Wall extensibility, wall pH and tissue osmolality: significance for auxin and ethylene-enhanced petiole growth in semi-aquatic plants

Abstract. Auxin and ethylene both enhance cell elongation in intact petioles of the semi-aquatic plants Regnellidium diphyllum and Nymphoides peltata. The authors now show that auxin but not ethylene increases the in vitro extensibility of cell walls. No response to ethylene occurs in auxin-depleted tissue. Neither hormone regulates cell expansion by direct control of internal osmolality (OS). During growth of segments, OS (and hence turgor) declines rapidly over the first 5–6 h with a net loss of osmotic solutes. Thereafter, an apparent threshold OS is maintained with net gains in osmostic solutes ( Nymphoides ) or further net losses ( Regnellidium ). Although wall extensibility determines initial rates of hormone-induced cell expansion, the primary control of wall loosening appears to differ in the two species. Nymphoides shows typical 'acid growth', and fusicoccin, auxin and ethylene (with auxin) all enhance proton secretion. In Regnellidium , neither low pH nor fusicoccin (FC) alters the rate of cell expansion, although proton secretion is stimulated by FC. Stress relaxation studies using low pH treatment of living or frozen-thawed segments show increases in the extensibility of walls in vitro for Nymphoides but not for Regnellidium. The authors propose that extensibility may be controlled by wall pH in Nymphoides but the availability of effective wall-loosening sites determines extensibility in Regnellidium.     

Calcium-Dependent Lamina Production of Nymphoides peltata (Gmel.) O. Kuntze (Menyanthaceae): Implications for Distribution

Nymphoides peltata (Gmel.) O. Kuntze, a nymphaeid macrophyte,occurs commonly in polder and fluviatile areas in large partsof Europe and Asia. In contrast to the nymphaeid macrophytesNymphaea alba L. and Nuphar lutea (L.) Sm., Nymphoides peltatais almost completely absent from poorly-buffered waters andis never found in acid water bodies. Transplantation experimentsin water bodies of varying alkalinity demonstrated that, irrespectiveof the sediment type, leaf production of Nymphoides did occurin poorly-buffered waters, but not in acid waters. Cultivation experiments showed that floating leaf developmentof Nymphoides peltata could only take place if sufficient calciumwas available in the water layer or in twice-demineralized water.Addition of calcium to an acid cultivation medium or to watercollected from an acid moorland pool resulted in leaf production.Growth of Nymphoides in acid waters is impossible due to insufficientcalcium concentrations in the water layer of such waters. Itis suggested that the absence of Nymphoides peltata in somepoorly-buffered water bodies is partly due to the spatial isolationfrom rivers and canals and the high frequence of desiccation.The restricted occurrence of Nymphoides peltata to well-bufferedalkaline waters is functionally more related to the calciumavailability than to the bicarbonate content. Key words: Aquatic macrophytes, distribution, Nymphoides peltata, leaf production, calcium, acid, poorly-buffered and alkaline water     

Ontogeny of stomata in some Gentianaceae

The mature epidermis and development of stomata in some species of Gentianaceae, viz. Canscora deccusata, C.diffusa, Hoppea dichotoma, Enicostema littorale, Nymphoides cristatum and Gentiana pedicellata , are described. The stomatal development is shown to be trilabrate mesogenous in Canscora deccusata, C.diffusa, Enicostema littorale and Hoppea dichotoma , mesoperigenous in Gentiana and perigenous in Nymphoides. It is suggested that adoption of an aquatic habit by Nymphoides may have resulted in suppression of divisions in its perigenous stomata.     

Germination requirements and seed banks of some nymphaeid macrophytes: Nymphaea alba L., Nuphar lutea (L.) Sm. and Nymphoides peltata (Gmel.) O. Kuntze

SUMMARY.

• 1 Germination experiments demonstrated that the innate dormancy of the seeds of Nymphaea alba L., Nuphar lutea (L.) Sm. and Nymphoides peltata (Gmel.) O. Kuntze could be overcome by a cold treatment. Light stimulated the germination of the three species. Hypoxic conditions stimulated the germination of Nymphaea alba and Nuphar lutea seeds but the seeds of Nymphoides peltata did not germinate under these conditions.
• 2 Experimental seed banks of Nymphaea alba, Nuphar lutea and Nymphoides peltata were laid out in three water bodies, varying in pH and alkalinity. Germination patterns indicated that Nymphaea alba and Nuphar lutea produce transient seed banks, but that Nymphoides peliata produces a persistent seed bank. Sampling of natural seed banks and subsequent germination tests were in concordance with the results of the seed bank experiment.
• 3 The experimental above-ground seed banks of Nymphaea alba, Nuphar lutea and Nymphoides peltata showed similar germination patterns in the three selected water bodies, despite the differences in pH and alkalinity between them. However, the distribution of Nymphoides peliata is restricted to well-buffered waters, so that its absence from soft and acid water bodies must be due to post-germination mechanisms and/or processes.
• 4 In aquatic systems where Nymphoides peltata co-exists with the other nymphaeid species studied, it is largely restricted to a bell between the helophytes and the vegetation at deeper sites. The deeper sites were dominated by Nuphar lutea and Nymphaea alba. Germination requirements and seedling emergence from buried seeds of Nymphaea alba, Nuphar lutea and Nymphoides peliata play an important role in the establishment of the zonation pattern of these nymphaeid macrophytes.

     

Floral morph composition and pollen limitation in the seed set of <Emphasis Type="Italic">Nymphoides indica</Emphasis> populations

We investigated the floral morph composition and seed set of the populations in a heterostylous aquatic plant, Nymphoides indica O. Kuntze (Menyanthaceae). The bias of floral morph ratio in the populations was negatively correlated with the seed set. In the populations with biased floral morph ratio, the pollen on the stigma of the common morphs were dominated by pollen of the same morph, resulting in reduced seed set. The addition of pollen of the opposite morph on the stigmas of the common morph resulted in an increase of fruit and seed set rates. The intermorph pollination rate and the fruit and seed set rates of Nymphoides indica were highest when two floral morphs were close in the populations. Various insect species including Lepidoptera, Diptera and Hymenoptera visited the flowers. However, geitonogamous pollination was common because of the short flying distance of the pollinators. These results indicate that the seed set of Nymphoides indica is limited by the shortage of compatible pollens and that the success of intermorph pollination is influenced by spatial distribution of the two floral morphs. As Nymphoides indica is endangered, conservation of the populations with high fecundity is proposed by us.     

水生植物的生态敏感度研究

水生生态系统是否稳定 ,总是以构成系统的各生物种群的结构及其相互关系的动态变化为判定标准。研究水生植物种类对环境因子的生态效应 ,确定不同种类的生态敏感度 ,为水生生态系统的敏感区域划定与保护和评价受破坏的生态系统的功能状况提供理论依据和方法。据美国和瑞典的资料报道 [4 ] ,生态敏感度概念已被引入生态规划 ,作为重要的规划工具之一 ,这是一种新的规划思路和方法。1　确定生态敏感度的原则1生态敏感度是指生物体对来自生存环境中的各种因子的影响所作出的反应。环境因子可以是单个的 ,也可以是复合的 ,因此生态敏感度表现为…     

Modeling Nymphoides architecture: A morphological analysis of Nymphoides aquatica (Menyanthaceae)

? Premise of the study: Species in the aquatic genus Nymphoides have inflorescences that appear to arise from the petioles of floating leaves. The inflorescence-floating leaf complex can produce vegetative propagules and/or additional inflorescences and leaves. We analyzed the morphology of N. aquatica to determine how this complex relates to whole plant architecture and whether whole plant growth is sympodial or monopodial. ? Methods: We used dissections, measurements, and microscopic observations of field-collected plants and plants cultivated for 2 years in outdoor tanks in south Florida, USA. ? Key results: Nymphoides aquatica had a submerged plagiotropic rhizome that produced floating leaves in an alternate/spiral phyllotaxy. Rhizomes were composed of successive sympodial units that varied in the number of leaves produced before the apex terminated. The basic sympodial unit had a prophyll that subtended a renewal-shoot bud, a short-petioled leaf (SPL) with floating lamina, and an inflorescence; the SPL axillary bud expanded as a vegetative propagule. Plants produced either successive basic sympodial units or expanded sympodia that intercalated long-petioled leaves between the prophyll and the SPL. ? Conclusions: Nymphoides aquatica grows sympodially, forming a rhizome composed of successive basic sympodia and expanded sympodial units. Variations on these types of sympodial growth help explain the branching patterns and leaf morphologies described for other Nymphoides species. Monitoring how these two sympodial phases are affected by water depth provides an ecologically meaningful way to assess N. aquatica's responses to altered hydrology.     

水生维管束植物对水体铅污染的反应、抗性和净化作用

重金属铅污染环境的一个主要方面是对水体的污染。主要的污染源是某些矿山(如铅锌矿等)、有色金属冶炼厂、铅制品加工厂等的废水和废气以及汽车尾气等。按金属的生物效应来分类,铅为有毒金属。大量的铅直接或者通过食物链的富集进入人体会造成严重的危害。目前,国内外对于铅污染的环境对陆生植物的影响以及陆生植物对铅污染的反应开展了一些     

A study on annual changes in surface cover of floating-leaved plants in a lake using aerial photography

A simple method of aerial photography was used to investigate annual and between-year changes in surface cover of the floating-leaved plants, Nymphoides peltata, Nelumbo nucifera and Trapa natans, in natural stands in Lake Kasumigaura. Oblique photographs of vegetation were taken with a 35-mm camera with real color film from airplanes between 1983 and 1989. Vegetation maps were traced on the positions after projective transformation to vertical photography. The distribution of the stand of Nymphoides peltata seemed to be restricted to a region of less than 1.5 m water depth. A rise of 1.0 m in the water level after a typhoon in 1986 caused a subsequent decrease in vegetation area of Nelumbo nucifera. The vegetation recovered in shallower regions after one year in 1987. The expansion rate of Nymphoides peltata and Nelumbo nucifera in one direction was in the range 3.8–10.0 m yr^-1 and 4.1–20.8 m yr^-1, respectively. Nelumbo nucifera, Nymphoides peltata and Trapa natans, having high leaf biomass to total biomass may therefore be more vulnerable to large flood and waves. Fluctuation in water level and spatial difference in water depth may also be important factors determining the vegetation area and survival of Nymphoides peltata, Nelumbo nucifera and Trapa natans.     

Cytochemistry and Ultrastructure of the Mucilage Secreting Trichomes of Nymphoides peltata (Menyanthaceae)

The young developing leaves in the buds of Nymphoides peltataare covered by a hyaline mucilage. The mucilage contains freesugars, polysaccharides and proteins. The most abundant monosaccharidesof the polysaccharide fraction are arabinose and galactose.Therefore, the major component of the mucilage is probably anarabinogalactan or arabinogalactan protein. The mucilage issecreted by glandular trichomes. It is suggested that both thepolysaccharide and the protein fraction of the mucilage aretransported to the plasmamembrane by vesicles of the Golgi apparatus(granulocrine secretion). Secretory proteins are probably synthesizedin the rough endoplasmic reticulum and transported to the Golgiapparatus via transition vesicles. Polysaccharides were localizedin Golgi vesicles by ultracytochemistry. After exocytosis thesecretion is accumulated between the cell wall and the cuticle;this leads to the formation of protrusions on the outer wallsof the glandular cells. Finally, the cuticle is ruptured andthe secretion is released. The biological function of the mucilageis not known. Possibly the mucilage is a lubricant or a protectionfrom desiccation. Nymphoides peltata (S.G. Gmel.) O. Ktz., trichomes, mucilage secretion, cytochemistry, ultrastructure     

西藏睡菜科一新记录属——莕菜属

报道西藏一个新记录属,是睡菜科(Menyanthaceae)的莕菜属(Nymphoides),并报道该地区的一个新记录种,即莕菜(Nym phoides peltatum(Gmel.)O.Kuntze).     

Aquatic macroinvertebrate community structure of aNymphoides peltata-dominated and macrophyte-free site in an oxbow lake

The aquatic macroinvertebrates in two freshwater biotopes,viz. aNymphoides peltata-dominated site and a macrophyte-free site, were studied quantitatively in a shallow alkaline oxbow lake of the river Waal, the main branch of the river Rhine in The Netherlands. The research comprised the analysis of water, sediment and macrophyte samples.In the macrophyte-free site Oligochaeta and Nematocera, particularly of the collector gatherer functional feeding group, dominated the prevailing benthic community. The total macroinvertebrate biomass ranged here from 0.3 to 0.9 g ash-free dry weight per m² of biotope.Species richness, densities, and biomass of macroinvertebrates were considerably higher in the biotope dominated byNymphoides peltata. Many taxa were found associated with the aboveground macrophyte. The sediment compartment, however, contributed most to the total density and biomass of macroinvertebrates. Nematocera and Oligochaeta were the most abundant fauna groups, whereas the largest share in total biomass was provided by clams (Mollusca). The biomass of the total macroinvertebrate community in theNymphoides-dominated site ranged from 6.2 to 7.5 g ash-free dry weight per m² of biotope. The biomass of the aboveground phytophilous fauna ranged from 0.1 to 0.6 g ash-free dry weight per m² of biotope. In September, when theNymphoides peltata vegetation was in its senescent phase, the largest numbers and the highest biomass of phytophilous macroinvertebrates were observed. The contribution of the shredder functional feeding group was high in this period. This, and the overall high abundance of fauna with a detritivorous mode of life, indicates the importance of macrophyte detritus as input to food chains.     

Reproductive consequences of interactions between clonal growth and sexual reproduction in Nymphoides peltata: a distylous aquatic plant

Distyly is a sexual polymorphism in which plant populations contain two floral morphs differing in morphology and physiology. The dimorphism serves to promote animal-mediated cross-pollination between the floral morphs. Clonal propagation can interfere with the functioning of distyly by compromising intermorph pollinations, resulting in reduced fertility. Here, we investigate the relations between clonal growth and sexual reproduction in the aquatic macrophyte Nymphoides peltata (Menyanthaceae). Surveys of morph representation in 30 populations from five regions of China revealed that most populations exhibited strongly biased morph ratios and 30% contained a single floral morph. Experimental pollinations indicate that N. peltata possesses a strong dimorphic incompatibility system preventing self and intramorph fertilizations. An experiment involving the manipulation of morph ratios in an experimental population and an investigation in a natural population with strong morph substructure both provided evidence that compatible pollen dilution limits fertility. Despite constraints on the functioning of distyly in N. peltata we found no evidence for evolutionary changes to the heterostylous syndrome, as reported in Nymphoides, including populations of N. peltata in other parts of its geographical range.     

Phenols and phenol oxidases are involved in cadmium accumulation in the water plants Nymphoides peltata (Menyanthaceae) and Nymphaeae (Nymphaeaceae)

This comparative study investigates the mechanism of cadmium accumulation in the semiaquatic plant Nymphoides peltata (Menyanthaceae) and the aquatic plant Nymphaea (Nymphaeaceae). It was conducted as part of an ongoing study of the use of water plants for phytoremediation. Epidermal structures, known as hydropotes, are located on the abaxial epidermis of the leaf laminae of Nymphoides peltata and are shown to contain phenols, peroxidase and polyphenol oxidase activities. When plants are subjected to 50 mg/l of cadmium in the growth medium, these hydropotes accumulate cadmium. Cadmium-induced increases in phenols, peroxidase and polyphenol oxidase activities were determined in plant extracts. Cadmium binding by polymerized phenols was demonstrated in vivo. In comparison with Nymphaeae epidermal glands, N. peltata hydropotes are larger, open, and create bigger crystal, the latter principally composed of calcium and, proportionally, less cadmium. Although both plants showed similar levels of cadmium accumulation, N. peltata was sensitive while Nymphaeae was resistant to this cadmium level. It is suggested that in these water plants the main mechanism for cadmium accumulation is based on the trapping of cadmium crystals by polymerized phenols in specialized epidermal structures and this is due to peroxidase and polyphenol oxidase activities. Nymphaeae, with greater peroxidase activity and more polyphenols, is more resistant to this heavy metal than N. peltata.     

Isolation and characterization of highly polymorphic microsatellites in the aquatic plant,Nymphoides peltata (Menyanthaceae)

Ten microsatellite loci were described for conservation design of a threatened clonal aquatic plant, Nymphoides peltata. The microsatellite loci obtained through the construction of an enriched library were polymorphic (2–6 alleles per locus) and exhibited high levels of observed (0.333–0.889) and expected (0.284–0.765) heterozygosities. All microsatellite loci were expected to be useful for identification of genets and evaluation of genetic diversity of this species.     

Cell Expansion and Wall pH in the Fern Regnellidium diphyllum, A Plant Lacking Acidinduced Growth

Petioles of the semi-aquatic fern Regnellidium diphyllum donot show acid growth but low wall pH is a necessary conditionfor maximum rates of IAA-induced cell expansion. Measurementsof wall pH by two indirect methods indicate an unusually lowvalue, in the range pH 4 to 5. This is one to two pH units belowthat estimated for petioles of the semi-aquatic dicotyledonNymphoides peltata, a species in which IAA aand ethylene causegrowth responses very similar to those in Regnellidium but whereacid growth occurs. Having shown previously that fusicoccinenhances proton secretion in both Regnellidium and Nymphoides,we now show that although it causes a reduction in the estimatedapoplast pH to below 4·0 in Regnellidium, cell expansionis not promoted. The FC-induced reduction in pH in Nymphoidesis less and occurs more slowly, but growth is promoted significantly;when IAA and fusicoccin are present together, growth promotionis approximately additive for Nymphoides A model is proposed for Regnellidium in which equilibrium wallpH is maintained at a low value that is optimal for acid growth,the availability of acid-labile sites in the wall being thechief limitation to cell extension. We suggest that this controlmechanism may be widespread for organs without a cuticle, includingroots and the gametophytes of lower plants growing in acidicconditions. Key words: Acid growth, wall pH, fusicoccin, Regnellidium diphyllum, Nymphoides peltata     

Phylogenetics and seed morphology of African Nymphoides (Menyanthaceae)

We evaluated the boundaries among Nymphoides (Menyanthaceae) species in Africa, a region where the genus has received relatively little attention. We gathered morphological data from seeds using light and scanning electron microscopy, and we conducted molecular phylogenetic analyses using nuclear sequence data from the internal transcribed spacer (ITS) region. Morphological and molecular features distinguished six species and affirmed their respective geographic ranges. No African specimens were attributable to Nymphoides indica, even though this paleotropical species previously had been understood to grow in Africa. We establish the new combination Nymphoides senegalensis (G. Don) Tippery, based on an established African basionym, to accommodate the specimens formerly identified as N. indica. Phylogenetic analyses resolved two distinct clades containing African species. The majority of African species are closely related to neotropical species, with which they share similar petal ornamentation. The morphologically distinct N. ezannoi shares floral and phylogenetic similarity with species from North America and Asia. Results presented here support prior hypotheses that allopolyploid species in the Americas may have originated from one or more parental lineages in Africa. Seed morphological characters remain some of the most reliable features for identifying species, particularly for herbarium specimens lacking observable floral characters.     

Nitrogen and phosphorus budgets of a nymphaeid-dominated system

Summary Communities dominated byNymphoides peltata (Gmel.) O. Kuntze (Menyanthaceae) have been studied within the framework of the project on structure, dynamics and function of nymphaeid-dominated systems. VAN DER VELDE, 1980). In this summary some remarks on the quantitative distribution of N and P in an old river bed are presented with special emphasis on sites dominated by the floating-leaved macrophyteNymphoides peltata. The study area was the Bemmelse Strang, an old branch of the river Waal situated in the northern river forelands between Lent and Bemmel, north of Nijmegen. Two sampling sites were chosen,viz. one dominated byNymphoides peltata and another without macrophytes (the open water). Several compartments were distinguished in these sites,viz., water, seston, sediment and macrophytes. Water and seston samples were collected biweekly and sediment and macrophyte samples monthly from January until December 1980. In the sediment most nutrients seemed to be stored in the upper layers, so cores to a depth of 10 cm were taken. The percentage of water in the sediment, the specific mass of the sediment, the biomass of seston andNymphoides (expressed as dry weight) and the water depth were recorded at each sampling data. The dried samples of sediment, seston andNymphoides were analysed for total N and total P. In the water and interstitial water (soil extracts) total N, NH₄ ⁺, NO₃ ^––NO₂ ^–, total P and PO₄ ^3– were determined. By means of these data the quantitative distribution of nitrogen and phosphorus in the system could be established.     

The Fine Structure of the Sieve Tubes of the Petiole of Nytnphoides peltatum (Gmel.) O. Kunze

A study has been made by electron microscopy of the fine structureof the peti-olar sieve tubes of the water plant Nymphoides peltatum.These are found to have very well-developed nacreous walls.The pores of the sieve plates appear to be filled in functioningsieve tubes with densely staining cytoplasm. The peripheralcytoplasm of the sieve tubes seems to contain an extensive developmentof the endoplasmic reticulum, whose elements become finer nearthe plates and crowd together through the pores. These findingsappear to be compatible with more than one theory of translocation,including the electro-osmotic theory of mass flow.     

Hg^2＋对金银莲花根和叶片的伤害

研究了汞离子胁迫下金银莲花〔Ｎｙｍｐｈｏｉｄｅｓｉｎｄｉｃａ（Ｌ．）Ｋｕｎｔｚｅ〕根受害情况、根部过氧化物酶活性和新叶叶绿素含量的变化。根部受害程度随汞离子浓度升高和处理时间的延长而加重。低浓度Ｈｇ２＋短时间处理后根过氧化物酶活性升高,而高浓度Ｈｇ２＋长时间处理后根过氧化物酶活性下降,且随着Ｈｇ２＋浓度的升高或处理时间的延长过氧化物酶活性均呈下降趋势。新叶叶绿素含量对Ｈｇ２＋胁迫的反应与根过氧化物酶活性的变化相似。