Gas Exchange between Equisetum limosum and its Environment

Equisetum limosum grows in standing water. Its perennating rhizomesramify in the bottom mud, which is a markedly oxygen-deficientmedium. During the summer months aerial shoots are present andextend above the surface of the water to a height of about 1m. The concentration of oxygen in the internal atmosphere ofplants in the field decreases progressively from the stem tothe perennating rhizomes. There is a reverse gradient of carbondioxide concentrations. The structure of the nodal diaphragms of the aerial stem isdescribed and the progressive occlusion of the pores of thediaphragms between May and November is demonstrated. This occlusionof the diaphragms has been correlated with a reduction in thecase of gaseous diffusion along lengths of aerial stems. Oxygen is present in the rhizomes throughout the winter, despitethe fact that the dead aerial stems have broken off and thestumps have been submerged by the rise in water level, thusisolating the rhizome from atmospheric oxygen. The results are discussed in relation to the ability of theplant to grow in an oxygen-deficient environment.     

Rhizome development ofPhragmites australis in a reed community

Variations in the height, shoot density, biomasses of above- and below-ground parts and rhizome distributions ofPhragmites australis were investigated along a line-transect in a reed community at Yufutsu Mire, Hokkaido. Relationships of performance of the reed plants to soil conditions and species compositions were also examined. Three types of rhizome development were recognized in reed plants; (1) the central part of the reed community, characterized by well developed rhizomes and dense aerial shoots, (2) the intermediate part, characterized by development of rhizomes along both the peat and surface layers and very dense aerial shoots, and (3) the marginal part, characterized by development of rhizomes only along the peat layer and sparse aerial shoots. Observation showed that rhizomes in the surface layer actively produced aerial shoots, whereas rhizomes in the peat layer contributed to the spreading of their distribution range. With the growth of rhizomes, organic debris originating from dead rhizomes and roots accumulated in the mineral soil to promote organic soil formation. In dense parts of the reed stand, species composition was poor because of the shading and litter accumulation by reed plants. The effects of microtopography and water level on the establishment of reed seedlings were also considered.     

A thermal time model for predicting time to aerial shoot elongation in Variegated Solomon's Seal

Aerial shoot development in Variegated Solomon's Seal (Polygonatum odoratum‘Variegatum’) was studied under warm (mean 18°C) conditions after dormant rhizomes had been stored at a range of temperatures. After chilling at 0.8–5.5°C for 21–77 days, all rhizomes produced elongated aerial shoots, with mean lengths from 33 cm to 44 cm. Exposure of rhizomes to 15°C or 20°C for 21–77 days resulted in 17% to 50% of buds emerging as shoots, but these either aborted or failed to extend beyond a rosette. The earliest aerial shoot elongation was observed after 7–13 days at 18°C in rhizomes that had been chilled at 0.8–2°C treatments for 59–72 days. The base, optimum and maximum temperatures during pre‐planting storage for subsequent aerial shoot elongation were derived respectively as ?1.5°C, 1.9°C and 15.8°C. A thermal time of storage was calculated from these cardinal temperatures and the rate of progress to normal aerial shoot clongation was shown to increase linearly with increasing thermal time to c. 150°Cd. The thermal time procedure for predicting time to aerial shoot elongation constructed from growth room/chamber data was validated using rhizomes that had been exposed to varying temperatures in shadehouse conditions during the winter in Taiwan.     

Nitrogen translocation via rhizome systems in monoclonal stands ofReynoutria japonica in an oligotrophic desert on Mt Fuji: Field experiments

Mechanisms which enableReynoutria japonica, a dominant pioneer herb, to be successful in maintaining large stands in an oligotrophic volcanic desert on Mt Fuji were investigated with special reference to its nitrogen acquisition.Reynoutria japonica forms circular stands, each of which comprises only one genet. As a stand develops outwards, the number of aerial shoots per unit area decreases in the center. Shoots grow vigorously in the peripheral area where the available nitrogen from soil and precipitation (about 2.4 g m⁻² year⁻²) was much less than total nitrogen in the shoots (6.1–9.1 g m⁻²). Leaf nitrogen content per unit mass was also greater in the leaves of the peripheral shoots. When rhizomes extending radially from the center to the periphery were severed, the dry mass of shoots in the periphery diminished by 75% on a ground area basis. In the periphery, leaf nitrogen content also reduced significantly and no flowers were produced. When fertilizer was applied to the peripheral shoots with severed rhizomes, neither growth, survival nor flower production of the shoots was significantly smaller than the control levels. In these shoots, it is also found that the nitrogen content in the youngest leaves decreased for about 1 month and then increased to above that in the control leaves. These results suggest that (i) nitrogen accumulated in the central part is translocated to peripheral shoots via rhizomes, and that long-distance translocation enables the stands to develop outwards, and (ii) aerial shoots in the periphery utilize the nitrogen translocated by rhizomes in the beginning of the growth season, whereas once the shoots have established, they begin to take up nitrogen with their own roots. Since the peripheral shoots are in sunnier environments than the shoots inside the stand, the acropetal nitrogen translocation via rhizomes will raise the production efficiency of a whole stand.     

Translocation of Photoassimilates Between Sister Ramets in Two Rhizomatous Forest Herbs

Autoradiography and liquid scintillation techniques were usedto trace the pattern of photoassimilate translocation in twoperennial forest herbs, Aster acuminatus Michx. and Clintoniaborealis (Ait.) Raf. Vegetative shoots of C. borealis emergeearly each spring from the growing apices of a long-lived rhizomesystem. Vegetative shoots of A. acuminatus emerge in late springfrom rhizomes that decay within 2 years. In both species rametssurvive for only one growing season. Any connected plants aresisters. Mature leaves of these two species were exposed for1 h periods to ¹⁴CO₂ during spring and summer. Radioactivitycould subsequently be detected in exposed leaves (16–84per cent, depending on the season), adjacent above ground plantparts (0.7–23 per cent), roots and rhizomes (4.9–84per cent) and, when present, flowers (4–8.5 per cent).Old rhizomes of C. borealis are prominent storage sites forphotosynthate. In A. acuminatus, no significant translocationbetween sister ramets (i.e. above ground shoots connected bya common rhizome) was observed. In C. borealis, there was small,but consistent translocation between sister ramets (0.2–4per cent). Disturbance of unexposed sister ramets by defoliation,shading or herbivory increased the flow of photoassimilatesto disturbed parts in C. borealis, but not in A. acuminatus.Based on the absence of translocation flow, ramets of A. acuminatusmay be regarded as physiologically independent. Connected rametsof C. borealis show physiological integration. These resultsare correlated with ecological differences between the two species. Aster acuminatus, Clintonia borealis, translocation, ramet, vegetative reproduction, forest herb     

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.     

OCCURRENCE OF INTERCALARY AND UNINTERRUPTED MERISTEMS IN EQUISETUM

Quantitative data on cell length and mitotic index in elongating internodes of seven species of Equisetum are presented as evidence for the occurrence of two patterns of internode development in rhizomes of different species. In rhizomes of three species of subgenus Equisetum (E. arvense, E. diffusum, E. telmateia) uninterrupted meristems are present, characterized by acropetal internode maturation. In rhizomes of four species of subgenus Hippochaetae (E. hyemale, E. variegatum, E. scripoides, and E. laevigatum) intercalary meristems are present. The number and locations of intercalary meristems are described for an aerial shoot of E. diffusum. The absence or diminution of intercalary meristems from rhizomes of a variety of vascular plants with intercalary meristems in aerial shoots is discussed from the standpoint of adaptive significance.     

Underground Metameric Complex in the Source-Sink System of Rhizome-Forming Perennial Cereals <Emphasis Type="Italic">Bromopsis inermis</Emphasis> and <Emphasis Type="Italic">Phalaroides arundinacea</Emphasis>

Morphophysiological characteristics of rhizomes and growth relationships between underground shoots and aboveground orthotropic shoots were studied in two species of perennial monocotyledonous plants—Hungarian brome (Bromopsis inermis (Leyss.) Holub.) and reed canary-grass (Phalaroides arundinacea (L.) Rauschert.). The underground metameric complex was shown to be comparable with the aerial shoots in terms of the number, biomass, and metabolic activity of the shoots. The role of the underground metameric complex in the source-sink system of perennial rhizome-forming cereals is determined by a significant proportion of rhizomes in plant biomass (30–50%), formation of a large amount of meristems in the underground stock of vegetative reproduction (more than 1000 per plant), a comparatively high respiration rate (1.5 mg CO₂/(g dry wt h)), and a high nitrogen content (3.5%). No pronounced growth response was found in the rhizome upon plant treatment with growth regulators (GA and chlorocholine chloride) and upon decapitation of plant shoots. It is concluded that the underground metameric complex of the perennial monocotyledonous herbaceous plants is relatively autonomous from the orthotropic shoots.     

Oxygen demand and supply in Zizania latifolia and Phragmites australis

The oxygen demand of underground organs in tall emergent plants was assumed to be supplied with the oxygen flux through aerial shoots. Measurements were made of the respiratory activity of underground organs and oxygen flux through aerial shoots in Zizania latifolia (Griseb.) Stapf and Phragmites australis (Cav.) Trin. ex Steud.Oxygen uptake rates of Z. latifolia roots and rhizomes decreased as the supply of oxygen decreased. Phragmites australis, however, did not respond to decreases in the supply of oxygen until the oxygen supply was relatively low. The values for old rhizomes of P. australis were very low. Most of the oxygen flux for P. australis was derived from standing dead shoots. The important role of dead shoots of P. australis in plant aeration is suggested. The degree of sufficiency of oxygen supply for Z. latifolia was 179% in May and 92% in August. The values for P. australis were 45% both in May and August. Furthermore, estimation of mean oxygen concentration of inner underground organs for P. australis showed very low values, 1.6% both in May and August, even in shallow water.     

Effects of post-chilling temperature on growth and variegation in Solomon's Seal

Effects of temperature were studied on the current and following season's growth of shoots from chilled rhizomes of Variegated Solomon's Seal. The rate of progress to completed elongation of the aerial shoot in chilled plants increased linearly with increasing temperature up to 28°C (24 h mean). A post‐chilling thermal time of 658 ± 47°Cd (> ‐1.3°C) was required for aerial shoots to become fully extended. Temperatures of 28°C and 33°C accelerated aerial shoot senescence and decreased rhizome and root dry weights, as compared with 18°C and 23°C treatments. Leaf number and variegation were not affected by temperature treatments during current growth season and all plants produced 12–13 leaves with between 7% and 9% leaf area variegated. Leaf variegation, however, was significantly increased in plants that had been grown after chilling at 28°C during the preceding growing season. Proteins of approximately 26, 32 and 62 kDa were present in the green parts of leaves but not in the white parts.     

Effects of self-thinning of shoots on the nutrient budgets of Zizania latifolia

This experimental study focused on the seasonal changes and mobilization of nutrients between plant parts to understand the implications of self-thinning in the ecology of Zizania latifolia (Griseb.) Turcz. ex Stapf. The observations of shoot density, above- and belowground biomass, and total nitrogen and total phosphorus concentrations were conducted from February 2002 to August 2003. The biomass of shoots that died during the period and daily uptake of nutrients were determined. The shoot density sharply increased until mid-April and thereafter decreased significantly due to self-thinning of shoots. Total nitrogen and phosphorus concentrations of rhizomes decreased initially due to translocation to new shoots; however, the nutrients of rhizomes were slightly replenished from dead shoots during self-thinning. In contrast to other species, self-thinning of Z. latifolia shoots reallocates some of the minerals contained in the dead shoots back to the rhizomes, which can be regarded as a strategy to replenish the reduced resources of the rhizomes. The initial intensive growth of shoots can be regarded as a strategy to maintain resource competition.     

Propagation mechanisms in Agave macroacantha (Agavaceae), a tropical arid-land succulent rosette

Agave macroacantha can sexually reproduce by seeds and propagate vegetatively by aerial bulbils and ground-level basal shoots and rhizomes. It forms compact patches apparently generated by the multiplication of ground-level offshoots. We experimentally evaluated the establishment and survival of bulbils and seedlings of A. macroacantha in the Tehuacán Valley, Mexico, between 1991 and 1994 and studied comparatively the effectiveness of sexual reproduction against vegetative propagation.Seedlings showed low survival rates. Cohorts placed outside nurse plants died in less than 1 yr, while 1-10% of cohorts under nurse plants survived for more than 2 yr. Herbivores negatively affected seedling survival in non-nursed plots. In rainy years, survival rates increased. Bulbils showed higher survival rates than seedlings.The excavation of rosettes showed that most are derived from vegetative shoots, as indicated by remains of rhizomes in their base. Most rosettes had ground-level vegetative offspring totaling almost three shoots per rosette.In A. macroacantha, the establishment of seedlings and bulbils is a rare event that possibly only occurs under nurse plants in rainy years, while ground-level cloning is highly effective as a propagation mechanism. These results are consistent with the aggregated spatial pattern of the species.     

Effects of Photoperiod and Temperature on Shoot, Root and Rhizome Growth in ThreeLotus uliginosusSchkuhr Populations

The growth of three populations of greater lotus (Lotus uliginosusSchkuhrsyn.L. pedunculatusCav.) was compared at photoperiods of 10,12 and 14 h at a maximum day/minimum night temperature of 21/16°C and at maximum day/minimum night temperatures of 27/22,21/16, 18/13 and 15/10 °C at a photoperiod of 12 h. Shortdays (10 h) favoured root and rhizome development compared tolong days (14 h). A temperature regime of 15/10 °C restrictedrhizome development compared to the 18/13 and 21/16 °C regimes.Shoot growth was restricted at the highest temperature regime(27/22 °C). The cultivar Sharnae had fewer, but heavier,rhizomes than Grasslands Maku; this may indicate adaptationto the dry summers at its site of origin (Algarve, Portugal).The response of rhizome growth to temperature and photoperiodexplains part of the performance of greater lotus in the fieldat a wide range of latitudes. Grazing management to encouragethe persistence ofL. uliginosusin pasture in temperate environmentsmay include the exclusion of grazing livestock in autumn. Inthe sub-tropics, monitoring of rhizome production in the fieldwould be required before deciding the appropriate time intervalbetween grazing.Copyright 1998 Annals of Botany Company Lotus uliginosus(Schkuhr); greater lotus; temperature; daylength; shoots; roots; rhizomes.     

Stand Density Effects on Branching in an Annual Legume (Senna obtusifolia)

Effects of proximity to nearest neighbours on shoot morphologyand branch growth were examined for Senna obtusifolia (L.) I.& B., an annual legume species that displays wide variationin branching. Periodic surveys described location and type ofgrowth at all nodes on plants in regularly spaced monocultures.Stands with interplant distances of 15-50 cm (51-5 plants m^-2)formed closed canopies with similar amounts of leaf area andbiomass. Number of lower primary branches, their degree of curvature,and location of branch apices relative to the main stem respondedto interplant distance relatively early in shoot growth, beforecanopy closure. The final effects of ten-fold differences inplanting density on the formation of these ascending lower brancheswere only two-fold differences in number of vertical shoot axeswithin the stand. An additional study examined response of lowerbranches to an adjacent gap in the stand. Initial number anddirection of branch growth were not affected by location ofthe gap. The first effect of crowding by neighbours was on number ofbranches initiated by developing shoots. Differences in branchingsubsequently increased through differentiaI survival and directionof branch growth. Such alterations principally affected lateralexpansion of individual shoots, with little effect on radialsymmetry or vertical distribution of leaves.Copyright 1994,1999 Academic Press Branch morphology, canopy, competition, light quality, Senna obtusifolia, shoot growth, sicklepod     

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.     

Tuber Formation in Excised Stem Tips of Stachys sieboldii

The excised stem-tips of the germinating tubers of Stachys sieboldii Miq. were cultivated under the laboratory condition. As the room temperature decreased, the formation of tubercles on the tips of the explanted stems was observed. The excised tips of the rhizome at its different stages of development, when cultured in vitro, demonstrated variations in tuber formation. When the excised tips were taken from the rhizomes which had just emerged from the base of the aerial shoots, they usually failed to grow. However, existence of normal growth was observed in the cultured excised tips of the rhizomes that had become elongated at which time flowering occurred on the aerial shoot. The tips became swollen as the environmental temperature decreased. In early September, as the process of underground tuberization initiated, cultures taken from the nontuberized tips again grew poorly and usually failed to form tubercles.     

Transformation of the Root System by Agrobacterium rhizogenes Changes Rhythmic Growth of the Shoot of Ericaxdarleyensis In Vitro

Erica x darleyensis grows rhythmically when plants are propagatedin vitro on a medium with nitrate nitrogen. The aim of thisstudy was to determine if transforming the root system withAgrobacterium rhizogenes (strain A4) alters the rhythmic developmentof the aerial shoots of Erica x darleyensis. Morphological analysisof the chimeric plants compared to the control show that thetransformed genetic information not only modifies the root systembut also dampens the morphogenetic rhythm of the apical bud.Copyright1994, 1999 Academic Press Erica x darleyensis, Agrobacterium rhizogenes, root, rhythmic growth, tissue culture     

Seasonality of Rhizome and Shoot Production and Nitrogen Fixation in Lotus uliginosus under Upland Conditions in South-west Scotland

Development, growth and nitrogen fixation were monitored over3 years in marsh birdsfoot trefoil (Lotus uliginosus Schkuhr),at an upland site in south-west Scotland Plants exhibited markedseasonality of growth, with production of both aerial shootsand rhizome reaching a peak in September and October Renewedaerial shoot growth in spring was initiated mainly from nodeson rhizomes formed in the previous autumn Nitrogen fixationwas measured in soil cores using the acetylene reduction techniqueFixation commenced in early June, coinciding with increasingsoil temperature, and thereafter rose to a peak in mid-summerThe subsequent decline in fixation was associated with increasingrainfall, decreasing soil temperature, flowering and the onsetof rhizome production Lotus uliginosus Schkuhr, marsh birdsfoot trefoil, seasonality, rhizome production, nitrogen fixation, acetylene reduction     

川西北高寒沙区切断根茎对赖草和沙生苔草克隆生长的影响

为了研究高原亚寒带沙化生境中切断根茎对克隆植物基株扩展能力和分株定居能力的影响,在川西北若尔盖高原沙化区内,对根茎禾草赖草和沙生苔草进行了以切断根茎为处理的野外实验。结果表明,赖草和沙生苔草基株的幼小部分(观测单元)地上枝总长度增量、主根茎长度增量和根茎总长度增量显著减少,而对根茎数增量、主根茎节增量和根茎节总数增量影响不显著;赖草观测单元地上枝数增量显著减少,而对沙生苔草地上枝数增量无影响;赖草地上枝与根茎的相关性质发生逆转。这表明．在高原亚寒带半湿润沙化生境中．克隆整合效应显著促进基株幼小部分地上枝和根茎的伸长生长,但对新生根茎的产生和根茎节分化没有影响;切断根茎处理导致赖草、沙生苔草生殖生长与营养生长间竞争加剧,同时使赖草地上部分与地下部分间竞争加剧;观测单元在缺少与基株(或上级株系)的克隆整合作用时,赖草受到的影响大于沙生苔草。     

The development of shoots in Phragmites communis Trin.

In aerial shoots of Phragmites communis Trin., final height,and often growth-rate, depend on the width of the emergent budwhich determines the basal diameter of the mature stem. In theearly stages there is internal competition for food reservesand smaller shoots may be harmed. Growth is faster during the early phase of growth, and it isincreased in wider buds, warmer weather, and more humid habitats.Any of these factors may be of over-riding importance, e.g.a summer-emerged shoot grows faster than a spring one of equalwidth. Minor fluctuations in environment have more effect earlythan later. New nodes develop more rapidly late in the yearthan early.