莲草直胸跳甲取食对空心莲子草和莲子草克隆整合的影响

很多外来入侵植物都具有克隆生长习性,探究克隆整合特性与外来克隆植物入侵性间的关系对阐明其生态适应性及入侵机制具有重要的意义。本研究以入侵植物空心莲子草及其本地同属种莲子草为研究对象,比较在生防昆虫莲草直胸跳甲的取食下,克隆整合对两种植物先端分株、基端分株及整个克隆片段生长和生物量分配的影响。结果表明: 在莲草直胸跳甲取食下,有克隆整合的空心莲子草先端分株的叶片数、茎长、分株数及整个克隆片段的地径均显著高于无克隆整合植株,其基端分株及整个克隆片段的地下生物量和总生物量相较于无克隆整合植株分别降低了78.2%、60.9%和48.7%、37.2%;有克隆整合的莲子草先端分株的地径及整个克隆片段的叶片数与无克隆整合植株相比显著增加,其基端分株数显著降低了21.7%,而其先端分株、基端分株及整个克隆片段的生物量均无显著差异。耗益分析表明,在莲草直胸跳甲取食下,空心莲子草先端分株的分株数与生物量及莲子草先端分株的分株数均能通过克隆整合显著受益,而两种植物基端的分株数、生物量的耗益则不受克隆整合处理的影响。这些结果表明,克隆整合虽能在一定程度缓解莲草直胸跳甲对于两种植物先端分株的取食压力,且空心莲子草的克隆整合作用要强于莲子草,但在整个克隆片段水平上,两种植物并不能通过克隆整合显著获益。     

Clonal growth of sagebrush (Artemisia tridentata) (Asteraceae) and its relationship to volatile communication

To increase systemic resistance to herbivory, some clonal plants transmit internal cues among clonal ramets to prevent widespread damage to genets. Sagebrush (Artemisia tridentata) (Asteraceae) is known to use volatile cues to induce resistance within and between plants (so‐called volatile communication). In the present study, we observed the extent and frequency of clonal growth in a natural sagebrush population in western North America to understand the influence of clonal growth on volatile communication. We used genetic analysis involving microsatellite markers and excavation of the root systems. In addition, we characterized the volatile profiles from the headspace of sagebrush ramets. Excavation of the root system of sagebrush plants revealed that sagebrush propagates clonally below ground and that daughter ramets grow near the mother stem. Volatiles were variable among genetically different ramets, although clonal ramets (genetically identical ramets) released similar volatiles, suggesting a genetic basis for volatile similarity. Sagebrush has been shown to be most responsive to volatiles released from artificially produced clones and suffers less herbivore damage as a result. Therefore, these results, taken into consideration together, imply that volatile communication may occur among genetically identical ramets under natural conditions, and that volatile similarity between the releaser and receiver may be recognized by the receiver and increase resistance against herbivory.     

Investigating the community consequences of competition among clonal plants

Although clonal plants comprise most of the biomass of several widespread ecosystems, including many grasslands, wetlands, and tundra, our understanding of the effects of clonal attributes on community patterns and processes is weak. Here we present the conceptual basis for experiments focused on manipulating clonal attributes in a community context to determine how clonal characteristics affect interactions among plants at both the individual and community levels. All treatments are replicated at low and high density in a community density series to compare plant responses in environments of different competitive intensity. We examine clonal integration, the sharing of resources among ramets, by severing ramets from one another and comparing their response to ramets with intact connections. Ramet aggregation, the spacing of ramets relative to each other, is investigated by comparing species that differ in their natural aggregation (either clumped growth forms, with ramets tightly packed together, or runner growth forms, with ramets loosely spread) and by planting individual ramets of all species evenly spaced throughout a mesocosm. We illustrate how to test predictions to examine the influence of these two clonal traits on competitive interactions at the individual and community levels. To evaluate the effect of clonal integration on competition, we test two predictions: at the individual level, species with greater clonal integration will be better individual-level competitors, and at the community level, competition will cause a greater change in community composition when ramets are integrated (connected) than when they are not. For aggregation we test at the individual level: clumped growth forms are better competitors than runner growth forms because of their ability to resist invasion, and at the community level: competition will have a greater effect on community structure when ramets are evenly planted. An additional prediction connects the individual- and community-level effects of competition: resistance ability better predicts the effects of competition on relative abundance in a community than does invasion ability. We discuss additional experimental design considerations as revealed by our ongoing studies. Examining how clonal attributes affect both the individual- and community-level effects of competition requires new methods and metrics such as those presented here, and is vital to understanding the role of clonality in community structure of many ecosystems.     

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.     

Herbivory and ramet performance in the clonal herb Trientalis europaea L.

1 The intensity of leaf damage caused by invertebrate herbivores and grazing by vertebrates and their effects on clonal growth, survival and reproduction were examined in a population of the forest herb Trientalis europaea during 4 years.
2 Levels of herbivory were low and varied between years in the studied population. Larger ramets were exposed to a greater risk of both grazing and leaf defoliation. Ramets that suffered leaf damage in 1 year experienced an increased probability of leaf damage in succeeding years. The probability of suffering herbivore attack was independent among the ramets belonging to the same clonal fragment, suggesting that clonal propagation might operate as a risk-spreading strategy in this species.
3 Leaf damage did not affect any measure of plant performance, probably due to the low amounts of leaf area removed by invertebrate herbivores. In contrast, vertebrate grazing affected all phases of the pseudo-annual life cycle of T. europaea . Grazing prevented flowering and fruiting, increased ramet mortality during summer and decreased tuber production. Furthermore, grazed ramets produced shorter stolons and smaller tubers, which in turn had a lower winter survival and produced smaller ramets in the following growing season. The large impact of grazing was due to the consumption of the whole of the single shoot of ramets of T. europaea . Although regrowth was possible, secondary shoots were significantly smaller and assimilation was delayed.
4 Tubers originating from grazed ramets were placed shallower than tubers from ungrazed ramets. This could be due to an alteration in the growth pattern of stolons of the grazed ramets.     

Interspecific variation in clonality in temperate lianas revealed by genetic analysis: Do clonal proliferation processes differ among lianas?

Lianas are important components of forest communities with a considerable impact on overall forest structure and function. Lianas are characterized by extensive clonal reproduction on the forest floor, which is important for their establishment and growth. Previous studies have suggested that clonal reproduction strategies vary substantially among liana species; however, few studies have quantitatively evaluated the clonality strategy in multiple co-occurring liana species. The primary objective of the present study was to evaluate the relative contribution of clonal reproduction and to understand the clonal proliferation processes in co-occurring liana species by assessing both small stems on the forest floor and mature stems that climbed trees. The clonal reproduction strategy in four common liana species (stem twiner: Wisteria floribunda; root climbers: Schizophragma hydrangeoides, Euonymus fortunei and Rhus ambigua) in a 6-ha plot and a belt transect within an old-growth temperate forest in central Japan was evaluated using genetic analysis. The contribution of clonal reproduction was smaller in root climbers than in W. floribunda. All W. floribunda genets with small ramets in the understory had genetically identical ramets that climbed trees, whereas few such ramets were found in root climbers. This indicates that W. floribunda mature ramets laterally produce small ramets via clonal proliferation, whereas seedlings of root climbers grow horizontally to climb trees. The results indicate that the clonal reproduction processes differ greatly among lianas and the clonal growth in lianas plays a contrasting role in their life-history strategy.     

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     

Combined effects of resource heterogeneity and simulated herbivory on plasticity of clonal integration in a rhizomatous perennial herb

Previous lines of investigation assuming potential advantage of clonal integration generally have neglected its plasticity in complex heterogeneous environments. Clonal plants adaptively respond to abiotic heterogeneity (patchy resource distribution) and herbivory‐induced heterogeneity (within‐clone heterogeneity in ramet performance), but to date little is known about how resource heterogeneity and simulated herbivory jointly affect the overall performance of clones. Partial damage within a clone caused by herbivory might create herbivory‐induced heterogeneity in a resource‐homogeneous environment, and might also decrease or increase the extent of heterogeneity under resource‐heterogeneous conditions. We conducted a greenhouse experiment in which target‐ramets of Leymus chinensis segments within homogeneous or heterogeneous nutrient treatments were subject to clipping (0% or 75% shoot removal). In homogeneous environments with high (9:9) nutrient availability, ramet biomass of L. chinensis with intact or severed rhizomes is 0.70 or 0.69 g. Conversely, target‐ramet biomass with intact rhizomes is obviously lower than that of the severed target‐ramets in the homogeneous environments with medium (5:5) and low (1:1) nutrient availability. High resource availability and the presence of herbivory can alleviate negative effects of rhizome connection under homogeneous conditions, by providing copious resource or creating herbivory‐induced heterogeneity respectively. Herbivory tolerance of clonal fragments with connected rhizomes was higher than that of fragments with severed rhizomes under heterogeneous conditions. These findings confirmed the unconditional advantage of clonal integration on reproduction under the combined influence of resource heterogeneity and simulated herbivory. Moreover, our results made clear the synergistically interactive effects of resource heterogeneity and simulated herbivory on costs and benefits of clonal integration. This will undoubtedly advance our understanding on the plasticity of clonal integration under complex environmental conditions.     

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.     

酸雨和采食模拟胁迫下克隆整合对空心莲子草生长的影响

研究表明克隆整合可以显著提升异质环境中克隆植物的生长,然而当克隆植物遭受均质环境压力时,整合对植物生长影响的研究相对较少。本文以典型入侵克隆植物空心莲子草（Alternanthera philoxeroides）为例,研究均质环境压力酸雨和采食模拟胁迫对空心莲子草生长的影响,以及克隆整合在空心莲子草适应不利环境过程中所起的作用。酸雨设3种浓度梯度：pH值3.5 、pH值4.5和 pH值6.5（对照）;采食设3种水平：不去叶、去叶50%和去叶90%;整合水平：匍匐茎切断和连接。结果表明：无论保持或切断匍匐茎的连接,酸雨处理都不影响空心莲子草生物量。当保持匍匐茎连接时,pH值4.5酸雨处理增加了空心莲子草匍匐茎长度和分株数目,因此,低度酸雨可能对空心莲子草生长有一定的促进作用。同样,无论匍匐茎是否被切断,采食处理都显著降低了空心莲子草克隆片段生物量,而显著增加了叶片数目。当切断匍匐茎连接时,采食处理使空心莲子草分株数目显著增加。本文得出的结论是：空心莲子草能较好地适应酸雨和采食的环境压力,当空心莲子草全部克隆分株遭受均质环境胁迫时,克隆整合并不能显著改善它的生长。     

Effects of resource sharing directionality on physiologically integrated clones of the invasive Carpobrotus edulis

资源共享的方向性对入侵植物海榕菜生理整合的影响 与植物克隆生长相关的一个关键性状是生理整合能力,它允许在一个克隆内相连接的分株之间共享资源。资源传输遵循源-汇关系：从生长在资源丰富斑块上的分株到生长在贫瘠斑块上的分株。然而,一些实验结果表明,向顶运输(从后端到前端)通常超过基部运输(从前端到后端)。在本研究中, 我们旨在确定入侵植物海榕菜(Carpobrotus edulis)生理整合模块的资源运输方向。我们开展了两个模拟实 验,研究了在不同位置(后端、中部、前端)的不同营养水平对海榕菜连接和断开克隆系统的影响。比较了分株生物量分配模式和最终生物量,以阐明生理整合的作用是否受资源运输方向性的影响。研究结果表明,海榕菜资源的向顶运输是普遍存在的,其发育分工是后端分株专注于获取土壤资源,前端分株专注于地上生长。虽然在最严酷的生长条件下的前端分株所获得的效益最高,但这种生物量分配模式不受后端或前端分株生长的营养条件的影响。这种发育程序化的分工被认为可以促进海榕菜的横向生长,从而对这种入侵物种的扩展具有重要意义。     

资源异质性环境中三种匍匐茎草本植物克隆内资源共享和分株功能特化

研究了3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜 (Potentilla reptans L. var. sericophylla Franch.)、鹅绒委陵菜 (P. anserina L.) 和金戴戴 (Halerpestes ruthenica (Jacq.) Qvcz.) 对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策.当生长于高光照低养分条件下分株 (HL分株) 与生长于低光照高养分条件下分株 (LH分株) 之间的匍匐茎连接时, 3种克隆植物HL分株、LH分株以及整个分株对系统 (HL分株 + LH分株) 的生物量均得到显著提高.同时, LH分株根冠比显著增加, 而HL分株根冠比显著下降.这表明, 当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时, 3种植物克隆分株均发生了环境诱导的功能特化.克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取, 从而能够缓解资源交互斑块性生境对克隆植物的不利影响.     

资源异质性环境中三种匍匐茎草本植物克隆内资源共享和分株功能特化(英文)

研究了 3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜 (PotentillareptansL .var.sericophyllaFranch .)、鹅绒委陵菜 (P .anserinaL .)和金戴戴 (Halerpestesruthenica (Jacq .)Qvcz .)对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策。当生长于高光照低养分条件下分株(HL分株 )与生长于低光照高养分条件下分株 (LH分株 )之间的匍匐茎连接时 ,3种克隆植物HL分株、LH分株以及整个分株对系统 (HL分株 LH分株 )的生物量均得到显著提高。同时 ,LH分株根冠比显著增加 ,而HL分株根冠比显著下降。这表明 ,当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时 ,3种植物克隆分株均发生了环境诱导的功能特化。克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取 ,从而能够缓解资源交互斑块性生境对克隆植物的不利影响     

Effects of clonal fragmentation on intraspecific competition of a stoloniferous floating plant

Disturbance is common and can fragment clones of plants. Clonal fragmentation may affect the density and growth of ramets so that it could alter intraspecific competition. To test this hypothesis, we grew one (low density), five (medium density) or nine (high density) parent ramets of the floating invasive plant Pistia stratiotes in buckets, and newly produced offspring ramets were either severed (with fragmentation) or remained connected to parent ramets (no fragmentation). Increasing density reduced biomass of the whole clone (i.e. parent ramet plus its offspring ramets), showing intense intraspecific competition. Fragmentation decreased biomass of offspring ramets, but increased biomass of parent ramets and the whole clone, suggesting significant resource translocation from parent to offspring ramets when clones were not fragmented. There was no interaction effect of density x fragmentation on biomass of the whole clone, and fragmentation did not affect competition intensity index. We conclude that clonal fragmentation does not alter intraspecific competition between clones of P. stratiotes, but increases biomass production of the whole clone. Thus, fragmentation may contribute to its interspecific competitive ability and invasiveness, and intentional fragmentation should not be recommended as a measure to stop the rapid growth of this invasive species.     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Life-history monographs of Japanese plants 14: Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae)

The life-history characteristics of Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae) are described. The species is an herbaceous perennial that favors open but relatively moist habitats. It is distributed from Kyushu to Hokkaido in Japan but also occurs in Korea, Mongolia, China and the Russian Far East. In southwestern Japan, shoots start sprouting from mid- to late April, reaching approximately 30–70 cm in height, with 5–10 compound leaves. Ramets simultaneously produce one or more stoloniferous rhizomes that elongate until new ramets are formed at the tips. Cardamine leucantha has a pseudo-annual life cycle, in which mother ramets wither at the end of each season and only daughter ramets appear aboveground in the next year. As a result, ramet positions change annually. In a study population, the number of flowers averaged 23.9 ± 21.0 per ramet and fruit set was 44.2 ± 24.8% (10.4 ± 10.1 fruits per ramet). Ramets produced 3.8 ± 2.3 rhizomes that were 22.0 ± 15.6 cm long. The species sometimes forms large populations. A single genet develops into a group of disconnected ramets spreading via clonal growth. Reproductive characteristics (e.g., fruit set and numbers of flowers and rhizomes) vary among populations, resulting in interpopulation differences in genet structure. Because the reference genome became available recently, established molecular tools will be applied effectively for the investigations of C. leucantha as a model clonal plant.     

RAMET DYNAMICS FOR THE CLONAL SEAWEED PTEROCLADIELLA CAPILLACEA (RHODOPHYTA): A COMPARISON WITH CHONDRUS CRISPUS AND WITH MAZZAELLA CORNUCOPIAE (GIGARTINALES)

Little is known about the dynamics and the ecological interactions among ramets (fronds) from populations of clonal red seaweeds. Small ramets are very difficult to tag, so their growth cannot be monitored directly. The temporal variation of the relationship between stand biomass and ramet density offers information on ramet performance. We calculated this relationship for an intertidal population of Pterocladiella capillacea (Gmelin) Santelices et Hommersand (Gelidiales) from Baja California, Mexico. Biomass and density were positively correlated on an annual basis, indicating that biomass accumulated without involving self-thinning among ramets. This contrasts with nonclonal seaweeds, for which self-thinning among individuals occurs during growth, but agrees with other clonal red seaweeds, such as Chondrus crispus Stackhouse and Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (both Gigartinales). The growth pattern for these members of the Gelidiales and of the Gigartinales holds despite differences in holdfast morphology and ramet branching degree and despite differences in the capacity of coalescence during early stages, known only for the Gigartinales. The positive slope for the dynamic biomass–density relationship, on a bilogarithmic scale, was statistically steeper for M. cornucopiae than for P. capillacea and for C. crispus. This suggests that the addition of new ramets during the growth season may be relatively more beneficial for biomass accumulation rates for M. cornucopiae. This would be expected for high-intertidal species subjected to strong abiotic stress, for which ramet crowding constitutes a key protection. Pterocladiella capillacea occurs at the mid-intertidal zone and C. crispus at the subtidal zone, so ramets would be relatively less important in that respect.     

Shifting effects of physiological integration on performance of a clonal plant during submergence and de-submergence

Background and Aims

Submergence and de-submergence are common phenomena encountered by riparian plants due to water level fluctuations, but little is known about the role of physiological integration in clonal plants (resource sharing between interconnected ramets) in their adaptation to such events. Using Alternanthera philoxeroides (alligator weed) as an example, this study tested the hypotheses that physiological integration will improve growth and photosynthetic capacity of submerged ramets during submergence and will promote their recovery following de-submergence.

Methods

Connected clones of A. philoxeroides, each consisting of two ramet systems and a stolon internode connecting them, were grown under control (both ramet systems untreated), half-submerged (one ramet system submerged and the other not submerged), fully submerged (both ramet systems submerged), half-shaded (one ramet system shaded and the other not shaded) and full-shaded (both ramet systems shaded) conditions for 30 d and then de-submerged/de-shaded for 20 d. The submerged plants were also shaded to very low light intensities, mimicking typical conditions in turbid floodwater.

Key Results

After 30 d of submergence, connections between submerged and non-submerged ramets significantly increased growth and carbohydrate accumulation of the submerged ramets, but decreased the growth of the non-submerged ramets. After 20 d of de-submergence, connections did not significantly affect the growth of either de-submerged or non-submerged ramets, but de-submerged ramets had high soluble sugar concentrations, suggesting high metabolic activities. The shift from significant effects of integration on both submerged and non-submerged ramets during the submergence period to little effect during the de-submergence period was due to the quick recovery of growth and photosynthesis. The effects of physiological integration were not found to be any stronger under submergence/de-submergence than under shading/de-shading.

Conclusions

The results indicate that it is not just the beneficial effects of physiological integration that are crucial to the survival of riparian clonal plants during periods of submergence, but also the ability to recover growth and photosynthesis rapidly after de-submergence, which thus allows them to spread.     

Relative costs and benefits of clonal integration of Zoysia japonica under various N:P ratios

Clonal plants adaptively respond to abiotic stress, but to date little is known about under what circumstanses clonal integration is beneficial, or costly. To study the costs and benefits of clonal integration on clonal growth, we cultivated Zoysia japonica under three ratios of N:P (N:P ≈ 7, 14:1 and 21:1), and four types of stolon severing treatments (connected stolon CS, light severing LS, moderate severing MS, serious severing SS). The results showed that Z. japonica performed best at a low ratio of N:P (N:P ≈ 7:1). When the stolons were connected (CS), the growth of primary A‐ramets, multiple‐nodes and stolons benefited from clonal integration; however, the growth of primary B‐ramets on the primary stolons, A and B ramets was significantly reduced (p < 0.05). In the moderate stolon severing treatment MS_7:1 (N:P ≈ 7:1), clonal integration appeared more cost effective than in all other treatments. On the whole, with increasing stolon severing intensity, the cost of clonal integration increased and the clonal growth of Z. japonica declined significantly. The pattern of biomass allocation may be useful for Z. japonica to adapt to the various environments, and clonal integration plays a significant role under adverse environmental conditions.     

Ramet performance in two tussock plants — Do the tussock-level parameters matter?

Studies in plant demography are primarily done at the level of ramets and typically collect ramet-related parameters such as ramet size, type and history. This approach ignores possible effects of factors associated with higher levels, such as genet or tussock. This is particularly important in perennial resprouting herbs with persistent root that consists of many ramets as interaction between ramets, both by competition and by resource sharing, are likely to be intense in these plants. This study investigates effects of tussock-level parameters (age, size and ramet position within tussock and ramet density) on performance of individual ramets in two tussock-forming resprouting herbs (Tanacetum vulgare and Centaurea jacea).The results show that position of a ramet within tussock did not affect ramet growth, but had significant effects on flowering and survival in both species. The direction of the effect differed between the two species; marginal ramets were more successful in T. vulgare, while central ramets were more successful in C. jacea. In addition, tussock age had a significant effect on ramet flowering and survival in T. vulgare. Both these effects are likely to be due to the more competitive life form of T. vulgare, which is restricted to temporary habitats with intense competition. C. jacea is a species of mown or grazed grasslands with lower productivity, which are more stable and where competition is weaker. The effects of the number of neighboring ramets and of the tussock size were significant, but often locality-specific.The results indicate that although many important effects of tussock-related parameters exist, their direction and magnitude differ between species or even populations and are thus not easily predictable. Neglecting these effects, however, is likely to make the demographic models weaker.