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。     

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     

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。     

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。     

典型黄帚橐吾型退化草地形成过程中土壤和植被特征

为深入了解典型毒草型退化草地形成的过程，本文对青藏高原原生草地上黄帚橐吾逐渐扩张形成典型毒草型退化草地过程中植被和土壤特征进行研究。结果表明：黄帚橐吾型退化草地形成过程中，黄帚橐吾种群密度、株高、盖度和生物量均增加；相比原生草地，退化草地的总地上生物量增加113.9%,土壤全氮、铵态氮、有机碳和速效磷含量分别增加61.0%、77.9%、45.3%和78.8%,土壤微生物生物量碳、氮分别增加42.1%和47.4%,而土壤硝态氮含量和物种丰富度分别减少40.1%和28.5%,除黄帚橐吾以外其他植物的生物量减少45.7%。黄帚橐吾极强的种间抑制能力、形态可塑性、高效的养分蓄积能力和利用效率是其成功扩张的关键，促进了黄帚橐吾型退化草地的形成。     

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。     

Effects of the spreading of Ligularia virgaurea on soil physicochemical property and microbial functional diversity

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。     

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60 d,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。     

密度制约下尕海湿地黄帚橐吾叶绿素与叶面积、叶厚度间的关系

密度制约下植物叶性状间的关系,对于认识植物叶片的物理构建与生理代谢的关系具有重要的意义。本研究采用标准化主轴估计分析方法,按黄帚橐吾(Liguaria virgaurea)种群数设置I(171～240株·m~(-2))、II(101～170株·m~(-2))和III(30～100株·m~(-2))3个密度,分析了尕海湿地不同种群密度黄帚橐吾叶绿素含量与叶面积、叶厚度的关系。结果表明:随着黄帚橐吾种群密度的降低,湿地群落的密度呈先减小后增大的U型趋势、盖度逐渐降低、高度和土壤水分逐渐增加,黄帚橐吾的叶面积和叶绿素含量逐渐增加、叶厚度和群落内部光合有效辐射逐渐减小、净光合速率和蒸腾速率呈先减小后增大的U型趋势;叶绿素含量与叶面积呈显著负相关(P0.05)、与叶厚度呈显著正相关(P0.05)。生长在高密度湿地的黄帚橐吾选择增加叶绿素含量且构建小而厚的叶片,在低密度湿地选择减少叶绿素含量且构建大而薄的叶片,反映了植物适应光照条件等异质性环境因子的资源利用策略和光合产物积累模式。     

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。     

Variation in floral sex allocation, female success, and seed predation within racemiform synflorescence in the gynomonoecious Ligularia virgaurea (Asteraceae)

Studies of diclinous species have showed that floral sex allocation and female reproductive success were quite variable within inflorescences. However, little attention has been paid to gynomonoecious species, in which individuals produce both female and bisexual flowers. The purpose of this study was to investigate the variations in reproductive patterns at different capitulum positions within racemiform synflorescence in Ligularia virgaurea, and to determine selective mechanisms of variations in reproductive patterns. We conducted observational and experimental studies in natural populations of the gynomonoecious composite L. virgaurea. Floral sex allocation, seed production and pre-dispersal seed predation were quantified in the field. The results showed several patterns of variation from top to bottom capitula, including an increase in bisexual flowers and flower number per capitulum, but a decrease in seed set and size. Removing earlier capitula during bud stage did not change floral sex allocation in later capitula. And no effect was found on seed set under supplemental pollination. Thus, although it has been reported many times in previous studies, the variation of floral sex allocation in L. virgaurea may not result from architectural effect or mating environment, and the variation of seed production could not be fully explained by pollination success. Additionally, our results showed that L. virgaurea was susceptible to high levels of bisexual biased predation, which was greater for top capitula. We therefore suggest that these variations may help to enhance reproductive success of L. virgaurea in the face of bisexual-biased seed predation.     

入侵植物剑叶金鸡菊的繁殖特征及其与入侵性之间的关系

入侵植物剑叶金鸡菊(Coreopsis lanceolata)既可进行有性繁殖, 又能进行克隆繁殖, 其繁殖特征可能对其入侵性具有重要影响, 然而目前相关研究较少。通过观测剑叶金鸡菊的花序开花动态、交配系统、种子萌发特性等有性繁殖特征及克隆繁殖特征, 研究了上述繁殖特性及其与入侵性之间的关系。结果表明, 剑叶金鸡菊每年4月下旬开始现蕾, 5-11月为花果期。单个花序的花期为5-6天, 至果实成熟开裂约18天。单个花序自交及同株异花授粉不亲和, 为异株异花授粉的异交型, 其花粉-胚珠比(P/O比)为103 549.40 ± 7 162.44。花序高花粉活力及本地蜂传粉行为保障了花粉资源, 因而结实率高达12 000粒∙株^-1。自然条件下其种子的萌发率高达50%以上, 且2-3天即可萌发。剑叶金鸡菊克隆繁殖能力非常强, 整个生长季节均可见从根蘖长出的密集型克隆分株, 8月下旬开始出现游击型克隆生长, 通过密集型和游击型克隆生长, 剑叶金鸡菊排斥其他物种形成单优种群。上述研究结果说明, 剑叶金鸡菊的入侵性与其繁殖特征密切相关。     

Seasonal changes in response ofSenecio ovatus to grazing by the chrysomelid beetleChrysomela speciosissima

Summary The response ofSenecio ovatus to herbivory byChrysomela speciosissima (Coleoptera, Chrysomelidae) was studied in the Krušné hory Mountains, Czechoslovakia. The following questions were addressed: (1) what is the impact of natural levels of insect herbivory on the performance of individual plants, (2) how do the levels of herbivory change during the growing period, and (3) what is the relative importance of direct and indirect effects of herbivory? Ten experimental plots sprayed with metathion and ten controls were established. Damage to plants was quantified in the period of maximum herbivore activity (June) and again at the end of the growing season (August). In the first half of the season, total leaf area in grazed ramets was reduced by 30.3% compared to ungrazed ramets; half of this reduction was due to beetle consumption and half was due to indirect effects. Individual leaf area was not significantly different between treatments. Insecticide-treated ramets were taller and heavier than the controls. During the second half of the growing period the differences in ramet weight and leaf area disappeared because of compensatory growth. However, the insecticide-treated ramets remained taller and had higher seed numbers per capitulum. Neither number of capitula per ramet nor seed weight were significantly different between treatments. In the grazed population, the leaf area reduction (30.3%) in June resulted in 36.5% reduction in fecundity at the end of August.     

Safflower susceptibility and response to feeding by grasshoppers

Feeding by three grasshopper species, Camnula pellucida, Melanoplus packardii and Melanoplus sanguinipes, on three safflower (Carthamus tinctorius) lines for a 6-wk period from anthesis was monitored under field conditions. Ratings of feeding damage to different plant parts (leaves, floral parts, capitula, and peduncles) and measurements after termination of feeding (dry weight, seed yield, seed weight, seeds per capitulum, and capitula per row) were compared among grasshopper species and safflower lines. The Melanoplus species fed preferentially on leaves, floral parts, and capitula, while C. pellucida exhibited only peduncle feeding, which resulted in head clipping. Defoliation of 20 to 30% was associated with significant increases in total dry matter, seed yield, and number of capitula. Further defoliation resulted in decreases. The safflower lines differed in response to grasshopper feeding. S-208 was most susceptible to defoliation by grasshopper feeding, exhibiting decreased dry weight, seed yield, and capitula number. Lesaf 34C-00 was most tolerant and only M. packardii caused significant dry weight and seed yield reductions. Feeding by C. pellucida on this line resulted in an overall seed yield increase. Feeding by M. sanguinipes on Seedtec-5 resulted in yield increases of up to 16%. It appears that certain grasshopper species can increase seed yield in some safflower lines by stimulating the production of additional capitula. Therefore, moderate populations of such grasshoppers in fields of appropriate safflower cultivars do not necessarily require control.     

Influence of achene heteromorphism on life-cycle traits in the annual weed gallant soldier (Galinsoga parviflora Cav.)

Galinsoga parviflora (gallant soldier), Asteraceae, produces two morphologically distinct achenes in a single capitulum: peripheral and central. The morphological, phenological and generative reproduction traits of the progeny derived from peripheral and central achenes that were cultivated in greenhouse conditions were analyzed. Differences between the progeny of various morph types were manifested at different stages of life. The plants of both morph types developed at a similar rate and they entered key phenological phases at the same time except for the flowering stage. The average height of the studied plants was similar on the same days of the experiment. The study showed that plants from peripheral achenes realize the generative reproduction in other pattern than plants from central achenes. The plants from central achenes produce more capitula per plant at the beginning of the fruiting stage, whereas plants from peripheral seeds achieve the highest number of capitula per plant at later stages. Finally, on day 130 of the experiment, the number of capitula per plant of both morphs equalized. At the early fruiting stage, the number of achenes per capitulum of plants from the two morphs did not differ significantly. The number of achenes within one capitulum decreases with plant age in both populations, but the rate of that decline is greater in the progeny of peripheral seeds. The number of peripheral achenes per capitulum in plants from both achene types decreases at a comparable rate. In contrast, the number of central achenes is reduced at a faster rate in plants germinated from peripheral achenes. After day 83, the individuals from peripheral diaspores intensified their reproductive effort; they produced more capitula but with fewer number of achenes than in individuals from central diaspores. On day 130, the number of dispores per plant of the two morphs equalized. The changes of fecundity of peripheral and central progeny with the age of the plant (at different stages of the life cycle of a single plant) contribute to an intensive seed rain throughout the fruiting period. By producing a large number of less fertile capitula with a stable number of peripheral achenes, plants derived from peripheral achenes are able to supply a higher share of peripheral achenes than if they would produce fewer but more fertile capitula. This strategy increases the pool of peripheral achenes.     

淫羊藿种子产量与生境的关系

以四川省南充市金城山的淫羊藿为研究对象,对小檗科淫羊藿产果数量,种子生产和种子质量与生境的关系进行了研究。研究表明:不同生境的淫羊藿植株高度和花序长度不同,单株结果率也不同,单果产种量差异较小,出种量一般在1~8粒之间,但每个生境单果主要出种量不同。对千粒重和不同果枝结果率单因素方差分析表明,不同生境的淫羊藿种子千粒重有显著差异,不同生境的淫羊藿植株的主次果枝结果率也有显著差异。不同生境中的淫羊藿果实的虫食率也不同。淫羊藿种子的低种子数,小粒种子,而不同生境的结实率虽有差异,但是总体偏低,是导致淫羊藿种子难以收集的主要内因,也导致种群受干扰后不易恢复。     

Growth and production characters of elymus cylindricus in different shortening heading time

Ramet modules in a certain population differ in terms of functions, which accounts for different contributions of the same ramets. Shortening heading time brings about different contributions of such modules. Ramets heading one after another were treated as a continuum in respective cohorts of Elymus cylindricus aged two. The reproductive ramets that head earlier were marked with tags every four days during the whole heading stage from the beginning to the end, after which all the labeled ramets at the maturity period were gathered. The results showed that, the height and biomass of ramets, the length and biomass of inflorescences, percentage of inflorescence length to ramet height, percentage of inflorescence biomass to ramet biomass, the number and biomass of seeds, seed-setting rate, and percentage of seed biomass to ramet biomass declined with the increasing intensity of heading time shortening as displayed with linear or quadratic function. Ramet characteristics weakened remarkably when shortened heading time added up to 17 days. The biomass distribution in relation to inflorescence and seed maintain a stable rate at the early heading stage and dwindled quickly at the near-end stage, but the biomass of ramets remain constant throughout the entire heading stage. The ramets with earlier heading time make greater contribution to the survival of population than the shortened heading time in this species of bunchgrass.     

Relationship between capitulum size and pre-dispersal seed predation by insect larvae in common Asteraceae

The evolution of a showy floral display as an advertisement to pollinators could simultaneously advertise the availability of resources to pre-dispersal seed-predators. The hypotheses tested here are that the incidence of seed predation by bud-infesting insect larvae in capitula of Asteraceae is positively related to (1) capitulum size among species, (2) capitulum size within species, (3) capitulum lifespan, and (4) the degree of flowering asynchrony on individual plants. Three populations of each of 20 common herbaceous species of Asteraceae from disturbed ground and grassland habitats were monitored for the presence of pre-dispersal, seed-eating insect larvae. Mean capitulum size (receptacle width) of each species was measured. In a sub-set of eight species, individual capitula were tagged to determine their flowering phenology and lifespan (from anthesis to seed shedding). From these data an index of flowering synchrony on individual plants was derived. Among species, the incidence of larval infestation increased with capitulum size. Small-flowered species such as Achillea millefolium were largely free of bud-infesting larvae, whilst large-flowered species such as Arctium minus were heavily infested. In three cases investigated in greater detail, bud infestation was found to increase with capitulum size within species, suggesting a potential for natural selection to favour smaller capitula. No relationship was found between infestation levels and either capitulum lifespan or degree of flowering synchrony, and there was no evidence that the relationship between capitulum size and infestation was confounded by correlations with these other features. The results support hypotheses 1 and 2, but not 3 and 4. It is suggested that the characteristic capitulum size of each species may represent a trade-off between the opposing selection pressures of pollinators and pre-dispersal seed predators.