Leaf anatomical responses to light in five tropical moraceae of different successional status

We characterized the leaf anatomical characteristics and maximum assimilation rates of five neotropical Moraceae of different genera and successional positions. Plants were grown under different light levels and transferred to high light, simulating canopy openings. Total blade thickness increased with irradiance among all species, and thicker blades were developed when plants were switched. However, blade thickness, and the extent to which it was modified, was independent of the species’ successional position and did not predict photosynthetic performance. Palisade thickness was a good predictor of maximum photosynthetic rate, but only on a species-specific basis. Overall, leaf thickening with increasing irradiance was associated more with structural than with photosynthetic changes. The early successionals Cecropia obtusifolia and Ficus insipida exhibited similarly high photosynthetic plasticity and acclimation values, but differed in their leaf anatomical traits. The late successional Poulsenia armata produced the most anatomically plastic leaves, but failed to acclimate either anatomically or photosynthetically when transferred to higher light levels.     

Steady and dynamic photosynthetic responses of seedlings from contrasting successional groups under low‐light growth conditions

To test the hypothesis that leaf‐level photosynthetic‐related traits might confer late successionals a competitive advantage over early successionals in low‐light growth conditions, steady photosynthetic assimilation‐ and dynamic photosynthetic induction‐related traits were examined in low‐light‐grown seedlings with contrasting successional status. Compared with the early successionals, late successionals as a group significantly exhibited lower leaf gas exchange rates. While late successionals required a longer time to respond to simulated sunflecks, they had lower rates of induction losses after sunflecks. Such photosynthetic induction traits allowed late successionals to more effectively utilize subsequent sunflecks. It was observed that plants with lower gas exchange rates responded more slowly to simulated sunfelcks, but they had lower rates of induction losses after sunflecks. In addition, the rate of response to sunflecks was positively correlated with the rate of induction loss after sunflecks across the successional status of species. A principal components analysis (PCA) demonstrated that early and late successionals were separated along the first axis of the PCA, and that early successionals were grouped on the right and were associated with higher gas exchange rates, fast responses to sunflecks, and rapid rates of induction loss after sunflecks; late successionals held an opposite pattern. Overall, our results suggest that smaller respiratory carbon losses and lower metabolic costs give late successionals a competitive advantage in low‐light growth conditions, that late successionals have an advantage over early successionals in utilizing sunflecks, and thus that the successional status of species are mainly associated with the leaf‐level photosynthetic‐related traits.     

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.     

Is hemiepiphytism an adaptation to high irradiance? Testing seedling responses to light levels and drought in hemiepiphytic and non‐hemiepiphytic Ficus

The epiphytic growth habit in many Ficus species during their juvenile stages has commonly been hypothesized to be an adaptation for avoiding deep shade in the forest understory, but this has never been tested experimentally. We examined growth and ecophysiology in seedlings of three hemiepiphytic (Hs) and three non‐hemiepiphytic (NHs) Ficus species grown under different irradiance levels. Both Hs and NHs exhibited characteristics of high light requiring species, such as high plasticity to growth irradiance and relatively high maximum photosynthetic assimilation rates. Diurnal measurements of leaf gas exchange showed that Hs have much shorter active photosynthetic periods than NHs; moreover, leaves of Hs have lower xylem hydraulic conductivity but stronger drought tolerance as indicated by much lower rates of leaf diebacks during the drought treatment. Seedlings of NHs had 3.3‐ and 13.3‐fold greater height and biomass than those of Hs species after growing in the nursery for 5 months, indicating a trade‐off between growth and drought tolerance due to the conflicting requirements for xylem conductivity and cavitation resistance. This study does not support the shade‐avoidance hypothesis; rather, it suggests that the canopy regeneration in Hs is an adaptation to avoid alternative terrestrial growth‐related risks imposed to tiny Ficus seedlings. The NHs with terrestrial regeneration reduce these risks by having an initial burst of growth to rapidly gain relatively large seedling sizes, while in Hs seedlings more conservative water use and greater drought tolerance for surviving the canopy environment are intrinsically associated with slow growth.     

Leaf ontogenetic dependence of light acclimation in invasive and native subtropical trees of different successional status

In the Bonin Islands of the western Pacific where the light environment is characterized by high fluctuations due to frequent typhoon disturbance, we hypothesized that the invasive success of Bischofia javanica Blume (invasive tree, mid‐successional) may be attributable to a high acclimation capacity under fluctuating light availability. The physiological and morphological responses of B. javanica to both simulated canopy opening and closure were compared against three native species of different successional status: Trema orientalis Blume (pioneer), Schima mertensiana (Sieb. et Zucc.) Koidz (mid‐successional) and Elaeocarpus photiniaefolius Hook.et Arn (late‐successional). The results revealed significant species‐specific differences in the timing of physiological maturity and phenotypic plasticity in leaves developed under constant high and low light levels. For example, the photosynthetic capacity of T. orientalis reached a maximum in leaves that had just fully expanded when grown under constant high light (50% of full sun) whereas that of E. photiniaefolius leaves continued to increase until 50 d after full expansion. For leaves that had just reached full expansion, T. orientalis, having high photosynthetic plasticity between high and low light, exhibited low acclimation capacity under the changing light (from high to low or low to high light). In comparison with native species, B. javanica showed a higher degree of physiological and morphological acclimation following transfer to a new light condition in leaves of all age classes (i.e. before and after reaching full expansion). The high acclimation ability of B. javanica in response to changes in light availability may be a part of its pre‐adaptations for invasiveness in the fluctuating environment of the Bonin Islands.     

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。     

热带雨林蒲桃属3个树种的幼苗光合作用对生长光强的适应

 测定了生长于3种光强下（100％、42％和14％自然光强） 热带雨林演替早期（思茅蒲桃Syzygium latilimbum）、中期（乌墨S.cumini）和后期（阔叶蒲桃S. szemaoense）出现的蒲桃属 （Syzygium）3个树种幼苗叶片气体交换参数和叶绿素荧光参数的日变化、比叶重和叶绿素含量。发现演替早期树种光合能力和光合可塑性最大、中期树种次之,后期树种光合能力最弱且在强光下受到显著抑制。但是,生长在强光下,3个树种均未发生长期光抑制和光破坏。随生长光强增加,3个树种热耗散速率都升高。不过在强光下,中期和后期树种通过热耗散消耗的过剩光能较多,通过叶黄素循环的热耗散可能对于这两个树种在强光下避免光合机构的光破坏起到了重要作用。14％光强下,3个树种最大净光合速率、光合作用的光饱和点、光补偿点、暗呼吸速率、比叶重降低,叶片单位干重叶绿素含量提高,对低光环境有了较好的形态学和生理学适应。但是,在4％光强下思茅蒲桃和乌墨幼苗全部死亡,只有阔叶蒲桃幼苗仍能存活,说明阔叶蒲桃幼苗适应弱光环境的能力高于思茅蒲桃和乌墨,与它们的演替状态一致。     

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。     

The effects of phenotypic plasticity on photosynthetic performance in winter rye, winter wheat and Brassica napus

The contributions of phenotypic plasticity to photosynthetic performance in winter (cv Musketeer, cv Norstar) and spring (cv SR4A, cv Katepwa) rye (Secale cereale) and wheat (Triticum aestivum) cultivars grown at either 20°C [non‐acclimated (NA)] or 5°C [cold acclimated (CA)] were assessed. The 22–40% increase in light‐saturated rates of CO₂ assimilation in CA vs NA winter cereals were accounted for by phenotypic plasticity as indicated by the dwarf phenotype and increased specific leaf weight. However, phenotypic plasticity could not account for (1) the differential temperature sensitivity of CO₂ assimilation and photosynthetic electron transport, (2) the increased efficiency and light‐saturated rates of photosynthetic electron transport or (3) the decreased light sensitivity of excitation pressure and non‐photochemical quenching between NA and NA winter cultivars. Cold acclimation decreased photosynthetic performance of spring relative to winter cultivars. However, the differences in photosynthetic performances between CA winter and spring cultivars were dependent upon the basis on which photosynthetic performance was expressed. Overexpression of BNCBF17 in Brassica napus generally decreased the low temperature sensitivity (Q₁₀) of CO₂ assimilation and photosynthetic electron transport even though the latter had not been exposed to low temperature. Photosynthetic performance in wild type compared to the BNCBF17‐overexpressing transgenic B. napus indicated that CBFs/DREBs regulate not only freezing tolerance but also govern plant architecture, leaf anatomy and photosynthetic performance. The apparent positive and negative effects of cold acclimation on photosynthetic performance are discussed in terms of the apparent costs and benefits of phenotypic plasticity, winter survival and reproductive fitness.     

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.     

Photosynthetic acclimation in eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings following transfer of shade-grown seedlings to high light

 We studied photosynthetic acclimation of eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings in the first month after sudden exposure of shade-grown seedlings to full sunlight. In a greenhouse experiment, seedlings were grown under full sun or 80% shade, and after 7 months, a sample of the shaded trees was transferred to full sun in the greenhouse. Photosynthetic responses of shaded, transferred, and sun trees were followed over the course of 26 days to track short to medium-term acclimation responses. A partial acclimation of photosynthesis at high light occurred in pre-existing (formed in the previous environment) and new foliage of transferred seedlings. This was associated with non-stomatal limitations to photosynthesis. Pre-existing foliage of transferred plants had a prolonged reduction in the ratio of variable to maximal fluorescence, and a limited capacity to adjust photochemical quenching or photosystem II quantum yield in the light to increasing light intensity compared to sun foliage, and apparently had some difficulty sustaining non-photochemical quenching. Seedling survival was only 58% among transferred seedlings, compared to 80% and 100% in the shade or sun groups, respectively. Photosystem II quantum yield in the light, and photochemical and non-photochemical quenching were similar between newly formed foliage of transferred and sun plants. These findings indicate that eastern hemlock depends strongly on the production of new foliage for photosynthetic adjustments to high light, and that development of photosynthetic competence may be a gradual process that occurs over successive foliar production cycles. Received: 12 May 1998 / Accepted: 27 July 1998     

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.     

Growth patterns of early versus late successional multipurpose trees of the western Himalaya

In the mid-western Himalaya (altitude 1350 m, rainfall 1100 mm), multipurpose trees found as escapees in agricultural fields or naturally growing in the forests, play an important role in providing fuel, fooder and small timber to the farmers. Shoot elogation, and tree architecture of 4 year old trees of Grewia optiva, Robinia pseudoacacia and Celtis australis (early successionals), and Quercus leucotrichophora, Q. glauca and Ilex odorata (late successionals), were analyzed. All the late successional species showed a proleptic type of bud and branch production, while the early successional trees made growth through syllepsis. The shoot elongation differed significantly (P <0.05) with the crown position, and ranged from 11 to 30 cm in different species. Early successional species tended to maintain a comparatively narrow crown and showed a significantly (P <0.05) higher ramification ratio, and multilayered canopy. The late successionals, in contrast, showed a wide crown with monolayered canopy, adapted to the weak light intensity. There was only one flush of leaves in Q. leucotrichophora and Q. glauca while in the rest of the species there were two distinct flush periods. The results are important for the management of agroforestry trees.     

The effect of light quantity and quality during development on the photosynthetic characteristics of six Australian rainforest tree species

Summary Seedlings of six subtropical rainforest tree species representing early (Omalanthus populifolius, Solanum aviculare), middle (Duboisia myoporoides, Euodia micrococca) and late (Acmena ingens, Argyrodendron actinophyllum) successional stages in forest development were grown in a glasshouse, under four levels of neutral shade (60%, 15%, 5%, 1% of photosynthetically active radiation (PAR) in incident sunlight) and three levels of selectively filtered shade (producing 15%, 5%, 1% of PAR). This design served to analyse the interactions between reduced photon flux density (PFD) and reduced red/far-red (R/FR) ratio in their effects on selected photosynthetic characteristics of each species. The light-saturated rate of photosynthesis was significantly influenced by growth irradiance in five of the six species, with all of these showing a non-linear decrease in maximum assimilation rate from 60% down to 1% PAR. The degree of acclimation to this range was not clearly related to the successional status of the species. Dark respiration was more sensitive to growth irradiance in the early- and mid-stage species than in the late-stage species. Although levels of dark respiration were clearly greater in leaves of early- and mid-stage species from the highest light levels, differences between successional groups were negligible at 1% PAR. Growth in filtered shade, typical of that beneath a closed canopy, resulted in lower photosynthetic capacities and quantum yields in those species which did respond. Although dark respiration rates were more sensitive to filtered shade in the early-stage than in the late-stage species, there was no evidence from other gas exchange characteristics to suggest that overall sensitivity to light quality (as characterised by the R/FR ratio) is greater in early successional-stage species.     

PHOTOPROTECTION OF SEA‐ICE MICROALGAL COMMUNITIES FROM THE EAST ANTARCTIC PACK ICE1

All photosynthetic organisms endeavor to balance energy supply with demand. For sea‐ice diatoms, as with all marine photoautotrophs, light is the most important factor for determining growth and carbon‐fixation rates. Light varies from extremely low to often relatively high irradiances within the sea‐ice environment, meaning that sea‐ice algae require moderate physiological plasticity that is necessary for rapid light acclimation and photoprotection. This study investigated photoprotective mechanisms employed by bottom Antarctic sea‐ice algae in response to relatively high irradiances to understand how they acclimate to the environmental conditions presented during early spring, as the light climate begins to intensify and snow and sea‐ice thinning commences. The sea‐ice microalgae displayed high photosynthetic plasticity to increased irradiance, with a rapid decline in photochemical efficiency that was completely reversible when placed under low light. Similarly, the photoprotective xanthophyll pigment diatoxanthin (Dt) was immediately activated but reversed during recovery under low light. The xanthophyll inhibitor dithiothreitol (DTT) and state transition inhibitor sodium fluoride (NaF) were used in under‐ice in situ incubations and revealed that nonphotochemical quenching (NPQ) via xanthophyll‐cycle activation was the preferred method for light acclimation and photoprotection by bottom sea‐ice algae. This study showed that bottom sea‐ice algae from the east Antarctic possess a high level of plasticity in their light‐acclimation capabilities and identified the xanthophyll cycle as a critical mechanism in photoprotection and the preferred means by which sea‐ice diatoms regulate energy flow to PSII.     

Acclimation responses to high light by Guazuma ulmifolia Lam. (Malvaceae) leaves at different stages of development

• The re‐composition of deforested environments requires the prior acclimation of seedlings to full sun in nurseries. Seedlings can overcome excess light either through the acclimation of pre‐existing fully expanded leaves or through the development of new leaves that are acclimated to the new light environment. Here, we compared the acclimation capacity of mature (MatL, fully expanded at the time of transfer) and newly expanded (NewL, expanded after the light shift) leaves of Guazuma ulmifolia Lam. (Malvaceae) seedlings to high light.
• The seedlings were initially grown under shade and then transferred to full sunlight. MatL and NewL were used for chlorophyll fluorescence and gas exchange analyses, pigment extraction and morpho‐anatomical measurements.
• After the transfer of seedlings to full sun, the MatL persisted and acclimated to some extent to the new light condition, since they underwent alterations in some morpho‐physiological traits and maintained a functional electron transport chain and positive net photosynthesis rate. However, long‐term exposure to high light led to chronic photoinhibition in MatL, which could be related to the limited plasticity of leaf morpho‐anatomical attributes. However, the NewL showed a high capacity to use the absorbed energy in photochemistry and dissipate excess energy harmlessly, attributes that were favoured by the high structural plasticity exhibited by these leaves.
• Both the maintenance of mature, photosynthetically active leaves and the production of new leaves with a high capacity to cope with excess energy were important for acclimation of G. ulmifolia seedlings.

     

Effects of nitrogen supply on photosynthetic and anatomical changes in current-year needles of <Emphasis Type="Italic">Pinus koraiensis</Emphasis> seedlings grown under two irradiances

We investigated the responses of photon-saturated photosynthesis rate (P _sat) and its simultaneous acclimation of anatomy and nitrogen use patterns of current needles of Korean pine (Pinus koraiensis) seedlings grown under factorial combinations of two nitrogen levels and irradiances. Although N supply resulted in a significant increase of N content in needles under both irradiances, the increase of P _sat tended to be suppressed only in shade (S). The significant increase of P _sat in full sunlight (O) was associated with the increase of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) and chlorophyll (Chl) contents. In contrast, small increase of Chl content and no increase of RuBPCO content were found in S (90 % cut of full irradiance), which would result in a small increase of P _sat. This result suggests that extra N is stocked in needles under shade for the growth in next season. With N supply, a significant decrease of specific leaf area (SLA) was detected only in O. This decrease of SLA was due to the increase of density of needle. Furthermore, the increase of needle density was not due to the increased number and size of mesophyll cells, but the increased density of each mesophyll cell. Therefore, although SLA changed in O, the change did not involve anatomical adaptation to use increased N effectively, at least observable by light microscopy. Hence, even though the SLA would change, N deposition will improve the photosynthetic capacity of Korean pine seedlings, not through the development of needle anatomy but through improvement of the allocation of N in both irradiances.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

Changes in pools of depsidones and melanins, and their function, during growth and acclimation under contrasting natural light in the lichen Lobaria pulmonaria

This study analysed relationships between secondary chemistry, lichen growth rates and external habitat factors for two groups of UV-B-absorbing secondary compounds in the lichen Lobaria pulmonaria in order to test some hypotheses on their formation and function. Medullary depsidones and cortical melanins were quantified in thalli transplanted to three successional forest stands (shaded young forest, open old forest, sun-exposed clear-cut area) and subjected to different watering regimes (spraying with water, water + nitrogen, no spraying). Growth rates were already known. The total concentration of all seven depsidones was constant across the entire range of growth rates and sun exposures, showing that these depsidones serve functions other than photoprotection. Thalli from the well-lit transplantation sites had the highest synthesis of melanins. Within each forest type there was a trade-off between growth and melanin synthesis. Melanins and photosynthetic acclimation enhanced survival on a subsequent exposure to high light intensity, despite excessive temperatures resulting from higher absorption of solar energy in melanic thalli relative to pale thalli. In conclusion, the highly responsive melanic pigments play a photoprotective role in light acclimation, whereas the constant amount of depsidones across a wide spectrum of growth ranges and irradiances is consistent with herbivore defence functions.     

Plastic responses of 4 tree species of successional subalpine coniferous forest serals to different light regimes

The morphological and physiological plasticity to 6 light conditions was investigated for seedlings of 4 tree species dominating at different successional serals in subalpine coniferous forests in eastern Qinghai-Tibet Plateau, China. Abies faxoniana is a late successional species, while Betula albo-sinensis is a pioneer tree species, with Picea asperata and Acer davidii among other mid-late successional species. To compare the responses of photosynthetic characters to different growth light conditions, the seedlings were potted and placed in artificially shaded chambers with gradients of 100%, 55%, 40%, 25%, 15% and 7% of the full sunlight, respectively. During two and a half years' cultivation, various morphological and photosynthetic parameters were measured and analyzed. The results were: 1) all seedlings of the four species under the low growth light conditions showed decrease in root collar diameter, relative growth rate, leaf thickness, root mass ratio, leaf area-based photosynthetic capacity, dark respiration rate, light saturation point and light compensation point, while showed increase in specific leaf area, above-ground to under-ground mass ratio, specific stem length, leaf mass ratio and stem mass ratio; 2) under most light conditions, A. faxoniana of the two conifers revealed lower values in both leaf area-based photosynthetic capacity and respiration rates than P. asperata; the same fact held true for the two broad-leaved trees with lower values observed in B. albo-sinensis; 3) under higher light conditions, relative growth rates of P. asperata and B. albo-sinensis got higher values than those of A. faxoniana and A. davidii, while contrary results were obtained under lower light regimes; 4) the means of phenotypic plastic indices of the eleven morphological and physiological parameters of P. asperata and B. albo-sinensis were higher than those of A. faxoniana and A. davidii, respectively. The findings indicate that A. faxoniana has better adaptation to low light regimes, but as a shade-tolerant species, it is not so adapted to low light regimes as the trees of earlier successional serals, especially P. asperata and B. albo-sinensis. Of the four tree species, physiological plastic indices were higher than morphological plastic indices, suggesting that morphological plasticity plays an important role in their adaptation to different growth light conditions. The results also support the hypothesis that ecophysiological traits of tree species determine their successional status and associate habitats of their seedlings.