紫茎泽兰和飞机草的形态和光合特性对磷营养的响应

比较研究了紫茎泽兰和飞机草的生长、形态、生物量分配和光合特性对磷营养的可塑性反应,及与其入侵性的关系.结果表明,两种入侵植物对磷营养变化表现出很强的可塑性和适应性.低磷时,两种植物的根生物量比增大,利于养分吸收;高磷时,两种植物的比叶面积、最大净光合速率、光饱和点、单位面积的叶绿素和类胡萝卜素含量增大,同化面积和同化能力增强,利于碳积累.相比之下,紫茎泽兰对磷的适应性更强.随供磷量的增加,紫茎泽兰和飞机草的相对生长速率、总生物量、株高、分枝数、叶面积指数和最大净光合速率均显著增大,过量磷素对上述参数抑制较小,表明两种入侵植物偏好较高的磷环境,土壤磷含量升高有利于其入侵,并在高磷时,通过增大株高、分枝数和叶面积指数荫蔽排挤本地种.在本地种基本停止生长的干季,紫茎泽兰和飞机草仍维持较高的相对生长速率.这也与其入侵性密切相关.     

光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响

 比较研究了不同光强下生长的（透光率分别为12.5%、36%、50%、100%）两种入侵性不同的外来种——紫茎泽兰（Eupatorium adenophorum）和兰花菊三七（Gynura sp.）的生物量分配、叶片形态和生长特性。结果表明: 1）两种植物叶片形态对光环境的反应相似。弱光下比叶面积（SLA）、平均单叶面积（MLS）和叶面积比（LAR）较大,随着光强的升高,SLA、MLS、LAR和叶根比（LARMR）降低。2）100%光强下紫茎泽兰叶生物量比（LMR）、叶重分数（LMF）和叶面积指数高于低光强下的值,也高于兰花菊三七,支持结构生物量比（SBR）则相反。强光下紫茎泽兰叶片自遮荫严重,这可能是其表现入侵性的重要原因之一;兰花菊三七分枝较多,避免了叶片自遮荫,较多的分枝利于种子形成对其入侵有利。3）随生长环境光强的升高,两种植物的净同化速率（NAR）、相对生长速率（RGR）和生长对NAR的响应系数均升高（但100%光强下兰花菊三七RGR降低）,平均叶面积比（LARm）和生长对LARm的响应系数均降低,但不同光强下LARm对生长的影响始终大于NAR。4）随着光强的减弱,两种植物都增加高度以截获更多光能,但它们的生物量分配策略不同,紫茎泽兰根生物量比（RMR）降低,SBR增大,而兰花菊三七SBR降低,RMR增大。紫茎泽兰的生物量分配策略更好的反应了弱光环境中的资源变化情况。结论：紫茎泽兰对光环境的适应能力强于兰花菊三七。     

氮肥和种植密度对紫茎泽兰生长和竞争的影响

紫茎泽兰（Ageratina adenophora）是我国西南热带亚热带地区危害最严重的外来入侵植物之一。本研究在野外设置不同氮肥供给水平和种植密度下的紫茎泽兰、拔毒散（Sida szechuensis）、伏生臂形草（Brachiaria decumbens）和非洲狗尾草（Setaria anceps）的单种和紫茎泽兰与其他3种植物混种的人工群落,从植被-土壤相互作用的角度分析紫茎泽兰的入侵机制。结果表明：单种时4种受试植物均表现了对氮营养的响应,随氮肥水平的升高,根冠比降低,比叶面积升高;通过比较总生物量增幅,氮响应能力非洲狗尾草>伏生臂形草>紫茎泽兰>拔毒散。混种时受试植物与紫茎泽兰的生物量比和株高也表明非洲狗尾草的竞争能力>伏生臂形草>紫茎泽兰>拔毒散;与拔毒散混种的紫茎泽兰以及与 紫茎泽兰混种的伏生臂形草和非洲狗尾草的种内竞争随种植密度的增大而增大,导致种间竞争能力下降,在高种植密度下对混种物种的抑制作用减弱。紫茎泽兰在氮响应能力、竞争能力和表型可塑性方面均强于本地种,可能与其入侵机制有关;牧草的氮响应能力和竞争能力强于紫茎泽兰,其中非洲狗尾草又强于伏生臂形草,可能更适宜于入侵地的替代控制。     

西双版纳外来入侵植物及其共存种叶片氮、磷化学计量特征

诊断外来植物和本地种的资源利用方式是入侵生态学研究的热点问题。叶片氮(N)、磷(P)含量和化学计量特征可以反映受侵地区植物的N、P吸收能力和限制状况,为把握外来植物的入侵能力、为本地植物共存或消失的机制提供基础科学依据。该研究以我国西南地区典型外来入侵植物(飞机草(Chromolaena odorata)、紫茎泽兰(又叫破坏草,Ageratina adenophora))及其共存的本地植物为对象,探讨不同入侵条件下入侵种及其共存的本地植物的N、P利用策略。在云南西双版纳孔明山研究区,调查了无入侵条件下和飞机草与紫茎泽兰的不同入侵程度(按入侵种生物量比例划分)下的物种数量和生物量,分析了主要植物的叶片N、P含量和N:P值。结果显示:尽管群落地上生物量随飞机草和紫茎泽兰入侵程度增加而增加,但本地植物种数随飞机草和紫茎泽兰入侵程度增加而显著减少。飞机草与紫茎泽兰的叶片N、P平均含量无显著差异,但均显著高于无入侵条件下的本地植物以及与其共存的本地植物。两种入侵植物的N、P含量均随它们所占样方总生物量的比例增大而升高,本地植物N含量也有相似的变化趋势。当对比入侵和无入侵两类样方的同种植物N、P和N:P变化时发现多数本地种叶片P含量呈降低趋势,N含量和N:P呈升高趋势。根据叶片N、P绝对含量和N:P值及其随入侵的变化规律,推测入侵可能提高了植物的N可利用性,但本地植物仍然受N限制;入侵植物N:P值总体小于10与其具有相对于N吸收的较高的P吸收能力有关。该研究揭示了西南典型外来入侵植物具有较强的N、P吸收富集能力。     

人工群落中苗期紫茎泽兰的化感作用和对光环境的适应

在有、无活性炭的条件下分别构建紫茎泽兰（Ageratinaa denophora）与4种受体植物——飞机草（Chromolaena odorata）、鬼针草（Bidenspilosa）、胜红蓟（Ageratum conyzoides）和兰花菊三七（Gynura sp.）混种的人工群落,研究了群落中紫茎泽兰的化感作用和对群落光环境的适应,探讨了化感作用和光适应特性与其入侵性的关系。结果表明,活性炭处理对4种受体植物的生长、生理特性影响不显著,说明苗期紫茎泽兰化感作用不明显,推测入侵初期化感作用不是紫茎泽兰排挤本地种的主要原因。4种受体植物可以通过化感作用对紫茎泽兰产生某些影响,但群落的光环境对其影响更大。紫茎泽兰能很好地适应群落中不同的光环境。苗期紫茎泽兰处于群落下层,叶片受光指数低,此时它能长期忍耐并缓慢生长;随着叶片受光指数的升高,其最大净光合速率、超氧化物歧化酶活性、叶绿素a／b比、总生物量、总叶面积、地茎、叶片数和分支数升高,比叶面积和比茎长降低,这有利于它维持叶片能量平衡并导致对邻近植物的严重遮荫。紫茎泽兰强的光适应能力、强光下对其它物种的遮荫效应与其入侵性密切相关。     

氮、磷养分对飞机草营养器官表型可塑性的影响

采用温室盆栽试验研究了不同氮、磷水平对入侵植物飞机草(Chromolaena odorata)营养器官表型可塑性的影响。结果表明:随着氮、磷水平的上升,飞机草的分枝数量、分枝长度、叶片数、总叶面积、总生物量以及茎、叶器官生物量显著增加。飞机草的根生物量比、根冠比随着氮、磷水平的升高显著下降;茎生物量比在供氮(磷)量达0.05 g·kg-1时显著增加,之后保持稳定;叶生物量比随氮水平的增加先降后升,但其受磷水平变化的影响较小。叶面积比、叶根比、比叶面积和平均相对生长速率随着氮、磷水平的上升显著增加,但叶面积比、叶根比和比叶面积在供磷量≥0.05 g·kg-1时的差异不明显。飞机草的分枝数量、分枝长度、叶片数、总叶面积、根生物量比、根冠比、叶根比以及茎、叶与植株总生物量等指标的可塑性指数较高,并且对氮素的响应更强。表明氮、磷水平能够显著影响飞机草的植株生长,飞机草亦能够通过植株形态、结构以及生物量积累与分配的调整来适应多变的养分环境,并表现出较高的可塑性。     

两种入侵植物与三种本地植物根系特征的比较研究

以菊科2种入侵植物飞机草(Chromolaena odorata(L.)R.M.King&H.Rob)和紫茎泽兰(Eupatorium adenophorum Speng)以及生活型相似的3种本地植物异叶泽兰(Eupatorium heterophyllum DC.)、佩兰(Eupatorium fortune Turcz.)和白头婆(Eupatorium japonicum Thunb.)为研究对象,以单种为对照,分析5种植物的株高、生物量以及细根形态和结构特征对混种(竞争)的响应。结果显示,在单种情况下,2种入侵植物的株高和地上生物量比3种本地植物高。2种入侵植物的株高和地上生物量在单种和混种下无显著差异,但在混种下3种本地植物的株高、地上生物量、根生物量、细根长度、细根表面积和体积与单种相比显著降低。在混种情况下,2种入侵植物的根组织密度显著降低,但比根长和比根面积仅在紫茎泽兰中显著增加。5种植物在单种和混种下的株高与细根长度、根生物量呈显著正相关,地上生物量与细根体积、表面积呈显著正相关。主成分分析结果表明入侵植物位于株高和地上生物量更高的一端。说明在与本地植物竞争的过程中,飞机草和紫茎泽兰能通过调整根系碳投资策略实现更强的竞争力。     

不同养分水平对飞机草生长与生物量分配的影响

采用温室盆栽模拟实验研究了不同养分水平对外来入侵植物飞机草(Chromolaena odorata)的生长性状、生物量积累以及生物量分配格局的影响。实验共设置5种养分浓度处理,分别为Hoagland标准营养液的10%、25%、50%、100%和200%溶液。结果表明:养分水平对飞机草植株的生长性状以及生物量的积累、分配产生显著影响。随着养分水平的增加,飞机草的总分枝数量、长度以及一级分枝数量、长度持续增加,并且100%、200%处理还能促进二级分枝的萌发生长。飞机草的叶片数、总叶面积、总生物量以及茎、叶两器官生物量随养分水平的上升显著增加,但株高、根生物量不受养分浓度变化的影响。根生物量比、根冠比随养分水平的提高显著下降,叶生物量比则显著上升,但茎生物量比在各养分浓度保持稳定。叶面积比、叶根比、RGR亦随养分含量的上升显著增加。说明养分资源丰富的环境将促进飞机草的地上部生长,而生物量分配格局的变化可能是其在入侵蔓延过程中适应养分异质性生境的重要生态策略。     

外来入侵植物飞机草和本地植物异叶泽兰对大气CO_2浓度升高的响应

大气CO2浓度升高影响外来植物入侵,研究外来入侵植物和本地植物对大气CO2浓度升高响应的差异,有助于准确预测和管理外来植物入侵。基于封顶式CO2生长室,模拟大气CO2浓度变化(对照和700μmol/mol),比较研究了外来入侵植物飞机草(Chromolaena odorata)和本地植物异叶泽兰(Eupatorium heterophyllum)形态、生长、生物量分配和光合特性对大气CO2浓度升高响应的差异。结果表明:(1)在当前大气CO2浓度下,飞机草总生物量、株高、基径和总叶面积高于异叶泽兰,分枝数低于异叶泽兰;CO2浓度升高,飞机的总生物量、株高、基径、分枝数和总叶面积分别增加了92%、41%、60%、325%和148%,高于异叶泽兰的32%、14%、30%、64%和79%,飞机草生长优势进一步提高。(2)无论在高或低CO2浓度下,飞机草根生物量分数(RMF)都低于异叶泽兰,叶生物量分数(LMF)和茎生物量分数(SMF)都高于异叶泽兰;CO2倍增两种植物RMF均降低,LMF和SMF均升高,但这2个参数对CO2倍增响应的种间差异不显著。(3)无论在高或低CO2浓度下,飞机草和异叶泽兰的净光合速率差异均不显著,CO2倍增对两种植物的净光合速率的促进作用相似。上述结果表明,在未来大气CO2浓度升高的条件下,飞机草的入侵性可能提高,入侵危害将加剧。     

光强对三叶鬼针草生长特征的影响

利用温室盆栽试验,研究了不同光强100%、40%、20%和5%下三叶鬼针草营养生长期和繁殖期的生长特征。结果表明:中度遮荫有利于三叶鬼针草支持结构的生长和营养期的物质积累,重度遮荫下仍能生长良好。在两个生长时期,相对生长速率(RGR)、净同化速率(NAR)均在100%光强下最大,5%光强下最小;总生物量在营养期和繁殖期分别在40%光强和全光照下最大,在3%光强下最小;株高、总叶面积(TLA)、根生物量比(RMR)、根冠比(R/C)、叶生物量比(LMR)、比叶面积(SLA)、叶面积比(LAR)、平均叶面积比(LARm)在3%光强时均大于全光照下的;支持结构生物量比(SBR)在40%光强、20%光强下大于3%光强和全光照处理。这说明三叶鬼针草在形态、生物量分配及生长特性上对光因素具较强的可塑性,这可能是其分布范围广、具有强入侵性的因素之一。     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Net Assimilation Rate Determines the Growth Rates of 14 Species of Subtropical Forest Trees

Growth rates are of fundamental importance for plants, as individual size affects myriad ecological processes. We determined the factors that generate variation in RGR among 14 species of trees and shrubs that are abundant in subtropical Chinese forests. We grew seedlings for two years at four light levels in a shade-house experiment. We monitored the growth of every juvenile plant every two weeks. After one and two years, we destructively harvested individuals and measured their functional traits and gas-exchange rates. After calculating individual biomass trajectories, we estimated relative growth rates using nonlinear growth functions. We decomposed the variance in log(RGR) to evaluate the relationships of RGR with its components: specific leaf area (SLA), net assimilation rate (NAR) and leaf mass ratio (LMR). We found that variation in NAR was the primary determinant of variation in RGR at all light levels, whereas SLA and LMR made smaller contributions. Furthermore, NAR was strongly and positively associated with area-based photosynthetic rate and leaf nitrogen content. Photosynthetic rate and leaf nitrogen concentration can, therefore, be good predictors of growth in woody species.     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.     

Contribution of root growth responses to leaf traits and relative growth rate of Populus alba under different water-table conditions

Water-table depth variations alter root growth response and may affect whole-plant growth in arid and semi-arid regions. We examined how root biomass allocation and root morphological traits affect the leaf physiological and morphological traits and whole-plant growth of Populus alba growing under different water tables. We exposed 1-year-old P. alba cuttings to contrasting soil–water conditions via water table changes in a greenhouse for 90 days. We examined relationships among net assimilation rate (NAR) and other growth components obtained from our published data for trees harvested every 30 days. Strongly negative correlations were found between RMR and root morphological traits. Root mass ratio had a strong negative relationship with LMR, and proportion of fine-root biomass per total root biomass was positively correlated with SLA and NAR. Both NAR and leaf area ratio were important determinants of variation in relative growth rate (RGR). Leaf mass ratio (LMR) and specific leaf area (SLA) were positively correlated with RGR; the correlation was stronger in the case of LMR. Along a water-table gradient, negative relationships between root growth responses are likely to indirectly influence RGR through changes in NAR, LMR, and SLA.     

Seedling growth strategies in Bauhinia species: comparing lianas and trees

BACKGROUND AND AIMS: Lianas are expected to differ from trees in their growth strategies. As a result these two groups of woody species will have different spatial distributions: lianas are more common in high light environments. This study determines the differences in growth patterns, biomass allocation and leaf traits in five closely related liana and tree species of the genus Bauhinia. METHODS: Seedlings of two light-demanding lianas (Bauhinia tenuiflora and B. claviflora), one shade-tolerant liana (B. aurea), and two light-demanding trees (B. purpurea and B. monandra) were grown in a shadehouse at 25% of full sunlight. A range of physiological, morphological and biomass parameters at the leaf and whole plant level were compared among these five species. KEY RESULTS: The two light-demanding liana species had higher relative growth rate (RGR), allocated more biomass to leaf production [higher leaf mass fraction (LMF) and higher leaf area ratio (LAR)] and stem mass fraction (SMF), and less biomass to the roots [root mass fraction (RMF)] than the two tree species. The shade-tolerant liana had the lowest RGR of all five species, and had a higher RMF, lower SMF and similar LMF than the two light-demanding liana species. The two light-demanding lianas had lower photosynthetic rates per unit area (A(area)) and similar photosynthetic rates per unit mass (A(mass)) than the trees. Across species, RGR was positively related to SLA, but not to LAR and A(area). CONCLUSIONS: It is concluded that the faster growth of light-demanding lianas compared with light-demanding trees is based on morphological parameters (SLA, LMF and LAR), and cannot be attributed to higher photosynthetic rates at the leaf level. The shade-tolerant liana exhibited a slow-growth strategy, compared with the light-demanding species.     

Relative growth rate and biomass allocation in ten woody species with different leaf longevity using phylogenetic independent contrasts (PICs)

In this study, we compare the relative growth rate (RGR) and biomass allocation of 10 woody species (5 deciduous and 5 evergreen) from the Mediterranean region using phylogenetic independent contrasts (PICs) to test if these two functional groups differ in these traits. In general, the results were similar when using PICs or without taking into account phylogenetic relations. Deciduous species had a higher RGR than evergreen species, due to the higher net assimilation rate (NAR). Deciduous species had a higher specific leaf area (SLA) but a lower leaf mass ratio (LMR), resulting in a similar LAR for deciduous and evergreen species (LAR = SLA x LMR). In some cases, the use of PICs revealed patterns that would not have appeared if phylogeny had been overlooked. For example, there was no significant correlation between RGR and final dry mass (after 4 months of growth) but PICs revealed that there was a positive relation between these two variables in all deciduous-evergreen pairs. In general, RGR decreased with time and this temporal variation was due primarily to NAR variations (r = 0.79, p < 0.01), and also to variations in LAR (r = 0.69, p < 0.05). Considering the phylogeny, the only variable constantly different for all deciduous-evergreen pairs was SLA. This result, and the fact that SLA was the best correlated variable with RGR (r = 0.81, p < 0.01), reinforce the value of SLA as a variable closely associated to growth and to the functional groups (deciduous vs. evergreen).     

Convergent succession of plant communities is linked to species’ functional traits

Functional traits reflecting the resource economy and growth strategy of plants vary widely both within and among ecosystems. Theory suggests that trait variation within a community may determine the relative abundance of species, though this idea requires more empirical support.We set up a long-term succession experiment in a nutrient-poor wetland, planting seedlings of twelve fenland species in different relative abundances and absolute densities, thereby creating 24 communities. The biomass of these species and the soil water and nutrient status of the system were monitored over ten years. Using these data, we could relate the changing relative abundance of species to five traits – leaf dry matter content (LDMC), leaf nitrogen concentration (LNC), specific leaf area (SLA), relative growth rate (RGR), and seed mass (SM).The initial communities converged after ten years to a common dominance–diversity structure, with two species accounting for 82% of total biomass. Soil water and nutrient conditions remained largely constant. By the end of the experiment, community trait structure had changed so that species functional traits were significantly related to their relative abundance. The most abundant species had high LDMC and SM, but low RGR and SLA, and varied little in LNC, suggesting that investment in leaf structure and retention of nutrients were most important for species dominance under low nutrient conditions. Our results provide experimental evidence that dominance–subdominance structures in plant communities are governed by functional traits.     

Consequences of early chilling stress in two Triticum species: plastic responses and adaptive significance

Phenotypic plasticity of two primitive wheat species (Triticum monococcum L. and Triticum dicoccum S.) was studied in response to early chilling stress. Selection pressure differentials, gradients and plasticity costs on plant morphogenesis, growth and reserve carbohydrate consumption were estimated. Regression analysis was applied to investigate differential developmental changes and patterns between treatments. Four‐day‐old seedlings of T. monococcum and T. dicoccum, differing in plant stature and reserve carbohydrates, were given an early chilling temperature (4 °C for 42 day) and compared with control plants grown at 23 °C. Early chilling stress resulted in a significant increase in leaf mass ratio (LMR) and relative growth rate (RGR), a reduction in flag leaf size, total biomass, specific leaf area (SLA) and reserve carbohydrate storage at flowering, together with advanced onset of flowering. Selection pressure within the early chilling environment favoured early flowering, smaller SLA, higher LMR and lower reserve carbohydrates, suggesting the observed responses were adaptive. Furthermore, a regression of daily cumulative plant biomass derived from a crop growth simulation model (CERES‐Wheat) on crop vegetation period revealed a divergent developmental pattern in early‐chilled plants. Using selection pressure gradient analysis, we found similar responses among these traits, except for SLA and sucrose, indicating that these two traits have indirect effects on fitness. Thus, the total effects of SLA and reserve sucrose on relative fitness seem to be buffered via the rapid growth rate in chilled plants. While lower SLA may reduce early chilling stress effects at an individual leaf level, a higher LMR and use of reserve carbohydrates indicated that compensatory growth of chilled plants during the recovery period relied on the concerted action of altered resource allocation and reserve carbohydrate consumption. However, a significant cost of plasticity was evident only for flowering time, LMR and fructan levels in the early chilling environment. Our results demonstrate that morphological and intrinsic developmental (ontogenetic) patterns in two Triticum species respond to early chilling stress.