模拟源株密度对蝴蝶花生长和克隆繁殖的影响

通过不同蝴蝶花(Iris japonica)起始源株密度的控制实验, 探讨源株对克隆植物蝴蝶花克隆繁殖、生长和生物量分配的影响。结果表明: 1)克隆数量特征: 1个起始源株处理(O)蝴蝶花新分株数显著高于2个起始源株(T)及4个起始源株处理(F), 而新分株死亡率显著低于后二者; 随着起始源株数增加(竞争增强), 一级与二级子株数显著降低。随着源株竞争增强, 克隆细根茎与根的长度、表面积、体积与根(茎)长密度逐渐降低。2)叶片特征: 随着源株竞争增强, 母株重度枯萎与总枯萎叶片数显著增加, 子株中度枯萎、重度枯萎与总枯萎叶片数显著增加; 源株竞争增强, 源株叶面积、源株与子株叶片数显著降低, 而源株叶面积比(leaf area ratio, LAR)显著增加。3)生物量及分配: 随着源株竞争增强, 细根茎、粗根茎、克隆繁殖、地上部分、地下部分及总生物量显著降低, 细根茎与地下部分分配降低, 母株粗根茎分配与地上部分分配显著增加。总的看来, 随着蝴蝶花源株竞争增强, 植株叶片生长状况受到更大的影响, 植株生长受限, 克隆繁殖减弱, 而其通过增加LAR, 提高叶片效率与增加母株粗根茎分配, 降低生长强度与消耗, 储备资源, 以待来年的生长与繁殖。     

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。     

内蒙古羊草草原群落主要植物的热值动态

对内蒙古羊草草原主要植物种群的干重热值动态研究表明,热值随植物种类,植物部位,取样时植物所处物候期及气候条件的不同而变化,羊草草原主要植物种群地上部分热值的变动范围在15703－18141J／g之间,其中灌木小叶锦鸡儿（Caragang microphylla)的热值最高,禾草类植物的热值多数较高,而大多数杂类草的热值相对较低,主要植物种群地下部分热值的分布范围为15051－16410J/g。其中根茎型草地下部分热值较高。不同种类植物地下部分热值差异并不与地上部分一致,根茎型禾草地上、地下部分热值差异较小,而须根型植物差异较在,不同种群的植物地上部分热值随植物候期的不同而波动,其变化规律是与植物种群本身的生物学特性相联系的,不同植物种群热值的年际小 规律有所不同,羊草（Leymus chinensis)、大针茅（Stipa grandis)和洽草（Koeloria cristata)的年际热值波动相关显著。但与生长季降水量和生长季累积日照时数之间无明显相关性,在某种程度上,植物热值的 种内变化反映了植物生长状况的差异。     

不同放牧率下羊草和芦苇可溶性碳水化合物和氮素含量的变化

在松嫩草原羊草草地上通过小区围栏放牧,对不同放牧率下羊草和芦苇可溶性碳水化合物和氮素含量的变化进行了分析。结果表明,羊草和芦苇在生长季初期茎基部的可溶性碳水化合物含量最低,分别为7．12％和3．95％,随着季节推移逐渐增加;叶片可溶性碳水化合物含量随季节推移的变化不大;羊草返青比芦苇早,从而造成5月份实验开始时两种禾草茎基部和叶片可溶性碳水化合物的差异;一定程度的放牧(本实验条件下为P4和P5小区)有利于牧草可溶性碳水化合物的提高,促进牧草再生氮素含量在生长季初期最大,随季节推移逐渐降低,与其物候期相一致;适当放牧能够刺激根对土壤中氮素的吸收,使其向地上部分转移,提高牧草的营养价值。     

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。     

CO2浓度升高对沉水克隆植物生长速率及营养元素积累的影响

 研究在不同CO2浓度下水生克隆植物刺苦草（Vallisneria spinulosa）整个生活周期中生长的动态变化及对营养元素积累情况。在不损伤植物体的前提下,采用刺苦草形态学指标组合史估计了植株生物量的动态变化。结果表明：刺苦草鳞茎的萌发不受CO2浓度变化的影响。在高浓度CO2即(1000±50) μmol·mol-1下,刺苦草源株地上部分生长速率在整个生长前期和中期都远远高于低浓度CO2即(400±20) μmol·mol-1,而在后期则出现相反的现象,其中一个原因是因为高浓度CO2下后期光合物质向地下大量转移形成鳞茎引起地上部分生长减慢。但高浓度CO2下克隆株中的初级和次级分株生长速率均高于低浓度CO2。在两种CO2浓度下相同克隆植株构件中的总碳含量没有明显差异;除鳞茎外,根、叶、匍匐茎中的总磷含量随CO2浓度升高显著增加;由于各构件生物量增加有明显差异,导致叶和鳞茎因为生物稀释作用而使其含氮量降低了12%～14%,但根和茎中含量基本保持不变。在高浓度CO2中植株总生物量显著升高,所以总碳、总氮和总磷吸收量均显著大于在低浓度CO2中的吸收量。研究结果揭示,大气CO2浓度升高对沉水克隆植物生长的促进,有利于提高水生克隆植物在群落中的竞争能力;水生植物克隆生长将增加水生生态系统中碳的沉积;水环境中N、P含量将直接影响到水生克隆植物生长。     

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     

冬春季海岸滨麦碳水化合物变化差异性与其环境异质性的关系

以烟台海岸不同生态断带自然生长的滨麦(Leymus mollis(Trin.)Hara)为试验材料,通过在晚秋气温降低和冬季冷冻和春季气温回升过程中滨麦叶片和地下各器官可溶性糖、淀粉和纤维素含量的测定以探讨不同生态断带滨麦在其含量上的差异与环境异质性关系及在滨麦抗盐抗风中作用。结果表明,处在近高潮线(10m)高盐、高水分、强风和低温环境的滨麦根茎和芽粗壮、地上枝叶低矮,春季返青晚;随远离高潮线(50m)土壤盐浓度和海风风速降低,温度增高,滨麦根茎变细,地上枝叶细长,春季返青早,其不同生态断带滨麦形态可塑性与环境异质性相关。随晚秋气温下降不同生态断带滨麦地上部非结构碳水化合物向地下转移,地下繁殖器官根茎和芽成为了非结构性碳水化合物的"库"。在茎和根中可溶性糖向顶芽转移同时淀粉下降,纤维素含量增加。但近高潮线10m处的滨麦,冬季地上非结构性碳水化合物向地下转移较早,芽中储存了较多的非结构碳水化合物,春季返青晚但叶中积累较多的纤维素与其抗冻和抗风相关;而生活在远离高潮线50m处的滨麦晚秋生活期略有加长,冬季转移到地下部的非结构碳水化合物较少,储存量较低,返青较早,其叶片和地下部非结构碳水化合物含量较高与其环境低盐、弱海风及其快速生长相关。因此,不同生态断带滨麦在入冬和春季地上和地下碳水化合物转移和转化上的差异在其适应异质环境、产生形态可塑性、形成多抗逆性中起重要作用。     

羊草种群克隆分株之间光合产物的转移

羊草 (L eymuschinensis)是具有克隆生长习性的多年生根茎植物。其分株系统能存活 7a,其根茎甚至可在地上部分死亡后继续存活。羊草分株可分为 7个年龄 (AC) ,有 :AC0、AC1、AC2、AC3、AC4、AC5和 AC6 ;4个世代 ,即祖代开花分株世代 (G(1) )或祖代营养分株世代 (G1)、母代开花分株世代 (G(2 ) )或母代营养分株世代 (G2 )、当代开花分株世代 (G(3) )或当代营养分株世代 (G3)和子代营养分株世代 (G4 )。揭示了羊草克隆分株之间存在 3种光合产物的转移格局 ,它们是向项性转移、向基性转移和水平性转移。还观察到 ,不同类型分株可调节不同类型分株之间的光合产物转移量。对于向项性转移 ,光合产物主要输出到营养分株 (VR) ,以低转移速率输出到开花分株 (FR)。对于向基性转移 ,光合产物主要输出到 FR,而以低转移速率输出到 VR。对于水平性转移 ,该种转移出现在同一世代的姊妹分株 (SR)之间 ,在从 FR到 VR的转移或在从 VR到 FR的转移 ,转移速率较高 ,但在同种类型分株 (即从 FR到 FR或从 VR到 VR)之间的转移速率较低。幼嫩的 VR比同一世代的 FR能保留更多的光合产物     

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。     

The role of carbohydrate reserves in the growth, resilience, and persistence of cabbage palm seedlings (Sabal palmetto)

Sabal palmetto (Walt.) Lodd. ex Schultes (cabbage palm) is an arborescent palm common in many plant communities throughout Florida, U.S.A., and the Caribbean. Although its seedlings grow very slowly in forest understories, they survive damage and defoliation well, and the species may increase in dominance following disturbances such as fire, logging, and hurricanes. We investigated the potential importance of total nonstructural carbohydrate (TNC) pools in the ability of cabbage palm seedlings to recover from the loss of aboveground tissue such as that caused by fire, grazing, or shallow burial by storm debris. TNC concentrations in belowground organs of seedlings from a forest understory were high, and TNC pools were sufficient to theoretically replace >50% of a seedling's canopy. The largest fraction of the belowground TNC pool was in stem tissue, where TNC in unclipped plants accounted for 26–54% of stem dry mass. Experimental reduction of TNC pools by repeated defoliation slowed seedling regrowth, and seedlings with inherently smaller pools (smaller seedlings) suffered higher mortality after repeated defoliation than did larger seedlings. Although regrowth and recovery after the loss of aboveground tissue was related to the size of the TNC pool in belowground organs, even the smallest seedlings with the smallest pools had sufficient stores to withstand at least two defoliations at frequent (7-week) intervals. Large belowground TNC pools in S. palmetto seedlings appear to enable them to survive all but the most frequent defoliations (e.g., frequent grazing or mowing). Allocation of resources to these stores, however, may contribute to the slow growth rates of S. palmetto seedlings in natural communities. Received: 13 April 1998 / Accepted: 28 August 1998     

Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie

Many clonal plants consist of many connected individual ramets, allowing them to share water and nutrients via physiological integration. Integration among ramets may also improve the ability of clonal plants to tolerate abiotic stress or improve the competitive ability of individual ramets. Here I use a field experiment to determine whether clonal integration improves ramet performance for a widespread clonal tree species invading into native prairie. Aspen (Populus tremuloides) dominates the southern treeline in western Canada, has long-lived belowground connections between mother and daughter ramets, and reproduces vegetatively via resprouting rhizomes after disturbance. I applied two competition treatments (neighbors present or absent) and two clonal integration treatments (belowground rhizomes between mother and daughter ramets either severed or left intact) to 12 replicate Populus daughter ramets at each of three sites. Neighbors improved the survivorship of Populus ramets by 25-35% after 2 yr, but decreased growth by ～20%. Clonal integration tended to improve ramet survival and growth, but these trends were often not significant. Clonal integration did not alter the effects of competition from neighboring vegetation, suggesting that connections between ramets do not necessarily improve the competitive ability of Populus invading into native prairie.     

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.     

Effect of defoliation upon root growth,phosphate absorption and respiration in nutrient-limited tundra graminoids

Summary Moderate experimental defoliation stimulated root respiration and phosphate absorption in two tundra graminoids, Eriophorum vaginatum and Carex aquatilis, growing under nutrient-limited field conditions in northern Alaska. The increase in phosphate absorption rate following defoliation of Eriophorum was associated with a decrease in root phosphate and available carbohydrate contents per unit root length but a constant root nitrogen content. Only after four repeated defoliations did phosphate absorption rate decrease below control levels. We suggest that the stimulation of root respiration and phosphate absorption immediately following defoliation resulted from lowered root phosphorus status as nutrient reserves were reallocated to support shoot regrowth. Root growth was affected more severely by defoliation than was root activity. Two or more defoliations reduced root elongation, initiation and weight per unit length, but root mortality increased only after four defoliations. Carex aquatilis, a species with large belowground biomass, was less sensitive to defoliation than Eriophorum. Phosphate absorption rate increased only after four defoliations in this species, and root elongation, initiation and mortality were affected only by the most severe clipping regimes. Responses of plants to repeated defoliation over two growing seasons were consistent with results of short-term studies.     

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.     

No evidence for nutrient foraging in root-sprouting clonal plants

Clonality is defined as vegetative reproduction via the production of ramets, which are, at least initially, connected by spacers. In general, there are three types of spacers of two origins. Whereas stolons are aboveground spacers, rhizomes are belowground spacers; however, both of stem origin. The third type of spacers are roots in root-sprouting plants. The possibility of foraging in clonal plants has attracted broad interest among ecologists but has been experimentally documented only for stoloniferous clonal plants foraging for light. Foraging for belowground resources has yet to be demonstrated, perhaps because tests of foraging have focused on clonal plants that spread laterally via stolons or rhizomes, i.e. stem organs. Lateral spread based on sprouting roots has not been considered even though, in addition to functioning as conduits between ramets, root spacers are able to sense and take up nutrients. We therefore hypothesized that root-sprouting clonal plants may be able to directly react to environmental heterogeneity and exhibit nutrient foraging. To test this hypothesis, we conducted two experiments with root-sprouters in nutrient-heterogeneous and -homogeneous environments. We found that plants produced more biomass when growing in a heterogeneous environment than in a homogeneous environment and that root biomass was greater in the nutrient-rich patches than in nutrient-poor patches. However, the number of ramets did not differ between patches in the heterogeneous environment. We conclude that plants whose clonality is based on roots, similarly as plants whose clonality is based on stolons or rhizomes, do not exhibit accumulation of ramets in nutrient-rich patches. Foraging at the organ level, i.e. by roots, seems to be more probable in this clonal group. To analyse how clonal plants with different clonal strategies perceive and react to environmental heterogeneity, researchers must account for the high variability in clonal growth forms and in scales of environmental heterogeneity.     

Seasonal allocation of carbohydrates between above‐ and below‐ground organs of Typha domingensis

Seasonal carbohydrates allocation by Typha domingensis was evaluated to identify the potential physiological weaknesses in the growth cycle of this plant in Lake Burullus, Egypt. Monthly plant samples (February–October 2014) were separated into shoots, roots and rhizomes to evaluate the seasonal changes in water‐soluble carbohydrates (WSC), starch and total non‐structural carbohydrates (TNC) for each plant organ. The present study indicated that rhizomes are strong carbohydrates sink during the life cycle of T. domingensis. Starch represented the greatest part of the TNC pool, surpassing the concentration of WSC 1.8–4.3 times. The WSC, starch and TNC concentrations of T. domingensis below‐ground organs (rhizomes and roots) were high at the beginning of the vegetative period (February); they reached their minima in March to support the shoots growth, then were followed by a gradual increase due to the translocation from shoots. The time when T. domingensis is expected to be most susceptible to a management technique is at the point in the seasonal cycle when the stored carbohydrates are at the lowest (in March).     

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 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.