Effects of nitrogen addition on clonal integration between mother and daughter ramets of Moso bamboo: a 13C-CO2 pulse labeling study

用¹³C-CO₂脉冲标记法研究氮添加对毛竹母子分株克隆整合的影响 相连分株间的资源共享(即克隆整合)是克隆植物的显著特征。克隆整合使毛竹(Phyllostachys pubescens)对多种环境条件具有较强的适应性。但毛竹通过克隆整合获得性能提升的机制尚不明确。本 研究区分并分析了毛竹光合碳的顶向整合和基向整合,以探究毛竹克隆片段如何在土壤氮异质性条件下 提高整体收益。本研究以由两株不同年龄毛竹分株组成 的克隆片段为研究对象,分株间通过根状茎连接。 每个分株设置氮添加或无氮添加两种处理,并通过单株母株或子株的¹³C-CO₂脉冲标记对光合碳的顶向整合和基向整合进行区分。研究结果表明,子株氮添加显著促进了光合碳由母株向子株的顶向整合,无论母株是否有氮添加,均向氮添加子株转运更多的光合碳。转运自无氮添加母株的光合碳主要分配至氮添加子株的 叶片,而转运自氮添加母株的光合碳则主要分配至根系。与无氮添加处理的子株相比,氮 添 加子株转运更多的光合碳至母株,且该过程不受母株氮添加的影响。转运自氮添加子株的光合碳主要 分配至无氮添加母株的根系和氮添加母株的叶片。这些结果说明,母株优先将更多的资源投入到具有高 养分可利用性的子株,随后子株作为更高效的资源采集点,根据母株的养分状况针对富集资源进行获取。通过这种方式,克隆植物可以降低资源获取成本并提高资源获取效率,最大限度地提升其整体表现。     

Water translocation between ramets of strawberry during soil drying and its effects on photosynthetic performance

To explore the mechanisms underlying water regulation in clonal plants and its effects on carbon assimilation under water stress, we studied the responses of water status, gas exchange and abscisic acid (ABA) contents to water stress in leaves of pairs of strawberry ramets that consist of mother and daughter ramets. There was a greater decrease in photosynthetic rates (P_n) and stomatal conductance (G_s) in the disconnected mother ramets than the connected mother ramets upon exposure to water stress, indicating that water stress in mother ramets was alleviated by water translocation from the well‐watered daughter ramets. Conversely, the connected mother ramets displayed enhanced symptoms of water stress when the connected daughter ramets were exposed to water deficit. The mother ramets had lower water potential (ψ_w) due to their stronger osmotic adjustment than in well‐watered daughter ramets; this resulted in water flow from the connected daughter ramets to mother ramets, thus alleviating water stress of mother ramets. During soil drying, there was a striking increase in ABA concentrations in leaves of the disconnected mother ramets, whereas leaf bulk ABA was much lower in the connected and water‐stressed mother ramets than that in the drought‐affected mother ramets in the disconnected group. In this study, though G_s was linearly correlated with leaf bulk ABA and ψ_w, G_s in water‐stressed mother ramets in disconnected group exhibited less sensitivity to the variation in leaf bulk ABA and ψ_w than that in connected and water‐stressed mother ramets. Taken together, these results indicate that: (1) the flux of water translocation between the connected ramets is determined by a water potential gradient; (2) water translocation between connected ramets helps to keep sensitivity of G_s to ABA and ψ_w in drought‐affected ramets, thereby benefit to effectively maintain the homeostasis of leaf water status and (3) the improvements in P_n in water‐stressed ramets due to water translocation from well‐watered ramets suggest the advantages of physiological integration in clonal plants in environments with heterogeneous water distribution.     

CLONAL INTEGRATION: NUTRIENT SHARING BETWEEN SISTER RAMETS OF SOLIDAGO ALTISSIMA (COMPOSITAE)

To test whether sharing of resources occurs among connected ramets of the tall goldenrod, Solidago altissima, we examined the extent of clonal integration for nutrients. In a greenhouse experiment, two-ramet clones were grown in a triad of connected pots so that nutrients could be supplied to either sister ramet or to their old rhizome (mother rhizome). Mother rhizomes and their associated roots shared nutrients with daughter ramets; however, any nutrient sharing that occurred between sister ramets was too little to significantly affect their growth. In addition, sister ramets not only competed for nutrients through parental connections, but larger ramets inhibited the growth of smaller ramets. We suggest that, for tall goldenrod, a clonal growth strategy in which nutrients are not shared among sister ramets may increase genet fitness by reducing the rhizome production of ramets in poor-nutrient microsites. Consequently, the genet would produce relatively fewer ramets in unfertile areas and make better use of heterogeneous nutrient resources.     

疏叶骆驼刺母株与子株间的水分整合

在未灌溉的土地上, 疏叶骆驼刺(Alhagi sparsifolia)通常不能进行有性繁殖, 克隆繁殖是其种群维持和延续的唯一方式。因此, 克隆性及其相关克隆性状(如水分整合)在疏叶骆驼刺自然种群的维持过程中可能扮演了极其重要的角色。该文通过疏叶骆驼刺母株和子株之间的间隔子切断和给母株补充水分的方法, 研究了母株和子株在各处理下的水势、叶形态和植株生长变化情况。结果表明: (1)间隔子切断后, 疏叶骆驼刺母株和子株正午水势均明显增大(p < 0.01), 说明间隔子切断使得母株和子株水分亏缺值都增大。(2)给母株补水后, 间隔子切断组和间隔子相连组中的母株水势均有明显增加, 同时间隔子相连组的子株水势明显增加(p < 0.01), 而间隔子切断组子株水势没有明显变化(p > 0.05)。(3)间隔子切断组的子株叶片含水率明显低于间隔子相连组子株, 而其株高、冠幅、分枝数和基径的增长量都明显小于间隔子相连组的子株(p < 0.01)。疏叶骆驼刺母株和子株间存在水分整合, 母株会通过根系向子株传输水分。研究成果对塔克拉玛干沙漠南缘的植被恢复以及水资源的合理利用有着重要的意义。     

极端干旱生境下胡杨克隆水分整合特征及其生态意义

克隆水分整合是克隆植物有效利用异质性生境资源的重要对策,也是在恶劣生境下得以定植、存活的重要生存策略。本研究以塔里木河下游的胡杨（Populus euphratica Oliv.）为对象,结合野外调查、稳定同位素技术、生理生态监测实验,调查胡杨的克隆水分整合特征并分析其生态意义。结果显示,胡杨母株与其克隆幼株间存在显著水分生理整合,水分整合以顶向传输为主,水分整合过程与整合水量受到母株生理节律及母株到幼株间隔子长度的影响;克隆幼株因水分整合而能够获取与母株相似的深层土壤水分,比同一区域实生幼株拥有更好的水分获取能力,并使胡杨克隆幼株比实生幼株保持相对更高的水力导度、叶片水分含量及叶水势;在极端干旱生境下的这种水分获取及利用策略使胡杨克隆幼株比实生幼株具有更高的光化学效率与光合性能,有助于克隆幼株在不利生境下定植、存活并保持更高的生存优势。     

毛乌素沙地根茎禾草拂子茅对异质性水分供应的表型可塑性

拂子茅(Calamagrostis epigejos（L.)Roth)为根茎型多年生禾草,具细长根茎。为了探讨拂子茅在异质性水分环境中的表型差异,在内蒙古鄂尔多斯高原的毛乌素沙地对拂子茅由母株、子株组成的分株对给予了高水、低水两种不同的异质性土壤水分处理。实验结果表明：土壤水分状况显著地影响着拂子茅分株的生长表型。在高土壤水分条件下,拂子茅的分株产生的根茎、新生后代分株较多,并使生物量主要分配于地上部分,地上生物量积累多;在低土壤水分条件下,拂子茅分株产生较少的根茎与新生后代分株,并且分配到根系的生物量明显增大。在具有一定对比度的异质性土壤水分环境中,拂子茅分株并不因相连的其他分株所处的土壤水分状况而在根茎生长、新生后代分株的产生和生物量分配等特征上,与同质环境中的具有相同土壤水分状况的分株相比,有明显差异。这些结果揭示：拂子茅仅以分株的形式对异质性水分供应发生表型反应;相连的克隆分株在向顶向和向基向这两个基本方向上,不能对另一分株的土壤水分状况在生K表型上发生反应,它们在水分关系上可能是相互相对独立的。分株的相对独立可能有利于在气候干旱、扰动强烈的沙地环境中实现风险分摊,提高基株的存活几率。     

水分胁迫条件下草莓克隆分株间水分调控及其对光合功能的影响

以盆栽草莓(Fragaria×ananassa)为材料研究了水分胁迫下克隆植物草莓母株和子株间的水分调控机制及其与碳同化、光系统Ⅱ激发能分配的关系.实验材料分为匍匐茎连接和剪断两个大组,进行两步实验.第1步实验,对连接组和剪断组的所有母株控水,子株充分供水;4d后进入第2步实验,把连接组分为两小组,对其中一组充分供水子株开始控水,另一组保持不变.结果表明,土壤干旱引起母株叶片失水,并使其净光合速率和气孔导度显著降低.但是连接组中供水良好的子株能有效缓解缺水母株的水分胁迫.当供水良好的子株也开始受到干旱处理的时候,则会加剧与之相连母株的水分胁迫.受胁迫母株可以通过加强渗透调节能力和降低水势从相连子株获取水分.虽然土壤干旱会造成受胁迫母株叶片脱落酸(abscisic acid, ABA)含量的大幅度增加,但是与之相连子株的叶片ABA含量并没有增加;并且气孔导度与ABA变化趋势一致.(1)草莓母株和子株间的水分运输是由二者的水势差驱动的;(2)ABA不会通过匍匐茎在母株和子株间传递并影响相邻子株气孔导度;(3)在水分异质性较大情况下,生理整合可明显提高克隆系统的碳同化能力和光系统Ⅱ激发能利用效率.     

毛乌素沙地根茎禾草拂子茅对异质性水分供应的表型可塑性

拂子茅(Calamagrostis epigejos(L.)Roth.)为根茎型多年生禾草,具细长根茎.为了探讨拂子茅在异质性水分环境中的表型差异,在内蒙古鄂尔多斯高原的毛乌素沙地对拂子茅由母株、子株组成的分株对给予了高水、低水两种不同的异质性土壤水分处理.实验结果表明:土壤水分状况显著地影响着拂子茅分株的生长表型.在高土壤水分条件下,拂子茅的分株产生的根茎、新生后代分株较多,并使生物量主要分配于地上部分,地上生物量积累多;在低土壤水分条件下,拂子茅分株产生较少的根茎与新生后代分株,并且分配到根系的生物量明显增大.在具有一定对比度的异质性土壤水分环境中,拂子茅分株并不因相连的其他分株所处的土壤水分状况而在根茎生长、新生后代分株的产生和生物量分配等特征上,与同质环境中的具有相同土壤水分状况的分株相比,有明显差异.这些结果揭示:拂子茅仅以分株的形式对异质性水分供应发生表型反应;相连的克隆分株在向顶向和向基向这两个基本方向上,不能对另一分株的土壤水分状况在生长表型上发生反应,它们在水分关系上可能是相互相对独立的.分株的相对独立可能有利于在气候干旱、扰动强烈的沙地环境中实现风险分摊,提高基株的存活几率.     

Hydraulic conductivity and PAT determine hierarchical resource partitioning and ramet development along Fragaria stolons

Co-ordination of metabolic and physiological activity between plant parts is key to the control of growth and development. Here the movement of resources and their allocation between mother plants and daughter ramets along Fragaria stolons was quantified with respect to hierarchy. Gradients of internodal ramet leaf water potential (ψ) and stolon and ramet hydraulic conductivities (L) were measured together with apparent stolon IAA movement via the polar auxin transport pathway (PAT). These processes are linked with measurements of stolon vascular development. The pattern of tissue differentiation and lignification in sequential stele sections of stolons demonstrated the rapid acquisition of the capacity for water transport, with transpiration potentially varying systematically with stolon lignification and the acropetal decline in stolon xylem ψ. Stolon and ramet L declined acropetally, with L across older ramets being significantly lower than that of the connecting stolons. The capacity for polar IAA transport increased with stolon age; this was due to increased transport intensity in older tissues. The partitioning of dry matter was strongly hierarchical with younger ramets smaller than older ramets, while foliar concentrations of N, P, and K were greater for the younger ramets. The results show that stolon anatomy develops rapidly at the apical end, facilitating hierarchical ramet development, which is evident as a basipetal increase in L. The rapid development of transport tissue functionality enables young unrooted ramets to acquire water, in order to supply an expanding leaf area, as well as mineral ions disproportionally with respect to older ramets. This facilitates colonization and self-rooting of apical ramets. The unidirectional increase in basipetal PAT along stolons facilitates hierarchical ramet development.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

To share or not to share: clonal integration in a submerged macrophyte in response to light stress

The ability of clonal plant species to share resources has been studied in many experiments. The submerged macrophyte Potamogeton perfoliatus produces interconnected ramets within short time intervals and hence may or may not share resources with ramets growing in less favourable microhabitats. From a genet point of view, sharing with ramets growing under less favourable conditions might not be an optimal strategy when photosynthates could be used to establish other ramets growing under more favourable conditions. To analyse the plasticity in clonal integration of P. perfoliatus, we set up a factorial aquaria experiment with unshaded or shaded recipient ramets (offspring), which were connected to or separated from donor ramets (parents). Increased biomass production of offspring in parent–offspring systems compared with severed offspring in both light and shade showed that ramets share resources through clonal integration. The relative translocation to the first- and second-offspring generation was influenced by habitat quality: If first-offspring ramets grew in a shaded microhabitat, second-offspring ramets clearly profited. This may be at least partially because of the fact that resources are shifted from first-offspring to second-offspring ramets, indicating controlled senescence of the first-offspring. This complex sharing behaviour might be relevant when plants produce ramets within a dense patch of macrophytes, where support of a shaded ramet might not pay off.     

Clonal integration in Ranunculus reptans: by‐product or adaptation?

We studied fitness consequences of clonal integration in 27 genotypes of the stoloniferous herb Ranunculus reptans in a spatially heterogeneous light environment. We grew 216 pairs of connected ramets (eight per genotype) with mother ramets in light and daughter ramets in shade. In half of the pairs we severed the stolon connection between the two ramets at the beginning of the experiment. During the experiment, 52.7% of the ramet pairs with originally intact connection physically disintegrated. We detected significant variation among genotypes in this regard. Survival of planted ramets was 13.3% higher for originally connected pairs. Moreover, there was significant variation among genotypes in survival, in the difference in survival between plant parts developing from mother and daughter ramets, and in the effect of integration on this difference. In surviving plants connection between ramets decreased size differences between mother and daughter parts. Variation among genotypes was significant in growth and reproduction and marginally significant in the effect of physiological integration on growth and reproduction. Connected daughter ramets had longer leaves and internodes than daughters in severed pairs indicating that integration stimulated plant foraging in both the vertical and the horizontal plane. Observed effects of integration on fitness components in combination with genetic variation in maintenance and effects of connection indicate that clonal integration in R. reptans has the capability to evolve, and therefore suggest that clonal integration is adaptive. If genetic variation in integration is common, future studies on clonal integration should always use defined genetic material and many clones to allow extrapolation of results to population and wider levels.     

Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie

Many clonal plants consist of many connected individual ramets, allowing them to share water and nutrients via physiological integration. Integration among ramets may also improve the ability of clonal plants to tolerate abiotic stress or improve the competitive ability of individual ramets. Here I use a field experiment to determine whether clonal integration improves ramet performance for a widespread clonal tree species invading into native prairie. Aspen (Populus tremuloides) dominates the southern treeline in western Canada, has long-lived belowground connections between mother and daughter ramets, and reproduces vegetatively via resprouting rhizomes after disturbance. I applied two competition treatments (neighbors present or absent) and two clonal integration treatments (belowground rhizomes between mother and daughter ramets either severed or left intact) to 12 replicate Populus daughter ramets at each of three sites. Neighbors improved the survivorship of Populus ramets by 25-35% after 2 yr, but decreased growth by ～20%. Clonal integration tended to improve ramet survival and growth, but these trends were often not significant. Clonal integration did not alter the effects of competition from neighboring vegetation, suggesting that connections between ramets do not necessarily improve the competitive ability of Populus invading into native prairie.     

High levels of inter-ramet water translocation in two rhizomatous Carex species,as quantified by deuterium labelling

We studied water trnaslocation between interconnected mother and daughter ramets in two rhizomatous Carex species, using a newly developed quantitative method based on deuterium tracing. Under homogeneous conditions, in which both ramets were subjected either to wet or dry soil, little water was exchanged between the ramets. When the ramet pair was exposed to a heterogeneous water supply, water translocation became unidirectional and strongly increased to a level at which 30–60% of the water acquired by the wet ramet was exported towards the dry ramet. The quantity of water translocated was unrelated to the difference in water potential between the ramets, but highly correlated to the difference in leaf area. In both species, the transpiration of the entire plant was similar under heterogeneous and homogeneous wet conditions. This was a direct result of an increase in water uptake by the wet ramet in response to the dry conditions experienced by the interconnected ramet. In C. hirta, the costs and benefits of integration in terms of ramet biomass paralleled the responses of water consumption. This species achieved a similar whole-plant biomass in heterogeneous and homogeneous wet treatments, and water translocation was equally effective in the acropetal and basipetal directions. In C. flacca, responses of biomass and water consumption did not match and, under some conditions, water translocation imposed costs rather than benefits to the plants of this species. It is concluded that enhanced resource acquisition by donor ramets may be of critical importance for the net benefits of physiological integration in clonal plants.     

异质水分环境下野牛草相连分株间光合同化物的生理整合及其调控

异质环境下,克隆植物通过生理整合机制使资源在分株间实现共享,提高了其对异质性环境的适应能力,具有重要的生态进化意义,研究生理整合机制及其调控机理可为进一步发掘克隆植物应用潜力提供理论依据。以野牛草3个相连分株为材料,对其中一个分株用30%聚乙二醇6000(PEG-6000)模拟水分胁迫,通过Hoagland营养液培养试验,研究了异质水分环境下光合同化物在野牛草相连分株间的生理整合及分株叶片与根系内源激素ABA与IAA含量的变化规律。结果表明,14C-光合同化物在克隆片断内存在双向运输,但以向顶运输为主,异质水分环境下,受胁迫分株光合同化物的输出率明显降低,而与其相邻分株合成的光合同化物向受胁迫分株方向运输率明显增加;异质水分环境下,各分株ABA含量均明显增加,但以受胁迫的分株叶片及根系ABA的含量增加幅度最大,各分株IAA含量较对照均显著下降(P0.05),且以受胁迫分株IAA含量下降幅度最大;各分株叶片与根系ABA/IAA均显著提高(P0.05),相邻分株ABA/IAA增加幅度低于受胁迫分株。异质水分环境影响野牛草克隆分株间光合同化物的生理整合,且ABA与IAA在分株间光合同化物运输与分配过程中具有重要的调节作用。     

Effects of inoculation with native arbuscular mycorrhizal fungi on clonal growth of Potentilla reptans and Fragaria moschata (Rosaceae)

Many clonal plants live in symbiosis with ubiquitous arbuscular mycorrhizal (AM) fungi, however, little is known about their interaction with respect to clonal reproduction and resource acquisition. The effects of arbuscular mycorrhiza on the growth and intraclonal integration between ramets of two stoloniferous species were studied experimentally in a nutritionally homogenous soil environment. Two species coexisting at the same field site, Potentilla reptans and Fragaria moschata, were selected as model plants for the study. Pairs of their ramets were grown in neighbouring pots with each ramet rooted separately. Four inoculation treatments were established: (1) both mother and daughter ramets remained non-inoculated, (2) both ramets were inoculated with a mixture of three native AM fungi from the site of plant origin, (3) only mother or (4) daughter ramet was inoculated. The stolons connecting the ramets were either left intact or were disrupted. Despite the consistent increase in phosphorus concentrations in inoculated plants, a negative growth response of both plant species to inoculation with AM fungi was observed and inoculated ramets produced fewer stolons and fewer offspring ramets and had lower total shoot dry weights as compared to non-inoculated ones. A difference in the extent of the negative mycorrhizal growth response was recorded between mother and daughter ramets of P. reptans, with daughter ramets being more susceptible. Due to AM effect on ramet performance, and thereby on the source-sink relationship, inoculation also significantly influenced biomass allocation within clonal fragments. Physiological integration between mother and daughter ramets was observed when their root systems were heterogeneous in terms of AM colonization. These results hence indicate the potential of mycorrhizal fungi to impact clonal growth traits of stoloniferous plant species, with possible consequences for their population dynamics.     

Differential influence of clonal integration on morphological and growth responses to light in two invasive herbs

Background and aims

In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions.

Methods

In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85% shade and their morphological and growth responses were assessed.

Key results

The influence of clonal integration on the light reaction norm (connection×light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection×light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mother plants invest in daughter ramets prior to their own branches.

Conclusions

Clonal integration modifies light reaction norms of morphological and growth traits in a species-specific manner for A. philoxeroides and P. canescens, but it improves the establishment of juvenile ramets of both species in light-limiting environments by promoting their growth during early development. This factor may be partially responsible for their ability to successfully colonize native plant communities.     

青藏高原东缘过路黄在资源交互斑块性生境中的克隆内资源共享

在青藏高原和四川盆地过渡带,分别于618m和1800m两个海拔高度上研究匍匐茎克隆植物过路黄(Lysimachia christinae)在资源交互斑块性生境中的克隆内资源共享及其对生长的影响.结果显示, 在海拔1800m处,与资源的空间同质性处理(Ⅰ) 和(Ⅱ)相比, 资源的空间异质性处理(Ⅲ)和(Ⅳ)下过路黄整个克隆片段的生物量和分株数均获得显著增加;在海拔618m处,与资源的空间同质性处理(Ⅰ) 和(Ⅱ)相比,资源的空间异质性处理(Ⅲ)和(Ⅳ)下过路黄整个克隆片段生物量显著增加.在海拔618m和1800m处,生长在低光高养条件下的远端分株, 若与高光低养的近端分株相连, 相比连接到低光高养的近端分株, 它们分配更多的生物量到地下部分;在海拔1800m处,生长在高光低养条件下的远端分株, 若与低光高养的近端分株相连, 相比连接到高光低养的近端分株, 它们分配更多的生物量到地上部分.在海拔618m和1800m处,生长在高光低养条件下的近端分株, 若与低光高养的远端分株相连, 相比连接到高光低养的远端分株, 它们分配更多的生物量到地上部分.处于资源交互斑块性生境中的过路黄发生了克隆内分工,依靠相连分株间的功能分化, 克隆植物能有效的利用异质性分布的资源, 缓解资源交互斑块性分布对克隆植物生长的不利影响.通过间隔子(匍匐茎或根状茎),相连分株间能够相互传递和共享由不同分株获得的资源,这种资源共享能够提高克隆植物在异质性生境中的存活与生长.同时,方差分析显示环境异质性和海拔的交互作用显著影响克隆片段的生物量和分株数.相比于海拔618m,在海拔1800m处克隆内资源共享对克隆植物生长表现的影响更大.     

Strawberry plant relationship through the stolon

Stolon is an elongated, two-node, vegetative, axillary shoot, which supports the ramet (rooted rosette) until it is completely independent on its own roots. The reciprocal capacity of the ramets, in a single runner chain, to sustain the growth and share locally abundant resources or to tolerate a local stress, is still in debate. This capacity may play an important role for improving nursery plant production and for better understanding the natural clonal multiplication. To describe strawberry stolon action, in plant-to-plant relationship, bare-rooted Camarosa ramets, joint in couples by their own stolons (generally, second and third ramet in a runner chain) were transplanted in two pots. The couples of ramets were treated in a factorial experiment with decortication (peeling a 2-mm ring of bark from the stolon), removal of root system or glyphosate application to one of the two ramets. In the studied system, the older ramet was referred as mother and the other as daughter. The two ramets were very similar in age and seem to act with a very limited hierarchic prevalence of the mother. When the root system of one ramet was eliminated, leaf number and chlorophyll content had a very slight decrease, independently in the mother ramet or in the daughter. The decortication did not reduce water integration, in any group of plants, but limited assimilate allocation towards the daughter ramet when the mother ramet had a severe root cut (not vice versa). The glyphosate action resulted localized in the sprayed ramet, which reduced chlorophyll content within 2 days and expired after 4 days.     

WATER SHARING AMONG RAMETS IN A DESERT POPULATION OF DISTICHLIS SPICATA (POACEAE)

Clonal growth enables plants to transport resources among separately rooted but connected ramets, a potential advantage in patchy or unpredictable habitats. Nevertheless, clonal plants are relatively scarce in deserts. To test whether clonal integration of water relations can increase plant performance under natural conditions in a desert species, water movement was traced and connection among ramets was manipulated in the rhizomatous grass Distichlis spicata in Death Valley, California. To examine potential costs of clonal growth form, connections were mapped and analyzed for dry mass and nitrogen content. Movement of dye showed potential transport of water among five ramets up to 1.4 m apart. Severance of connecting rhizomes increased mortality and decreased water potential of individual ramets within 36 hr, indicating that water sharing among ramets could be of significant benefit. However, plants had a high investment of mass and nitrogen in underground organs, which might be a cost of clonal growth associated with desert environments.