根状茎型植物扁秆荆三棱对土壤养分异质性尺度和对比度的生长响应

土壤养分的空间异质性在自然界普遍存在, 而克隆植物被认为能很好地适应和利用土壤养分异质性。尽管尺度和对比度是异质性的两个重要属性, 但有关土壤养分异质性的尺度和对比度及其交互作用对克隆植物生长和分株分布格局影响的研究仍比较缺乏。在一个温室实验中, 根状茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis) (异名扁秆藨草(Scirpus planiculumis))被种植在由高养分斑块和低养分斑块组成的异质性环境中。实验为两种尺度处理(大斑块和小斑块)和两种对比度处理(高对比度和低对比度)交叉组成的4种处理组合。在每个处理中, 高养分和低养分斑块的总面积相同; 在所有4种处理中, 土壤养分的总量也完全相同。无论在整个克隆(植株)水平, 还是在斑块水平, 尺度、对比度及其交互作用对扁秆荆三棱的生物量、分株数、根状茎长和块茎数的影响均不显著。然而, 在斑块水平, 扁秆荆三棱在高养分斑块中的生物量、分株数、根状茎长和块茎数均显著高于低养分斑块, 而在高养分斑块中相邻分株间的距离(间隔物长)小于低养分斑块, 并且这种效应均不依赖于斑块尺度的大小和对比度的高低。因此, 在土壤养分异质性环境中, 扁秆荆三棱可以通过缩短间隔物长, 并可能通过提高根状茎的分枝强度, 把较多的分株和潜在分株放置在养分条件好的斑块中。这种响应格局体现出克隆植物的觅食行为, 有利于整个克隆对异质性资源的吸收和利用。然而, 该实验中的尺度和对比度对扁秆荆三棱分株的放置格局均没有显著效应。作者推测, 在一个更大的斑块尺度和(或)对比度范围内, 扁秆荆三棱对土壤养分异质性的响应可能不同。因此, 下一步的研究应涉及更广泛的尺度和对比度。     

土壤养分水平影响绢毛匍匐委陵菜匍匐茎生物量投资

为研究匍匐茎草本植物对基质养分供应水平的生物量分配格局的可塑性,在一盆栽实验中对绢毛匍匐委陵菜(Potentilla reptans var. sericophylla)进行了8种不同的养分处理。绢毛匍匐委陵菜植株生物量、匍匐茎数、分株数以及匍匐茎节间长在中等养分条件下最大。随土壤养分的降低,绢毛匍匐委陵菜对叶片和叶柄的生物量投资减小,而对根系的生物量投资增加。在中等养分条件下,绢毛匍匐委陵菜对匍匐茎的生物量投资倾向于最大,而在更高或更低的养分条件下倾向于减少。此生物量分配格局与de Kroon和Schieving的模型模拟结果相符合,结果表明在中等资源水平下增加对匍匐茎的生物量投资是克隆植物增加资源获取的对策之一。     

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。     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

外部干扰对根茎型克隆植物甘草自然种群植株生长及主要药用成分含量的影响

克隆植物, 特别是游击型克隆植物, 其分株在一定时间段往往通过连接结构(即间隔子)连接在一起形成克隆网络, 基于克隆网络的克隆植物特性显著提高了克隆植物应对外部干扰的能力。长期封育禁牧使得甘草(Glycyrrhiza uralensis)自然种群免受外部干扰, 却并没有促使受破坏的甘草自然种群快速恢复。因为克隆整合、克隆储存及克隆分株的选择性放置等克隆植物特性的存在, 适度的外部干扰有利于根茎型克隆植物甘草的生长与繁衍。但外部干扰与克隆植物特性在甘草种群恢复中的作用及其机理目前并不清楚。该研究以甘草自然种群为研究对象, 通过野外控制实验进行模拟动物采食和人工采挖干扰, 探讨不同程度干扰对甘草自然种群生长和主要药用成分含量的影响。结果表明在中度放牧采食干扰或者人工采挖干扰下, 甘草自然种群的分株密度、高度和地上生物量与对照相比均没有显著差异, 甘草表现出等补偿生长; 而在不同类型的重度干扰下, 甘草产生的响应不一致。重度放牧采食干扰下甘草表现出欠补偿生长, 而重度人工采挖干扰下甘草表现出超补偿生长。克隆植物特性在甘草应对外部干扰过程中发挥着重要作用。外部干扰一定程度上提高了甘草中甘草苷和甘草酸的含量, 从而提升了药材的品质。进一步的研究应该关注于甘草自然种群与环境因子之间的关系, 解析甘草自然种群退化的原因, 提出相应的复壮技术, 探讨可持续利用的管理模式。     

切断根茎对根茎禾草沙鞭和赖草克隆生长的影响

为了研究半干旱内陆沙化生境中植物克隆整合对克隆植物基株扩展能力和对克隆植物分株定居逆境能力的贡献,在内蒙古鄂尔多斯高原毛乌素沙地对根茎禾草沙鞭（Rsammochloavillosa（Trin．）Bor）和赖草（Leymussecali－nus（Georgi．）Tzvel．）进行了以切断根茎为处理的野外实验。实验结果表明,切断根茎处理使根茎禾草沙鞭基株的幼小部分（观测单元）在实验期间根茎数量增量、地上技数量增量、地上枝总长度增量、主根茎节数增量、根茎节总数增量、主根茎长度增量和根茎总长度增量显著地减少。然而,切断根茎处理对另一根茎禾草赖草基株幼小部分（观测单元）的上述7个特征在实验期间的增量没有影响。这些结果指示着在克隆整合及其对克隆生长贡献方面的种间差异。沙鞭的克隆整合特性和克隆生长特性对它在水分短缺、蒸散剧烈、养分贫瘠、生境斑块化和扰动经常的沙化草地环境中的生存能力有重要贡献。     

分蘖型克隆植物黍分株和基株对异质养分环境的等级关系

为揭示1年生分蘖草本黍对异质养分环境的等级反应,比较了4种异质养分环境中黍分株和基株的生长,繁殖特征。在分株水平,异质养分环境显著影响源株高度,叶面积,繁殖分配,千粒重和种子数,而对生物量无影响;异质养分环境显著影响分蘖株的这6个指标。在基株水平,生物量,分蘖数和种子数随环境中养分总量的提高而逐渐增大,根系比好相反;繁殖分配不受异质养分环境的影响;除一种处理外,其它3种环境中的千粒重不受养分状况的影响。这些结果表明：黍分株和基株对异质养分环境的反应具有等级性,源株和分蘖株对相同环境的反应存在差异。     

伞形科两种植物幼苗生长对光照强度的可塑性响应

将明党参（Changium smyrnioides Wolff）和峨参（Anthriscus sylvestris Hoffm．）幼苗置于按全光照百分率为100％（S100）、65％（S65）和25％（S25）的人工控制光环境下处理。结果表明,光照强度显著影响明党参和峨参幼苗的生长：明党参在S6,生长最好,株高、冠幅、叶长、叶宽、地上地下生物量等均达到最大值,且S65的明党参生物量是S100的2倍、S25的3倍左右;而峨参地上部生长随着光照强度的减弱而增强,S25生长最好,但地下部生长则在S65生长最好,表现出与明党参同样的响应规律。明党参在不同光照强度下生长速率差异显著,S65显著高于其余两个处理;峨参虽然也在S65生长速率最大,但在不同光照强度间无差异。明党参生长速率明显小于峨参,总生物量约为峨参的1／3—1／8,相对于峨参而言明党参是一个在幼苗阶段生长缓慢的物种。     

杉木种子萌发及幼苗生长对光强的响应

设置不同光强梯度(透光率分别为100%、40%、20%、10%和5%,光照强度PPFD分别为201.3、77.0、37.5、19.2、9.8 μmol·m^-2·s^-1),研究光对杉木种子萌发和幼苗早期生长的影响,分析杉木种子萌发、幼苗存活、生长、形态响应、生物量积累及其分配格局对不同光环境的响应策略.结果表明: 杉木种子的萌发率、存活率、建植率和萌发指数在不同光强梯度下均有显著差异,且40%的透光率是种子最适萌发条件,萌发率最高,而全光照下存活率和建植率最高;随光照强度的减弱,杉木幼苗茎长增大,根长、子叶长、子叶厚、真叶数呈降低趋势,而基径在各光照强度间无显著差异;总生物量、根生物量、茎生物量、叶生物量均表现为全光照下最大.随着光照强度的减弱,光合组织与非光合组织生物量比、叶生物量比呈降低趋势,茎生物量比呈增加趋势,根冠比和根生物量比无显著差异.弱光环境促进杉木种子萌发,不利于杉木幼苗存活和生长.在弱光环境下,杉木幼苗通过增大茎生物量来提高对弱光环境的耐受力.     

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.     

不同种源黄花蒿生长及生物量分配

对都安、崇左、阳朔和融安县四个不同种源黄花蒿的生长发育状况和生物量分配进行比较,结果表明:黄花蒿的生长旺盛期是在5月底至7月初,6月中旬为生长的高峰期,这段时期内应施以重肥;不同种源黄花蒿各功能构件生物量平均值具有相同的规律:茎构件>叶构件>根构件;都安种源的叶构件生物量和青蒿素含量均比其它种源的高,综合表现最优,是引种栽培的好种源。     

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.     

Counteractive biomass allocation responses to drought and damage in the perennial herb Convolvulus demissus

Herbivory and water shortage are key ecological factors affecting plant performance. While plant compensatory responses to herbivory include reallocation of biomass from below‐ground to above‐ground structures, plant responses to reduced soil moisture involve increased biomass allocation to roots and a reduction in the number and size of leaves. In a greenhouse study we evaluated the effects of experimental drought and leaf damage on biomass allocation in Convolvulus demissus (Convolvulaceae), a perennial herb distributed in central Chile, where it experiences summer drought typical of Mediterranean ecosystems and defoliation by leaf beetles and livestock. The number of leaves and internode length were unaffected by the experimental treatments. The rest of plant traits showed interaction of effects. We detected that drought counteracted some plant responses to damage. Thus, only in the control watering environment was it observed that damaged plants produced more stems, even after correcting for main stem length (index of architecture). In the cases of shoot : root ratio, relative shoot biomass and relative root biomass we found that the damage treatment counteracted plant responses to drought. Thus, while undamaged plants under water shortage showed a significant increase in root relative biomass and a significant reduction in both shoot : root ratio and relative shoot biomass, none of these responses to drought was observed in damaged plants. Total plant biomass increased in response to simulated herbivory, apparently due to greater shoot size, and in response to drought, presumably due to greater root size. However, damaged plants under experimental drought had the same total biomass as control plants. Overall, our results showed counteractive biomass allocation responses to drought and damage in C. demissus. Further research must address the fitness consequences under field conditions of the patterns found. This would be of particular importance because both current and expected climatic trends for central Chile indicate increased aridity.     

Adaptive plasticity in response to light and nutrient availability in the clonal plant Duchesnea indica

克隆植物蛇莓对光照强度和养分条件的适应性可塑性 表型可塑性可帮助植物缓冲环境压力并使其表型与当地环境相匹配,但目前仅少数性状的可塑性被广泛认为是适应性的。为充分理解可塑性的适应性意义,仍需进一步研究更多的植物功能性状及其环境因子。本研究将匍匐茎克隆植物蛇莓(Duchesnea indica)的21个基因型种植于不同的光照和养分条件下,并利用选择梯度分析检测了形态和生理可塑性对光照强度和养分有效性变化的适应性值。在遮荫条件下,蛇莓适合度(果实数、分株数和生物量)降低,节间缩短变细,成熟叶叶绿素含量降低,但叶柄长度、比叶面积、老叶叶绿素含量均增加。在低养分条件下,植株叶柄缩短,叶面积缩小变厚,叶绿素含量降低,但果实数量和根冠比增加。选择梯度分析表明,叶柄长度和老叶叶绿素含量对光照变化的可塑性是适应性的,老叶和成熟叶叶绿素含量对养分变化的可塑性也是适应性的。因此,不同性状的可塑性适应值取决于特定的生态背景。该研究的发现有助于理解克隆植物表型可塑性响应环境变化的适应性意义。     

Competitive ability of two Brassica varieties in relation to biomass allocation and morphological plasticity under varying nutrient availability

Green cabbage (Brassica campestris, leafy variety) and turnip (Brassica campestris var. rapifera, rooty variety) were grown in both monocultures and mixtures at three nutrient levels to investigate their responses to nutrient availability with respect to biomass allocation, morphological plasticity and competitive ability. Their allocation parameters and leaf morphological traits were affected by both nutrient availability and developmental stage. Both of the varieties had a smaller biomass allocation to leaf blades, but a greater allocation to petioles at high nutrient levels. Root:shoot ratio (RSR) of green cabbage decreased with increasing nutrient availability, whereas that of turnip increased. Turnip had a smaller leaf blade weight ratio (LBWR) than cabbage, being compensated for by larger leaf area ratio (LAR) and specific leaf area (SLA). Leaf area ratio and SLA of both the varieties increased with increasing nutrient availability as did their mean dry weights. The mean dry weight of turnip was slightly greater than that of green cabbage in their respective monocultures, while that of green cabbage was greater than that of turnip in their 1:1 mixture. Therefore, green cabbage, having inherently greater biomass allocation to leaves, was generally more competitive than turnip with more biomass allocation to roots, especially at higher nutrient levels. However, within a variety, morphological plasticity (variation in LAR and SLA) was more important than the plasticity in biomass allocation (e.g. variation in RSR and LBWR) in determining competitive ability. The implication of our results is that competition models based on biomass allocation pattern alone may fail to predict competitive outcomes and that such models should also take morphological plasticity into full account.     

Plasticity in bud demography of a rhizomatous clonal plantLeymus chinensis L. in response to soil water status

We surveyed the bud demography ofLeymus chinensis L plants along a soil-moisture gradient that was caused by a flood in 1998 on the Song-nen Plain in northeastern China. The number of vegetative buds per ramet was influenced by soil water content, with regression curves being quadratic and the opening of the parabola pointing downward. In addition, the optimum regression models for the numbers of rhizomatous buds and tiller buds relative to soil water resulted in a quadratic parabola and exponential curve, respectively. Vegetative buds flourished between August and October, with plants producing more of those buds on flooded plots than on control sites. The number of rhizomatous buds per ramet was much higher than for tiller buds throughout most of the growing season, and production of the former was more apt to be affected by soil water status. This observed superiority of rhizomatous bud production was thought to be a consequence of the whole-plant adjustment that was stimulated by an abnormally high moisture content. It could also be interpreted as a strategy for “escape” from disadvantageous overly wet conditions. Moreover, the position-based preference for bud emergence along the ramets could be an underlying mechanism for selective ramet placement.     

钩梢强度对麻竹生物量分配和生理特性的影响

以不钩梢为对照,对笋期内麻竹进行轻度、中度、重度钩梢处理,研究各钩梢处理立竹构件生物量分配、当年生枝条和商品叶生长,以及净光合速率(P_n)、蒸腾速率(T_r)、水分利用效率(WUE)和叶片水势(LWP)等的变化,分析麻竹对营林措施中钩梢强度的适应机制和生存策略.结果表明: 与对照相比,钩梢处理对麻竹生物量及分配影响显著,3个处理麻竹立竹的叶/枝、叶/秆、枝/秆生物量比均显著高于对照.轻度钩梢和中度钩梢处理当年生枝条数量、基径、长度均显著大于对照,其中,中度钩梢处理当年生枝条基径、数量、长度和商品叶数量分别比对照增加43.0%、53.3%、29.8%和39.5%.各钩梢处理麻竹P_n、T_r、WUE、LWP在笋期内大小为出笋盛期>末期>初期;钩梢后麻竹的P_n、T_r和WUE显著提高,其中,各处理出笋盛期和末期的P_n大小为中度钩梢>轻度钩梢>重度钩梢>对照,且出笋盛期和末期的P_n分别是对照的1.4和1.3倍.各处理出笋初期、盛期和末期的T_r均为重度钩梢>中度钩梢>轻度钩梢>对照.中度钩梢处理对麻竹立竹的生物量分配、商品叶产量和生理特性均有明显优势,并能有效提高麻竹立竹的水分可获得性.