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。     

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。     

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。     

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.     

Effects of lead contamination on the clonal propagative ability of Phragmites australis (common reed) grown in wet and dry environments

Clonal propagation is important for the survival and maintenance of the common reed Phragmites australis. Pot culture experiments were conducted to investigate the effects of lead (Pb) concentration (0, 500, 1500, 3000, 4500 mg·kg^?1) and water stress on the clonal reproductive ability of this species. The Pb concentration found in plant organs, in decreasing order, was roots >shoots >rhizomes. There was a negative relationship between the growth of clonal propagative modules (excluding axillary shoot buds) and Pb concentrations, which caused a decrease in biomass, rhizome growth and number of axillary and apical rhizome buds. Daughter axillary shoots exhibited a tolerance strategy, with no significant change in their number; the axillary and apical rhizome buds, daughter apical rhizome shoots and rhizomes exhibited compensatory growth during the late stage of Pb (excluding 4500 mg·kg^?1) treatment in a wet environment. Pb applications above 500 mg·kg^?1 reduced these parameters significantly in the drought treatment, except for the number of axillary shoot buds, which did not change. Our results indicate that clonal propagative resistance to Pb contamination can occur via tolerance strategies, compensatory growth and a Pb allocation strategy, enabling these reeds to maintain population stability in wet environments. However, clonal modular growth and reproductive ability were inhibited significantly by the interaction between drought and Pb, which would cause a decline in P. australis populations in a dry environment. Lead concentrations of 4500 and 500 mg·kg^?1 in soils might meet or exceed the Pb tolerance threshold of clonally propagated reeds in wet and dry environments, respectively.     

Integrated growth patterns of turtle grass,Thalassia testudinum Banks ex König

Indications of physiological integration in the clonal plant Thalassia testudinum Banks ex König from the Puerto Morelos reef lagoon, Mexican Caribbean, were deduced from synchronisation in the formation of ‘inactive shoots’, lateral rhizomes and inflorescences. ‘Inactive shoots’ (i.e. bare, pointed shoots with live roots attached) on a rhizome section often had similar numbers of leaf scars. Lateral rhizome sections were generally found in similar positions when a rhizome had more than one shoot bearing a lateral rhizome. Additionally, the position of lateral rhizomes and the number of leaf scars on inactive shoots were often similar when encountered on the same rhizome section. Synchronisation of flowering events was suggested by the similar position of inflorescence scars on different shoots on the same rhizome section. It is suggested that inactive shoots can play a role in density regulation of the clonal population, and that they possibly constitute a ‘dormant meristem bank’ analogous to ‘seed banks’, or ‘dormant bud banks’.     

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。     

克隆整合对无芒雀麦在异质性盐分环境中存活和生长的影响

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在异质性盐分环境中存活和生长的影响.结果表明,克隆整合显著提高了无芒雀麦分株在高盐环境中的存活能力,耗-益分析表明无芒雀麦在高盐斑块中分株的生物量、分株数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的非盐分斑块中的分株却没有产生显著的损耗.因而,克隆整合特性是无芒雀麦对异质性环境形成的重要适应对策,它使无芒雀麦能够扩展到不适合植物生长的高盐分斑块中,从而增加了无芒雀麦在浑善达克沙地中的存活和生长,提高了其在半干旱沙化地区的适合度.     

慈竹母株大小对克隆生长的影响

 克隆生长在慈竹种群更新中占有重要地位。本文以基径和生物量两个指标同时衡量慈竹(Neosinocalamus affinis)的母株大小,并分析了母株大小对新竹笋产生、竹笋大小和根茎大小的影响。结果表明：1）慈竹的成竹基径与一级母株基径、二级母株基径以及与2龄和3龄母株平均单株生物量呈正相关,但是成竹基径随一级母株基径增加比随二级母株基径增加而增加的速率快得多;2）慈竹母株大小与各样方中出笋数、活笋数和死笋数无显著的正相关;3）发笋母株的基径显著大于未发笋母株,但发一个笋母株与发两个以上笋母株的基径无显著差异;4）母株基径、2龄和3龄母株平均单株生物量都与根茎的长度和粗度呈显著正相关,且竹笋基径与根茎粗度也呈显著的正相关。母株大小对慈竹竹笋的影响是通过影响根茎实现的。总之,慈竹的克隆生长在个体水平受母株大小制约。     

Below-ground spatial pattern of rhizomes in a grassland community and its relevance to above-ground spatial pattern

Studies of spatial patterns in grassland plant communities have focused on above-ground patterns, ignoring the fact that in clonal plant communities, such as those found in grasslands, above-ground spatial structure must reflect below-ground horizontal growth. The present study examines, at both a fine and a coarse spatial scale, relationships between rhizome and ramet distribution. At the coarse scale, the dominance of species differed between above- and below-ground; some species dominated only above- or below-ground, and others dominated in both layers. At the fine scale, a single species' ramet aggregation above-ground significantly differed from its rhizome aggregation below-ground, for many species. Even for a given species, quantitative relationships between above- and below-ground dominance varied among localities. The variation in spatial pattern among species can be explained by attributes of clonal growth form, including rhizome size, rhizome origin and pattern of above-ground ramet aggregation. Using these parameters of clonal growth, four major space occupation patterns were described for mountain grassland species. For species with a high abundance of evenly distributed rhizomes, ramets either i) reflect rhizome distribution, or ii) do not fully reflect rhizome distribution, but rather are spatially aggregated, and rhizomes are typically developed below-ground. For species with a low abundance of rhizomes, ramets either iii) reflect rhizome distribution and rhizomes are typically of above-ground origin, or iv) do not reflect rhizome distribution and are aggregated only at the growing tips of rhizomes. Spatial correlation above-ground among species was the same as below-ground for some pairs of species but was significantly different for other pairs. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phragmites australis at an extreme altitude: Rhizome architecture and its modelling

In central EuropePhragmites australis is a lowland plant, occurring rarely up to the tree line. In the Velká Kotlina cirque (Jeseníky mountains, NE Czech Republic), where it reaches its maximum altitude at about 1350 m a.s.l., its culms are 0.5–0.7 m high and the plants flower only in some years. During the last decade no germinable seeds have been observed. The architecture ofPhragmites rhizomes from this site was studied on seven randomly selected clonal fragments. They consisted of 3 to 10 partial tussocks (clumps) and 4 to 17 green shoots. The total length of the rhizomes was 9.7 to 50 m per plant. The number of nodes per plant was 96 to 431 and the longest internodes were 83 mm long. The number of side branches was 31 to 105 per plant. The branching angle depended on the type of branched rhizome. The mean angles of horizontal rhizomes, which connect individual tussocks, were relatively wide (modus 45°, arithmetic mean 37°), whereas within a tussock much sharper angles of branching prevailed (modal value 5°, arithmetic mean 15°). The mean internode-to-internode angle on continuing rhizomes was about 8°, with a wide variation. An architectural, spatially-explicit model ofPhragmites rhizome growth has been developed, showing that thePhragmites population in the studied locality can be maintained by vegetative multiplication, and seedling recruitment is not needed for its long-term persistence.     

Growth of clonal modules on <i>Agropyron michnoi</i> in the Songnen Plain of Northeast China

Spatial expansion of clonal plants and growth of their modules are of concern in the field of plant ecology. After measuring a large number of samples, we analyzed the module components and the growth patterns of vegetatively propagated Agropyron michnoi clones in the Songnen Plain on Northeast China. The results showed that the plasticity of clonal growth was large; the coefficients of variation of both extensive areas and the quantitative characters of modules were more than 20%. The numbers of ramets, seedlings, and buds and the cumulative length of the rhizomes showed exponentially and linearly increasing patterns with increases of the area and the total number of modules. The biomass of each module, total number of modules and total biomass showed an allometric growth pattern, which was best described by power functions. For A. michnoi, there was a relatively stable investment to sexual reproduction; it showed a priority for allocating biomass to reproductive ramets, and also to rhizomes and buds formation.     

Allozyme Variance and Clonal Diversity in the Rhizomatous Grass Psammochloa villosa(Gramineae)

Psammochloa villosa (Trin.) Bor (Gramineae) is a rhizomatous grass with characteristic sand fixation. Ten enzymes encoded with 15 allozymic loci were used to examine the genetic structures of four natural populations in Inner Mongolia from different habitats. Electrophoretie data indicated that P. viUosa had lower genetic variation (He = 0.104) and lower clonal diversity (D = 0.764) than other similar plants. Genetic compositions of the 4 populations were different. In Shilongmiao mobile dune and Shihuimiao fixed dune there were more genotypes and higher levels of variation than in Shilongndao fixed dune and Shihuimiao semi-mobile dune. Just hke other clonal plants, most genetic variation of P. villosa resided among populations ( Gst = 62,16% ). Most genotypes were local, and there were no spread genotypes. Factors contributed to the genetic structure are herein discussed.     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

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.     

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。     

Effects of resource availability on integration and clonal growth inMaianthemum bifolium

The aim of this study was to investigate whether resource availability affects the degree of physiological integration and the growth pattern of interconnected ramets in the clonal plantMaianthemum bifolium (L.)F.W. Schmidt (Liliaceae), a rhizomatous herb of European forests, by studying it at two contrasting South Swedish beech forest sites termed “poor” and “rich”. The degree of physiological integration was studied by tracing the pattern of¹⁴C translocation and in a cutting experiment involving rhizome severing and defoliation treatments. The size of the plants, growth of new rhizomes, branching frequency, distance between shoots and the internode length were compared. The plants were larger, rhizomes had greater specific mass (mg mm^?1), internodes were shorter and branching frequency higher at the rich site. The cutting treatments reduced the growth of new rhizomes at both sites, and new rhizome segments had lower specific mass in treated plants than in controls, showing the importance of physiological integration for new growth. Translocation of¹⁴C in May showed that the young rhizome tip was a strong sink for carbon. Basipetal translocation to older portions of the rhizome system was greater at the rich site than at the poor site. In September, four months after labelling, the rhizome tips were still strongly labelled with¹⁴C and basipetal translocation had increased at both sites. Plants at the rich site appeared to translocate larger amounts of¹⁴C basipetally than plants at the poor site. It is concluded thatM. bifolium shows a plastic response to resource availability by varying rhizome growth and branching frequency, but the degree of physiological integration is probably only indirectly affected through an increased number of sinks (new rhizome branches) along the ramet system at the rich site.     

Clonal integration increases performance of ramets of the fern Diplopterygium glaucum in an evergreen forest in southeastern China

Physiological integration is a major ecological advantage of clonal growth in angiosperms. Clonal growth is also common in pteridophytes, but almost no study has tested whether clonal integration increases performance in ramets of pteridophytes in natural populations. To test this hypothesis and also whether the positive effect of integration is greater on smaller ramets, we severed the connecting rhizomes of individual ramets of the common, understory fern Diplopterygium glaucum in an evergreen, broadleaf forest in southeastern China. In another experiment, we severed rhizomes around the edges of small plots each containing several ramets. After 19.5 weeks, survival was 100% in intact individual ramets but only 27% in severed ones. Among surviving ramets, final dry mass and lamina mass were also less in severed than in intact ramets, though stalk, rhizome, and root mass and maximum quantum yield of PSII (F_v/F_m) were not reduced. Individual ramets with fewer stalk nodes had lower dry mass but were not more affected by severing than ramets with more stalk nodes. Severance around the edge of plots did not significantly affect the combined final mass of the ramets within a plot. We conclude that clonal integration can have significant positive effects on both survival and growth of individual ramets of ferns in natural populations.