南京地区落叶栎林主要木本植物的展叶动态研究

 植物的展叶过程是由自身遗传因子决定的,同时又受到多种生态因子的调节,反映了植物的生活史对策和群落物种多样性的维持机制。在2001和2002年的3～6月间,不定期记录了南京地区三个落叶栎(Quercus spp.)林中主要木本植物的展叶情况,包括被标记标准枝的叶数、叶的长度、宽度、叶面积、叶干重等参数。结果表明在所调查的落叶栎林中,林冠层物种的成熟叶面积和单位叶面积干重都显著大于林下层物种;最早展叶的物种为林下层物种,但林冠层展叶顺序与林下层无显著差异。叶面积越大、单位叶面积干重越小的物种展叶越晚;林冠层物种展叶较林下层快,物种成熟叶面积越大,展叶速率越大。最后对展叶顺序和展叶速度的生态学意义作了讨论。     

南京地区落叶栎林木本植物叶物候研究

叶物候参数长期以来被认为与植物的碳获取的最大化有关,能反映物种的资源利用策略。温带地区因为寒冷冬天的限制,延长叶寿命成为一些物种进行生长发育和繁衍的基础。为探讨叶寿命延长的可能途径(早出叶、晚落叶,或两者兼有),该研究以南京地区两个落叶栎(Quercus spp.)林为研究对象,观测了其中木本植物的出叶物候、落叶物候,并分析了它们与叶寿命之间的关系。结果发现:1)不同物种的出叶开始时间相差较大,出叶早的物种早结束出叶过程;2)不同物种的落叶开始时间相差较大,早开始落叶的物种,落叶持续时间较长,落叶结束时间则相对集中。3)相关分析和回归分析都表明,叶寿命与出叶时间和落叶时间显著关联,但早出叶对叶寿命的延长可能更为重要,因为早出叶相对于晚落叶在物种资源利用上比较具有优势。4)不同物种的出叶时间和落叶时间没有显著相关,可能因为出叶过程和落叶过程是由不同的启动因子引起。这说明延长叶寿命不一定同时通过早出叶和晚落叶来达到。     

Leaffall Phenology in a Subtropical Wet Forest in Puerto Rico: From Species to Community Patterns

Leaffall phenology is an important periodical event in forests, contributing to mobilization of organic matter from primary producers to soil. For seasonal forests, leaffall periodicity has been related to rainfall regime and dry season length. In weakly seasonal forests, where there is no marked dry season, other climatic factors could trigger leaf shed. In this study, we aimed to determine if other climatic variables (wind speed, solar radiation, photosynthetic photon flux density [PPFD], day length, temperature, and relative humidity) could be better correlated with patterns of litter and leaffall in a weakly seasonal subtropical wet forest in Puerto Rico. Leaffall patterns were correlated mainly with solar radiation, PPFD, day length, and temperature; and secondarily with rainfall. Two main peaks of leaffall were observed: April–June and August–September, coinciding with the periods of major solar radiation at this latitude. Community leaffall patterns were the result of overlapping peaks of individual species. Of the 32 species analyzed, 21 showed phenological patterns, either unimodal (16 species), bimodal (three species), or multimodal (two species). Lianas also presented leaffall seasonality, suggesting that they are subject to the same constraints and triggering factors affecting trees. In addition to solar radiation as a main determinant of leaffall timing in tropical forests, our findings highlight the importance of interannual variation and asynchrony, suggesting that leaffall is the result of a complex interaction between environmental and physiological factors.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

The vertical foliage distributions of six understory tree species in a Chamaecyparis obtusa Endl. forest

Summary The relationships between the amounts of foliage and heights of trees were studied for the dominant understory tree species, including three evergreen and three deciduous species, in a secondary forest of Chamaecyparis obtusa Endl. The relationships showed two phases: leaf increasing and stationary phases. In the leaf-increasing phase, the height growth allowed these species to expand the canopy by increasing the number of leaves. In the stationary phase, the number of leaves was relatively constant number irrespective of tree height from 160 to 400 cm. The number of leaves in the stationary phase represents the maximum number of leaves that can be supported by trees under shady conditions. From the analyses of vertical distributions of leaves in six species, mono- and multi-layer foliage distributions were detected. Two evergreen species, Eurya japonica and Cleyera japonica, showed multi-layer foliage distributions, whereas three deciduous species, Lyonia ovalifolia, Rhododendron reticulatum and Vaccinium hirtum, and one evergreen species, Pieris japonica, showed mono-layer foliage distributions. The relationships between the weights of non-photosynthetic and photosynthetic organs of the six species were examined. The proportion of non-photosynthetic organs increased with tree height. The understory species attained the stationary phase and were maintained by minimizing their investment in non-photosynthetic organs, i.e. their height growth was arrested by the shady conditions under the crown trees.     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.     

Responses of Tropical Understory Plants to a Severe Drought: Tolerance and Avoidance of Water Stress1

Shade-tolerant understory shrubs and subcanopy trees constitute most of the woody species in Neotropical moist forest, but studies demonstrating physiological differences among these species are few. Shade-tolerant species that coexist in the forest understory exhibit differences in leaf life span that have been associated with variation in physiological traits. We hypothesized that water relations of understory species with widely divergent leaf life spans differ in response to drought. Although severe drought is infrequent in Neotropical moist forest, we studied the water relations of shade-tolerant understory species with short or long leaf life spans during the severe 1991-1992 dry season on Barro Colorado Island, Panama. The predawn leaf water potential declined to -2.8 and -3.6 MPa during the dry season in Hybanthus prunifolius and Psychotria horizontalis, respectively, two species with short leaf life spans, but remained above -1.3 MPa in two species with long leaf life spans, Swartzia simplex and Ouratea lucens. The midday leaf water potential dropped as low as -3.4 and -4.5 MPa for H. prunifolius and P. horizontalis, respectively. The osmotic potential of H. prunifolius and P. horizontalis and another species with short leaf life span, Alms blackiana, decreased early in the dry season, a period during which all three had substantially negative predawn water potential. In contrast, the osmotic potential of S. Simplex, O. lucens, and Licania platypus, a third species with long leaf life span, declined late in the dry season, even though we observed little change in predawn water potential for S. simplex and O. lucens. We conclude that the variable and potentially severe dry season in Neotropical moist forest can be sufficiently intense to severely limit soil moisture availability for understory plants. H. prunifolius and P. horizontalis tolerated dehydration, whereas S. simplex and O. lucens postponed dehydration.     

Ecological distribution of homobaric and heterobaric leaves in tree species of Malaysian lowland tropical rainforest

Tree species can generally be classified into two groups, heterobaric and homobaric leafed species, according to whether bundle-sheath extensions (BSEs) are found in the leaf (heterobaric leaf) or not (homobaric leaf). In this study, we study whether the leaf type is related to the growth environment and/or life form type, even in a tropical rain forest, where most trees have evergreen leaves that are generally homobaric. Accordingly, we investigated the distribution of leaf morphological differences across different life forms of 250 tree species in 45 families in a tropical rainforest. In total, 151 species (60%) in 36 families had homobaric leaves, and 99 species (40%) in 21 families had heterobaric leaves. We found that the proportion of heterobaric and homobaric leaf species differed clearly across taxonomic groups and life form types, which were divided into five life form types by their mature tree heights (understory, subcanopy, canopy, and emergent species) and as canopy gap species. Most understory (94%) and subcanopy (83%) species such as Annonaceae had homobaric leaves. In contrast, heterobaric leaf trees appeared more frequently in the canopy species (43%), the emergent species (96%) (such as Dipterocarpaceae), and the canopy gap species (62%). Our results suggest that tree species in the tropical rainforest adapt to spatial differences in the environmental conditions experienced at the mature height of each tree species, such as light intensity and vapor pressure difference, by having differing leaf types (heterobaric or homobaric) because these types potentially have different physiological and/or mechanical functions.     

沈阳城市森林主要树种叶片形态生态特征与生长规律

利用统计和多元分析方法研究了沈阳城市森林10个主要树种叶片的形态生态特征和生长规律.结果表明,影响树木叶片生长发育节律的主要气象因子是≥5 ℃积温、累积日照时数和各物候期中旬平均温度.不同物候期树木所需要的气象因子有所变化,树木萌动期要求低温和寒冷指数,而在生长期需要一定的温暖指数和湿润指数.10个树种的叶片形态数量特征差异较大,按叶形指数依次为短序胡枝子＞华北丁香＞国槐＞银白杨＞红瑞木＞金银忍冬＞水腊＞水曲柳＞稠李＞黄菠萝;按叶面积依次为华北丁香＞银白杨＞黄菠萝＞稠李＞水曲柳＞红瑞木＞短序胡枝子＞金银忍冬＞国槐＞水腊.在10个树种叶长度与宽度、周长、叶面积之间相互关系中,基本属于异速生长类型,符合y=ax^k模型,但在叶长度与宽度之间关系中,除银白杨k＞1外,其它9个树种均k＜1;在叶长度与周长关系中,黄菠萝k＞1, 银白杨k≈1,呈线性关系,其它树种均k＜1;在叶长度与叶面积关系中,10个树种k＞1,而 银白杨k=2.1028,叶面积增长速率近似于叶长的2倍.建立了10个树种的叶面积最优回归估测模型,并对模型进行了估测检验.     

Ecophysiology of exotic and native shrubs in Southern Wisconsin

Summary We compared seasonal trends in photosynthesis of two naturalized exotic shrubs (Rhamnus cathartica and Lonicera X bella) and two native shrubs (Cornus racemosa and Prunus serotina) in open and understory habitats in southern Wisconsin. We examined the relationships between resource availability and leaf photosynthetic performance in these four species. All four species had similar relationships between leaf nitrogen (N) content and photosynthetic rate, but the species differed in absolute leaf N content and therefore in photosynthetic rates. Maximum daily photosynthetic rates of all species were significantly correlated with leaf N content in the open habitat, but not in the understory, where low light availability was the major limitation to photosynthesis. Extended leaf longevity was important in the forest understory because it allowed shrubs to take advantage of high light availability at times when the overstory canopy was leafless. Early leaf emergence was more important than late senescence: from 27% to 35% of the annual carbon gain of P. serotina, R. cathartica, and L. X bella occurred prior to leaf emergence of C. racemosa, the species with the shortest leaf life span. Extended leaf longevity of exotic shrubs may help explain their persistence in the understory habitat, but it contributed relatively less to their annual carbon gain in the open habitat.     

Foliar demand and resource economy of nutrients in dry tropical forest species

Important phenological activities in seasonally dry tropical forest species occur within the hot‐dry period when soil water is limiting, while the subsequent wet period is utilized for carbon accumulation. Leaf emergence and leaf area expansion in most of these tree species precedes the rainy season when the weather is very dry and hot and the soil cannot support nutrient uptake by the plants. The nutrient requirement for leaf expansion during the dry summer period, however, is substantial in these species. We tested the hypothesis that the nutrients withdrawn from the senescing leaves support the emergence and expansion of leaves in dry tropical woody species to a significant extent. We examined the leaf traits (with parameters such as leaf life span, leaf nutrient content and retranslocation of nutrients during senescence) in eight selected tree species in northern India. The concentrations of N, P and K declined in the senescing foliage while those of Na and Ca increased. Time series observations on foliar nutrients indicated a substantial amount of nutrient resorption before senescence and a ‘tight nutrient budgeting’. The resorbed N‐mass could potentially support 50 to 100% and 46 to 80% of the leaf growth in terms of area and weight, respectively, across the eight species studied. Corresponding values for P were 29 to 100% and 20 to 91%, for K 29 to 100% and 20 to 57%, for Na 3 to 100% and 1 to 54%, and for Ca 0 to 32% and 0 to 30%. The species differed significantly with respect to their efficiency in nutrient resorption. Such interspecific differences in leaf nutrient economy enhance the conservative utilization of soil nutrients by the dry forest community. This reflects an adaptational strategy of the species growing on seasonally dry, nutrient‐poor soils as they tend to depend more or less on efficient internal cycling and, thus, utilize the retranslocated nutrients for the production of new foliage biomass in summer when the availability of soil moisture and nutrients is severely limited.     

Survival curves and longevity of the leaves of Alnus japonica var. arguta in Kushiro Marsh

The seasonal changes in leaf emergence and leaf-fall in a Japanese alder stand of the fen in Kushiro Marsh were studied, and survival curves for the leaves were drawn. Leaves collected in litter traps were dried and weighed to study the seasonal changes, peaks in mid-August and late September to October suggested a bimodal annual curve. Study of the seasonal changes in the number of emerged and fallen leaves per shoot revealed a third peak about one month before the August peak, showing a trimodal annual curve. First and second leaves had a longevity of about 40 and 50–60 days, respectively. The longevity increased until the fifth leaf. With the sixth and following leaves, longevity decreased. Leaf size increased with leaf rank, with the first leaf being the smallest. The first leaf had only about 10% and the second leaf only 20% of the area of the fifth leaf. On this basis, the early to mid-July peak in number of fallen leaves was composed of first and second leaves which were smaller and short-lived. The early August and September/October peaks were high in both number and mass of fallen leaves. Compared to reports on Japanese alder of other mountainous districts in Hokkaido, the alder trees of Kushiro Marsh had about the same number of leaves per shoot, but had a season of leaf emergence which was about 6 weeks shorter. In addition, the longevity of the longest-lived fifth leaf was about 30–40 days shorter. The short life span of the leaves could be considered as an adaptive strategy of this species to environmental constraints of its habitat.     

东灵山地区辽东栎的叶群体统计

 辽东栎是我国暖温带落叶阔叶林主要优势种之一。本研究中,我们采用随机枝取样法,研究了东灵山地区辽东栎的叶群体的数量动态。结果表明：(1)不同个体,同一个体内部不同层次,叶数量动态趋势都基本一致。现叶期和叶落半衰期很短,现叶方式为爆发型。(2)不同个体间叶期差异很大,暗示在辽东栎种群内部或许存在有一定的遗传分化。(3)叶数量动态同气候因子变化相吻合,有利于辽东栎的碳获取。     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

The basis for variation in leaf longevity of plants

Any theory of leaf phenology must predict leaf longevity, leaf habit, leaf expansion and its timing among other variables. These phenological traits may be important keys to understand the response of trees to climatic change. Here I concentrate on and review two of these critical phenological traits, leaf longevity and leaf habit. Theories of leaf longevity were re-evaluated and leaf longevity is concluded to be optimized to maximize plant carbon gain. From this perspective, three points are predicted. Leaf longevity is short when the photosynthetic rate of the leaf is high, when the photosynthetic rate decreases rapidly through time, or when the construction cost of the leaf is small. These predictions are well supported by empirical as well as experimental results on various plant species. The theory, which is extended to seasonal environments, is general and applicable to seasonal as well as aseasonal environments. The theory simulated the bimodal geographic distribution of evergreenness.     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

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.     

Phenology of the shrub strata of successional sub-tropical humid forests of north-eastern India

The phenology of 70 shrub species occurring in 5-, 25- and 60-yr old forest fallows, developed after slash and burn agriculture in north-eastern India, was studied. The early successional shrubs were largely deciduous with predominant leaf fall during the dry winter, while late successional shrubs were evergreen and with a less pronounced leaf fall pattern; the species in the 25-yr old fallow were largely of the leaf-exchanging-evergreen type. Peak leaf production, flowering and fruiting are delayed by one month in the 60-yr old fallow. The phenological separation of the growth types in the three fallows is discussed and related to possible micro-environmental differences.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.