松嫩平原异质生境羊草种群营养繁殖特征

多年生无性系禾草主要依靠营养繁殖实现种群的持续更新,而各类芽和由芽形成的苗为其种群进行营养繁殖的潜在种群。通过单位面积挖掘取样,利用营养繁殖世代数划分分株、根茎和各类型芽和苗的龄级,对松嫩平原封育草甸、长期割草草甸、封育积沙草甸和林间草地的羊草潜在种群组成和大小进行了比较研究。结果表明: 在生长季末期,羊草的潜在种群均由根茎芽和各龄级分蘖节向上生长的芽以及根茎苗和分蘖节苗组成。在4种生境条件下,羊草分株潜在种群由3～4个龄级组成,1～4龄级根茎均可形成潜在种群,其中,根茎潜在种群占整个羊草潜在种群数量比例为68.3%,占绝对优势。1龄分株和根茎产生的潜在种群占总体的60.2%,是潜在种群组成的主要部分,是种群更新的主要来源。分株潜在种群数量以长期割草草甸最高,根茎潜在种群数量以封育积沙草甸最高。营养繁殖力均以年轻龄级分株和根茎最高,根茎的营养繁殖力是分株的10.0倍,在羊草种群营养繁殖过程中始终占据优势。羊草的潜在种群构成在不同生境条件下变异较大,但体现出对生境差异的趋异适应。     

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     

不同群落中米氏冰草和羊草的年龄结构动态

米氏冰草和羊草都是根茎型禾草,是典型的无性系植物。采取单位面积挖掘取样法,对呼伦贝尔沙地植被次生演替过程中的米氏冰草和羊草种群分株年龄结构进行调查。结果表明,在单生和混合群落中,米氏冰草分株分别由3和4个年龄级组成,羊草均由2个年龄级组成;米氏冰草分株数量和生物量年龄结构变化基本一致,即随着生长年限的延长,呈减少低龄级和增加高龄级比例的趋势,使单生群落中始终为增长型,但接近于稳定型,混生群落中由增长型变为稳定型。羊草分株数量和生物量年龄结构在连续2a均为明显的增长型。羊草各龄级平均单株生产力均高于米氏冰草,平均是米氏冰草的5.2倍。米氏冰草和羊草种群分株年龄结构的变化,蕴含着米氏冰草种群的优势地位将被羊草种群取代的趋势。     

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     

濒危植物朝鲜崖柏克隆种群可塑性及其对土壤条件的响应

朝鲜崖柏为长白山植物区系特有树种,是国家Ⅱ级重点保护的珍稀濒危植物。通过调查长白山区天然分布朝鲜崖柏种群的生境条件和克隆构件的形态学指标,分析了朝鲜崖柏克隆种群特征,并基于构件理论,采用灰色关联及通径分析方法,对种群构件与土壤因子的关系进行研究。结果表明:1)朝鲜崖柏克隆种群具有较强的表型可塑性,分布于暗针叶林下的朝鲜崖柏克隆种群分株间隔子长、地上扩张域、分株间距高于朝鲜崖柏纯林群落,种群构件变异性较高。2)朝鲜崖柏克隆生长格局介于密集型和游击型之间,具有快速克隆扩张能力和明显的斑块状。在土壤贫瘠的纯林群落内,种群更趋近于密集型分布;在土壤条件较好的暗针叶林下,更趋于游击型分布。3)朝鲜崖柏克隆种群生长对土壤异质性的响应具有特殊性,土壤水分是影响朝鲜崖柏种群扩张的最关键因子,土壤p H值、土壤腐殖酸、土壤速效磷、土壤速效钾是影响朝鲜崖柏克隆种群构件较重要的因子。4)土壤养分直接影响克隆植物构件特征及生态适应对策,各克隆分株构件受土壤养分影响的排序为速效磷速效钾有机质全氮。     

松嫩平原水淹恢复演替过程中羊草无性系种群构件的年龄结构

对松嫩平原羊草草甸水淹恢复演替过程中不同大小羊草无性系斑块的种群构件年龄结构进行了研究。结果表明 ,羊草种群分蘖株在斑块中心由 4个龄级组成 ;最外圈层由 3个龄级组成。根茎在斑块中心和中间圈层均由 2～ 4个龄级组成 ,最外层由1～ 2个龄级组成。潜在种群冬性苗和分蘖节芽总量各圈层均由 4个龄级组成。斑块中心的分蘖株种群为稳定型年龄结构 ,向外发展至增长型年龄结构。根茎长度和潜在种群在各圈层均呈增长型年龄结构。在水淹羊草草甸的恢复过程中 ,不同大小羊草无性系斑块均呈不断扩展的趋势。在生存空间充足时 ,羊草的分蘖节在一个生长季里可以繁殖多个世代。通过羊草根茎的年龄结构可预测演替的进程     

羊草种群密度与生长动态研究

 通过人为控制种群密度的栽培实验,从基株和构件水平上观察羊草无性系种群的生长动态,分析不同密度水平上羊草种群通过调节种群的生长、发育、增殖以及死亡过程来适应密度制约的途径。研究结果表明：1)羊草种群密度可在不同的构件水平间得到调节。当基株密度得到适当调节后便可减缓构件密度(无性系枝条、根茎芽等)的变化幅度和影响构件的发育进程。2)无性系枝条是羊草种群的主体,常在生长季后期出现大量增长,都处于生长旺盛的幼苗期,为翌年春季的返青积累能量和物质。3)单位面积总生物量随密度增加而提高,虽然尚未出现产量恒值现象,但存活枝条平均单株重(地上生物量)随密度增加呈下降趋势,这已反映出密度制约的作用。4)不同密度下羊草种群的生殖过程表现为：低密度时以无性繁殖为主,根茎芽的存活力较高;高密度时以有性繁殖为主,生殖分配值较大。     

松嫩草原水淹恢复演替过程中羊草无性系种群构件的物质生产与贮藏

在松嫩平原羊草草甸,对水淹恢复演替过程中半径为0.5、1、3、和10 m的羊草无性系种群斑块分蘖株的物质生产和根茎的物质贮藏进行了研究.结果表明,0.5、1、3、 m斑块各圈和10 m中间及边缘根茎的物质贮藏以1龄级、不同斑块各圈分蘖株的物质生产以1、2龄级对种群的贡献最大,其物质贮藏和生产所占比例均逐圈增加.整体上,年轻龄级分蘖株和根茎具有旺盛的物质生产力和贮藏力.无性系斑块由中心至外围的物质生产和贮藏规律与斑块扩展趋势具有一致性.0.5～10 m斑块分蘖株物质生产力和根茎贮藏力的变异以最外圈最大.在水淹羊草草甸的恢复过程中,不同大小羊草无性系斑块种群构件的物质生产和贮藏策略为种群不断扩展和群落进展演替奠定了物质基础.     

基因型多样性对羊草种群年龄结构的影响

基因型多样性作为遗传多样性的重要组成部分,不仅在群落水平,而且在种群水平均可以发挥重要生态作用。以羊草（Leymus chinensis）为研究对象,设置5种基因型多样性梯度试验控制小区,进行基因型多样性对羊草种群年龄结构的影响研究。结果表明：随基因型多样性梯度的逐步增加,2a分蘖株生物量和数量均显著增加（P<0.05）;1-4龄级羊草根茎生物量、长度和干物质积累量多数呈极显著性增加的变化趋势（P<0.01）。在1、2、4、8和12种基因型多样性梯度时,分蘖株生物量及数量年龄结构均以1a或2a分蘖株所占比例为最高,年龄结构表现为增长型或稳定型;而羊草种群根茎生物量和长度也以2a根茎所占比例为最高,年龄结构表现为稳定型。随基因型多样性水平的增加,羊草种群分蘖株数量和生物量的年龄结构从增长型向稳定型过渡,但根茎长度和生物量的年龄结构均为稳定型。12基因型多样性水平的4龄级分蘖株和根茎的特征高于其他基因型多样性水平。因此,在不同基因型多样性水平羊草种群分蘖株和根茎具有增长型或稳定型的年龄结构,适当的基因型多样性数量显著促进了各龄级羊草分蘖株和根茎的生物量和数量特征的升高,但高度的基因型多样性水平会增加羊草种群中高龄级的分蘖株和根茎所占的比例,种群发展表现出衰退信号。     

淹水和干旱生境下铅对芦苇生长、生物量分配和光合作用的影响

芦苇(Phragmites australis)作为典型的根茎型多年生湿地植物, 具有广泛的环境耐受性。该研究采用盆栽实验, 采取裂区实验设计, 水分处理为主区, 包括淹水和干旱两个水平, 铅(Pb)为副区, 包括0、500、1500、3β000、4β500 mg·kg^-1 5个水平, 共10个处理, 每个处理12个重复, 研究淹水和干旱条件下Pb污染对芦苇生长、生物量分配及光合作用的影响, 以期明确不同生境下芦苇适应或忍耐重金属污染而采取的策略, 为芦苇应用于湿地恢复和污染修复提供理论依据。结果表明, 在淹水处理中, Pb显著抑制地下芽形成和根茎生长, 但对子株数没有影响; 与母株相比子株具有高的日生长速率、光合速率和生物量(母株的3-7倍)。在干旱环境中, Pb显著抑制根、地下芽和根茎生长, 母株和子株生物量积累及光合作用, 且这些指标均小于淹水处理的。无论在淹水还是干旱环境中, 芦苇体内绝大部分Pb积累在根中, 根茎和子株中Pb含量较少, 被转运至母株中的Pb大约是子株的3倍。淹水条件下子株体内Pb含量小于干旱处理的。结果表明, 干旱和Pb的协同作用显著抑制芦苇生长、生物量积累和光合作用, 可能导致子株生产力和种群密度减小甚至种群衰退。但淹水芦苇能够采取相应的Pb分配策略减缓Pb污染对芦苇生长、生理和繁殖的负面影响, 有利于芦苇种群的繁衍和稳定。     

Growth forms of Leymus chinesis (Poaceae) at the different developmental stages of the natural population

The manner in which the density of Leymus chinensis increases from a single plant to a dominant population can be understood by tracing the development of a population from early to late stages. Parent shoot density, above‐ground dry weight, spike density, heading rate and spike dry weight, density of spreading shoots (buds/daughter shoots in apical/axillary rhizomes) and clumping shoots (buds/daughter shoots in axillary parent shoots), and young rhizome length and weight were investigated in the same quadrats for a low density/early stage (LE) population and a high density/late stage (HL) population. Clonal growth (buds/daughter shoots formation) and sexual reproduction (spikes formation) increased while rhizome storage (young rhizome weight) decreased during the transition from LE to HL. In a LE population an outward occupation strategy was employed, with a high proportion of spreading shoots. As the population density gradually increased until HL, an inward consolidation strategy increasing shoot amount in previously occupied areas, was adopted. This was characterized by a high proportion of clumping shoots. Interestingly, the trade‐off between spreading and clumping shoots can be adjusted by the duration of young rhizome elongation during a growth season. In other words, compared with a HL population, a LE population shortened the duration of young rhizome elongation during the growth season, which resulted in more time for the production of axillary spreading shoots along the rhizomes, and high amounts and proportions of total spreading shoots. The special growth patterns, that is, trade‐offs among growth forms, allow L. chinensis to establish dominant populations throughout the eastern Eurasian Steppe.     

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。     

Demography of Carex brevicuspis (Cyperaceae) rhizome populations: a wetland sedge that produces both elongated and shortened rhizomes

The wetland sedge Carex brevicuspis reproduces vegetatively by producing short rhizomes to form clumping ramets phalanx) and long rhizomes to form spreading ramets (guerrilla), resulting in a combined growth form. As an initial step towards understanding the adaptation of Carex growth strategies to seasonal fluctuations in wetland habitats, we investigated the density and composition of C. brevicuspis rhizome populations immediately after flooding (November), in winter (January), in spring (March), and before flooding (May) in the Dongting Lake wetlands, China. The total rhizome density peaked in winter and was lowest before flooding. A large rhizome population in winter may enable C. brevicuspis to survive the seasonal cold weather and recruit a shoot population in the spring. A small rhizome population before flooding may optimize reproductive allocations and be a strategy for enduring the long flooding season. Regardless of date, short rhizomes comprised the majority of the rhizome population (73.0% in March to 98.2% in May). This indicates that C. brevicuspis primarily uses a phalanx growth strategy to utilize locally abundant resources in wetlands. The percentage of long rhizomes in the rhizome population varies significantly between seasons (1.8% in May to 27.0% in March), indicating that growth form also changes with seasonal fluctuation of wetland habitats. The results show that C. brevicuspis may adapt to seasonal changes in wetland habitats through changes in demography of rhizome populations.     

Quantitative analysis of major axis development inViburnum dilatatum andV. wrightii (Caprifoliaceae)

The architectural development ofViburnum dilatatum andV. wrightii was investigated quantitatively. In both species, the major axis is developed from terminal buds of vegetative shoots and from axillary buds on the most distal nodes of reproductive shoots. The architecture of the two species is formed mainly by four branching patterns: a monopodial pattern (M), a sympodial pattern producing a pair of opposite daughter shoots (SP), a sympodial pattern producing a single daughter shoot (SS), and a pattern terminated with a dormant or dead bud (D). In the process of the architecture formation, four successive stages are recognized: 1) height growth, where the M pattern is dominant; 2) crown formation, in addition to the M pattern, the SP pattern occurs frequently; 3) crown expansion, the M and SP patterns are also frequent; 4) over mature, the M pattern becomes dominant again. These four stages are common to the two species, butViburnum wrightii proceeds with the crown formation stage more rapidly and stays in the crown expansion stage for a longer time thanV. dilatatum. The crown ofViburnum wrightii is thus more branched than that ofV. dilatatum.     

A change from phalanx to guerrilla growth form is an effective strategy to acclimate to sedimentation in a wetland sedge species Carex brevicuspis (Cyperaceae)

The rhizomatous sedge Carex brevicuspis can produce clumping ramets from shortened rhizomes (phalanx) and spreading ramets from elongated rhizomes (guerrilla) to form a combined clonal growth form. In this paper, changes in clonal growth and biomass allocation pattern of C. brevicuspis in response to sedimentation were studied. Four sedimentation depths (0, 3, 6, and 9 cm) were applied to 48 ramets in a randomized block design. Plants were harvested after 20 weeks. With increasing sedimentation depth, the proportion of spreading ramets to total ramets increased from 19.6% in 0 cm to 92.9% in 9 cm sedimentation treatments, whereas that of clumping ramets decreased from 80.4% to 7.1%, indicating a change of clonal growth form from phalanx to guerrilla as a response to sedimentation. With increasing sedimentation depth, biomass allocation to shoots and roots did not change, but rhizome mass ratio increased from 2.7% in 0 cm to 7.2% in 9 cm sedimentation treatments, suggesting that production of long rhizomes changes biomass allocation pattern. The results show that plasticity of clonal growth forms, by which more spreading ramets are produced, is an effective strategy to avoid sedimentation stress under our experimental conditions.     

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.     

羊草与大针茅根系构型对水分梯度响应的比较研究

羊草(Leymus chinensis)与大针茅(Stipa grandis)是内蒙古锡林郭勒典型草原的两大建群种,也是内蒙古草原的重要优良牧草。选取锡林郭勒草原以大针茅和羊草为优势种的围封草场为研究样地,通过原状土柱移栽,进行了两年的水分梯度控制实验(150、300、450、600 mm),分别模拟当地年降雨量由干旱年到是湿润年的变化情况,分析比较羊草种群和大针茅种群地上高度和根系构型对水分梯度的响应情况。结果表明:相比大针茅种群,羊草种群对水分梯度的响应更敏感,随着水分梯度的增加,羊草地上高度和根系直径显著增加,根系长度和根系深度显著减少;而大针茅的地上高度和根系特征各项指标均没有显出与水分梯度的相关性,显然大针茅比羊草更能适应干旱生境。在干旱条件(模拟年降雨量150 mm处理)下,大针茅种群地上高度达到峰值,生长状况良好;羊草种群则采用地上个体小型化,地下主根变细,分叉,向土壤深层扩展的生长策略。因此,在气候变化背景下,干旱化的气候将导致大针茅种群在群落中的优势地位逐渐增加,反之羊草种群则会随着气候湿润化而占据更大的优势。     

Nitrogen translocation via rhizome systems in monoclonal stands ofReynoutria japonica in an oligotrophic desert on Mt Fuji: Field experiments

Mechanisms which enableReynoutria japonica, a dominant pioneer herb, to be successful in maintaining large stands in an oligotrophic volcanic desert on Mt Fuji were investigated with special reference to its nitrogen acquisition.Reynoutria japonica forms circular stands, each of which comprises only one genet. As a stand develops outwards, the number of aerial shoots per unit area decreases in the center. Shoots grow vigorously in the peripheral area where the available nitrogen from soil and precipitation (about 2.4 g m⁻² year⁻²) was much less than total nitrogen in the shoots (6.1–9.1 g m⁻²). Leaf nitrogen content per unit mass was also greater in the leaves of the peripheral shoots. When rhizomes extending radially from the center to the periphery were severed, the dry mass of shoots in the periphery diminished by 75% on a ground area basis. In the periphery, leaf nitrogen content also reduced significantly and no flowers were produced. When fertilizer was applied to the peripheral shoots with severed rhizomes, neither growth, survival nor flower production of the shoots was significantly smaller than the control levels. In these shoots, it is also found that the nitrogen content in the youngest leaves decreased for about 1 month and then increased to above that in the control leaves. These results suggest that (i) nitrogen accumulated in the central part is translocated to peripheral shoots via rhizomes, and that long-distance translocation enables the stands to develop outwards, and (ii) aerial shoots in the periphery utilize the nitrogen translocated by rhizomes in the beginning of the growth season, whereas once the shoots have established, they begin to take up nitrogen with their own roots. Since the peripheral shoots are in sunnier environments than the shoots inside the stand, the acropetal nitrogen translocation via rhizomes will raise the production efficiency of a whole stand.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.