Terminal Motility in Elongating Stolons of Proboscidactyla flavicirrata

Proboscidactyla flavircirrata is a commensal hydroid whose stolonselongate rapidly in the absence of its host. Growth pulsationscomparable to those of other hydroids originate endogenouslyin each stolon tip, although influenced by the to and fro movementsof the hydroplasm. Neither growth pulsations nor elongationappear to depend upon cellular proliferation, because the distalends of stolons lacked mitotic figures and since irradiationsufficient to halt mitosis failed to stop stolon elongation.Cellular migration is also apparently unessential for elongationbecause severed stolon tips, isolated from potential sourcesof reusable cells, nevertheless elongated. Marking experimentssuggested that cells changed shape rather than location as severedtips advanced. In old preparations the stolons became attenuatedbecause the tips continued to progress without adequate recruitmentof cells. Stolons deprived of their tips failed to elongatealthough they were still connected to the remainder of the colonyand contained mitotic cells. Cytochalasin B simultaneously haltedgrowth pulsations and stolon elongation in a reversable manner.Apparently stolon tip locomotion in Proboscidactyla is essentialto the elongation process.     

Mechanics of growth pulsations as the basis of growth and morphogenesis in colonial hydroids

Growth and shaping in colonial hydroids (Hydrozoa, Cnidaria) are realized due to the functioning of special colony elements, growing tips located at the terminuses of branched colony body. Unlike in plants, the growing tips of colonial hydroids are sites of active cell movements related to morphogenesis and lacking proliferation. The activity of hydroid growing tips is expressed as growth pulsations: cyclic repetitions of their apex extensions and retractions. The parameters of growth pulsations are species specific and related to the shape of a forming element. Here, the succession of cell movements and changes in mutual arrangement within the growing tip are described in detail at all pulsation phases. The role of the inner cell layer in the tip activity was demonstrated for the first time. Relationships between the growing tip parameters, length and diameter, and pulsations are discussed. A scheme is proposed for cyclic processes in both epithelial layers. An explanation is provided for the two-step mode of growth pulsations with relative independence of the main phases. It was proposed that successive activities of the tip ecto- and endoderm serve as driving forces provided there is a hard outer skeleton. This scheme makes it possible to explain some patterns of growth and morphogenesis in colonial hydroids, such as gradually increasing growth rate of a new tip and its maximum growth rate, differences in the parameters of growth pulsations between shoot and stolon tips, shoot base inclination towards the stolon tip, etc., and provides a basis for further improvement of the model of morphogenesis in hydroids.     

Mechanics of growth pulsations as the basis of growth and morphogenesis in colonial hydroids

Growth and shaping in colonial hydroids (Hydrozoa, Cnidaria) are realized due to the functioning of special colony elements, growing tips located at the terminuses of branched colony body. Unlike in plants, the growing tips of colonial hydroids are sites of active cell movements related to morphogenesis and lacking proliferation. The activity of hydroid growing tips is expressed as growth pulsations: cyclic repetitions of their apex extensions and retractions. The parameters of growth pulsations are species specific and related to the shape of a forming element. Here, the succession of cell movements and changes in mutual arrangement within the growing tip are described in detail at all pulsation phases. The role of the inner cell layer in the tip activity was demonstrated for the first time. Relationships between the growing tip parameters, length and diameter, and pulsations are discussed. A scheme is proposed for cyclic processes in both epithelial layers. An explanation is provided for the two-step mode of growth pulsations with relative independence of the main phases. It was proposed that successive activities of the tip ecto-and endoderm serve as driving forces provided there is a hard outer skeleton. This scheme makes it possible to explain some patterns of growth and morphogenesis in colonial hydroids, such as gradually increasing growth rate of a new tip and its maximum growth rate, differences in the parameters of growth pulsations between shoot and stolon tips, shoot base inclination towards the stolon tip, etc., and provides a basis for further improvement of the model of morphogenesis in hydroids.     

The role of polyp-stolon junctions in the redox signaling of colonial hydroids

An encrusting colonial hydroid can be regarded as a network of polyps or ‘mouths’ connected by tube-like stolons. The success of the colony crucially depends on putting these mouths where the available food is. Feeding-related perturbations may provide important signals in this regard. After feeding, polyps contract regularly, dispersing food throughout the colony via the gastrovascular fluid. Mitochondrion-rich epitheliomuscular cells concentrated near polyp-stolon junctions likely drive these contractions. Putatively, the redox state of these cells may influence colony-level form. For instance, the metabolic demand associated with feeding-related contractions results in mitochondria that have relatively oxidized electron carriers and produce lesser amounts of reactive oxygen species (ROS). ROS or other redox-sensitive molecules emitted from polyp-stolon junctions into the gastrovascular fluid may provide stolons with signals influencing elongation, branching, and regression. Treatments of colonies with anti-oxidants cause peripheral stolon tips to rapidly regress. This regression appears to be an active process involving a flux of locally produced peroxides and cell and tissue death. At the same time, polyps and stolon tips in the center of treated colonies remain healthy. ‘Sheet-like’ growth of short, branched stolons ensues. Signals that inhibit the outward growth of stolons may lead by default to the concentrated growth of stolons and polyps in food-rich areas. ROS may mediate signaling mechanisms involving nitric oxide, programmed cell death, a variety of redox-regulated proteins, or all of these.     

The growth and degeneration of the hydroid Sertularia perpusilla,an obligate epiphyte of leaves of the seagrass Posidonia oceanica

The hydrorhizae of Sertularia perpusilla colonies that originate from stolon attachments initially grow up and down Posidonia leaves. After a short time the distal hydroid tissue degenerates concurrently with the downward growth of the proximal hydrorhiza onto younger sections of the leaves. Consequently the hydroids move down the leaves which grow upward from a basal meristem. In situ inversion of leaves had no effect on either the orientation or the rates of growth and degeneration. This indicates that orientation may be in response to age related features of leaf tissue rather than to light, gravity or water movement, the directions of which were reversed by inverting the leaves. However, orientation of growth of new hydroids from attached stolons is immediate, probably before the hydroid colony is long enough to be able to detect an age gradient. This paradox could be explained if initial orientation were based on the direction of gravity or light but subsequent growth were directed along the age gradient. For hydroids in the size range studied there was a relation between growth rate and number of hydranths. Thus growth rate may be limited by food rather than by there being only one growing tip of the hydrorhizae.     

Relay-ray way of hydroplasm movement in hydroid colonies

In this study, we continued the investigation of the distribution system of colonial hydroids in the course of its development, starting with its emergence during the planula metamorphosis and ending with the formed colony. The hydroplasmic stream system of two species of colonial hydroids—Perigonimus abyssi G.O. Sars, 1874, and Stauridia producta Wright, 1858—was studied. We found that the main principle by which hydroplasma moves in these colonies, which form no shoots, is the relay-race from one hydranth to another through a stolon fragment connecting them or it moves past the neighbouring hydranth to the next one, etc. We show that the efficiency of the conducting (distribution) system does not depend on the level of complexity of the colonial structure. The results of this study confirm the absence of general colonial processes of integration and self-regulation in hydroid polyps.     

Cell Movements Associated with Terminal Growth in Colonial Hydroids

A series of studies into the nature or leimiual growth movementsin hydroids (chiefly the thecate Campanularia) is reviewed.Both stolon and pedicel tips of all hydroids observed grow byan endless repetition of "growth cycles." In thecate stolons,the geometry of the tip's excursions is relatively simple andsuccessive cycles are predictable in their duration and percycle growth. Conversely, athecate stolons are much more complexor variable in each of these respects. In Campanularia bothcycle time and per cycle growth differ among genetic stocks. Such environmental or environment-related factors as temperature,organic contaminants in the sea water, and the nutritive conditionof the colony each affect cycle time and/or per cycle growth.Also, ingestion of a large meal temporarily inhibits the retractionsthat characterize most growth cycles. Growth movements are a property of the stolon tip, since isolatedtips less than 0.5 mm long move in basically normal fashion.However, two aspects of the cycle are, modified by more distantregions: the cycle is accelerated by a pacemaker lying 0.5 to2.0 mm from the tip, and the extent of each cycle's retractionphase is determined by intrastolonic pressure generated by hydroplasmicflow. Certain neuroinhibitor drugs duplicate in intact stolonsthe cycle changes seen in surgical isolates. Synchronized with each growth cycle is a pattern of epidermalthickening and thinning at the stolon tip, this layer beingthinnest just after the crest and thickest about two-thirdsof the way through the cycle. Cells behind the tip also movein relation to the growth cycle, generally approaching the tipduring epidermal thickening and retreating during epidermalthinning. Several types of metabolic inhibitors markedly affect the stolonicgrowth movements, indicating the need of oxidative metabolismand of protein synthesis using a short-lived mRNA for the tip'sactivities. Campanularia pedicel growth cycles differ from stolon growthcycles in having a shorter and more variable duration, a lesserper cycle growth, and a shallower retraction phase, with changesin these features along the length of the pedicel correlatedwith changes in the type of growth occurring. These studies and those of other investigators are related,their contributions to our understanding of elongation are discussed,and persisting problems are highlighted.     

Anchoring cells (Desmocytes) in the hydrozoan polyp Cordylophora.

Desmocytes or anchoring cells are present on the upright stolons of the athecate hydroid Cordylophora caspia and function to support the soft coenosarc within the rigid tube of perisarc by linking the perisarc with the mesoglea. These cells are characterized by accumulations of 70 A filaments which aggregate into dense rods at the apical end and contact the perisarc. At the base of the desmocytes the filaments are distributed within large cytoplasmic processes which interdigitate with an extension of the mesoglea. Desmocytes in Cordylophora are temporally and spatially formed in sequence as the upright elongates. Depending on their location and structure they can be categorized as forming, functional, or remnant desmocytes. The youngest, forming desmocytes are found in the distal end of the stolon 0.5-1.0 mm from the base of the hydranth. In this region coenosarc is just beginning to separate from the perisarc. Functional desmocytes are scattered 1-3 mm from the base of the hydranth and are associated with perpendicular extensions of the mesoglea. Remnants have lost their mesogleal connection and are located in more proximal, older regions of upright stolon. Support provided by the desmocytes to the upright stolon is limited by three factors that characterize the athecate hydroid: distribution of perisarc, pattern of growth, and extent of movement. The distal location of forming desmocytes is coincident with the hardening of new perisarc. The temporary nature of attachment sites is directly related to upright elongation. It is probable that the orientation of filaments within the cell and the mesogleal extension provide an addition feature of flexibility necessary to permit feeding, growth, and rhythmic pulsation movements characteristic of these hydroids.     

The Selective Myosin II Inhibitor Blebbistatin Reversibly Eliminates Gastrovascular Flow and Stolon Tip Pulsations in the Colonial Hydroid Podocoryna carnea

Blebbistatin reversibly disrupted both stolon tip pulsations and gastrovascular flow in the colonial hydroid Podocoryna carnea. Epithelial longitudinal muscles of polyps were unaffected by blebbistatin, as polyps contracted when challenged with a pulse of KCl. Latrunculin B, which sequesters G actin preventing F actin assembly, caused stolons to retract, exposing focal adhesions where the tip epithelial cells adhere to the substratum. These results are consistent with earlier suggestions that non-muscle myosin II provides the motive force for stolon tip pulsations and further suggest that tip oscillations are functionally coupled to hydrorhizal axial muscle contraction.     

Shoot Dynamics and Adaptive Morphology of Ipomoea phillomega (Vell.) House (Convolvulaceae), a Tropical Rainforest Liana

Ipomoea phillomega (Vell.) House (Convolvulaceae) is a commonliana in undisturbed and disturbed patches of a primary tropicalrain forest in Veracruz, Mexico. In this study a demographic-morphologicalapproach to an understanding of its growth dynamics was attempted.Clonal spread via stolons accounts for nearly all populationrecruitment. The liana can produce a variety of shoot types.The balance among these is influenced by light intensity suchthat shaded conditions favour the production of creeping shoots(stolons) with high elongation rates, long internodes and short-livedrudimentary leaves, and in full sunlight most shoots twine,elongate more slowly, have short internodes and produce largelong-lived leaves. The correlation between light intensity andshoot type is not absolute. Even in deep shade of primary forestsome twiners are produced and in full sunlight some stolonsare produced. This suggests that over a span of light intensityranging from 0.4–100 per cent of full sunlight, the lianainvests energy both in vertical climbing and horizontal spread.Different shoot types showed different patterns of survivorshipof plant parts. Twiner leaves have higher mortality rates thanstolon leaves, but twiner apices have higher mortality ratesthan stolon apices. Exploration by essentially leafless stolonsrepresents growth subsidized by insolated canopies elsewherein the shoot system. The ‘costs’ of this patternof growth are unknown since the physiological relationshipsamong the shoot types have not been studied.     

Multicellular redox regulation in an early-evolving animal treated with glutathione

Redox signaling has emerged as a unifying theme in many seemingly disparate disciplines. Such signaling has been widely studied in bacteria and eukaryotic organelles and is often mediated by reactive oxygen species (ROS). In this context, reduced glutathione (GSH) acts as an important intracellular antioxidant, diminishing ROS and potentially affecting redox signaling. Complementing this cell-level perspective, colonial hydroids can be a useful model for understanding organism-level redox signaling. These simple, early-evolving animals consist of feeding polyps connected by tubelike stolons. Colonies treated exogenously with GSH or reduced glutathione ethyl ester (GEE) were expected to show a morphological change to sheetlike growth typical of low levels of ROS. Contrary to expectations, diminished stolon branching and polyp initiation was observed. Such runnerlike growth is associated with higher levels of ROS, and surprisingly, such higher levels were found in GSH- and GEE-treated colonies. Further investigations show that GSH triggered a feeding response in hydroid polyps, increasing oxygen uptake but at the same time relaxing mitochondrion-rich contractile regions at the base of polyps. Diminished gastrovascular flow and increased emissions of mitochondrial ROS also correlated with the observed runnerlike growth. In contrast to cell-level, "bottom-up" views of redox signaling, here the phenotype may arise from a "top-down" interaction of mitochondrion-rich regions and organism-level physiology. Such multicellular redox regulation may commonly occur in other animals as well.     

Advanced Cambrian hydroid fossils (Cnidaria: Hydrozoa) extend the medusozoan evolutionary history

Primitive cnidarians are crucial for elucidating the early evolution of metazoan body plans and life histories in the late Neoproterozoic and Palaeozoic. The highest complexity of both evolutionary aspects within cnidarians is found in extant hydrozoans. Many colonial hydrozoans coated with chitinous exoskeletons have the potential to form fossils; however, only a few fossils possibly representing hydroids have been reported, which still require scrutiny. Here, we present an exceptionally well-preserved hydroid found in the Upper Cambrian Fengshan Formation in northern China. It was originally interpreted as a problematic graptolite with an uncertain systematic position. Based on three characteristic morphological traits shared with extant hydroids (with paired hydrothecae, regular hydrocaulus internodes and special intrathecal origin pattern of hydrocladium), we propose this fossil hydroid as a new genus, Palaeodiphasia gen. nov., affiliated with the advanced monophyletic hydrozoan clade Macrocolonia typically showing loss of the medusa stage. More Macrocolonia fossils reviewed here indicate that this life strategy of medusa loss has been achieved already as early as the Middle Devonian. The early stratigraphical appearance of such advanced hydroid contrasts with previous molecular hypotheses regarding the timing of medusozoan evolution, and may be indicative for understanding the Ediacaran cnidarian radiation.     

Structure and signaling in polyps of a colonial hydroid

Abstract. After feeding, polyps of colonial hydroids contract regularly, dispersing food throughout the colony via the gastrovascular fluid. Such contractions may trigger signaling pathways that allow colonies to grow in an adaptive manner, i.e., to initiate development of more polyps in food‐rich areas and to suppress polyp development in food‐poor areas. In this context, we investigated the structure and potential signaling of the junction between polyps and stolons in colonies of the hydroid Podocoryna carnea. Using transmission electron microscopy, we found that the density of mitochondrion‐rich epitheliomuscular cells was low in polyp and stolon tissues except at or near the polyp‐stolon junction, where many of these mitochondrion‐rich cells occur in ectodermal tissue. In vivo fluorescence microscopy suggests that these mitochondria are a principal source of the metabolic signals of the colony. Both native fluorescence of NAD(P)H and fluorescence from peroxides (visualized with H₂DCFDA) co‐localize to this region of the polyp. Rhodamine 123 fluorescence suggests that both these metabolic signals emanate from mitochondria. To test whether such metabolic signals may be involved in colony pattern formation, inbred lines of P. carnea were used. Colonies of a runner‐like inbred line grow with widely spaced polyps and long stolonal connections, much like wild‐type colonies in a food‐poor environment. Colonies of a sheet‐like inbred line grow with closely spaced polyps and short stolonal connections, similar to wild‐type colonies in a food‐rich environment. Polyp‐stolon junctions in runner‐like and sheet‐like colonies were imaged for the fluorescence of H₂DCFDA. Densitometric analysis of this signal indicates that the mitochondria in epitheliomuscular cells of runner‐like polyps emit greater amounts of peroxides. Because peroxides and other reactive oxygen species are frequently intermediaries in metabolic signaling pathways, we suspect that such signaling may indeed occur at polyp‐stolon junctions, affecting colony pattern formation in these inbred lines and possibly in hydroid colonies in general.     

Tissue compatibility in regenerating explants from the colonial marine hydroid Hydractinia echinata (Flem.)

Artificially maintained colonies of the colonial marine hydroid Hydractinia echinata are endogenously limited in lateral growth and usually develop in a circular pattern with a closed periderm. Further lateral growth consists of free stolons. Preliminary studies of tissue compatibility in regenerating explants indicate that fusion generally fails to occur if the opposing explants are of differing sex, from different individuals of the same sex, or if peripheral explant contact occurs after free stolons have begun to form. Of the three aforementioned causes of fusion failure, the most important factor appears to be the timing of peripheral explant contact. If regenerating explants contact each other before the endogenous circular growth pattern is achieved, fusion will occur regardless of sex and/or individuality. Incompatibility between explants from the same individual may be because of periderm development inhibition in some instances. However, the failure of free stolon fusion between explants from the same individual suggests the development of “temporal specificity” once the endogenous limit of lateral growth is achieved.     

Quantitative measures of gastrovascular flow in octocorals and hydroids: toward a comparative biology of transport systems in cnidarians

Using microscopy, the gastrovascular systems of four hydroids (Eirene viridula, Cordylophora lacustris, Hydractinia symbiolongicarpus, and Podocoryna carnea) and two distantly related alcyonacean octocorals (Acrossota amboinensis and Sarcothelia sp.) were examined and compared within a phylogenetic framework. Despite a range of stolon widths (means 53–160 μm), the hydroid species exhibited similar patterns of gastrovascular flow: sequentially bidirectional flow in the stolons, driven by myoepithelial contractions emanating from the center of the colony. Unlike the hydroids, the gastrovascular system of A. amboinensis (mean stolon widths for 5 colonies, 0.57–1.21 mm) exhibited simultaneously bidirectional flow with incomplete, medial baffles (width 4–20 μm) separating the flow. Baffles visualized with transmission electron microscopy consisted of endoderm, mesoglea, and occasionally another layer of tissue. Mean flow rates of the gastrovascular fluid for seven stolons ranged from 125 to 275 μm s^?1, with maximum rates of 225–700 μm s^?1. In Sarcothelia sp., stolons were of comparable width (means for 13 colonies 0.55–1.4 mm) to those of A. amboinensis. These stolons, however, were divided by several partitions (width 8–25 μm), both complete and incomplete, which were spaced every 100.5±5.1 μm (mean±SE; range 27.1–283.7 μm) and appeared structurally similar to baffles. In lanes defined by these partitions, ciliary motion was visible in image sequences, and flow was unidirectional. Within a single stolon, flow moved in different directions in different lanes and changed direction by moving from lane to lane via occasional spaces between the partitions. Mean flow rates for 30 stolons ranged from 75 to 475 μm s^?1, with maximum rates of 85–775 μm s^?1. For both octocorals, flow rates of the gastrovascular fluid were not correlated with the width of the stolon lumen. While octocoral gastrovascular systems probably exhibit differences based on phylogenetic affinities, in all species studied thus far, gastrovascular flow is entirely driven by cilia, in contrast to the hydroid taxa.     

Nutritional physiology and colony form in Podocoryna carnea (Cnidaria: Hydrozoa)

Abstract. We compared growth rates and final morphological states of the athecate colonial hydroid Podocoryna carnea in two nutritional environments: one varying the quantity of food provided at a fixed interval and the second varying the time between feedings of a fixed quantity. In both environments, replicate colonies were either fed uniformly, or fed on only one side and starved on the other. In addition, we fed colonies fluorescence-labeled cultures of Artemia salina and documented the subsequent distribution of label. We found that both the growth rates and the final morphological state varied logarithmically with food supply. Heterogeneous feeding had a marked effect on colony morphology, with a sharp boundary in polyp number, stolon density, and polyp size forming at the fed–unfed interface. The distribution of fluorescence was correlated with sites of colony growth. These results confirm and extend early work on the priority of growth zones in colonial hydroids, and present new challenges for understanding the relationship among energy metabolism, gastrovascular circulation, and colony form.     

Influence of Ethylene and Kinetin on Tuberization and Enzyme Activity in Solanum tuberosum L.: Stolons Cultured in vitro

The response of potato stolons, cultured in vitro, to ethylenewas investigated utilizing 2-chloroethyl-phosphonic acid asthe source of ethylene. Concentrations of 0.067, 0.67, 6.7,and 67 µm did not promote tuber formation which occurredin the presence of 16 µm kinetin. In the presence of 2-chloroethyl-phosphonicacid stolon branching was promoted and they maintained a diageotropicgrowth habit in contrast to the negatively geotropic growthhabit of kinetin-treated cultures. The amount of soluble sugars accumulating at the stolon tipwas similar in the presence of kinetin and 2-chloroethylphosphonicacid. However, the level decreased in kinetin-treated stolonsas starch synthesis increased whereas a high specific activitywas maintained in the presence of 2-chloroethylphosphonic acidwhere no starch synthesis occurred. Higher levels of sucroseoccurred in kinetin-treated cultures. Both 2-chloroethylphosphonic acid and kinetin decreased invertaseactivity at the stolon tip. Peroxidase activity increased withtime in response to 2-chloroethylphosphonic acid whereas activityincreased appreciably in kinetin-treated stolons only afterday 5. Generally higher levels of 3'nucleotidase activity existedin 2-chloroethylphosphonic acid-treated stolons. RNase activitydecreased in kinetin treated stolons while activity increasedin the presence of 2-chloroethylphosphonic acid. It is suggestedthat ethylene may influence stolon growth but may not be directlyinvolved in tuber initiation.     

Mitochondria as integrators of information in an early-evolving animal: insights from a triterpenoid metabolite

Mitochondria have the capacity to integrate environmental signals and, in animals with active stem cell populations, trigger responses in terms of growth and growth form. Colonial hydroids, which consists of feeding polyps connected by tube-like stolons, were treated with avicis, triterpenoid electrophiles whose anti-cancer properties in human cells are mediated in part by mitochondria. In treated hydroids, both oxygen uptake and mitochondrial reactive oxygen species were diminished relative to controls, similar to that observed in human cells exposed to avicins. While untreated colonies exhibit more stolon branches and connections in the centre of the colony than at the periphery, treated colonies exhibit the opposite: fewer stolon branches in the centre of the colony than at the periphery. The resulting growth form suggest an inversion of the normal pattern of colony development mediated by mitochondrial and redox-related perturbations. An as-yet-uncharacterized gradient within the colony may determine the ultimate phenotypic effects of avicin perturbation.     

Aspects of the Comparative Physiology of Ranunculus bulbosus L. and Ranunculus repens L.: II. Carbon Dioxide Assimilation and Distribution of Photosynthates

Detailed analysis of the interrelationships between sourcesof photosynthate production and sites of utilization in thetaxonomically closely related species Ranunculus bulbosus L.and R. repens L. showed that leaves whether present on rosette,stem, or stolon had similar levels of ¹⁴CO₂-fixation but thepattern of distribution of radiocarbon to the rest of the plantdiffered. Fruits of R. bulbosus had a lower fixation rate thanleaves but were characterized by total retention of the fixedradiocarbon. Rosette leaves of R. bulbosus supplied the youngleaves, developing apices in the rosette, roots, and corms,whereas the labelled assimilates from cauline leaves were evenlydistributed between reproductive and vegetative parts. The cormwas the major sink both at the flowering and fruiting stages.When plants were treated with ¹⁴CO₂ in the field even higherlevels of radiocarbon moved into the corm than in comparableexperiments under greenhouse conditions. The rosette leaf ofR. repens exported mainly to actively growing stolons in plantswith many stolons bearing rooted ramets although growth of astolon was also substantially supported by photosynthates producedby its own ramets. A proportion of the radiocarbon fixed byleaves of mature ramets was exported and moved in a predominantlyacropetal direction into the stolon apex, stolon axis, and youngramets of the same stolon. The stock in R. repens had a muchlower demand for assimilates than the corm in R. bulbosus. The results are consistent with the concept that R. bulbosusoperates a conservative policy involving the replacement ofthe parent in situ by a daughter from the corm, coupled withextensive fruit production. In R. repens the emphasis is onlateral spread and exploitation of substantial areas of groundby vegetative spread and replacement of the parent by daughtersmany of which may occupy sites some distance from the parent.