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     

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, (14)C labeling and experimental defoliation were employed to test the photosynthate translocation within the interconnected parent-daughter ramet pairs. A proportion of (14)C-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 (14)C-photosynthate translocation into defoliated daughter ramets when compared to intact daughter ramets. The daughter ramets transported significantly more (14)C-photosynthates to the defoliated parent ramets than to the intact parent ramets. A portion of (14)C-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.     

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

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

Partial mechanical stimulation facilitates the growth of the rhizomatous plant Leymus secalinus: modulation by clonal integration

Background and Aims

Mechanical stimulation (MS) often induces plants to undergo thigmomorphogenesis and to synthesize an array of signalling substances. In clonal plants, connected ramets often share resources and hormones. However, little is known about whether and how clonal integration influences the ability of clonal plants to withstand MS. We hypothesized that the effects of MS may be modulated by clonal integration.

Methods

We conducted an experiment in which ramet pairs of Leymus secalinus were subjected to three treatments: (1) connected ramet pairs under a homogeneous condition [i.e. the proximal (relatively old) and distal (relatively young) ramets were not mechanically stressed]; (2) connected ramet pairs under a heterogeneous condition (i.e. the proximal ramet was mechanically stressed but the distal ramet was not); and (3) disconnected ramet pairs under the same condition as in treatment 2. At the end of the experiment, we harvested all plants and determined their biomass and allocation.

Key Results

Clonal integration had no significant influence on measured traits of distal L. secalinus ramets without MS. However, under MS, plants with distal ramets that were connected to a mother ramet produced more total plant biomass, below-ground biomass, ramets and total rhizome length than those that were not connected. Partial MS exerted local effects on stimulated ramets and remote effects on connected unstimulated ramets. Partial MS increased total biomass, root/shoot ratio, number of ramets and total rhizome length of stimulated proximal ramets, and increased total biomass, root weight ratio, number of ramets and total rhizome length of connected unstimulated ramets due to clonal integration.

Conclusions

These findings suggest that thigmomorphogenesis may protect plants from the stresses caused by high winds or trampling and that thigmomorphogenesis can be strongly modulated by the degree of clonal integration.     

Evolutionary impacts of the goldenrod ball gallmaker on Solidago altissima clones

Summary Three ramet clones of Solidago altissima were grown under greenhouse conditions to determine the effects of varying levels of attack by the goldenrod ball gallmaker (Eurosta solidaginis) on biomass allocation, leaf senescence rate and rhizome connections among ramets. The results, examined at both the individual ramet level and the clone level, showed that galled ramets became isolated from their clone through deterioration of rhizome connections. Gall effects were only observed at the ramet level although rhizome connection effects were detected at both the ramet and clone levels. The goldenrod ball gallmaker may therefore have little evolutionary impact on large clones but could appreciably affect newly established clones.     

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.     

CLONE STRUCTURE IN FOUR SOLIDAGO ALTISSIMA (ASTERACEAE) POPULATIONS: RHIZOME CONNECTIONS WITHIN GENOTYPES

We studied clone structure and degree of genotypic mixing of Solidago altissima L. (Asteraceae) clones in four old fields near Ithaca, New York. The fields differed in time from agricultural abandonment and were approximately 1, 5, 20, and 35 years old. In the three older fields, three 0.75 m² plots were excavated intact and rhizome connections among ramets were mapped. In the youngest field 30 ramets were dug up singly. The genotype of all ramets was determined using electrophoresis of four polymorphic enzyme systems. Fields differed in the number and dispersion of genotypes within plots, and the degree of connection among ramets in the same clone. The one-year-old field was composed of single ramet genotypes which had probably established from seed the previous year. The five-year-old field contained many small contiguous clones of S. altissima with highly interconnected ramets. In the oldest two fields clones were highly intermixed and ramets of the same genotype were not extensively interconnected. These results demonstrate that clones of S. altissima display considerable phenotypic variability between fields and patterns of clone development may differ. The causes of this variability remain to be identified. We suggest that either selection for different genotypes or changing habitat conditions during succession may lead to changes in clone form.     

Water integration patterns in two rhizomatous dune perennials of different clonal fragment size

In clonal plants, a clonal fragment is a basic unit. There may exist clonal integration that is often regarded adaptive. By definition, a larger clonal fragment comprises more interconnected ramets and/or occupies a larger area of the habitat. When the habitat is so heterogeneous in terms of essential resources that the resources the whole clonal fragment needs can be captured only in a limited number of microhabitats, a large clonal fragment may require extensive and/or intensive clonal integration. Therefore, we hypothesize that in an environment where the resources are distributed in a highly heterogeneous way, a species forming large clonal fragments possesses more extensive and/or intensive clonal integration gaining essential resources than that forming small ones.This hypothesis was tested in a field experiment with two rhizomatous species growing in inland-dune habitats with high heterogeneity of water resource, one (Psammochloa villosa) forming large clonal fragments and the other (Hedysarum laeve) forming small ones. In P. villosa, tracer-(water-soluble acid fuchsin)-labeled water could be transported along the rhizome at a velocity of about 0.93 m per hour and 4 ramets per hour, which was much faster than that in H. laeve. Similarly, the acid fuchsin-labeled water was transported to a longer length in the rhizomes (3.96 m vs. 1.12 m) and to more ramets (14 vs. 3) in P. villosa than in H. laeve. In P. villosa, the acid fuchsin-labeled water reached the ends of the rhizomes and all ramets along the rhizomes were dyed purple. In H. laeve, however, the acid fuchsin-labeled water could not reach the rhizome tips and along the rhizomes only some ramets were labeled. The results suggest that P. villosa is an extensive and intensive integrator, whereas H. laeve is a restrictive and less intensive integrator, supporting the hypothesis. The possible mechanisms and the ecological implications of the findings were discussed.     

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.     

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.     

Physiological integration among ramets of Lathyrus sylvestris L.

Summary Lathyrus sylvestris is a pioneer legume often found in disturbed habitats. Mainly reproduced through vegetative propagation, this clonal species presents a system of ramets that remain connected for several years. The existence of carbon transfer among ramets within a clone has been studied using ¹⁴C in situ. Assimilate translocation from primary to secondary ramets was observed in all clones when the primary ramet was exposed to ¹⁴CO₂. The amount of transfer ranged from trace up to 90% of the total ¹⁴C incorporated. However, in only half of the clones there was consistent enrichment of the secondary ramet (5 to 89%) suggesting that interramets transfer of carbon may be facultative. Furthermore, when significant export occurred from the primary ramet, it was always principally towards only one ramet even when the clone included more than one. The transfer of ¹⁴C from secondary to primary ramets was shown to be significant only when photosynthesis of the latter was decreased by shading. In this case import of carbon was never more than 60% of the incorporated ¹⁴C.No correlation was found between age or size of the ramets and the intensity of transfer. The shading effect let suppose that transfers are mainly driven by carbon limitation due to changing environmental conditions and not to the state of ramet maturity. The adaptative advantage of such facultative physiological integration between ramets of a clone is discussed.     

Reproductive demography of ramets and genets in a rhizomatous clonal plant <Emphasis Type="Italic">Convallaria keiskei</Emphasis>

Clonal growth occurring below the ground makes it difficult to identify individuals and demonstrate the demographic features of a focal plant species. In this study, genotypically identified ramets of a rhizomatous clonal herb, Convallaria keiskei Miq., were monitored for their growth, survival, and reproduction from 2003 to 2006. After the monitoring period, their subterranean organs were excavated to explore the underground connections of established ramets and the direction of clonal growth. We then combined data on the fate of the monitored ramets with the information of rhizome connections, clarifying reproductive demography at both the ramet and genet levels. Although each ramet initiated both sexual reproduction (via flowering) and clonal growth, clonal growth tended to precede sexual reproduction. In a surveyed genet, 51.0% of ramets produced flowers and 29.6% generated clonal offspring during the study period. Consequently, we clarified the reproductive demography of C. keiskei: clonal growth tended to precede flowering in a ramet, and a genet can keep reproducing every season at the genet level, despite a ramet not having inflorescence every year.     

Carbon allocation during fruiting in Rubus chamaemorus

Background and Aims

Rubus chamaemorus (cloudberry) is a herbaceous clonal peatland plant that produces an extensive underground rhizome system with distant ramets. Most of these ramets are non-floral. The main objectives of this study were to determine: (a) if plant growth was source limited in cloudberry; (b) if the non-floral ramets translocated carbon (C) to the fruit; and (c) if there was competition between fruit, leaves and rhizomes for C during fruit development.

Methods

Floral and non-floral ramet activities were monitored during the period of flower and fruit development using three approaches: gas exchange measurements, ¹⁴CO₂ labelling and dry mass accumulation in the different organs. Source and sink activity were manipulated by eliminating leaves or flowers or by reducing rhizome length.

Key Results

Photosynthetic rates were lower in floral than in deflowered ramets. Autoradiographs and ¹⁴C labelling data clearly indicated that fruit is a very strong sink for the floral ramet, whereas non-floral ramets translocated C toward the rhizome but not toward floral ramets. Nevertheless, rhizomes received some C from the floral ramet throughout the fruiting period. Ramets with shorter rhizomes produced smaller leaves and smaller fruits, and defoliated ramets produced very small fruits.

Conclusions

Plant growth appears to be source-limited in cloudberry since a reduction in sink strength did not induce a reduction in photosynthetic activity. Non-floral ramets did not participate directly to fruit development. Developing leaves appear to compete with the developing fruit but the intensity of this competition could vary with the specific timing of the two organs. The rhizome appears to act both as a source but also potentially as a sink during fruit development. Further studies are needed to characterize better the complex role played by the rhizome in fruit C nutrition.Key words: Allocation pattern, ¹⁴C labelling, carbon translocation, carbon reserves, cloudberry, defoliation, fruit production, gas exchange, Rubus chamaemorus, source–sink relationship, flowering     

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.     

Integration in the clonal plant <Emphasis Type="Italic">Eriophorum angustifolium</Emphasis>: an experiment with a three-member-clonal system in a patchy environment

A clonal plant in heterogeneous environments is usually expected to profit from resource exchange via a clonal network where ramets placed in contrasting environments can specialise so to acquire the most abundant resources. An experiment was designed using the three member clonal system of Eriophorum angustifolium, which consisted of one parent ramet growing in a resource poor environment and two offspring: one was limited in growth by nutrients while the other was light limited; the contrast in availability of limited resources between the offspring ramets was high, medium or none, with the system either connected or severed. The total resource availability was the same in all treatments. We proposed four possible scenarios for the system: offspring ramets will share resources via the deficient parent ramet, and the whole clone will profit from the contrasting environment (scenario 1); offspring ramets will support exclusively the parent ramet, and the whole clone will profit from a homogeneous environment (scenario 2); offspring ramets will stop the export of the limiting resource to the parent ramet, with split and connected treatments not differing (scenario 3); and offspring ramets will exhaust the carbon stored in the biomass of the parental ramet; offspring ramet will profit from connection (scenario 4). In the experiment, the limiting resources were sent to the strongest sink (scenario 2). The parent ramet growing in a deficient environment received the highest support in the treatment where both offspring ramets were growing in the same conditions (no-contrast treatment). Production of new shoots, but not biomass of whole clone, was supported in a homogenous environment. The experiment revealed that multiple stresses might prohibit free exchange of limiting resources via the clonal network and supports the idea that experimental studies on more complex clones are essential for understanding the costs and benefits of clonal growth.     

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.     

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.     

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

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

Effects of resource availability on integration and clonal growth inMaianthemum bifolium

The aim of this study was to investigate whether resource availability affects the degree of physiological integration and the growth pattern of interconnected ramets in the clonal plantMaianthemum bifolium (L.)F.W. Schmidt (Liliaceae), a rhizomatous herb of European forests, by studying it at two contrasting South Swedish beech forest sites termed “poor” and “rich”. The degree of physiological integration was studied by tracing the pattern of¹⁴C translocation and in a cutting experiment involving rhizome severing and defoliation treatments. The size of the plants, growth of new rhizomes, branching frequency, distance between shoots and the internode length were compared. The plants were larger, rhizomes had greater specific mass (mg mm^?1), internodes were shorter and branching frequency higher at the rich site. The cutting treatments reduced the growth of new rhizomes at both sites, and new rhizome segments had lower specific mass in treated plants than in controls, showing the importance of physiological integration for new growth. Translocation of¹⁴C in May showed that the young rhizome tip was a strong sink for carbon. Basipetal translocation to older portions of the rhizome system was greater at the rich site than at the poor site. In September, four months after labelling, the rhizome tips were still strongly labelled with¹⁴C and basipetal translocation had increased at both sites. Plants at the rich site appeared to translocate larger amounts of¹⁴C basipetally than plants at the poor site. It is concluded thatM. bifolium shows a plastic response to resource availability by varying rhizome growth and branching frequency, but the degree of physiological integration is probably only indirectly affected through an increased number of sinks (new rhizome branches) along the ramet system at the rich site.     

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.