Spatial and seasonal variations in mobile carbohydrates in Pinus cembra in the timberline ecotone of the Central Austrian Alps

To test whether the altitudinal limit of tree growth is determined by carbons shortage or by a limitation in growth we investigated non structural carbohydrates and their components starch and total soluble sugars in Pinus cembra trees along an elevational gradient in the timberline ecotone of the Central Austrian Alps. NSC contents in needles, branches, stems, and coarse roots were measured throughout an entire growing season. At the tissue level NSC contents were not significantly more abundant in treeline trees as compared to trees at lower elevations. Along our 425 m elevational transect from the closed forest to the treeline we failed to find a stable elevational trend in the total NSC pool of entire trees and observed within season increases in the tree's NSC pool that can be attributed to an altitudinal increase in leaf mass as needles contained the largest NSC fraction of the whole tree NSC pool. Furthermore, whole tree NSC contents were positively correlated with net photosynthetic capacity. Although our observed NSC characteristics do not support the hypothesis that tree life at their upper elevational limit is determined by an insufficient carbon balance we found no consistent confirmation for the sink limitation hypothesis.     

青海云杉休眠前后非结构性碳水化合物含量随海拔变化

[目的]冬季低温是决定高山林线树种能否生存的重要生态因素。休眠前后树木体内碳水化合物的变化能够反映出其安全越冬的策略。但目前高山林线树种休眠前后体内碳水化合物的变化及机理尚不清楚。为了探讨林线树种越冬期间的生存策略,[方法]本文以祁连山林线青海云杉为对象,研究了不同海拔(2 900、3 000、3 100、3 200、3 300 m)成年树枝、叶休眠前后非结构性碳水化合物含量变化,包括可溶性糖(葡萄糖和果糖)、淀粉以及糖/淀粉比值的变化。[结果]表明:(1)休眠前青海云杉当年及1年生叶在高海拔处NSC含量高于中、低海拔处NSC含量,而在枝条中高、中、低海拔间没有显著差异。(2)休眠后叶中NSC含量最高值在中海拔3 100 m处,显著高于高海拔;不同组织中NSC含量高于休眠前,叶片中尤为明显。(3)与休眠前相比,休眠后可溶性糖及淀粉含量增加,淀粉含量增加幅度显著,引起可溶性糖/淀粉比值降低。[结论]青海云杉休眠前贮存足够的碳水化合物保证越冬期间的细胞渗透调节及能量代谢,并且休眠后气温升高时叶片和枝条的NSC能够得到迅速补充,为新枝条萌发提供充足的碳源,反映了青海云杉在越冬期间的生存策略。     

Minimum cuticular conductance and cuticle features of Picea abies and Pinus cembra needles along an altitudinal gradient in the Dolomites (NE Italian Alps)

Winter desiccation is believed to contribute to stress in coniferous trees growing at the treeline because cuticular conductance increases with altitude. To test whether winter desiccation occurs in high-altitude conifers of the Dolomites (NE Italian Alps), we measured minimum cuticular conductance (g(min)), needle wettability (contact angle) and cuticle thickness in Picea abies (L.) Karst. and Pinus cembra L. needles from December to August. Samples were collected from adult trees along an altitudinal gradient from valley bottom (1050 m a.s.l.) to the treeline (2170 m a.s.l.). The treeline site is one of the highest in the area and is characterized by a generally low wind exposure. Altitude had no effect on g(min) in either species. In P. abies, large seasonal variations in g(min) were recorded but no changes were related to needle age class. Pinus cembra had a low g(min) and appeared to be efficient in reducing needle water losses. There was a significant increase in g(min) with needle aging in P. cembra growing at low altitude that could be related to a shorter needle longevity compared with P. abies. High contact angles (> 110-120 ) suggested the presence of tubular epicuticular waxes on needles of both species. Contact angles were higher (low wettability) in high-altitude needles than in low-altitude needles. By the end of winter, there was no difference in contact angles between needles in the windward and leeward positions. Wax structures transformed toward planar shapes as demonstrated by the decrease in contact angle from winter to summer. In both species, the cuticle was thicker in needles of high-altitude trees than in needles of low-altitude trees and there was no correlation between g(min) and cuticle thickness. Because desiccation resistance did not decrease with altitude in either species, we conclude that they are not susceptible to winter desiccation at the tree line.     

峨眉山不同海拔冷杉径向生长对气候变化的响应

【目的】研究峨眉山不同海拔冷杉Abies fabri径向生长对气候变化的响应规律。【方法】以冷杉为研究对象,用树轮生态学的方法对峨眉山不同海拔的冷杉进行年轮采样、处理与分析,比较不同海拔冷杉径向生长特征及其与各气候因子之间的关系。【结果】年表统计数据表明,4个年表均具有较高的可靠性,可以用于树木生长与气候关系的研究。不同海拔冷杉径向生长与气候的关系具有明显的海拔差异,海拔2 400 m处冷杉径向生长与当年7月的温度呈显著正相关;海拔2 800 m处冷杉径向生长与当年4-9月的帕尔默干旱指数(PDSI)呈显著正相关;海拔2 900 m处冷杉径向生长与前一年9月和当年4月的温度呈显著负相关,与当年1-9月的PDSI均呈显著正相关;海拔3 000 m处冷杉径向生长和当年3-4月温度呈显著负相关,与当年6-7月的PDSI和当年4月平均相对湿度呈显著正相关。【结论】峨眉山高海拔的冷杉包含有更多的环境信息,对气候变化的响应也更为敏感;高海拔区域冷杉径向生长受到水分的强烈限制,而低海拔区域冷杉径向生长主要受温度影响;冷杉衰亡与气候因子的变化密切相关,随着区域气候的变暖,高海拔地区冷杉可能会进一步衰退。该研究揭示了峨眉山不同海拔冷杉生长动态及其对气候变化响应的敏感性差异,为全球气候变化背景下冷杉林的保护与适应性管理提供科学参考。     

Seasonal changes of C and N non-structural compounds in the stem sapwood of adult sessile oak and beech trees

We assessed the pools of non-structural nitrogen compounds (NSNC) through a year, thereby addressing the question of whether mature sessile oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.), which differ in wood anatomy and growth patterns, exhibit contrasting seasonal dynamics of NSNC pools as previously shown for non-structural carbohydrate (NSC) pools. Seasonal fluctuations of NSNC (amino acids and soluble proteins) and NSC (starch and soluble sugars) pools were analyzed in the inner and the outer stem sapwood. In oak, NSC showed marked seasonal variation within the stem sapwood (accumulation during winter and decrease during bud burst and early wood growth), whereas in beech seasonal fluctuations in NSC were of minor amplitude. Even if the distribution and intensity of the NSNC pools differed between the two species, NSNC of the stem sapwood did not show seasonal variation. The most significant change in NSNC pools was the seasonal fluctuation of protein composition. In both species, two polypeptides of 13 kDa (PP13) and 26 kDa (PP26) accumulated during the coldest period in parallel with starch to sugar conversion and disappeared with the onset of spring growth. The absence of seasonal changes in total soluble protein concentration suggests that the polypeptides are involved in the internal nitrogen (N) cycling of the stem rather than in N storage and remobilization to the other growing organs of the tree.     

Winter water relations of New England conifers and factors influencing their upper elevational limits. I. Measurements

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.     

Interactions among cutting frequency, reserve carbohydrates, and post-cutting biomass production in Gliricidia sepium and Leucaena leucocephala

The ability of trees and shrubs to coppice vigorously after being cut or pruned is of great importance in the management of agroforestry systems. In a study conducted in the seasonally dry climate of Ibadan, Nigeria, frequent cutting progressively decreased concentrations of starch and total reserve carbohydrates in Gliricidia sepium (Jacq.) Walp. and Leucaena leucocephala (Lam.) De Wit. In frequently cut trees, starch was severely reduced in the stumps of both species. In contrast, soluble sugar concentrations in roots and stems of both species were consistently maintained at or above the levels in uncut control trees, suggesting that sugar levels were maintained through the hydrolysis of starch reserves. No starch replenishment occurred during the six weeks after cutting. In trees that were cut and then allowed to grow undisturbed during the wet season, replenishment of stem starch began within three months after cutting. Shoot regrowth after cutting decreased starch levels first in stems and, only after additional cuts, in roots. Dry-season cuts had little effect on reserve carbohydrates in G. sepium but quickly reduced stem starch in L. leucocephala. Frequent cutting decreased dry matter production, and this decrease was correlated with reduced levels of reserve carbohydrates.This revised version was published online in November 2005 with corrections to the Cover Date.     

Winter water relations of New England conifers and factors influencing their upper elevational limits. II. Modeling

We used the water relations model, WINWAT, to model winter water relations of three conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.) and red pine (P. resinosa Ait.)-growing at their upper elevational limits on Mt. Ascutney, Vermont, USA, in the winters of 1997 and 1999. Modeled relative water contents remained above 60% in the two youngest foliar age classes of all three species during both winters, indicating that desiccation stress in winter is not responsible for setting the upper elevational limits of these species at this site under present climatic conditions. WINWAT indicated that winter water relations of these low-elevational species were sensitive to low relative humidity, which increased transpiration rates, and low temperatures, which inhibited recharge, but are much less sensitive to summer climate than in the case of subalpine conifers in Colorado. Our results indicate that summer and winter temperatures and relative humidities (or precipitation/potential evapotranspiration ratios) should be incorporated into climate change models designed to simulate future tree distributions.     

Impacts of competition and nitrogen addition on plant stoichiometry and non-structural carbohydrates in two larch species

Previous research has shown that competition between plants can have differential effects on leaf stoichiometry and non-structural carbohydrate(NSC) in different environments.However,little attention has been given to understanding these effects on non-photosynthetic organs,particularly of deciduous tree species.Here we assess the impact of competition on below and aboveground biomass,stoichiometry,nutrient composition and NSC in pure and mixed forests of two Larch species,Larix kaempferi and L.olgensis under nitrogen(N) addition.Nitrogen enrichment did not result in stronger intraspecific competition for both species and L.olgensis benefited from the presence of L.kaempferi under different N levels.Stems kept relatively stable C/N compared to roots and branches in response to competition,while N addition imposed stronger impacts on N/P of different organs rather than competition.In contrast to stable C concentrations,starch and soluble sugar concentrations were more easily impacted by competition and the addition of nitrogen.Competition forced L.kaempferi and L.olgensis to allocate more carbon into storage by increasing their starch concentration and starch/soluble sugar of stems under competition.However,no significant differences in stoichiometry and NSC concentration between intra-and interspecific competition were found.NSC and nutrient pools of L.kaempferi stems,branches and coarse roots consistently declined due to competition regardless of N addition.Coarse and fine roots of L.kaempferi accumulated more N when in competition with L.olgensis than with a conspecific neighbor under N addition.Our results show that NSC was more sensitive to competition relative to stoichiometric traits(N and P) of non-photosynthetic organs.     

Divergent growth responses and increasing temperature limitation of Qinghai spruce growth along an elevation gradient at the northeast Tibet Plateau

Divergent responses between tree growth and climate factors have been widely reported at high latitudes in the northern hemisphere. Here we show variable climate-growth relationships and divergent growth responses of Qinghai spruce (Picea crassfolia) along an elevation gradient at a mid-latitude site at the northeastern Tibetan Plateau, China. Trees from higher elevations, limited mainly by temperature, show divergent growth trends over time and two responses to climate. Some trees show increasing positive and some increasing negative responses to growing season temperature during the last decades. Trees from lower treeline show a strengthening drought stress signal over time and no divergent growth trends within sites. Our results indicate that single tree analysis might be a worthwhile tool to (1) uncover spatial–temporal changes in climate-growth relationships of trees, (2) better understand future growth performance and (3) help overcome current limitations of tree ring based climatic reconstructions.     

Carbohydrates in individual poplar fine roots: effects of root age and defoliation

Late-summer starch accumulation in fine roots of poplars (Populus x canadensis Moench.) defoliated by gypsy moth (Lymantria dispar L.) lagged behind that in fine roots of undefoliated trees. If starch concentration declines with age, defoliation-induced changes in root system age structure could be partly responsible for this difference. To test this hypothesis, we measured fine-root starch and soluble sugar concentrations in roots of known age from trees in defoliated and undefoliated plots. There was a significant interaction between the effects of defoliation and root type (white, brown, or dead) on fine root soluble sugar concentration because of the high concentration of soluble sugars in white roots from trees in undefoliated plots. Both root starch and soluble sugar concentrations were variable among individuals of each age class. The population frequency distributions for starch and soluble sugar concentrations were both right-skewed, and fit by exponential functions. These data are most consistent with direct defoliation effects on a labile and dynamic pool of carbohydrates in poplar fine roots, rather than indirect defoliation effects on root system age structure.     

Cellular basis for limitation of poplar leaf growth by water deficit

During the summers of 1986 and 1987, stem and leaf growth were measured on coppiced plants of Populus trichocarpa Torr. & A. Gray, P. deltoides Bartr. ex Marsh, and P. trichocarpa x deltoides growing in the field in Puyallup, WA. The trees were either irrigated periodically throughout the season, or grown without irrigation. In both treatments, stem volume at the end of the growing season was directly proportional to total leaf area in all three genotypes. The rate of individual leaf growth was reduced by lack of irrigation more in the parental species than in the hybrid. Only in the parental species did unirrigated trees have lower leaf water potentials (predawn and midday) than irrigated trees. However, stomatal conductances of all three genotypes were lower in unirrigated trees than in irrigated trees. Osmotic potentials of growing leaves of all three genotypes were also lower in unirrigated trees than in irrigated trees. As a consequence, turgor of growing leaves was as great in unirrigated trees as in irrigated trees, which indicates that turgor differences cannot explain the lower rates of leaf growth in the unirrigated trees. However, cell wall extensibility of leaves was lower in unirrigated trees than in irrigated trees, and the difference was greater in the parental species than in the hybrid. Unlike its effect on leaf area growth, irrigation increased stem volume growth of the hybrid and the parental species by a similar amount (12-16%).     

Diurnal changes in gas exchange and carbon partitioning in needles of fast- and slow-growing families of loblolly pine (Pinus taeda)

We investigated diurnal and seasonal changes in carbon acquisition and partitioning of recently assimilated carbon in fast- and slow-growing families of loblolly pine (Pinus taeda L.) to determine whether fast-growing families exhibited greater carbon gain at the leaf level. Since planting on a xeric infertile site in Scotland County, NC, USA in 1993, five Atlantic Coastal Plain (ACP) and five "Lost Pines" Texas (TX) families have been grown with either optimal nutrition or without fertilization (control). In 1998 and 1999, gas exchange parameters were monitored bimonthly in four families and needles were analyzed bimonthly for starch and soluble sugar concentrations. Although diurnal and seasonal effects on net photosynthesis (A(net)) and maximum rate of light-saturated photosynthesis (A(max)) were significant, few family or treatment differences in gas exchange characteristics were observed. The A(net) peaked at different times during the day over the season, and A(max) was generally highest in May. Instantaneous water-use efficiency (WUE(i)), derived from gas exchange parameters, did not differ among families, whereas foliage stable isotope composition (delta(13)C) values suggested that TX families exhibited lower WUE than more mesic ACP families. Although there were no diurnal effects on foliar starch concentrations, needles exhibited pronounced seasonal changes in absolute concentrations of total nonstructural carbohydrates (TNC), starch and soluble sugars, and in partitioning of TNC to starch and sugars, mirroring seasonal changes in photosynthesis and shoot and root growth. In all families, foliar starch concentrations peaked in May and decreased to a minimum in winter, whereas reducing sugar concentrations were highest in winter. Some family and treatment differences in partitioning of recently assimilated carbon in needles were observed, with the two TX families exhibiting higher concentrations of TNC and starch and enhanced starch partitioning compared with the ACP families. We conclude that growth differences among the four families are not a function of differences in carbon acquisition or partitioning at the leaf level.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.     

Seasonal patterns of carbohydrate reserves in red spruce seedlings

We studied seasonal dynamics of carbohydrate storage in red spruce (Picea rubens Sarg.) seedlings by measuring starch and sugar concentrations of old needles (>/= one year old), new needles (< one year old), stems, and roots in two stands in the Green Mountains of Vermont. Although the two stands differed in many site characteristics including percent slope, aspect, soil type, drainage, and 564 m in elevation, concentrations and seasonal patterns of carbohydrates were similar for the two stands. For all tissues, starch concentrations peaked in late spring, declined through summer, and reached a minimum in winter. Sugar concentrations were greater than starch concentrations in all months except May and June. Sugar concentrations peaked in winter, and old needles showed a significant increase in sugar concentration between February and March. This increase in sugar concentration occurred without any reduction in localized starch concentrations or reductions in sugar or starch concentrations in new needles, stems or roots. Because March measurements were made toward the end of a prolonged thaw, a time when increases in photosynthesis have been documented for red spruce, it is likely that the March increase in sugar concentrations resulted from photosynthesis during the thaw. Compared with stems and roots, needles generally contained the highest concentration of carbohydrates and exhibited the greatest seasonal change in carbohydrate concentration. Needles were also the largest reservoir of carbohydrates throughout the year, especially during winter. Because of the critical roles of needles in photosynthesis and storage of carbohydrates, we conclude that any factors that disrupt the accumulation or availability of carbohydrates in red spruce needles will greatly alter plant carbon relations.     

Seasonal biochemical changes in coniferous forest canopies and their response to fertilization

Seasonal changes in concentrations of total nitrogen, free amino acids, chlorophyll, starch and sugar were measured in foliage from fertilized and unfertilized conifer forests in New Mexico and Oregon. In the New Mexico Douglas-fir (Pseudotsuga menziesii var glauca (Beissn.) Franco) forest, fertilization resulted in elevated foliar nitrogen concentrations on all dates, from an average of 9 mg g(-1) in unfertilized trees to 14 mg g(-1) in fertilized trees. In the Oregon western hemlock (Tsuga heterophylla (Raf.) Sarg.) forest, fertilization increased total N by only 15%, from 13 mg g(-1) in unfertilized trees to 15 mg g(-1) in fertilized trees. Foliar nitrogen concentrations on a weight basis were lowest in winter and spring, but did not vary seasonally when expressed on a leaf area basis. Chlorophyll concentrations increased with fertilization and had greater seasonal variation than did total nitrogen concentrations. Chlorophyll concentrations were significantly higher during the growing season than in the winter and spring months. Fertilization did not result in major changes in the proportion of total nitrogen in chlorophyll at either the Oregon or the New Mexico site. Concentrations of free amino acids varied with date and fertilization treatment; in New Mexico, amino acids were highest in the winter sample, whereas in Oregon, they were lowest in winter and spring. At both sites, amino acid concentrations were significantly higher in fertilized trees than in control trees on most dates and the ratios of amino acid-N to total N were also significantly higher in fertilized trees. For both sites, starch concentrations were nearly zero for most of the year, but increased sharply just before bud break and initiation of new growth in the spring. Although fertilization resulted in increased nitrogen concentrations in foliage at both sites, the response in New Mexico was much greater than in Oregon. These results are in agreement with forest productivity data that suggest that growth in the New Mexico site is limited by nitrogen, whereas in the Oregon site it is not.     

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

Wood specific gravity of some tree species in the Garhwal Himalayas,India

Estimation of terrestrial biomass depends critically on reliable information about wood specific gravity of forest trees. In recent years, wood specific gravity has become more important when exploring the universality of functional traits of plants and estimating their global carbon stocks. To estimate their specific gravity, wood samples were collected from a total of 34 tree species, 30 from lower elevations and 4 from upper elevations in the Garhwal Himalayas, India. The results show that the average wood specific gravity was 0.631 (ranging between 0.275 ± 0.01 and 0.845 ± 0.03) for the species at lower elevations and 0.727 (ranging between 0.628 ± 0.02 and 0.865 ± 0.02) for the upper elevations. The average wood specific gravity for the upper elevation species was 9.6% greater than that for the species at lower elevations. Aegle marmelos among the lower elevation species and Quercus leucotrichophora among the upper elevation species had the highest wood specific gravity, which were 0.845 ± 0.03 and 0.865 ± 0.02, respectively.     

卧龙自然保护区油竹子（Fargesia angustissima）生理生态特征的海拔变化

全球气候变化（如温度升高）迅速而强劲地影响着高海拔地区的生态环境,致使植物生长发生变化,因此高山地区成为研究植物环境适应性及其对全球气候变化响应的理想区域。为了认清气候变化如何影响竹子的生长和生理生态特性,本文系统调查研究了卧龙自然保护区内大熊猫典型主食竹种油竹子（Fargesia angustissima（Mitford）T.P.Yi）,从其天然分布下限至上限,在卧龙自然保护区沿海拔梯度研究了分株的比叶面积（SpecificLeaf Area,SLA）、基于单位叶面积和单位叶质量的叶氮含量（Narea,Nmass）,以及组织非结构性碳水化合物（NSC）含量及其组分。研究结果表明,各调查因子对海拔的响应均是非线性的,表现为随着海拔升高,各调查值先降后升,高峰值出现在分布上限区域（1 810 m）,而最低值出现在中海拔区域（1 620 m）,反映了环境因子随海拔的非线性变化。分析认为,油竹子的生长受不同海拔环境影响较大。     

Dynamics of nonstructural carbohydrates and biomass yield in a fodder legume tree at different harvest intensities

Tropical tree fodder is harvested by frequent prunings, and resprouting depends on nonstructural carbohydrate reserves in the remaining tree parts. We studied the effects of three pruning intensities (removal of all leaves and branches leaving 1 m of stem once a year (T-12), or every 6 months (T-6), and about 50% pruning every 2 months (P-2)) on regrowth and the dynamics of soluble sugars and starch in the legume tree Gliricidia sepium (Jacq.) Walp. growing under humid tropical conditions in Guadeloupe, Lesser Antilles. Carbohydrates were sampled in roots, stems and branches. Among pruned trees, trees in the T-6 harvest regime had the highest leaf fodder yield (0.73 kg tree(-1) year(-1)). High litter loss reduced leaf yield of T-12 trees, but compared with the other treatments, T-12 trees produced the most branch biomass (3.43 kg tree(-1)). Among treatments, P-2 trees had an intermediate leaf fodder yield and the lowest branch production. Sucrose, glucose and fructose were the most common sugars in all biomass compartments. Mannose, pinitol and an unidentified cyclitol were relatively abundant in branches. Root sugar and starch concentrations were unaffected by harvest regime. There was a significant interactive effect of harvest intensity and regrowth time on stem sugar concentration. Stem starch concentration was highest in T-12 trees. After a year of fodder harvesting, whole-tree reserves of nonstructural carbohydrates were highest in T-12 trees; however, a larger proportion of reserves were located in roots and stems of T-6 and P-2 trees. These reserves, which were not lost in pruning and contributed to regrowth of G. sepium after pruning, may explain the relatively small effects of harvesting regime on soluble sugar and starch concentrations.