扎龙湿地不同生境芦苇分株的形态特征及生物量分配的异速关系

不同环境条件下的植物个体可以表现出形态特征的分异和物质分配的权衡与调整。采用大样本抽样调查与统计分析方法,比较研究扎龙湿地不同生境芦苇(Phragmites Australis)生殖株和营养株的形态特征以及生物量分配的异速关系。结果表明：在9月末,盐碱生境、旱生生境、湿生生境和水生生境芦苇分株的生长表现出较大的生态可塑性,株高和株重均以盐碱生境最小,水生生境最大,最大值与最小值的比值分别为1.3—3.3和1.8—5.1,分株生长在种群间的变异度高于种群内,与营养株相比,生殖株的变异度较低;分株的支持分配与生产分配的比值为1.8—4.2,生产分配以盐碱生境最高,以水生生境最低,而支持分配和生殖分配表现与生产分配相反的序位;生殖株的花序长和花序重与株高间呈直线函数形式增长,株高和株重低于种群平均值的20%和35%的分株不进行有性生殖;叶重、叶鞘和茎重以及分株重与株高间呈幂函数形式的异速生长关系。植物通过改变个体的形态特征以及调整构件间生物量分配适应不同环境,而受遗传因素控制的构件间生长关系却相对稳定。     

Responses of rhizomatous grass Phragmites communis to wind erosion: effects on biomass allocation

Background and aims

The ability of modifying biomass allocation to deal with different environmental stress is an important mechanism for plant population expansion and maintenance in the unstable dune environment where wind erosion persists. However, how biomass is partitioned between horizontal rhizome extension and vertical ramet growth in response to wind erosion has not been fully understood. The objective of this study was to explore how wind erosion affected the relationship between horizontal rhizome extension and vertical ramet growth using a common rhizomatous perennial grass, Phragmites communis.

Methods

We dug 300 cm?×?200 cm, 80 cm deep pits in a garden experiment plot. Clonal fragments of P. communis were planted individually at a depth of 40 cm in these pits for 4 weeks before treatments. Surface sand was gradually removed to the final depth of 0 (control), 10, 20, 30 and 40 cm (maximum sand removal). Ramet emergence time, rhizome-based and tiller-based ramet number, rhizome number and length, biomass of vertically and horizontally oriented structures were monitored at the end of the experiment.

Results

With increasing erosion depth, the proportion of tiller-based ramets (in total number of ramets) increased, whereas that of rhizome-based ramets decreased. With increasing erosion depth, the percentage of vertically oriented structures biomass in total biomass increased significantly, whereas that of horizontally oriented structures biomass decreased.

Conclusions

The changes in biomass allocation (i.e., more allocation in vertical than horizontal biomass) together with a trade-off in tiller-based and rhizome-based ramets may enable P. communis to make better use of the resources in erosion conditions and maximize plant population expansion and maintenance.     

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。     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

Ramet demography and ecological attributes of the perennial river corridor plant Cnidium dubium (Schkuhr) Thell. (Apiaceae)

Cnidium dubium is a highly endangered, perennial river corridor plant in Central Europe. Here we summarise a 4-year field study of its ramet demography, growing in two different flood meadows in the Lower Havel River Valley in northeastern Germany, emphasising the influence of site-specific abiotic factors, mowing and the summer flood of 2002.We recorded 1658 ramets at densities between 2 and 180 ramets m⁻², which varied over area and time, indicating a high small-scale turnover of this species within grasslands. The half-life of the different ramet cohorts was estimated to range from 0.1 to 1.3 years, but single ramets of more than 4 years old were also found. Ramets are monocarpic: they propagate only once in their life. The highest probability of a vegetative ramet becoming a flowering stalk was 11.6%. Ramet density, cumulative number of newly produced ramets per year, leaf length, leaf age, reproductive growth and recovery from the summer flood of 2002 varied, depending on site. C. dubium had a higher performance at sites in intermediate zones of the overall flooding gradient compared with higher zones. Flooding stress is the controlling factor for the lower limit of its performance, while there was no evidence for an upper limit controlled by summer drought. Mowing slightly increased the cumulative number of leaves per ramet, but it decreased the size of new leaves and also the summer above-ground biomass per ramet. As a consequence, the most marked effect was that fewer ramets entered the generative life stage, but the plants were able to keep their overall reproductive performance on a level independent of mowing. Knowledge of basic growth patterns of C. dubium in natural populations could help to explain the species rareness, and may also suggest appropriate strategies to manage its survival.     

Ramet dynamics in a centrifugally expanding clonal sedge: a matrix analysis

Carex humilis is a clonal sedge that can form distinct rings of densely aggregated ramets. We hypothesize that rings form because both production of new ramets and ramet dispersal are positively correlated to ramet size. This would lead to an overrepresentation of fast-moving and large ramets with high ramet production at the periphery, whereas slow-moving and small ramets with low ramet production would mainly be found in the interior of rings. We use matrix models to analyse how ramet populations both at the periphery and in the interior develop in the absence of ramet dispersal. We found that the stable size class distributions of ramets predicted by the models were not different from the distributions found in the field. Also, the asymptotic ramet population growth rates (λ₁) were the same. Hence, we conclude that rings would form even in the absence of a link between ramet dispersal and ramet production. Further analysis of the matrix models showed that the ramet population increases at the periphery but decreases in the interior of rings because medium and large ramets produce fewer large ramets in the interior than at the periphery. We also found that the temporal variance in λ₁ and transitions rates during the four study years was much higher at the periphery than in the interior. Our results suggest that rings may form because C. humilis ramets use below-ground resources from a much larger area than the one covered by the shoots. As the clone grows larger, the soil volume available to the ramets in the interior decreases because their access to soil outside the ring is cut-off by the ramets at the periphery. Ramet density in the interior is therefore decreasing.     

松嫩平原碱化草甸星星草种群分蘖株延长生殖生长的表型可塑性调节

在松嫩平原碱化草甸,采用大样本随机取样的方法,研究了不同时间到达抽穗初期、抽穗期、开花期和乳熟期的星星草种群生殖分蘖株数量性状的可塑性及其调节规律.结果表明：除在开花期存在一定的波动外,在每5 d的时间里,随着生殖生长时间的延长, 星星草种群在抽穗初期、抽穗期和乳熟期的生殖分蘖株高、分蘖株生物量、花序长和花序生物量均依次显著增加.各生育期的星星草种群生殖分蘖株高与花序生物量呈显著正相关,而与生殖分配呈显著负相关.随着生殖生长时间的延长,抽穗初期、开花期和乳熟期的星星草花序生物量随着分蘖株高增加,其幂函数的增长速率均呈增大趋势.生殖生长时间延长10 d,其抽穗初期和抽穗期的生殖分配直线下降速率分别降低了43.2%和44.31%;延长5 d,乳熟期的下降速率降低了130%.表明星星草种群分蘖株生殖生长的表型可塑性调节遵循着一定的规律.     

Demography of the clonal palm Geonoma brevispatha in a Neotropical swamp forest

Abstract We expected that in clonal plants with the phalanx growth form, prereproductive ramets function as a meristem bank that helps buffer ramet demography against environmental fluctuations. Specifically, we evaluated four expectations for the forest understorey clonal palm Geonoma brevispatha in a swamp forest in south‐eastern Brazil. Our expectations were are follows: (i) the vital rates of prereproductive ramets are more responsive to genet‐related characteristics (genet size, distance and size of the nearest genet neighbour, and reproductive ramet survivorship) than to environmental variables (canopy openness, basal area of small trees, flooding, and distance and size of the nearest large tree neighbour), whose effects are expected to affect ramet dynamics indirectly through physiological integration; (ii) the progress of prereproductive ramets to later stages is dependent on the death of reproductive ramets; (iii) the population dynamics of G. brevispatha is resistant to significant interannual changes in the vital rates of ramets, (i.e. the occurrence of good and bad years); and (iv) there are large negative covariances among demographic transitions within a given year, as well as between years, reflecting the flexibility of G. brevispatha to adjust to environmental variability. Expectations 1, 2 and 4 were confirmed, while expectation 3 was only partially confirmed. Matrix models predicted that the growth rate of the overall ramet population was significantly greater than unity in the first study year, but it was significantly smaller than 1.0 in the second study year. Ramet population growth rates at the level of individual genets were greater than unity in the majority of genets in the first year, but in the second year genets with growth rates smaller than unity prevailed. Ramet population dynamics seems to be resistant to the succession of good and bad years; in effect, the presence of a meristem bank of ramets helps stabilize λ.     

天津沿海滩涂互花米草种群生殖分株数量特征及生殖分配研究

通过对天津沿海滩涂单优群落的取样调查,研究了互花米草种群生殖分株的数量特征和生殖分配规律。结果表明,在籽实成熟期,互花米草种群单个生殖分株重为15.62±9.26 g,穗和种子重分别为2.68±2.08和1.39±1.12 g,小穗和种子数分别为537.7±362.2和490.2±376.3个,生殖分配Ⅰ和生殖分配Ⅱ分别为15.02±5.83和7.62±3.8%,结实率为81.9 ±28.6%。穗和种子的形成分别需要分株积累3.2和3.6 g以上的生物量。穗长、穗重、种子数、小穗数、生殖分配Ⅰ和生殖分配Ⅱ分别与分株高度呈极显著(p<0.01)的正相关关系。穗重、小穗数、种子重、种子数、生殖分配Ⅰ和生殖分配Ⅱ分别与分株重呈极显著(p<0.01)的线性正相关关系。种子数、小穗数、生殖分配Ⅰ和生殖分配Ⅱ分别与茎和叶鞘生物量分配呈极显著(p<0.01)的负相关关系。穗各构件在空间上的分布也存在较强的规律性。     

不同水分下准噶尔无叶豆分株种群特征和生物量分配差异

准噶尔无叶豆是豆科无叶豆属小半灌木,在中国仅片断化分布于新疆古尔班通古特沙漠局部区域.该种自然种群种子萌发率极低(<3%),主要靠克隆繁殖维持种群.为研究克隆生长在异质性生境中的可塑性特征及其生态适应意义,对生境土壤含水量存在差异的两个准噶尔无叶豆自然种群(河边种群,沙漠种群)的分株种群特征、生物量及其分配比重进行了比较,研究发现:①在分株种群特征上,河边种群的分株显著高于沙漠种群,分株间距也显著大于沙漠种群.沙漠种群分株密度和根的密度显著大于河边种群,但每一分株拥有根的概率显著小于河边种群;②在生物量分配方面,河边种群具有比沙漠种群大的根生物量,但根茎生物量则要显著的小于沙漠种群;③在植株各部分生物量比重随植株大小的变化趋势上,河边种群各部分的比重随植株大小变化的趋势平缓,而沙漠种群各部分的比重变化趋势则较为急剧,表明河边准噶尔无叶豆分株种群随着植株的大小变化,资源分配比例并没有发生很明显的变化,而沙漠种群变化较大.结果表明,准噶尔无叶豆种群对所处生境水分条件存在可塑性响应,并通过分株种群特征变化和生物量分配差异应对水分条件的变化,形成自身的适应对策.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

扎龙湿地芦苇分株生态可塑性及其对土壤因子的响应

扎龙湿地的芦苇既可形成大面积的单优群落,也可形成不同群落斑块。采用大样本抽样调查与统计分析方法,对湿地内水生生境、湿生生境、旱生生境和盐碱生境芦苇种群分株高度和生物量进行比较。结果表明,6—10月份,4个生境芦苇种群分株高度及生物量均以水生生境最高,盐碱生境最低,水生生境株高为盐碱生境的1.5—2.3倍,分株生物量为2.0—5.1倍,生境间的差异性以及差异序位均相对稳定。4个生境株高生境间变异系数(19.45%—31.56%)均高于生境内变异系数(8.07%—17.61%),分株高度在生境间的可塑性更大;分株生物量中水生生境、湿生生境和盐碱生境3个生境间的变异系数(33.43%—55.61%)均低于生境内变异系数(44.85%—79.82%),分株生物量在生境内的可塑性更大。不同生境条件下芦苇种群分株,在生长和生产上均存在较大的生态可塑性,表现出明显的环境效应,其中土壤含水量是该地区芦苇分株生态可塑性变异的主要驱动因子(R0.80),为正向驱动。     

慈竹母株大小对克隆生长的影响

 克隆生长在慈竹种群更新中占有重要地位。本文以基径和生物量两个指标同时衡量慈竹(Neosinocalamus affinis)的母株大小,并分析了母株大小对新竹笋产生、竹笋大小和根茎大小的影响。结果表明：1）慈竹的成竹基径与一级母株基径、二级母株基径以及与2龄和3龄母株平均单株生物量呈正相关,但是成竹基径随一级母株基径增加比随二级母株基径增加而增加的速率快得多;2）慈竹母株大小与各样方中出笋数、活笋数和死笋数无显著的正相关;3）发笋母株的基径显著大于未发笋母株,但发一个笋母株与发两个以上笋母株的基径无显著差异;4）母株基径、2龄和3龄母株平均单株生物量都与根茎的长度和粗度呈显著正相关,且竹笋基径与根茎粗度也呈显著的正相关。母株大小对慈竹竹笋的影响是通过影响根茎实现的。总之,慈竹的克隆生长在个体水平受母株大小制约。     

松嫩平原全叶马兰种群不同龄级分株的生长策略

全叶马兰为根蘖性多年生草本菊科植物,种群由3个龄级无性系分株组成,在松嫩平原6月份,是全叶马兰营养生长的旺盛期,研究结果表明,分株的生长与生产力以1龄级最低,2、3龄级依次增高,3个龄级分株的生物量分配均为茎>叶>主根,叶的生物量分配为1龄级>2龄级>3龄级,茎和主根的均为1龄级<2龄级<3龄级,3个龄级分株总重与分株高度、分株叶重与茎重、分株主根重与枝条重之间均为幂函数异速生长规律,主根的生物量分配则随着分株高度的增加呈线性减少,全叶马兰无性系分株的生长与分配策略是在分株幼小时将较多的物质分配给叶,以保证物质生产器官叶的建造上,随着分株生长逐渐将生产的物质分配到茎和主根的生长与贮藏;分株不向主根投入较多的能量,但在分株幼小时向主根中分配的物质相对较多,随着分株的生长则按比例较小,并保持相对稳定的分配比率     

Effects of Heterogeneous Competitor Distribution and Ramet Aggregation on the Growth and Size Structure of a Clonal Plant

Spatially heterogeneous distribution of interspecific competitors and intraspecific aggregation of offspring ramets may affect the growth and size structure of clonal plant populations, but these have been rarely studied. We conducted a greenhouse experiment in which we grew a population of eight offspring ramets (plants) of the stoloniferous clonal plant Hydrocotyle vulgaris aggregately or segregately in two homogeneous treatments with or without a competing grass Festuca elata and a heterogeneous treatment with a patchy distribution of the grass. In patchy grass treatments, H. vulgaris produced markedly more biomass, ramets and stolons in open patches (without grasses) than in grass patches, but displayed lower size variations as measured by coefficient of variation of biomass, ramets and stolons among the eight plants. In open areas, H. vulgaris produced statistically the same amounts of biomass and even more stolons and showed higher size variations in patchy grass treatments than in open (no grass) treatments. In grass areas, H. vulgaris grew much worse and displayed higher size variations in patchy grass treatments than in full grass treatments. Ramet aggregation decreased the growth of H. vulgaris in open treatments and in both open and grass patches in patchy grass treatments, but had little effect in full grass treatments. Ramet aggregation had little effect on size variations. Therefore, heterogeneous distribution of competitors can affect the growth and size structure of clonal plant populations, and ramet aggregation may decrease population growth when they grow in open environments or heterogeneous environments with a patchy distribution of interspecific competitors.     

外部干扰对根茎型克隆植物甘草自然种群植株生长及主要药用成分含量的影响

克隆植物, 特别是游击型克隆植物, 其分株在一定时间段往往通过连接结构(即间隔子)连接在一起形成克隆网络, 基于克隆网络的克隆植物特性显著提高了克隆植物应对外部干扰的能力。长期封育禁牧使得甘草(Glycyrrhiza uralensis)自然种群免受外部干扰, 却并没有促使受破坏的甘草自然种群快速恢复。因为克隆整合、克隆储存及克隆分株的选择性放置等克隆植物特性的存在, 适度的外部干扰有利于根茎型克隆植物甘草的生长与繁衍。但外部干扰与克隆植物特性在甘草种群恢复中的作用及其机理目前并不清楚。该研究以甘草自然种群为研究对象, 通过野外控制实验进行模拟动物采食和人工采挖干扰, 探讨不同程度干扰对甘草自然种群生长和主要药用成分含量的影响。结果表明在中度放牧采食干扰或者人工采挖干扰下, 甘草自然种群的分株密度、高度和地上生物量与对照相比均没有显著差异, 甘草表现出等补偿生长; 而在不同类型的重度干扰下, 甘草产生的响应不一致。重度放牧采食干扰下甘草表现出欠补偿生长, 而重度人工采挖干扰下甘草表现出超补偿生长。克隆植物特性在甘草应对外部干扰过程中发挥着重要作用。外部干扰一定程度上提高了甘草中甘草苷和甘草酸的含量, 从而提升了药材的品质。进一步的研究应该关注于甘草自然种群与环境因子之间的关系, 解析甘草自然种群退化的原因, 提出相应的复壮技术, 探讨可持续利用的管理模式。     

松嫩平原不同生境芦苇种群分株的生物量分配与生长分析

在松嫩平原8月中旬,4个旱地生境芦苇种群分株的生长和生产力以季节性临时积水的低洼地为最高,以碱斑地为最低,两个生境之间分株高度相差2.8倍,分株生物量相差4.4倍,其它2个生境沙地和草甸混生 少也达到了显著或极显著水平,不同生境芦苇种群分株生物量分配的差异,蕴涵着重要的生长调节和物质分配策略。在分株株较小而又有充分生长空间的沙地生境和三斑生境,以及具有种间竞争的草句生境,分株将50％以上的物质分配给叶的建造上,以保证充分的物质生产;在环境条件相对较好的低洼地生境,分株将近40％的物质分配给茎的伸长生长,以提高个体对空间和光资源的竞争力。4个生境芦苇种群在分株增高与增重,以及茎、叶与分株的相对增重均具有相同的幂函数异速生长规律。     

Differential effects of water depth and sediment type on clonal growth of the submersed macrophyte <Emphasis Type="Italic">Vallisneria natans</Emphasis>

The response of clonal growth and ramet morphology to water depth (from 60 to 260 cm) and sediment type (sand versus organic clay) was investigated for the stoloniferous submersed macrophyte Vallisneria natans in an outdoor pond experiment. Results showed that water depth significantly affected clonal growth of V. natans in terms of clone weight, number of ramets, number of generations, clonal radius and stolon length. V. natans showed an optimal clonal growth at water depths of 110–160 cm, but at greater depths clonal growth was severely retarded. A high allometric effect was exhibited in ramet morphology. Along the sequentially produced ramet generations, ramet weight and plant height decreased while stolon length and the root:leaf weight ratio increased. When using ramet generations as covariate, sediment type rather than water depth more strongly affected the ramet characteristics. For plants grown in clay, ramet weight, ramet height and stolon length were greater, and plants exhibited lower root:leaf weight ratio. These data suggest that water depth and sediment type have differential effects on clonal growth of V. natans: Water depth appears primarily to affect numerical increase in ramets and spatial spread, whereas sediment type mainly affects biomass accumulation and biomass allocation. Handling editor: S. M. Thomaz     

两种践踏胁迫下克隆整合对入侵植物空心莲子草生长的影响

研究了两种践踏胁迫下克隆整合对入侵植物空心莲子草生长的影响.结果表明:(1)切断分株间的匍匐茎连接,会降低先端分株的生物量、分株数、总匍匐茎长度和总叶片数,但会显著增强基端分株的生物量.(2)对先端分株的践踏胁迫会显著降低先端分株叶片的叶绿素相对含量,对基端分株的践踏胁迫会显著降低基端分株的生物量和总匍匐茎长度.(3)对于基端分株的分株数、总匍匐茎长度和总叶片数来说,当进行基端分株践踏胁迫时,匍匐茎连接对其影响不大,而当进行先端分株践踏胁迫时,则明显对其不利.(4)对于整个克隆片段,践踏胁迫的差异和匍匐茎是否切断对其生长没有显著影响.     

Clonal integration affects growth and sediment properties of the first ramet generation,but not later ramet generations under severe light stress

克隆整合影响严重光胁迫下第一分株世代的生长和沉积物特征但不影响 后续分株世代的生长和沉积物特征 克隆整合通过缓冲环境压力和提高资源获取效率使克隆植物受益。然而,在一个克隆系统中,受益于克隆整合的连接分株世代的数量很少受到关注。我们进行了一个盆栽实验来评估沉水植物苦草 (Vallisneria natans)克隆系统内的生理整合程度,该克隆系统由一个母株和3个依次连接的后代分株组成。 母株生长在正常光照下,而后代分株被严重遮荫。母株与后代分株间的匍匐茎被切断或保持连接,但3个后代分株之间的连接仍然存在。与遮荫的后代分株连接时,苦草未遮荫的母株的光合能力显著增强,但其生物量积累大大减少。克隆整合显著增加了第一分株世代(相邻分株)的生物量积累和土壤的碳氮可用性、胞外酶活性和微生物生物量,但没有增加后续分株世代的这些特征。我们的结果表明,在严重光胁迫下,来自苦草母株的支持可能仅限于克隆系统中相邻的后代分株,这暗示着一个分株世代的效应。我们的结果有助于更好地理解克隆植物的层次结构和分段化。这些发现表明克隆整合程度在分株种群的生态相互作用中起着至关重要的作用。