Ephemeral clonal integration in Calathea marantifolia (Marantaceae): Evidence of diminished integration over time

A major advantage of clonal growth forms is the intergenerational transfer of resources through vascular connections (clonal integration). Connections linking ramets can be persistent or ephemeral. For species with ephemeral connections, whether the extent of clonal integration changes over time is unclear. To address this issue, we tracked water movement using an isotopic label and assessed the demographic performance of parent and offspring ramets over time in a severing experiment. Our study system was the understory herb Calathea marantifolia, which has parent ramets that produce vegetative bulbils (clonal offspring) that pass through distinct pre- and post-rooting stages. Little water was transported between parents and offspring, and the direction of movement was primarily from parent to pre-rooting offspring. Anatomical observations of inter-ramet connections showed that vascular bundles were twice as abundant in parent stems compared to inter-ramet connections. Severing inter-ramet connections reduced the growth of offspring ramets but not parents. Survival of pre-rooting offspring was reduced by 10% due to severing, but post-rooting offspring were not affected. Our results suggest that offspring ramets of C. marantifolia are weaned from their parent as they progress from pre- to post-rooting stages.     

Effects of the loss of clonal integration on four sedges that differ in ramet aggregation

Although clonal growth is a dominant mode of plant growth in wetlands, the importance of clonal integration, resource sharing among ramets, to individual ramet generations (mother and daughter) and entire clones of coexisting species has not been well investigated. This study evaluated the significance of clonal integration in four sedge species of varying ramet aggregations, from clump-forming species (Clumpers –Carex sterilis, Eleocharis rostellata), with tightly aggregated ramets (rhizomes<1cm), to runner species (Runners –Schoenoplectus acutus, Cladium mariscoides), with loosely aggregated ramets. We manipulated clonal integration by either severing connections between target mother and daughter ramets or leaving connections intact, and then planted them in an intact neighborhood of a fen in Michigan, USA. We measured growth parameters of original and newly produced ramets over two growing seasons and conducted a final biomass harvest, to address four hypotheses. First, we expected integrated clones to accumulate more biomass than severed clones. However, final clone-level biomass and ramet production were the same for both treatments in all species although severing initially stimulated ramet production by Schoenoplectus and produced a more compact ramet aggregation in Cladium. Second, we hypothesized that mother ramets would experience a cost of integration, through reduced ramet or biomass production, while daughters would experience a benefit, through increased resource availability from mothers. Mother ramets of Cladium suffered a cost from integration, while Schoenoplectus mothers suffered a slight cost and Carex daughters saw a slight benefit. Finally, we hypothesized that integration would be more active in runner species than in clumper species. Indeed, we documented more active integration in runners than clumpers, but none of the study species were dependent upon integration for growth or survival once daughter ramets were established with their own roots and shoots. This study demonstrates that integration between established ramets may not be the most important advantage to clonal growth in this wetland field site. The loss of integration elicited varied responses among coexisting species in their natural habitat, somewhat but not completely related to their growth form, suggesting that a combination of plant life history traits contributes to the dependence upon clonal integration among established ramets of clonal species.     

Physiological integration enhanced the tolerance of Cynodon dactylon to flooding

Many flooding‐tolerant species are clonal plants; however, the effects of physiological integration on plant responses to flooding have received limited attention. We hypothesise that flooding can trigger changes in metabolism of carbohydrates and ROS (reactive oxygen species) in clonal plants, and that physiological integration can ameliorate the adverse effects of stress, subsequently restoring the growth of flooded ramets. In the present study, we conducted a factorial experiment combining flooding to apical ramets and stolon severing (preventing physiological integration) between apical and basal ramets of Cynodon dactylon, which is a stoloniferous perennial grass with considerable flooding tolerance. Flooding‐induced responses including decreased root biomass, accumulation of soluble sugar and starch, as well as increased activity of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in apical ramets. Physiological integration relieved growth inhibition, carbohydrate accumulation and induction of antioxidant enzyme activity in stressed ramets, as expected, without any observable cost in unstressed ramets. We speculate that relief of flooding stress in clonal plants may rely on oxidising power and electron acceptors transferred between ramets through physiological integration.     

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。     

克隆整合提高淹水胁迫下狗牙根根部的活性氧清除能力

虽然国内外已开展大量关于克隆整合影响植物抗逆生理的研究,但迄今未见克隆整合是否会影响逆境下不同分株清除活性氧过程的报道。以河岸带适生克隆植物狗牙根(Cynodon dactylon)为例,研究克隆植物的抗氧化生理响应,检测了狗牙根在先端淹水/不淹水、先端与基端匍匐茎连接/切断两个因素的交互作用下的根部主要抗氧化酶:超氧化岐化酶(Superoxide dismutase, SOD)、抗坏血酸过氧化物酶(Ascorbate peroxidase, APX)、过氧化氢酶(Catalase, CAT)的活力以及生物量的变化。结果显示,淹水环境中狗牙根先端的生物量和根部SOD酶活力在匍匐茎连接处理下显著高于切断处理组,同一处理的生物量以及根部APX、CAT酶活力总体上表现出不同程度的提高趋势;与受淹先端连接的基端分株根部抗氧化酶活力均低于切断处理组,且SOD和CAT受连接处理影响显著;淹水和切断处理显著降低先端分株的生物量,但对基端和克隆片段影响不明显。这表明淹水胁迫下克隆整合提高了其根部活性氧清除能力,显著改善了先端分株的表现。     

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.     

The effects of clonal integration on morphological plasticity and placement of daughter ramets in black locust (Robinia pseudoacacia)

We studied the field response of Robinia pseudoacacia L. to light, total soil nitrogen, available soil phosphorus and soil pH. Results indicated that there was very strong clonal integration between mother and daughter ramets. Mother ramets can provide nitrogen and phosphorus to daughter ramets sufficient for their continued growth through strong clonal integration, but cannot provide enough photosynthate. With clonal integration, soil nitrogen and phosphorus availability had no effect on biomass allocation to roots, number of ramets and length of connection roots. Biomass allocation to roots increased markedly and responded to nitrogen and phosphorus availability, when the connections were severed. Light had a significant effect on the percent of biomass allocation to leaves and number of ramets, but no effect on the length of connection roots. Daughter ramets allocated more resources to leaves, and clones placed more daughter ramets in high light patches than in low light patches. Soil pH had a significant effect on ramet number and connection root length. Clones concentrated in alkaline patches and escaped from acid patches through selective placement of daughter ramets and changing the length of connection roots. We suggest that the clonal integration may be very strong and provide sufficient soil resources to daughter ramets, then affect the daughter ramets’ morphology and placement, if the size of a specific ramet is significantly larger than the other ramets in an arbor clone.     

Effects of physical connection and genetic identity of neighbouring ramets on root-placement patterns in two clonal species

Root-placement patterns were examined in the clonal species Glechoma hederacea and Fragaria vesca when grown with different types of neighbours. Three different patterns were predicted as consequences of different types of interactions between roots: the avoidance pattern if root growth decreases in the presence of neighbouring roots; the intrusive pattern if root growth increases towards neighbouring roots; and the unresponsive pattern if root growth is unaffected by neighbouring roots. Experiments were conducted in which physical connection between ramets, and the genetic identity of neighbouring ramets, were manipulated. The patterns of distribution of entire root systems and elongation rates of individual roots were measured. Root systems and individual roots of G. hederacea avoided contact with roots of neighbouring ramets, irrespective of connection to the neighbour and its genetic or specific identity. In contrast, F. vesca roots grew equally towards and away from intraspecific ramet neighbours and their elongation was stimulated by contact with roots of G. hederacea ramets. These results demonstrate that root-placement patterns of plants grown with different types of neighbours vary between species, and suggest that factors additional to resource depletion could be involved in their development.     

Do Physiological Integration and Soil Heterogeneity Influence the Clonal Growth and Foraging of <Emphasis Type="Italic">Schoenoplectus pungens</Emphasis>?

Physiological integration and foraging behavior have both been proposed as advantages for clonal growth in heterogeneous environments. We tested three predictions concerning their short- and long-term effects on the growth of the clonal perennial sedge Schoenoplectus pungens (Pers.) Volk. ex Schinz and R. Keller: (1) growth would be greatest for clones with connected rhizomes and on heterogeneous soil, (2) clones would preferentially place biomass in the nutrient-rich patches of a spatially heterogeneous environment, and (3) physiological integration would decrease a clone’s ability to forage. We tested our predictions by growing S. pungens clones for 2 years in an experimental garden with two severing treatments (connected and severed rhizomes) crossed with two soil treatments (homogeneous and heterogeneous nutrient distribution). Severing treatments were only carried out in the first year. As predicted, severing significantly decreased total biomass and per capita growth rate in year one and individual ramet biomass both in year one and the year after severing stopped. This reduction in growth was most likely caused by severing damage, because the total biomass and growth rate in severed treatments did not vary with soil heterogeneity. Contrary to our prediction, total biomass and number of ramets were highest on homogeneous soil at the end of year two, regardless of severing treatment, possibly because ramets in heterogeneous treatments were initially planted in a nutrient-poor patch. Finally, as predicted, S. pungens concentrated ramets in the nutrient-rich patches of the heterogeneous soil treatment. This foraging behavior seemed enhanced by physiological integration in the first year, but any possible enhancement disappeared the year after severing stopped. It seems that over time, individual ramets become independent, and parent ramets respond independently to the conditions of their local microsite when producing offspring, a life-history pattern that may be the rule for clonal species with the spreading “guerrilla” growth form.     

Molecular and dendrochronological analysis of natural root grafting in Populus tremuloides (Salicaceae)

Trembling aspen (Populus tremuloides) is a clonal tree species, which regenerates mostly through root suckering. In spite of vegetative propagation, aspen maintains high levels of clonal diversity. We hypothesized that the maintenance of clonal diversity in this species can be facilitated by integrating different clones through natural root grafts into aspen's communal root system. To verify this hypothesis, we analyzed root systems of three pure aspen stands where clones had been delineated with the help of molecular markers. Grafting between roots was frequent regardless of their genotypes. Root system excavations revealed that many roots were still living below trees that had been dead for several years. Some of these roots had no root connections other than grafts to living ramets of different clones. The uncovered root systems did not include any unique genotypes that would not occur among stems. Nevertheless, acquiring roots of dead trees helps to maintain extensive root systems, which increases the chances of clone survival. Substantial interconnectivity within clones as well as between clones via interclonal grafts results in formation of large genetically diverse physiological units. Such a clonal structure can significantly affect interpretations of diverse ecophysiological processes in forests of trembling aspen.     

Clonal segmentation – The development of physiological independence within stolons of Glechoma hederacea L. (Lamiaceae)

The youngest parts of clonal plants benefit from substantial physiological support from older parts, but the extent to which this physiological dependence persists through time is poorly understood. The development of autonomy among connected subunits was therefore analysed in the clonal species Glechoma hederacea. The stolons of a series of clonal fragments with differing numbers of primary ramets were severed at a fixed point relative to the four oldest primary ramets. The subsequent growth of both parts of the severed fragments was compared with that of a series of intact fragments.The growth of apical stolon portions that included five or more rooted primary ramets at the time of severing was unaffected by severing. Apical portions with three or fewer rooted ramets at the time of severing produced fewer new primary ramets than equivalent parts of intact fragments, while apical portions with four or fewer rooted ramets produced less above-ground mass than equivalent apical portions of intact clonal fragments. Basal portions of clonal fragments severed when there were one or two rooted ramets in the apical portion produced more secondary ramet mass than equivalent parts of intact fragments. The gain in mass of secondary ramets in the basal portions of severed fragments matched the reduction in mass of secondary ramets in the apical portions. However, severing caused an overall loss of mass when apical portions had three or fewer rooted ramets at the time of severing, because the mass of primary ramets in basal portions did not increase following severing. Severing had little impact on the allometry of the apical portions. The relationship between mass in secondary ramets and mass in primary ramets was similar in the apical portions of severed and intact clonal fragments. None of the severing treatments increased the total mass of secondary ramets, suggesting that apical dominance in this species only affects branches very close to the apex.These observations, combined with existing knowledge of vascular architecture in G. hederacea, demonstrate that, whether or not physical connections persist between ramets, growing stolons rapidly develop into physiologically autonomous segments. This may be a characteristic of species that exploit disturbed, spatially heterogeneous habitats through rapid multiplication of ramets connected by long, aerial runners or stolons.     

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。     

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.     

克隆水分整合有助胡杨无性系幼株在极端干旱生境下保持更高生存优势

以塔里木河下游的胡杨为对象,研究胡杨母株与其克隆幼株间的克隆水分整合特征及对克隆幼株生理机能的影响.结果表明: 胡杨母株与其克隆幼株间存在明显的以顶向传输为特征的水分整合,母株对克隆幼株的日平均水分整合流量为同区域长势相近的实生幼株通过自身根系日平均获取水量的3倍,克隆幼株因水分整合而能够获取与母株相似且较实生幼株水分来源相对较深层的土壤水,拥有更好的水分获取能力.这使胡杨克隆幼株较实生幼株保持显著更高的叶片水分含量,且黎明前和正午叶水势分别显著增加10.0%和29.7%.更好的水分生理状态使克隆幼株的光合电子传递速率与光适应下实际光化学效率显著高出实生幼株23.4%和11.5%,并能够保持显著更低的光系统Ⅱ过剩激能比例和光抑制风险.这可能让胡杨克隆幼株在极端干旱生境中较实生幼株保持更高的生存优势.     

Ecological benefits of integration of Calamagrostis epigejos ramets under field conditions

We established two independent experiments to estimate the ecological consequences of artificial severing on ramets of the competitively strong perennial grass Calamagrostis epigejos. We compared the responses of mature ramets of different size growing in different environments. Finally, we tested whether the response of young ramets to the severing depends on the density of surrounding vegetation.Severing decreased biomass and number of rhizomes of young ramets but did not affect their competitive tolerance. It decreased survival of mature ramets (probably due to traumata caused by cutting attached rhizomes) but did not influence total biomass of survived ramets. The response of total biomass of mature ramets to severing changed with size of the ramets. Further, biomass allocation to rhizomes changed differently after ramet severing in the two environments.The results suggest that field grown young ramets of Calamagrostis benefit from physiological integration. In contrast, mature ramets seem to be independent units according to the resource economy. Ecological benefits of integration depended on size of ramet clumps: ramets in clump had larger survival probability than control ramets. They also depended on environmental conditions: severing increased formation of new rhizomes at a sand dune subsrate, but it had a negative effect on plants in the forest experimental site. This intra-specific variation should be taken into account when trying to explain ecological patterns of integration benefits of clonal plants.     

Fitness Implications of Clonal Integration and Leaf Dynamics in a Stoloniferous Herb, Nelsonia canescens (Lam.) Spreng (Nelsoniaceae)

Very little has been published on the life-history significance of clonal plants inhabiting southern African savanna environments. This study investigated the fitness implications of clonal integration, resprouting behaviour and growth phenology in a stoloniferous herb, Nelsonia canescens (blue pussyleaf) at a savanna site in Zambia, central Africa. Census data on growth and survival were obtained regularly on permanently marked ramets over a 4-year period, from 2001 to 2005, and analyzed to assess how physiological integration and module demography contribute to fitness in Nelsonia. Above ground and below ground growth occurred during the dry and rainy seasons, respectively. Dry season growth was characterized by resprouting and production of stolons that bore small pubescent leaves with high mortality (30–80% month⁻¹). Deep roots and high leaf turnover appear to contribute to sustained growth during the dry season when topsoil moisture and nutrient availability are low. The interaction between maximum temperature and precipitation explained a significant proportion (59%, p<0.01) of the monthly variation in leaf size and increasing evapo-transpiration levels appeared to trigger the shift in leaf size from a large wet season type to a small dry season one. During the dry season Nelsonia resprouted from dormant buds buried at the time of root development in daughter ramets in the rainy season. Temporal integration significantly (p<0.05) enhanced survival of daughter ramets. However, daughter ramets with severed mother–daughter ramet inter-connectors experienced high initial mortality that was caused by both early stolon severing and drought stress during the root development phase. The majority of ramets lived for 5–10 months and 25% were still alive at the age of 3.5 years. The study showed that although the growth phenology of Nelsonia has serious ecological implications for accessing scarce resources during the dry season, the species utilizes a number of strategies to overcome resource limitations in a seasonally heterogenous environment. Co-ordinating editor: G. P. Cheplick     

Clonal integration increases performance of ramets of the fern Diplopterygium glaucum in an evergreen forest in southeastern China

Physiological integration is a major ecological advantage of clonal growth in angiosperms. Clonal growth is also common in pteridophytes, but almost no study has tested whether clonal integration increases performance in ramets of pteridophytes in natural populations. To test this hypothesis and also whether the positive effect of integration is greater on smaller ramets, we severed the connecting rhizomes of individual ramets of the common, understory fern Diplopterygium glaucum in an evergreen, broadleaf forest in southeastern China. In another experiment, we severed rhizomes around the edges of small plots each containing several ramets. After 19.5 weeks, survival was 100% in intact individual ramets but only 27% in severed ones. Among surviving ramets, final dry mass and lamina mass were also less in severed than in intact ramets, though stalk, rhizome, and root mass and maximum quantum yield of PSII (F_v/F_m) were not reduced. Individual ramets with fewer stalk nodes had lower dry mass but were not more affected by severing than ramets with more stalk nodes. Severance around the edge of plots did not significantly affect the combined final mass of the ramets within a plot. We conclude that clonal integration can have significant positive effects on both survival and growth of individual ramets of ferns in natural populations.     

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。     

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.     

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.