异质性重金属镉胁迫下克隆整合对匍匐茎草本植物积雪草生长的影响

 采用盆栽试验研究了异质性重金属镉胁迫下, 克隆整合对匍匐茎草本植物积雪草(Centella asiatica)生长的影响。将远端分株(相对年幼的分株)分别置于对照和镉胁迫处理下, 并对远端分株与近端分株(相对年长的分株)之间的匍匐茎进行切断或保持连接处理。研究结果显示: 镉胁迫处理显著降低了积雪草远端分株的净光合速率(P_n)、最大光量子产量(F_v/F_m)、叶绿素含量、叶面积、分株数和生物量; 克隆整合缓解了镉胁迫对远端分株生长的不利影响; 克隆整合不仅未导致相连近端分株的损耗, 而且相连近端分株的光合效率也没有表现出补偿性增加; 克隆整合降低了远端受胁迫分株的根冠比, 从而使之减少了对土壤中重金属镉的吸收; 匍匐茎切断和镉胁迫处理对近端分株、远端分株的叶柄长没有显著的影响。结果表明: 克隆整合提高了积雪草遭受镉胁迫的远端分株的生长, 改变了其生物量分配格局, 并有助于整个克隆片段在异质性重金属胁迫下的生长。该研究对于丰富和发展异质性环境胁迫下克隆整合的生态适应对策具有重要意义。     

异质性光照下匍匐茎草本狗牙根克隆整合的耗益

在一盆栽实验中,将匍匐茎草本植物狗牙根克隆分株对进行异质性光照处理,对远端分株(幼年)分别进行不同程度的遮荫,并对分株对之间的匍匐茎进行保持连接或是切断处理。结果显示:在中度遮荫时,切断匍匐茎连接显著降低了远端分株的生物量和净光合速率(Pn);在重度遮荫时,切断匍匐茎连接显著降低了远端分株的分株数、叶片数、生物量、净光合速率(Pn)、最大光量子产量(Fv/Fm)和实际光量子产量(ФPSⅡ),克隆整合缓解了遮荫胁迫对远端分株生长的影响。在中度遮荫时,保持匍匐茎连接显著提高了近端分株的净光合速率(Pn);在重度遮荫时,保持匍匐茎连接显著提高了近端分株的最大光量子产量(Fv/Fm)、实际光量子产量(ФPSⅡ)和净光合速率(Pn),克隆整合使得相连未受胁迫近端分株的光合效率表现出补偿性增加。相比于匍匐茎切断处理,当远端分株遭受重度遮荫胁迫时,克隆整合引起近端未受胁迫分株生物量显著降低,而整个克隆片段生物量没有显著变化。因此,当远端分株遭受重度遮荫胁迫时,匍匐茎草本狗牙根可能采取一种风险分摊策略以降低基株死亡风险,这样一种策略对于维持克隆植物在异质性生境中基株适合度具有重要意义。     

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。     

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。     

两种践踏胁迫下克隆整合对入侵植物空心莲子草生长的影响

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

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.     

Effects of clonal integration,nutrients and cadmium on growth of the aquatic macrophyte Pistia stratiotes

许多湿地同时遭受养分和有毒金属的污染,其植被多数为克隆植物。我们假设,富养化和克隆整合可以通过增加植物的生长来提高对有毒金属污染的植物修复能力,即使在毒性胁迫环境下也是如此。为验证此假说,我们将常见、广布的匍匐茎漂浮植物大薸(Pistia stratiotes L.)的单个分株种植在两种不同的养分条件下和两种镉污染处理(无镉或含0.6 mg L⁻¹ 的镉)中42天。通过保持或切断母株与其后代分株之间的连接来维持或阻止克隆整合,并通过保留或移除切断后的后代分株来控制克隆内竞争的有无。由于高养分条件下克隆后代分株的数量增加了一倍,因此镉处理下大薸的生物量在高养分条件下几乎是在低养分条件下的两倍。切断匍匐茎连接对整个克隆(母株和后代分株的总和)的生物量没有影响。切断连接后去除后代分株对镉污染下母株的生物量没有显著的影响,但却显著增加了无镉污染下母株的生物量。这些研究结果支持富养化可以提高水生植物对有毒金属污染的修复能力这一假说,但并不支持克隆整合有利于植物修复的假说。因此,水生植物(如大薸)可能有助于对同时遭受有毒金属和养分污染的湿地进行修复,但克隆片段化对植物的这种修复能力可能没有显著影响。     

资源交互斑块性生境中两种不同分枝型匍匐茎植物的克隆内分工

以青藏高原和四川盆地过渡带两种不同分枝型匍匐茎植物野草莓 (Fragaria vesca)和过路黄 (Lysimachia christinae)为对象 ,研究它们在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中的克隆内分工。结果显示 ,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)中野草莓和过落黄的近端、远端和整个克隆片段的生物量和分株数均获得显著增加。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 ;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 ;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。实验结果表明 ,资源交互斑块性生境中野草莓和过路黄均发生了克隆内分工。通过克隆内分工 ,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     

不同土壤养分条件下克隆整合对薇甘菊忍耐埋藏能力的影响

克隆植物可通过克隆整合较好地适应逆境胁迫,克隆整合对克隆植物忍耐逆境胁迫能力的促进作用可能随环境条件而异。本文以海南恶性入侵植物薇甘菊为对象,通过温室盆栽实验,研究不同土壤养分条件下(低或高)克隆整合对薇甘菊匍匐茎克隆片段(具2个节)忍耐部分分株被埋藏(仅远端分株被埋藏)能力的影响。结果表明:当远端分株被埋藏3 cm时,无论有无克隆整合,其均有较高的出苗率;当远端分株被埋藏入土壤6 cm时,克隆整合显著提高了其出苗率(从不足17%提高到67%),且这些结果不受土壤养分条件的影响。对于近远端均出苗的克隆片段,埋藏深度、是否保持克隆连接对克隆片段的总生物量未见显著影响,尽管近端或远端分株生物量可能受到影响。即便是对远端分株埋藏6cm的处理,克隆整合也可使克隆片段的总生物量维持在和远端分株没有被埋藏时克隆片段总生物量相当的水平,且这些结果也不受土壤养分条件的影响。因此,克隆整合是薇甘菊对局部埋藏胁迫的生态适应对策之一,它能够提高薇甘菊被埋藏分株的存活,提高薇甘菊在局部埋藏胁迫下的适合度。不同土壤养分条件下,克隆整合对出苗率和生物量的影响差异不大,可能是因为不同土壤养分条件下,克隆分株间的资源输送并没有差别,或埋藏深度(6 cm)不足以使植物生长表现出差别。     

异质性光照条件下匍匐茎草本野草莓的克隆整合

通过盆栽实验,研究了匍匐茎草本野草莓在异质性光照条件下的克隆整合。结果显示克隆整合显著增强了野草莓胁迫分株段的生长,损—益分析表明未受胁迫分株没有显著损耗,整个克隆片段的生长得到显著提高。在局部遮荫处理,克隆整合对克隆形态可塑性的修饰作用没有观察到。最后,讨论了克隆植物对环境的生态适应意义。     

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.     

Effects of soil heterogeneity and clonal integration on Alternanthera philoxeroides populations with a radial ramet aggregation

Some clonal plants can spread their ramet populations radially, and soil heterogeneity and clonal integration may greatly affect the establishment of these types of populations. We constructed Alternanthera philoxeroides populations with a radial ramet aggregation, allowing old ramets of clonal fragments to concentrate in central pots and younger ramets to root in peripheral pots. The peripheral pots were supplemented either with three different levels (high, medium and low) of soil nutrients to simulate a heterogeneous soil environment, or only one medium level of soil nutrients to simulate a homogeneous environment. Stolon connections between the central older ramets and the peripheral younger ramets were left intact or severed to test the effect of clonal integration. The maintenance of stolon connection could induce the division of labor between different‐aged ramets, by increasing the root investment in central ramets and the above‐ground growth in peripheral ramets. The maintenance of stolon connection could improve the growth of the central and peripheral ramets, clonal fragments and even the whole population. However, the positive consequence in peripheral ramets and whole fragments was only detected in the high‐nutrient patch of heterogeneous treatment. In sum, in the population with the radial ramet aggregation, clonal integration can play a key role in the rapid recruitment of young ramets of A. philoxeroides fragments, as well as the expansion of the whole population. The magnitude of clonal integration also became more obvious in the peripheral young ramets and whole fragments that experienced high‐nutrient patches.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

Effects of stolon severing on the expansion of Alternanthera philoxeroides from terrestrial to contaminated aquatic habitats

Few studies have examined the effects of clonal integration (translocation of resources between interconnected ramets) during the expansion of amphibious clonal plants from terrestrial to aquatic habitats. We conducted a greenhouse experiment to simulate the expansion of plants from terrestrial to contaminated aquatic habitats in the amphibious stoloniferous herb Alternanthera philoxeroides (alligator weed). The proximal ramets (i.e. relatively old) of clonal fragments grown in uncontaminated soils were connected to (allowing clonal integration) or disconnected from (preventing clonal integration) distal ramets (i.e. relatively young) grown either in uncontaminated water (control, no CuSO₄) or in four copper‐contaminated water treatments containing 31.25, 62.5, 125 and 250 mg/L CuSO₄, respectively. When a stolon connection was severed, all distal ramets grown in the contaminated water died. When the stolon connection was intact, however, the survival rate of the distal ramets was 85–100% when they were grown at the three lower levels of contamination and 43.75% at the highest level. Moreover, the survival rate and growth of the distal ramets grown in the three lower levels of contamination treatments did not differ from those in the control (uncontaminated water). These results suggest that clonal integration could greatly improve the survival and growth of alligator weed subjected to moderate levels of copper stress. Although clonal integration could also increase the survival rate of the connected distal ramets subjected to the highest level of copper stress (250 mg/L CuSO₄) compared with that of disconnected distal ramets, the survival rate and growth measures were still significantly lower than those in the control. This suggests that clonal integration plays a limited role in the survival and growth of alligator weed when it is subjected to severe stress by high levels of copper contamination.     

Clonal integration ameliorates the carbon accumulation capacity of a stoloniferous herb,Glechoma longituba,growing in heterogenous light conditions by facilitating nitrogen assimilation in the rhizosphere

Background and Aims Enhanced availability of photosynthates increases nitrogen (N) mineralization and nitrification in the rhizosphere via rhizodeposition from plant roots. Under heterogeneous light conditions, photosynthates supplied by exposed ramets may promote N assimilation in the rhizosphere of shaded, connected ramets. This study was conducted to test this hypothesis.Methods Clonal fragments of the stoloniferous herb Glechoma longituba with two successive ramets were selected. Mother ramets were subjected to full sunlight and offspring ramets were subjected to 80 % shading, and the stolon between the two successive ramets was either severed or left intact. Measurements were taken of photosynthetic and growth parameters. The turnover of available soil N was determined together with the compostion of the rhizosphere microbial community.Key Results The microbial community composition in the rhizosphere of shaded offspring ramets was significantly altered by clonal integration. Positive effects of clonal integration were observed on NAGase activity, net soil N mineralization rate and net soil N nitrification rate. Increased leaf N and chlorophyll content as well as leaf N allocation to the photosynthetic machinery improved the photosynthetic capability of shaded offspring ramets when the stolon was left intact. Clonal integration improved the growth performance of shaded, connected offspring ramets and whole clonal fragments without any cost to the exposed mother ramets.Conclusions Considerable differences in microbial community composition caused by clonal integration may facilitate N assimilation in the rhizosphere of shaded offspring ramets. Increased N content in the photosynthetic machinery may allow pre-acclimation to high light conditions for shaded offspring ramets, thus promoting opportunistic light capture. In accordance with the theory of the division of labour, it is suggested that clonal integration may ameliorate the carbon assimilation capacity of clonal plants, thus improving their fitness in temporally and spatially heterogeneous habitats.     

Physical connection decreases benefits of clonal integration in Alternanthera philoxeroides under three warming scenarios

Physical connection between ramets usually allows clonal plants to perform better but can have the opposite effects in some cases. Clonal integration and the effects of climate warming have been extensively studied, but to date little is known about how climate warming affects the benefits of clonal integration. We conducted a field experiment in which Alternanthera philoxeroides segments with connected and severed stolons were subject to four climate regimes (ambient, day warming, night warming and daily warming), and measured final biomass, number of ramets and total length of stolons. Across the three warming treatments, temperature rise suppressed growth of clonal fragments with connected stolons but increased growth of fragments with severed stolons; temperature rise affected the biomass of distal ramets but not proximal ramets, and had similar effects on the numbers of proximal and distal ramets. When the three warming treatments were considered separately, they had contrasting consequences for the benefits of clonal integration. Specifically, when fragments were exposed to day and night warming, physical connection evened out the advantages of clonal integration that occur under ambient conditions; when fragments were exposed to daily warming, physical connection led to smaller clonal plants. These findings suggest that physical connection between ramets may be disadvantageous to overall performance of A. philoxeroides fragments under climate warming, and also indicate that the net consequences of daily warming outweigh those of day or night warming.     

Effects of clonal integration on allelopathy of invasive plant Wedelia trilobata under heterogeneous light conditions

异质光照条件下克隆整合对入侵植物南美蟛蜞菊化感作用的影响 植物入侵已成为全球生态系统最严重的威胁之一。当具有克隆生长能力的入侵植物入侵或定殖到新的生境时,它们相互连接的分株可能受到异质光照的影响。在异质光照条件下,克隆整合对入侵植物化感作用的影响尚不清楚。为研究异质光照条件下克隆整合对入侵植物南美蟛蜞菊(Wedelia trilobata) 化感作用的影响,采用两个连续分株的克隆片段进行了盆栽试验。较老的分株暴露在全光下,而年轻的分株则受到20%的全光照。同时,每个克隆片段的年轻分株与目标植株(一个番茄苗)在盆栽中相邻生长。南美蟛蜞菊的两个连续分株之间的匍匐茎设置切断和不切断两种处理。另外,两株番茄幼苗 (一株作为目标植株)在盆栽中相邻生长作为对照。研究结果表明,当与目标植物相邻生长的南美蟛蜞菊分株之间的匍匐茎保持完整时,目标植株的生物量积累、叶片叶绿素和氮含量、叶绿素荧光参数和净光合速率及其根长和活性相较于匍匐茎切断处理显著降低。异质光照条件下连续两个分株之间碳水化合物的运输或共享可以增强20%全光处理下年轻分株的化感作用。克隆整合在异质光照条件下对具有克隆生长能力的入侵植物的入侵或定殖具有重要意义。     

克隆整合对无芒雀麦在异质性盐分环境中存活和生长的影响

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在异质性盐分环境中存活和生长的影响.结果表明,克隆整合显著提高了无芒雀麦分株在高盐环境中的存活能力,耗-益分析表明无芒雀麦在高盐斑块中分株的生物量、分株数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的非盐分斑块中的分株却没有产生显著的损耗.因而,克隆整合特性是无芒雀麦对异质性环境形成的重要适应对策,它使无芒雀麦能够扩展到不适合植物生长的高盐分斑块中,从而增加了无芒雀麦在浑善达克沙地中的存活和生长,提高了其在半干旱沙化地区的适合度.