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     

Phenological regulation of opportunities for within-inflorescence geitonogamy in the clonal species,iris versicolor (iridaceae)

Opportunities for selfing through geitonogamy are possible if more than one flower within the same clone, inflorescence, or floral unit is open at the same time. In a total of 200 inflorescences in two natural populations of Iris versicolor, flowers were observed and classified daily on the basis of anther dehiscence and stigma receptivity. Analysis of the flowering phenology demonstrated that (1) protandry limits opportunities for autogamy, (2) flowers mature sequentially within a floral unit (defined as a cluster of flowers borne on a single branch within an inflorescence), preventing the opportunity for geitonogamous fertilization between flowers of the same floral unit, and (3) 77% of all flowers had no opportunity to be pollinated by another flower within the same inflorescence. Both the number and the proportion of flowers with opportunities for geitonogamy increased with the number of flowers open in each population, and opportunities for geitonogamy also increased with the number of floral units within inflorescences. These morphological and phenological controls suggest that when selfing occurs in this species, it is most likely to occur between flowers on different inflorescences within the same clone. Since the organization of whole inflorescences in space is determined primarily by rhizome placement, clonal architecture may play an important role in mating system regulation in this species.     

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.     

Regulation of clonal growth and dynamics of Panicum virgatum (Poaceae) in tallgrass prairie: effects of neighbor removal and nutrient addition

Clones of the perennial grass Panicum virgatum were studied on the Konza Prairie in northeast Kansas to determine the effects of neighbors, nutrient availability, and physiological integration on ramet population dynamics and clonal growth and architecture. Opposite halves of established clones in the field were subjected differentially to treatments including neighbor removal and nitrogen addition, with intact or severed rhizome connections between halves. Neighborhood competition strongly influenced clone architecture and expansion rates. Removal of neighbors resulted in a >;95% increase in radial clone expansion, intraclonal ramet densities, ramet population growth rates, ramet biomass, and percent of stems flowering, averaged over a 4-year period relative to halves or clones with intact competitors. Plant responses suggest that effects of interclonal neighbors are mediated through alteration of the light environment in the clone canopy and water availability. Addition of nitrogen did not affect lateral spread or clone structure, but resulted in significant increases in ramet size, flowering, and seed production. ANOVA revealed no significant effect of rhizome severing or treatment × severing interactions, suggesting that the size of the integrated physiological unit is much smaller than clone size and/or that physiological integration had no effect on clone responses to environmental heterogeneity at the scale of the diameter of established clones.     

Within population genetic differentiation in traits affecting clonal growth: Festuca rubra in a mountain grassland

Festuca rubra , a clonal grass of mountain grasslands, possesses a considerable variability in traits related to spatial spreading (rhizome production, length and branching; tussock architecture). Since these traits highly influence the success of the species in a spatially heterogeneous system of grasslands, a combined field and growth chamber approach was adopted to determine the within-population variation in these parameters. Clones were sampled in a mountain grassland (The Krkono?e Mts., Czech Republic); the environment (mean neighbour density) of individual clones varied highly. Before the clones were collected, shoot demography and tussock architecture within these clones were recorded in the field for four seasons. Their clone identity was determined using DNA RAPD. Vegetatively propagated plants from these clones were cultivated in a common garden experiment to demonstrate variation in tussock growth and architecture. Their response to change in red/far red light ratio was determined in the growth chamber. Highly significant variation among clones was found in almost all parameters. In the common garden, the clones differed in tussock growth (mean tillering rate), architecture (mean shoot angle, mean tussock density) and proportion of flowering shoots. In the growth chamber, both the aboveground parameters and parameters of the rhizome system were strongly affected by red/far red ratio; among-clone variation was also almost always significant. The genotype × environment interaction was significant for tillering rate and rhizome architecture. The structure of the rhizome system (which is the major component of clonal spread in space) is a complex result of several components whose inter-clone variations differ: (i) genetically determined mean rhizome system size, (ii) overall plasticity in rhizome system size (with no significant genetic variation in plasticity), and (iii) genetically determined plasticity in rhizome architecture. Because of the variation in plasticity in rhizome architecture, some c lones seemed to possess the ability to exploit a favourable habitat patch by producing short branches when there; whereas the remaining clones appeared to possess only a simple escape mechanism from unfavourable patches. Environmental variation in the light levels in the studied grassland is fine grained; horizontal growth rates of F. rubra are sufficient to make genets experience different patches in their lifetime. The high variation in both genotype means and plasticities is likely to be due to selection early in genet life in an environment which is heterogeneous at a fine scale.     

Hormonal control of the outgrowth of axillary buds in <Emphasis Type="Italic">Alstroemeria</Emphasis> cultured <Emphasis Type="Italic">in vitro</Emphasis>

We study apical dominance in Alstroemeria, a plant with an architecture very different from the model species used in research on apical dominance. The standard explant was a rhizome with a tip and two vertically growing shoots from which the larger part had been excised leaving ca. 1 cm stem. The axillary buds that resumed growth were located at this 1-cm stem just above the rhizome. They were released by removal of the rhizome tip and the shoot tips. Replacement of excised tips by lanolin with indole-3-butyric acid (IBA) restored apical dominance. The auxin transport inhibitors 2,3,5-triiodobenzoic acid (TIBA) and N-1-napthylphthalamic acid (NPA) reduced apical dominance. 6-Benzylaminopurine (BAP) enhanced axillary bud outgrowth but the highest concentrations (> 9 μM) caused fasciation. Thidiazuron (TDZ) did not show improvement relative to BAP. Even though the architecture of Alstroemeria and the model species are very different, their hormonal mechanisms in apical dominance are for the greater part very similar.     

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.     

Auxin Depletion from the Leaf Axil Conditions Competence for Axillary Meristem Formation in Arabidopsis and Tomato

The enormous variation in architecture of flowering plants is based to a large extent on their ability to form new axes of growth throughout their life span. Secondary growth is initiated from groups of pluripotent cells, called meristems, which are established in the axils of leaves. Such meristems form lateral organs and develop into a side shoot or a flower, depending on the developmental status of the plant and environmental conditions. The phytohormone auxin is well known to play an important role in inhibiting the outgrowth of axillary buds, a phenomenon known as apical dominance. However, the role of auxin in the process of axillary meristem formation is largely unknown. In this study, we show in the model species Arabidopsis thaliana and tomato (Solanum lycopersicum) that auxin is depleted from leaf axils during vegetative development. Disruption of polar auxin transport compromises auxin depletion from the leaf axil and axillary meristem initiation. Ectopic auxin biosynthesis in leaf axils interferes with axillary meristem formation, whereas repression of auxin signaling in polar auxin transport mutants can largely rescue their branching defects. These results strongly suggest that depletion of auxin from leaf axils is a prerequisite for axillary meristem formation during vegetative development.     

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.     

Effects of endogenous hormones on variation of shoot branching in a variety of non-heading Chinese cabbage and related gene expression

Shoot branching (tillering) primarily determines plant shoot architecture and has been studied in many plants. Shoot branching is an important trait in non-heading Chinese cabbage (Brassica rapa ssp. chinensis Makino). The B. rapa ssp. chinensis var. multiceps exhibits unique and multiple shoot branching characteristics. Here, we analyzed the variation in shoot branching between ‘Maertou,’ with multiple shoot branching, and ‘Suzhouqing,’ a common variety. The levels of endogenous indole-3-acetic acid (IAA), zeatin riboside and active gibberellins in the shoot meristem tissues of the two cultivars were quantified by enzyme-linked immunosorbent assay during the vegetative growth stage. High levels of IAA maintained axillary bud dormancy and repressed axillary bud outgrowth allowing shoot branching to form in the vegetative stage in ‘Suzhouqing.’ In contrast, low levels of IAA did not inhibit axillary buds in ‘Maertou,’ while a high level of cytokinin promoted axillary bud growth and branch shoot development. Exogenous hormone (rac-GR24 and 6-benzylaminopurine) treatment showed that ‘Maertou’ was relatively sensitive to cytokinin, because the fold changes of cytokinin-responsive genes in ‘Maertou’ were significantly more frequent than those in ‘Suzhouqing’. Cytokinin was the direct regulator for axillary bud growth of ‘Maertou’. Compared with ‘Suzhouqing’, ‘Maertou’ was sensitive to cytokinin and this weakened the strigolactone–cytokinin branching pathway.     

Nonlinear processes in seagrass colonisation explained by simple clonal growth rules

The development of single clones of the seagrass Cymodocea nodosa was analysed using a growth model based on the formation of structures limited by diffusive aggregation. The model implemented the measured clonal growth rules (i.e. rhizome elongation and branching rates, branching angle, and spacer length between consecutive shoots) and shoot mortality rate for C. nodosa at Alfacs Bay (Spain). The simulated patches increased their size nonlinearly with time displaying two different domains of growth. Young patches showed a rapid increase with time of the length of rhizome network and the number of living shoots, which depended on rhizome branching rate, and increased the radial patch size (R_g) algebraically with the number of living shoots as R_g ∝ N_s^1/Df, being D_f the fractal dimension of the patch structure. Patches older than 4 years increased the production of rhizome network and the number of living shoots much more slowly, while their radial patch size behaved as R_g ∝ N_s^0.5 resulting from an internal patch compactation. Moreover, the linear growth rate of the simulated patches changed up to 30 fold during patch development, increasing with increasing patch size until patches reached an intermediate size. The modelled patch development was found to closely reproduce the observed patch structure for the species at the Alfacs Bay (Spain). Hence, the growth of C. nodosa patches initially proceeds with a growth mode controlled by the branching pattern (branching frequency and angle) of the species, producing sparse and elongated patches. Once patches exceed 4–5 years of age and contained >500 shoots, becoming dense and circular, they shifts to a growth model typical of compact structures. These results explain previously unaccounted evidence of the emergence of nonlinear patch growth from simple clonal growth rules, and highlight the importance of branching frequency and angles as critical determinants of the space occupation rate of seagrasses and probably other clonal plants.     

Spatial genotypical diversity of <Emphasis Type="Italic">Sesleria albicans</Emphasis> (Poaceae) in a dry grassland community

Spatial genotypical diversity of Sesleria albicans Kit. ex Schultes was studied in a dry grassland community by isozyme analysis. The aim was to identify the genetical individuals within the studied population and to assess the species’ clonal growth parameters. Vegetative mobility and branching intensity were measured in field for the sake of the correct interpretation of the results. Five isozyme systems were analyzed and interpreted (MDH, MNR, 6-PGDH, SkDH, LAP). Altogether, 98 distinct isozyme profiles were identified within an area of 2 m². Average genotype identity rapidly decreased with distance. Several very remote ramets (more than 1 m) belonging to the same genet were identified. The longest distance between ramets of the same genotype was 153 cm. With average annual rhizome increment of 13.59 mm assessed for the studied population the age of genet with the most remote ramets exceeded 56 years by a bi-directional growth and 112 years by a growth in single direction. Number of daughter tillers produced by a tiller per year was 0.67 (branching intensity, median) and the median of tiller life span was 2.5 years. The high genotypical diversity of S. albicans in the studied population could be a result of both, regular and continuous seed production with subsequent seedling recruitment and long life span of genets.     

The molecular and genetic regulation of shoot branching

The architecture of flowering plants exhibits both phenotypic diversity and plasticity, determined, in part, by the number and activity of axillary meristems and, in part, by the growth characteristics of the branches that develop from the axillary buds. The plasticity of shoot branching results from a combination of various intrinsic and genetic elements, such as number and position of nodes and type of growth phase, as well as environmental signals such as nutrient availability, light characteristics, and temperature (Napoli et al., 1998; Bennett and Leyser, 2006; Janssen et al., 2014; Teichmann and Muhr, 2015; Ueda and Yanagisawa, 2019). Axillary meristem initiation and axillary bud outgrowth are controlled by a complex and interconnected regulatory network. Although many of the genes and hormones that modulate branching patterns have been discovered and characterized through genetic and biochemical studies, there are still many gaps in our understanding of the control mechanisms at play. In this review, we will summarize our current knowledge of the control of axillary meristem initiation and outgrowth into a branch.

The key regulatory genes and the role of multiple plant hormones coordinate the process of axillary meristem initiation and subsequent growth into a branch.     

Sind Farne Kormophyten? Eine Alternative zur Telomtheorie

Modern ideas concerning cormophyte phylogeny are strongly influenced by the telome theory. Erect growth, radial symmetry, dichotomy, protostely and eusporangial structure are considered to be primitive features. Yet, the cormus needs redefinition. TheHypolepidaceae (s. str.) are shown to have rather a two-dimensional thallus than a three-dimensional cormus. Their so-called rhizome develops a continuous areophore margin which is connected with the margin of the fronds. The longitudinally inserted fronds and branches are produced by the terminal rhizome meristem which itself grows in many respects like the apical frond meristem. The rhizomes ofDennstaedtiaceae and theDavallia-type of rhizomes are intermediate between thisHypolepis-type and the true cormus-type. TheHypolepis-type is compared with fern prothalli. Under perhumid tropical conditions higher land plants may have evolved from small, creeping, two-dimensional fern prothallium-like progenitors with isomorphous gameto- and sporophytes.

Vorgetragen auf der Morphologentagung in Zürich 1975.     

植物茎分枝的分子调控

植物茎分枝结构决定了不同植物的不同形态结构.本文从腋生分生组织的发生、腋芽的生长两个方面综述了近年来植物分枝发生发育相关的分子机理研究及其进展.发现在不同植物中腋分生组织形成的基本机制是相似的,LS（lateral suppressor）及其同源基因在不同植物中都参与腋生分生组织的形成,而BL(blind)及其同源基因也参与调控腋生分生组织的形成.腋生分生组织的形成可能也是受激素调控的.目前,对腋芽生长的分子调控机制的认识主要集中于生长素通过二级信使的作用调控腋芽的生长.而生长素调控腋芽生长的机制已经较为清楚的有两条途径：一是生长素通过抑制细胞分裂素合成来调控腋芽的生长;另一途径是一种类胡萝卜素衍生的信号物质参与生长素的运输调控(MAX途径)来调控腋芽的生长.最新研究表明,TB1的拟南芥同源基因在MAX途径的下游负调控腋芽的生长.此外,增强表达OsNAC2也促进腋芽的生长.     

Clonal Growth of White Clover: Factors Influencing the Viability of Axillary Buds and the Outgrowth of a Viable Bud to Form a Branch

The development of an axillary bud of white clover to form abranch depends on (1) the bud being viable, vegetative and non-dormant,and (2) suitable conditions for outgrowth of the bud. Foragingtheory emphasises the second of these requirements. Glasshousestudies with white clover rarely result in a loss of bud viability.In contrast, in field populations over 50% of the buds reachingthe stage of maturity when branching can occur are not in aviable, vegetative, non-dormant condition. We examined whethernon-viability could be induced in a glasshouse experiment byapplying treatments in a factorial design. The factors were:defoliation, phosphorus supply, soil moisture status, simulatedtreading and grass competition. In addition, we measured theeffects of the treatments on the outgrowth of viable buds inorder to assess whether the same factors were determining viabilityand outgrowth. Defoliation significantly reduced bud viability(by 44%) but no other factors, either singly or in combination,had a significant effect. A greater variety of factors and combinationsof factors influenced bud outgrowth; these were defoliation,phosphorus status and interactions involving phosphorus andgrass; defoliation, phosphorus and soil moisture; and soil moisture,grass and treading. For white clover it is relevant to includethe state of the axillary meristem in any model of foraging. Trifolium repens ; white clover; axillary bud; viability; clonal growth; foraging; defoliation     

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

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

Histochemical Study of Xylem Cells in In Vitro Culture of Iris sibirica L.

In this study, lignin content data are presented for annual regenerant Iris sibirica plants, comparable to those in six-year-old intact plants. The structure of the shoots of Iris sibirica grown on artificial nutrient media was studied by the histochemical method. Features of the formation of the xylem in Iris sibirica on artificial nutrient media were revealed. Regenerants very quickly developed a complex system consisting of vascular bundles containing sieve tubes, vessels and tracheids, and hydrocyte systems. Hydrocytes of Iris sibirica were tracheids with lignified thickening, but, in contrast to tracheids and vessels of xylem (they are formed based on procambium or cambium—special lateral primary or secondary meristem), hydrocytes differentiated from the cells of permanent tissues (like phellogen), which probably possessed meristematic activity at the time of differentiation. In Iris sibirica hydrocytes covered the vascular bundle by the thick layer and strung along it up to a certain height. High lignin content in young regenerant Iris sibirica plants was due to the formation of the dense tissue from lignified tracheal elements. The study of the differentiation of xylem elements under controlled conditions can serve as a model for our understanding of wood formation processes.