Rhizome structure and evolution in the ant-associated epiphytic fern Lecanopteris Reinw. (Polypodiaceae)

GAY, H., 1993. Rhizome structure and evolution in the ant-associated epiphytic fern Lecanopteris Reinw. (Polypodiaceae). The Lecanopteris rhizome is expanded or hollow, and is used as a nest by ants of the genera Iridomyrmex and Crematogaster. The 13 species of Lecanopteris display six rhizome forms, unequally distributed between two subgenera. Subgenus Myrmecopteris comprises four species, each possessing a characteristic rhizome: L. mirabilis has a solid, arched rhizome, with the domatium between the rhizome underside and host trunk; L. sarcopus displays dimorphism between solid frond-bearing axes and hollow, frondless side branches; the rhizome of L. Crustacea is hollow but phyllopodia are solid; L. sinuosa has hollow rhizomes and phyllopodia. The architecture of L. mirabilis, L. sarcopus and L. Crustacea results in a compact, many-layered domatium, but L. sinuosa has a tittle-branched habit. Members of subg. Lecanopteris are completely hollow and have a compact architecture: six species typified by L. pumila have a central gallery and hollow phyllopodia, and three species typified by L. darnaedii have two gallery and chamber systems. The genus Lecanopteris is unlikely to be monophyletic; its nearest relative is Phymalodes. Phylogeny in subg. Myrmecopteris is unclear; no gradation of rhizome complexity exists. In subg. Lecanopteris, L. curtisii is considered most similar to the ancestral species, giving rise to the L. pumila group, which engendered the L. darnaedii group.     

Ant specificity and behaviour in mutualisms with epiphytes: the case of Lecanopteris (Polypodiaceae)

Seven species of the fern Lecanopteris: L. sinuosa, L. sarcopus, L. mirabilis, L. curtisii, L. pumila, L. celebica and L. damaedii are regularly inhabited by ants of five species: Iridomyrmex cordatus, I. murinus, Crematogaster treubi, C. difformis and Camponotus pallidas. Inhabitation is not obligate; either party can survive without the other, but ferns without ants are rare in their natural habitat. The total recorded diversity of ants in Lecanopteris is 31 species, 20 of which were only recorded once.
All five regularly inhabiting ant species kept larvae in the domatium (82% of Lecanopteris specimens examined). The major ant species inhabiting Lecanopteris deposited debris in the domatium, segregated from their brood (90% of Lecanopteris specimens). Iridomyrmex murinus did not build carton around the rhizomes, but the other major inhabitants constructed runways in 79% of Lecanopteris specimens. Goccid cultivation was variable within ant and plant species: I murinus tended them in 50% of specimens, and homopterans were also recorded with Crematogaster treubi (39%), C. bomeensis (27%) and I cordatus (16%). A high frequency of ant colonies inhabiting Lecanopteris , keeping larvae, depositing debris, building carton runways, and occasionally keeping coccids has been established. Specificity of ant species is high within a population of Lecanopteris , or a given habitat or geographical area, but outside the ranges of regular inhabitants other ant occupants are found. Comparable data from other genera of ant-epiphytes with domatia show similar diversity of inhabitance over geographical areas; no data are available within single plant populations.     

Animal-fed plants: an investigation into the uptake of ant-derived nutrients by the far-eastern epiphytic fern Lecanopteris Reinw. (Polypodiaceae)

The epiphytic fern genus Lecanopteris (Polypodiaceae) is regularly inhabited by five species of Iridomyrmex and Crematogaster, which nest and deposit debris in the hollow rhizomes. These epiphytes gain nutrients from ants in two ways: by root absorption from carton runways which surround plants, and by uptake of solutes from ant faeces and debris through the inner rhizome walls. The rhizome cavity surface is black, minutely pitted and bears no specialized absorptive structures. Nutrients containing 14-glucose, 86-rubidium and 32-phosphorus injected into the rhizome cavity were translocated through the plant. Ants inhabiting Lecanopteris were fed glycine and urea containing 15-nitrogen, and this label was incorporated into the fern tissues. Thus ant-derived nutrients can be incorporated into Lecanopteris tissues. The ferns may gain other benefits from the ants such as defence against herbivores, nurture of juveniles or spore dispersal.     

Vegetative propagation and sexual reproduction in the woodland understorey pseudo-annual Circaea lutetiana L.

On the forest floor of deciduous woodlands, relative light intensity gradually declines during the early growing season. The woodland understorey pseudo-annual Circaea lutetiana L. completes its life-cycle at the end of summer. These pseudo-annuals are clonal plants which survive the winter only as seeds and as hibernacles produced by the rhizome apices. In this paper, we asked several questions related to the life-history of C. lutetiana. It was found that shoot formation in early spring did not exhaust the old hibernacle. A trade-off between rhizome + hibernacle number and weight might be expected when plants grow under resource limitation. It was hypothesised that both number and weight of rhizomes and hibernacles will be affected by light availability. Since the effect of resource supply on the size number trade-off will depend on the developmental pattern of the rhizome system, rhizome development was studied as well. Soon after the shoots emerged, 1st order rhizomes were formed in May on the nodes of the old hibernacle. First-order rhizomes branched in June and 2nd order rhizomes (side-branches) were continuously produced throughout the growing season. The phenology and developmental rate of plants growing in different light treatments were plastic. On average 30% of rhizome biomass was formed during the vegetative phase, and rhizome and fruit production were only partly separated in time. The ratio of total rhizome biomass to total fruit biomass was not affected by light. Also flower bud removal did not lead to an increase in rhizome production, which suggests that division of biomass to both reproductive modes is rather rigid. The number of 1st order rhizomes was not affected by the light treatments. Under light limitation, both rhizome number and weight of single rhizomes were reduced. In contrast, fruit number, but not weight of single fruits, was limited.     

Morphogenesis and its relation to architecture in the dimorphic clonal fern Lomagramma guianensis (Aublet) Ghing (Dryopteridaceae)

HEBANT-MAURI, R. & GAY, H., 1993. Morphogenesis and its relation to architecture in the dimorphic clonal fern Lomagramma guianensis (Aublet) Ching (Dryopteridaceae). The complex growth cycle of Lomagramma guianensis results from comparatively simple and uniform morphogenesis. The juvenile sporophyte is terrestrial, and assumes adult morphology upon becoming scandent. A single frondless runner is produced from the base of each monocarpic scandent rhizome base, and ramifies extensively on the forest floor. Moribund runners produce reiterative ramifications identical to the juvenile form, and these repeat the life cycle. All rhizomes share radial apical symmetry which becomes bilateral in differentiated tissues. Phyllotaxy is orthodistichous in sporelings and juveniles, and tristichous in adults. The vascular system of differentiated tissues is a variably stretched, perforated solenostele. The sporeling and inception of adult ramifications have solid protosteles, which become bilateral, vitalized and pierced by gaps resulting from departure of lateral meristem and frond traces. This transition from solid protostele to perforated solenostele facilitates reiteration and shows that a delay exists between formation of lateral meristems and their development. Cryptic extra-axillary meristems occur on adult rhizomes, which remain dormant on scandent rhizomes whilst the apex is alive, and subsequently develop fleetingly before axis death. They develop concomitantly with apex expansion in runners. A single type of lateral meristem produces the variety of adult and reiterated forms, regulated by its position on the axis, parent rhizome vigour and axis orientation.     

Mathematical inference of the underground clonal growth of Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae)

Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae) can spread by sexual or vegetative propagation. The choice of strategy likely depends on the environmental conditions. The rhizome is the organ of vegetative reproduction; hence, it is crucial to understand its development. Unfortunately, it is hardly possible to investigate rhizome morphology directly, since E. helleborine is a protected species in most European countries. The goal of our investigation was to infer the growth patterns of underground parts of an orchid population from long-term annual observations of its aboveground shoots. We implemented the Minimum Spanning Tree method to determine a likely set of underground connections between shoots and to simulate the annual growth of new rhizomes. Furthermore, we modelled the spatial distribution of shoots with a density kernel estimator to compare the density gradients with the direction of growth of the rhizomes. Observed shoot numbers fluctuated between 72 and 183 from year to year. Our results suggest that (1) vegetative reproduction prevails in the studied population, (2) the population consists of about a dozen clones with a diameter of up to 6 m, (3) rhizomes produce up to five new shoots at one branch end per year, (4) rhizomes develop in the direction of decreasing population density, and (5) nodes of rhizomes may produce new offshoots after up to 7 years of dormancy.     

The effects of 1-naphthaleneacetic acid and N6-benzyladenine on the growth of Cymbidium forrestii rhizomes in vitro

An Asiatic orchid, Cymbidium forrestii, was clonally propagated using seed-derived rhizomes as explants. The rhizomes were cultured and proliferated on Murashige and Skoog medium supplemented with various growth substances. Auxins stimulated rhizome growth by increasing branching and fresh weight of the explant, with 1-naphthaleneacetic acid (NAA) being the most effective auxin. All auxins tested suppressed normal shoot formation. The apical meristem of the rhizome reacted to exogenously applied auxin by reducing the cytoplasmic zone of the apical meristem and causing meristem derivatives to rapidly differentiate into vacuolated parenchyma cells. Leaf formation and development was retarded in the presence of auxin. Cytokinins generally reduced rhizome growth and the number of branches, but benzyladenine (BA) can induce shoot formation in vitro. BA induced the cytoplasmic zone of the apical meristem to enlarge and enhanced leaf development. A 5% (w/v) sucrose concentration was most effective in shoot induction when combined with 5 mg1^-1 BA. Activated charcoal promoted rhizome growth; however, shoot formation was inhibited.     

盾叶薯蓣实生苗根状茎的形态发生及薯蓣皂甙积累的研究

对盾叶薯蓣实生苗根状茎的形态发生、发育过程及薯蓣皂甙积累与分布进行了研究。种子萌动后,节部膨大形成球状体,其直径约1．5cm。其胚芽生长锥先后形成4个突起,分别发育形成芽的原分生组织。按其出现的先后分别称为第1芽、第2芽、第3芽和第4芽。第1芽呈剑指形,以后发育为地上缠绕茎,其余3个芽呈丘状突起都分别发育为地下根状茎。有的芽的原分生组织以后还可以形成2个芽的原分生组织,从而使根状茎形成分枝。根状茎顶端的原分生组织由鳞片包被,顶端下方的原表皮内存在初生增厚分生组织。初生增厚分生组织细胞不断向内分裂和其衍生细胞的体积增大,是根状茎能迅速增粗的主要原因。分化完成的根状茎由周皮、基本组织和散生的维管束构成。经组织化学测定,根状茎中薯蓣皂甙主要存在于基本组织的薄壁细胞中,呈液滴状。原分生组织不含薯蓣皂甙,近顶端的基本分生组织细胞内不形成含薯蓣皂甙的液滴。其中,有小型维管束分布的基本组织中薯蓣皂甙的积累与分布最丰富,两年生根状茎中薯蓣皂甙的含量比一年生的高。     

CONTROL OF SHOOT-RHIZOME DIMORPHISM IN THE WOODY MONOCOTYLEDON,CORDYLINE (AGAVACEAE)

In Cordyline terminalis negatively geotropic leafy shoots and positively geotropic rhizomes develop from single axillary buds on either shoots or rhizomes. All axillary buds have similar morphogenetic potential when released from apical dominance. Experiments in which the orientation of the apex is changed, organs removed, or growth regulators applied indicate that after a rhizome is initiated, it is maintained as a rhizome by auxin originating in the leafy shoot. When auxin levels are lowered by changes in the orientation of the axis or shoot removal, the rhizome apex becomes a shoot apex, which appears to be the stable state of the actively growing apex. Benzyl adenine when applied exogenously to the apex or lateral buds has the same effect as lowering the auxin level. Gibberellic acid has no effect on the apex or lateral buds. High levels of exogenous naphthaleneacetic acid cause bud release and development of rhizomes from previously inhibited axillary buds of the shoot. However, it was not possible to convert a shoot apex into a rhizome apex by auxin treatment. It is suggested that the release of buds on the lower side of horizontal branches and of buds directly above a stem girdle is caused by high auxin levels on the lower side or distal to the girdle. The experimental results are discussed in relation to naturally occurring shoot-rhizome dimorphism.     

Some aspects of the extreme anoxia tolerance of the sweet flag,Acorus calamus L.

Acorus calamus L. is a neophyte in Europe with remarkable properties. Among other things, it is the most anoxial tolerant species and a competitive invader at eutrophic sites. The following overview presents the most recent work on these subjects. Carbohydrates of the rhizomes sustain anaerobic ATP production for very long periods. Ethanolic fermentation naturally occurs in winter and produces rather low, but sufficient amounts of ATP for survival, as shown by adenylate energy charge and total adenylate content. Fermentation energy is mainly used for the synthesis and preservation of essential macromolecules, such as proteins and membrane lipids. The extent of these processes is unique. Moreover, ammonia and sulphide uptake is maintained during the cold season. Both ions are detoxified to alanine and thiols which are translocated into the rhizome, where the nitrogen of alanine is used to form arginine. Overwintering leaves contain asparagine instead of arginine. Recycled nitrogen compounds from the rapidly degrading summer leaves return into the rhizomes. Therefore, the nitrogen nutrition consists of an external and internal cycle. The abundance of carbohydrates and nitrogen compounds allows spring shoot growth earlier than other species. These strategies could contribute markedly to the competitive power ofA. calamus at its natural site.     

Phragmites australis at an extreme altitude: Rhizome architecture and its modelling

In central EuropePhragmites australis is a lowland plant, occurring rarely up to the tree line. In the Velká Kotlina cirque (Jeseníky mountains, NE Czech Republic), where it reaches its maximum altitude at about 1350 m a.s.l., its culms are 0.5–0.7 m high and the plants flower only in some years. During the last decade no germinable seeds have been observed. The architecture ofPhragmites rhizomes from this site was studied on seven randomly selected clonal fragments. They consisted of 3 to 10 partial tussocks (clumps) and 4 to 17 green shoots. The total length of the rhizomes was 9.7 to 50 m per plant. The number of nodes per plant was 96 to 431 and the longest internodes were 83 mm long. The number of side branches was 31 to 105 per plant. The branching angle depended on the type of branched rhizome. The mean angles of horizontal rhizomes, which connect individual tussocks, were relatively wide (modus 45°, arithmetic mean 37°), whereas within a tussock much sharper angles of branching prevailed (modal value 5°, arithmetic mean 15°). The mean internode-to-internode angle on continuing rhizomes was about 8°, with a wide variation. An architectural, spatially-explicit model ofPhragmites rhizome growth has been developed, showing that thePhragmites population in the studied locality can be maintained by vegetative multiplication, and seedling recruitment is not needed for its long-term persistence.     

The Hexagonal Branching Pattern of Rhizomes of Alpinia speciosa L. (Zingiberaceae)

Alpinia speciosa L. (Zingiberaceae) is a striking example ofa plant in which an organized architectural structure fulfilsa basic functional requirement - the economical explorationand exploitation of the substrate. This rhizomatous plant spreadslaterally in a predictable manner by vegetative growth to forma clone of separated ramets. A morphological study establishes‘rules of growth’ governing the success and failureof lateral branches (sympodial units), their lengths, and theirrelative angles. The basic branching pattern of this rhizome system approximatesto that of a hexagonal grid - the most economical structuralshape that uniformly ‘samples’ a plane surface.It is shown that the significant deviation of branching anglefound in Alpinia from that required to develop an exact hexagonalnetwork will virtually eliminate aerial shoot ‘clashes’in which the placing of one aerial shoot coincides with thatof another aerial shoot, whilst at the same time enhancing thebenefits of the hexagonal system. The broader implications ofthe organised architecture of plants are emphasised. rhizome, branching pattern, architecture, productivity     

Spatial and temporal distribution of the alkaloid sanguinarine in Sanguinaria canadensis L. (Bloodroot)

Acclimated and non-acclimated potted plants of Sanguinaria canadensis L. were harvested at early and late dormancy, anthesis, and immature and mature fruiting stages. Sanguinarine content and concentration were determined for rhizomes (distal, proximal, and middle sections), roots, leaves, flower, and fruit. Rhizomes had highest sanguinarine content and concentrations, and exhibited decreasing concentration gradients from the distal to proximal third. Concentrations in roots were a tenth of rhizome concentration. Concentrations in leaves, flowers, and fruit were one-thousandth of rhizome Sanguinarine content in whole acclimated plants was constant. Content in whole nonacclimated plants increased as the plant became physiologically active, but was constant during fruit maturation: content in roots, leaves, and fruit did not change. The substantial increase in whole-plant dry weight coupled with the unchanging sanguinarine content during fruit maturation suggests either a shift in photosynthate allocation from defense to growth, or a constant turnover of sanguinarine.     

Seasonal variations in the carbohydrate content of bracken

The amounts of dry matter, mobile carbohydrate and reserve carbohydrate have been determined in bracken fronds and buds, frond-bearing rhizomes and storage rhizomes over a period of 13 months. Dry matter and reserve carbohydrate levels are closely linked, especially in the rhizomes, falling from May to the beginning of July and rising until complete frond death at the end of September; but variations occur between frond-bearing and storage rhizomes. The carbohydrate reserve of frond-bearing rhizome is rapidly exhausted by the developing fronds and further carbohydrate is drawn from the storage rhizomes, but a concurrent fall in mobile carbohydrate suggests that the rate of conversion of reserve carbohydrate is slower than the rate of translocation to the fronds. By August, there is an accumulation of carbohydrate in the frond-bearing rhizome, which is followed by a fall in September and this again suggests low enzyme activity in the storage rhizome, but also that sink strengths in bracken are in the order (1) developing buds, (2) storage rhizome, (3) frond-bearing rhizome.
The responses of bracken to cutting and herbicide-application are discussed in relation to sink strengths and a low level of enzyme activity in the storage rhizome.     

Flower removal increases rhizome mass in natural populations of Alstroemeria aurea (Alstroemeriaceae)

Plant architecture and phenotypic plasticity under natural conditions remain little known for many rhizomatous species. This study evaluates, in situ, the plastic responses of Alstroemeria aurea plants from three Patagonian populations to flower or flowering-shoot removal. The size and architecture of treated and untreated plants were assessed. Nutrient contents (N, P and K) were evaluated for rhizomes and roots developed in two successive years. Those plants that were deprived of their inflorescences developed, on average, a heavier rhizome than both control plants and plants from which flowering shoots had been removed. Neither of the two treatments applied altered the number of metamers or the branching pattern of the rhizomes. The contents of N, P and K were higher in rhizomes than in roots. In summer, nutrients were more concentrated in inflorescences and the new rhizome segment than in the rhizome segment developed in the previous year. The idea that fruiting failure in A. aurea promotes resource re-assignment from aerial shoots to rhizomes without altering the architecture of plants is supported. The development of the underground portion of aerial shoots in late summer-autumn allows A. aurea plants to take full advantage of short growth periods, but would impose a limit to plasticity.     

Do spatial patterns of clonal fragments and architectural responses to defoliation depend on the structural blue-print? An experimental test with two rhizomatous Cyperaceae

Clonal architecture is involved in performance of clonal fragments, as it determines spatial distribution of ramets. It is expected to rely on the species-specific expression of several architectural traits (structural blue-print). However, in contrasting environments, realized clonal architectures may differ, due to phenotypic plasticity. In this paper, we compared clonal architectures between two rhizomatous ecologically close Cyperaceae (Carex divisa and Eleocharis palustris) in non-defoliated and defoliated conditions. Two questions were addressed. (1) How much do the structural blue-print and resulting colonization and occupation of space differ between both species? (2) Does the structural blue-print constrain plastic responses of clonal architecture to defoliation? Traits related to performance, spatial pattern, architecture and biomass allocation of clonal fragments were monitored through an original non-destructive mapping method. In non-defoliated conditions, both species showed similar biomass but contrasting architectures and patterns of biomass allocation to rhizomes that resulted in different spatial patterns. The rhizome network of C. divisa, which consisted in only two primary rhizomes but several branches, was involved in resource storage rather than in spatial colonization. Conversely, E. palustris produced on average six primary rhizomes that grew in the whole horizontal plane, maximizing both occupation and colonization of space. These differences in structural constraints coupled with allometric relationships, resulted in differential responses to defoliation. In C. divisa, the costs associated to defoliation caused a decrease in branching, limiting the area occupied and number of ramets produced by clonal fragments, but increasing ramet density. Conversely, the weakly branched rhizome network of E. palustris was not affected by defoliation. Both spatial strategies (consolidation vs. colonization) are likely to provide ecological advantages allowing their coexistence in grazed meadows.     

Experimental demography of rhizome populations of establishing clones of Solidago altissima

1 Solidago altissima forms large compact below-ground rhizome systems. The rhizome systems of 30 genets in a population that had colonized an old-field in 1984 were mapped yearly in situ from 1988 to 1992. Branching angles, lengths and survival of rhizomes, and production of daughter rhizomes, were calculated from digitized maps.
2 We hypothesized that genets of S. altissima would prevent intraclonal crowding by producing longer but fewer rhizomes as they increased in size. In addition, we tested the effects of environmental factors on rhizome dynamics by (i) mowing in late summer, (ii) cutting of rhizome connections, and (iii) removal of close neighbour plants.
3 Most rhizomes were initiated at an angle of 60°−65° relative to the parent rhizome. Subsequent growth tended towards the parent axis and towards more centrifugal growth. Rhizome angles were not influenced by any treatment.
4 In control genets, rhizome density increased 1.3-fold from March 1988 to March 1992. The number of newly produced rhizomes per genet decreased and their length increased from 1988 to 1992. Longer rhizomes had a higher survival probability than shorter rhizomes did. The number of newly produced rhizomes per parent rhizome was positively correlated with mean rhizome length per genet.
5 Mowing strongly decreased rhizome density from 1988 to 1992, mainly due to reduced production of new rhizomes.
6 Rhizome cutting led to a slight decline in the rhizome population and to shorter lengths of rhizomes in their second year compared with controls.
7 The removal of close neighbour plants led to increased rhizome densities. These rhizomes grew longer than controls in unmown plots and shorter in mown plots and had higher probabilities of surviving into their second year.
8 Yearly mowing could successfully prevent the further spread of S. altissima because it acts on rhizome and shoot dynamics as well as on seed output.     

Rhizome Growth and Clone Development in Anemone nemorosa L.

Studies of natural populations indicate that clones grow radially,soon developing a hollow centre which may be invaded by newgenotypes. In the populations studied, individual rhizomes elongate byan average of 2.5 cm per year. Growth is sympodial, the terminalbud forming an aerial shoot and further elongation occurringfrom an axillary bud. The oldest rhizome segment found was 7 years with most rhizomesbeing 3 or 4 years old. Because there is no apparent senescence,individual clones may persist for very long periods, possiblyhundreds of years. A computer model of rhizome growth was used to postulate theextent of intermixing of old clones. It indicated that old clonesbecome very fragmentary and totally inmixed with their neighbours,one ‘clump’ often representing several clones. To supplement the model, plants from a woodland with an unbrokencarpet of Anemone nemorosa were collected, 27 morphologicalcharacters measured and the data analysed using principal componentsanalysis and cluster analysis. These supported the hypothesisthat old clones are large and fragmentary with complete intermixing. The possible role of soil factors in influencing rhizome extensionis discussed and the strategy of A.nemorosa is compared withthose of its close associates in the pre-vernal woodland herblayer. Anemone nemorosa L., wood anemone, rhizome growth, clone development, computer model, vegetative reproduction     

The architecture of a dimorphic clonal fern, Lomagramma guianensis (Aublet) Ching (Dryopteridaceae)

Lomagramma guianensis possesses a dimorphic rhizome and trimorphic fronds. The juvenile genet consists of a slender, sinuous, unbranched, terrestrial rhizome which bears pinnatifid fronds. The adult rhizome form is initiated when the apex becomes scandent. It is stout, exhibiting rhythmic production of pinnate, adult-type fronds, and probably rhythmic growth. The scandent rhizome is ultimately monocarpic, producing fertile acrostichoid fronds and subsequendy dying. Terrestrial runners are produced from proleptic buds at the scandent rhizome base. These are largely similar in morphology to the scandent rhizomes, but display different behaviour. The runners are relatively ephemeral structures which produce no fronds. They branch proleptically from their base to produce scandent rhizomes, and also along their length to give rise to a complex system of runnerS. Senescent runners which have lost contact with their parent scandent rhizome produce reiterative complexes which represent a return to the juvenile form. These reiterated rhizomes behave identically to the juvenile genets, and repeat the entire cycle of development. The architectural history of the clonal pteridophyte Lomagramma is compared to that of angiosperm trees; interesting similarities and differences are found.     

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.