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。     

青藏高原东部橐吾属三种植物的繁育系统

研究青藏高原东缘分布的箭叶橐吾、黄帚橐吾和掌叶橐吾的繁育系统,箭叶橐吾和黄帚橐吾各自居群中有性生殖和克隆繁殖的权衡关系,分析三种植物的的克隆繁殖与生态适应,探讨橐吾属植物对高寒环境胁迫条件下的适应与进化。结果显示三种橐吾的花果期在6月中旬到10月初,花序为头状花序组成的总状排列。三种橐吾的杂交指数(OCI)均为4,繁育系统以远交为主、部分自交亲和、虫媒、需要传粉者;影响有性生殖成功的因素除昆虫侵食以外,黄帚橐吾和箭叶橐吾低的结实率主要原因是传粉媒介限制。限制结实的主要因素是高寒多风的环境限制了昆虫的活动,访花昆虫数量减少,从而使花粉转移效率降低,导致了低的结实率。植物采取的适应对策:大量开花,延长雌雄配子体的功能期和传粉媒介的泛化等,可见三种橐吾有性生殖的潜力依然很大。箭叶橐吾和黄帚橐吾在自然条件下主要进行有性生殖,在保护草场时,应针对其有性生殖器官采取措施。     

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。     

黄帚橐吾对高寒草甸凋落物分解及氮释放的影响

受过度放牧等干扰因素的影响,黄帚橐吾(Ligularia virgaurea)在高寒草甸逐渐扩张成为优势种而被称为“退化指示种”,从而威胁生态系统功能和养分循环。然而,黄帚橐吾扩张引起的凋落物比例增多,对分解及养分释放的影响尚不清晰。凋落物分解连结着生态系统碳、氮循环,对于系统功能维持具有重要意义。本试验在三江源区黄帚橐吾为优势种的高寒草甸中,设置黄帚橐吾和其他物种凋落物的多梯度混合分解试验,测定黄帚橐吾和其他物种凋落物的重量损失和氮释放,分析黄帚橐吾生物量增多对凋落物分解的影响。结果表明：(1)分解1年后,黄帚橐吾重量损失及氮素释放较其他物种更缓慢;(2)随着黄帚橐吾生物量比例的增加,凋落物整体重量损失及氮素释放显著降低;(3)混合凋落物中黄帚橐吾能限制其他物种的分解,这种拮抗效应进一步延缓了混合凋落物的分解。综上,高寒草甸黄帚橐吾扩张引起的凋落物生物量增多通过延缓凋落物分解及氮释放,阻碍土壤养分输入,进而对草地碳氮循环过程产生不利影响。     

高寒草场优势杂草黄帚橐吾水浸液对牧草的化感作用

研究了高寒草场中主要毒杂草——黄帚橐吾水浸液对同域分布的5种牧草的化感作用.将反应指数(RI)相加平均后分为一、二、三级敏感指数,分别评价不同测定指标、不同发育期和物种水平对化感作用的敏感性;从供体角度对相应RI值做同样处理,以综合评价不同水浸液及其不同浓度的化感效应.结果表明,5种牧草对黄帚橐吾化感作用的敏感性从强到弱依次为早熟禾>大雀麦>中华羊茅>羊茅>垂穗披碱草;牧草种子萌发期的敏感性强于幼苗生长期,尤以萌发指数受化感作用的影响明显.黄帚橐吾根、叶水浸液对牧草生物测定结果大多达到显著差异水平,物种水平的化感效应均表现为抑制作用,说明雨水淋溶是黄帚橐吾向环境中释放化感物质的途径之一.黄帚橐吾根部化感效应强于叶片,可能与其地下种间激烈的资源竞争有关.化感作用在增强黄帚橐吾生存竞争力、扩大种群和入侵草场中起着不容忽视的作用,可能是黄帚橐吾单优势种群落形成和草场退化的原因之一.     

青藏高原东缘同域分布的2种橐吾属杂草的合子前生殖隔离

生殖隔离是阻止种间杂交、保持物种完整性和独立性的关键,对生物多样性的产生和维持有重要意义.不同物种间的生殖隔离形成阶段、方式和强度不完全相同,因而对物种间隔离机制的研究有助于揭示物种形成和维持机制.对青藏高原东缘同域分布的2种橐吾属杂草黄帚橐吾和箭叶橐吾的花期物候观察、访花昆虫观察及花粉-柱头亲和性进行研究,分析了2种橐吾多种合子前生殖隔离机制的特点及其在生殖隔离中的作用.结果表明: 黄帚橐吾种群花期比箭叶橐吾早20 d左右,虽然盛花期存在差异,但是2种群花期存在10 d左右的重叠期,并且在重叠期内2种橐吾花粉和柱头均具有活性.2种橐吾共享传粉昆虫,为泛化传粉模式,并且传粉者对2种植物的访花行为相似,存在2种植物间交叉访问.尽管2种橐吾花期不存在完全的时间隔离,也不存在完全的传粉者隔离,但2种橐吾柱头对异源花粉的亲和性差,表现为柱头对异源花粉有明显的排斥反应,异源花粉管在柱头上的生长过程明显受阻.这种花粉-柱头排异形式很有可能是2种橐吾属植物重要的合子前隔离机制,从而避免了种间的自然杂交,更好地维持了自然种群的多样性和遗传结构的稳定性.     

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。     

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。     

黄帚橐吾挥发物的化感作用及其主要成分分析

研究了青藏高原东部高寒草场上广布的菊科毒杂草--黄帚橐吾(Ligularia virgaurea)挥发物的化感作用,并运用气相色谱-质谱方法对其挥发物主要化学成分进行了分析.结果表明,黄帚橐吾挥发物对所有5种受试牧草种子的萌发速度指数和最终萌发率均产生抑制作用,其中对垂穗披碱草和大雀麦的种子萌发抑制较为显著,而对早熟禾和羊茅影响较小.黄帚橐吾挥发物中主要含有2-甲基-庚烷(9.84%)、3-甲基-庚烷(8.25%)、庚烷(7.93%)、4-甲基-1-异丙基-双环[3,1,0]己-2-烯(7.79%)、3-甲基-己烷(6.38%)、2-甲基-己烷(5.54%)、D-苧烯(.70%)等18种化合物,占总挥发物的68.24%,其中单萜类化合物含量占总挥发物的16.58%,因此萜类化合物是其主要的化感活性物质,并通过挥发途径释放后影响到受体植物种子的萌发.黄帚橐吾挥发物的化感作用对增强其生存竞争力和种群增长起着重要作用.     

Reproductive modes of three Ligularia weeds (Asteraceae) in grasslands in Qinghai-Tibet Plateau and their implications for grassland management

Ligularia virgaurea, L. sagitta and L. przewalskii are noxious weeds that are widely distributed in psychro-grasslands in the east of Qinghai-Tibet Plateau of China. However, little is known about reproductive modes and genetic diversity in the populations of L. virgaurea and L. sagitta under different environmental conditions, although such information is available for L. przewalskii. In this study, two experiments were conducted to examine the ex situ clonal growth of these weeds, and to determine the effects of reproductive modes on the diversity of their populations, using random amplified polymorphic DNA (RAPD) analysis. Transplanted seedlings of L. sagitta and L. virgaurea showed considerable clonal growth by forming rhizomes and ramets. The RAPD analysis showed that all individuals collected from natural populations were genetically different, indicating that no individuals originated from clonal growth. It is concluded that L. virgaurea and L. sagitta have the capacity for clonal growth, but the expression of their clonality is constrained by their growing conditions in natural grasslands. Sexual reproduction plays the major role in their population diversity in natural habitats, which requires that the control measures for these weeds should be targeted at the sexually reproducing populations.     

高寒草甸黄帚橐吾种群根际/非根际土壤可培养微生物群落特征

黄帚橐吾（Ligularia virgaurea）是高寒草甸常见的毒杂草,被认为是指示一个地区草地植被退化的重要物种,研究其根际/非根际土壤微生物在不同海拔梯度上的群落特征具有重要意义。以甘南州高寒草甸不同海拔梯度黄帚橐吾根际/非根际土壤可培养微生物为研究对象,采用稀释涂布平板法和最大可能数法（MPN）测定了土壤微生物的数量及土壤理化因子的变化。结果表明：细菌在微生物总数中占比最大,根际微生物数量随海拔升高呈先增加后减小的变化,非根际则表现为递增的趋势,微生物功能群在根际和非根际土壤中均逐渐增加;根际土壤的微生物和功能群数量均高于非根际土壤。RDA分析发现,土壤温度、有机碳、电导率、pH、全氮、全磷、速效氮及脲酶对根际/非根际土壤微生物数量及功能群变化影响较大。通径分析可知：根际土壤中,细菌和真菌受速效氮和有机碳影响较大,放线菌主要受土壤温度和电导率的影响;根际土壤固氮菌和氨化细菌决策系数速效氮 > 有机碳 > 全氮;根际和非根际土壤中硝化细菌的影响因子各不相同,根际土壤决策系数最大和最小分别为全磷和全氮,非根际则是pH和脲酶。     

Arbuscular mycorrhiza regulate inter-specific competition between a poisonous plant, <Emphasis Type="Italic">Ligularia virgaurea</Emphasis>, and a co-existing grazing grass, <Emphasis Type="Italic">Elymus nutans</Emphasis>, in Tibetan Plateau Alpine meadow ecosystem

In order to investigate the function of arbuscular mycorrhizae in Tibetan Plateau Alpine meadow ecosystems, experiments were carried out to evaluate the influence of arbuscular mycorrhizal (AM) fungi on inter-specific competition between the poisonous plant, Ligularia virgaurea, and the grazing grass, Elymus nutans, at different relative densities. Our results showed that the biomass of L. virgaurea significantly declined in AM treatments while the biomass of E. nutans increased in our inter-specific competition system. Relative yields and root/shoot ratios of these two species indicated that AM fungi had a significant influence on the growth of L. virgaurea and E. nutans. This implies that arbuscular mycorrhiza do not benefit both plant species equally but rather contribute to the growth of E. nutans more than L. virgaurea. We thus provide evidence that AM fungi could regulate plant interactions and influence the plant community structure in the Tibetan Plateau Alpine meadow ecosystem.     

Seed-to-seed potential allelopathic effects between <Emphasis Type="Italic">Ligularia virgaurea</Emphasis> and native grass species of Tibetan alpine grasslands

Allelopathy is an important process in plant communities, but the role of seed allelopathy in natural ecosystems remains poorly understood. In the present study, we examined the potential allelopathic effects of Ligularia virgaurea (a dominant species in degraded Tibetan grasslands) seeds on the germination of four native grass species (Festuca sinensis, Agrostis gigantean, Bromus inermis, and Elymus nutans). The results showed that L. virgaurea seeds can have potential allelopathic effects on seed germination, mean time to germination and root growth rates of native grass species. We further demonstrate that these effects are driven by a water-soluble seed leachate. Species with smaller seeds were generally more sensitive than larger seeded species. The results suggest that seed-to-seed allelopathic potential may be an important mechanism driving the dominance of L. virgaurea in degraded alpine grasslands on the Tibetan Plateau. Further studies are required to demonstrate effects of seed-to-seed potential allelopathy in a field setting.     

植物替代控制3种入侵杂草技术的研究与应用进展

植物替代控制是利用一种或多种植物的生长优势控制入侵杂草的方法,它是控制外来杂草危害的有效途径之一;因其既可控制入侵杂草危害又能取得一定的经济效益及生态效益而被人们广泛接受。本文简要介绍了我国3种入侵杂草植物替代控制技术研究与应用现状,系统总结了植物替代控制技术阻截和修复豚草、紫茎泽兰及黄顶菊扩散危害的技术模式和效果,提出了植物替代控制技术未来研究的方向和重点。     

Relative Importance of Deterministic and Stochastic Processes in Driving Arbuscular Mycorrhizal Fungal Assemblage during the Spreading of a Toxic Plant

Both deterministic and stochastic processes are expected to drive the assemblages of arbuscular mycorrhizal (AM) fungi, but little is known about the relative importance of these processes during the spreading of toxic plants. Here, the species composition and phylogenetic structure of AM fungal communities colonizing the roots of a toxic plant, Ligularia virgaurea, and its neighborhood plants, were analyzed in patches with different individual densities of L. virgaurea (represents the spreading degree). Community compositions of AM fungi in both root systems were changed significantly by the L. virgaurea spreading, and also these communities fitted the neutral model very well. AM fungal communities in patches with absence and presence of L. virgaurea were phylogenetically random and clustered, respectively, suggesting that the principal ecological process determining AM fungal assemblage shifted from stochastic process to environmental filtering when this toxic plant was present. Our results indicate that deterministic and stochastic processes together determine the assemblage of AM fungi, but the dominant process would be changed by the spreading of toxic plants, and suggest that the spreading of toxic plants in alpine meadow ecosystems might be involving the mycorrhizal symbionts.     

Changes in soil biochemical properties associated with Ligularia virgaurea spreading in grazed alpine meadows

Ligularia virgaurea, a toxic perennial weed, has become a dominant species in the heavily-grazed alpine meadows of the eastern Qinghai?CTibetan Plateau over recent decades. We investigated changes in soil biochemical properties associated with L. virgaurea spreading in grazed alpine meadows at three sites. Soil and root biomass samples were taken at depths of 0?C8?cm, 8?C16?cm and 16?C24?cm from patches where L. virgaurea was dominant and from areas between L. virgaurea patches, with only other native species. Across sites, root biomass, acid-extracted carbohydrate C, organic C and N mineralization, and microbial biomass C and N concentrations in the top 8-cm layer were significantly higher while nitrate N concentration was significantly lower in L. virgaurea patches than in areas between L. virgaurea patches. The increased activities of dehydrogenase, ???Cglucosidase, urease and phosphatase in the top 8-cm layer under L. virgaurea were associated with enhanced soil microbial biomass. Our results indicate a close association between changes in soil biochemical properties in the top 8-cm layer and the spread of L. virgaurea in grazed alpine meadows of the eastern Qinghai?CTibetan Plateau.     

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.     

青藏高原生态屏障生态系统时空演变及驱动机制

青藏高原生态屏障是我国生态安全战略格局的重要组成部分,对我国乃至亚洲生态安全具有重要的屏障作用。研究青藏高原生态屏障生态格局的演变规律,探究影响生态系统变化的驱动机制,对青藏高原生态屏障建设和保障国家生态安全具有重要的意义。利用多源遥感数据,分析了2000-2015年国家屏障区生态格局演变规律,选取了自然和社会经济等驱动要素,采用冗余分析等方法解析了青藏高原生态屏障生态系统演变机制。结果表明：（1）2000-2015年间,青藏高原生态屏障生态系统格局发生明显变化,草地生态系统减少了1792 km²,主要转化为荒漠生态系统和河流生态系统,转换面积分别为1088 km²和832 km²;河流生态系统增加了1600 km²,主要由荒漠生态系统和草地生态系统转换而来,分别转换了1152 km²和832 km²。（2）2000-2015年青藏高原生态屏障生态系统格局演变驱动机制中以社会经济因子为主。冗余分析结果显示国内生产总值（83%）和人口数量（72%）贡献率最大;方差分析结果中国内生产总值和人口数量的共同贡献率达到了77%;地理加权回归结果显示生态系统演变受社会经济因子主导的区域面积比例为60.41%。（3）青藏高原生态屏障内人口数量的总贡献率达到了72%,在人口数量作用下生态系统呈正向转换的面积占比达到了86.24%,且人口数量与河流、森林生态系统的相关程度较高,综合说明了青藏高原生态屏障内生态保护政策的实施、生态工程的建设卓有成效。     

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.