QUANTITATIVE PHENOLOGY AND LEAF SURVIVORSHIP OF RHODODENDRON MAXIMUM IN CONTRASTING IRRADIANCE ENVIRONMENTS OF THE SOUTHERN APPALACHIAN MOUNTAINS

Seasonal plant growth patterns were compared for Rhododendron maximum L. in two contrasting subcanopy environments. The two subcanopy, above ground environments differed only in their quantity of irradiance by virtue of the relative dominance of evergreen or deciduous trees in the canopy. A third site had no canopy influence. Overall growth (shoot elongation, woody increment, leaf production) was maximized under the open (BMO) and deciduous dominated canopy (PCD). The leaf pool was significantly smaller under the evergreen dominated canopy (PCE) but average leaf area per leaf was slightly larger at PCE. Individual age-specific leaf cohorts, identified from shoot morphology, indicated increased leaf survivorship with a decreased irradiance environment. Leaf production was synchronous and rapid (1 week), followed by three weeks of leaf expansion, which created the even-aged leaf cohorts. Wood growth (diameter increment), in contrast, continued through the beginning of the winter. Reproductive effort increased with increasing irradiance environment. Significant variation in growth was observed between canopy shoot types at all three research sites. The significance of these phenological patterns is discussed in view of the variable subcanopy environment of southwestern Virginia.     

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST AND SOUTHWEST SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. I. IRRADIANCE TOLERANCE

Deciduous and evergreen species are segregated on northeast and southwest slopes of the southern Appalachian Mountains. The segregated distributions of three ericaceous shrubs (Rhododendron maximum valley positions; Rhododendron periclymenoides on northeast slopes; Kalmia latifolia on southwest slopes) were compared to the respective irradiance environments. Growth patterns of field plants, and photosynthetic acclimation of each species to three irradiance treatments in a phytotron were studied. Rhododendron maximum, an evergreen species, was found to be most sensitive to high radiation. In phytotron experiments, quantum yield, light saturated photosynthetic capacity, photosynthesis per chlorophyll, and water use efficiency decreased at high ambient irradiance for R. maximum. These characteristics limit the growth of R. maximum on high irradiance southwestern slopes. Both K. latifolia and R. periclymenoides were able to improve their photosynthetic performance at high ambient irradiance. Rhododendron periclymenoides, a deciduous species, was found to continue increasing leaf conductance at high irradiance without an increase in photosynthesis indicating a possible limitation by water in high light environments such as southwest slopes. Kalmia latifolia, an evergreen species, had reduced photosynthetic capacity and reduced water use efficiency when grown in low irradiance conditions which coincides with the higher K. latifolia abundance on high light, southwestern slopes.     

Influence of water relations and temperature on leaf movements of rhododendron species

Rhododendron maximum L. and R. Catawbiense L. are subcanopy evergreen shrubs of the eastern United States deciduous forest. Field measurements of climate factors and leaf movements of these species indicated a high correlation between leaf temperature and leaf curling; and between leaf water potential and leaf angle. Laboratory experiments were performed to isolate the influence of temperature and cellular water relations on leaf movements. Significant differences were found between the patterns of temperature induction of leaf curling in the two species. Leaves of the species which curled at higher temperatures (R. catawbiense) also froze at higher leaf temperatures. However, in both cases leaf curling occurred at leaf temperatures two to three degrees above the leaf freezing point. Pressure volume curves indicated that cellular turgor loss was associated with a maximum of 45% curling while 100% or more curling occurred in field leaves which still had positive cell turgor. Moisture release curves indicated that 70% curling requires a loss of greater than 60% of symplastic water which corresponds to leaf water potentials far below those experienced in field situations. Conversely, most laboratory induced changes in leaf angle could be related to leaf cell turgor loss.     

The relationship between freezing tolerance and thermotropic leaf movement in five Rhododendron species

Summary Leaf movement kinetics in five species of Rhododendron were studied in response to leaf temperature, leaf freezing point, and leaf water deficit. There was a gradient in the degree of leaf curling among species in the following order from the greatest curling to the least curling: Rhododendron catawbiense, R. maximum, R. minus, R. macrophyllum, R. ponticum. Those species found to be tolerant of winter conditions had the most intense leaf movements (both curling and angle) while those species with minimal cold tolerance had limited or no leaf movements. Leaf curling occurred at leaf temperatures above the tissue freezing points in all species. Athough leaf angle was influenced by leaf turgor, general tissue desiccation was not the ultimate cause for thermotropic leaf curling in any species tested. Those species with the greatest leaf curling and angle movements had the highest osmotic potential, the lowest water deficit at the turgor loss point, and the lowest symplastic water fraction. These data suggest that there is a trade off in Rhododendron leaf physiology between cold tolerance (due to leaf movements) and water stress tolerance (due to turgor maintenance mechanisms).     

Thermonastic leaf movements: a synthesis of research with Rhododendron

Thermonastic leaf movements in Rhododendron L. occur in response to freezing temperatures. These movements are composed of leaf curling and leaf angle changes that are distinct leaf movements with different responses to climatic factors. Leaf angle is controlled by the hydration of the petiole, as affected by soil water content, atmospheric vapour pressure, and air temperature. In contrast, leaf curling is a specific response to leaf temperature, and bulk leaf hydration has little effect. The physiological cause of leaf curling is not well understood, but the mechanism must lie in the physiology of the cell wall and/or regional changes in tissue hydration. Available evidence suggests that intercellular freezing is not a cause of leaf curling.
Manipulation experiments demonstrate that changes in leaf orientation in Rhododendron most likely serve to protect the leaves from membrane damage due to high irradiance and cold temperatures. In particular, the pendent leaves protect the chloroplast from photoinhibition. Leaf curling may serve to slow the rate of thaw following freezing, a common phenomenon in the Appalachian mountains of the U.S. The thermonastic leaf movements have a greater importance to plants in a dim environment because the potential impact to canopy carbon gain is greater than in high light environments.
These leaf movements have several implications for horticultural management. There seems to be a trade-off between water stress tolerance and freezing stress tolerance by leaf movements. Thermonastic leaf movements may be a major mechanism of cold stress tolerance in Rhododendron species. The actual physiological cause of leaf movement has not been elucidated and many more species need to be evaluated to verify the general importance of leaf movements to Rhododendron ecology and evolution.     

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST- AND SOUTHWEST-FACING SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. II. WATER RELATIONS

Three species of shrubs (Ericaceae) were found to segregate upon the northeast and southwest slopes of spur ridges on Brush Mountain, in southwestern Virginia. Rhododendron maximum was found only in valleys and lower northeast slopes, Rhododendron periclymenoides = R. nudiflorum) was found on northeast slopes while Kalmia latifolia was most abundant on southwest slopes. Previous vegetation studies indicated that these partially segregated distributions were related to irradiance and water availability. In field studies of water potential, R. periclymenoides had the lowest diurnal leaf water potentials and the largest seasonal variation in midday leaf water potential. Kalmia latifolia had the highest leaf conductance in field and phytotron experiments. Rhododendron maximum had the greatest seasonal osmotic adjustment followed by R. periclymenoides and K. latifolia. In phytotron experiments, the photosynthetic capacity of R. maximum was the most sensitive to water stress followed by R. periclymenoides and K. latifolia. Kalmia latifolia was able to modify its conductance rates to reduce water loss and maintain constant leaf water potential minimizing photosynthetic inhibition. Rhododendron periclymenoides showed extreme luxury spending of water indicated by high conductance and low photosynthesis. The ecophysiological responses to water and irradiance provided an explanation for the distributions of the three species. For example, R. maximum leaves are sensitive to elevated irradiance, and carbon gain is strongly influenced by water stress. Thus, R. maximum will perform best in low irradiance environments with ample water, such as valley sites. Each species had a unique set of adaptations for performing best in their optimum habitat.     

Plant diversity patterns in semi-natural grasslands along an elevational gradient in southern Norway

Leaf demography and growth of six common, co-occurring woody plant species that varied in stature (tree vs. shrub) and leaf texture (sclerophyllous, coriaceous, malacophyllous) were examined in a subtropical savanna parkland in southern Texas, USA. We tested the hypotheses that, (a) leaves of plants with evergreen canopies would have longer life spans than those of deciduous species; (b) supplementation of soil moisture would decrease leaf life span in both evergreen and deciduous species; (c) species responses to increased soil moisture availability would be inversely related to leaf longevity; and (d) deciduous growth forms would exhibit a greater growth response to increased soil moisture availability than their evergreen counterparts.A variety of seasonal leaf habits (evergreen, winter-deciduous and summer-deciduous canopies) and leaf life spans (median = 66 to 283 days) were represented by the targeted species, but there was no clear relationship between seasonal leaf habit and leaf longevity. Among species with evergreen canopies, median leaf longevity ranged from short (Zanthoxylum fagara = 116 days; Condalia hookeri = 158 days) to long (Berberis trifoliolata = 283 days) but did not exceed 1 yr. In fact, leaf longevity in evergreen shrubs was often comparable to, or shorter than, that of species with deciduous canopies (Ziziphus obtusifolia = 66 days; Diospyros texana = 119 days; Prosopis glandulosa = 207 days). Augmentation of surface soil moisture had no detectable effect on median leaf life span in any species and there was no clear relationship between leaf longevity and species growth responses to irrigation. Contrary to expectations, species with evergreen canopies responded to irrigation by producing more leaf biomass, longer shoots and more leaf cohorts/year than did deciduous species.Species differences in the annual cycle of leaf initiation, leaf longevity and canopy development, combined with contrasts in root distributions and a highly variable climate, may allow for spatial and temporal partitioning of resources and hence, woody species coexistence and diversity in this system. However, the lack of expected relationships between leaf longevity, leaf habit and plant responses to resource enhancement suggests that structure-function relationships and functional groupings developed in strongly seasonal environments cannot be applied with confidence to these subtropical savannas and thorn woodlands.     

The responses of light interception,photosynthesis and fruit yield of cucumber to LED‐lighting within the canopy

Mathematical models of light attenuation and canopy photosynthesis suggest that crop photosynthesis increases by more uniform vertical irradiance within crops. This would result when a larger proportion of total irradiance is applied within canopies (interlighting) instead of from above (top lighting). These irradiance profiles can be generated by Light Emitting Diodes (LEDs). We investigated the effects of interlighting with LEDs on light interception, on vertical gradients of leaf photosynthetic characteristics and on crop production and development of a greenhouse‐grown Cucumis sativus‘Samona’ crop and analysed the interaction between them. Plants were grown in a greenhouse under low natural irradiance (winter) with supplemental irradiance of 221 µmol photosynthetic photon flux m^?2 s^?1 (20 h per day). In the interlighting treatment, LEDs (80% Red, 20% Blue) supplied 38% of the supplemental irradiance within the canopy with 62% as top lighting by High‐Pressure Sodium (HPS)‐lamps. The control was 100% top lighting (HPS lamps). We measured horizontal and vertical light extinction as well as leaf photosynthetic characteristics at different leaf layers, and determined total plant production. Leaf mass per area and dry mass allocation to leaves were significantly greater but leaf appearance rate and plant length were smaller in the interlighting treatment. Although leaf photosynthetic characteristics were significantly increased in the lower leaf layers, interlighting did not increase total biomass or fruit production, partly because of a significantly reduced vertical and horizontal light interception caused by extreme leaf curling, likely because of the LED‐light spectrum used, and partly because of the relatively low irradiances from above.     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。     

Aspects of tissue water relations and seasonal changes of leaf water potential components of evergreen and deciduous species coexisting in tropical dry forests

Summary This study compared the tissue water relations and seasonal changes in leaf water potential components of an evergreen tree,Morisonia americana, and two evergreen shrubs,Capparis verrucosa andC. aristiquetae, with two deciduous trees,Humboltiella arborea andLonchocarpus dipteroneurus, and the deciduous vineMansoa verrucifera. All these species coexist in a tropical dry forest in Venezuela. Leaves of the evergreen species are sclerophyllous, while those of the deciduous species are mesophytic. Leaf area to leaf weight ratios of fully mature leaves were about 75 and 170 cm² g^–1 in evergreen and deciduous species, respectively. Seasonal fluctuations of leaf water content per unit of dry weight, water potential, and turgor pressure were smaller in evergreen than in deciduous species. The analysis of tissue water relations using pressurevolume curves showed that evergreen species could develop a higher leaf turgor and lose turgor at lower leaf water potentials than deciduous species. This was related to a lower osmotic potential at full turgor in evergreen (-3.0 MPa)_than in deciduous (-2.0 MPa) species, rather than to the elastic properties of leaf tissue. The volumetric modulus of elasticity was 14 MPa in evergreen compared with 7–10 MPa in deciduous species. Thus, leaf characteristics are important in determining the drought resistance of evergreen species of this tropical dry forest.     

Some observations of phenology and ecophysiology ofDaphne kamtschatica Maxim.var.jezoensis (Maxim.) Ohwi, a shade deciduous shrub, in the forest of northern Japan

The phenology and leaf traits ofDaphne kamtschatica Maxim. var.jezoensis (Maxim.) Ohwi, the only summer deciduous shrub (20–40 cm) in the temperate forest of northern Japan, are examined. This plant carries through the winter mature leaves and well formed flower buds. It flowers in early spring during snowmelt and begins photosynthesis under relatively high irradiance under an open forest canopy. Our results show that there is significant carbon gain during the period when new leaves and fruit maturation also take place. Beginning in June, as the forest canopy closes, leaves onDaphne shoots senesce acropetally and the plants become completely bare in mid-July. After a period of 20-day dormancy, the shoots begin to resprout. Leaves become mature in early October and remain on the stem over winter. Leaf traits and photosynthesis measurements suggest as follows. 1) By becoming summer deciduous,D. kamtschatica avoids the cost of maintaining leaves inefficient under deep shade. 2) The onset and breaking of the summer dormancy is triggered by photoperiod since plants at the forest edge also become dormant even when light remained relatively high. However, the decreased duration of dormancy with higher light levels suggests that there is a tendency towards shorter dormancy where summer shade is absent and this could eventually lead to an evergreen habit such as that found in the alpine speciesDaphne miyabeana.     

藏东南色季拉山林线过渡带7种灌木植物的叶氮回收潜力

在湿润的青藏高原东南部, 为什么常绿灌木广泛占据高海拔的林线过渡带及以上的高山带, 而落叶灌木只能零星分布?未来气候变暖对该区不同功能群物种的影响是否相同?通过测定西藏东南部色季拉山林线过渡带7种灌木凋落叶的氮含量, 比较了极端高海拔地区灌木不同表达单位的叶氮回收潜力在不同功能群间的差异, 以及不同海拔、不同坡向间的差异, 试图从养分限制的角度为解答上述科学问题提供基础数据。研究结果表明: 1)从基于单位质量叶氮含量(N_mass)的叶氮回收潜力来看, 常绿灌木裂毛雪山杜鹃(薄毛海绵杜鹃) (Rhododendron aganniphum var. schizopeplum)显著高于其他6种落叶灌木, 但由于受比叶重的影响, 基于单位面积叶氮含量(N_area)的叶氮回收潜力则表现为落叶灌木总体较高; 2)落叶灌木山生柳(Salix oritrepha)和拉萨小檗(Berberis hemsleyana)的叶氮回收潜力在不同海拔或不同坡向间均无显著差异, 但裂毛雪山杜鹃基于N_mass的叶氮回收潜力在高海拔地段明显偏高。在极端高海拔的林线过渡带, 通过降低凋落叶中的氮含量(增加叶氮回收潜力)以达到高效的养分利用可能是常绿灌木裂毛雪山杜鹃适应高寒胁迫环境的重要策略。与落叶灌木相比, 常绿灌木裂毛雪山杜鹃叶氮回收潜力对未来气候变暖可能更敏感。     

冠层结构和光环境的时空变化对紫耳箭竹种群特征的影响

选择金佛山国家自然保护区内落叶阔叶林、常绿落叶阔叶混交林、常绿阔叶林3种典型群落类型,研究冠层结构和光环境特征,以及林下优势种--紫耳箭竹的种群特征.结果表明: 随着落叶阔叶林→常绿落叶阔叶混交林→常绿阔叶林的演替,Shannon多样性指数、Simpson优势度指数和Pielou指数呈增加趋势,表明群落趋于稳定发展的状态;冠层结构特征也发生了显著改变,冠层开度和平均叶倾角减小,叶面积指数增加,冠层的消光能力增强,林下光照水平降低.上层林冠是造成林型郁闭的主要原因,其中冠层厚度和冠层面积是2个主要的影响因素.冠层结构与林下光照指标显著相关,对林下散射光的影响最大.冠层开度、林下光照条件均随着生长季的到来而下降,而叶面积指数呈现增长的趋势,峰值出现在6、7月;平均叶倾角在春季达到最大值,在夏季为最小值.紫耳箭竹的生长与冠层结构和光环境密切相关,其在光照适中的常绿落叶阔叶混交林中生长得最好,分株粗壮、密度大(29.69±1.68株·m^-2),地下茎拓展能力强;落叶阔叶林中的强光环境可能造成土壤水分缺失,从而对其生长产生影响;而在常绿阔叶林的低光环境下紫耳箭竹分株矮小,密度小(5.80±1.16株·m^-2),克隆扩展能力降低.在森林结构演变的过程中,冠层结构发生了明显的改变,显著影响林下光环境,过度的低光环境对紫耳箭竹种群的更新和发展有限制作用.     

The impact of subcanopy light environment on the hydraulic vulnerability of Rhododendron maximum to freeze-thaw cycles and drought

The vulnerability of xylem to embolism development in Rhododendron maximum L., an evergreen diffuse-porous shrub, was investigated in relation to the frequency of winter freeze–thaw cycles in high and low light sites of the Eastern US. Though the frequency of freeze–thaw cycles during the winter was lower in North Carolina than in Virginia, the hydraulic conductivity of 3-year-old branches was reduced by up to 60% by winter embolism development in North Carolina compared to less than 30% in Virginia. Generally, small vessel diameters and volumes were associated with a significant resistance to embolism formation resulting from repeated freeze–thaws of xylem sap. In stems grown in high light sites (gaps), larger vessel volumes, and greater diameter growth of stems were associated with a significantly higher degree of freeze–thaw embolism development than in those grown in the low light sites. Thus, the growth patterns of R. maximum stems, under conditions of higher light availability, rendered them more susceptible to freeze–thaw-induced embolisms. Vulnerability to drought-induced embolism in stems was not affected by light environment. Rhododendron maximum was relatively sensitive to drought-induced embolism because 50% loss of hydraulic conductivity occurred at a water potential of -2.2 MPa. The distribution and gas exchange of R. maximum are constrained by the dual effects of freeze-thaw cycles and drought on vascular function.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Carbon dioxide concentrations within forest canopies—variation with time, stand structure, and vegetation type

Vertical CO₂ profiles (between 0.02 and 14.0 m) were studied in forest canopies of Pinus contorta, Populus tremuloides, and in a riparian forest with Acer negundo and Acer grandidentatum during two consecutive growing seasons. Profiles, measured continuously during 1- to 13-day periods in four to five stands differing in overstorey canopy area index (CAI < 4.5; including leaves, branches and stems), were well stratified, with highest [CO₂] just above the forest floor. Canopy [CO₂] profiles were influenced by stand structure (CAI, presence of understorey vegetation), and were highly dependent on vegetation type (deciduous and evergreen). A doubling of CAI in Acer spp. and P. tremuloides stands did not show an effect on upper canopy [CO₂], when turbulent mixing was high. However, increasing understorey biomass in Acer spp. stands had a profound effect on lower canopy [CO₂]. In open stands with a vigorous understorey layer, higher soil respiration rates were offset by increased understorey gas exchange, resulting in [CO₂] below those of the convective boundary layer (CBL). Midday depletions up to 20 ppmv below CBL values could be frequently observed in deciduous canopies. In evergreen canopies, [CO₂] stayed generally above the CBL background values, [CO₂] profiles were more uniform, and gradients were smaller than in deciduous stands with similar CAI. Seasonal changes of canopy [CO₂] reflected changes in soil respiration rates as well as plant phenology and gas exchange of both dominant tree and understorey vegetation. Seasonal patterns were less pronounced in evergreen than in deciduous forests.     

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

Photosynthetic acclimation to simultaneous and interacting environmental stresses along natural light gradients: optimality and constraints

There is a strong natural light gradient from the top to the bottom in plant canopies and along gap-understorey continua. Leaf structure and photosynthetic capacities change close to proportionally along these gradients, leading to maximisation of whole canopy photosynthesis. However, other environmental factors also vary within the light gradients in a correlative manner. Specifically, the leaves exposed to higher irradiance suffer from more severe heat, water, and photoinhibition stresses. Research in tree canopies and across gap-understorey gradients demonstrates that plants have a large potential to acclimate to interacting environmental limitations. The optimum temperature for photosynthetic electron transport increases with increasing growth irradiance in the canopy, improving the resistance of photosynthetic apparatus to heat stress. Stomatal constraints on photosynthesis are also larger at higher irradiance because the leaves at greater evaporative demands regulate water use more efficiently. Furthermore, upper canopy leaves are more rigid and have lower leaf osmotic potentials to improve water extraction from drying soil. The current review highlights that such an array of complex interactions significantly modifies the potential and realized whole canopy photosynthetic productivity, but also that the interactive effects cannot be simply predicted as composites of additive partial environmental stresses. We hypothesize that plant photosynthetic capacities deviate from the theoretical optimum values because of the interacting stresses in plant canopies and evolutionary trade-offs between leaf- and canopy-level plastic adjustments in light capture and use.     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval. This revised version was published online in June 2006 with corrections to the Cover Date.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型（常绿与落叶）亚热带木本植物不同冠层高度（上下冠层）当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计（standardized major axis estimation,SMA）方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示：（1）当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异（P>0.05）,叶面积在常绿和落叶植物中有显著差异（P<0.05）,常绿和落叶植物的比叶重存在显著差异（P<0.05）,而落叶植物的比叶重在不同冠层高度存在显著差异（P<0.05）,同一冠层,常绿植物比叶重显著高于落叶植物（P<0.05）;（2）69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;（3）不同冠层和生活型植物的叶面积与叶生物量呈等速生长关系,但其异速生长常数在不同冠层高度与生活型间存在差异。这些结果表明冠层高度和生活型未改变叶面积-生物量之间的等速生长关系,不支持"收益递减"假说。