Expansion Kinematics are an Intrinsic Property of Leaf Development and are Scaled from Cell to Leaf Level at Different Nutrient Availabilities

Abstract: Leaves of Nicotiana tabacum L. and Ricinus communis L. develop with a wide range of different final sizes due to leaf position and nutrient availability. The aim of this study was to investigate, on different organizational levels of leaf growth, which parameters are affected by external nutrient availability and which parameters are not affected and might thus be intrinsic, general patterns of growth that govern plant architecture. We found that leaf size and final cell size was larger with higher external nutrient availability, and that hexose concentrations in N. tabacum were lower with higher nutrient availability. Despite these differences, several dynamic parameters of leaf development were not affected by the nutrient treatment. Leaves of all sizes within a species exhibited the same relationship between relative leaf growth rate and relative leaf area (RLA), which is defined as the ratio of momentary and final leaf area. External nutrient availability did not affect chlorophyll concentration per parenchyma cell, which increased linearly with leaf development. Leaves of identical RLA exhibited identical cell density patterns within their interveinal tissue layers. This indicates a close connection between the kinematics of cell expansion and RLA and, hence, reveals that kinematics are an intrinsic property of growing leaves that can be scaled from the cell to the leaf level.     

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.     

不同光照梯度的遮荫处理对绒毛番龙眼幼苗生长的影响

在不同光照梯度,即100%自然全光照(natural sunlight,NS)、37.3%NS、15.5%NS、4.2%NS、1.6%NS和0.6%NS的人工遮荫条件下,研究了西双版纳季节雨林冠层树种绒毛番龙眼(Pometia tomentosa)幼苗的早期生长和定居后的生长特点。结果表明,光照是影响幼苗生长的重要环境因子。生长早期的幼苗基径和复叶数随遮荫程度的增加而降低;主根长、根冠比、总干重和单株叶面积均以37.3%NS处理最大;比叶面积随遮荫程度的增加而增大,而相对生长率则降低;幼苗株高在0.6%NS下增长最快,表明种子中贮藏的营养物质对幼苗的早期生长可能具有重要作用。37.3%NS处理对定居后绒毛番龙眼幼苗的生长最有利,幼苗的株高、基径、复叶数、叶轴长、复叶最多小叶数、单株叶面积、相对生长率和净同化率均在37.3%NS处理下获得最大增长;幼苗总干重随光照强度的减弱而降低;比叶面积在15.5%NS处理时最大。幼苗比叶面积和根冠比在生长过程中的波动可能是光照和土壤水分共同作用的结果。     

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。     

Genetic modification of photosynthesis with E. coli genes for trehalose synthesis

Improvement in photosynthesis per unit leaf area has been difficult to alter by breeding or genetic modification. We report large changes in photosynthesis in Nicotiana tabacum transformed with E. coli genes for the trehalose pathway. Significantly, photosynthetic capacity (CO2 assimilation at varying light and CO2, and quantum yield of PSII electron transport) per unit leaf area and per leaf dry weight were increased in lines of N. tabacum transformed with the E. coli gene otsA, which encodes trehalose phosphate synthase. In contrast, transformation with otsB, which encodes trehalose phosphate phosphatase or Trec, encoding trehalose phosphate hydrolase, produced the opposite effect. Changes in CO2 assimilation per unit leaf area were closely related to the amount and activity of Rubisco, but not to the maximum activities of other Calvin cycle enzymes. Alterations in photosynthesis were associated with trehalose 6-phosphate content rather than trehalose. When growth parameters were determined, a greater photosynthetic capacity did not translate into greater relative growth rate or biomass. This was because photosynthetic capacity was negatively related to leaf area and leaf area ratio. In contrast, relative growth rate and biomass were positively related to leaf area. These results demonstrate a novel means of modifying Rubisco content and photosynthesis, and the complexities of regulation of photosynthesis at the whole plant level, with potential benefits to biomass production through improved leaf area.     

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients?Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (M D), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-to-shoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.Important findings Significant interactions for irradiance and N availability for majority of growth attributes indicates that growth and biomass allocation of seedlings were more responsive to N availability under high irradiance. However, species responded differentially to N addition and they did not follow successional status. Slow growers (B. retusa, a shade-tolerant species and L. parviflora, a light demander) exhibited greater response to N enrichment than the fast growers (A. catechu, D. sissoo and T. arjuna). However, N-mediated increment in growth traits was greater in non-legumes (B. retusa, L. parviflora and T. arjuna) compared with that of legumes (A. catechu and D. sissoo). Allocation of biomass to root was strongly suppressed at the highest N supply across species; however, at high irradiance and high N availability, a greater suppression in R:S ratio was observed for B. retusa. NAR was a stronger determinant of RGR relative to LAR, suggesting its prominent role in increased RGR along increasing irradiances. Overall, a higher growth response of slow-growing species to elevated N levels, particularly the non-pioneers (B. retusa and L. parviflora) suggests that future N deposition may lead to perturbations in competition hierarchies and species composition, ultimately affecting community dynamics in nutrient-poor tropical dry forests.     

INTERSPECIFIC VARIATION IN GROWTH,BIOMASS PARTITIONING,AND DEFENSIVE CHARACTERISTICS OF NEOTROPICAL MANGROVE SEEDLINGS: RESPONSE TO LIGHT AND NUTRIENT AVAILABILITY

Seedlings of three mangrove species—Rhizophora mangle L., Avicennia germinans (L.)Stearn., and Laguncularia racemosa (L.) Gaertn. f.—were subjected to different light and nutrient regimes in two separate growth chamber experiments. At higher nutrient or light availability, relative growth rate, leaf production, and branch growth differed significantly among species in the following order: Rhizophora < Avicennia < Laguncularia. At lower nutrient or light levels, however, species’ differences were greatly minimized. Lower nutrient or light levels caused greater investment in root biomass by all species, whereas higher nutrient availability resulted in greater investment in leaf area and maximized species’ differences in total leaf area, number of leaves, and leaf area ratio. Mangrove leaves also differed among species in quantity and composition of secondary compounds that may protect seedlings against herbivores or stress factors such as excessive solar radiation. Relative amounts of condensed tannins, gallotannins, and nitrogen were significantly affected by light and nutrient regimes, but patterns of response differed among species. The results indicate that these sympatric species differ substantially in their potential for growth, acquisition of resources, stress tolerance, and susceptibility to herbivores during the seedling stage, but that these characteristics are significantly modified by availability of resources.     

Growth and nutrient dynamics of beech (Fagus sylvatica L.) seedlings in acid soils

The relative uptake rates of N, P, K, S, Ca, Mg, Fe, Mn, Zn, Cu, and Al were estimated in beech seedlings pot cultured in the field in six acid soils (treatments). The relative uptake rates were compared with the relative growth rates. The relative uptake rates of N, K and Ca agreed well with the growth rates of the seedlings irrespective of widely differing soil conditions (acid sand-clayey till, pH 4–6). The relative uptake rates of P, Fe, and Al differed from that predicted by the growth rate. The uptake rates of Fe and Al were highest at the lowest growth rates, and the P uptake rate was lower than the growth rate in these treatments. Thus the P availability probably limited growth in an eluvial (E) horizon of a podzol, and possibly in the illuvial (B) horizon of a podzol and in an acid clayey till (Dystric Cambisol). Low P uptake was associated with a tendency towards higher relative root growth rates. In terms of the concept of steady state nutrition the high relative root growth rate in some treatments may be interpreted as an acclimation to low P supply. The P limitation seemed to be related to interactions among Fe, Al and organic compounds of the soil solution.FAX no: +4646104423     

Response of Growth Dynamics of Two Spring Geophytes to Light Regime in a Lime-Beech Forest

Blomaee accumulation, leaf longevity and growth rate of two spring forest geophytes, Scllla blfolla L. and Arum maculatum L. were estimated separately for three size groups within each population of these species. Despite the differences in leaf longevity, both species showed a similar pattern of blomass accumulation In relation to their phenologles and reproductive demands. Eco-physlological acclimation to changing light environment was assumed through photosynthetic parameters and dynamics of leaf area Index In the predominant size group of each species. A light response curve was measured under natural light for each species through the continuum of Its phenology to quantify the photosynthetic photon flux density at light saturation, light-saturated photosynthetic rate, light compensation point, and dark respiration. Light-saturated assimilation per leaf area basis, dark respiration rate and light compensation points were significantly higher in S. blfolla relative to A. maculatum. However, the acclimation of photosynthesis that would respond to light changes in environment was not found in S. bifolla. In contrast, In A. maculatum a marked shift In the light dependence of photosynthesis through the season was noticed, which resulted In a strong photosynthetic acclimation to the low-light conditions. Accompanied by significant leaf area Index, this efficient low-light photosynthesis enabled greater leaf longevity, and consequently longer accumulative period to A. maculatum. From the different parameters that we determined （both photosynthetic acclimation and growth strategy） it would appear that these species belong to two distinct subgroups： S. blfolla to the early and A. maculatum to the late vernals.     

Effects of a 60 Hz magnetic field on photosynthetic CO2 uptake and early growth of radish seedlings

Photosynthetic CO2 uptake rate and early growth parameters of radish Raphanus sativus L. seedlings exposed to an extremely low frequency magnetic field (ELF MF) were investigated. Radish seedlings were exposed to a 60 Hz, 50 microT(rms) (root mean square) sinusoidal magnetic field (MF) and a parallel 48 microT static MF for 6 or 15 d immediately after germination. Control seedlings were exposed to the ambient MF but not the ELF MF. The CO2 uptake rate of ELF MF exposed seedlings on day 5 and later was lower than that of the control seedlings. The dry weight and the cotyledon area of ELF MF exposed seedlings on day 6 and the fresh weight, the dry weight and the leaf area of ELF MF exposed seedlings on day 15 were significantly lower than those of the control seedlings, respectively. In another experiment, radish seedlings were grown without ELF MF exposure for 14 d immediately after germination, and then exposed to the ELF MF for about 2 h, and the photosynthetic CO2 uptake rate was measured during the short-term ELF MF exposure. The CO2 uptake rate of the same seedlings was subsequently measured in the ambient MF (control) without the ELF MF. There was no difference in the CO2 uptake rate of seedlings exposed to the ELF MF or the ambient MF. These results indicate that continuous exposure to 60 Hz, 50 microT(rms) sinusoidal MF with a parallel 48 microT static MF affects the early growth of radish seedlings, but the effect is not so severe that modification of photosynthetic CO2 uptake can observed during short-term MF exposure.     

两种光强下木质藤本与树木幼苗的竞争关系

 为了探讨木质藤本和树木幼苗的相互作用关系,对两种光强(4%和35%的光强)、4种竞争处理下(全竞争、地上竞争、地下竞争和无竞争),一种 需光木质藤本(刺果藤(Byttneria grandifolia))和3个树种(耐荫种：五桠果木姜子(Litsea dilleniifolia)和绒毛番龙眼(Pometia tomentosa);需光种：羊蹄甲(Bauhinia variegata))幼苗的地上部分和地下部分的竞争关系进行了研究。结果表明：木质藤本的竞争显著影响 着3种树木幼苗的光合能力、形态特征和生长,但生长环境的不同光强影响地上部分竞争和地下部分竞争的相对强度。在低光下,地上部分竞争 比地下部分竞争对3种树木幼苗的相对生长速率(Relative growth rate,RGR)和光合能力造成更大的影响;而高光下,地下竞争对树木幼苗的 生长有更强的抑制作用。不同的竞争处理和光强对树木幼苗的生物量积累造成显著的影响。光强对3种树种的比叶面积(Specific leaf area, SLA)和叶面积比(Leaf area ratio,LAR)有显著的抑制作用,但竞争只对需光的羊蹄甲的SLA和LAR有显著影响。不同的光照和竞争处理之间, 同种植物表现出不同的表型特征。由于竞争的影响,苗木在形态上较为矮小、叶片数目较少、叶面积减小,但是长细比改变较少 。     

Effects of above- and below-ground competition from shrubs on photosynthesis, transpiration and growth in Quercus robur L. seedlings

For a tree seedling to successfully establish in dense shrubbery, it must maintain function under heterogeneous resource availability. We evaluated leaf-level acclimation in photosynthetic capacity, seedling-level transpiration, and seedling morphology and growth to gain an understanding of the effects of above- and below-ground competition on Quercus robur seedlings. Experimental seedlings were established in a typical southern Swedish shrub community where they received 1 of 4 competition levels (above-ground, below-ground, above- and below-ground, or no competition), and leaf-level responses were examined between two growth flushes. Two years after establishment, first-flush leaves from seedlings receiving above-ground competition showed a maximum rate of photosynthesis (A_max) 40% lower than those of control seedlings. With the development of a second flush above the shrub canopy, A_max of these seedlings increased to levels equivalent to those of seedlings free of light competition. Shrubby competition reduced oak seedling transpiration such that seedlings exposed to above- and below-ground competition showed rates 43% lower than seedlings that were not exposed to competition. The impaired physiological function of oak seedlings growing amid competition ultimately led to a 60-74% reduction in leaf area, 29-36% reduction in basal diameter, and a 38-78% reduction in total biomass accumulation, but root to shoot ratio was not affected. Our findings also indicate that above-ground competition reduced A_max, transpiration and biomass accumulation more so than below-ground competition. Nevertheless, oak seedlings exhibited the ability to develop subsequent growth flushes with leaves that had an A_max acclimated to utilize increased light availability. Our findings highlight the importance of flush-level acclimation under conditions of heterogeneous resource availability, and the capacity of oak seedlings to initiate a positive response to moderate competition in a shrub community.     

Development and plasticity of endangered shrub Lindera melissifolia (Lauraceae) seedlings under contrasting light regimes

Lindera melissifolia (Walt.) Blume seedlings were raised in a growth chamber to determine the effects of light availability on shoot growth pattern, and basic leaf and stem growth. Lindera melissifolia seedlings exhibited a sympodial shoot growth pattern for 3 months following emergence from the soil medium, but this pattern was characterized by a reduction in leaf blade area approximately 30 days after emergence, followed by increases in leaf blade area. Seedlings receiving low light were 76% taller than seedlings receiving high light. Seedlings receiving low light also had larger leaf blade dimensions, blade area, seedling leaf area, and greater mass. Seedlings raised in high light had a greater proportional distribution of biomass in the roots, suggesting possible water stress from greater vapor pressure deficits. Furthermore, these seedlings displayed sharp angles of blade inclination and blade folding – acclimation that reduces exposure to light and subsequent higher leaf temperatures in open environments. These differences in morphological response to light resulted in high phenotypic variability in L. melissifolia seedlings. Lindera melissifolia seedling development showed a brief period of phenotypic plasticity, followed by ontogenetic plasticity. The short period of phenotypic plasticity may, however, have profound ecological implications for the conservation and recovery of this federally endangered shrub. Further experimentation should take into account the development of ontogenetic standards for comparisons of plant traits in addition to temporal standards.     

Dependence of starch storage on nutrient availability and photon flux density in small birch Betula pendula Roth)

Abstract Small birch plants (Betula pendula Roth) were grown in a climate chamber at different levels of nutrient availability and at two photon flux densities. The extent to which starch storage was dependent upon nutrient availability and photon flux density was investigated. Acclimated values of starch concentration in leaves were highest at low nutrient availability and high photon flux density. Starch storage in roots was only found at the lowest nutrient availability. However, the relative rate of starch storage (starch stored per unit plant dry weight and time) was higher in plants with good nutrition. The data suggest that, at sub-optimal nutrient availability, the momentary rate of net shoot photosynthesis is unlikely to limit the structural (as opposed to carbon storage) growth of the plant. Although photosynthetic rate per unit leaf area (as measured at the growth climate) was slightly lower in plants with poor nutrient availability, photosynthetic rate per unit leaf nitrogen was higher. These data suggest a priority of leaf nitrogen usage in photosynthesis, with limiting amounts of leaf nitrogen (and possibly other nutrients) for subsequent growth processes. This argument is consistent with the higher concentrations of starch found in plants with poor nutrient availability.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

光照对鄂东南2种落叶阔叶树种幼苗生长、光合特性和生物量分配的影响

通过设置4个光照梯度(25%、12%、6%和3%自然光)模拟鄂东南低山丘陵地区落叶阔叶林林下的光环境,研究了2种耐荫性不同的树种幼苗--麻栎(Quercus acutissima)和化香(Platycarya strobilacea)不同光强下的存活率、光合特性、生长和生物量分配,探讨了低光环境中耐荫性不同的树种幼苗维持自身碳平衡的机制和权衡"存活-生长"选择的生活史策略。结果表明:(1)低光下的2个树种幼苗的生长、光合特性和生物量分配具有显著性差异。(2)各个光照梯度下麻栎幼苗都生长良好,存活率保持在35%以上,而化香幼苗遭遇高的死亡率,80d后3%和6%自然光下的幼苗全部死亡;低光环境中麻栎幼苗比化香幼苗具有更大的表观光量子(AQY)和最大净光合效率(Pmax),更低的光补偿点(LCP)和暗呼吸效率(Rd),即耐荫性较强的麻栎幼苗比耐荫性较弱的化香幼苗具有更高的低光碳同化率和碳捕获能力。(3)2个树种幼苗的成活率与RGR呈负相关关系,各个光照梯度下耐荫性较弱的化香幼苗的相对生长率(RGR)显著高于耐荫性较强的麻栎幼苗,而两个树种幼苗的净同化率(NAR)无明显差异。相对于麻栎幼苗较高的根生物量比(RMR),化香幼苗将更多的生物量分配给叶部,因而具有较高的叶生物量比(LMR)、叶面积比(LAR)和比叶面积(SLA)。不同耐荫性的幼苗生长及生物量分配方式的差异是植物"存活-生长"权衡后的结果,耐荫性弱的化香幼苗具有较高的生长潜力和较弱的自我保护能力,而耐荫性强的麻栎幼苗具有更高的低光碳储量,能够维持更好的低光碳平衡,具有竞争优势。     

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。     

Combined effects of light and water availability on photosynthesis and growth of Arisaema heterophyllum in the forest understory and an open site

Photosynthetic characteristics, leaf longevity and biomass accumulation of a threatened herb species, Arisaema heterophyllum, were studied in the understory of a riparian forest and at a neighboring deforested open site for 3 years in order to understand the combined effects of light and water availability. Light availability was 2- to 4-fold higher at the deforested than at the forest site during the growing season of the species, and precipitation varied considerably over the 3 years. Despite the difference in water availability among the years (dry in 1994 and 1996, and wet in 1995), the species showed a strong acclimation to the different light environments. Light-saturated assimilation rate on a leaf area basis, leaf mass area ratio (LMA), and relative growth rate (RGR) were all higher at deforested site. While a positive correlation between individual RGR and microsite light availability was found in the wet year, no correlation was found in the dry years, and mean RGR was significantly lower in the dry year at both sites. Leaf longevity, photosynthetic capacity on a leaf mass basis, dark respiration rate, and leaf conductance, varied considerably from year to year, especially in the plants at the open site, probably depending on water availability. In the dry years plants at the deforested sites showed a lower photosynthetic rate and leaf conductance under unwatered than under watered conditions. These results suggest that the water availability in a given year may strongly affect light acclimation and annual RGR of the herb species in natural habitats, even under mesic climate conditions. Received: 15 February 1997 / Accepted: 20 May 1997     

Effect of two natural light regimes and nutrient addition on the forest herb Begonia decandra (Begoniaceae)

The effect of two natural light-growing conditions (understory versus light gaps) and the interaction with nutrient availability (through fertilization) were studied in the understory herb Begonia decandra, in the Luquillo Experimental Forest in Puerto Rico. Sixteen potted plants obtained from cuttings were randomly chosen and distributed in each of eighth forest environments (four light gaps and four understories), for a total of 128 plants. Fertilizer was applied to half of the plants in each site. After seven months in the two given microenvironments, increased light and fertilization resulted in greater growth and some changes in the biomass allocation patterns. All measured variables responded similarly to reported changes for tree seedlings and saplings from other tropical and subtropical regions. Total growth parameters (height, biomass and leaf area) were very sensitive to increases in the main resource (light). The addition of nutrients was less important in producing changes in the allocation variables (root to shoot ratio, leaf area ratio, and specific leaf mass) under conditions of high light availability. Changes due to nutrient levels were relatively greater on plants grown under under-story conditions. Also, small light differences among sites can cause significant changes in the variables related to total growth. Lastly, plant mortality in the nutrient treatments was found to be independent of mortality in two forest light environments. Some hypotheses about resource acquisition and plant growth are not supported by this data.