核桃叶δ13C、δ15N和碳氮含量对海拔变化的响应

【目的】探明海拔对核桃碳和氮代谢的影响,评估不同海拔范围核桃生长与环境气候因子的关系。【方法】以青藏高原东南部不同海拔(2 500～3 868 m)样地的核桃Juglans regia L.为材料,测定了叶片的稳定碳同位素组成(δ¹³C)、稳定氮同位素组成(δ¹⁵N)和碳氮含量及其比值,进行了各指标间的相关性分析及其与气候因子的逐步回归分析。【结果】核桃叶片δ¹³C值在-27.06‰～-30.03‰之间,平均值为-28.45‰;δ¹⁵N值在2.35‰～7.41‰之间,平均值为5.35‰。随海拔的升高,δ¹³C值呈先升高后降低的趋势,而叶C∶N值呈先降低后增加的趋势,碳含量变化趋势不明显。δ¹⁵N、单位面积氮含量和单位质量氮含量随海拔的升高均呈先升高后降低的趋势,拐点在海拔3 100 m。δ¹⁵N与单位面积氮含量显著正相关(P <0.01),而δ¹³C与碳含量无显著相关性(P> 0.01)。逐步回归分析...     

千年桐雌雄株间形态及生理学差异研究

为了研究千年桐群落的生态结构及不同性别植株的生理学、形态学差异,以期为宜春市千年桐的保护和人工林合理配置提供理论依据。以江西省宜春市袁州区的6个天然千年桐群落植株为试验对象,对其种群的性比格局和雌雄株间生理、形态学差异进行研究。结果表明：本次踏查了6个地区,处于花期的千年桐单株共计316株,其中雌株占比35.75%;雄株占比62.24%,总群体雄株显著偏多(χ²=18.00,P<0.01);且每个千年桐天然林样地中的雌雄比也显著偏离1∶1,雄株偏多。同一踏查地的千年桐的胸径、树高、冠幅、树龄在雌雄植株间差异不显著(P>0.05);但雌雄不同植株的形态指标间两两具有显著相关性;雌株的数量仅与海拔有相关性。千年桐叶片中雌雄不同植株的养分含量均差异显著(P<0.05),且叶片中各类酶的酶活性均为雌株>雄株。韧皮部养分中仅可溶性蛋白含量存在显著差异(P<0.05)。     

罗汉松雌雄植株叶片生理特征比较

以广西南宁市的罗汉松(Podocarpus macrophyllus)为材料,对罗汉松雌雄植株叶片部分生理指标在不同生长季节的差异进行比较。结果表明,所测的各项生理指标在4个生长季节中均产生变化。雌株叶片可溶性糖和可溶性蛋白含量在秋季和冬季均高于雄株,在春季则低于雄株;雌株叶片中的脯氨酸含量在秋季和冬季显著小于雄株,但在春季显著高于雄株;除夏季外,雌株叶片的丙二醛(MDA)含量在其他3个季节均显著高于雄株,过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性在各季节基本表现出小于雄株;雌株叶片中叶绿素含量在春、夏和冬季,以及类胡萝卜素含量在春、秋和冬季均高于雄株叶片。综合分析表明,在反映同化能力和营养物质贮藏的指标上雌株高于雄株,而常用适应性指标表现为雄株优于雌株。研究结果可为罗汉松目标植株在非繁殖季节的选择、应用及抚育管理措施的制定提供理论依据。     

贵州皱叶膏桐和普通膏桐营养转运和光合特征

利用便携式光合仪Li-6400,并结合稳定碳(~(13)C)同位素标记技术,研究分析皱叶膏桐和普通膏桐不同开花期的δ~(13)C值及各光合指标,并分析相互关系。结果表明:不同生长期的枝条叶片δ~(13)C值、Pn、Tr、Gs和Ci的差异均显著,皱叶膏桐叶片δ~(13)C值、Pn、Tr和Gs表现为叶芽期花芽期,Ci表现为花芽期叶芽期,普通膏桐δ~(13)C值、Tr、Gs和Ci均表现为花芽期叶芽期,Pn表现为叶芽期花芽期;皱叶膏桐不同时期δ~(13)C值表现出花芽小花序成熟花序,普通膏桐则正相反。物质能量的转运变化表明,皱叶膏桐在叶芽期将大量同化营养物向叶芽转运用于叶芽向花芽分化发育,普通膏桐在叶芽期将大量同化营养物向花芽转运用于雄花分化发育及花序形态建成。两树种不同时期营养物质的转运变化显示,叶片和花序δ~(13)C值含量的差异与雌雄花开花量相关,较高δ~(13)C值的花序具有较高雌雄花比例。初步推测膏桐控制雌花分化的阶段可能发生在叶芽向花芽转化期,控制开花量的阶段可能发生在花序发育时期。     

中华猕猴桃雌雄株可溶性糖含量的差异

用蒽酮比色法对中华猕猴桃雌株和雄株,分别进行了叶片中可溶性糖含量的测定。结果表明,雄株的可溶性糖含量极显著地高于雌株,并证明在未开花的实生苗中也存在同样的现象,从而为中华猕猴桃苗期的雌雄鉴别找到了一种依据。     

短期增温对亚热带杉木幼林碳氮同位素组成的影响

[目的]利用稳定同位素手段,研究增温对氮循环和水分利用效率的影响,揭示亚热带杉木幼苗水分利用效率和氮饱和状态对气候变暖的响应。[方法]在福建省三明市森林生态系统与全球变化研究站,布设发热电缆对土壤进行增温,设置对照(CT)和增温(W)两种实验小区。分别采集不同处理杉木叶片和0～10 cm表层土壤,测定植物-土壤碳、氮稳定同位素,以及碳和氮含量。[结果]显示:增温处理叶片δ~(15)N值(0. 40‰)比对照处理(-2. 79‰)显著增加了3. 19‰;但增温后叶片氮含量只是略有上升,与对照无显著差异;增温后,表层土壤δ~(15)N值显著上升,叶片15N的富集指数比对照处理更接近于0;增温与对照处理叶片δ~(13)C值分别为-29. 35‰和-29. 08‰,无显著差异;叶片δ~(13)C与δ~(15)N之间存在显著正相关关系。[结论]温度是中亚热带地区氮循环的一个重要影响因素,增温促使杉木叶片和表层土壤δ~(15)N值显著提高,碳代谢与氮代谢相辅相成。     

山茶属植物叶片δ~(13)C和δ~(15)N与养分含量及光合作用的关系

以11个山茶属植物为试材,测定其叶片稳定碳氮同位素比率(δ13C和δ15N)、光合参数(净光合速率、气孔导度、蒸腾速率、胞间CO2浓度、水分利用效率)以及叶片C、N、P和K元素含量等指标,分析叶片δ13C和δ15N与光合参数、养分含量之间的关系。结果表明:11个山茶属植物δ13C值在-26.38‰~-29.44‰之间,其中多齿红山茶显著高于其它10个物种;δ15N在-3.01‰~4.27‰之间,其中以攸县油茶最高,石果红山茶最低;普通油茶净光合速率最高,是其它10个茶属植物种的124%~496%;11个山茶属植物叶片C含量在47.72%~53.07%之间,叶片N、P含量分别以攸县油茶和普通油茶最高,各物种叶片K含量差异不大;叶片δ13C与C和大量元素含量、光合速率均呈负相关关系,其中与N、P含量以及胞间CO2浓度呈极显著负相关,与水分利用效率呈正相关;δ15N与N、P含量显著正相关,与K含量显著负相关,与光合参数呈正相关,与水分利用效率呈负相关。     

两种彗星兰应对干旱胁迫的光合碳同化途径响应

为更加科学和高效地对两种彗星兰进行繁育工作,开展了叶片厚度、稳定碳同位素分馏值(δ¹³C)、昼夜净CO₂吸收动态和可滴定酸含量检测,探索该两物种应对干旱胁迫的光合碳同化途径响应。结果表明：长距彗星兰(Angraecum sesquipedale)的叶片厚度为1.07 mm、δ¹³C值为-17.24‰;拟长距彗星兰(A.sororium)的叶片厚度为0.58 mm、δ¹³C值为-27.74‰;长距彗星兰仅在暗期吸收CO₂并随着干旱胁迫的加深其净CO₂吸收速率也逐渐降低;拟长距彗星兰在未干旱条件下主要在昼期吸收CO₂,但在干旱处理后的第2～5天暗期净CO₂吸收速率转变为正值并逐渐增加;长距彗星兰的早晚可滴定酸含量差值由未干旱的47 mmol·g^-1降低至干旱胁迫的22 mmol·g^-1,而拟长距彗星兰的由39 mmol·g^-1增加至60 mmol·g<...     

林龄、叶龄对亚热带杉木人工林碳氮稳定同位素组成的影响

[目的]研究5个不同林龄(3,8,14,21,46年)、不同叶龄(当年生、1年生、2年生、3年生)杉木人工林叶片的碳、氮稳定同位素组成,并根据它们对氮循环等过程的指示作用来探索不同林龄、叶龄杉木人工林氮循环过程及氮饱和程度的差异,从而为不同生长阶段杉木人工林制定施肥措施提供科学依据.[方法]以福建南平峡阳林场5块相互毗邻的不同林龄杉木人工林为研究对象,在每个林龄的林分内分别设置4个20 m ×20 m的试验小区.分别采集不同林龄杉木活叶并根据“主干法”将采集的杉叶分为不同叶龄,然后在每个小区内采集0 ～ 10 cm深度土层的土样,用同位素质谱仪测定它们的碳、氮稳定同位素组成(δ13C,δ15N),用碳氮元素分析仪测定叶片氮含量,叶片15N富集指数由叶片δ15N值减去相应的土壤δ15N得到.[结果]叶片δ15N值的变化范围为-2.52‰～2.81‰,叶片氮含量的变化范围为7.72％～13.5％,二者在不同林龄间均具有极显著差异,并均呈现出幼林和老林较高、处于速生期的林分较低的趋势,且叶片δ15N值与叶片氮含量之间存在显著的相关性,但不同叶龄叶片δ15N值间则不具有显著差异;不同林龄叶片的15N富集指数存在显著差异,呈现出幼林与老林叶片15N富集指数较接近于0的趋势;叶片δ13C的变化范围为-29.93‰～-27.88‰,不同林龄间差异不显著,但同一林龄不同叶龄叶片的δ13C则有显著差异,且有随着叶龄增大而减小的趋势.[结论]不同林龄叶片δ13C差异不显著但呈现幼年较低的趋势,可能是不同树高导致不同林龄杉木水分利用效率间的差异所致,而同一林龄不同叶龄杉木δ13C间的显著差异则可能是光合作用效率不同造成的.不同林龄在叶片δ15N、叶片氮含量、叶片15N富集指数间的显著差异均指示出处于速生期的林分氮饱和程度显著低于幼林和老林,这说明虽然我国亚热带地区氮沉降现象严重,但氮素仍是限制处于速生期杉木人工林生长的因素.     

美洲黑杨及其杂种F1不同生长势无性系叶片δ13C和氮素利用效率

以美洲黑杨不同生长势的杂种F1及其亲本为研究对象,通过测定不同冠层叶片的稳定碳同位素组成和碳、氮含量,分析不同生长势美洲黑杨叶片水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明:不同生长势杂种F1的δ13C值及碳、氮含量均差异显著,超亲杂种F1在每个冠层的叶片δ13C值和碳、氮含量均显著大于低亲杂种Ft;美洲黑杨叶片δ13C值与氮含量呈极显著正相关(R2=0.941);叶片δ13C和碳、氮含量存在空间分布差异,树冠上层显著大于中层,下层最小;超亲杂种F1的水分利用效率(WUE)高于低亲杂种F1,但低亲杂种F1的氮素利用效率(NUE)极显著大于超亲杂种F1,WUE和NUE之间呈现显著负相关(R2=-0.652),说明美洲黑杨叶片的水分利用效率和氮素利用效率存在一定的制约关系.     

Impact of vegetation succession on leaf-litter-soil C:N:P stoichiometry and their intrinsic relationship in the Ziwuling Area of China's Loess Plateau

Long-term natural vegetation succession plays a substantial role in the accumulation and distribution of plant and soil C:N:P stoichiometry.However,how plant and soil C:N:P relationships or ratios change along with successional stages over a century in the severely eroded areas remain unclear.These were measured over a 100-year natural succession in five successional stages from annual grasses to climax forests.The results show that natural succession had significant effects on carbon(C),nitrogen(N) and phosphorous(P) concentrations in leaf-litter-soil and their ratios in severely eroded areas.Nitrogen concentrations and N:P ratios in leaf and litter increased from annual grasses to the shrub stage and then decreased in the late successional forest stages.Leaf P levels decreased from annual grasses to shrub stages and did not significantly change during late successional stages.Litter P concentration decreased in the early successional stages and increased during late successional stages,with no overall significant change.Soil C and N concentrations and C:N,C:P and N:P ratios increased with successional stages.Soil C and N concentrations decreased with the increasing soil depth.Both were significantly different between any successional stages and controls(cropland) in the upper 10 cm and 10-20 cm soil layers.Leaf N:P ratios may be used to indicate nutrient limitations and this study suggests that plant growth during the grass stages was limited by N,during the shrub stage,by P,and during the forest stages,by both of N and P.In addition,there were close correlations between litter and leaf C:N:P ratios,soil and litter C and N levels,and C:P and N:P ratios.These results show that long-term natural vegetation succession is effective in restoring degraded soil properties and improving soil fertility,and provide insights into C:N:P relationships of leaf,litter and soil influenced by vegetation succession stage.     

Gas exchange characteristics of mangosteen (Garcinia mangostana L.) leaves

We used the plastochron index to study the relationship between plant age, leaf age and development, and net photosynthesis of black cherry (Prunus serotina Ehrh.) seedlings. Leaf area and net photosynthesis were measured on all leaves >/= 75 mm of plants ranging in age from 7 to 20 plastochrons. Effects of plant developmental stage on leaf area and net photosynthesis were evaluated for leaves of differing age (horizontal series), leaves on plants of constant age (vertical series), and leaves of constant age (oblique series). Regression techniques were used to estimate leaf area from leaf blade dimensions. The best equations for predicting leaf area had R(2) values of 0.991-0.992 and used linear or logarithmic functions of both leaf length and width. Suitable, but less precise, equations with R(2) values of 0.946-0.962 were developed from either leaf length or leaf width. Leaf area development in black cherry seedlings was similar to that in other indeterminate species. Leaves of young plants reached full expansion at a lower leaf plastochron age than leaves of older plants. Maximum net photosynthesis per unit leaf area occurred 2-3 plastochrons before full leaf expansion. There was strong ontogenetic drift in net photosynthesis with leaf age; net photosynthesis decreased as plant age increased in leaves of the same plastochron age. Plots of the oblique series were particularly useful in providing information about interaction effects.     

Variation in Leaf Shapes of Nitraria Species and Effect on Leaf δ13C

Leaf shapes are not only the useful indicators in plant taxonomy,but also the important factors affecting energy and material exchange in leaves.In this paper,we collected and scanned the leaves of Nitraria tangutorum in Dengkou of Inner Mongolia Autonomous Region（the mean annual precipitation 145 mm） and Minqin of Gansu Province （the mean annual precipitation 115 mm） and N. sphaerocarpa in Dunhuang,and then analyzed leaf shape parameters with Image - Pro Plus6.0 image processing software and leafδ¹³C values in the isotope laboratory of the Chinese Academy of Forestry.The result showed that:1) as leaf area increased with increasing water availability the increases in the leaf length and width were asynchronously;2) with the same leaf width,the 1 eaves of N.tangutorum and N.sphaerocarpa were significantly longer in high water available conditions;and 3) although there were significantly differences in water availability between Dengkou and Minqin,as well as between the bottom and middle of the alluvial fan near the East Lake in Dunhuang,the leafδ¹³C values of N.tangutorum or N.sphaerocarpa were similar in different water conditions（P>0.05）.Our results suggested that the ratio of leaf perimeter to area would be an important factor which linked leaf shape to plant water physiology.During growing procedure of leaf area,leaf length increase was prior to its width to alleviate the reduction in ratio of perimeter to area and maintain water use efficiency of the plant.     

Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

磷施肥水平对油茶植株营养生长的影响

为了提供科学合理的施肥依据,在氮、钾肥最佳用量的基础上,依次设置5个磷水平处理,对油茶幼林开展施磷肥试验。结果表明:低于每年900 g/株磷肥水平时,油茶植株的株高、地径、冠幅乘积增幅、叶绿度、叶面积及叶片养分N、P、K随着磷素供应水平的增加而增大;在每年900 g/株磷肥水平下,各项指标达到最大值;大于每年900 g/株磷肥水平时,各项生长指标没有增加反而降低了。说明过度施入磷肥,导致植株磷肥利用率的降低,影响植株体内干物质积累,不利于植株生长;不同磷水平处理中,对油茶各指标的影响呈先增大后减少的趋势。叶片养分与油茶树体指标间呈极显著相关性,施磷肥与各项指标关系密切。油茶植株营养生长对磷水平的响应,说明合理施用磷肥有利于油茶植株树体的生长和叶片养分的累积。     

Morphological and physiological plasticity response to low nitrogen stress in black cottonwood(Populus deltoides Marsh.)OA

It is important to evaluate nitrogen use efficiency and nitrogen tolerance of trees in order to improve their productivity. In this study, both were evaluated for 338 Populus deltoides genotypes from six provenances. The plants were cultured under normal nitrogen(750 μM NH₄ NO₃) and low nitrogen(5 μM NH₄ NO₃) conditions for 3 months. Growth, chlorophyll content and glutamine synthetase activity of each genotype were measured. Under low nitrogen, height...     

霞浦县阔叶混交林生态系统碳、氮储量格局

依据全国碳汇专项调查的理论和方法,对福建省霞浦县不同林龄阔叶混交林生态系统各组分的碳、氮含量及碳、氮储量格局进行调查分析,结果表明:灌木层各器官碳含量从大到小依次为枝干根叶,氮含量为叶干根枝;草本层碳、氮含量从大到小均为地上部分地下部分;土壤碳、氮含量均随土层深度增加而降低,随林龄的增大而上升;系统各组分C/N从大到小依次为枝根干枯落物叶土壤;12年生、19年生和28年生阔叶混交林生态系统的碳储量分别为164.066、231.751和290.985t!hm-2,氮储量分别为15.011、23.503和31.236t!hm-2,其中,土壤层碳储量所占比重分别为60.27%、46.50%和39.50%,氮储量所占比重分别为45.94%、33.09%和28.67%;乔木层、灌木层、枯落物层和土壤层碳、氮储量均随林龄的增大而增加。     

Effect of elevated CO2 on monoterpene emission of young Quercus ilex trees and its relation to structural and ecophysiological parameters

We investigated growth, leaf monoterpene emission, gas exchange, leaf structure and leaf chemical composition of 1-year-old Quercus ilex L. seedlings grown in ambient (350 microl l(-1)) and elevated (700 microl l(-1)) CO2 concentrations ([CO2]). Monoterpene emission and gas exchange were determined at constant temperature and irradiance (25 degrees C and 1000 micromol m(-2) s(-1) of photosynthetically active radiation) at an assay [CO2] of 350 or 700 microl l(-1). Measurements were made on intact shoots after the end of the growing season between mid-October and mid-February. On average, plants grown in elevated [CO2] had significantly increased foliage biomass (about 50%). Leaves in the elevated [CO2] treatment were significantly thicker and had significantly higher concentrations of cellulose and lignin and significantly lower concentrations of nitrogen and minerals than leaves in the ambient [CO2] treatment. Leaf dry matter density and leaf concentrations of starch, soluble sugars, lipids and hemi-cellulose were not significantly affected by growth in elevated [CO2]. Monoterpene emissions of seedlings were significantly increased by elevated [CO2] but were insensitive to short-term changes in assay [CO2]. On average, plants grown in elevated [CO2] had 1.8-fold higher monoterpene emissions irrespective of the assay [CO2]. Conversely, assay [CO2] rapidly affected photosynthetic rate, but there was no apparent long-term acclimation of photosynthesis to growth in elevated [CO2]. Regardless of growth [CO2], photosynthetic rates of all plants almost doubled when the assay [CO2] was switched from 350 to 700 microl l(-1). At the same assay [CO2], mean photosynthetic rates of seedlings in the two growth CO2 treatments were similar. The percentage of assimilated carbon lost as monoterpenes was not significantly altered by CO2 enrichment. Leaf emission rates were correlated with leaf thickness, leaf concentrations of cellulose, lignin and nitrogen, and total plant leaf area. In all plants, monoterpene emissions strongly declined during the winter independently of CO2 treatment. The results are discussed in the context of the acquisition and allocation of resources by Q. ilex seedlings and evaluated in terms of emission predictions.     

猕猴桃雌雄植株过氧化物酶同工酶的研究

对猕猴桃(Actinidia)雌雄植株叶片中过氧化物酶同工酶的电泳分析测定,结果表明,雌雄株之间同工酶的酶谱存在着明显差异。酶谱的变化与叶片生长量、植株的生育期、不同种类有关。其酶带变化的总趋势是,在开花期雌株在负极一端比雄株多1条酶带,植株进入结果期,其结果相反,雄株比雌株多1条酶带。中华猕猴桃(Actinidia chinensis planch)。同工酶的酶谱还表现出,靠正极,在结果期比开花期多1条酶带。同时用未开花植株进行测定,其酶谱变化与上述雌雄株有相似结果,在靠负极一端雌株多1条酶带,有时雄株靠正极端不出现酶带。     

Sex-related differences in leaf morphological and physiological responses in Hippophae rhamnoides along an altitudinal gradient

In most woody plants, leaf morphological and physiological characteristics are extremely variable across environmental gradients, particularly across altitudinal gradients. Hippophae rhamnoides L., a dioecious and deciduous shrub species, occupies a wide range of habitats in the Wolong Nature Reserve, southwest China. We measured growth, sex ratio and morphological and physiological characteristics of leaves in male and female H. rhamnoides individuals along an altitudinal gradient. Shoot height (HT), leaf N concentration per unit dry mass (N(mass)), leaf N concentration per unit area (N(area)) and leaf carbon isotope composition (delta(13)C) were higher in males than in females, whereas females had higher specific leaf area (SLA), stomatal length (SL) and stomatal index (SI) (i.e., total stomatal length per unit leaf area) than males along the altitudinal gradient. Females also had higher values of stomatal density (SD) at all altitudes except 2800 m. The male:female ratio (MFR) was biased toward males at all altitudes except at 2800 m. Changes in HT, MFR, SLA, SD, SL, SI, N(mass), N(area) and delta(13)C along the altitudinal gradient were nonlinear. Below 2800 m, HT, SLA, SD, SL and SI increased with increasing altitude, but above 2800 m they decreased with increasing altitude. In contrast, MFR, N(mass), N(area) and delta(13)C showed the opposite patterns with altitude. Consequently, we confirmed our hypotheses: (1) stressful environments have a more negative impact on females than on males in a variety of ways; (2) under optimal growth conditions the sex ratio is even, but becomes male-biased as resources become limited; and (3) there is an optimum altitudinal range at around 2800 m for the growth of H. rhamnoides in the Wolong Nature Reserve.