内蒙古锡林河流域不同生境中羊草的克隆构型和分株种群特征

 克隆植物的形态可塑性在基株和种群水平上分别表现为克隆构型和分株种群特征的变化。研究对象为内蒙古锡林河流域草地、林地、沙地3种生境下的羊草(Leymus chinensis)种群,通过对羊草根茎节间长度、间隔子长度、分枝强度、分枝角度、株高和分株密度等指标的测定和分析,对这3种不同生境中羊草的克隆构型及分株种群特征进行了研究。结果表明羊草克隆构型相关特征,如,根茎节间长度,根茎节间长度频次分布格局、间隔子长度、间隔子长度频次分布格局,在不同生境差异较大。同时,羊草的分枝角度在不同生境间差异显著。而每间隔子所     

吉首蒲儿根的繁育系统及克隆构型

吉首蒲儿根是分布于武陵山区的稀有植物,通过观察其花部特征和开花进程,检测花粉活力、柱头可授性以及套袋实验对其繁育系统进行了研究,并根据地下根茎构筑型、形态特征和地下根茎间的夹角分析了其克隆构型。结果显示:(1)吉首蒲儿根种群花期一般为45~50d,有不明显的二次开花现象,单株花期为10~22d,单个头状花序花期约为3~5d;散粉时柱头高于花药筒,形成花冠、药筒、柱头三者在空间上的分离。(2)吉首蒲儿根的杂交指数为5,P/O值为3 188,繁育类型属于专性异交,异花授粉为主,部分自交亲和,需要传粉者。(3)吉首蒲儿根地上平均含有3.61个分株和4.74个根茎数,根茎间的分枝角度以30°为主,地下根茎构筑型归属于密集型,根状茎断裂是由于组织破碎引起的。研究认为,为适应特殊的峡谷生境,吉首蒲儿根形成了独特的适应策略,先以克隆繁殖来占据生境中的有利地位,后通过"集中开花模式"、雌雄异熟、产生大量花粉等方式来提高异交率,以获得高质量的种子来拓展生境。     

不同生境中淫羊藿克隆构型和分株种群特征

为探求淫羊藿资源的保护策略,通过野外样地调查、室内测定和统计分析等方法,对林缘旷地、林缘和林下三种不同生境中淫羊藿(Epimedium brevicornum)克隆构型及其分株种群特征进行了研究,并初步分析环境因子对淫羊藿克隆构型和分株种群特征的影响.结果表明:分枝强度、间隔子长度、分枝高度、根状茎长度及分株种群密度在3种生境中差异显著,且与生境相对光强和土壤含水量有较强的回归关系.群落相对光强、土壤含水量在淫羊藿的克隆生长过程中起着重要作用.淫羊藿以地下根茎为克隆器官进行克隆生长,有着较强的克隆构型可塑性.结合克隆植物对资源的利用策略,讨论不同生境中淫羊藿克隆构型和分株种群特征可塑性的生态适应意义及淫羊藿保护机制.     

根茎草本披针叶黄华自然分株种群多尺度分布格局

通过在鄂尔多斯高原毛乌素沙地的实地调查 ,应用相邻格子样方和空间自相关分析方法 (Moran′sI) ,研究了在 0 .2m、0 .4m、0 .6m、0 .8m和 1.0m等 5个尺度上根茎豆科草本披针叶黄华 (ThermopsislanceolataR .Br.)自然分株种群的分布格局。同时 ,研究了基株特征和其他分株种群特征。研究显示 ,披针叶黄华自然分株种群具有 4个尺度的非随机分布格局 ;其中 0 .2m尺度的聚集分布格局频率最高。该植物的根茎分枝类型为合轴型(sympodial) ,克隆构型接近于密集型 (phalanx) ,根茎分枝角度为 10°～ 30° ;分株种群密度为 35～ 131株·m-2 ;种群平均分株高度为 11.0～ 2 5 .9cm。在所调查的 2个样方中 ,种群生物量平均分别为 2 6 3.6 3g·m-2 和 30 6 .19g·m-2 ;分株种群的根茎生物量分配分别为 33.71%和 44 .97% ,根生物量分配分别为 2 9.91%和 2 9.95 % ,地上枝叶生物量分配分别为 2 5 .12 %和 36 .35 % ;根冠比均为 0 .44 ,地下 /地上生物量比分别为 2 .12和 3.5 9。     

沙丘坡面异质性小生境中准噶尔无叶豆对水分条件变化的响应

以多年生克隆植物准噶尔无叶豆(Eremosparton songoricum(Litv.)Vass.)为材料,选择河边(A种群)和沙漠腹地(B种群)两个沙丘,研究从沙丘底部至顶部,沿着水分条件连续变化的梯度,准噶尔无叶豆在分株种群和克隆片段水平的形态变化特征,以期能揭示其在异质性小生境内利用水分资源的对策,并为准噶尔无叶豆的资源保护、培育和利用提供有意义的参考.研究发现:①在分株种群水平,A种群分株高度及地上部生物量显著高于B种群,而B种群地下部(根)的生物量则显著高于A种群;②在克隆片段水平,随着沙丘底部至顶部,A种群与B种群克隆片段高度和地上生物量都减小,而分株密度都增加,但升高或降低的强度不同;A种群根的生物量和长度增加,主要是水平的位于地下0～10 cm层面的直径10mm以下的根长度增加,而B种群根的生物量减小,但长度却在增加,主要是水平的位于地下0～10 cm层面的直径6mm以下的细根长度增加.水平细根的长度增加,更利于无性系进行广泛觅食,同时促进无性系尽快越过不利生境斑块和提高分株在有利生境中的生长概率.结果表明,准噶尔无叶豆对沙丘坡面水分条件连续变化的异质性小生境存在分株种群及克隆片段两个等级的可塑性响应,并通过可塑性变化适应了沙丘坡面水分条件的分异.     

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。     

珠芽蓼种群克隆多样性及克隆结构的初步研究

珠芽蓼(Polygonum viviparum)是青藏高原东缘广泛分布的克隆植物,具有有性和无性(根状茎和珠芽)两种生殖方式。该研究采用RAPD技术对分布于不同海拔的珠芽蓼7个自然种群进行了克隆结构和克隆多样性(是单克隆种群还是多克隆种群)以及克隆多样性与海拔因子之间的相关性研究,为了解高山克隆植物对环境的适应性策略及揭示克隆植物的繁殖和分布特点提供科学依据。研究结果表明:1)采用13条RAPD引物对珠芽蓼7个种群共140个样本进行扩增分析,共扩增到117个位点,其中多态性位点84个,多态位点百分率PPL达到71.79%,检测到43个基因型,且全部为局限型基因型;2)与Ellstrand和Roose(1987)总结的克隆植物的克隆多样性平均值相比(PD=0.17,D=0.62),珠芽蓼种群克隆多样性水平稍高,Simpson指数平均为0.639,基因型比率PD平均为0.307;3)克隆结构分析表明,珠芽蓼种群内克隆之间的镶嵌明显,这可能与珠芽蓼过渡型的克隆构型有关。研究中珠芽蓼种群的构型有游击型、密集型以及这两者之间的过渡类型;4)采用SPSS软件对珠芽蓼种群的克隆多样性与海拔高度进行相关性分析,结果显示它们之间并无明显的相关性。     

根茎型木本克隆植物准噶尔无叶豆的种群数量动态

 根据根茎型木本克隆植物的特征, 不以种群的分株数量代表种群大小, 而尝试以不同茎级的根茎长度代表种群大小, 运用种群静态生命表、存活曲线、生殖力表和Leslie矩阵模型, 研究了准噶尔无叶豆(Eremosparton songoricum)的两个种群——A种群(46°31.09′ N, 88°33.06′ E, 紧邻乌伦古湖)和B种群(46°28.07′ N, 88°33.07′ E, 位于沙漠腹地)的种群数量动态。结果表明: 种群存活表现为Deevey-I型。A种群在中龄阶段受到的人为干扰较大, 死亡率出现高峰, 种群的净增长率(R₀)、内禀增长率(r_m)和周限增长率(λ)较低, 表现为衰退型种群, Leslie矩阵模型的模拟结果表明, 15 a内种群呈现下降趋势; B种群所受到的压力主要是干旱贫瘠的荒漠环境所导致的系统压力, 种群的R₀、r_m和λ值适中, 表现为缓慢增长型种群, Leslie矩阵模型的模拟结果表明, 15 a内种群呈现先下降、再上升的趋势。此外, 研究结果验证了Leslie矩阵模型可以扩展应用到根茎型木本克隆植物这类特殊生活型植物的种群数量动态研究上。     

扎龙湿地不同生境芦苇种群根茎构件的年龄结构

芦苇是典型的长根茎型克隆植物,天然种群主要依靠根茎的营养繁殖进行更新。采用单位土体挖掘取样,按实际生活年限划分根茎龄级的方法,对扎龙湿地芦苇种群不同龄级根茎进行调查。结果表明,6月份,4个生境芦苇种群根茎构件均由2—6a的5个龄级组成,7—10月份均由1—6a的6个龄级组成。6—10月份,1a根茎长度比率逐渐增加;除盐碱生境略有增加外,其他3个生境2a、3a比率均小幅减少;4—6a比率逐渐减小,根茎长度比率均以3a最大,依次是2a,4a,1a,5a,最高的6a最小。1—3a根茎生物量比率逐渐增加;4—6a比率逐渐减小,均以3a根茎生物量比率最大,依次是4a,2a,5a,6a,以最低1a最小。不同龄级根茎长度和生物量比率与返青后实际生长时间之间均较好地符合直线函数关系(R~20.91,P0.01;R~20.81,P0.05),6—9月份,根茎长度呈衰退型年龄结构,10月份又转为稳定型结构,整个生长季根茎生物量均为衰退型结构。不同龄级根茎构件在芦苇种群中的地位和作用不同,根茎构件的年龄结构蕴含着种群调节的重要信息。     

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     

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.     

新疆伊犁河谷平原狗牙根无性系构件的生长

采用随机挖掘整个无性系的方法,在新疆伊犁河谷平原2年无人工管理的葡萄园撂荒地,对狗牙根无性系进行了大样本取样,定量分析了构件生长的规律.结果表明：在30个狗牙根无性系中,分株数平均为（272.6±186.6）个,其中营养分株占总分株数的82.3%,是生殖分株的4.3倍;无性系总生物量为(45.4±40.0) g,其中根茎占总生物量的54.4%,营养分株占21.0%,匍匐茎占14.8%,生殖分株占9.4%;根茎与匍匐茎的累计长度分别为(5.1±4.7) m和(3.3±3.4) m、着生的芽数分别为(291.5±246.8)和(78.8±87.4)个.根茎和匍匐茎芽数与无性系的营养分株、生殖分株、根茎和匍匐茎的数量性状之间均呈极显著的正相关关系(P<0.01),表明在新疆伊犁河谷平原,狗牙根无性系主要以根茎芽实现其持续更新.     

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.     

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。     

Life-history monographs of Japanese plants 14: Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae)

The life-history characteristics of Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae) are described. The species is an herbaceous perennial that favors open but relatively moist habitats. It is distributed from Kyushu to Hokkaido in Japan but also occurs in Korea, Mongolia, China and the Russian Far East. In southwestern Japan, shoots start sprouting from mid- to late April, reaching approximately 30–70 cm in height, with 5–10 compound leaves. Ramets simultaneously produce one or more stoloniferous rhizomes that elongate until new ramets are formed at the tips. Cardamine leucantha has a pseudo-annual life cycle, in which mother ramets wither at the end of each season and only daughter ramets appear aboveground in the next year. As a result, ramet positions change annually. In a study population, the number of flowers averaged 23.9 ± 21.0 per ramet and fruit set was 44.2 ± 24.8% (10.4 ± 10.1 fruits per ramet). Ramets produced 3.8 ± 2.3 rhizomes that were 22.0 ± 15.6 cm long. The species sometimes forms large populations. A single genet develops into a group of disconnected ramets spreading via clonal growth. Reproductive characteristics (e.g., fruit set and numbers of flowers and rhizomes) vary among populations, resulting in interpopulation differences in genet structure. Because the reference genome became available recently, established molecular tools will be applied effectively for the investigations of C. leucantha as a model clonal plant.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.Co-ordinating editor: J. Tuomi     

光强对比度对大米草克隆整合作用的影响

通过温室控制试验,分析不同光强及光强对比度处理下克隆植物大米草生长性状的差异,研究同质异质光强条件下克隆整合对大米草响应遮阴能力的修饰作用.结果表明： 在同质条件下,大米草在无遮阴(高光强：温室内自然光照强度)条件下的生物量显著大于中度遮阴(中光强：光照强度为高光强的70%)和深度遮阴(低光强：光照强度为高光强的30%).在低对比度异质性光强条件下(分株对的一个分株不遮阴,另一个分株中度遮阴),大米草遮阴分株的叶片数、根长和生物量均显著高于同质中度遮阴处理,而无遮阴分株各生长指标与同质无遮阴处理相比均无显著差异.因此,在低对比度异质性光强下,大米草受体(遮阴)分株通过克隆整合显著受益;同时,对供体(非遮阴)分株没有显著的耗损.然而,在高对比度处理下(分株对的一个分株不遮阴,另一个分株深度遮阴),克隆整合对受体(遮阴)分株的效应不显著.大米草的克隆整合并不随着光强对比度的增加而增加.在自然生境中度遮阴情况下,克隆整合可以提高大米草的生长和克隆繁殖能力,但在深度遮阴情况下,克隆整合对大米草适应性的作用可能很小.