Regeneration Capacity of Small Clonal Fragments of the Invasive Mikania micrantha H.B.K.: Effects of Burial Depth and Stolon Internode Length

The perennial stoloniferous herbaceous vine Mikania micrantha H.B.K. is among the most noxious exotic invaders in China and the world. Disturbance can fragment stolons of M. micrantha and disperse these fragments over long distances or bury them in soils at different depths. To test their regeneration capacity, single-node stolon fragments with stolon internode lengths of 0, 3, 6 and 12 cm were buried in soil at 0, 2, 5 and 8 cm depths, respectively. The fragments were growing for nine weeks, and their emergence status, growth and morphological traits were measured. The results indicated that increasing burial depth significantly decreased survival rate and increased the emergence time of the M. micrantha plants. At an 8-cm burial depth, very few fragments (2.19%) emerged and survived. Burial did not affect the total biomass and root to shoot ratio of the surviving M. micrantha plants that emerged from the 0- and 2-cm burial depths. Increasing internode length significantly increased survival rate and growth measures, but there was no interaction effect with burial depth for any traits measured. These results suggest that M. micrantha can regenerate from buried stolon fragments, and thus, disturbance may contribute to the spread of this exotic invader. Any human activities producing stolon fragments or facilitating dispersal should be avoided.     

How internode length,position and presence of leaves affect survival and growth of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> after fragmentation?

Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in stoloniferous plants. After fragmentation storage in stolon internodes and leaves may enhance survival and growth of stoloniferous plants. We hypothesize that (1) increasing length of the internode attached to the ramet and (2) presence of leaves will increase ramet survival and growth, and that (3) internode positions (before or after the ramet or both) will also play a role. We tested these hypotheses with the stoloniferous, invasive herb Alternanthera philoxeroides. In one experiment, we measured survival and growth of the ramets either without stolon internode (0 cm in length) or attached with internodes of 2, 4, 6 and 8 cm and either with or without leaves. In the other experiment, we measured survival and growth of the ramets attached with a proximal internode (before the ramet), a distal internode (after the ramet) or both. Increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower. These results support the hypotheses and suggest that storage in stolons and leaves may be of great significance for clonal plants in frequently disturbed habitats and may contribute greatly to the invasiveness of A. philoxeroides.     

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

Clonal integration enhances survival and performance of Potentilla anserina, suffering from partial sand burial on Ordos plateau, China

On Ordos plateau, a semi-arid, desertified area in China, sand burial is a common stress factor for plants. The extent to which sand burial occurs is heterogeneous and unpredictable in space and in time. Therefore, clonal fragments (i.e., interconnected ramets of a clonal plant) often experience partial sand burial, with some ramets buried in sand while the rest may remain unburied. It was hypothesized that clonal fragments are able to benefit from clonal integration, in case they experience partial sand burial. A pot experiment was conducted with Potentilla anserina, a stoloniferous herb often found on Ordos plateau. We used clonal fragments consisting of four interconnected ramets. In the experiment, the two proximal (older) ramets were unburied while the two distal (younger) ramets were either unburied (control) or buried with a 2, 4 or 6 cm deep layer of sand (burial treatments). The stolon connection between the proximal and the distal ramets was either severed or left intact. Stolon severing dramatically decreased the survival of buried ramets. Stolon severing and sand burial had significant effects on plant performance in terms of biomass production, number of leaves and leaf area. A cost–benefit analysis based on performance measures shows that the proximal ramets supported their connected distal ramets and did not incur any cost from this resource export. These results suggest that clonal integration, which is one of the functionally most important consequences of clonal growth, contributes significantly to our test species' capacity to withstand partial sand burial on Ordos plateau, a semi-arid and desertified area of China.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

Differential effects of clonal integration on performance in the stoloniferous herb Duchesnea indica, as growing at two sites with different altitude

Clonal integration may be adaptive and enhance the genet performance of clonal plants. Degree of clonal integration may differ between different environments . Here, a container experiment was used to determine how clonal integration affected the performance of the stoloniferous herb Duchesnea indica at two sites with different altitude along the transitional zone between the Qinghai-Tibet plateau and the Sichuan basin of Southwest China. In the experiment, the stolon between partially shaded two ramets experienced severing and intact treatments.We predicted that clonal integration would increase performance of whole clonal fragments and their shaded clonal parts at both sites. In both arctic and alpine environments, clonal plants may form highly integrated plant units (group of ramets).We predicted again that the reduction due to stolon severing in performance of whole clonal fragments and their shaded clonal parts would be greater at the site with high altitude than one with low altitude. The results indicated that the benefit for the shaded clonal parts and whole clonal fragments due to clonal integration was only observed at the site with high altitude. The results suggest that the performance of Duchesnea indica tends to be more responsive to the stolon severing at the site with high altitude than one with low altitude and support the second prediction. In addition, the effects of conditions of the sites and clonal integration on local morphological traits of ramets may be adaptive, five morphological traits of ramet-level (length of petiole, mean stolon internode length, specific petiole weight, specific stolon internode weight and specific leaf area) were investigated. Altogether, the results suggest that clonal integration might help D. indica plants to successfully inhabit the high-altitude habitat of the Qinghai-Tibet plateau of Southwest China, providing new evidences for the notion that clonal integration is an adaptive trait in stressful environments.     

海滨沙地单叶蔓荆匍匐茎对沙埋适应的生长对策

单叶蔓荆(Vitex trifolia var.simplicifoli)是一种耐盐、耐旱固沙地被植物。依据海滨沙地自然沙埋特点对单叶蔓荆匍匐茎进行了不同厚度(半埋和全埋)和不同长度交叉沙埋处理,研究探讨了单叶蔓荆沙埋适应生长对策,为其开发利用、科学管理和海滨环境修复提供指导。结果表明,正常情况下,单叶蔓荆匍匐茎基部和中部生长缓慢,顶部生长快。轻度(沙埋匍匐茎基部)和中度(沙埋匍匐茎基部和中部)半埋和全埋使匍匐茎顶部生长加速,茎长增长量较对照高出1.5到3.1倍;但重度(沙埋整个匍匐茎)半埋和全埋使匍匐茎顶部净增长量减少12%和13%。在20d沙埋中,对照整个匍匐茎各段均无不定根长出,但不同程度半埋和全埋沙埋处理下沙下匍匐茎上均长出不定根,重度半埋使不定根生长受抑;同时匍匐茎上各段茎生物量上升,枝叶生物量下降,且随着沙埋程度的增加而增减幅度提高,在重度半埋和全埋达到最大。在轻度和中度半埋和全埋下,匍匐茎上未沙埋部位枝条生长加速。研究表明,在自然环境中,单叶蔓荆匍匐茎顶端是一个对环境变化反应敏感的部位,并与沙埋后单叶蔓荆茎延伸生长和植株能否生存密切相关。当匍匐茎顶部没被沙埋时,沙埋促进沙埋部位匍匐茎和枝叶中物质转移,加速匍匐茎顶部快速生长和物质积累以弥补沙埋带来的损伤维持物质和能量的代谢平衡。沙埋后,单叶蔓荆以茎顶端快速生长、形成不定根、枝条生长维持茎水分平衡和能量和物质代谢平衡,以快速生长摆脱沙埋影响的生长方式为其对沙埋环境的重要适应对策。因此,在海岸沙地单叶蔓荆种群管理和维护中,在强风移沙引起的重度沙埋后,及时剥离匍匐茎顶部沙子对维护单叶蔓荆种群的延续生存和扩散均有重要作用。     

Shade avoidance in the clonal herb Trifolium fragiferum/: a field study with experimentally manipulated vegetation height

In herbaceous canopies light availability can show high degrees of spatial variability in a vertical and also in a horizontal direction. Stoloniferous plants are hence likely to encounter differences in light availability during their ontogenetic development. Different mechanisms, such as petiole elongation, plasticity in internode length and branching, and an enhanced allocation to sexual reproduction have been suggested to represent viable shade-avoidance mechanisms for clonal plants.In a field experiment we tested the response of the stoloniferous herb Trifolium fragiferum L. to experimentally manipulated vegetation heights. Naturally occurring clonal fragments were exposed to four different vegetation heights ranging from 0 cm (high light availability created by clipping the surrounding natural vegetation at ground level) to 20 cm (natural shading in closed canopy). The growth and development of individual clones was followed for two months. At the end of the experiment above-ground plant parts were harvested. Growth-related and morphological parameters (e.g., petiole and internode length) as well as patterns of meristem utilization (i.e., flowering, branching) were recorded.Neither primary stolon growth and biomass accumulation nor branching and flowering were significantly affected by treatments. However, increased vegetation height resulted in a reduced number of secondary ramets and also had strong positive effects on petiole length, leading to marked changes in the architecture of plants growing in canopies of different heights. In addition, the average weight of individual ramets on the primary stolon was markedly higher in plants exposed to taller vegetation as compared to shorter vegetation.The results of this study suggest the occurrence of a trade-off between clonal expansion (i.e., secondary ramet production) and the average size of clonal offspring. If grown under higher vegetation plants invested more into the size of individual ramets, especially into elongating petioles, and less into the growth and development of lateral branches. Placing leaf laminae higher up in the canopy results in an enhanced light interception which has apparently buffered negative effects of increased vegetation height on whole-clone biomass. Plants grown under shorter vegetation invested more into lateral spread by producing more, but smaller ramets.     

不同光照条件下聚花过路黄的克隆构型和分株种群特征

以都江堰灵岩山常绿阔叶林林下、林缘和林缘旷地3种不同光照环境中匍匐茎草本聚花过路黄(Lysimachla congestiflora)为对象,对其匍匐茎节间长、分枝强度、分枝角度、分株种群密度、分株种群生物量等指标进行了测定和分析．结果表明,聚花过路黄的匍匐茎节间长、分枝强度、分株种群密度和分株种群生物量在3种生境间差异显著．Kruskal Wallis检验表明,匍匐茎节间长度和分枝角度的频次分布在3种生境间差异显著．3种生境中匍匐茎节间长度分布偏斜度(skewness)的大小分别为：林缘旷地>林缘>林下．林缘旷地与林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异显著,而且林缘旷地生境中分枝角度分布偏斜度最小．林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异不显著．从林缘旷地、林缘到林下,聚花过路黄的克隆构型和分株种群特征发生了相应改变．结合克隆植物对资源的利用对策,讨论了不同生境中聚花过路黄克隆构型和分株种群特征可塑性的生态适应意义．     

匍匐茎草本金戴戴对盐分梯度的表型可塑性

 研究了匍匐茎型克隆草本金戴戴(Halerpestes ruthenica) 4种基株（基因型）对不同盐分处理（0,85.5, 171.0, 256.5和342.0 mM NaCl）的表型可塑性。随着盐分浓度的增加,实验植物与生长相关的性状指标 (如植株干重、总叶面积、分株数和总匍匐茎长度) 显著减小。植株干重、总叶面积和总匍匐茎长度具有显著的基株间差异。实验植物与形态相关的性状指标 (如平均叶柄长和根冠比) 对盐分梯度具有可塑性并具有显著的基株间差异;而其它形态指标 (如平均节间长、比节间长和比叶柄长)     

克隆整合对无芒雀麦(Bromus inermis)忍受沙埋能力的影响

无芒雀麦是浑善达克沙地植物群落中占优势的多年生根茎禾草。研究了克隆整合特性对无芒雀麦忍受沙埋能力的影响。结果表明,克隆整合显著提高了远端完全沙埋分株的存活,耗-益分析表明远端沙埋分株的生物量、分株数、叶片数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的近端非沙埋分株却没有产生显著的损耗,并且随着沙埋程度增加时,远端沙埋分株的收益有增大的趋势。因而,克隆整合特性是无芒雀麦对严酷沙埋环境形成的重要适应对策,它能够缓解沙埋对无芒雀麦存活、生长的胁迫,提高其在半干旱沙化地区的适合度。     

匍匐茎草本蛇莓对基质养分条件的克隆可塑性

A pot experiment with three levels of nutrient (N, P, K) supply was carried out to investigate clonal plasticity of stoloniferous herb Duchesnea indica Focke in response to nutrient availability. The plants had greater biomass at higher levels of nutrient availability. The root/shoot ratio of the plants changed with the nutrient availability in the following order: low level > high level > medium level. They had the largest biomass allocation to stolon at the medium level of nutrient availability. The biomass allocation to petiole did not respond to the treatments. The plants formed more stolons and ramets at the high and medium levels than the low level of nutrient availability. The petiole length, specific petiole weight（mg/cm）and stolon internode length of the plants did not respond to the treatments, while the specific stolon weight (mg/cm）of the plants was greater at the high and medium levels than the low level of nutrient availability. The results have been discussed in the context of adaptation of clonal plants to environmental heterogeneity.     

Effects of orientation on survival and growth of small fragments of the invasive, clonal plant Alternanthera philoxeroides

Background

The ability of small clonal fragments to establish and grow after disturbance is an important ecological advantage of clonal growth in plants and a major factor in the invasiveness of some introduced, clonal species. We hypothesized that orientation in the horizontal position (typical for stoloniferous plants) can increase the survival and growth of dispersed clonal fragments, and that this effect of orientation can be stronger when fragments are smaller and thus have fewer reserves to support initial growth.

Methodology/Principal Findings

To test these hypotheses, we compared performance of single-node pieces of stolon fragments of Alternanthera philoxeroides planted at angles of 0, 45 or 90° away from the horizontal position, with either the distal or the proximal end of the fragment up and with either 1 or 3 cm of stolon left attached both distal and proximal to the ramet. As expected, survival and growth were greatest when fragments were positioned horizontally. Contrary to expectations, some of these effects of orientation were stronger when attached stolons were longer. Orientation had smaller effects than stolon length on the performance of fragments; survival of fragments was about 60% with shorter stolons and 90% with longer stolons.

Conclusions/Significance

Results supported the hypothesis that orientation can affect establishment of small clonal fragments, suggested that effects of orientation can be stronger in larger rather than smaller fragments, and indicated that orientation may have less effect on establishment than amount of stored resources.     

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     

Burial depth and diameter of the rhizome fragments affect the regenerative capacity of a clonal shrub

Clonal plants in highly disturbed habitats are often broken into small fragments of various sizes and buried at various soil depths. As a storage organ, rhizome fragments play an important role in enabling plants to survive in such habitats. But few studies have been concerned about the regenerative capacity of rhizome fragments of clonal shrubs of different rhizome diameter and at different burial depths. Here, we investigated whether deeper burial decreased, and diameter of the rhizome fragment increased, the regenerative capacity of a clonal shrub. Research samples of rhizome fragment (rhizome diameters of 2, 5, 10, 15, and 20 mm) of the clonal shrub Calligonum arborescens were buried at different depths (0, 1, 5, 10, and 20 cm). Increasing the diameter of the rhizome fragments significantly increased the survival rate of fragments, and increased the above-ground, below-ground and total biomass production of fragments. Vegetative reproduction ability also increased with an increase in diameter of the rhizome fragments. With an increase in sand burial depth, above-ground, below-ground, total biomass production and vegetative reproduction ability first decreased and then increased, and no fragments survived at the 0 cm burial depth. These results indicate that sand burial depth and diameter of the rhizome fragments significantly affected the regeneration capacity of C. arborescens. Sand burial is one of the essential prerequisites for C. arborescens rhizome fragments’ survival. Moderate burial depth (5 cm) and larger fragment diameter (20 mm diameter) were more suitable for biomass production and vegetative reproduction. These results indicate that reserves stored in rhizome fragments can contribute greatly to the regeneration capacity of the C. arborescens—responses that are very important for C. arborescens survival and establishment in frequently disturbed habitats.     

克隆整合有助于狗牙根抵御水淹

尽管国内外开展了大量的克隆整合对克隆植物抵御逆境能力影响的研究, 但整合对植物抵御水淹能力的影响研究仍比较缺乏。该文从克隆整合的角度探讨多年生草本植物狗牙根(Cynodon dactylon)对水淹胁迫的响应。试验模拟了先端分株(相对年幼的分株)分别处于0、5和15 cm三种水淹胁迫环境, 并在每个水淹梯度下实施先端分株与基端分株(相对年长的分株)之间匍匐茎连接或切断处理, 调查水淹一个月后基端分株和先端分株以及整个克隆片段在形态和生理上的表现。研究发现: 切断匍匐茎连接显著降低了狗牙根先端分株的生长, 表现在生物量下降、匍匐茎长度减短和分株数减少等方面; 水淹显著抑制了先端分株的生长, 但对基端分株的生长并未造成显著影响; 在5 cm水淹处理下, 匍匐茎保持连接时, 先端分株和整个克隆片段的生长显著增加; 连接或切断处理在不同水淹梯度下对匍匐茎平均节间长没有显著影响, 对先端分株或基端分株在光化学转化效率上也未表现显著性差异。结果表明: 克隆整合效应促进了狗牙根在水淹胁迫下分株的生长, 并有助于整个克隆片段抵御水淹胁迫。     

匍匐茎草本活血丹(Glechomalongituba)在不同养分条件下的克隆形态

通过人工施肥的方法,对活血丹无性系实验种群在不同养分供应情况下的形态特征进行了研究。研究结果表明：施肥可以显著改变克隆植株的形态,无怀系分株产生能力增强,个体生长旺盛;匍匐茎总长增长,生物量增大,节间短,匍匐茎较粗壮;相对叶面积小,叶片厚,叶柄较粗短。在未施肥条件下,克隆植株形态表现则正好相反。养分供应的差异对叶片的形态以及根重和根量的影响较其对隔离者和叶柄的特征的影响更大。     

The role of vegetative spread and seed dispersal for optimal life histories of clonal plants: a simulation study

Sexual and vegetative reproduction of clonal plants phenotypically differ in dispersal distance, in the phenology of offspring production and establishment, and in the success of establishment. We applied a combination of analytical and of spatially explicit individual-based simulation modelling to calculate long-term fitness. We predicted optimal clonal plant life histories in a parameter space spanned by stolon number, stolon internode length, and relative allocation to sexual and vegetative reproduction. For a given allocation to sexual reproduction and number of stolons, fitness was optimised for rather short internode lengths under small disturbances, and for the longest possible internodes under larger disturbances. A trade-off between length and number of vegetative spacers drew parameters away from their unrestricted optima. Now, intermediate length and number of spacers led to maximum fitness under large disturbances. Simultaneous trade-offs between sexual and vegetative reproduction and between the length and number of spacers could also lead to fitness optima at intermediate parameter values, depending on the success of seedling establishment. We demonstrated that spatial habitat structure (1) selects for an efficient use of available space either by optimum internode length or by investment into seeds, which disperse farther than vegetative spacers, and (2) leads to an interaction between trade-offs. We conclude, that dispersal distance, i.e. a spatial life-history component, and trade-offs must be included in considerations on adaptive evolution of clonal life histories.     

匍匐茎草本活血丹(Glechoma longituba)在不同养分条件下的克隆形态

通过人工施肥的方法 ,对活血丹无性系实验种群在不同养分供应情况下的形态特征进行了研究。研究结果表明 :施肥可以显著改变克隆植株的形态 ,无性系分株产生能力增强 ,个体生长旺盛 ;匍匐茎总长增长 ,生物量增大 ,节间短 ,匍匐茎较粗壮 ;相对叶面积小 ,叶片厚 ,叶柄较粗短。在未施肥条件下 ,克隆植株形态表现则正好相反。养分供应的差异对叶片的形态以及根重和根量的影响较其对隔离者和叶柄的特征的影响更大。