不同光照处理下青藏高原克隆植物黄帚橐吾(Ligulariavirgaurea)种子大小对其幼苗生长的影响

以高寒草甸克隆植物黄帚橐吾为实验材料,通过遮荫网模拟植被遮荫,研究种子大小与萌发及幼苗生长能力的关系和幼苗对光照条件的反应。结果表明:(1)在自然光照下,黄帚橐吾种子大小对种子萌发的影响显著,大种子的萌发率高于小种子。遮荫生境下,大、小种子萌发率有所降低,但遮荫对小种子萌发的影响比大种子显著。小种子的萌发率下降了近1/8,而大种子的萌发率仅下降了1/11。(2)黄帚橐吾种子大小对幼苗生物量积累影响显著,大种子幼苗总生物量(TB)大于小种子幼苗的。但生物量的分配与播种时间相关,播种后60d,在自然光照条件下,大种子幼苗对根生物量的分配大于小种子幼苗,而对叶生物量的分配则正好相反。在遮荫环境中,大、小种子幼苗普遍对根的生物量分配增加,大种子幼苗根冠比(R/S)大于小种子幼苗。(3)黄帚橐吾种子大小对幼苗的生长也有明显影响。在自然光照下,小种子幼苗的相对生长速率(RGR)较大于大种子幼苗,但叶面积比率(LAR)、叶面积干质量比(SLA)、叶干质量(LWR)差别不明显。在遮荫条件下,幼苗的LAR、SLA、LWR显著增加,但大、小种子幼苗间差异不显著,幼苗的RGR减小,小种子幼苗的减小趋势大于大种子幼苗。     

高寒草甸不同海拔梯度下多年生黄帚橐吾的克隆生长特征

通过对多年生植物黄帚橐吾(Ligularia virgaurea)在四个不同海拔梯度下的克隆生长行为进行研究,结果表明：(1)资源水平(土壤养分)、干扰和群落性质影响间隔物(spacer)长度的变化。在第一和第四海拔梯度中(土壤养分较丰富)间隔物长度较短,而在第二和第三海拔梯度中(土壤养分较贫乏)间隔物长度较长,说明其能对资源水平和生境优劣作出反应。(2)分枝强度(branching intensity)随资源水平的增加而上升。(3)在高海拔、寒冷和资源较丰富的生境中,其生物量的投资偏向于地下部分生物量,说明黄帚橐吾的资源分配方式受到环境资源条件和群落性质的影响。     

不同养分条件下和不同生境类型中根茎草本黄帚橐吾的克隆生长

 对多年生植物黄帚橐吾(Ligularia virgaurea)在不同氮肥梯度和不同生境类型中的克隆生长行为进行研究。结果表明：1)资源水平与群落性质影响着间隔子(Spacer)长度的变化,氮肥处理使之明显变短;自然群落中的间隔子长度远远长于人工群落,说明黄帚橐吾间隔子具有很强的形态可塑性,能对资源水平和生境优劣作出反应。2)分枝强度(Branching intensity)具有随资源水平的增高而上升的趋势,表明无性系分株(Ramet)的形成和生长受环境资源状况和源株(Genet)生长状况的影响;3)黄帚橐吾的分枝角度为0˚或180˚,资源水平与环境差异对其影响不大,可塑性低;4)环境资源贫乏或植株比较拥挤时,黄帚橐吾个体根系资源分配比例加大,说明黄帚橐吾的资源分配(Resource allocation)模式更多地受环境资源条件和群落性质的影响。     

不同生境下黄帚橐吾（Ligularia Virgaurea）个体大小依赖的繁殖分配

 研究了不同生境和密度下黄帚橐吾(Ligularia virgaurea)个体大小依赖的繁殖分配。结果表明无论在种群水平还是在个体水平上,黄帚橐吾都表现出：1) 在不同的生境和种群密度中,个体大小与繁殖体大小均不相同,其排列顺序依次为沙地＞坡地＞滩地、裸地＞非裸地、低密度＞高密度;2) 繁殖分配和单株种子均重在不同生境和种群密度中,其差异程度均从不显著(p>0.05)到极显著(p<0.01)不等;3) 繁殖体大小与个体大小在不同生境和种群密度中均呈极显著的正相关关系(p<0.01);4) 单株种子均重与个体     

影响黄帚橐吾种子生产的因素I．生境和花序结构

通过对黄帚橐吾(Ligularia virgaurea)自然种群在不同生境和种群密度下的种子生产的研究,结果显示:1)黄帚橐吾的种子生产无论是在种群水平,还是在总花序上同一部位头状花序水平,均受到生境条件的影响,其种子数量、种子均重和种子总重量等特征在不同生境问表现出沙地>滩地>坡地,在同一生境内则为高密度种群>低密度种群和植被盖度较低>植被盖度较高的趋势(滩地例外);2)种子生产中的种子数量和种子总重与总花序大小、头状花序数量以及无性系株大小多呈显著正相关,与个体密度无关,但种子均重与上述因素(除无性系株大小外)的关系不显著;3)在总花序内不同部位头状花序问表现出的种子生产变化趋势(顶部>基部)不受生境条件、种群密度等因素的影响;4)种子生产在总花序上所表现出的这种部位依赖性可能是由于不同部位头状花序间开花时序上的差异引起种子从母体获取资源上的不同所致。     

黄帚橐吾种子生产的花序位置效应及其对幼苗建植的影响

关于开花植物花序内变异原因的研究有很多,而关于花序内资源分配的位置效应产生影响的研究并不常见。通过对青藏高原高寒草甸退化草地一种花序轴基向发育的优势毒杂草黄帚橐吾(Ligularia virgaurea)的野外调查和方差分析发现,其种子生产存在显著的位置效应。结实率和种子平均重表现出顶部头状花序基部头状花序的变化趋势,胚珠平均数则基部顶部,而中部种子平均数最多。通过室内萌发实验和野外盆栽实验研究了位置效应对幼苗建植能力的影响,发现基部种子的萌发开始时间比顶部种子早,萌发率和出苗率在部位间的差异不显著,而部位间幼苗存活率表现一致;幼苗的根冠比和光合速率表现出顶部基部的变化趋势,而幼苗总生物量的差异不显著,这些幼苗特性在两种肥力条件下表现一致,这说明不同花序部位种子的差异是该种植物所固有的特征,且部位间种子的萌发和幼苗生长对策不同。但相对于基部种子,顶部种子扩散距离更远,拓展到新生境的能力更强,相对较高的幼苗根冠比和净光合速率增强了其幼苗建植能力,对自然种群的更新补充有着更重要的生态学意义,对扩大种群和植被更新有更大的贡献。     

影响黄帚橐吾种子生产的因素Ⅰ.生境和花序结构

 通过对黄帚橐吾(Ligularia virgaurea)自然种群在不同生境和种群密度下的种子生产的研究,结果显示：1）黄帚橐吾的种子生产无论是在种群水平,还是在总花序上同一部位头状花序水平,均受到生境条件的影响,其种子数量、种子均重和种子总重量等特征在不同生境间表现出沙地＞滩地＞坡地,在同一生境内则为高密度种群＞低密度种群和植被盖度较低＞植被盖度较高的趋势（滩地例外）;2）种子生产中的种子数量和种子总重与总花序大小、头状花序数量以及无性系株大小多呈显著正相关,与个体密度无关,但种子均重与上述因素（除无性系株大小外）的关系不显著;3）在总花序内不同部位头状花序间表现出的种子生产变化趋势（顶部＞基部）不受生境条件、种群密度等因素的影响; 4）种子生产在总花序上所表现出的这种部位依赖性可能是由于不同部位头状花序间开花时序上的差异引起种子从母体获取资源上的不同所致。     

黄帚橐吾花序结构的资源配置与环境的关系

 为了系统地了解黄帚橐吾(Ligularia virgaurea) 自然种群的繁殖对策,在其繁殖分配研究的基础上,通过统计不同部位头状花序的生物量投入,进一步分析了存在于总状花序内资源分配上的结构效应及其对不同生境条件的反应。结果表明：1）总状花序、不同部位头状花序在资源投入上受其栖息生境各种生态因子的影响而在不同生境和平均种群密度间存在着差异,并表现出一定的趋势;2）可代表个体水平的总状花序大小、头状花序大小、头状花序数量和头状花序平均重量等特征,都与种群密度呈现出程度不同的负相关关系,而总状花序内不同部位的头状花序大小则与种群密度变化无关;3）总状花序内的资源分配存在着位置依赖性,这种显著差异表现为顶部头状花序比基部和中部头状花序有较大的资源分配;4）总状花序顶部、中部和基部头状花序的资源分配与总状花序大小表现出极显著或显著的负相关关系。     

土著入侵种黄帚橐吾的替代防控展望

黄帚橐吾是青藏高原高寒草场中分布最广、危害极大的优势毒草,严重影响了该地区群落结构和草地畜牧业的发展。在介绍土著入侵种和替代防控概念及相关研究进展的基础上,针对黄帚橐吾目前缺乏高效经济生态防控措施的现状,结合国内外对入侵植物进行替代防控的经验与原则,提出了对黄帚橐吾进行替代防控的生态防控设想和黄帚橐吾替代植物选用的基本原则,初步划定了防控黄帚橐吾的替代植物范围。通过引种或者驯化乡土草种,在青藏高原培育禾本科、豆科植物或者建植禾豆混播人工草地,使其具备替代防控黄帚橐吾的潜力。     

黄帚橐吾种群扩张对土壤理化特性与微生物功能多样性的影响

黄帚橐吾(Ligularia virgaurea)是高寒草甸退化的指示物种, 其种群扩张已严重影响了草地生态系统的经济服务功能, 但目前仍不明确土壤微生物是否参与了黄帚橐吾的种群扩张。该研究依托兰州大学高寒草甸试验站, 选择了4个不同密度的黄帚橐吾斑块, 分析了黄帚橐吾种群扩张对该草甸土壤微生物功能多样性的影响。结果显示: 黄帚橐吾种群扩张虽然提高了土壤微生物活性, 但降低了土壤速效氮浓度。各斑块间土壤微生物Shannon指数、碳源利用种类、均一度指数均无显著差异, 但高密度斑块的土壤微生物碳源利用结构与对照斑块有显著差异。表明黄帚橐吾分布地土壤微生物功能多样性的改变所引起的土壤氮素限制是黄帚橐吾种群数量急剧增加的机制之一。     

Seed size effect on seedling growth under different light conditions in the clonal herb Ligularia virgaurea in Qinghai-Tibet Plateau

We studied the influence of seed size on germination, seedling growth and seedling responses to light in Ligularia virgaurea, a clonal herb native to the Qinghai-Tibet Plateau. (1) Under unshaded conditions, large seeds had significantly (P < 0.001) higher rates of germination than did small seeds. Both large and small seeds showed significantly reduced levels of germination under shaded conditions. The magnitude of this effect was greater for small seeds than for large seed. (2) Seedlings from large seeds had significantly higher rates of biomass accumulation (g · day⁻¹) than did seedlings from small seeds. The total biomass of seedlings from larger seeds is larger than that from smaller ones. And seedlings from large and small seeds also differed in biomass allocation. (3) Seedlings from small seeds have higher relative growth rates (RGR; g · g⁻¹ · day⁻¹) than do seedlings from large seeds under both shaded and unshaded conditions. In contrast, there was no significant difference in leaf area ratio (LAR), specific leaf area (SLA) or leaf weight ratio (LWR) between seedlings from small and large seeds. RGR, LAR, SLA and LWR were all significantly higher in seedlings grown under shaded conditions than under unshaded conditions.     

Effects of light on the growth and clonal reproduction of Ligularia virgaurea

Ligularia virgaurea is a perennial herb that is widely distributed in the alpine meadow on the eastern Qinghai-Tibet plateau.We investigated the patterns of growth and reproduction of L.virgaurea under two contrasting levels of light conditions for two continuous growing seasons.Our results showed that the light affects on the maximum relative growth rate,the shoot weight ratio and the root weight ratio differed between the two growing seasons.L.virgaurea reproduced initially through rhizome in the second growing season,rather than sexual reproduction.The proportion of genets with clonal reproduction decreased under shaded conditions.A minimum genet size should be attained for clonal reproduction to begin under the shaded conditions.There was a positive linear relationship between clonal reproduction and genet size.Light level affected the allocation of total biomass to clonal structures,with less allocation under the full natural irradiance than under the shaded conditions.There seemed to be a trade-off between vegetative growth and clonal reproduction under the full natural irradiance,in terms of smaller relative growth rates of genets with clonal reproduction than those without clonal reproduction.L.virgaurea emphasized clonal reproduction under the full natural irradiance,while the plant emphasized vegetative growth under the shaded conditions.     

Intraspecific competition and light effect on reproduction of Ligularia virgaurea,an invasive native alpine grassland clonal herb

The relationship between sexual reproduction and clonal growth in clonal plants often shows up at the ramet level. However, only a few studies focus on the relationship at the genet level, which could finally account for evolution. The sexual reproduction and clonal growth of Ligularia virgaurea, a perennial herb widely distributed in the alpine grasslands of the Qinghai‐Tibetan Plateau of China, were studied under different competition intensities and light conditions at the genet level through a potted experiment. The results showed that: (1) sexual reproduction did not depend on density or light, and increasing clonal growth with decreasing density and increasing light intensity indicated that intraspecific competition and light intensity may affect the clonal life history of L. virgaurea; (2) both sexual reproduction and clonal growth show a positive linear relationship with genet size under different densities and light conditions; (3) a threshold size is required for sexual reproduction and no evidence of a threshold size for clonal growth under different densities and light conditions; (4) light level affected the allocation of total biomass to clonal and sexual structures, with less allocation to clonal structures and more allocation to sexual structures in full sunlight than in shade; (5) light determined the onset of sexual reproduction, and the genets in the shade required a smaller threshold size for sexual reproduction to occur than the plants in full sunlight; and (6) no evidence was found of trade‐offs between clonal growth and sexual reproduction under different densities and light conditions at the genet level, and the positive correlation between two reproductive modes indicated that these are two integrated processes. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Effects of temperature and storage length on seed germination and the effects of light conditions on seedling establishment with respect to seed size in Ligularia virgaurea

Ligularia virgaurea is widely distributed in the alpine meadows of the eastern Qinghai-Tibet plateau. We studied the effects of temperature and seed storage length on seed germination in a laboratory experiment, the effects of seed mass and light intensity on seed emergence in a pot experiment, and the effect of meadow disturbance intensity on seed emergence in a field experiment. Our results showed that seeds of L. virgaurea germinated well under a wide range of temperatures. Germination percentage decreased with increased seed storage length. When seeds were stored for either 6 or 12 months the germination percentage increased with decreasing temperature. The emergence percentage of large seeds was higher than that of small seeds. Seedling survival of large seeds was greater than that of small seeds in 75% and 50% of natural irradiance. Large seeds of L. virgaurea were more successful colonizers in the alpine meadows. Seeds of L. virgaurea germinated in both disturbed meadows and non-disturbed meadows, but seed emergence and seedling establishment were higher in the disturbed meadows, that is, disturbance could facilitate the recruitment and establishment of L. virgaurea .     

Seed size variation and its effect on germination and seedling performance in the clonal herb Convallaria majalis

Variation in seed size is common both within and among plant species. This study examined within-species variation in seed weight, and its implications for some components of fitness in the clonal herb Convallaria majalis. This species produces berries containing 4.1 seeds on average. The average seed weight was 16.5 mg, with a coefficient of variation of 32.7%. Seed packaging in fruits was on average 12.5%, and showed a slight tendency to increase with fruit weight. A trade-off was found between seed weight and seed number both within fruits and within ramets. The probability and timing of germination was not influenced by seed size. A field experiment and indirect evidence suggested that post-dispersal seed predation was not related to seed size. Increasing seed weight conferred an advantage to developing seedlings. This advantage was enhanced if a seedling was growing in the close vicinity of a seedling of another species. It is suggested that seed size variation in C. majalis primarily is the result of resource variation during fruit development. A conflict between parents and offspring may however contribute to increase seed size variation.     

青藏高原特有属——固沙草属(禾本科)2个类群的修订

固沙草属(Orinus)是禾本科(Poaceae)中一个具有重要经济价值的高山特有属,共记载6个物种,它们均具有较强的抗逆特性,是农业良种繁育、畜牧业牧草改良利用的重要资源,但固沙草属物种形态差异不显著,种间界限较模糊。该研究通过标本研究、野外考察、形态性状分析以及实验的观察研究,对固沙草属2个类群进行了分类修订,将长颖固沙草(O.longiglumis)和西藏固沙草(O.tibeticus)以及高秆固沙草(O.alticulmus)和鸡爪草(O.anomala)分别处理为固沙草(O.thoroldii)和青海固沙草(O.kokonorica)的新异名。     

青藏高原特有属——固沙草属表型变异及其对环境因子的响应

该研究通过测量固沙草属(Orinus)3个物种(青海固沙草、固沙草和居间固沙草)的40个天然居群145个个体的22个表型性状,依据VIF膨胀系数筛选得到年平均降雨(MAP)等9个气候因子和海拔等因子,并采用冗余分析(db-RDA)考察不同物种、不同居群植物表型多样性与多元地理气候环境的关系,探讨该属植物表型多样性驱动机理。结果显示:(1)基于22个植物表型性状,固沙草属天然居群在组内最小方差(Ward)55时可归为2类,居群间表型性状不存在显著的空间自相关性(P 0.05),植物表型多样性高(Shannon-Wiener index,H:1.045～2.734),17个定量的表型性状变异系数(CV)平均值47.84%,而居群间小穗颜色多样性最低(H:0.170),证明固沙草属植物表型性状具有独特的环境响应模式。(2)巢式方差分析显示,固沙草属植物表型在居群间和居群内均存在丰富的变异(F 10,P0.01),旗叶、颖片和外稃分别是主成分分析中表型总变异的重要组成部分,表型分化系数居群间变异(71.10%)大于居群内(28.90%),固沙草属植物表型变异来源于居群间。(3)冗余分析结果表明,海拔(ALT)和年平均风速(Wind)是决定青海固沙草(O.kokonoricus)和固沙草(O.thoroldii)居群间表型差异性的主要影响因子;极端气候条件[最冷季平均温度(Bio9)和最干季降雨量(Bio14)]对固沙草居群间表型相似性起促进作用,相反干燥度指数(AI)与水气压(Vapr)对其差异性起促进作用;居间固沙草(O.intermedius)居群间表型差异受年均降雨(MAP)、潜在蒸散(PET)和水气压(Vapr)影响。研究认为,固沙草属植物天然居群之间表型变异反映出明显的地理气候变化趋势,这些表型变异是高山植物环境适应性的重要标志。     

不同营养条件下24种高寒草甸菊科植物种子重量对幼苗生长的影响

以青藏高原高寒草甸中常见的24种菊科植物为材料,在去离子水、河沙加蒸馏水、河沙加半强度霍格兰营养液和河沙加标准霍格兰营养液四种营养基质中培养幼苗,在10d、20d和30d三个幼苗生长期研究种子重量对幼苗生长的影响,结果表明：(1)在10d、20d和30d三个不同的幼苗生长期,在不同培养基质中种子重量与幼苗重量均呈显著的正相关,但R值在去离子水中变化不大,而在其它3种培养基质中持续减少。(2)种子重量与相对生长率负相关,这种负相关性在去离子水中10d和20d生长期表现较弱,在其它3种培养基质中同一生长期的幼苗相对生长率变化不明显。(3)除在10d去离子水中种子重量与幼苗根冠比率呈显著的正相关外,其余均没有显著的相关关系。