Growth, Allocation and Water Relations of Shade-grownQuercus rubraL. Saplings Exposed to a Late-season Canopy Gap

For understory saplings to exploit canopy gaps successfully,carbon gain must increase in the gap environment. We predictedthat total biomass of shade-grown red oak saplings would increaseafter exposure to a late-season canopy gap, and that increasedwater and nutrient demand within the canopy gap would drivechanges in the allocation of this carbon. Shade-grown red oaksaplings acclimated to gaps by increasing biomass during theseason of gap formation and increasing the potential for carbongain in the following summer. Within-season carbon gain didnot result from greater production of leaf area, so it mostlikely arose from higher photosynthetic rates of existing shade-developedfoliage, which may be linked to accumulation of leaf nitrogen.During the season of gap formation, shade-gap plants increasedallocation to storage of total non-structural carbohydrates(TNC), and to root growth. The increase in TNC storage suggeststhat shade-developed saplings exposed to gaps were also primedfor fast growth and carbon gain in the following summer. Theincrease in root growth suggests that higher nutrient and waterdemand drove allocation shifts to enhance the capacity for nutrientand water uptake in the gap. Plant hydraulic conductivity (K_a)of shade-grown plants was limited upon exposure to the gap,possibly because of embolism formation resulting from the abruptincrease in water demand. Greater water potential gradientscompensated for limitations to K_a, allowing saplings to maintainhigh transpiration rates, suggesting that actual water uptakeof shade-gap plants was unaffected by gap exposure. Acclimation; canopy gaps; carbon allocation; hydraulic conductivity; nitrogen allocation; non-structural carbohydrates;Quercus rubra L.; red oak; stomatal conductance; transpiration; water-relations     

Photosynthetic responses to light in seedlings of selected Amazonian and Australian rainforest tree species

Summary Seedlings of the Caesalpinoids Hymenaea courbaril, H. parvifolia and Copaifera venezuelana, emergent trees of Amazonian rainforest canopies, and of the Araucarian conifers Agathis microstachya and A. robusta, important elements in tropical Australian rainforests, were grown at 6% (shade) and 100% full sunlight (sun) in glasshouses. All species produced more leaves in full sunlight than in shade and leaves of sun plants contained more nitrogen and less chlorophyll per unit leaf area, and had a higher specific leaf weight than leaves of shade plants. The photosynthetic response curves as a function of photon flux density for leaves of shade-grown seedlings showed lower compensation points, higher quantum yields and lower respiration rates per unit leaf area than those of sun-grown seedlings. However, except for A. robusta, photosynthetic acclimation between sun and shade was not observed; the light saturated rates of assimilation were not significantly different. Intercellular CO₂ partial pressure was similar in leaves of sun and shade-grown plants, and assimilation was limited more by intrinsic mesophyll factors than by stomata. Comparison of assimilation as a function of intercellular CO₂ partial pressure in sun- and shade-grown Agathis spp. showed a higher initial slope in leaves of sun plants, which was correlated with higher leaf nitrogen content. Assimilation was reduced at high transpiration rates and substantial photoinhibition was observed when seedlings were transferred from shade to sun. However, after transfer, newly formed leaves in A. robusta showed the same light responses as leaves of sun-grown seedlings. These observations on the limited potential for acclimation to high light in leaves of seedlings of rainforest trees are discussed in relation to regeneration following formation of gaps in the canopy.     

Leaves of Japanese oak (Quercus mongolica var. crispula) mitigate photoinhibition by adjusting electron transport capacities and thermal energy dissipation along the intra-canopy light gradient

We investigated the morphological and physiological acclimation of leaves grown within a canopy of Japanese oak tree (Quercus mongolica var. crispula) in terms of the susceptibility to photoinhibition under various growth light conditions. The maximum rates of photosynthesis (P(max) ) and electron transport (ETR(max) ) were higher in mature leaves grown under stronger light with higher area-based leaf nitrogen (N) content closely associated with higher leaf mass per area. The net photosynthetic (P(n) ) and electron transport (ETR) rates corresponding to the daily peak photosynthetic photon flux density (PPFD(max) ) during leaf maturation were almost comparable to P(max) and ETR(max) , respectively. Conversely, P(n) and ETR at the daily average PPFD (PPFD(avg) ) were substantially low in shade-grown leaves when compared with P(max) and ETR(max) . The susceptibility to photoinhibition at PPFD(max) , i.e. at sunflecks for the shade-grown leaves, was assessed by the rate of excess energy production. Although sun leaves showed higher rates of electron transport and thermal energy dissipation than shade leaves under PPFD(max) conditions, the rate of excess energy production was almost constant across shade to sun leaves. The shade leaves of the Japanese oak grown within a crown were suggested to adjust their N investment to maintain higher photosynthetic capacities compared with those required to maximize the net carbon gain, which may facilitate the dissipation of the excessive light energy of sunflecks to circumvent photoinhibition in cooperation with thermal energy dissipation.     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. I. Interactions between photosynthetic acclimation and photoinhibition during simulated canopy gap formation

Natural regeneration of European beech (Fagus sylvatica L.) establishes under shade, but sudden exposure to high irradiance may occur due to openings in the canopy. To elucidate ecophysiological mechanisms associated with survival of European beech seedlings, the gas exchange, chlorophyll concentrations, and chlorophyll a fluorescence parameters of two different beech populations were studied under changing light conditions. Plants were grown both in a growth chamber and at a natural site (one population) where the seedlings were raised in containers placed in understory and in simulated canopy gaps. Upon exposure to high light in the growth chamber, photosynthetic rates of shade-acclimated leaves of seedlings from both populations increased severalfold and then decreased over several days to the rates of the low-light control seedlings. High-light seedlings always had the highest photosynthetic rates. Initial fluorescence displayed a trend opposite that of photosynthesis; it increased over time, and relative fluorescence and half-time rise declined continuously until the end of experiment to very low values. Exposure to high light of shade-acclimated seedlings resulted in a shift in chlorophyll concentrations to levels intermediate between high-light and low-light seedlings. The light treatment effects were statistically greater than population effects; however, seedlings from the Abetone population were found to be more susceptible to changing light conditions than seedlings from Sicily. Reciprocal light treatments on plants growing at the natural site confirmed the results obtained in the growth chamber experiment. Overall, beech seedlings grown in the field appeared to have a fairly large acclimation potential achieved by plasticity in the photosynthetic apparatus. The lack of pronounced acclimation to high light in seedlings grown in the growth chamber was ascribed to a threshold-type relationship between the acclimation capacity and the level of damage. These observations on the limited potential for acclimation to high light in leaves of European beech seedlings which show a clear capability to exploit sunflecks, are discussed in relation to regeneration following canopy gap formation and reinforce the view of the central role of gap formation in forest dynamics. We conclude that small forest gaps (in which sunflecks play a major role) may present a favorable environment for survival and growth of beech because of their limited ability to acclimate to a sudden increase in irradiance and because of the moderate levels of light stress found in small gaps.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance

Dipterocarps dominate the canopy of lowland tropical rain forest in South‐east Asia. Seedlings of these species form diverse assemblages on the forest floor where low irradiance severely limits their growth. Further growth depends largely upon the increased irradiance that can occur with the creation of canopy gaps. However, the response of dipterocarp seedlings to increased irradiance and their subsequent establishment in the canopy may be influenced by the availability of other resources, such as nutrient availability. We investigated the influence of nitrogen supply on aspects of the photosynthetic physiology and growth of seedlings of four dipterocarp species (Shorea leprosula, Shorea johorensis, Shorea oleosa and Dryobalanops lanceolata) growing under low irradiance, during transfer from low to high irradiance, and during subsequent growth at high irradiance. All four species increased growth and photosynthetic capacity in response to N‐supply at high irradiances but not at low irradiance approximating that which can be expected to occur in the forest understorey. When seedlings grown at low irradiances and varying N‐supply were exposed to a large increase in irradiance, all species showed some degree of initial photodamage (measured through chlorophyll fluorescence), the extent of which was similar between species but differed markedly depending on the pre‐exposure growth irradiance and N‐supply. Greater photodamage occurred in seedlings grown at lower compared with higher N‐supply and irradiance. Despite these initial difference in the extent of this photodamage, all seedlings demonstrated a similar capacity to recover from damage. However, the alterations in the photosynthetic physiology of leaves during this recovery differed between species and depended on N‐supply. Under high N‐supply all species apart from S. oleosa increased photosynthetic capacity per unit chlorophyll following exposure to high irradiance by increasing photosynthetic capacity per unit leaf area while, under low N‐supply, an increase in photosynthetic capacity per unit leaf only occurred in D. lanceolata. Our results suggest that variations in N‐availability may have a much greater impact on the relative competitiveness of dipterocarp seedlings during the regenerative phase following canopy gap formation than physiological differences between seedlings. Our results demonstrate a potentially significant role for N‐availability in the regeneration dynamics and distribution of canopy‐dominating dipterocarp species.     

油樟幼苗对马尾松林窗面积的光合响应特征

为了解马尾松人工林窗对伴生树种的影响,为马尾松人工纯林的团块状混交提供科学依据,研究了10 m×10 m(T1)、15 m×15 m(T2)、20 m×20 m(T3)、25 m×25 m(T4)、30 m×30 m(T5)、35 m×35 m(T6)和40 m×40 m(T7)马尾松人工林窗中油樟(Cinnamomum longepaniculatum)幼苗叶片形态和光合生理特征的变化,探讨马尾松林窗斑块对混生树种生长的影响。结果表明:1)林窗面积低于20 m×20 m时,油樟幼苗叶片最大净光合速率显著低于旷地对照;2)叶片比叶重随着林窗面积的增大显著升高;林窗内油樟幼苗叶氮含量在小林窗中(10 m×10 m)显著低于旷地对照,但在大林窗中(如20 m×20 m)显著高于对照;林窗内幼苗叶磷含量则与旷地无显著性差异;3)叶氮在光合组分中的总分配系数随着林窗面积的增加而增大,其中叶氮在羧化组分中的分配系数升高尤为明显,而捕光组分的分配系数在林窗面积10 m×10 m—20 m×20 m范围内随林窗面积的增加而显著降低。可见,当马尾松林窗面积低于20 m×20 m时,林窗环境会显著影响油樟幼苗的光合能力,油樟幼苗可以通过调节比叶面积、叶氮含量以及叶氮在光合组分中的分配等形态、生理适应特征来适应林窗环境的变化。     

四种亚高山针叶林树种的表型可塑性对不同光照强度的响应

以青藏高原东缘亚高山针叶林群落演替后期种岷江冷杉、演替中后期种粗枝云杉和青榨槭、及先锋树种红桦为材料,研究了不同光强下生长的4种树苗生长、生物量分配、叶片形态和光合特性,探讨植物幼苗的形态和生理特征的表型可塑性与光适应的关系。结果表明：（1）弱光环境中生长的4种植物的基茎、相对生长速率、叶片厚度、根重比、最大净光合速率、光饱和点、光补偿点、暗呼吸速率较低,而比叶面积、地上／地下生物量、茎长／茎重、叶重比和茎重比较高。（2）大部分光环境下岷江冷杉幼苗的最大净光合速率和暗呼吸速率低于粗枝云杉,青榨槭幼苗的最大净光合速率和暗呼吸速率略低于红桦。（3）高光强下生长的粗枝云杉和红桦幼苗的相对生长速率分别大于岷江冷杉和青榨槭,但在低光强下则与之相反。（4）粗枝云杉和红桦幼苗的11种可塑性指数平均值则分别大于岷江冷杉和青榨槭。岷江冷杉适应弱光环境的能力略强于粗枝云杉和红桦,但适应强光的能力较差。生理适应的可塑性指数大于形态适应的可塑性指数,表明前者在4种植物幼苗光适应方面起到了重要的作用。研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。     

Photosynthetic plasticity of two rain forest shrubs across natural gap transects

Summary Photosynthetic plasticity of two congeneric shrub species growing under natural field conditions was compared along transects spanning two canopy gaps in a Costa Rican rain forest. Piper arieianum is a shadetolerant species common in successional and mature forests, whereas P. sancti-felicis is a pioneer species abundant in abandoned clearings and large gaps. Twenty potted cuttings of each species were placed at regular intervals along two east-west transects crossing a small branch-fall gap and a large tree-fall gap. Along the transects, the percent of full sun photon flux density varied from less than 2% to 45%. After six months of growth under these conditions, leaves were monitored for incident photon flux density, photographic measures of light availability, photosynthetic capacity (A_max), leaf nitrogen content, leaf chlorophyll content, and specific leaf mass. Although both species demonstrated considerable plasticity in A_max across gap transects, P. sancti-felicis leaves had a superior capacity to track closely variation in light availability, particularly in the larger gap. For regressions of A_max on measures of light availability, P. sancti-felicis consistently showed a 3.5 to 5-fold higher coefficient of determination (R²) and a 3 to 4-fold higher slope than P. arieianum. In both species leaf nitrogen content per leaf area increased significantly with light availability, although P. sancti-felicis, again, showed a much stronger relationship between these variables. Across the transects, mean chlorophyll content per unit leaf area did not differ significantly between the species, whereas mean chlorophyll content per unit leaf dry mass was 3-times greater in leaves of P. sancti-felicis. Piper arieianum exhibited highly significant increases in chlorophyll a:b ratio with increased light availability, whereas P. sancti-felicis lacked significant variation in this trait across a gradient of light availability. Mean specific leaf mass did not vary significantly between species across the gap transects. The nature of the light acclimatory response differs quantitatively and qualitatively between these species. An important constraint on light acclimation of the shade-tolerant P. arieianum is its inability to increase photosynthetic nitrogen-use efficiency under conditions of high light availability. The lack of plasticity in chlorophyll a:b ratios does not restrict light acclimation of A_max in P. sancti-felicis. Leaves of P. arieianum exhibited symptoms of chronic photoinhibition in exposed microsites within the large gap. Species differences in the capacity to finely adjust A_max across a wide range of light conditions may be attributed to their maximum growth potential. Light acclimation in species with low maximum growth potential may be constrained at the cellular level by rates of protein and chlorophyll synthesis and at the whole-plant level by low maximum rates of uptake and supply of nutrients and water. For P. arieianum, restriction of photosynthetic plasticity is likely to limit competitive abilities of plants in high-light conditions of large gaps and clearings, whereas observed habitat restrictions for P. sancti-felicis do not appear to depend upon the highly-developed capacity for adjustment of A_max observed in this species.     

The dynamics of photosynthetic acclimation to changes in light quanlity and quality in three Australian rainforest tree species

Photosynthetic acclimation was studied in seedlings of three subtropical rainforest species representing early (Omalanthus populifolius), middle (Duboisia myoporoides) and late (Acmena ingens) successional stages in forest development. Changes in the photosynthetic characteristics of pre-existing leaves were observed following the transfer of plants between deep shade (1–5% of photosynthetically active radiation (PAR), selectively filtered to produce a red/far-red (R/FR) ratio of 0.1) and open glasshouse (60% PAR and a R/FR ratio of 1.1–1.2), and vice versa. The extent and rate of response of the photosynthetic characteristics of each species to changes in light environment were recorded in this simulation of gap formation and canopy closure/overtopping. The light regimes to which plants were exposed produced significant levels of acclimation in all the photosynthetic parameters examined. Following transfer from high to low light, the light-saturated rate of photosynthesis was maintained near pre-transfer levels for 7 days, after which it decreased to levels which closely approximated those in leaves which had developed in low light. The decrease in photosynthetic capacity was associated with lower apparent quantum yields and stomatal conductances. Dark respiration was the parameter most sensitive to changes in light environment, and responded significantly during the first 4–7 days after transfer. Acclimation of photosynthetic capacity to increases in irradiance was significant in two of the three species studied, but was clearly limited in comparison with that of new leaves produced in the high light conditions. This limitation was most pronounced in the early-successional-stage species, O. populifolius. It is likely that structural characteristics of the leaves, imposed at the time of leaf expansion, are largely responsible for the limitations in photosynthetic acclimation to increases in irradiance.     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Invasion of broad-leaf tree species into a larch plantation: seasonal light environment, photosynthesis and nitrogen allocation

We have studied how tree seedlings with differing leaf phenological traits change their nitrogen allocation in seasonally changing light environments. Specifically, we have investigated seasonal changes in maximum leaf photosynthetic rates in situ, and changes in nitrogen, chlorophyll and ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco) content. We have also estimated nitrogen allocation characteristics from leaf nitrogen to photosynthetic proteins, light-harvesting protein and other proteins in four successional tree species growing in a larch plantation. The gap phase species, Magnolia hyporeuca Siebold et Zucc., displays higher photosynthetic rates to compensate for the short leafy period and allocates more nitrogen to Rubisco. In contrast, tree species that grow at the forest floor, Prunus ssiori F. Schmidt and Carpinus cordata Blume, use the leafless period of the canopy trees to change their allocation of nitrogen from Rubisco to chlorophyll in line with canopy closure. Quercus mongolica Fischer ex Ledeb. var. crispula (Blume) Ohashi is a mid-successional tree species, and maintains a high photosynthetic rate in September because it allocates more nitrogen to Rubisco. We conclude that phenological differences in tree seedlings are clearly reflected in the nitrogen allocation pattern at gaps and at the forest floor in larch plantations.     

Comparative life history and physiology of two understory Neotropical herbs

Summary Demography and physiology of two broad-leaved understory tropical herbs (Marantaceae) were studied in gaps and shaded understory in large-scale irrigated and control treatments during the dry season at Barro Colorado Island (BCI), Panama. Because photosynthetic acclimation potential may not predict light environments where tropical species are found, we studied a suite of physiological features to determine if they uniquely reflect the distribution of each species. Calathea inocephala and Pleiostachya pruinosa grow and reproduce in gaps, persist in shade, and have equivalent rates of leaf production. Calathea leaves survived 2 to 3 times as long as leaves of Pleiostachya and plants of Pleiostachya were 6 to 8 times more likely to die as plants of Calathea during 3.5 years of study. Pleiostachya had lowest survival in shade and when not irrigated during the dry season, while Calathea survived well in both habitats and both treatments. Pleiostachya had higher photosynthetic capacity and stomatal conductance than Calathea and acclimated to gaps by producing leaves with higher photosynthetic capacity. Calathea had lower mesophyll CO₂ concentrations than Pleiostachya. Both species had similar dark respiration rates and light compensation points, and water-use and nitrogen-use efficiencies were inversely related between species. Species showed no differences in leaf osmotic potentials at full turgor. Calathea roots were deeper and had tuberous swellings.Leaf-level assimilation and potential water loss are consistent with where these species are found, but photosynthetic acclimation to high light does not reflect both species' abilities to grow and reproduce in gaps. Pleiostachya's gap-dependent, rapid growth and reproduction require high rates of carbon gain in short-lived leaves, which can amortize their cost quickly. High rates of water loss are associated with reduced longevity during drought. Calathea's roots may confer greater capacitance, while its leaves are durable, long-lived and have lower water loss, permitting persistence long after gap closure.     

四种热带雨林树种幼苗比叶重,光合特性和暗呼吸对生长光环境的适应

雾凉季研究了西双版纳热带雨林4种植物幼苗对生长光环境的适应,其中两个树种幼苗喜光（团花和滇南插柚紫）,两个树种幼苗耐荫（滇南红厚壳和玉蕊）发现弱光环境中生长的4种植物比叶重、光合能力、光饱和点、光补偿点暗呼吸速度、叶绿素a/b比较低,叶绿素含量较高。玉蕊和滇南红厚壳幼苗的光合能力和呼吸速度 于团花树和滇南插柚紫。团花树和滇南插机紫的比叶重和光合作用的可塑性大于玉蕊和滇南红厚壳。高光强下生长的团花树和滇南插机紫增加叶氮分配给羧化酶的比较。减少分配给叶绿素的比例。滇南红厚壳和玉蕊适应弱光环境的能力略强于团花树和滇南插机紫,但适应强光的能力较差。研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说,单位干重叶的光合能力和呼吸速率并未表现出利于光适应的可塑性,表明4种植物生理适应能力较差,形态学上的适应在4种热带雨林树种幼苗光适应方面起到了重要的作用,叶氮分配也是它们光适应的策略之一。     

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.     

A field study of photosynthetic temperature acclimation in Carex eleocharis Bailey

Abstract. Seasonal patterns in photosynthetic temperature acclimation and growth were investigated in the sedge, Carex eleocharis Bailey, a species which has demonstrated a marked capacity for shifts in the photosynthetic temperature optimum in previous growth chamber studies. The seasonal production of new leaves was 90% complete by the earliest study date, June 3. Shifts in the photosynthetic temperature optimum of 10°C (from 15 to 25°C) were observed during the months of June and July. These results indicate that in situ acclimatory adjustments in C. eleocharis occur in existing leaf tissue, rather than new leaves which are produced as the season progresses. Despite the 10°C increase in the temperature optimum, mean mid-day leaf temperatures were higher than the optimum throughout the summer. A broad temperature response appeared to be more important than the acclimation adjustments in maintaining near-maximum photosynthesis rates during the mid-day period. Seasonal shifts in the photosynthetic temperature optimum were not as great as those previously observed in growth chamber studies. This discrepancy arises because of the capacity for growth chamber grown plants to produce new leaves with temperature response characteristics closely tuned to the growth temperature regime. In field-grown plants the production of 90% of the leaves during the cool portion of the season places limitations on the potential for acclimation to the warmer midsummer temperatures.     

Acclimation to changing light conditions of long-term shade-grown beech (Fagus sylvatica L.) seedlings of different geographic origins

 Effects of changing light conditions on the ecophysiological condition behind survival were examined on beech from two different populations. Plants were grown in a greenhouse under simulated understorey and canopy gap light conditions. Upon exposure to high light maximum photosynthesis of shade-acclimated leaves increased followed by a reduction over several days to between high- and low-light control rates. In the reciprocal transfer, the decrease in maximum photosynthesis was rapid during the first 2–3 days and then levelled off to values comparable to low-light controls. Seedlings from Sicily (Madonie) showed generally higher maximum photosynthetic rates than those from Abetone. Leaf conductance varied in the same direction as photosynthesis in high- to low-light seedlings but to a lesser degree. Leaves grown under low light and exposed to high light experienced photoinhibition. The Abetone population was more susceptible to photoinhibitory damage than the seedlings from Sicily. Exposure to high light of shade-acclimated seedlings resulted in intermediate chlorophyll concentrations between levels of the high-light and low-light seedlings. Carotenoid concentration was unaffected by treatments. Seedlings grew more in high light, but had a lower leaf area ratio. Light-limited seedlings showed a shift in carbon allocation to foliage. Leaves formed in the new light regime maintained the same anatomy that had been developed before transfer. Seedlings from Sicily had thicker leaves than those of seedlings from Abetone. Seedlings from Abetone were found to be more susceptible to changing light conditions than seedlings from Sicily. We conclude that small forest gaps may represent a favorable environment for photosynthesis and growth of beech regeneration as a result of the limited ability of seedlings to acclimate to sudden increases in high irradiance and because of the moderate levels of light stress in small gaps. Received: 11 April 1997 / Accepted: 11 December 1997     

Effects of nutrient supply on photosynthetic acclimation and photoinhibition of one-year-old foliage of Picea abies

The present study examined the effect of nutrient supply on acclimation potential and photoinhibitory susceptibility in one-year-old foliage of shade-grown Norway spruce ( Picea abies Karst.) seedlings transferred to high-light, as compared to seedlings grown in shade or in high-light. After the transfer, the photosynthetic response to intercellular CO₂ and chlorophyll fluorescence parameters were measured, and the allocation of leaf N within the photosynthetic apparatus was estimated. The effects of light and nutrient supply markedly differed. Light availability positively affected both photosynthetic capacity on an area basis (but not on a mass basis) and leaf N allocation to ribulose-1,5-biphosphate carboxylase/oxygenase (Rubisco), thereby increasing photosynthetic capacity per unit nitrogen. By contrast, nutrient supply in high-light affected photosynthetic capacity both on an area and a mass basis, but did not affect the allocation of N within the photosynthetic apparatus. Fluorescence parameters indicated that shade-grown seedlings transferred to high-light with low-nutrient supply, underwent prolonged photoinhibition. By contrast, transferred seedlings with high-nutrient supply were able to avoid photoinhibition, increasing their photosynthetic capacity on an area basis. On the whole, nutrient shortage was found to prevent the acclimation response and to increase the photoinhibitory susceptibility to changing light conditions in P. abies mature needles.     

林下与全光下地枫皮叶片形态和光合特性的比较

测量了林下与全光下地枫皮的叶片形态和光合-光响应曲线,探讨光强对地枫皮的形态和生理特性的影响。结果表明：林下与全光下地枫皮叶片净光合速率（Pn）、气孔导度（Gs）、蒸腾速率（Tr）和水分利用效率（WUE）对光强的响应趋势均基本一致,但全光下的Pn、Gs和Tr值较高,林下WUE值较高。全光下地枫皮的最大净光合速率、光饱和点和光补偿点均极显著高于林下,但弱光下的量子效率无显著差异;林下地枫皮的叶长、叶宽、干物质重、叶面积和比叶面积等叶片形态参数均极显著大于全光。推断地枫皮为耐阴性较弱的阳生植物,其光合能力和光饱和点较低,是对干旱环境的适应性反应;全光下地枫皮叶片狭小降低了吸光面积,有利于避免过高光强对叶光合器官的损伤。