中国三种水韭属植物的地理分布与生境特征

水韭属（Isoetes）是水韭科中的孑遗属,分类上属拟蕨类,对研究蕨类植物的系统演化具有重要价值。由于人类活动对其水生生境的破坏,水韭属植物在我国的分布范围及其种群数量日趋减少,濒临灭绝,其中,高寒水韭(I. hypsophila)、云贵水韭(I. yunguiensis)和中华水韭(I. sinensis)已被列为国家一级重点保护野生植物。对原产中国的3种水韭的地理分布与生境特征进行调查,将为合理保护该属植物提供相关基础数据和科学依据。本文野外实地考察了原产中国的3种水韭属植物的现存种群及历史记载曾经分布但现已绝迹的水韭属植物种群(绝迹种群)所在地。在每一调查点,测定了各样点的海拔高度、水体基底状况以及水体pH值等生境特征;在现存种群所在地,分别记载了所见水生植物种类、水韭种群规模及其生长状况,并采集和鉴定了3种水韭属植物及其伴生种。调查发现3种水韭属植物种与种之间为间断分布,且具有垂直梯度差异;种内各种群之间相互隔离,各种群的个体数均偏少;现存种群所生长的水体及其水体基底的pH特性无明显差异,为酸性或中性;绝迹种群所在的原生境已遭到严重破坏,一些地点的水体pH和基底pH与以往存在水韭的时期相比已明显升高。此外,调查还发现沼泽地带的水韭种群正面临着其他水生植物的激烈竞争而处于随时灭绝的危险之中,而淡水潮间带沿岸的水韭种群因周期性的水位波动而生长良好。调查结果表明3种水韭属植物的地理分布及其生长状况与海拔、水体pH值、基底pH值以及种间竞争和水位波动密切相关。本文还讨论了保护水韭属植物的基本策略和具体措施。     

The quillworts (Isoetes) of India: distribution, endemism and species radiation

The genus Isoetes L. in India is represented by 14species, of which eight species are recognized as being local endemics [confinedto one particular phyto-geographical division (PGD)] while the remaining sixoccur in more than one PGD and are described as endemic to a wider range. Thelocal endemic species are Isoetes dixitei,Isoetes panchganiensis and Isoetessahyadriensis in Western Ghats region; Isoetespantii, Isoetes bilaspurensis, Isoetesreticulata and Isoetes tuberculata inChotanagpur Malwa Vindhya Plateau and Isoetes debii innortheastern India. The wider endemic species are Isoetespanchananii, Isoetes sampathkumaranii,Isoetes rajasthanensis, Isoetesmahadevensis, Isoetes indica andIsoetes coromandelina. Our studies on the patterns ofendemism suggest that the radiation of quillworts advanced from dry lowlandareas to the rainy uplands and mountains. Isoetescoromandelina is the first Indian quillwort to colonize in lowlandsof the coastal zone (Coromandel), from where it spread to different parts of thesub-continent and gave rise to new species. Thus this species is a key Indianspecies which has played an important role in the radiation of quillwort in thecountry and appeared as the connecting link among the quillwort flora of variousPGDs. The centres of diversity for almost all the presently known Indianquillworts species are recognized.     

Studies of Diurnal Acid Fluctuations in British Isoetid-type Submerged Aquatic Macrophytes

British isoetid species are examined for the presence of diurnalfluctuations in tritratable acidity (to pH 6·4), in plantscollected directly from a small lake and in plants grown inconstant conditions in the laboratory. Wide diurnal fluctuationsare present in Isoetes lacustris and in both submerged and terrestrialpopulations of Littorella uniflora. They are absent in Lobeliadortmanna, Subularia aquatica, Eriocaulon septangulare, Ranunculusflammula and Pilularia globulifera. The significance of submerged CAM is discussed in relation toother carbon accumulating mechanisms in isoetids and in considerationof their general ecology. Crassulacean acid metabolism, photosynthesis, isoetid, oligotrophic lakes     

DISTRIBUTION OF DIURNAL ACID METABOLISM IN THE GENUS ISOETES

Previous studies have revealed a crassulacean acid metabolism-like pathway in the submerged aquatic Isoetes howellii (Isoetaceae). A survey of 11 other species in this largely aquatic genus showed all had diurnal fluctuations in titratable acidity and malic acid concentration. The magnitude of the diurnal fluctuation in titratable acidity varied from 119 (μeq g^–1 fresh weight) for I. bolanderi to 44 for I. engelmanni. Although these species were grown under similar conditions in a greenhouse, it is questionable whether or not any significance can be attached to observed species specific differences in magnitude of diurnal acid fluctuation. This is suggested by a single species comparison of plants in the field with greenhouse grown plants which indicated nearly as much variation in magnitude of acid fluctuations as observed across all species in the greenhouse. Much of this variation may be due to seasonal differences. Substrate appears to have very little effect on magnitude of acid fluctuations. Since this survey covered most of the range of the latitudinal, elevational, habitat, and morphological variation in the genus, it is likely that diurnal acid metabolism will prove to be common in the genus Isoetes.     

The structure of the apical meristem of Isoetes engelmannii, I. riparia and I. macrospora (Isoetales)

The shoot apex in three species of Isoetes (I. engelmannii, I. macrospora and I. riparia ) is differentiated into a cap-like initiation layer of somewhat columnar meristematic cells which divide most frequently anticlinally and a subadjacent central core of cells which divide irregularly, but with periclinal divisions predominating. The conflicting views of apical organization in Isoetes are discussed.     

中国水韭属植物的染色体数目及其分类学意义

利用细胞学方法观察统计了中国4种水韭属Isoestes植物的染色体数目。结果发现4个种的染色体基数均为x=11,其中高寒水韭I．hypsophila Hand.-Mazz．为2n=22,为首次报道;台湾水韭 I．taiwanensis DeVol为2n=22;中华水韭I．sinensis Palmer 2n=44为四倍体;而产于云南贵州一带并一直被当作是宽叶水韭I．japonica A．Br．的水韭属植物2n=22,与文献报道的宽叶水韭(2n=66,67,77,87,88,89)完全不同,应重新予以认识和评价。依据本文和其他相关的研究结果,对东亚水韭属植物进行了分类处理。     

Autecology and distribution of Isoetes longissimum in Europe

Isoetes longissimum is a rare aquatic pteridophyte known only from the northwest Spanish region of Galicia and from Algeria. We mapped the distribution of this species in Galicia, and investigated the relationship between its presence or absence and water quality variables. I. longissimum is present in sixteen 10 times 10 km grid squares in Galicia, largely in the western central part of the region. Within Galicia, I. longissimum shows a significant preference for sites with low sulphate, ammonium and phosphate concentrations.     

极濒危植物中华水韭屯溪居群调查研究报告

中华水韭（Isoetes sinensisPalmer）为水韭科（Isoetaceae）水韭属（Isoetes）多年生沼泽植物,由于人为干扰日益严重,现分布范围急剧缩小,种群数量稀少,处于濒危状态,已被列为国家一级重点保护野生植物。为科学合理的保护和抢救这一珍稀濒危植物资源提供理论依据,为此对中华水韭屯溪居群的生长环境、面积、株丛数量、水体的pH值、伴生植物种类和生存现状等的调查研究。中华水韭屯溪居群面积500m^2,株丛密度较大,水体pH值为6.55,伴生植物有40余种,其中10种有较高优势度,生存面临多重威胁,保护迫在眉睫。     

中国3种水韭属植物自然居群水体化学性质特征及差异性研究

 通过测定20个化学因子,利用平均值、变异系数和差异显著性检验,分析研究了中国3种水生水韭属植物中华水韭(Isoetes sinensis)、云贵水韭(I. yunguiensis)和高寒水韭(I. hypsophila)自然居群水体化学性质的特征和差异性。结果显示：这３种水韭属植物生活水体的20个化学因子整体上表现出Ni含量占绝对优势、电导率较低、受干扰较小的共同特征;虽然平均值和变异系数变化较大,但除了pH、P、NO3-N、NO3-和Al具有显著性差异以外,其余各因子在3个种的生活水体间相对稳定不具显著性差异。结合其它的研究结果分析表明：这3种水韭属植物自然居群水体化学性质的差异可能源于这3种水韭属植物生理特征的差异性,对3种水韭属植物的分布不具选择差异性。     

Evidence of crussulacean acid metabolism in two North American isoetids

The internal acidity levels of four common North American isoetids, or rosette-form aquatic macrophytes, were monitored under field conditions to determine the presence of crassulacean acid metabolism (CAM). The leaves of Littorella uniflora var. americana (Fern.) Gl. and Isoetes macrospora Durieu were found to have a diurnal fluctuation in titratable acidity characteristic of CAM plants, 124 and 142 μeq g^?1 fresh wt., respectively. No variations were detected in the acidity of stems and leaves of Myriophyllum tenellum Bigel. and Lobelia dortmanna L. Litorella and Isoetes were then grown in the laboratory where the diurnal acid rhythm was shown to be due to the accumulation and disappearance of malic acid.Based on the magnitude of the diurnal acid rhythm and existing information on the productivity of these plants, it appears that the carbon assimilated via crassulacean acid metabolism may contribute substantially to their net annual productivity. This appears to be a case for which CAM has been selected directly as a response to carbon stress.     

中华水韭雌配子体及胚胎发育的初步研究

中华水韭（Isoetes sinensis Palmer）属水韭科（Isoetaceae）草本拟蕨类,是长江中下游特有的古老物种,对环境污染敏感,为国家一级重点保护的极度濒危的野生植物。本研究通过人工培养,按不同发育阶段,用常规石蜡切片法连续观察了雌配子体及胚胎发育的全过程。结果显示：大孢子培养15～20 d萌发,雌配子体为球状、属壁内发育型,每个雌配子体着生8～20个颈卵器。成熟的颈卵器发育有多种可能,但只有颈沟细胞和腹沟细胞都退化,卵才有受精的机会。合子首先进行横向分裂,培养约50 d发育成数十个细胞的球形胚胎,约80 d雌配子体的细胞萎缩,第一叶及其叶舌、第一根结构更明显,出现第二叶。文中还初步探讨了中华水韭濒危的生殖生物学机制。     

Casparian Strips in the Leaf Intrastelar Canals of Isoetes duriei Bory, a Mediterranean Terrestrial Species

Anatomical observations, using light and fluorescence microscopy,were made on leaves of Isoetes duriei to verify the presenceof casparian bands around the intrastelar canals. This peculiaranatomical feature, previously reported for some Isoetes species,is confirmed. The possible role of this endodermis-like structure,in the transport of water due to root pressure or in the accumulationof certain metabolites, is discussed in relation to the ecologicaland anatomical features of the species. Copyright 2000 Annalsof Botany Company Isoetes duriei, Lycopsida, Mediterranean, leaf anatomy, endodermis, casparian strips, fluorescence microscopy     

Observations on Isoetes in Ghana

Three species of Isoetes, I. abyssinica Chiovenda, I. nigritiana A. Br. and I. tenuifolia A. C. Jermy, here described for the first time, are reported from Ghana. All species are found in seasonally saturated or flooded hollows and species lists of associate plants are given for the sites and are discussed in relation to water and soil analyses.
The morphology and anatomy of the sporophylls of all three species are compared, and the taxonomic value of the pseudo-ligule, idioblasts projecting into the four lacunae of the leaf ('internal hairs') and intrastelar canals is evaluated. The first chromosome count for a West African species of Isoetes is reported as 2 n = c . 58.     

Silicon in sporoderms of micro- and megaspores of Isoetes echinospora Durieu registered by EDS and EELS

Protoplasma - The present study reveals silica in sporoderms of micro- and megaspores of the modern quillwort Isoetes echinospora Durieu and homologizes layers of the sporoderm in spores of this...     

Ultrastructure, Origin, and Composition of the Protein Bodies in the Ligule of Isoetes lacustris L.

The basal cells in the ligule of Isoetes lacustris contain numerousprotein bodies, the contents of which can be digested enzymicallyby pronase and are stained red by treatment with ninhydrin Schiff'sreagent. Two types of protein bodies can be distinguished ultrastructurally:spherically-shaped bodies with granular contents and spindle-likebodies with fibrillar contents. Both are ensheathed by singlemembranes and do not show any solid inclusions within theirmatrix. The protein bodies probably arise from dilatation of the endoplasmicreticulum (ER) cisternae. This conclusion is based upon threeobservations: (a) The protein bodies occasionally show membranecontinuity with the ER; (b) ribosomes and polysomes are frequentlyattached to the protein-body membranes; (c) the contents ofthe protein bodies and of the dilated ER cisternae show similarultrastructural features. The dilatation of the ER cisternae is assumed to be a resultof protein accumulation in the intracisternal space. Based upon the results of polyacrylamide gel electrophoresis,it is likely that the spherically-shaped protein bodies storepredominately two proteins with molecular weights of 51300 and55800 D, while the spindle-like bodies store two proteins withmolecular weights of 92000 and 98000 D. The results presented do not permit a definite conclusion regardingthe function of the ligule of Isoetes lacustris but it is suggestedthat it may have a nutritive role. Isoetes lacustris L., ligule, protein bodies, endoplasmic reticulum, ultrastructure     

Macrophytes from lakes in the eastern Pyrenees: community composition and ordination in relation to environmental factors

1. Thirty-six species of macrophytes (fourteen flowering plants, two quillworts, sixteen mosses and liverworts, and two algae) were collected in an extensive survey of 116 high mountain lakes in the eastern Pyrenees. Seventy per cent of the lakes showed macrophyte development. 2. The isoetids (Isoetes lacusiris, Isoetes setacea and Subularia aquatica) were the dominant growth form, although the natopotamid Sparganium angusifolium was the most widespread species. Potamids (Potamogeton spp., Ranunculus spp.), the alga Nitella gr. opaca and some mosses (Warnstorfia exannulata, Sphagnum denticulatum) were often present. 3. A multivariate ordination analysis (RDA-redundancy analysis) revealed that water chemistry, altitude and vegetation cover of the catchment, and nutrient availability are major environmental factors associated with macrophyte distribution along the eastern Pyrenean lakes. Isoetids prevailed in softwater oligotrophic lakes, potamids in relatively hardwater oligotrophic lakes, and Potamogeton natans and Callitriche palustris in small and eutrophic water bodies affected by the presence of cattle.     

Root Cap Structure in Isoetes macrospora Dur

Root meristem cells of Isoetes macrosporausually have one plastidwhich is associated with the prominent nucleus, numerous ribosomesand mitochondria, and small vacuoles. During mitosis each plastidappears to replicate so that each daughter cell contains oneplastid. The cell walls of the meristem cells are traversedby numerous plasmodesmata. Central cells of the root cap lackdistally displaced plastids but have one or more amyloplastsassociated with the nucleus. These cells also contain largeprotein deposits. Peripheral root cap cells are characterizedby being vacuolated, and by possessing a few dictyosomes andprotein deposits. They appear to be sloughed infrequently. Isoetes macrospora Dur, root cap, protein bodies, ultrastructure     

基于核基因和叶绿体基因的台湾水韭遗传多样性和遗传结构的分析

利用核基因和叶绿体基因对台湾水韭（Isoetes taiwanensis Devol）2个居群20个样本的遗传多样性和遗传结构进行了分析,并对台湾水韭的保育提出了建议。核基因的检测结果显示,2个居群共存在18个单倍型,单倍型多样性为0.9842,核苷酸多样性为0.00215,居群间基因流N_m为4.26,居群间分化系数G_st值为0.05543;叶绿体DNA的检测结果显示,单倍型数目为6,单倍型多样性为0.6211,核苷酸多样性为0.00326。其中台北居群的单倍型多样性和核苷酸多样性均比金门居群高。核基因和叶绿体基因的AMOVA分析显示,居群内的遗传差异远高于居群间。台湾水韭的核苷酸多样性相对其他水韭属植物较低,可能与台湾水韭的染色体倍型以及狭域分布有关。居群结构的形成可能和基因流以及奠基者效应有关。宜采用原地和迁地保护的方法对居群进行保护。     

MEGASPORE WALL ULTRASTRUCTURE IN ISOETES

In Selaginella, megaspore wall ultrastructure (unit morphology and arrangement) is correlated with taxonomic position. In some Pennsylvanian lycopods there is a correlation with dispersal strategy. This study was designed to assess any correlation between habitat preference and wall ultrastructure in Isoetes. Except for a few minor structural correlations, wall ultrastructure appears to be constant in terrestrial, amphibious, and aquatic species of Isoetes. Several distinct sizes of megaspores occur in some megasporangia and correspond to 1) full-size megaspores with siliceous coatings, 2) small megaspores with siliceous coatings, and 3) full-size megaspores without siliceous coatings. The uniformity in wall ultrastructure within modem Isoetes together with the lack of uniformity of fossil isoetalean megaspore wall ultrastructure suggest that the modem species of Isoetes are closely related.