The effect of nutrient availability on biomass allocation patterns in 27 species of herbaceous plants

We investigated allocation to roots, stems and leaves of 27 species of herbaceous clonal plants grown at two nutrient levels. Allocation was analyzed as biomass ratios and also allometrically. As in other studies, the fraction of biomass in stems and, to a lesser extent, in leaves, was usually higher in the high-nutrient treatment than in the low-nutrient treatment, and the fraction of biomass in roots was usually higher under low-nutrient conditions. The relationship between the biomass of plant structures fits the general allometric equation, with an exponent 1 in most of the species. The different biomass ratios under the two nutrient conditions represented points on simple allometric trajectories, indicating that natural selection has resulted in allometric strategies rather than plastic responses to nutrient level. In other words, in most of the species that changed allocation in response to the nutrient treatment, these changes were largely a consequence of plant size. Our data suggest that some allocation patterns that have been interpreted as plastic responses to different resource availabilities may be more parsimoniously explained as allometric strategies.     

Size‐dependent resource allocation and sex allocation in herbaceous perennial plants

Individuals within a population often differ considerably in size or resource status as a result of environmental variation. In these circumstances natural selection would favour organisms not with a single, genetically determined allocation, but with a genetically determined allocation rule specifying allocation in relation to size or environment. Based on a graphical analysis of a simple evolutionarily stable strategy (ESS) model for herbaceous perennial plants, we aim to determine how cosexual plants within a population should simultaneously adjust their reproductive allocation and sex allocation to their size. We find that if female fitness gain is a linear function of resource investment, then a fixed amount of resources should be allocated to male function, and to post‐breeding survival as well, for individuals above a certain size threshold. The ESS resource allocation to male function, female function, and post‐breeding survival positively correlate if both male and female fitness gains are a saturating function of resource investment. Plants smaller than the size threshold are expected to be either nonreproductive or functionally male only.     

不同支持物对攀援植物--葎草雌雄株光合特性及生物量结构的影响

葎草[Humulus scandens(Lour.)Merr.]为草本雌雄异株攀援植物,采用野生种群人为控制的方法,设置分枝找到乔木支持物(高度(3±0.5)m)、灌木支持物(高度(1±0.5)m)和无支持物3种生长方式,测定雌(♀)、雄(♂)株分枝的叶面积参数、光合参数、叶绿素含量、可溶性糖含量及茎、叶、花在分枝、构件、单株水平下生物量分配比等指标,分析支持物对分枝光合特性和生物量结构的影响,探讨雌、雄分枝利用支持物的性别差异及响应支持物的生态适应差异。结果表明:(1)分枝的单叶面积、总叶面积、叶面积比率、比叶面积对支持物响应存在性别差异,♀株叶面积参数均大于♂株;支持物差异主要影响叶绿素a含量,对叶绿素b、叶绿素a/b影响不显著。(2)支持物差异对分枝叶片的光合参数均有显著的影响(P0.05),支持物对光合参数影响顺序为GsPnTrCi,性别差异影响顺序为GsTrPnCi;可溶性糖含量在性别间、支持物间均表现出极显著的差异(P0.01),♂株糖含量显著高于♀株。(3)在分枝水平下,♀株叶、茎及花生物量分配比都未受到支持物不同的影响(P≥0.05),而♂株在叶、茎的生物量分配比方面受影响显著(P0.05);在构件水平下,支持物差异显著影响了♂株分枝间的营养生长构件分配比,显著影响了♀株分枝间的生殖生长构件分配比;在单株水平下,叶分配比仅在支持物间差异显著(P0.05),而茎、花分配比在性别间、支持物间均有显著差异。(4)雌雄分枝的光合特性和生物量结构对不同支持物的响应差异明显,使葎草单株的生理整合性和适应性大幅度提高。研究以雌雄异株攀援植物为材料,从分枝水平分析支持物对雌雄株光合特性及生物量结构的影响,研究结果对探讨雌雄异株攀援植物的生态适应性具一定的价值。     

光照强度对8种红树植物幼苗生长和生物量分配的影响

选取无瓣海桑(Sonneratia apetala Buch-Ham.)、卤蕨(Acrostichum aureum Linn.)、桐花树(Aegiceras corniculatum Blanco.)、老鼠簕(Acanthus ilicifolius Linn.)、木榄(Bruguiera gymnorrhiza Lam.)、秋茄(Kandelia candel Druce.)、银叶树(Heritiera littoralis Ait.)和黄槿(Hibiscus tiliaceus L.)为研究对象,通过遮荫控制实验,研究这8种红树植物幼苗在不同光照强度(自然光强的100%、45%、30%、10%)下的生长和生物量分配特征。结果表明:(1)遮荫对无瓣海桑幼苗相对生长速率、生物量积累、基径的增长和叶片数的增长均有显著的抑制作用;(2)桐花和老鼠簕在遮荫处理下生长状况较好,遮荫对桐花生长的影响不显著,对老鼠簕幼苗的生长则有一定的促进作用;(3)当光照强度降低时,无瓣海桑、秋茄、老鼠簕、桐花树的幼苗都对地上部分投入了更多的生物量,其中秋茄、老鼠簕和桐花树通过提高叶生物量来获取更大的光能捕获能力。研究结果表明,从对光照的生长适应角度来看,桐花、老鼠簕和卤蕨适宜种植在郁闭度较高的林下;秋茄适宜在郁闭度较低的林下种植;真红树无瓣海桑和木榄以及半红树黄槿和银叶树则不适合在林下种植。     

放牧格局和生境资源对矮嵩草（Kobresia humilis）分株生物量分配和补偿性生长的影响

为研究密集型克隆植物对放牧扰动和生境资源变化的生物量分配和补偿生长响应特性,验证克隆植物的觅食模型和3个有关植物个体补偿反应的假说(①放牧优化假说,grazing optimization hypothesis,GOH;②反应连续谱假说,continuum of responses hypothesis,CRH;③增长率模型,growth rate model,GRM),在具有不同放牧利用格局和土壤养分水平的高寒矮嵩草草甸(Ⅰ.畜圈草地:重度放牧、资源丰富;Ⅱ.牧道草地:中度放牧、资源贫乏;Ⅲ.封育草地:不放牧、资源贫乏)中通过设置扣笼/无扣笼样方对其建群种矮嵩草(Kobresia humilis)进行了研究.结果表明不论当年解除家畜放牧与否,春季采摘率越高(畜圈草地),分株生物量向生长的投入越少,向贮藏器官的投入越多.繁殖分配在中度采摘下最高(牧道草地),扣笼内外分株各部分的生物量分配无差异.矮嵩草分株在中度采摘×资源贫乏条件下产生了超补偿响应,在重度采摘×资源丰富条件下为等量补偿,重度采摘导致分株密度显著减少.生物量分配格局与觅食模型的预测不符.补偿生长响应特性证实了GOH和GRM的预测,但与CRH的预测不符.这说明在研究地区放牧扰动格局对克隆植物矮嵩草分株的生物量分配和补偿生长具有重要影响,适度放牧利用更利于引起超补偿,而重度利用可能会对该种群的长期保持产生不利影响.     

喜马拉雅地区藤本植物多样性及其地理格局

喜马拉雅山地是生物地理学研究的热点地区之一。本文对喜马拉雅地区的藤本植物多样性及其与毗邻地区的联系进行了统计分析, 并对该地区与印度河-恒河平原地区藤本植物多样性的地理格局及其成因进行了研究。结果显示: (1)喜马拉雅地区总计有1,083种藤本植物, 分属72科309属; 其中木质藤本725种, 草质藤本358种; 攀援方式主要为缠绕攀援(51.3%)。(2)该区域的藤本植物组成受相邻区域植物区系的显著影响, 其1,083种藤本植物中有74.1% (802种)在东南亚地区有分布, 50.6% (548种)在南亚有分布, 48.9% (530种)在中国西南地区有分布。本区藤本植物缺乏特有性, 仅125种(11.5%)为本区所特有, 没有特有含藤属。(3)藤本植物多样性及其在植物区系中的比例均自东向西逐渐降低; 木质藤本比例和缠绕攀援藤本比例均自东向西略呈上升趋势; 大多数含藤属的藤本多样性由东往西递减, 仅极少数含藤属由东往西逐渐增加, 如野豌豆属(Vicia)和菟丝子属(Cuscuta)。(4)藤本植物多样性在喜马拉雅和印度河-恒河平原地区呈现出自东向西递减的相似格局, 由东往西方向上含藤属递减率分别为8.4属/100 km和6.3属/100 km, 但喜马拉雅地区藤本植物多样性更高。喜马拉雅和印度河-恒河平原地区均有分布的272个含藤属中有196属在中亚及伊朗高原不再有分布, 其中31.1% (61属)在喜马拉雅地区的分布显著更偏西, 仅4.1% (8属)在印度河-恒河平原的分布显著更偏西。综合分析表明, 喜马拉雅地区藤本植物的多样性及其地理格局的特点与其特殊的地理位置、气候条件和生境的梯度变化以及毗邻地区植物区系的多元化有关; 水分条件的东西向梯度变化可能是藤本植物在喜马拉雅和印度河-恒河平原地区形成相似格局的主要原因。     

The rates of shoot and root growth in intact plants of pea mutants in leaf morphology

Isogenic lines of pea (Pisum sativum L.) with the genetically determined changes in leaf morphology, afila (af) and tendril-less (tl), were used to study the relationship between shoot and root growth rates. The time-course of shoot and root growth was followed during the pre-floral period in the intact plants grown under similar conditions. The af mutation produced afila leaves without leaflets, whereas in the case of the tl mutations, tendrils were substituted with leaflets, and acacia-like leaves were developed. Due to the changes in leaf morphology caused by these mutations, pea genotypes differed in leaf area: starting from day 7, the leaf area was lower in the af plants and larger in the tl plants as compared to the wild-type plants. Such divergence was amplified in the course of plant development and reached its maximum immediately before the transition to flowering. Plants of isogenic lines did not notably differ in stem surface areas. In spite of significant difference in total leaf area, the wild type and tl plants did not differ in leaf dry weight. Starting from leaf 9, the af plants lagged behind two leaflet-bearing genotypes (wild type and tl) in leaf dry weight, whereas stem dry weight was similar in the wild type and tl forms and slightly lower in the af plants. Root dry weights were practically similar in the wild type and tl plants until flowering. The reduction of leaf area in the af plants drastically reduced root dry weight. In other words, the latter index was related to the total weight and total area of leaves and stems. The correlation analysis demonstrated an extremely low relationship between leaf and stem area and dry weight and those of roots early in plant development (when plants develop five to seven leaves). Later, immediately before flowering (nine to eleven leaves), root weight was positively related to leaf weight and area; however, stem area and root weight did not correlate. Thus, in three genotypes (wild type, af, and tl), at the end of their vegetative growth phase, leaf and root biomass accumulated in proportion, independently of leaf area expansion.     

Quantifying and understanding reproductive allocation schedules in plants

A plant's reproductive allocation (RA) schedule describes the fraction of surplus energy allocated to reproduction as it increases in size. While theorists use RA schedules as the connection between life history and energy allocation, little is known about RA schedules in real vegetation. Here we review what is known about RA schedules for perennial plants using studies either directly quantifying RA or that collected data from which the shape of an RA schedule can be inferred. We also briefly review theoretical models describing factors by which variation in RA may arise. We identified 34 studies from which aspects of an RA schedule could be inferred. Within those, RA schedules varied considerably across species: some species abruptly shift all resources from growth to reproduction; most others gradually shift resources into reproduction, but under a variety of graded schedules. Available data indicate the maximum fraction of energy allocated to production ranges from 0.1 to 1 and that shorter lived species tend to have higher initial RA and increase their RA more quickly than do longer‐lived species. Overall, our findings indicate, little data exist about RA schedules in perennial plants. Available data suggest a wide range of schedules across species. Collection of more data on RA schedules would enable a tighter integration between observation and a variety of models predicting optimal energy allocation, plant growth rates, and biogeochemical cycles.     

江西马头山自然保护区攀缘植物区系研究

马头山自然保护区位于江西省东部资溪县的马头山林场内,地理坐标为117°08’52”～117°18’00”E, 27°40’43”～27°53’52”N,总面积为21 570 hm2,有丰富的野生攀缘植物资源。通过野外采集及标本的整理鉴 定,初步确认该区共有攀缘植物40科92属222种(包括种下等级)。根据对马头山攀缘植物区系分析的结 果,指出本区攀缘植物区系具有以下特征:(1)种类组成丰富,地理成分复杂;(2)区系中热带性地理成分占主 导地位;(3)区系成分相对古老,特有类群丰富,多型性突出;(4)华东区系特征明显,南北区系过渡。     

Leaf damage induces twining in a climbing plant

Successful climbing by vines not only prevents shading by neighbouring vegetation, but also may place the vines beyond ground herbivores. Here we tested the hypothesis that herbivory might enhance climbing in a vine species, and that such induced climbing should be greater in the shade. We assessed field herbivory in climbing and prostrate ramets of the twining vine Convolvulus arvensis. We evaluated plant climbing after mechanical damage in a glasshouse under both sun and shade conditions, and determined whether control and damaged plants differed in growth rate or photosynthetic capacity. Plants experienced greater herbivory when growing prostrate than when climbing onto companion plants, in both an open habitat and a shaded understorey. Experimental plants increased their twining rate on a stake after suffering leaf damage, in both high- and low-light conditions, and this induced climbing was not coupled to an increase in growth rate. Increased photosynthesis was associated with enhanced twining rate only in the shade. Herbivory may be an ecological factor promoting the evolution of a climbing habit in plants.     

不同光照强度对入侵植物薇甘菊（Mikania micrantha）和飞机草（Chromolaena odorata）生长及生物量分配的影响

外来植物入侵严重威胁着当地生物多样性和生态系统功能,引发巨大经济损失,已成为日益严重的全球性问题。光是热带和亚热带森林生态系统植物生长最重要的限制性资源之一,对光的捕获和利用直接影响植物在自然生态系统中的生存和适合度。本研究以华南地区危害严重的入侵植物薇甘菊(Mikania micrantha)和飞机草(Chromolaena odorata)为研究对象,对比二者生长、生物量分配及叶片特征对光照强度变化的响应,从而验证假设:外来植物入侵力受有效光辐射的限制,光照是影响不同演替阶段林型具有不同可入侵性的重要原因之一。研究结果表明,3%—60%全光照下两种外来植物均可存活,并显示较灵活的生物量分配策略,可见其对光照具有广适性,为其广泛分布提供生态学基础。低光严重抑制了两种外来植物的生物量积累,这可能是导致演替后期季风林入侵现象少的主要原因之一。对比两种植物叶片特征对光照强度变化的响应,高光下薇甘菊SLA降低,LAR、LARMR减小,叶片变小变厚,有利于植株保持强光下的水分平衡;低光下薇甘菊增加SLA和LAR,叶片变薄变大,增大对光的截获面积,以实现对光强减弱的适应。相比薇甘菊,3种光照处理对飞机草的SLA、LAR、LARMR无显著影响。以上结果提示:(1)光照影响所在群落对外来植物入侵的抵抗力;(2)攀援植物薇甘菊较丛生型草本飞机草对变化的光环境在叶片水平上具有更高的形态可塑性,有效寻觅光资源。     

Steady-state nutrition and growth responses of Betula pendula to different relative supply rates of copper

In a series of experiments on the growth and nutrition of copper-limited birch (Betula pendula Roth) plants, growth was controlled by the relative addition rate of copper, R_Cu (d^–1). This was 0·05, 0·10, 0·15 or 0·20 d^–1 with free access to all other nutrients. An additional treatment provided free access to all nutrients. The pH in the nutrient solution was ≈ 4·5 and conductivity was 100 μS cm^–1. At steady-state growth, there was a linear relationship between the relative growth rate, R_G, and R_Cu. The [Cu] of the plants ranged from 2·4 to 2·7 μg g^–1 dry mass (DM) in all treatments with limiting R_Cu and was ≈ 28 μg g^–1 in the free access treatment. The plants showed specific copper deficiency symptoms at limitation. Total non-structural carbohydrate concentrations and the fraction of plant DM partitioned to roots was much less at copper limitation than at free access. The uptake rate of copper per unit root growth rate, dCu/dW_r (μmol g^–1 root DM) was unaffected by the copper supply. Low rates of plant growth at copper limitation were associated with high values of specific leaf area (SLA; 47 m² kg^–1) and leaf area ratio (LAR; 28 m² kg^–1 plant DM) but lower values of net assimilation rate (NAR; 2·5 kg m^–2 leaf DM d^–1) than were found at free access, 28 m² kg^–1 (SLA), 17 m² kg^–1 DM (LAR) and 14 kg m^–2 leaf DM d^–1 (NAR), respectively. It is not obvious from the present data how the growth response can help alleviate copper limitation in the field.     

光照对鄂东南2种落叶阔叶树种幼苗生长、光合特性和生物量分配的影响

通过设置4个光照梯度(25%、12%、6%和3%自然光)模拟鄂东南低山丘陵地区落叶阔叶林林下的光环境,研究了2种耐荫性不同的树种幼苗--麻栎(Quercus acutissima)和化香(Platycarya strobilacea)不同光强下的存活率、光合特性、生长和生物量分配,探讨了低光环境中耐荫性不同的树种幼苗维持自身碳平衡的机制和权衡"存活-生长"选择的生活史策略。结果表明:(1)低光下的2个树种幼苗的生长、光合特性和生物量分配具有显著性差异。(2)各个光照梯度下麻栎幼苗都生长良好,存活率保持在35%以上,而化香幼苗遭遇高的死亡率,80d后3%和6%自然光下的幼苗全部死亡;低光环境中麻栎幼苗比化香幼苗具有更大的表观光量子(AQY)和最大净光合效率(Pmax),更低的光补偿点(LCP)和暗呼吸效率(Rd),即耐荫性较强的麻栎幼苗比耐荫性较弱的化香幼苗具有更高的低光碳同化率和碳捕获能力。(3)2个树种幼苗的成活率与RGR呈负相关关系,各个光照梯度下耐荫性较弱的化香幼苗的相对生长率(RGR)显著高于耐荫性较强的麻栎幼苗,而两个树种幼苗的净同化率(NAR)无明显差异。相对于麻栎幼苗较高的根生物量比(RMR),化香幼苗将更多的生物量分配给叶部,因而具有较高的叶生物量比(LMR)、叶面积比(LAR)和比叶面积(SLA)。不同耐荫性的幼苗生长及生物量分配方式的差异是植物"存活-生长"权衡后的结果,耐荫性弱的化香幼苗具有较高的生长潜力和较弱的自我保护能力,而耐荫性强的麻栎幼苗具有更高的低光碳储量,能够维持更好的低光碳平衡,具有竞争优势。     

Growth,phenology, and biomass allocation of alien Solidago species in central Europe

Plant invasion is a major threat to the integrity of an ecosystem. Exceptionally successful invaders in Europe are the American species of Solidago genus. In this study we examined growth, reproduction, and phenology of Solidago species, of American origin, growing in central Europe (S. altissima, S. canadensis, S. gigantea, S. graminifolia). These taxa were compared with two native species: Solidago virgaurea and Tanacetum vulgare. We observed high differentiation in height, number of shoots, and biomass production between individuals within taxa. Generally, the invasive species produced substantially (two to five times) more biomass than the native ones, being statistically significant in the comparison of alien Solidago graminifolia and S. gigantea versus native S. virgaurea and T. vulgare. The ratio of biomass of reproductive parts to overall biomass varied considerably among years, but generally the lowest one was for Solidago altissima, and the highest for S. graminifolia. It shows a lack of a clear pattern of differentiation between alien and native species in terms of biomass investment in reproduction. We observed a general tendency of allocation of a major part of biomass in rhizomes by phalanx species (S. graminifolia and S. gigantea), while species with guerrilla strategy (S. altissima and S. canadensis) invested more biomass in stems and leaves. However, because of the high variability there was no clear, stable pattern of statistically significant differences between these two groups. The results suggest that S. graminifolia reveals a strong potential of invasion, in spite of its, so far limited, distribution in Europe.     

珍稀濒危植物张氏红山茶花的花发育形态及生理特性研究

以珍稀濒危植物张氏红山茶为材料,对其花发育形态及分化过程中生理指标动态变化进行了研究.采用定株调查的方法,通过观察、统计、解剖等确定张氏红山茶盛花期花芽着生位置并划分5个发育阶段,以分光光度法测定了5个发育阶段可溶性糖、可溶性蛋白和核酸含量等的动态变化.结果表明:盛花期花芽着生位置主要在枝条顶端,占70.0％;花芽和叶...     

北方地区藤本类忍冬叶表皮结构及其生态适应性比较研究

利用光镜和扫描电镜观察了8种在北京地区引种栽培成功的藤本类忍冬的叶表皮形态,观察指标包括气孔器、表皮毛、表皮细胞特征等.结合其在北京的栽培状况,发现叶片表皮形态和解剖结构与生态适应性之间有很强的相关性.自然分布广、适应性强的种在叶表皮形态上表现出气孔密度大、表皮细胞小、被毛或叶革质等特征.金银花、红白忍冬、淡红忍冬是8种忍冬中适应性最强的,具有推广应用价值.本研究为藤本类忍冬在北方地区推广应用提供了理论基础.     

Effects of water depth and substrate type on growth dynamics and biomass allocation of Oryza rufipogon

Oryzarufipogon is known as the ancestor of Asian cultivated rice (Oryzasativa) and the most important germplasm for rice improvement. In this study, we investigated its growth and biomass allocation in response to water depth and substrate type. The results demonstrated that individual plant had an average weight of 18.98 g, 9.36 g, and 10.59 g and produced 77.7 seeds, 35.5 seeds, and 41.6 seeds in deep-, fluctuated-, and shallow water, respectively. Root-to-shoot ratios were 0.37 in deep water, 0.61 in fluctuated water level, and 0.56 in shallow water, reflecting larger investment in roots under water-level fluctuation and shallow water conditions. However, biomass allocation for sexual reproduction was no significant different among the three water levels. In substrates experiment, total biomass and seed production were significantly different between sediment types, and the averages were 10.59 g, 3.56 g, and 0.76 g in biomass and 41.6 seeds, 13.8 seeds, and 0.5 seeds per plant in lake sediment, river sand and clay, respectively. The number of tillers was significantly fewer in sandy and clay substrates than in lake sediments. Individual plant in clay substrates was significantly shorter than those in sandy and in lake sediment substrates. Our study suggests that the plasticity in morphology and biomass allocation of O.rufipogon appears to be an adaptation to the variability in water level and substrate conditions.     

长期模拟增温对岷江冷杉幼苗生长与生物量分配的影响

川西亚高山针叶林是青藏高原东部高寒林区的重要组成部分,也是研究全球变化对森林生态系统影响的重要组成。长期模拟增温对川西亚高山森林树木的生长、物质积累及其分配格局的影响至今鲜有报道。以川西亚高山针叶林优势种岷江冷杉(Abies faxoniana)幼苗为研究对象,采用控制环境生长室模拟增温的方法,研究了模拟增温对岷江冷杉幼苗生长、物质积累及其分配格局的影响。结果表明,模拟增温(2.2±0.2)℃处理65个月后,岷江冷杉幼苗基径、株高、单株叶面积和比叶面积(SLA)均显著增加,比叶重(LMA)显著下降。增温对岷江冷杉幼苗的茎、侧枝、叶和总生物量具有显著的促进作用,对根生物量没有显著影响。岷江冷杉幼苗的叶重比(LMR)下降、枝重比(SMR)增加、根重比(RMR)无显著变化。长期增温能显著促进岷江冷杉幼苗的生长和物质积累,改变生物量分配格局,促使叶片物质向茎转移,降低光合物质投入。