荒漠土壤因子和DSE定殖对克隆植物入侵的响应

克隆植物,尤其是游击型克隆植物,具有很强的扩展能力,通过克隆扩展可侵入到不同生境斑块。克隆植物入侵可能会影响入侵地土壤营养状况和微生物群落。为了探明克隆植物入侵对DSE(dark septate endophytes)活动和土壤理化性质的影响,于2013年6月在克隆植物羊柴(Hedysarum laeve)和沙鞭(Psammochloa villosa)群落空地沿根状茎延伸方向设置样方,分别于6月、8月和10月在样方内分0—10、10—20、20—30、30—40、40—50 cm土层采集土样和根样,研究了不同采样时间羊柴和沙鞭群落空地DSE和土壤理化性质时空变化。结果表明,从6月到10月,随时间后延,克隆植物逐渐侵入群落空地,沙鞭入侵群落空地数和分株数高于羊柴。羊柴群落空地根系DSE定殖率随采样时间后延,逐渐降低,最大值在6月;沙鞭群落空地根系DSE定殖率随采样时间后延,逐渐升高,最大值在10月。随着克隆植物入侵,入侵地土壤中可利用的营养物质含量显著提高,羊柴入侵提高了入侵群落空地土壤碱解N、有效P和速效K含量,沙鞭入侵提高了入侵群落空地土壤碱解N和有效P含量。相关性分析表明,羊柴群落空地DSE定殖率与土壤p H值和电导率显著正相关,沙鞭群落空地DSE定殖率与土壤p H值极显著负相关,与电导率、碱解N和有效P极显著正相关。克隆植物入侵使得土壤环境更有利于克隆植物自身生长,为荒漠植被恢复提供了前提。     

Responses of Hedysarum Laeve , a guerrilla clonal semi-shrub in the Mu Us sandland, to local sand burial

In arid and semi-arid inland deserts, one of the environmental stresses for plants is recurrent sand burial, which can influence the physical and biotic microenvironments of the plants and soil. Previous studies have shown that different levels of sand burial have different effects on plants. Slight sand burial could increase the height increment, leaf biomass and the number of new ramets of the plants while heavy sand burial could impair the growth of the plants and even decrease their chances of survival. In other words, below a certain threshold level of burial, the growth of plants is stimulated probably because of multiple factors. However, as the level of burial increases, the positive response starts to decline until it becomes a negative value. Arid and semi-arid inland deserts are frequently colonized and stabilized by many rhizomatous clonal plants. Clonal physiological integration often helps clonal plants buffer local environmental stress encountered by ramets. A rhizomatous clonal semishrub, Hedysarum laeve (H. laeve), is the dominant plant species and important for vegetation restoration in the Mu Us sandland. To investigate whether clonal integration can increase the threshold of sand burial and help rhizomatous H. laeve tolerate heavy sand burial, we conducted a field experiment. The results showed that slight sand burial could accelerate ramet growth and enhance leaf biomass, stem biomass and shoot biomass, while heavy sand burial reducesed the biomass of the plant and impairs survival and growth of the ramets. Clonal integration increased the threshold of sand burial. Under heavy sand burial, ramets connected to other ramets not buried in sand were more in terms of height increment, stem biomass, leaf biomass and shoot biomass compared to the ramets encountering sand burial but disconnected from other ramets. It suggested that clonal physiological integration could help H. laeve ramets tolerate relatively heavy sand burial. We also discussed that clonal integration plays a role in H. laeve presence in the Mu Us sandland. __________ Translated from Journal of Plant Ecology (formerly Acta Phytoecologica Sinica), 2006, 30(2): 278–285 [译自: 植物生态学报]     

The significance of rhizome connection of semi-shrub Hedysarum laeve in an Inner Mongolian dune,China

Hedysarum laeve, a rhizomatous clonal semi-shrub, commonly dominates the inland dunes in semiarid areas of northern China. This species propagates vegetatively by extension of horizontal woody rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, the plants were experimentally manipulated by cutting rhizome connections and ¹⁴C-labelling techniques were employed to investigate the ecological significance of rhizome connections within the H. laeve clone. Severance of rhizome connections had a great effect on the performance of young ramets within a clone. Young ramets severed from their parent ramets experienced a significant reduction both in ramet growth and vegetative propagation, as compared with the intact young ramets. Within clonal fragments, consisting of three interconnected ramets including a mother ramet, a daughter ramet and a granddaughter ramet, ¹⁴C-photosynthates from the fed leaves of mother ramets were acropetally transported to all clonal component parts. The ¹⁴C-photosynthate translocation within the clonal fragment provides evidence that the young ramets were supported by their parent ramets. Our results suggest that the woody rhizome connections among the interconnected ramets are ecologically and strategically important for the species to grow in the sand dune habitat.     

Do rhizome severing and shoot defoliation affect clonal growth of Leymus chinensis at ramet population level?

Leymus chinensis (Trin.) Tzvel. is a perennial species of Gramineae, usually subject to defoliation from grazing and mowing. We examined whether shoot defoliation and rhizome severing affected rhizome and ramet growth, and vegetative bud outgrowth of L. chinensis ramet populations. We also tested the hypothesis that clonal growth of the ramets subject to defoliation would benefit from clonal integration between interconnected ramets besides from possible compensatory growth. To 48 experimental plots, we applied six treatments resulting from interactions between two rhizome connection states (unsevered/severed) and three defoliation regimes (non-defoliated, mildly-defoliated and heavily-defoliated). Defoliation affected rhizome growth and bud outgrowth, but had little effect on shoot growth. Mild and heavy defoliation exerted similar effects on rhizome growth. Only heavy defoliation significantly reduced bud outgrowth while mild defoliation did not. The fact that shoot growth did not change after defoliation and that the bud numbers remained unchanged after mild defoliation suggest that the compensatory response enable the species to tolerate grazing to some extent. Neither rhizome severing nor the interaction of rhizome severing and defoliation had effect on any tested variables. Lack of the effect of rhizome severing falsified the first half of our hypothesis, that is, clonal integration was unimportant in our experiment. The probable reasons were suspected to be the short duration of the experiment and/or the buffer effect of carbohydrate reserves in rhizomes for shoot growth and bud production in time of defoliation.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

Ramet phenology and clonal architectures of the clonal sedge <Emphasis Type="Italic">Schoenoplectus americanus</Emphasis> (Pers.) Volk. ex Schinz &; R. Keller

The clonal plant Schoenoplectus americanus shows variable belowground clonal architecture as a result of producing two types of ramets: those with very long rhizomes (long rhizome ramet, LRR) and those with very short ones (short rhizome ramet, SRR). In a previous study we demonstrated that the two types of ramets are functionally specialised. The production of SRRs results in the formation of consolidated clonal patches with densely packed shoots, while the production of LRRs results in a more diffuse network of connected rhizomes with widely spaced shoots. We hypothesised that the two types of ramets would be produced at different times during the growing season because of their functional differences. The production of LRRs throughout the growing season would enable the species to continuously explore new habitats while the production of SRRs early in the growing season would enable the species to occupy and consolidate resources in available open patches. We evaluated this hypothesis through field observations in different communities with S. americanus and indeed found that SRRs were produced early in the growing season while LRRs tended to be produced over an extended period of time. Plants in high-quality environments (i.e. higher light conditions) produced more SRRs, and these were formed early in the growing season. In contrast, plants in low-quality environments produced more LRRs, and these were formed continuously over the growing season. We also observed that the shoot longevity was greater for SRR. In high-quality patches, the production of the lower cost SRRs results in a more rapid occupancy of open spaces; in lower quality patches, the production of LRRs throughout the growing season enables plants to explore the immediate environment for higher quality patches.     

Clonal integration and its ecological significance in Hedysarum laeve, a rhizomatous shrub in Mu Us Sandland

Hedysarum laeve, a rhizomatous clonal half-shrub, commonly dominates in inland dunes in semiarid areas of northern China. This species propagates vegetatively by the extension of horizontal rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, ¹⁴C labeling and experimental defoliation were employed to test the photosynthate translocation within the interconnected parent–daughter ramet pairs. A proportion of ¹⁴C-photosynthates was transported from the parent ramet into the daughter ramet, the roots of the daughter ramet, and the rhizome; these three components showed more than 70% sink activity after 24-h translocation. On the other hand, the basipetal translocation (from daughter ramet into parent ramet) was relatively small with sink activity of less than 5%, but sink activity of the rhizome exceeded 10%. Defoliation had an influence on the photosynthate translocation between parent and daughter ramets. The intact parent ramets significantly increased their ¹⁴C-photosynthate translocation into defoliated daughter ramets when compared to intact daughter ramets. The daughter ramets transported significantly more ¹⁴C-photosynthates to the defoliated parent ramets than to the intact parent ramets. A portion of ¹⁴C-photosynthates was transported into the rhizome from both parent and daughter ramets, indicating that the rhizome is supported by both ramets for photosynthates. The clonal integration between ramets of the species through rhizome connection may confer benefit both to the ramets and the genet in adverse environments. Received: April 12, 2001 / Accepted: November 26, 2001     

The effect of resource integration on fitness related traits in a clonal dune perennial,Hydrocotyle bonariensis

Summary The costs and benefits of resource integration in Hydrocotyle bonariensis were examined by comparing severed and intact clones grown across multiple resource gradients. Basipetal movement of water, nitrogen and photosynthates was demonstrated to occur between two rhizome branch systems interconnecting hundreds of ramets within a clone. Hydrocotyle clones of this size and larger have been shown to span highly patchy and unpredictable resource conditions in coastal dune environments. The extensive movement of water and nitrogen to portions of a clone deficient in these resources, resulted in a significant net benefit to the clone in terms of fitness-related traits: total biomass, ramet proliferation and seed production. The translocation of photosynthates across light gradients allowed for sexual reproduction and clonal expansion in the shade although this incurred a net cost to the clone in terms of ramet and seed production.     

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.     

Clonal integration improves compensatory growth in heavily grazed ramet populations of two inland-dune grasses

Herbaceous species possess several mechanisms to compensate for tissue loss. For clonal herbaceous species, clonal integration may be an additional mechanism. This may especially hold true when tissue loss is very high, because other compensatory mechanisms may be insufficient. On inland dunes in northern China, we subjected Bromus ircutensis and Psammochloa villosa ramets within 0.5 m×0.5 m plots to three clipping treatments, i.e., no clipping, moderate (50% shoot removal) and heavy clipping (90% shoot removal), and kept rhizomes at the plot edges connected or disconnected. Moderate clipping did not reduce ramet, leaf or biomass density of either species. Under moderate clipping, rhizome connection significantly improved the performance of Psammochloa, but not that of Bromus. Heavy clipping reduced ramet, leaf and biomass density in the disconnected plots of both species, but such negative effects were negated or greatly ameliorated when the rhizomes were connected. Therefore, clonal integration contributed greatly to the compensatory growth of both species. The results suggest that clonal integration is an additional compensatory mechanism for clonal plants and may be important for their long-term persistence in the heavily grazed regions in northern China.     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

Effects of moisture availability on clonal growth in bamboo Pleioblastus maculata

The effects of moisture availability on clonal growth and biomass investment in the bamboo Pleioblastus maculata were investigated over a four-year period by transplanting Pleioblastus maculata clones into soils with different levels of moisture availability in the field. The results showed that: (1) The higher the moisture availability, the greater the total biomass of P. maculata clones. Although fewer culms are produced at the higher moisture levels, mean tiller biomass is greater. (2) Under different levels of moisture availability, obvious differences in the total rhizome length (p < 0.01), spacer length (p < 0.05) and the sizes of bamboo culms (height, p < 0.01; diameter, p < 0.01) were observed. Thus, the higher the moisture availability, the shorter the rhizomes and the larger ramets. (3) In microhabitats with low moisture availability, bamboo allocated more biomass to underground organs, which promotes elongation of rhizomes and increases root production, thereby helping to capture underground resources essential to growth. In microhabitats of high moisture availability, the biomass is primarily allocated to the aboveground growth of ramets. (4) We suggest that soil moisture availability effects the foraging strategies of bamboo, that bamboo plants growing with low moisture availability produce longer rhizomes (that is, more, although shorter, spacers) with more biomass allocation than plants in high moisture and have a better ability to forage to increase the probability of locating adequate moisture patches. Also, longer length distance between shoots (that is, longer spacers) in high soil moisture than in low is adapted to avoid intense competition from faster growing aboveground growth in high moisture patches.     

Clonal integration facilitates the proliferation of smooth brome clones invading northern fescue prairies

Predicting exotic invaders and reducing their impacts on the biodiversity and function of native ecosystems require understanding of the mechanisms that facilitate their success during key stages of invasion. We determined whether clonal growth, characteristic of the majority of successful invaders of natural areas, facilitates the proliferation of Bromus inermis (smooth brome), an exotic grass invading prairie ecosystems across the Great Plains. By manipulating the below-ground connections of proliferating rhizomes as well as the levels of soil nitrogen along the margins of clones invading northern fescue prairies in Manitoba, Canada, we hypothesized that physiological integration would most benefit ramets invading low resource environments. Severing clonal connections reduced the mass of smooth brome shoots invading native prairies and was exacerbated by the immobilization of soil nutrients with glucose. Clonal connections were equally important in the maintenance of smooth brome density and the horizontal proliferation of ramets. Our results demonstrate the role of physiological integration in the proliferation of a clonal exotic invader and may help explain the success of clonal invaders in other regions. Although integration among invading ramets suggests several possibilities for successful management, future research must continue to elucidate differences in the invasiveness of native versus exotic species as well as the persistence of clonal connections among exotic invaders.     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

Long-term suppression of insect herbivores increases the production and growth of Solidago altissima rhizomes

Summary Although insect herbivores have many well documented effects on plant performance, there are few studies that assess the impact of above-ground herbivory on below-ground plant growth. For a seven year period in which no large-scale herbivore outbreaks occurred, a broad spectrum insecticide was utilized to suppress herbivorous insects in a natural community dominated by Solidago altissima. Ramet heights, rhizome lengths, rhizome biomass, and the number of daughter rhizomes all were lower in the control plots than in the insecticidetreated plots. These effects should lead to a decrease in the fitness of genets in the control plots relative to the fitness of genets in the insecticide-treated plots. We also found that ramets in the control plots appear to have compensated for herbivory: the ratio of rhizome length to rhizome biomass was greatest in the control plots, which indicates that clones moved farther per unit biomass in these plots than in the insecticide-treated plots. Clonal growth models show that this shift in allocation patterns greatly reduced the magnitude of treatment differences in long-term clonal displacements.Previous work has shown, and this study verified, that clonal growth in S. altissima is well represented by random-walk and diffusion models. Therefore, we used these models to examine possible treatment differences in rates of clonal expansion. Although rhizome lengths were greater in the insecticide-treated plots, results from the models suggest that our treatments had little impact on the short- and long-term displacement of S. altissima ramets from a point of origin. This occurred because S. altissima ramets backtrack often, and thus, treatment differences in net displacements are less pronounced than treatment differences in rhizome lengths.     

Clonal integration enhances flood tolerance of Spartina alterniflora daughter ramets

Recently, considerable attention has been paid to the invasion of the clonal plant Spartina alterniflora into coastal wetlands at lower elevations. In this experiment, we tested whether clonal integration improved flood tolerance in S. alterniflora daughter ramets. Daughter ramets at two growth stages (young and old ramets) were flooded to water levels of 0, 9 and 18 cm above the soil surface, and the rhizomes between mother and daughter ramets were either severed or left intact. Biomasses of connected ramets grown in controls or in shallow and deep water treatments were 119%, 108% and 149% higher in the old ramet group than those of severed ramets, respectively, whereas they were 3.0, 3.3 and 11.2 times higher in the young ramet group, respectively. At the end of the experimental period, the shoot height, connected with young ramets, in shallow and deep water treatments increased by 19% and 26%, respectively, over that in the control treatments, whereas the old ramets increased by 11% and 39%, respectively. In contrast, the shoot height of the severed young ramets was 27% and 26% lower in shallow and deep water treatments than in the control treatment, respectively. However, the shoot height of the severed old ramets remained constant with increasing water depth. We conclude that clonal integration enhances the flood tolerance of S. alterniflora daughter ramets, and the trait of clonal integration plays more important roles in severe flooding stress conditions and at early growth stages.     

Physiological integration helps a clonal macrophyte spread into competitive environments and coexist with other species

Physiological integration may help clonal macrophytes invade or escape from existing communities. No studies have tested the above hypothesis in aquatic plants. In an outdoor pond experiment, we subjected clonal fragments of the submerged macrophyte Vallisneria spiralis L. to heterogeneous environments in which V. spiralis spread from bare habitats towards vegetated habitats occupied by Myriophyllum spicatum L. or V. spiralis spread from vegetated habitats towards bare habitats. V. spiralis stolons between ramets in bare habitats and in vegetated habitats were either intact or severed. We investigated the habitat selection of V. spiralis by examining the allocation of biomass and ramets to heterogeneous habitats during its vegetative spread phase. Results showed that the stolon connection had different effects on the habitat selection of V. spiralis with regard to invasion and escape. When V. spiralis spread from bare to vegetated habitats, in comparison to severing the stolon, the stolon connection eventually facilitated a 49% increase in biomass and a 27% increase in number of ramets allocated to vegetated habitats. However, when V. spiralis spread from vegetated to bare habitats, biomass and ramets allocated to bare habitats were not significantly changed by the stolon connection (only a 5% increase in biomass and a 6% increase in number of ramets). These results indicate that clonal integration facilitated V. spiralis not to escape from but invade into vegetated habitats. The study provides evidence that physiological integration is important for survival and tolerance of ramets in competitively stressful environments and can help clonal macrophytes coexist with other species.     

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.     

Density- and growth stage-dependent responses to defoliation in two rhizomatous grasses

Summary Responses to defoliation were studied in two tallgrass prairie perennials (Andropogon gerardii and Panicum virgatum) established from seed at three densities. P. virgatum was also grown from transplanted rhizomes of established clones. Plants of both species displayed a continuum of responses to defoliation, from large reductions in biomass, tillering and seed production to significant increases in one or more performance measures. In crowded populations, defoliation shifted plants into subordinate positions within the competitive hierarchy. Plants competing intraspecifically and those that were initially small suffered more from defoliation than either plants grown at low density or those that were larger than their neighbors. At the highest plant density, the effects of defoliation or initial plant size were overshadowed by the effects of crowding. When defoliated and grown at similar densities, P. virgatum and A. gerardii grown from seed showed large reductions in biomass, seed production, and new rhizome production, but established P. virgatum ramets grown from rhizomes showed increases in these performance measures. Thus, herbivory may be particularly detrimental to P. virgatum during juvenile stages before perennating organs have developed. Overcompensation of P. virgatum clones in response to defoliation only occurred if all ramets within the clone were defoliated. In clones containing both defoliated and undamaged ramets, there were no differences in their performance, suggesting that genets are capable of integrating the effects of differential defoliation among shoots. Defoliated P. virgatum clones allocated a smaller fraction of their total biomass to new rhizomes, indicating that the short-term regrowth response following defoliation may incur a longer-term cost associated with gradual reduction in biomass of the perennating organs and reduced genet success.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.