Foliar water supply of tall trees: evidence for mucilage-facilitated moisture uptake from the atmosphere and the impact on pressure bomb measurements

The water supply to leaves of 25 to 60 m tall trees (including high-salinity-tolerant ones) was studied. The filling status of the xylem vessels was determined by xylem sap extraction (using jet-discharge, gravity-discharge, and centrifugation) and by (1)H nuclear magnetic resonance imaging of wood pieces. Simultaneously, pressure bomb experiments were performed along the entire trunk of the trees up to a height of 57 m. Clear-cut evidence was found that the balancing pressure (P(b)) values of leafy twigs were dictated by the ambient relative humidity rather than by height. Refilling of xylem vessels of apical leaves (branches) obviously mainly occurred via moisture uptake from the atmosphere. These findings could be traced back to the hydration and rehydration of mucilage layers on the leaf surfaces and/or of epistomatal mucilage plugs. Xylem vessels also contained mucilage. Mucilage formation was apparently enforced by water stress. The observed mucilage-based foliar water uptake and humidity dependency of the P(b) values are at variance with the cohesion-tension theory and with the hypothesis that P(b) measurements yield information about the relationships between xylem pressure gradients and height.     

落葵薯粘液细胞分布及发育的解剖学研究

采用石蜡切片法对落葵薯(Anredera cordifolia(Tenore)Steenis)粘液细胞的分布及发育进行了研究。结果表明,粘液细胞普遍存在于落葵薯的茎、叶、叶柄中,粘液细胞单个散生分布于茎的髓及皮层组织中;叶的栅栏组织和海绵组织中都可见粘液细胞,且海绵组织中的数量明显多于栅栏组织中的;叶柄中的粘液细胞不多,主要分布在维管束四周的皮层组织中。粘液细胞的发育分为4个阶段:原始细胞阶段、液泡化阶段、成熟阶段和细胞质解体阶段。粘液细胞最早出现于第六叶原基,且其发育与茎、叶器官的组织分化不同步。     

落葵粘液细胞分布及发育的解剖学研究

采用石蜡切片法对落葵粘液细胞的分布及发育构造进行观察研究.结果表明:(1)除花药、子房及种子外,粘液细胞普遍存在于落葵植株的地上部分内.茎中的粘液细胞多单个散生分布于皮层、髓及髓射线;叶内的粘液细胞主要分布于海绵组织,栅栏组织中则很少见;叶柄中的粘液细胞主要沿叶柄"U"型皮层的两边分布;发育后期作为果实的花萼片中粘液细胞则散生分布很多.(2)根据发育过程的不同形态,可将粘液细胞的发育分为4个阶段:原始细胞阶段、液泡化阶段、成熟阶段和细胞质解体阶段;粘液细胞最早可见于第三叶原基,并且粘液细胞的发育与植株器官的发育不同步.     

油樟油细胞和粘液细胞的发育解剖学研究

利用薄切片法对油樟茎叶油细胞和粘液细胞发育的研究结果表明,油细胞最早发生于第二叶原基以及茎端皮层和髓的基本分生组织中。未出现油细胞以前,在上述器官的基本分生组织和原分生组织中,难以区分油细胞的原始细胞与周围细胞,当油细胞原始细胞呈现出体积较大,液泡化程度较低,细胞核大而明显的特征才明显可辨,以后经过液泡融合,油细胞成熟和油细胞的细胞质解体阶段而成为一贮油的囊,而且油囊连接在杯形构造上,粘液细胞的早期发育过程与油细胞的相同,而在细胞液泡化的后期,靠近大液泡的细胞质中产生粘液物质。并扩散到大液泡中,粘液物质不断产生,变浓,占据整个细胞腔,细胞质解体后而成为完全成熟的粘液细胞,因此可见,油细胞和粘液细胞是同源的,也可能粘液细胞是由油细胞转化而来的。     

Mucilages and polysaccharides in Ziziphus species (Rhamnaceae): localization, composition and physiological roles during drought-stress.

The drought-tolerant tree species Ziziphus mauritiana Lamk. and Z. rotundifolia Lamk. were shown to have similar high mucilage concentrations (7-10% dry weight) in their leaves, with large numbers of mucilage-containing cells in the upper epidermis and extracellular mucilage-containing cavities in the leaf veins and stem cortex. The main sugar constituents of the water-soluble mucilage extract were rhamnose, glucose and galactose. During drought-stress in two independent studies, foliar mucilage content was unaffected in both species, but glucose and starch contents declined significantly in crude mucilage extracts from droughted leaves. Enzymatic hydrolysis of the mucilage extract using alpha-amylase and amyloglucosidase released glucose, indicating that a mucilage-associated water-soluble glucan, with alpha-1,4- and alpha-1,6-linkages, may exist which was extracted together with the mucilage. From the current data, it is not possible to localize the glucan to determine whether or not it is associated with mucilage-containing cells. Data from pressure-volume analyses of drought-stressed and control leaves showed that, in line with their similar mucilage contents, the relative leaf capacitance isotherm (change in relative water content per unit change in water potential) was similar in both species. During drought-stress, reduced relative capacitance resulted from osmotic adjustment and decreased wall elasticity. Data suggest that in Ziziphus leaves, intracellular mucilages play no part in buffering leaf water status during progressive drought. In Ziziphus species, growing in environments with erratic rainfall, the primary role of foliar mucilage and glucans, rather than as hydraulic capacitors, may be as sources for the remobilization of solutes for osmotic adjustment, thus enabling more effective water uptake and assimilate redistribution into roots and stems prior to defoliation as the drought-stress intensified.     

Structure and bioactivity of the polysaccharides in medicinal plant Dendrobium huoshanense

Detailed structures of the active polysaccharides extracted from the leaf and stem cell walls and mucilage of Dendrobium huoshanense are determined by using various techniques, including chromatographic, spectroscopic, chemical, and enzymatic methods. The mucilage polysaccharide exhibits specific functions in activating murine splenocytes to produce several cytokines including IFN-gamma, IL-10, IL-6, and IL-1alpha, as well as hematopoietic growth factors GM-CSF and G-CSF. However, the deacetylated mucilage obtained from an alkaline treatment fails to induce cytokine production. The structure and bioactivity of mucilage components are validated by further fractionation. This is the first study that provides clear evidence for the structure and activity relationship of the polysaccharide in D. huoshanense.     

Leaf Structural Peculiarities in Sarcopoterium spinosum, a Seasonally Dimorphic Subshrub

Extensive investigations on the anatomy of the two leaf typesin a seasonally dimorphic subshrub revealed interesting variationsbetween summer and winter leaves. Summer leaves of Sarcopoteriumspinosum possess a thick epidermis composed of tannin-containingcells and large amounts of mucilage secreted through the innerpericlinal walls towards the mesophyll. A thick cuticle is alsopresent on the surface of the leaf. In winter leaves the epidermalcells produce no mucilage while phenolics are accumulated ingranular form only. Besides these, some other variations betweensummer and winter leaves are also discussed in respect of theability of the plant to withstand the unfavourable Mediterraneanconditions. Seasonal dimorphism, leaf anatomy, Sarcopoterium spinosum     

Development of mucilage cells of Araucaria angustifolia (Araucariaceae)

The roles of mucilage cells were investigated through morphological and cytological analysis during leaf development in young Araucaria angustifolia plants. Differentiation began in leaf primordia in the shoot apex, when the young cells underwent a greater increase in volume in comparison with other mesophyll cells. The mucilage polysaccharides were synthesized by dictyosomes, from where they were taken by large vesicles and released into a cavity formed by detachment of the tonoplast, which was separated from the cytoplasm. At the end of differentiation, the cell was completely filled with mucilage, a gel consisting of a denser reticular structure surrounding less dense regions. The nucleus and cytoplasm were degenerated in mature cells. The A. angustifolia mucilage cells presented some cytological resemblances to the mucilage cells of members of some dicotyledonous families; however, differences in the dictyosomes and the secretion route were observed. Translocation and water storage of solutes was suggested by the use of the hydroxy pyrenetrisulfonic acid tri-sodium salt apoplastic tracer. The tonoplast detachment, dechromatinization, nuclear condensation, and general degeneration of the membrane systems observed during maturity indicated a programmed cell death process, one not yet described for angiosperm mucilage cells.     

Presumed role of mucilage of plantain seeds in spread of tobacco mosaic virus

A strain of tobaoco mosaic virus(TMVbs) isolated fromPlantago major L. is not seed-borne with this plant species; however, plantain seeds and the mucilage on their surface also contain the virus. The mucilage shows a very low infectivity; the visoous mucilage inhibits infeotion in rubbed leaves both in a mixture with the virus and if applied before (but not after) the inoculation of the virus. Polysaccharides and hexuronic acids were detected in the great plantain seed mucilage. A mixture of mucilage and viras stored at room temperature showed s low infectivity until 33 days, but the infectivity increased oonspicuously on the 40th day, apparently simultaneously with the decay of mucilage. Polysaccharides are suggested as being responsible for the inhibition. They affect the inoculated leaf but not the virus, as ascertained by eleotron mioroscopy and serology. The seed mucilage being an important factor in the spread of plantain seeds may act as a vehicle of the virus as well.     

ULTRASTRUCTURE OF CARPOSPOROPHYTE DEVELOPMENT IN THE RED ALGA GLOIOSIPHONIA VERTICILLARIS (CRYPTONEMIALES,GLOIOSIPHONIACEAE)

The ultrastructure of carposporophyte development is described for the red alga Gloiosiphonia verticillaris Farl. The auxiliary cell produces gonimoblast initials, which divide to produce two types of gonimoblast cells—the nondividing vacuolate cells and terminal generative gonimoblast cells. The generative gonimoblast cells form clusters of carpospore initials, which eventually differentiate into carpospores. After gonimoblast filaments are formed, the auxiliary cell undergoes autolysis, causing degeneration of septal plugs between the auxiliary cell and adjacent cells, thus forming a fusion cell. Since this cell lacks starch and appears degenerate throughout carposporophyte development, a nutritive function cannot be ascribed to the fusion cell. Carpospore differentiation is simple and proceeds through three developmental stages. Young carpospores structurally resemble gonimoblast cells, because they contain undeveloped plastids, large quantities of floridean starch, and are surrounded by extensive mucilage instead of a distinct wall. In addition, dictyosomes form and begin to produce vesicles with fibrous contents representing carpospore wall material. During the intermediate stage, dictyosomes continue to produce vesicles that contribute additional carpospore wall material, thereby compressing the mucilage and creating a darker-staining layer outside the carpospore wall. Plastids form internal thylakoids by invaginations of the inner membrane of the peripheral thylakoid. The endoplasmic reticulum forms large granular vacuoles that appear to be degraded during subsequent stages of development. Mature carpospores form cored vesicles. They also contain mature chloroplasts, large amounts of floridean starch, and occasionally granular vacuoles. During this stage, interconnecting carpospore-carpospore and carpospore-gonimoblast cell septal plugs begin to undergo degeneration. This process may be mediated by tubular structures.     

ULTRASTRUCTURE OF AUXILIARY AND GONIMOBLAST CELLS DURING CARPOSPOROPHYTE DEVELOPMENT IN FAUCHEOCOLAX ATTENUATA (RHODOPHYTA)1

The ultra structure of post-fertilization development in Faucheocolax attenuata Setch. is described. Following fertilization and transfer of the diploid nucleus to the auxiliary cell, four gonimoblast initials usually are produced of the multinucleate auxiliary cell. Gonimoblast initials originally are uninucleate but undergo karyokinesis to form multinudeate gonimoblast cells. Terminal or generative gonimoblast cells cleave successively to form lobes of incipient carpospores, with each group of spores differentiating synchronously. Portions of the initial generative gonimoblast cells, however, remain to resume karyokinesis and repeat the process of cleavage into carpospores. Axial gonimoblast cells are transformed into secretory cells, which produce mucilage. Generative gonimoblast cells and auxiliary cells are similar in cellular structure. Both contain typical red algal proplastids, some dictyosomes, cytoplasmic concentric membranes, and numerous small vesicles. In addition, dark staining spherical masses, occurring in the cytoplasm of all cell types, may represent dehydrated haploid chromatin. Large septal plugs interconnect gonimoblast cells and the auxiliary cell. These plugs are small when first formed but increase dramatically in size during carposporophyte development.     

Ultrastructure of post-fertilization development in the red alga Nienburgia andersoniana (Delesseriaceae, Ceramiales, Rhodophyta)

The ultrastructure of post-fertilization development in Nienburgia andersoniana (J. Ag.) Kyl. is described. Above the auxiliary cell there is a group of four sterile cells. The presence of abundant storage products (starch granules, lipid bodies and protein crystals) in these cells indicates that the sterile cells function as nutrient suppliers to the young auxiliary and gonimoblast cells of the carposporophyte during its early steps of development. Following fertilization and transfer of the diploid nucleus to the auxiliary cell, the trichogyne disappears and large multinucleate gonimoblast initials are produced. These subsequently produce generative gonimoblast cells which cleave successively to form young carpospores. Those of the gonimoblast cells which will not differentiate into carpospores are transformed into cells producing mucilage. Both kinds of gonimoblast cells contain plastids, starch granules, cytoplasmic concentric membrane bodies and small vesicles. Dark-staining spherical masses occurring in the cytoplasm of the auxiliary and gonimoblast cells may represent degenerating haploid nuclei. Septal plugs interconnecting the auxiliary cell and gonimoblast cells increase considerably in size during carposporophyte development. The fusion cell at the late stage of carposporophyte development appears degenerative. Young carpospores have plastids and mitochondria, and concentric membrane bodies that will form mucilage sacs. Medium-aged carpospores have fully developed plastids, starch granules and fibrous vacuoles. Mature carpospores possess, in addition, cored vesicles. The inner pericarp cells contribute large amounts of mucilage to the cytostocarpic cavity and eventually are consumed. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 289–299.     

Epidermal Micromorphology and Mesophyll Structure of Populus euphratica Heteromorphic Leaves at Different Development Stages

Leaf epidermal micromorphology and mesophyll structure during the development of Populus euphratica heteromorphic leaves, including linear, lanceolate, ovate, dentate ovate, dentate rhombic, dentate broad-ovate and dentate fan-shaped leaves, were studied by using electron and light microscopy. During development of heteromorphic leaves, epidermal appendages (wax crystals and trichomes) and special cells (mucilage cells and crystal idioblasts) increased in all leaf types while chloroplast ultrastructure and stomatal characters show maximum photosynthetic activity in dentate ovate and rhombic leaves. Also, functional analysis by subordinate function values shows that the maximum adaptability to adverse stress was exhibited in the broad type of mature leaves. The 12 heteromorphic leaf types are classified into three major groups by hierarchical cluster analysis: young, developing and mature leaves. Mature leaves can effectively obtain the highest stress resistance by combining the protection of xerophytic anatomy from drought stress, regulation of water uptake in micro-environment by mucilage and crystal idioblasts, and assistant defense of transpiration reduction through leaf epidermal appendages, which improves photosynthetic activity under arid desert conditions. Our data confirms that the main leaf function is differentiated during the developing process of heteromorphic leaves.     

Studies on the Development of Glandular Hairs in Nymphoides peltatum and the Ultrastructure During Their Secretory P

Each glandular hair of Nyrnphoides peltaturn (Gmel.) O. Kuntz consisted of only one row of cylindar cells with secretory function. The hairs originated from the protoderm cells on the adaxial surface of the second leaf primordium from the shoot apex. Cells of the glandular hairs prossessed dense cytoplast during the secretory period, but the vacuoles were very small. There were not only abundant mitochondria, Golgi bodies and endoplasmic reticulum in the glandular hair cells, but also many plasmodesmata. The authors' research indicated that the mucilage was carried to the edge of the cells by the membranous multilamellar bodies and the vesicles from both Golgi bodies and endoplasmic reticulum. The mucilage was secreted extracellularly by either exocytosis or ecrine secretion. The side walls of the glandular hairs swelled because of mucilage mass accumulation in the walls. The mucilage, being tested to be composed of polysaccharides and a trace of protein, played an important role in protecting the development of the vegetative buds of N. peltatum.     

Cytochemical and Morphological Evidence for the Involvement of the Plasma Membrane and Plastids in Mucilage Secretion in Aloe arborescens

Most of the volume of Aloe arborescens leaves (volumetric density= 0.68±0.08) is occupied by a mucilage tissue. The mucilagesof Aloe species are glucomannans of great medical and pharmaceuticalimportance. An electron microscopical and histochemical studywas carried out, following the development of the tissue andsecretion processes. In contrast to other reports on polysaccharide secretion inplants, no observable activity was found in the Golgi apparatus.In the young stages of leaf development, positive histochemicalstaining for polysaccharides and structural changes were foundmainly in the plastids. In the mature leaves the mucilage cellsremain alive and intact and secretory activity appears to becentered in the plasma membrane. Aloe arborescens, cytochemistry, mucilage, plasma membrane, plastids, secretion     

Two new petrified cycad stems,Brunoa gen. nov. andWorsdellia gen. nov., from the Cretaceous of Patagonia (Bajo de Santa Rosa, Río Negro Province), Argentina

Polyxylic columnar stems covered by persistent leaf bases and found in sediments assignable to the Upper Cretaceous of Bajo de Santa Rosa, Río Negro Province, Argentina, are described as two new generic entities in the Cycadales. Anatomical characters are the basis for their being assigned to the Encephalartoideae of the Zamiaceae.Brunoa santarrosensis gen. et sp. nov. is characterized by the presence of polyxyly, cone domes, mucilage cavities, and uniseriate to triseriate araucaroid, scalariform, or bordered intervascular pitting.Worsdellia bonettiae gen. et sp. nov. has polyxyly, anastomosing medullary vascular bundles, centripetal xylem, mucilage canals, and concentric extraxylary bundles. Some characters (polyxyly, medullary vascular bundles, and cone domes) were used to determine the systematic position, while other characters (mucilage reservoirs and centripetal xylem) were used to establish the relationship between polyxylic and monoxylic forms.     

Structure and function of trichornes in the shoot tip of Ardisia crispa (Thunb.) A.DC. (Myrsinaceae)

MILLER, I. M., GARDNER, I. C. & SCOTT, A., 1984. Structure and function of trichomes in the shoot tip of Ardisia crispa (Thunb.) A.DC. (Myrsinaceae). The trichomes in the shoot tip of the myrsinaceous leaf-nodulated species Ardisia crispa (Thunb.) A.DC. have been studied at the ultrastructural level. Two distinct kinds of trichomes are found. Small peltate scales arise from the abaxial protoderrn of the developing leaves. Multicellular stellate trichomes on the adaxial surface of young leaves are uniseriate, rotate and consist of up to eight multicellular arms radiating from a central stalk. The multicellular arms terminate distally in swollen club-shaped tips. The stellate trichomes secrete large quantities of carbohydrate-containing mucilage. Colonies of leaf nodule bacteria are found resident in this mucilage. The role of the peltate scales and the stellate trichomes n the maintenance of the cyclic leaf nodule symbiosis is discussed.     

我国两种特有苏铁的生态环境及叶片结构

考查广西百色西林县叉孢苏铁和广西百色田东县锈毛苏铁的生态环境,结合观测其叶片的内部结构来探讨其与生境的适应性。结果表明叉孢苏铁群落乔木和灌木均9种,藤本8种,草本18种,蕨类2种,空间格局明显;锈毛苏铁群落乔木和灌木均5种,草本6种,蕨类4种,空间格局不明显。两种苏铁的羽片均由表皮、栅栏组织、海绵组织和副转输组织构成;叶轴均由表皮、厚壁组织、薄壁组织和维管组织构成。不同之处在于,锈毛苏铁的羽片、角质层、栅栏组织和叶轴的厚壁组织均较叉孢苏铁的厚,分别是叉孢苏铁的1．82倍、2．32倍、1．79倍和2．45倍左右;锈毛苏铁叶片的表皮之下有下皮层、各组织中有丰富的异细胞,而又孢苏铁的无或较少;锈毛苏铁的羽片中有黏液道,而叉孢苏铁的无。又孢苏铁的叶片表现为弱旱生异面叶,锈毛苏铁的则表现为强旱生异面叶。     

A mathematical model of mucilage expansion in myxospermous seeds of Capsella bursa-pastoris (shepherd's purse)

Background and Aims

Myxospermy is a term which describes the ability of a seed to produce mucilage upon hydration. The mucilage is mainly comprised of plant cell-wall polysaccharides which are deposited during development of those cells that comprise the seed coat (testa). Myxospermy is more prevalent among those plant species adapted to surviving on arid sandy soils, though its significance in determining the ecological fitness of plants is unclear. In this study, the first mathematical model of myxospermous seed mucilage expansion is presented based on seeds of the model plant species Capsella bursa-pastoris (shepherd''s purse).

Methods

The structures underpinning the expansion process were described using light, electron and time-lapse confocal micrographs. The data and experimental observations were used to create a mathematical model of myxospermous seed mucilage expansion based on diffusion equations.

Key Results

The mucilage expansion was rapid, taking 5 s, during which the cell mucilage volume increased 75-fold. At the level of the seed, this represented a 6-fold increase in seed volume and a 2·5-fold increase in seed surface area. These increases were shown to be a function of water uptake (16 g water g⁻¹ mucilage dry weight), and relaxation of the polymers which comprised the mucilage. In addition, the osmotic pressure of the seed mucilage, estimated by assessing the mucilage expansion of seeds hydrated in solutions of varying osmotic pressure, was –0·54 MPa (equivalent to 0·11 m or 6·6 g L⁻¹ NaCl).

Conclusions

The results showed that the mucilage may be characterized as hydrogel and seed-mucilage expansion may be modelled using the diffusion equation described. The potential of myxospermous seeds to affect the ecological services provided by soil is discussed briefly.     

Studies of protonemal morphogenesis in mosses. XII. Ephemeropsis,the zenith of morphological differentiation

Abstract

This light and scanning electron microscope study of freshly collected material confirms that the protonemal system of Ephemeropis is the most highly differentiated so far found in mosses and reveals hitherto unsuspected roles for its unique features. Whereas substrate attachment in the Hypnales, illustrated here in Homalothecium sericeum, is by clusters of irregular terminal mucilage-secreting ramifications on long otherwise unbranched rhizoids and in other mosses is via otherwise unmodified rhizoids, the anchoring structures or hapteres in Ephemeropsis are very regular side branch systems that also secrete copious mucilage. This last feature, only clearly visible in living specimens, also invests the bristle-like appendages similarly unique to Ephemeropsis. Abundant cyanobacteria in this mucilage may be an important nitrogen source for Ephemeropsis. Other features most likely linked to growth on twigs, the leaf bases of ferns and tree ferns and epiphylly are the absence of food-conducting cytology from the main protonemal axes and the rapid absorption of water by the much-branched chloronemata. Gemmae are produced at the tips of attenuated filaments that grow downwards in a spiral pattern and are rapidly anchored to the subjacent leaves by means of short side branches at their bases that also secrete copious mucilage.