切断根茎对根茎禾草沙鞭的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和赖草,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,地上枝数量增量,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     

川西北高寒沙区切断根茎对赖草和沙生苔草克隆生长的影响

为了研究高原亚寒带沙化生境中切断根茎对克隆植物基株扩展能力和分株定居能力的影响,在川西北若尔盖高原沙化区内,对根茎禾草赖草和沙生苔草进行了以切断根茎为处理的野外实验。结果表明,赖草和沙生苔草基株的幼小部分(观测单元)地上枝总长度增量、主根茎长度增量和根茎总长度增量显著减少,而对根茎数增量、主根茎节增量和根茎节总数增量影响不显著;赖草观测单元地上枝数增量显著减少,而对沙生苔草地上枝数增量无影响;赖草地上枝与根茎的相关性质发生逆转。这表明．在高原亚寒带半湿润沙化生境中．克隆整合效应显著促进基株幼小部分地上枝和根茎的伸长生长,但对新生根茎的产生和根茎节分化没有影响;切断根茎处理导致赖草、沙生苔草生殖生长与营养生长间竞争加剧,同时使赖草地上部分与地下部分间竞争加剧;观测单元在缺少与基株(或上级株系)的克隆整合作用时,赖草受到的影响大于沙生苔草。     

根茎禾草沙鞭的克隆基株及分株种群特征

 在内蒙古沙地站对根茎禾草沙鞭的观测实验发现,沙鞭具有规则的克隆生长、“游击型”克隆构型和相当快的克隆扩展。其地下根茎的寿命至少2年。这些发现指示着该植物种可能具有很强的克隆整合。对内蒙古沙地站和内蒙古草原站的单种沙鞭分株种群的比较和在各站对单种和混交沙鞭分株种群的比较发现,不同地点和在不同群落条件下的沙鞭分株种群在许多重要性状上都存在差异。这些结果暗示着克隆可塑性对沙鞭生态适应性的可能贡献。关于沙鞭克隆整合和克隆可塑性的进一步研究是必要的。     

毛乌素沙地克隆植物对风蚀坑的修复

通过在根茎型克隆植物沙鞭和赖草的种群中设置模拟风蚀坑,研究了根茎禾草沙鞭和赖草对风蚀坑的修复能力及其机理。实验结果表明:沙鞭和赖草都能通过克隆拓展在风蚀坑中产生比自然条件下更多的分株,以更好利用风蚀坑中充足的光照,从而具有很强的风蚀坑修复能力;但同时这些分株的生长也受到风蚀坑中养分条件的制约,生物量、叶片数量和分株高度都不如自然条件下形成的分株;赖草通过间隔子的可塑性反应,将更多的新生分株放置在风蚀坑内,从而具有更强的修复能力。赖草对风蚀坑的修复主要是通过周围根茎扩展进入坑中,然后产生新的分株;而沙鞭不仅可以通过周边根茎进入产生新的分株,同时也可以通过更深层次的根茎来产生新的分株。     

根茎禾草沙鞭的克隆生长在毛乌素沙地斑块动态中的作用

毛乌素沙地沙化景观是由众多不同类型和大小的斑块镶嵌而成的,许多生态学过程影响着这些斑块的动态,根茎禾草沙鞭的克隆生长就是其中之一.通过对有沙鞭生长的3块1hm2风蚀样地的野外调查和染料饲喂实验,研究了沙鞭的克隆生长在毛乌素沙化景观斑块动态中的作用.每块样地均被划分为625个4m×4m的格子,在3块样地的流沙斑块出现的植物中,沙鞭占据了最多的格子数.沙鞭根茎扩展的主要方向是从半固定斑块到流沙斑块,染料饲喂实验测量了沙鞭扩展到风蚀流沙斑块中的分株数、根茎分枝数、根茎长度和地上生物量.将不同斑块土壤含水量和沙鞭根茎在土壤中的分布进行比较发现,在沙鞭根茎分布集中的地下30～50cm处,流沙斑块的土壤含水量显著地高于半固定斑块.这一研究结果表明沙鞭的克隆生长在风蚀流沙斑块的固定和演变中起着非常重要的作用.     

克隆整合对无芒雀麦(Bromus inermis)忍受沙埋能力的影响

无芒雀麦是浑善达克沙地植物群落中占优势的多年生根茎禾草。研究了克隆整合特性对无芒雀麦忍受沙埋能力的影响。结果表明,克隆整合显著提高了远端完全沙埋分株的存活,耗-益分析表明远端沙埋分株的生物量、分株数、叶片数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的近端非沙埋分株却没有产生显著的损耗,并且随着沙埋程度增加时,远端沙埋分株的收益有增大的趋势。因而,克隆整合特性是无芒雀麦对严酷沙埋环境形成的重要适应对策,它能够缓解沙埋对无芒雀麦存活、生长的胁迫,提高其在半干旱沙化地区的适合度。     

克隆整合对无芒雀麦(Bromus inermis)忍受沙埋能力的影响

无芒雀麦是浑善达克沙地植物群落中占优势的多年生根茎禾草。研究了克隆整合特性对无芒雀麦忍受沙埋能力的影响。结果表明,克隆整合显著提高了远端完全沙埋分株的存活,耗-益分析表明远端沙埋分株的生物量、分株数、叶片数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的近端非沙埋分株却没有产生显著的损耗,并且随着沙埋程度增加时,远端沙埋分株的收益有增大的趋势。因而,克隆整合特性是无芒雀麦对严酷沙埋环境形成的重要适应对策,它能够缓解沙埋对无芒雀麦存活、生长的胁迫,提高其在半干旱沙化地区的适合度。     

毛乌素沙地游击型克隆半灌木羊柴对局部沙埋的反应

 不同程度的沙埋是生长在干旱和半干旱区内陆沙丘的植物经常遭遇的事件,沙埋可以改变植物所处的生物和非生物环境条件。已有研究表明不同程度的沙埋对于植物的影响不同 。轻微程度的沙埋可以增加植物高度、促进生物量的积累和新生分株的产生。如果沙埋强度不断增加,对植物的影响由正效应逐渐转变为负效应。即超过一定沙埋阈值后,沙埋会削弱植物的生长,甚至影响植物的存活。干旱和半干旱区内陆沙丘中常常生长着许多克隆植物, 克隆整合常常可以缓解克隆植物分株所遭受的局部环境胁迫。根茎型克隆植物羊柴（Hedysarum laeve）是毛乌素沙地的优势半灌木之一,也是当地重要的固沙植物。为了探讨克隆整合的作用是否可以提高沙埋阈值,并有助于羊柴忍受高强度的沙埋,以其为研究对象开展了野外实验。结果表明：轻微程度的沙埋（例如沙埋深度是原始羊柴分株高的10%～20%）可以加速羊柴分株的高生长,提高叶片生物量、茎生物量以及整个地上部分的生物量。高强度的沙埋（例如沙埋深度是原始羊柴分株高的80%～100%）会削弱羊柴分株的存活和生长。在与不遭受沙埋分株相连的情况下,羊柴分株遭受沙埋的阈值高于没有分株相连的,而且在高强度的沙埋下,前者（有分株相连的遭受沙埋的分株）比后者（没有分株相连的遭受沙埋的分株）在株高增量、茎生物量、叶片生物量以及地上分株生物量上都要显著高。这暗示着克隆整合提高了羊柴遭受沙埋的阈值并有助于羊柴分株忍受高强度的沙埋。     

匍匐茎草本绢毛匍匐委陵菜对局部遮荫的克隆可塑性

采自林窗和林内生境的绢毛匍匐委陵菜 (PotentillareptansL .var.sericophyllaFranch)“分株对”(即由一匍匐茎节间相连着的两个分株 ,其一为“目标分株” ,另一为“相连分株”)在一户外实验中经历了全不遮荫、全部遮荫和局部遮荫处理。该植物的基株生物量、匍匐茎总长度、分株数、匍匐茎比节间重、叶柄长、比叶柄重在遮荫条件下较小。匍匐茎节间长度没有对遮荫处理发生反应。在局部遮荫处理 ,遮荫斑块的分株的叶柄长度由于连着未遮荫斑块中分株而变得更长。这种克隆整合对克隆形态可塑性的修饰作用只在林窗生境来源的实验植物中观察到。其他克隆生长和克隆形态特征的可塑性在不同生境来源的实验植物间没有差异。     

毛乌素沙地根茎灌木羊柴的基株特征和不同生境中的分株种群特征

通过对毛乌素沙地常见的豆科根茎灌木羊柴的基株的分株种群的调查研究,揭示植物的克隆生长的生态适应意义。研究表明：１）根茎灌木羊柴具有合轴型分枝类型和游击型基株构型。多年生根茎在沙基质中形成多层密集网络结构。２）固定沙丘的羊柴分株种群与半固定沙丘比具有较大的密度和较长的根状茎节间,较小的平均株距,但两种生境中的分株高度是相似的。固定沙丘的种群生物量和对根茎与花果生物量的投资也高于半固定沙丘上的种群。本     

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.     

根茎禾草沙鞭的等位酶变异及克隆多样性

沙鞭（Ｐｓａｍｍｏｃｈｌｏａｖｉｌｏｓａ（Ｔｒｉｎ．）Ｂｏｒ）是禾本科的一种沙生旱生植物,靠根茎进行无性生殖。采用１０种酶系统共１５个基因位点对中国科学院植物研究所内蒙古伊克昭盟鄂尔多斯沙地草地生态研究站的４个天然居群进行了遗传结构和克隆多样性分析。结果表明,和同类植物相比,沙鞭的遗传变异水平和克隆多样性均较低。沙鞭４个居群的遗传组成明显不同。石龙庙流动沙丘和石灰庙固定沙丘的基株数目多,遗传变异水平较高,而石龙庙固定沙丘和石灰庙半流动沙丘的基株数目少,只有１株或２株,且多样性水平极低。环境差异在沙鞭的等位酶变异上未得到反映。和其它克隆植物相似,沙鞭的大部分变异存在于居群间（ＧＳＴ＝６２．１６％）,居群内变异所占比例较少。沙鞭９１．６７％的基因型属地方型,无广布基因型。     

松嫩平原两个趋异类型羊草无性系种群特征的比较研究

松嫩平原上羊草(Leymus chinensis(Tzvel.)Tzvel.)有两个趋异类型:灰绿型和黄绿型。两个类型羊草的分蘖节一般均存活2～4年,最多可存活5年;根茎一般存活2～3年,最多可存活4年。两个类型无性系种群的分蘖株均为增长型的年龄结构类型。种群根茎的累积长度,灰绿型为18035cm/m~2,黄绿型为21218cm/m~2,其中,均以1、2龄占绝对比重。两个类型均以1龄分蘖株生产力最大,至3龄分蘖株明显减小;各龄根茎的生物量随着年龄的增加呈直线下降;1龄根茎的营养繁殖力甚强,至8月中旬所形成的芽数均已远远多于地上全部分蘖株数;2龄根茎尚存在较小的营养繁殖潜力,3、4龄根茎均已丧失了营养繁殖力。两个类型羊草无性系种群都是通过根茎芽补充更新。     

Partial mechanical stimulation facilitates the growth of the rhizomatous plant Leymus secalinus: modulation by clonal integration

Background and Aims

Mechanical stimulation (MS) often induces plants to undergo thigmomorphogenesis and to synthesize an array of signalling substances. In clonal plants, connected ramets often share resources and hormones. However, little is known about whether and how clonal integration influences the ability of clonal plants to withstand MS. We hypothesized that the effects of MS may be modulated by clonal integration.

Methods

We conducted an experiment in which ramet pairs of Leymus secalinus were subjected to three treatments: (1) connected ramet pairs under a homogeneous condition [i.e. the proximal (relatively old) and distal (relatively young) ramets were not mechanically stressed]; (2) connected ramet pairs under a heterogeneous condition (i.e. the proximal ramet was mechanically stressed but the distal ramet was not); and (3) disconnected ramet pairs under the same condition as in treatment 2. At the end of the experiment, we harvested all plants and determined their biomass and allocation.

Key Results

Clonal integration had no significant influence on measured traits of distal L. secalinus ramets without MS. However, under MS, plants with distal ramets that were connected to a mother ramet produced more total plant biomass, below-ground biomass, ramets and total rhizome length than those that were not connected. Partial MS exerted local effects on stimulated ramets and remote effects on connected unstimulated ramets. Partial MS increased total biomass, root/shoot ratio, number of ramets and total rhizome length of stimulated proximal ramets, and increased total biomass, root weight ratio, number of ramets and total rhizome length of connected unstimulated ramets due to clonal integration.

Conclusions

These findings suggest that thigmomorphogenesis may protect plants from the stresses caused by high winds or trampling and that thigmomorphogenesis can be strongly modulated by the degree of clonal integration.     

Vegetative propagation and sexual reproduction in the woodland understorey pseudo-annual Circaea lutetiana L.

On the forest floor of deciduous woodlands, relative light intensity gradually declines during the early growing season. The woodland understorey pseudo-annual Circaea lutetiana L. completes its life-cycle at the end of summer. These pseudo-annuals are clonal plants which survive the winter only as seeds and as hibernacles produced by the rhizome apices. In this paper, we asked several questions related to the life-history of C. lutetiana. It was found that shoot formation in early spring did not exhaust the old hibernacle. A trade-off between rhizome + hibernacle number and weight might be expected when plants grow under resource limitation. It was hypothesised that both number and weight of rhizomes and hibernacles will be affected by light availability. Since the effect of resource supply on the size number trade-off will depend on the developmental pattern of the rhizome system, rhizome development was studied as well. Soon after the shoots emerged, 1st order rhizomes were formed in May on the nodes of the old hibernacle. First-order rhizomes branched in June and 2nd order rhizomes (side-branches) were continuously produced throughout the growing season. The phenology and developmental rate of plants growing in different light treatments were plastic. On average 30% of rhizome biomass was formed during the vegetative phase, and rhizome and fruit production were only partly separated in time. The ratio of total rhizome biomass to total fruit biomass was not affected by light. Also flower bud removal did not lead to an increase in rhizome production, which suggests that division of biomass to both reproductive modes is rather rigid. The number of 1st order rhizomes was not affected by the light treatments. Under light limitation, both rhizome number and weight of single rhizomes were reduced. In contrast, fruit number, but not weight of single fruits, was limited.     

Water integration patterns in two rhizomatous dune perennials of different clonal fragment size

In clonal plants, a clonal fragment is a basic unit. There may exist clonal integration that is often regarded adaptive. By definition, a larger clonal fragment comprises more interconnected ramets and/or occupies a larger area of the habitat. When the habitat is so heterogeneous in terms of essential resources that the resources the whole clonal fragment needs can be captured only in a limited number of microhabitats, a large clonal fragment may require extensive and/or intensive clonal integration. Therefore, we hypothesize that in an environment where the resources are distributed in a highly heterogeneous way, a species forming large clonal fragments possesses more extensive and/or intensive clonal integration gaining essential resources than that forming small ones.This hypothesis was tested in a field experiment with two rhizomatous species growing in inland-dune habitats with high heterogeneity of water resource, one (Psammochloa villosa) forming large clonal fragments and the other (Hedysarum laeve) forming small ones. In P. villosa, tracer-(water-soluble acid fuchsin)-labeled water could be transported along the rhizome at a velocity of about 0.93 m per hour and 4 ramets per hour, which was much faster than that in H. laeve. Similarly, the acid fuchsin-labeled water was transported to a longer length in the rhizomes (3.96 m vs. 1.12 m) and to more ramets (14 vs. 3) in P. villosa than in H. laeve. In P. villosa, the acid fuchsin-labeled water reached the ends of the rhizomes and all ramets along the rhizomes were dyed purple. In H. laeve, however, the acid fuchsin-labeled water could not reach the rhizome tips and along the rhizomes only some ramets were labeled. The results suggest that P. villosa is an extensive and intensive integrator, whereas H. laeve is a restrictive and less intensive integrator, supporting the hypothesis. The possible mechanisms and the ecological implications of the findings were discussed.     

Differences in allocation patterns in clonal and sexual offspring in a woodland pseudo-annual

We compared the patterns of allocation to reproduction among seed-derived and clonal offspring of a woodland pseudo-annual. Pseudo-annuals are clonal plants which survive the winter only as seeds and hibernacles produced by the rhizome system. Previous studies indicate that flowering is related to the size of these hibernacles. Since seedlings do not have a hibernacle, we did not expect that these plants would reproduce sexually. Assuming a trade-off between sexual and asexual reproduction, and assuming a linear relationship between vegetative plant weight and weight of all reproductive structures (i.e., rhizomes, hibernacles, inflorescences, and seeds), we expected that seed-derived plants would have a stronger biomass allocation to rhizomes and hibernacles. Since resource supply affects plant size, and thus hibernacle and seed production, we also subjected the plants to different levels of shade. At the start of the experiment seed-derived and clonal offspring hardly differed in total fresh weight. At the final harvest in September seed-derived and clonal offspring did not differ in vegetative plant weight (i.e., leaves, stems, and roots). Only light availability significantly affected these plant structures. As predicted, seed-derived plants did not flower in either of the light treatments. Seed-derived plants allocated more biomass to rhizomes and hibernacles, but this was only significant in the highest-light treatment. This result was due only to an increase in the number of hibernacles. Dry weight of single hibernacles was not affected by plant type. The ecological implications of this allocation pattern are discussed. Received: 2 October 1997 / Accepted: 8 March 1998     

Do rhizome severing and shoot defoliation affect clonal growth of Leymus chinensis at ramet population level?

Leymus chinensis (Trin.) Tzvel. is a perennial species of Gramineae, usually subject to defoliation from grazing and mowing. We examined whether shoot defoliation and rhizome severing affected rhizome and ramet growth, and vegetative bud outgrowth of L. chinensis ramet populations. We also tested the hypothesis that clonal growth of the ramets subject to defoliation would benefit from clonal integration between interconnected ramets besides from possible compensatory growth. To 48 experimental plots, we applied six treatments resulting from interactions between two rhizome connection states (unsevered/severed) and three defoliation regimes (non-defoliated, mildly-defoliated and heavily-defoliated). Defoliation affected rhizome growth and bud outgrowth, but had little effect on shoot growth. Mild and heavy defoliation exerted similar effects on rhizome growth. Only heavy defoliation significantly reduced bud outgrowth while mild defoliation did not. The fact that shoot growth did not change after defoliation and that the bud numbers remained unchanged after mild defoliation suggest that the compensatory response enable the species to tolerate grazing to some extent. Neither rhizome severing nor the interaction of rhizome severing and defoliation had effect on any tested variables. Lack of the effect of rhizome severing falsified the first half of our hypothesis, that is, clonal integration was unimportant in our experiment. The probable reasons were suspected to be the short duration of the experiment and/or the buffer effect of carbohydrate reserves in rhizomes for shoot growth and bud production in time of defoliation.