Overwintering leaves of a forest-floor fern, Dryopteris crassirhizoma (Dryopteridaceae): a small contribution to the resource storage and photosynthetic carbon gain

BACKGROUND AND AIMS: Dryopteris crassirhizoma is a semi-evergreen fern growing on the floor of deciduous forests. The present study aimed to clarify the photosynthetic and storage functions of overwintering leaves in this species. METHODS: A 2-year experiment with defoliation and shading of overwintering leaves was conducted. Photosynthetic light response was measured in early spring (for overwintering leaves) and summer (for current-year leaves). KEY RESULTS: No nitrogen limitation of growth was detected in plants subjected to defoliation. The number of leaves, their size, reproductive activity (production of sori) and total leaf mass were not affected by the treatment. The defoliation of overwintering leaves significantly reduced the bulk density of rhizomes and the root weight. The carbohydrates consumed by the rhizomes were assumed to be translocated for leaf production. Photosynthetic products of overwintering leaves were estimated to be small. CONCLUSION: Overwintering leaves served very little as nutrient-storage and photosynthetic organs. They partly functioned as a carbon-storage organ but by contrast to previous studies, their physiological contribution to growth was found to be modest, probably because this species has a large rhizome system. The small contribution of overwintering leaves during the short-term period of this study may be explained by the significant storage ability of rhizomes in this long-living species. Other ecological functions of overwintering leaves, such as suppression of neighbouring plants in spring, are suggested.     

Nitrogen and carbohydrate pools of two rhizome types of Phragmites australis (Cav.) Trin. ex Steudel

Carbohydrate and total nitrogen contents as well as free amino acid (FAA) concentrations were determined in distinct types of rhizomes of two genetically homogeneous stands of reed (Phragmites australis (Cav.) Trin. ex Steudel) differing in morphology, productivity and nutrient supply in order to evaluate the storage capacity of vertical rhizomes and expansion rhizomes. The expansion rhizomes possess significantly higher amounts of FAA and of total nitrogen but similar carbohydrate concentrations in comparison to the vertical rhizomes. However, no significant differences were found for total nitrogen, FAA and total carbohydrates between both investigated stands indicating a comparable storage capacity of rhizomes independent of nutrient availability. Only the composition of the FAA pool varied in the alanine/asparagine ratio probably influenced by the oxygen supply of the rhizome/root system.     

CHANGES IN STARCH DISTRIBUTION IN THE OVERWINTERING ORGANS OF TYPHA LATIFOLIA (TYPHACEAE)

Histochemical determinations for storage of carbohydrates in rhizomes, roots, and young shoots of Typha latifolia L. (Typhaceae) were conducted during the overwintering period from November to April. Early winter analysis showed that rhizomes and roots contained large amounts of starch (45.03% and 22.80% dry weight, respectively). The major storage tissue was parenchyma of the rhizome central core. From winter into spring a gradual decrease in storage starch in the rhizome and root occurred concurrently with starch accumulation near zones of rapid development in young shoots (buds), but the rhizome retained much starch (27.40% dry weight) into the start of its 2nd yr.     

Rhizome age structure of three populations ofPhragmites australis (Cav.) Trin. ex Steud.: Biomass and mineral nutrient concentrations

The rhizome age structure for populations ofPhragmites australis (Cav.) Trin. exSteud. was determined at three sites of the T?eboň Biosphere Reserve (Czech Republic). These sites were classified according to plant species composition and phosphorus availability as oligotrophic (Branná), mesotrophic (Ro?mberk East) and hypertrophic (Ro?mberk West).P.australis stands at these sites were expanding, stabilized and retreating, respectively. Rhizomes were sampled within the terrestrial parts of the reed stands (at a water depth of about 10 cm). The rhizomes were dated according to the position in the branching hierarchy and to supplementary morphological criteria. The analysis of the branching pattern revealed that at the sites studied, the rhizomes typically branch only once a year. The longest life span of rhizomes (six years) was found for the stable reed population at Ro?mberk East. The reduced rhizome life span at Ro?mberk West (four to five years), accompanied by a reduced branching frequency of horizontal rhizomes, was ascribed to a greater incidence of anaerobic conditions associated with the permanent flooding of the hypertrophic organic soil. The maximum life span of the Branná population (four years) probably corresponds to the total age of the reed population. The concentrations of nitrogen, phosphorus, and potassium in the tissue decreased with rhizome age while those of calcium and magnesium increased. The effect of rhizome age on mineral-nutrient concentrations was more pronounced than the effect of site on these concentrations. The mineral nutrient concentrations were not closely associated with reed performance.     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Latent Pathogenic Fungi in the Medicinal Plant Houttuynia cordata Thunb. Are Modulated by Secondary Metabolites and Colonizing Microbiota Originating from Soil

Latent pathogenic fungi (LPFs) affect plant growth, but some of them may stably colonize plants. LPFs were isolated from healthy Houttuynia cordata rhizomes to reveal this mechanism and identified as Ilyonectria liriodendri, an unidentified fungal sp., and Penicillium citrinum. Sterile H. cordata seedlings were cultivated in sterile or non-sterile soils and inoculated with the LPFs, followed by the plants’ analysis. The in vitro antifungal activity of H. cordata rhizome crude extracts on LPF were determined. The effect of inoculation of sterile seedlings by LPFs on the concentrations of rhizome phenolics was evaluated. The rates of in vitro growth inhibition amongst LPFs were determined. The LPFs had a strong negative effect on H. cordata in sterile soil; microbiota in non-sterile soil eliminated such influence. There was an interactive inhibition among LPFs; the secondary metabolites also regulated their colonization in H. cordata rhizomes. LPFs changed the accumulation of phenolics in H. cordata. The results provide that colonization of LPFs in rhizomes was regulated by the colonizing microbiota of H. cordata, the secondary metabolites in the H. cordata rhizomes, and the mutual inhibition and competition between the different latent pathogens.     

Targeted mapping of quantitative trait locus regions for rhizomatousness in chromosome SBI-01 and analysis of overwintering in a Sorghum bicolor × S. propinquum population

While rhizome formation is intimately associated with perennialism and the derived benefit of sustainability, the introduction of this trait into temperate-zone adapted Sorghum cultivars requires precise knowledge of the genetics conditioning this trait in order to minimize the risk of weediness (e.g., Johnsongrass, S. halepense) while maximizing the productivity of perennial sorghum. As an incremental step towards dissecting the genetics of perennialism, a segregating F4 heterogeneous inbred family derived from a cross between S. bicolor and S. propinquum was phenotyped in both field and greenhouse environments for traits related to over-wintering and rhizome formation. An unseasonably cold winter in 2011 provided high selection pressure, and hence 74.8 % of the population did not survive. This severe selection pressure for cold tolerance allowed the resolution of two previously unidentified over-wintering quantitative trait locus (QTL) and more powerful correlation models than previously reported. Conflicting with previous reports, a maximum of 33 % of over-wintering variation could be explained by above-ground shoot formation from rhizomes; however, every over-wintering plant exhibited rhizome growth. Thus, while rhizome formation is required for over-wintering, other factors also determine survival in this interspecific population. The fine mapping of a previously reported rhizome QTL on sorghum chromosome SBI-01 was conducted by targeting this genomic region with additional simple sequence repeat markers. Fine mapping reduced the 2-LOD rhizome QTL interval from ~59 to ~14.5 Mb, which represents a 75 % reduction in physical distance and a 53 % reduction in the number of putative genes in the locus.     

Metabolism of Monoterpenes : Evidence for the Function of Monoterpene Catabolism in Peppermint (Mentha piperita) Rhizomes

l-Menthone of peppermint leaves is reduced to d-neomenthol which is glucosylated and transported to the rhizome, whereupon the β-d-glucoside is hydrolyzed, the aglycone oxidized back to l-menthone, and this ketone converted to l-3,4-menthone lactone. l-[G-³H]-3,4-Menthone lactone and its labeled progenitors, when incubated with excised mint rhizomes, gave rise to nonvolatile lipids as well as polar metabolites. The lipids thus generated consisted of labeled squalene and phytosterols in the nonsaponifiable fraction and C₁₄-C₂₆ fatty acids in the saponifiable fraction. These results imply degradation of the terpenoid to acetylcoenzyme A and reduced pyridine nucleotide, and reincorporation of label via these products. Starch and soluble carbohydrates were also found to be labeled; however, chemical degradation of the [³H]glucose obtained on hydrolysis of starch indicated the presence of tritium only on interior carbons, suggesting that labeling had occurred via reduced pyridine nucleotides. Analysis of the labeled organic acids revealed the presence of several hydroxy methylacyl intermediates suggesting the operation of a modified β-oxidation pathway in the degradation of the acyclic terpenoid skeleton. The results indicate that monoterpenes transported to the rhizome are oxidized to yield acetyl-coenzyme A and reduced pyridine nucleotides, and suggest that metabolic turnover of monoterpenes in mint represents a mechanism for recycling carbon and energy from foliar terpenes into other metabolites of the rhizome.     

Comparison of nitrogen uptake in the roots and rhizomes of <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Leymus chinensis (Trin.) Tzvel is a rhizomatous grass species in the Eastern Eurasian steppe zone that is often limited by low soil nitrogen availability. Although a previous study showed that the rhizomes of L. chinensis have the capacity to take up nitrogen, the importance of such uptake for nitrogen nutrition is unclear. Moreover, little is known regarding the inorganic nitrogen uptake kinetics of roots and rhizomes in response to nitrogen status. Here, we first found that ammonium is preferred over nitrate and glycine for L. chinensis growth. Using the ¹⁵N-labelling method, we found that the rate of ion influx into roots was approximately five-fold higher than into rhizomes under the same nitrogen content, and the ion influxes into roots and rhizomes under 0.05 mM N were greater than in the presence of 3 mM N, especially in the form of NH₄⁺. Using a non-invasive micro-test technique, we characterised the patterns of NH₄⁺ and NO₃^– fluxes in the root mature zone, root tip, rhizome mature zone, and rhizome tip following incubation in the solution with different N compounds and different N concentrations. These results suggest a dynamic balance between the uptake, utilisation, and excretion of nitrogen in L. chinensis.     

Effect of Temperature, Photoperiod and Several Growth Substances on the Cold Hardiness of Chrysanthemum morifolium Rhizomes

The effect of 14 combinations of photoperiod, soil and air temperature, and growth substance applications on the cold hardiness of Chrysanthemum morifolium‘Astrid’ rhizomes was evaluated. Both triphenyl tetrazolium chloride and regrowth tests were used to determine the viability of the cold-stressed rhizome tissues. The rhizomes exhibited different degrees of cold hardiness under these environmental conditions. A combination of short photoperiod and low air and soil temperatures induced maximum cold hardiness. Low soil temperature accompanied by long photoperiods and warm aerial temperatures did not induce rhizome hardening, while some hardening in cool soils was evident under either short photoperiods or low aerial temperatures. Warm soils reduced rhizome hardening under the normally inductive short photoperiod-cool aerial conditions. Since the induction of rhizome hardening was dependent on the induction of the aerial organs, the involvement of translocatable hardiness promoters is indicated. Foliar applications of low levels of gibberllic acid (GA₃) or abscisic acid only slightly influenced rhizome hardiness.     

扎龙湿地不同生境芦苇种群根茎数量特征及动态

采用单位土体取样,计测长度和生物量的调查与统计方法,对扎龙湿地保护区4个生境芦苇种群根茎数量特征进行比较分析。结果表明,芦苇5月10日左右返青后进入营养生长期,根茎长度6—8月份缓慢增加,8—10月份显著增加,后期是前期的3.5—10.3倍,生长季中后期是种群新根茎补充和生长的主要时期,不仅实现了种群空间扩展,并为营养繁殖储备更多繁殖芽;根茎生物量和干物质贮量6—8月份逐渐减少,8—10月份又逐渐增加,均以生长季末期的10月份最大,并均显著地(P0.05)高于其他月份,种群根茎养分的消耗主要供给根茎芽的萌发和幼株生长,根茎养分的储藏又为翌年种群的更新及扩展提供物质保障,种群对地下根茎存在明显的养分"超补偿性"贮藏现象。种群根茎长度和生物量均以湿生生境最大,依次为旱生生境、水生生境,盐碱生境最小,根茎干物质贮量以旱生生境最大,依次为湿生生境、水生生境,盐碱生境最小。种群根茎长度与返青后实际生长时间之间均较好地符合直线函数关系,种群根茎生物量和干物质贮量与生长时间之间较好地符合二次曲线函数关系,R2在0.804—0.997之间,拟合方程均达到了显著或极显著(P0.01)水平。4个生境芦苇种群在根茎长度、生物量、干物质贮量等数量特征均表现出由遗传因素控制的比较稳定的季节动态规律,在生境间的差异及其差异序位又均基本稳定,均表现出明显的土壤因子环境效应,其中土壤含水量、有机质、速效氮为正向驱动,p H、速效磷为负向驱动,土壤含水量、p H对根茎数量特征的驱动作用更突出。     

Moderate abiotic stresses increase rhizome growth and outgrowth of axillary buds in Alstroemeria cultured in vitro

Alstroemeria is multiplied in vitro by forced outgrowth of lateral rhizomes from rhizome explants. The multiplication rate is very low because of strong apical dominance and poor rhizome growth. We report here that moderate abiotic stresses stimulate both rhizome growth and outgrowth of lateral rhizomes, and accordingly increase multiplication. Rhizome explants were exposed to heat by a hot-water treatment (HWT) or by a hot-air treatment. Both increased rhizome growth when applied for 1 or 2?h in the range of 30?C40?°C. The maximal enhancement was 75?%. Other abiotic stresses were also examined. Cold (0?°C) and partial anaerobiosis increased rhizome growth significantly. The increases brought about by drought and salinity were not statistically significant. Because underground storage tissues like rhizomes are adaptations to survive climatic stresses, we presume that the increased sink-strength of rhizomes induced by moderate stress is related to stress adaptation. Moderate heat stress (38?°C HWT, 1?h) also resulted in protection of Alstroemeria plantlets from severe heat stress (45?°C HWT, 1?C2?h) a few hours after the moderate stress. All abiotic stresses also increased the outgrowth of lateral rhizomes.     

青藏高原掌叶大黄和丽江大黄及其土壤的主要元素含量

采集掌叶大黄（Ｒｈｅｕｍ ｐａｌｍａｔｕｍ）和丽江大黄（Ｒｈｅｕｍ ｌｉｋｉａｎｇｅｎｓｅ）根茎、叶及根部土壤,应用ＩＣＰ测定主要元素含量特征,研究了２种大黄及其土壤的元素含量特征。结果表明,２种大黄土壤Ｐ含量远低于大黄根茎和叶,Ｆｅ含量则为土壤〉根茎〉叶,其中根茎与叶相差不大,但它们与土壤相差５０～１１０倍;Ｎａ、Ｍｎ、Ｃｕ含量在２种大黄中都表现为土壤〉叶〉根茎,Ｃａ含量在掌叶大黄中为土壤〈叶〈     

Tissue-Specific Transcriptomic Profiling of Sorghum propinquum using a Rice Genome Array

Sorghum (Sorghum bicolor) is one of the world''s most important cereal crops. S. propinquum is a perennial wild relative of S. bicolor with well-developed rhizomes. Functional genomics analysis of S. propinquum, especially with respect to molecular mechanisms related to rhizome growth and development, can contribute to the development of more sustainable grain, forage, and bioenergy cropping systems. In this study, we used a whole rice genome oligonucleotide microarray to obtain tissue-specific gene expression profiles of S. propinquum with special emphasis on rhizome development. A total of 548 tissue-enriched genes were detected, including 31 and 114 unique genes that were expressed predominantly in the rhizome tips (RT) and internodes (RI), respectively. Further GO analysis indicated that the functions of these tissue-enriched genes corresponded to their characteristic biological processes. A few distinct cis-elements, including ABA-responsive RY repeat CATGCA, sugar-repressive TTATCC, and GA-responsive TAACAA, were found to be prevalent in RT-enriched genes, implying an important role in rhizome growth and development. Comprehensive comparative analysis of these rhizome-enriched genes and rhizome-specific genes previously identified in Oryza longistaminata and S. propinquum indicated that phytohormones, including ABA, GA, and SA, are key regulators of gene expression during rhizome development. Co-localization of rhizome-enriched genes with rhizome-related QTLs in rice and sorghum generated functional candidates for future cloning of genes associated with rhizome growth and development.     

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。     

Fungal endophytes of turmeric (<Emphasis Type="Italic">Curcuma longa</Emphasis> L.) and their biocontrol potential against pathogens <Emphasis Type="Italic">Pythium aphanidermatum</Emphasis> and <Emphasis Type="Italic">Rhizoctonia solani</Emphasis>

Endophytic fungi have been isolated from the healthy turmeric (Curcuma longa L.) rhizomes from South India. Thirty-one endophytes were identified based on morphological and ITS–rDNA sequence analysis. The isolated endophytes were screened for antagonistic activity against Pythium aphanidermatum (Edson) Fitzp., and Rhizoctonia solani Kuhn., causing rhizome rot and leaf blight diseases in turmeric respectively. Results revealed that only six endophytes showed >?70% suppression of test pathogens in antagonistic dual culture assays. The endophyte T. harzianum TharDOB-31 showed significant in vitro mycelial growth inhibition of P. aphanidermatum (76.0%) and R. solani (76.9%) when tested by dual culture method. The SEM studies of interaction zone showed morphological abnormalities like parasitism, shriveling, breakage and lysis of hyphae of the pathogens by endophyte TharDOB-31. Selected endophytic isolates recorded multiple plant growth promoting traits in in vitro studies. The rhizome bacterization followed by soil application of endophyte TharDOB-31 showed lowest Percent Disease Incidence of rhizome rot and leaf blight, 13.8 and 11.6% respectively. The treatment of TharDOB-31 exhibited significant increase in plant height (85 cm) and fresh rhizome yield/plant (425 g) in comparison with untreated control under greenhouse condition. The confocal microscopy validates the colonization of the TharDOB-31 in turmeric rhizomes. The secondary metabolites in ethyl acetate extract of TharDOB-31 were found to contain higher number of antifungal compounds by high resolution liquid chromatograph mass spectrometer analysis. Thereby, endophyte T. harzianum isolate can be exploited as a potential biocontrol agent for suppressing rhizome rot and leaf blight diseases in turmeric.     

Harvesting reed sweetgrass (Glyceria maxima,poaceae): Effects on growth and rhizome storage of carbohydrates

This study was aimed at elucidating the general effects of harvesting on the growth of reed sweetgrass, Glyceria maximaPoaceae, an emergent aquatic macrophyte potentially suitable for recovery of nutrients from wastewater. In a natural stand significantly lower amounts of young rhizomes and a lower concentration of nonstructural carbohydrates were observed after the first harvest. This indicated a reduced translocation of carbohydrates to the rhizomes and a depressed rhizome growth. The concentration of carbohydrates increased to the same level as in an unharvested plot within 7 wk. Rhizome biomass at the end of that growing season was 380 g/m² in the plot harvested two times and 725 g/m² in the unharvested plot. More N was removed with two harvests (12.1 g/m²) than with one (8.0 g/m²). In the following spring the shoots harvested two times had the lowest N content, which suggests that in harvested stands a high input of external nutrients may be needed to maintain optimal growth.     

Storage carbohydrate production and overwintering strategy in a winter-green tussock grass on South Georgia (Sub Antarctic)

Summary Water-soluble and ethanol-soluble carbohydrates were extracted from leaves, shoots, roots, rhizomes and flowers ofParodiochloa flabellata (Lam.) Rasp. (=Poa flabellata (Lam.) Hook f.) throughout a growing season. Very high levels of soluble carbohydrates were found in the shoot, comprising up to 71% of the shoot dry weight. There was no significant storage in the rhizome. It is suggested that the production of storage fructans with a degree of polymerization greater than 14, rather than oligosaccharides or starch, may be related to the cold environment. Relationships between the onset of senescence and levels of fructans suggest that recovery of sugars from dying tissue is high. The sugar reserves together with preformation of the flowers should ensure production of seed however poor the summer or long the winter might be. The high levels of fructans may partly inhibit growth but may also offer some degree of freezing resistance.     

Effects of simulated feeding by snow geese on Scirpus americanus rhizomes

Summary We simulated the feeding of Greater Snow Geese (Chen caerulescens atlantica) on the rhizomes of three-square bulrush (Scirpus americanus) in a tidal marsh along the St. Lawrence River estuary in Québec. During the spring staging period, aboveground biomass is unavailable and geese feed solely on rhizomes and overwintering buds. An experiment was designed to test the effect of three factors on subsequent growth of Scirpus: the intensity of removal (3 to 77% removal of belowground biomass), the number of bites (1, 2 or 3 sections removed) and the number of adventitious buds removed (1, 2 or 3). Rhizomes were dug out in May, treated and transplanted into 85-1 basins sunk in the marsh and filled with marsh soil freed of all plant material. Growth was observed weekly until the end of the growing season in August. Shoots and rhizomes were then collected, dried and weighed to obtain biomass estimates. The net above- and belowground production of Scirpus was inversely related to the initial rhizome biomass removed. At a high level of removal (>35%), the cumulative number of shoots was significantly reduced as early as two weeks after transplantation. The relative reduction in production of the treated rhizomes compared to the control plants was also related to the intensity of removal. An increased number of bites reduced production and the removal of an increased number of adventitious buds further amplified the effect of removal on rhizome production. These experimental results show that even low intensity of feeding by Snow Geese can reduce the production of Scirpus marshes.     

Seasonal changes in the amount of carbohydrate and crude protein in the rhizome ofMiscanthus sacchariflorus

In the herbaceous perennials, seasonal changes in the amount of reserve substances play an important role in the production processes of whole plants. Mutohet al. (1968) indirectly estimated the amount of reserve substances stored in the rhizome ofMiscanthus sacchariflorus by measuring the bulk density of the rhizome, and found aarked seasonal changes. In the present study seasonal changes in the carbohydrate and crude protein content of the rhizome were determined and the results obtained were compared with the changes in the total amount of reserve substances estimated by means of changes in the bulk density. A good parallel seasonal relationship was confirmed between the amount of consumable carbohydrate determined chemically and the quantity of reserve substances estimated from the changes in bulk density. The total yields of carbohydrate and crude protein were ca. 75% of the available reserve substances estimated from the changes of bulk density for the old rhizomes and ca. 65% for the new rhizomes in the middle of December.