Chemical composition of salt-marsh plants from Ynys Môn (Anglesey): the concept of physiotypes

Abstract The chemical compositions of a number of halophytes from salt marshes on Ynys Môn (Anglesey), Wales, and of some related mesophytes and sand dune plants have been determined. Analyses of the inorganic ions broadly confirmed the existence of a characteristic chemical composition of many monoco-tyledonous salt-marsh plants in that they contain high levels of potassium and relatively low levels of sodium. In contrast to most dicotyledonous halophytes, especially members of the Chenopodiacease, the monocots restrict the entry of inorganic ions and use high levels of soluble sugars to maintain an adequate solute potential. Low calcium levels were not found to be a feature of these plants, as was previously reported. The large amounts of sugars found in the monocotyle-donous plants suggested that they must be located mainly in the vacuoles, in contrast to glycinebetaine which is thought to accumulate principally in the cytoplasm of the salt accumulating Chenopodiaceae. The monocotyledonous halophytes which accumulate proline differ from the normal monocotyledonous physiotype in the accumulation of larger quantities of sodium. Triglochin maritima is one species of this type, and Puccinellia maritima a less extreme example. Spartina spp. accumulating glycinebetaine and β-dimethyl-sulphoniopropionate also have unusually high inorganic ion contents for monocots. Several salt marsh plants contained large quantities of amino acids other than proline. As with ionic composition, the nature of the organic solutes broadly followed taxonomic lines. The usefulness of the physiotype concept is discussed.     

Macroevolutionary patterns of salt tolerance in angiosperms

Background Halophytes are rare, with only 0·25 % of angiosperm species able to complete their life cycle in saline conditions. This could be interpreted as evidence that salt tolerance is difficult to evolve. However, consideration of the phylogenetic distribution of halophytes paints a different picture: salt tolerance has evolved independently in many different lineages, and halophytes are widely distributed across angiosperm families. In this Viewpoint, I will consider what phylogenetic analysis of halophytes can tell us about the macroevolution of salt tolerance.Hypothesis Phylogenetic analyses of salt tolerance have shown contrasting patterns in different families. In some families, such as chenopods, salt tolerance evolved early in the lineage and has been retained in many lineages. But in other families, including grasses, there have been a surprisingly large number of independent origins of salt tolerance, most of which are relatively recent and result in only one or a few salt-tolerant species. This pattern of many recent origins implies either a high transition rate (salt tolerance is gained and lost often) or a high extinction rate (salt-tolerant lineages do not tend to persist over macroevolutionary timescales). While salt tolerance can evolve in a wide range of genetic backgrounds, some lineages are more likely to produce halophytes than others. This may be due to enabling traits that act as stepping stones to developing salt tolerance. The ability to tolerate environmental salt may increase tolerance of other stresses or vice versa.Conclusions Phylogenetic analyses suggest that enabling traits and cross-tolerances may make some lineages more likely to adapt to increasing salinization, a finding that may prove useful in assessing the probable impact of rapid environmental change on vegetation communities, and in selecting taxa to develop for use in landscape rehabilitation and agriculture.     

Spatial and halophyte-associated microbial communities in intertidal coastal region of India

Microbial communities in intertidal coastal soils respond to a variety of environmental factors related to resources availability, habitat characteristics, and vegetation. These intertidal soils of India are dominated with Salicornia brachiata, Aeluropus lagopoides, and Suaeda maritima halophytes, which play a significant role in carbon sequestration, nutrient cycling, and improving microenvironment. However, the relative contribution of edaphic factors, halophytes, rhizosphere, and bulk sediments on microbial community composition is poorly understood in the intertidal sediments. Here, we sampled rhizosphere and bulk sediments of three dominant halophytes (Salicornia, Aeluropus, and Suaeda) from five geographical locations of intertidal region of Gujarat, India. Sediment microbial community structure was characterized using phospholipid fatty acid (PLFA) profiling. Microbial biomass was significantly influenced by the pH, electrical conductivity, organic carbon, nitrogen, and sodium and potassium concentrations. Multivariate analysis of PLFA profiles had significantly separated the sediment microbial community composition of regional sampling sites, halophytes, rhizosphere, and bulk sediments. Sediments from Suaeda plants were characterized by higher abundance of PLFA biomarkers of Gram-negative, total bacteria, and actinomycetes than other halophytes. Significantly highest abundance of Gram-positive and fungal PLFAs was observed in sediments of Aeluropus and Salicornia, respectively than in those of Suaeda. The rhizospheric sediment had significantly higher abundance of Gram-negative and fungal PLFAs biomarkers compared to bulk sediment. The results of the present study contribute to our understanding of the relative importance of different edaphic and spatial factors and halophyte vegetation on sediment microbial community of intertidal sediments of coastal ecosystem.     

北疆荒漠几种盐生(耐盐)植物抗逆附属结构的初步研究

利用石蜡切片、扫描电镜、临时装片等方法,对北疆荒漠的3种藜科植物—灰绿藜、费尔干猪毛菜、蒙古猪毛菜和1种菊科植物—花花柴的表皮附属结构进行了显微和超微观察研究。结果表明:(1)对四种植物的解剖结构观察显示,其叶片都含有角质层;气孔器下陷;茎中含有大量的维管束;多数种的细胞中含有簇状晶体结构;(2)四种植物表皮附属结构研究表明:花花柴表皮具有多细胞组成的盐腺和表皮毛结构;灰绿藜表皮有大量囊泡结构;蒙古猪毛菜叶表皮有短硬毛和乳突状结构;费尔干猪毛菜表皮具大量表皮毛,且表皮毛有节。上述结构和特征反映出不同植物对干旱、盐碱土生境适应的多样性,也为旱生和盐生植物的生理学研究提供了新的实验依据。     

In vitro salt tolerance of cell wall enzymes from halophytes and glycophytes

The halophyte Suaeda maritima grows optimally in high concentrations(40–60% seawater) of salt. In these conditions the concentrationof salt in the apoplast of the leaves is at least 500 mM, aconcentration which severely inhibits the activity of cytoplasmicenzymes of both glycophytes and halophytes. The in vitro salttolerance of a number of cell wall enzymes was assayed in thepresence of a range of concentrations of NaCl. There was nosignificant inhibition of the activity of galactosidase, glucosidase,peroxidase or xyloglucan endo-transglycosylase extracted fromSuaeda maritima by in vitro concentrations of NaCl up to atleast 1 M. In vitro salt tolerance of cell wall enzymes wasnot restricted to the halophyte, similar enzymes from the non-halophilicrelative Kochia tricophylla, and from the glycophytes Vignaradiata and Cicer arietinum, were inhibited little, or not atall, by the same concentrations of salt. Pectin esterase wassomewhat less tolerant, but activity at 500 mM NaCl was stillgreater than at 0 mM NaCl in both Suaeda and Vigna. It is concludedthat these enzymes of the cell wall compartment are much moresalt-tolerant than cytoplasmic enzymes of higher plants. Theresults are discussed in relation to conditions thought to pertainin the apoplast. Key words: Apoplast, cell wall enzymes, halophyte, salt tolerance, Suaeda maritima     

Structural,physiological, and biochemical aspects of salinity tolerance of halophytes

Modern concepts on structural, physiological, and biochemical aspects of salt tolerance of higher plants were considered. Integral physiological processes, such as growth and photosynthesis of glycophytes and halophytes in the context of their ecological plasticity, variety of their adaptive strategies developed in the course of their evolution, and natural selection, were discussed. Analysis of the known anatomical and morphological adaptations of halophytes (succulence, special salt-excreting structures, features associated with special tissues growth, leaf kranz-anatomy and mesostructure) providing their salt tolerance was conducted. The most important physiological and biochemical adaptations of such plants to salinity related to uptake, accumulation and excretion of Na⁺ and Cl^–, peculiarities of membrane composition and the pigment system, and protection against osmotic and oxidative stresses were described. The association of physiological and biochemical peculiarities of halophytes with ecological salt tolerance strategy was discussed.     

An efficient autofluorescence method for screening Limonium bicolor mutants for abnormal salt gland density and salt secretion

With the spread of saline soils worldwide, it has become increasingly important to understand salt-tolerant mechanisms and to develop halophytes with increased salt tolerance. Limonium bicolor is a typical recretohalophyte and has a typical salt excretory structure in the epidermis called the salt gland. A method that can be used to screen a large population of L. bicolor mutants for altered salt gland density and altered salt secretion is needed but is currently unavailable. Leaves of 1-month-old L. bicolor seedlings were processed by three traditional methods [epidermal peel, nail impression, and clearing/differential interference contrast microscope (clearing/DIC) method] and a fluorescence method (fluorescence microscopic examination of cleared leaves). With the fluorescence method, the autofluorescence of salt glands under UV excitation (330–380 nm) was easily distinguished with the least labor and time. The fluorescence method was used to screen ~ 10,000 seedlings (which grew from gamma-irradiated seeds). Four mutants with reduced salt gland density and 15 with increased salt gland density were obtained. Both kinds of mutants will be useful for the isolation of genes involved in salt gland development and salt secretion in L. bicolor and other halophytes. The fluorescence method was also successfully used to observe the salt glands of Aegialitis rotundifolia and the stomata and trichomes of Arabidopsis. The fluorescence method described here will be useful for examining plant epidermal structures that have autofluorescence under UV or other wavelengths.     

Halophyte Improvement for a Salinized World

It is more important to improve the salt tolerance of crops in a salinized world with the situations of increasing populations, declining crop yields, and a decrease in agricultural lands. Attempts to produce salt-tolerant crops have involved the manipulation of existing crops through conventional breeding, genetic engineering and marker-assisted selection (MAS). However, these have, so far, not produced lines growing on highly saline water. Hence, the domestication of wild halophytes as crops appears to be a feasible way to develop agriculture in highly saline environments. In this review, at first, the assessment criteria of salt tolerance for halophytes are discussed. The traditional criteria for the classification of salinity in crops are less applicable to strong halophytes with cubic growth curves at higher salinities. Thus, realistic assessment criteria for halophytes should be evaluated at low and high salinity levels. Moreover, absolute growth rather than relative growth in fields during a crop's life cycle should be considered. Secondly, the use of metabolomics to understand the mechanisms by which halophytes respond to salt tolerance is highlighted as is the potential for metabolomics-assisted breeding of this group of plants. Metabolomics provides a better understanding of the changes in cellular metabolism induced by salt stress. Identification of metabolic quantitative trait loci (QTL) associated with salt tolerance might provide a new method to aid the selection of halophyte improvement. Thirdly, the identification of germplasm-regression-combined (GRC) marker-trait association and its potential to identifying markers associated with salt tolerance is outlined. Results of MAS/linkage map-QTL have been modest because of the absence of QTLs with tight linkage, the non-availability of mapping populations and the substantial time needed to develop such populations. To overcome these limitations, identification by GRC-based marker-trait association has been successfully applied to many plant traits, including salt tolerance. Finally, we provide a prospect on the challenges and opportunities for halophyte improvement, especially in the integration of metabolomics- and GRC-marker-assisted selection towards new or unstudied halophyte breeding, for which no other genetic information, such as linkage maps and QTL, are available.     

Predicting species’ tolerance to salinity and alkalinity using distribution data and geochemical modelling: a case study using Australian grasses

Background and Aims Salt tolerance has evolved many times independently in different plant groups. One possible explanation for this pattern is that it builds upon a general suite of stress-tolerance traits. If this is the case, then we might expect a correlation between salt tolerance and other tolerances to different environmental stresses. This association has been hypothesized for salt and alkalinity tolerance. However, a major limitation in investigating large-scale patterns of these tolerances is that lists of known tolerant species are incomplete. This study explores whether species’ salt and alkalinity tolerance can be predicted using geochemical modelling for Australian grasses. The correlation between taxa found in conditions of high predicted salinity and alkalinity is then assessed.Methods Extensive occurrence data for Australian grasses is used together with geochemical modelling to predict values of pH and electrical conductivity to which species are exposed in their natural distributions. Using parametric and phylogeny-corrected tests, the geochemical predictions are evaluated using a list of known halophytes as a control, and it is determined whether taxa that occur in conditions of high predicted salinity are also found in conditions of high predicted alkalinity.Key Results It is shown that genera containing known halophytes have higher predicted salinity conditions than those not containing known halophytes. Additionally, taxa occurring in high predicted salinity tend to also occur in high predicted alkalinity.Conclusions Geochemical modelling using species’ occurrence data is a potentially useful approach to predict species’ relative natural tolerance to challenging environmental conditions. The findings also demonstrate a correlation between salinity tolerance and alkalinity tolerance. Further investigations can consider the phylogenetic distribution of specific traits involved in these ecophysiological strategies, ideally by incorporating more complete, finer-scale geochemical information, as well as laboratory experiments.     

不同类型盐生植物适应盐胁迫的生理生长机制及Na+逆向转运研究进展

盐生植物是指能在离子浓度至少200 mmol/L以上的生境中生长并完成生活史的植物。盐生植物可分为稀盐盐生植物、泌盐盐生植物、拒盐盐生植物三类。本文从生长形态、生理和分子3个方面总结三类盐生植物响应盐胁迫的不同策略及研究进展,发现盐生植物在分子水平上主要通过Na⁺转运蛋白和为其提供能量的两类基因应对体内过高Na⁺,这可能是引起盐生植物生理和生长形态异于非盐生植物的重要因素。其中稀盐盐生植物主要通过液泡离子区隔化应对盐胁迫,并表现出肉质化生长形态;泌盐盐生植物通过将体内盐分排出体外应对盐胁迫,并进化出特有的生理结构——盐腺或盐囊泡;拒盐盐生植物通过将盐离子积累在皮层细胞液泡和根部木质部薄壁细胞中减少向上运输Na⁺,同时根部多栓质化减少Na⁺吸收。本综述旨在为今后研究盐生植物及其耐盐机制提供相关依据,为植物耐盐分子育种奠定基础。     

碱蓬属植物耐盐机理研究进展

碱蓬属(Suaeda)植物是一类典型的真盐生植物,属于重要的盐生植物资源,全球广泛分布.人们已经对20种碱蓬属植物进行了观察和盐胁迫实验,研究了不同器官或组织的生理生化特征及其对盐胁迫的反应,并基于这些研究分析了盐胁迫的应答机制.叶片肉质化、细胞内离子区域化、渗透调节物质增加和抗氧化系统能力增强是碱蓬属植物响应和适应盐胁迫的重要方式和途径.但迄今为止的研究工作尚有一定的局限性,主要包括:研究工作主要集中在植物地上部分,而对植物地下部分的研究较少;多是少数生物学指标或生理学现象的单独观察,而缺乏对生理代谢过程的整体和综合分析;针对某种碱蓬的独立分析较多,而与近缘种的比较研究较少;植物对中性盐胁迫的反应研究较多,而对碱性盐的研究较少.为进一步系统阐明碱蓬属植物的耐盐机制,今后的工作应注重碱蓬属植物响应和适应盐胁迫的信号网络和调控机制研究,基于系统生物学研究思路,采用现代组学技术探索该属植物响应盐胁迫的由复杂信号网络调控的特殊生理特征和特异代谢途径.     

Halophytes--an emerging trend in phytoremediation

Halophytic plants are of special interest because these plants are naturally present in environments characterized by an excess of toxic ions, mainly sodium and chloride. Several studies have revealed that these plants may also tolerate other stresses including heavy metals based on the findings that tolerance to salt and to heavy metals may, at least partly, rely on common physiological mechanisms. In addition, it has been shown that salt-tolerant plants may also be able to accumulate metals. Therefore, halophytes have been suggested to be naturally better adapted to cope with environmental stresses, including heavy metals compared to salt-sensitive crop plants commonly chosen for phytoextraction purposes. Thus, potentially halophytes are ideal candidates for phytoextraction orphytostabilization of heavy metal polluted soils and moreover of heavy metal polluted soils affected by salinity. Some halophytes use excretion processes in order to remove the excess of salt ions from their sensitive tissues and in some cases these glandular structures are not always specific to Na+ and Cl- and other toxic elements such as cadmium, zinc, lead, or copper are accumulated and excreted by salt glands or trichomes on the surface of the leaves--a novel phytoremediation process called "phytoexcretion". Finally, the use of halophytes has also been proposed for soil desalination through salt accumulation in the plant tissue or dissolution of soil calcite in the rhizosphere to provide Ca2+ that can be exchanged with Na+ at cation exchange sites.     

The effect of salt on protein synthesis in the halophyte Suaeda maritima

Summary An amino acid-incorporating microsomal fraction has been isolated from the leaves of the halophyte Suaeda maritima and the characteristics of the incorporation described. There were no differences in the properties of the microsomes isolated from plants grown in saline and non-saline conditions. The incorporation was severely inhibited by high concentrations of sodium or potassium ions. The results are discussed in relation to the mechanism of salt tolerance in halophytes and the localization of salt in the cells.     

Amaranthaceae halophytes from the French Flanders coast of the North Sea: a review of their phytochemistry and biological activities

Halophytes are plant species that tolerate high salinity levels. To adapt to these particular abiotic conditions, they develop multiple physiological, biochemical and molecular mechanisms, including the biosynthesis of osmolytes, enzymes and specialized metabolites. The French Flanders coast of the North Sea is an ideal environment for this kind of plant. Amaranthaceae is one of the most represented botanical families of halophytes present on this coast, with 15 species belonging to 7 genera, namely Atriplex, Beta, Halimione, Kali, Oxybasis, Salicornia and Suaeda. Some of these species are well known as wild edible plants, and some are used in traditional medicine. This review examines the chemistry of these species and their potential for human health.

     

Microtuboli in Cellule Parenchimatiche ed Epidermiche di Coleottile di Avena

Abstract

Microtubules in parenchyma and epidermis cells of avena coleoptile. — The fine structure of differentiating parenchyma and epidermis cells of the oat coleoptile, fixed in glutaraldehyde-osmium tetroxide, or glutaraldehyde-potassium permanganate, was investigated. Tubular structures have been observed aligned in the peripheral cytoplasm, between the cell wall and the plasma membrane, embedded in the cell wall and inside the tonoplast in the vacuoles.

The nature and function of these structures are yet unknown. Microtubular structures, localized beneath and above the plasma membrane, have been associated to the wall development; the function of the microtubules observed in vacuoles results, anyhow, of far problematic interpretation.     

Agricultural production of halophytes irrigated with seawater

Summary Growing agricultural crops with direct seawater irrigation has progressed within the past few years from the conceptual to the experimental phase. This has been accomplished by selecting halophytes with inherently high salinity tolerance for use as crop plants rather than by increasing the ability of traditional crop plants to tolerate seawater. Some of the halophytes being investigated for use as crops in seawater irrigation scenarios have high nutritional value as forage or fodder crops. Most of them also have high digestibility. The limiting factor in such use is their high salt content, but this limitation can be moderated. However, since seeds of halophytes do not accumulate salt any more than do those of glycophytes, the greatest promise for seawater-irrigated halophytes probably will be as seed crops. The seeds of many halophytes have high protein and oil contents and compare favorably with traditional oilseed crops. Sustained high yields of seed and biomass already have been obtained from some halophytes irrigated with seawater, and within the next few years seawater agriculture should proceed from the experimental to the operational phase.