal ETR and NPQ values in HL plants in at least one season; however, no link between quantitative measures of plasticity of these traits and shade tolerance was found. Plasticity of nine anatomical traits (including palisade cell length, which was reduced in LL) showed no relationship with shade tolerance, but was less in conifers than in deciduous trees, suggesting that leaf life span may be a significant correlate of plasticity. When LL-acclimated plants were exposed to HL conditions, the degree and duration of photoinhibition (measured as a decline in maximum quantum yield) was greatest in F. sylvatica, much lower in P. abies and A. alba, and lowest in A. pseudoplatanus. Thus, as with the other traits studied, vulnerability to photoinhibition showed no relationship with shade tolerance.     

Chlorophyll a fluorescence analysis along a vertical gradient of the crown in a poplar (Oxford clone) subjected to ozone and water stress

An experiment in open-top chambers was carried out in summer 2008 at Curno (Northern Italy) in order to study the effects of ozone and mild water stress on poplar cuttings (Oxford clone). In this experiment direct fluorescence parameters (JIP-test) were measured in leaves from different sections of the crown (L: lower; M: medium; U: upper parts of the crown). The parameters considered were calculated at the different steps of the fluorescence transient, and include maximum quantum yield efficiency in the dark-adapted state (F(v)/F(M)); the L-band, at 100?∝?s, that expresses the stability of the tripartite system reaction centre-harvesting light complex-core antenna; the K-band, at 300?∝?s, that expresses the efficiency of the oxygen-evolving complex; the J-phase, at 2 ms, that expresses the efficiency with which a trapped exciton can move an electron into the electron transport chain from Q(A)(-) to the intersystem electron acceptors; the IP-phase, which expresses the efficiency of electron transport around the photosystem 1 (PSI) to reduce the final acceptors of the electron transport chain, i.e., ferredoxin and NADP; and finally the performance index total (PItot) for energy conservation from photons absorbed by PSII to the reduction flux of PSI end acceptors. The main results are: (i) different dynamics were observed between leaves in the lower section, whose PItot decreased over time, and those in the upper sections in which it increased, with a dynamic connected to the leaf age; (ii) ozone depressed all the considered fluorescence parameters in basal leaves of well-watered plants, while it had little or no damaging effect on medium-level or upper-section leaves; (iii) PItot and IP-phase increased in upper leaves of plants subjected to ozone stress, as well as the net photosynthesis; (iv) water stress increased PItot of leaves in all levels of the crown. The results suggest that ozone-damaged poplar plants compensate, at least partially, for the loss of photosynthesis with higher photosynthetic rates in young leaves (in the upper section of the crown), more efficient to fix carbon.