基于复合节间组分关系的结缕草克隆植株形态可塑性特征分析

结缕草Zoysia japonica克隆植株的基本形态学单元是复合节间。分析复合节间组分关系,可以进一步揭示其形态可塑性的细微特征和资源传输与分配的差异,同时也可以为形态可塑性成因提供合理的生理生态学解释。本文对温室内、外两种环境条件下生长的结缕草克隆植株的复合节间组分进行测定和分析,结果表明：温室内、外植株主匍匐茎的长度和生物量的波动比一级分枝大,缩短节间长度比伸长节间长度稳定,但缩短节间生物量较伸长节间生物量变化幅度大。伸长、缩短节间长度、生物量分别与复合节序数呈二次函数关系。伸长、缩短节间生物量和分株生物量在一个生长季的生长方程亦均呈二次函数关系。主匍匐茎根数量、生物量随复合节数递增基本呈递增趋势。在两种环境下,A分株根数、生物量均大于B分株根数、生物量（在结缕草匍匐茎上,着生于每个复合节的基端和梢端缩短节间上的分株分别称为A、B分株）,且差异明显。温室内植株,在分株发育形成后,伸长、缩短节间的长度、生物量分别与匍匐茎根、A分株根数、B分株根数、生物量间相关性显著,温室外的相关性格局较复杂,且总体显著性低于温室内。结缕草克隆植株基本形态结构单元的复合节上的各个组成部分之间存在的密切相关关系,是导致结缕草克隆植物形成形态可塑性的细微特征格局的主要原因。     

克隆整合提高了入侵植物空心莲子草对北美车前的竞争力

选取北美车前Plantago virginica L.为竞争背景草,以空心莲子草Alternanthera philoxeroides（Mart.）Griseb.为研究对象,通过切断或保持其先端分株与后端分株间的匍匐茎连接,研究了克隆整合对先端分株生长以及竞争力的影响。结果显示：不管是否存在竞争,克隆整合显著提高了空心莲子草先端分株的生物量、总匍匐茎长度、叶片数目和分株数目,并降低了其对茎的生物量投资。此外,克隆整合显著提高了空心莲子草先端分株对北美车前的竞争力。种间竞争显著降低了空心莲子草的生长,但并没有显著影响其生物量分配。上述结果表明,克隆整合在一定程度上能够促进入侵克隆植物的生长和竞争力,从而可能潜在影响其入侵性。     

5个腰果无性系株形研究初报

比较了5个腰果无性系株形之间的差异,分析了各株形性状之间及其与产量性状之间的相关性。结果表明：5个无性系之间的主干高度、主干直径、2级枝梢长度、3级枝梢直径、4级枝梢长度、4级枝梢直径均存在显著差异,1级枝梢长度、1级枝梢直径、2级枝梢直径以及3级枝梢长度之间的差异均不显著;5个无性系之间的结果枝叶片数量、叶片长度、叶片宽度、株高、结果枝条数、结果数以及株产量均存在显著差异,冠幅之间的差异不显著。5个无性系的2、3、4级枝梢直径以及主杆直径之间的相关性达显著或极显著水平;株高与2、3级枝梢直径以及主杆直径之间具有显著的相关性;1级枝梢长度与1级枝梢直径之间呈显著的负相关关系;叶片长度和宽度之间呈极显著的正相关关系。产量与结果枝条数、结果数以及冠幅之间呈显著或极显著的正相关关系。5个无性系中,HL2-21、FL30分别适宜作抗风固沙、水土保持的生态经济林优良树种。     

克隆整合在空心莲子草抗病原真菌中的作用

入侵克隆杂草——空心莲子草在其入侵地给自然环境及社会经济造成严重危害,研究发现其野外种群很少感染病害。克隆整合作用在空心莲子草的生长发育中起着重要的作用,然而克隆整合作用对其抗病性影响的研究却很少。因此,设置实验以验证克隆整合在空心莲子草的抗病原真菌中是否起重要的作用。试验结果表明,空心莲子草的抗病原真菌能力显著强于同科植物苋菜,在接种病原真菌立枯丝核菌40 h后,苋菜叶片的侵染面积达到50%以上,而空心莲子草叶片的侵染面积不到10%;且克隆整合作用促进了空心莲子草顶端分株的地上部分的生长,茎段相连处理下,顶端的茎长、茎节数、茎的生物量、叶片数量、叶片生物量都显著高于基端,而茎段剪切处理下,顶端分株的地上部分生长并不显著大于基端分株;叶片接种病原真菌立枯丝核菌后,相连的顶端分株的总酚含量显著高于切断分株的顶端分株的含量。且二次接种病原菌后,相连分株的病原真菌侵染叶片面积不到0.1 cm~2,显著小于剪切分株的叶片侵染面积(0.8 cm~2)。结果表明,克隆整合作用不仅促进了空心莲子草相连分株的地上部分的生长,也提高了其抗病原真菌的能力。     

模拟采食下克隆整合对虉草生长及生理特征的影响

为了解克隆整合对克隆植物适应均质性采食环境胁迫的作用,以江西省鄱阳湖-乐安河段河岸带优势根状茎克隆物种虉草(Phalaris arundinacea)为对象,研究均质性采食条件下虉草的生理生态响应以及克隆整合在虉草应对模拟采食两月后发挥的作用.模拟采食(去叶)强度设置4组水平:不去叶(CK对照组)、去叶25%、去叶50%和去叶75%;克隆整合采用2种方式:茎切断和连接.结果显示,无论是否进行断茎处理,模拟采食均显著(P5%)降低虉草的叶片数、地上枝总长度、分株数和生物量,去叶处理下断茎的虉草分株相对茎连接分株具有更强生长优势.断茎和去叶处理对虉草叶绿素含量无显著影响,断茎的虉草分株在25%、50%去叶水平下叶绿素a(Chl a)、叶绿素b(Chl b)和总叶绿素(Chlt)含量均高于茎连接的分株,所有去叶处理强度下断茎的虉草分株各叶绿素指标均高于对照组.去叶显著影响了虉草分株各光合指标,断茎分株在50%、75%去叶水平上的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、胞间CO2浓度(Ci)等指标均高于茎连接分株,25%去叶处理组则相反;去叶处理的虉草分株的Pn、Tr、Ci均低于对照组.因此,采食引起虉草分株光合速率下降,使营养物质积累不足,导致生长状态下降;虉草能够在一定程度上适应均质性采食处理带来的压力,但是克隆整合不能显著改善均质性采食条件下虉草的生长.     

空心莲子草在不同生境中氮素迁移和表型可塑性差异

为探究克隆繁殖性入侵植物空心莲子草在不同生境下的生长特性,以更好地了解其整体的生存策略,利用15N同位素标定技术评估空心莲子草在陆生(Ⅰ)、湿生(Ⅱ)、水生(Ⅲ)3种不同生境相互转换中的氮素迁移效率,并对不同生境中的空心莲子草生物量分配和表型可塑性差异进行分析研究。结果表明,在不同生境中,空心莲子草氮素迁移量和各构件的生物量均存在显著差异。氮素迁移量存在显著差异(P0.05),具体生境体现为ⅡⅢⅠ;Ⅱ较Ⅲ平均增长23.32%,较Ⅰ平均增长54.49%。空心莲子草植株总生物量Ⅱ最大,Ⅰ最小。地下生物量投资与生境含水量呈显著负相关,体现为ⅠⅡⅢ,茎生物量投资比叶生物量投资平均高15.57%。在不同生境中,空心莲子草株高和茎节长度均体现为ⅡⅢⅠ,陆生、湿生、水生3种不同生境的母株处理下,Ⅱ株高分别较Ⅲ和Ⅰ增长39%、189%,52%、116%,56%、127%;茎节长度分别增长36%、150%,63%、117%,57%、94%。空心莲子草在不同生境下的氮素迁移量、各构件的生物量以及分配格局表现出的显著差异显示了其在湿生生境下具备更强的氮素迁移利用、表型可塑性和对异质环境的适应能力,揭示了克隆繁殖植物空心莲子草在湿地生境中具有更强的入侵性。该研究可为空心莲子草防控提供理论指导依据。     

九寨沟自然保护区4种水深梯度下芦苇分株地上生物量的分配与生长

芦苇是世界自然遗产--九寨沟自然保护区的一种重要的湿地植物.本文对该地区芦苇海4个不同水深梯度下芦苇的无性系地上分株的生物量分配、生长与繁殖策略进行了比较研究.结果表明,在47 cm (水在土层表面上)水深环境中,芦苇单株的平均生物量(4.2 g)最大.在-15 cm (水在土层表面下15 cm)的生境中,叶生物量百分比(叶生物量占单株总生物量的百分比, 46.1%)最大.芦苇地上分株高度、地上分株有分枝的单株百分比(茎上有分枝的分株占样地总分株的百分数)也存在明显的差别.茎的生物量分配百分比和生长速率随水深的增加而增加.在滩地生境中,开花率、花序的生物量百分比明显大于水较深的生境.分株株高与分株生物量、茎生物量与叶生物量都表现出较明显的幂指数异速生长规律.在-15 cm水深的生境中,叶生物量的生长快于茎的增长;在其他水深梯度的生境中,则是茎生物量的增加快于叶生物量的增加.     

基因型及播种密度对冬小麦分蘖期生长、生物量分配及产量的影响

采用双因素随机区组设计,研究了分别来自干旱与湿润区的4个小麦品种6个播种密度对冬小麦生长、生物量分配及产量的影响.结果表明,在本试验条件下,生长指标、生物量积累各指标及籽粒产量均主要受基因型和播种密度互作(g×d)影响,而成穗数、穗粒数主要受播种密度(d)的影响,基因型(g)则是引起小麦籽粒产量、千粒重及穗长变化的主要因素.白粒三号和川麦39最佳播种密度为315×104 hm-2,籽粒产量达到5 863.8 kg hm-2和5 882.1 kg hm-2;西农2000与陕麦139产量最佳播种密度为195×104 hm-2,籽粒产量分别为6 422.4 kg hm-2和7 062.4 kg hm-2.陕西小麦品种籽粒产量高出四川小麦约12.90%~18.62%.相关分析表明,籽粒产量与成穗数、千粒重呈极显著正相关(r=0.859 7**,r=0.499 3**),而与穗长、地上部干重及根系长度呈极显著负相关,与穗粒数及其它生长、生物量指标无显著相关.研究表明,干旱区小麦在湿润地区种植表现出较大的增产潜力,此外,在生产中应重视基因型和播种密度二者互作所表现出来的优势.表5参31     

淀山湖沉水植物——苦草群落恢复影响因子研究

在淀山湖0.5、1.0、1.5、2.0 m水层进行苦草(Vallisneria natans)现场生长实验,试图找出淀山湖苦草植被恢复的适宜环境条件。试验期间每天测量光照强度和水体溶解氧等环境因子,每隔10 d监测苦草的根长、株高、鲜重等生长指标的变化情况。结果表明:水深对苦草的生长具较显著影响,光照强度、溶氧分别和苦草日相对生长率具显著相关性;苦草的株高、叶片数、根长等生物学生长指标在0.5、1.0 m水层生长良好,1.5 m与2.0 m均出现不同程度的负增长状况。光照强度与苦草日相对生长率之间具有显著相关性(r=0.905),0.5、1.0 m水层的光照强度下苦草的相对生长率分别为0.14和0.11,而1.5、2.0 m水层的光照强度下苦草出现负增长的情况。水中的溶解氧含量与苦草成活率之间具有显著相关性(r=0.935)。随着水深的增加,溶氧逐渐降低,苦草的成活率也逐渐下降。在2.0 m水层处,溶氧均值为2.76 mg.L-1,苦草的成活率为46.5%,在1.0 m水层处,溶氧为5.66 mg.L-1,苦草成活率为86.5%,因此,淀山湖苦草群落恢复宜先在1.0 m以浅的沿岸带开展。     

淀山湖沉水植物——苦草群落恢复影响因子研究

在淀山湖0.5、1.0、1.5、2.0 m水层进行苦草（Vallisneria natans）现场生长实验,试图找出淀山湖苦草植被恢复的适宜环境条件。试验期间每天测量光照强度和水体溶解氧等环境因子,每隔10 d监测苦草的根长、株高、鲜重等生长指标的变化情况。结果表明：水深对苦草的生长具较显著影响,光照强度、溶氧分别和苦草日相对生长率具显著相关性;苦草的株高、叶片数、根长等生物学生长指标在0.5、1.0 m水层生长良好,1.5 m与2.0 m均出现不同程度的负增长状况。光照强度与苦草日相对生长率之间具有显著相关性（r=0.905）,0.5、1.0 m水层的光照强度下苦草的相对生长率分别为0.14和0.11,而1.5、2.0 m水层的光照强度下苦草出现负增长的情况。水中的溶解氧含量与苦草成活率之间具有显著相关性（r=0.935）。随着水深的增加,溶氧逐渐降低,苦草的成活率也逐渐下降。在2.0 m水层处,溶氧均值为2.76 mg.L-1,苦草的成活率为46.5%,在1.0 m水层处,溶氧为5.66 mg.L-1,苦草成活率为86.5%,因此,淀山湖苦草群落恢复宜先在1.0 m以浅的沿岸带开展。     

松嫩草地羊草克隆构型特征在不同种群密度下的可塑性

对不同密度羊草(Leymuschinensis)种群的克隆构型特征进行了调查,比较在资源水平相对一致时,羊草克隆构型特征,如根茎节间长度、间隔子长度、分枝距离和分枝角度,对种群密度变化的可塑性.结果表明:随着种群密度增加,羊草根茎节间长度和间隔子长度逐渐减小,分枝角度增大,且分枝角度在不同密度间差异显著.分枝角度的频率分布格局在不同密度间也存在显著差异,种群密度高时,羊草根茎具有更多的大分枝角.此外,羊草根茎节间长度、间隔子长度和分枝角度与其种群密度具有较强的冥函数关系.羊草根茎克隆特征在不同密度间具有较强的可塑性,当种群密度低时,羊草克隆特征更趋于“游击型”;随着种群密度的增加,其“游击型”特征减弱. 图2表2参19     

Size and estimated age of genets in eelgrass, Zostera marina, assessed with microsatellite markers

We examined the spatial distribution of genotypes in a perennial population of eelgrass, Zostera marina L., at two spatial scales. We mapped and sampled 80 ramets in a subtidal area of 20 × 80 m, and an additional 15 ramets in two 1-m² sub-quadrats. Ramets were genotyped for seven polymorphic microsatellite loci. Among a total number of 54 genotypes detected, 12 occurred more than once. The ramets of ten of these genotypes occurred in clusters and represented genets based on their expected multi-locus genotype frequencies. The size distribution of genets was uneven with estimated ramet numbers ranging from 2 to 5000. Whereas some areas displayed a high genet diversity, which may indicate past disturbances, large genets (≥10 m²) predominated in other locations of the sampled plot. Spatial heterogeneity in clone distribution was also obvious at the smaller sampling scale (15 ramets sampled within 1 m²) where the clonal diversity (genets identified/ramets sampled) was 0.24 in one quadrat, and 0.077 in the other. Ramets belonging to the largest clone were maximally 17 m apart, which corresponds to a genet age of 67 yr based on annual rhizome growth rates. We conclude that the spatial arrangement of clones in seagrasses allows inferences about the past demography and the disturbance regime at a given site which may prove useful for coastal zone management of ecologically valuable seagrass meadows. Received: 15 September 1998 / Accepted: 14 November 1998     

Evidence that reef-wide patterns of coral bleaching may be the result of the distribution of bleaching-susceptible clones

The hypothesis that intraspecific variation in coral bleaching is a result of the distribution of bleaching-susceptible clonal genotypes (genets) was addressed using photoquadrats recorded during the 1987 Caribbean bleaching event on a reef dominated by Montastraea annularis (Morphotype I), together with manipulative experiments with Porites porites. Nearest-neighbor analysis showed that bleached colonies (ramets) of M. annularis at 10 m depth had a high probability (0.80) of having a nearest bleached neighbor of the same genet rather than a bleached ramet of a different genet. Furthermore, the frequency distributions of bleached ramets of M. annularis in the photoquadrats was significantly different from a Poisson distribution, suggesting that bleached ramets were aggregated on the reef. Manipulative experiments with P. porites from 15 m depth showed that some genets were more susceptible to thermal bleaching than others, since three genets had significantly different rates of zooxanthellae loss when exposed to elevated temperatures in tanks receiving irradiances similar to those found in situ. These results suggest that the in situ patchy distribution of bleached ramets could correspond to the distribution of certain genets, and that adjacent genets can exhibit sufficiently different phenotypes to account for intraspecific variation in bleaching. Further studies of genet-specific coral bleaching may provide valuable insights into the causes and consequences of bleaching.     

Large scale parameter study of an individual-based model of clonal plant with volunteer computing

Understanding clonal strategies (i.e. the ability of plants to reproduce vegetatively) is particularly important to explain species persistence. A clonal individual may be considered as a network of interconnected ramets that colonizes space. Resources in this network can be shared and/or stored. We developed an individual-based model (IBM) to simulate the growth of an individual clonal plant. Typically a realistic IBM requires a large set of parameters to adequately represent the complexity of the clonal plant growth. Simulations in the literature are often limited to small subsets of the parameter space and are guided by the a priori knowledge and with heuristic aims of the researcher. The aim of this paper was to demonstrate the benefit of volunteer computing in computational ecology to systematically browse the parameter space and analyze the simulation results in order to draw rigorous conclusions. To be specific, we simulated clonal plant growth using nine growth rules related to the metabolic process, plant architecture, resource sharing and storage and nineteen input parameters. We chose 2-4 values per input parameter which corresponded to 20 millions of combinations tested through volunteer computing. We used three criteria to evaluate plant performance: plant total resource, ramet production and maximum length of one branch. The 1% top-performing plants were sorted according to these criteria. Plant total resource and ramet production were correlated while considering the top-performing plants. The maximum length of one branch was independent from the other two performance traits. We detected two processes promoting at least one of the plant performance traits: (i) a relatively high metabolic gain (high photosynthetic activity and low production cost for new growth units), a low resource storage and long integration distance for resource sharing; (ii) short spacer lengths and the predominance of elongation of existing branches over branching. Interactive effects between parameter values were demonstrated for more than half of the input parameters. Best performance was reached for plants with slightly different combinations of values for these parameters (i.e. different strategies) rather than a single one (i.e. unique strategy). This modeling approach with volunteer computing enabled us to proceed to large-scale virtual experiments which provided a new quality of insight into ecological processes linked with clonal plant growth.     

Multiple disturbances accelerate clonal growth in a potentially monodominant bamboo

Organisms capable of rapid clonal growth sometimes monopolize newly freed space and resources. We hypothesize that sequential disturbances might change short-term clonal demography of these organisms in ways that promote formation of monotypic stands. We examined this hypothesis by studying the clonal response of Arundinaria gigantea (giant cane, a bamboo) to windstorm and fire. We studied giant cane growing in both a large tornado-blowdown gap and under forest canopy, in burned and unburned plots, using a split-block design. We measured density of giant cane ramets (culms) and calculated finite rates of increase (lamda) for populations of ramets over three years. Ramet density nearly doubled in stands subjected to both windstorm and fire; the high ramet densities that resulted could inhibit growth in other plants. In comparison, ramet density increased more slowly after windstorm alone, decreased after fire alone, and remained in stasis in controls. We predict that small, sparse stands of giant cane could spread and amalgamate to form dense, monotypic stands (called "canebrakes") that might influence fire return intervals and act as an alternative state to bottomland forest. Other clonal species may similarly form monotypic stands following successive disturbances via rapid clonal growth.     

克隆植物的碳素生理整合及其生态学效应

碳素整合是克隆植物生理整合过程中非常重要的一部分,是克隆植物生存、生长、繁殖的物质基础.克隆植物碳素整合的基本机制具有一定的共同性或相似性,但克隆植物的种类不同、所处生境以及发育阶段不同或所受到的干扰、胁迫的程度不同,其碳素整合的范围、强度、方向、过程等也有所差异.本文对克隆植物碳素生理整合的基本特征、生态效应以及主要的影响因素进行了详细分析,并对迄今有关克隆植物碳素整合的最新研究进展进行了系统总结.同时,对碳素整合的主要研究方法进行了评价与展望,认为在今后研究中,应加强对克隆植物碳素整合的遗传学基础的研究,同时对于碳素整合的生理学、解剖学和生态学机制的研究也亟待深入.     

Clonal extent,apical dominance and networking features in the phalanx angiosperm <Emphasis Type="Italic">Zostera noltii</Emphasis> Hornem.

Disaggregating seagrass meadows and studying its components separately (clones, ramets, shoots) can provide us insights on meadow dynamics and growth patterns. The clonal growth, dependent upon clonal rules may regulate and impose constraints to plant architecture and, therefore, determine how individual clones evolve into the environment. In order to investigate the relationship between clonal growth rules and clone architecture, the belowground network architecture of single-clones of the seagrass Zostera noltii was studied. Networks were traced in situ after washing out the overlying sediment, and network characteristics were measured using digital analysis: area covered by clone, total rhizome length, type of rhizomatic axes (main, secondary, tertiary, quaternary), number and length of the internodes, branching angles and branching frequencies. This approach revealed that Z. noltii is able to develop into large clones integrating up to 300 internodes, 676 cm of rhizome, 208 shoots and 4,300 cm² of plant area. Internodal length depended on both, the distance to the apical shoot (time effect) and the axes type (apical dominance effect). However, average branching angle was independent of axis type (average 58.3 ± 0.75), but varied significantly depending on the distance from the apical shoot. This average branching angle allows Z. noltii maximize the rate of centrifugal expansion, maintaining a high density in colonized areas to produce close stands but also minimizing the investment in belowground biomass and ramets overlapping. The clonal architecture of Z. noltii seems to be regulated by the interaction of both, apical dominance strength and clonal integration distance. Moreover, clonal growth rules and growth pattern seem to constrain clonality through (clonal) plant architecture regulations (i.e. branching is restricted in secondary axes, similar average branching angles regardless the axes, the higher the distance to the apex the higher the number of internodes in secondary axes, shorter internodes in secondary and tertiary axes). Future research efforts should focus on how these complex relationships between apical dominance and clonal integration interact to elucidate the temporal (seasonal) and spatial scales of both processes and the outcome at the plant architectural level.     

Disturbance by ice and life-history adaptations of the seagrassZostera marina

The formation of winter ice, and its movement with the tides, has had a major influence on the life-history parameters of shallow-water populations of the rhizomatous marine angiosperm eelgrass (Zostera marina) in Nova Scotia, Canada. In this region, and annual form of eelgrass inhabits intertidal and shallow subtidal mudflats and a perennial form is common in subtidal areas. Where lowtide water depth was greater than winter ice thickness (25 cm), ice movement removed much of the aboveground biomass from perennial ramets, but did not influence the density of ramets. Measurements of primary production showed that perennial plants allocated a greater proportion of their total production to below-ground structures than measured in all previous studies on eelgrass. Thus perennials are well anchored in the sediment, are more able to withstand removal by ice, and have reserves available for production of new leaves when ice melts. In the spring, seed germination was greatest in areas where ice had removed whole plants (or their aboveground parts) than among mature ramets which had survived winter intact. Survival of seedlings (new genets) was not affected by shading from the adult canopy, but a shading experiment showed that competition for light with mature ramets had a significant negative effect on morphology, growth and the allocation of seedling biomass to below-ground parts, thus reducing the ability of new genets to survive ice disturbance in the next winter. The annual form ofZ. marina was restricted to areas where low tide water depth was much less than winter ice thickness. Annual plants did not survive winter, had small investment in below-ground parts and high reproductive effort, and overwintered as seeds. The genetic status of the two forms and the restriction of the annual to very shallow water are discussed in the light of previous work on eelgrass