Cavitation Resistance in Seedless Vascular Plants: The Structure and Function of Interconduit Pit Membranes

Plant water transport occurs through interconnected xylem conduits that are separated by partially digested regions in the cell wall known as pit membranes. These structures have a dual function. Their porous construction facilitates water movement between conduits while limiting the spread of air that may enter the conduits and render them dysfunctional during a drought. Pit membranes have been well studied in woody plants, but very little is known about their function in more ancient lineages such as seedless vascular plants. Here, we examine the relationships between conduit air seeding, pit hydraulic resistance, and pit anatomy in 10 species of ferns (pteridophytes) and two lycophytes. Air seeding pressures ranged from 0.8 ± 0.15 MPa (mean ± sd) in the hydric fern Athyrium filix-femina to 4.9 ± 0.94 MPa in Psilotum nudum, an epiphytic species. Notably, a positive correlation was found between conduit pit area and vulnerability to air seeding, suggesting that the rare-pit hypothesis explains air seeding in early-diverging lineages much as it does in many angiosperms. Pit area resistance was variable but averaged 54.6 MPa s m⁻¹ across all surveyed pteridophytes. End walls contributed 52% to the overall transport resistance, similar to the 56% in angiosperm vessels and 64% in conifer tracheids. Taken together, our data imply that, irrespective of phylogenetic placement, selection acted on transport efficiency in seedless vascular plants and woody plants in equal measure by compensating for shorter conduits in tracheid-bearing plants with more permeable pit membranes.Water transport in plants occurs under tension, which renders the xylem susceptible to air entry. This air seeding may lead to the rupture of water columns (cavitation) such that the air expands within conduits to create air-vapor embolisms that block further transport. (Zimmermann and Tyree, 2002). Excessive embolism such as that which occurs during a drought may jeopardize leaf hydration and lead to stomatal closure, overheating, wilting, and possibly death of the plant (Hubbard et al., 2001; Choat et al., 2012; Schymanski et al., 2013). Consequently, strong selection pressure resulted in compartmentalized and redundant plant vascular networks that are adapted to a species habitat water availability by way of life history strategy (i.e. phenology) or resistance to air seeding (Tyree et al., 1994; Mencuccini et al., 2010; Brodersen et al., 2012). The spread of drought-induced embolism is limited primarily by pit membranes, which are permeable, mesh-like regions in the primary cell wall that connect two adjacent conduits. The construction of the pit membrane is such that water easily moves across the membrane between conduits, but because of the small membrane pore size and the presence of a surface coating on the membrane (Pesacreta et al., 2005; Lee et al., 2012), the spread of air and gas bubbles is restricted up to a certain pressure threshold known as the air-seeding pressure (ASP). When xylem sap tension exceeds the air-seeding threshold, air can be aspirated from an air-filled conduit into a functional water-filled conduit through perhaps a large, preexisting pore or one that is created by tension-induced membrane stress (Rockwell et al., 2014). Air seeding leads to cavitation and embolism formation, with emboli potentially propagating throughout the xylem network (Tyree and Sperry, 1988; Brodersen et al., 2013). So, on the one hand, pit membranes are critical to controlling the spread of air throughout the vascular network, while on the other hand, they must facilitate the efficient flow of water between conduits (Choat et al., 2008; Domec et al., 2008; Pittermann et al., 2010; Schulte, 2012). Much is known about such hydraulic tradeoffs in the pit membranes of woody plants, but comparatively little data exist on seedless vascular plants such as ferns and lycophytes. Given that seedless vascular plants may bridge the evolutionary transition from bryophytes to woody plants, the lack of functional data on pit membrane structure in early-derived tracheophytes is a major gap in our understanding of the evolution of plant water transport.In woody plants, pit membranes fall into one of two categories: the torus-margo type found in most gymnosperms and the homogenous pit membrane characteristic of angiosperms (Choat et al., 2008; Choat and Pittermann, 2009). In conifers, water moves from one tracheid to another through the margo region of the membrane, with the torus sealing the pit aperture should one conduit become embolized. Air seeding occurs when water potential in the functional conduit drops low enough to dislodge the torus from its sealing position, letting air pass through the pit aperture into the water-filled tracheid (Domec et al., 2006; Delzon et al., 2010; Pittermann et al., 2010; Schulte, 2012; but see Jansen et al., 2012). Across north-temperate conifer species, larger pit apertures correlate with lower pit resistance to water flow (r_pit; MPa s m⁻¹), but it is the ratio of torus-aperture overlap that sets a species cavitation resistance (Pittermann et al., 2006, 2010; Domec et al., 2008; Hacke and Jansen, 2009). A similar though mechanistically different tradeoff exists in angiosperm pit membranes. Here, air seeding reflects a probabilistic relationship between membrane porosity and the total area of pit membranes present in the vessel walls. Specifically, the likelihood of air aspirating into a functional conduit is determined by the combination of xylem water potential and the diameter of the largest pore and/or the weakest zone in the cellulose matrix in the vessel’s array of pit membranes (Wheeler et al., 2005; Hacke et al., 2006; Christman et al., 2009; Rockwell et al., 2014). As it has come to be known, the rare-pit hypothesis suggests that the infrequent, large-diameter leaky pore giving rise to that rare pit reflects some combination of pit membrane traits such as variation in conduit membrane area (large or small), membrane properties (tight or porous), and hydrogel membrane chemistry (Hargrave et al., 1994; Choat et al., 2003; Wheeler et al., 2005; Hacke et al., 2006; Christman et al., 2009; Lee et al., 2012; Plavcová et al., 2013; Rockwell et al., 2014). The maximum pore size is critical because, per the Young-Laplace law, the larger the radius of curvature, the lower the air-water pressure difference under which the contained meniscus will fail (Jarbeau et al., 1995; Choat et al., 2003; Jansen et al., 2009). Consequently, angiosperms adapted to drier habitats may exhibit thicker, denser, smaller, and less abundant pit membranes than plants occupying regions with higher water availability (Wheeler et al., 2005; Hacke et al., 2007; Jansen et al., 2009; Lens et al., 2011; Scholz et al., 2013). However, despite these qualitative observations, there is no evidence that increased cavitation resistance arrives at the cost of higher r_pit. Indeed, the bulk of the data suggest that prevailing pit membrane porosity is decoupled from the presence of the single largest pore that allows air seeding to occur (Choat et al., 2003; Wheeler et al., 2005 Hacke et al., 2006, 2007).As water moves from one conduit to another, pit membranes offer considerable hydraulic resistance throughout the xylem network. On average, r_pit contributes 64% and 56% to transport resistance in conifers and angiosperms, respectively (Wheeler et al., 2005; Pittermann et al., 2006; Sperry et al., 2006). In conifers, the average r_pit is estimated at 6 ± 1 MPa s m⁻¹, almost 60 times lower than the 336 ± 81 MPa s m⁻¹ computed for angiosperms (Wheeler et al., 2005; Hacke et al., 2006; Sperry et al., 2006). Presumably, the high porosity of conifer pits compensates for the higher transport resistance offered by a vascular system composed of narrow, short, single-celled conduits (Pittermann et al., 2005; Sperry et al., 2006).Transport in seedless vascular plants presents an interesting conundrum because, with the exception of a handful of species, their primary xylem is composed of tracheids, the walls of which are occupied by homogenous pit membranes (Gibson et al., 1985; Carlquist and Schneider, 2001, 2007; but see Morrow and Dute, 1998, for torus-margo membranes in Botrychium spp.). At first pass, this combination of traits appears hydraulically maladaptive, but several studies have shown that ferns can exhibit transport capacities that are on par with more recently evolved plants (Wheeler et al., 2005; Watkins et al., 2010; Pittermann et al., 2011, 2013; Brodersen et al., 2012). Certainly, several taxa possess large-diameter, highly overlapping conduits, some even have vessels such as Pteridium aquilinum and many species have high conduit density, all of which could contribute to increased hydraulic efficiency (Wheeler et al., 2005; Pittermann et al., 2011, 2013). But how do the pit membranes of seedless vascular plants compare? Scanning electron micrographs of fern and lycopod xylem conduits suggest that they are thin, diaphanous, and susceptible to damage during specimen preparation (Carlquist and Schneider 2001, 2007). Consistent with such observations, two estimates of r_pit imply that r_pit in ferns may be significantly lower than in angiosperms; Wheeler et al. (2005) calculated r_pit in the fern Pteridium aquilinum at 31 MPa s m⁻¹, while Schulte et al. (1987) estimated r_pit at 1.99 MPa s m⁻¹ in the basal fern Psilotum nudum. The closest structural analogy to seedless vascular plant tracheids can be found in the secondary xylem of the early-derived vesselless angiosperms, in which tracheids possess homogenous pit membranes with r_pit values that at 16 MPa s m⁻¹ are marginally higher than those of conifers (Hacke et al., 2007). Given that xylem in seedless vascular plants is functionally similar to that in vesselless angiosperms, we expected convergent r_pit values in these two groups despite their phylogenetic distance. We tested this hypothesis, as well as the intrinsic cavitation resistance of conduits in seedless vascular plants, by scrutinizing the pit membranes of ferns and fern allies using the anatomical and experimental approaches applied previously to woody taxa. In particular, we focused on the relationship between pit membrane traits and cavitation resistance at the level of the individual conduit.     

Antitranspirant Activity in Xylem Sap of Maize Plants

Xylem sap from unwatered maize plants was collected and testedfor antitranspirant activity. Two assays were used. These werea transpiration assay with detached wheat leaves and a stomatalbio-assay involving the direct microscopic observation of epidermisof Commelina communis. The reduction in transpiration of detached wheat leaves promotedby xylem sap could be duplicated almost exactly by the applicationof solutions of ABA of equivalent concentration to that foundin the xylem sap. Removal of virtually all the ABA from thexylem sap, using an immunoaffinity column, removed virtuallyall the antitranspirant activity in both assays. These results are discussed in the context of other resultswhich suggest the presence of as-yet unidentified inhibitorsin the xylem sap of unwatered plants. We suggest that with maize plants at least, stomatal responsesto soil drying can be entirely explained by enhanced concentrationof ABA in the xylem stream. Key words: Antitranspirant activity, ABA, ABA bio-assay, xylem sap     

Gap Junctions: Structure, Functions, and Regulation

The review presents data on the role of gap junctions in intercellular communication. The review includes information on history of the appearance of this problem. Data are presented on ultrastructure and function of gap junctions as well as on the mechanisms providing for their activity. A part of the review deals with the problem of regulation of intercellular communication realized by the gap junctions.     

Effect of Macerase, Oxalic Acid, and EGTA on Deep Supercooling and Pit Membrane Structure of Xylem Parenchyma of Peach

The object of this study was to determine if calcium cross-linking of pectin in the pit membrane of xylem parenchyma restricts water movement which results in deep supercooling. Current year shoots of `Loring' peach (Prunus persica) were infiltrated with oxalic acid or EGTA solutions for 24 or 48 hours and then either prepared for ultrastructural analysis or subjected to differential thermal analysis. The effect of 0.25 to 1.0% pectinase (weight/volume) on deep supercooling was also investigated. The use of 5 to 50 millimolar oxalic acid and pectinase resulted in a significant reduction (flattening) of the low temperature exotherm and a distinct swelling and partial degradation of the pit membrane. EGTA (10 millimolar) for 24 or 48 hours shifted the low temperature exotherm to warmer temperatures and effected the outermost layer of the pit membrane. A hypothesis is presented on pectin-mediated regulation of deep supercooling of xylem parenchyma.     

Pit Membrane Degradation and Air-Embolism Formation in Ageing Xylem Vessels of Populus tremuloides Michx

Air-embolism formation in xylem vessels of Populus tremuloidesMichx. was quantified by its reduction of hydraulic conductivityin branch segments. Embolism was induced by increasing xylemtension in drying stems, or by inserting one end of a hydratedstem in a pressure bomb and increasing air pressure in the bomb.Both treatments produced the same response suggesting that embolismby water stress was caused by air entering water-filled vessels,presumably through inter-vessel pits. In rapidly-growing P.tremuloides branches, the vessels of the outer growth ring werefunctional whereas vessels in older xylem were mostly embolized.This selective embolizing of older vessels was associated witha marked increase in permeability of their inter-vessel pitsto air, relative to pits of younger vessels. Air-injection pressuresless than 1·0 MPa completely embolized older vesselsthat had been re-filled in the laboratory, whereas pressuresover 4·0 MPa were required to embolize young vessels.Greater permeability of old vessels was due to degradation oftheir pit membranes as seen in the scanning electron microscope;large openings were present that were not seen in pit membranesof young vessels. These holes would allow air to penetrate vesselends at low pressure differences causing embolism. Degradationof pit membranes causing the selective dysfunction of oldersapwood may be a general phenomenon initiating heartwood formationin many species. Key words: Xylem embolism, hydraulic conductivity, heartwood formation, cavitation, Populus tremuloides, Michx     

The Relationships between Xylem Safety and Hydraulic Efficiency in the Cupressaceae: The Evolution of Pit Membrane Form and Function

Water transport in conifers occurs through single-celled tracheids that are connected to one another via intertracheid pit membranes. These membranes have two components: the porous margo, which allows water to pass through the membrane, and the impermeable torus, which functions to isolate gas-filled tracheids. During drought, tracheids can become air filled and thus hydraulically dysfunctional, a result of air entering through the pit membrane and nucleating cavitation in the water column. What are the hydraulic tradeoffs associated with cavitation resistance at the pit level, and how do they vary within the structural components of the intertracheid pit? To address these questions, we examined pit structure in 15 species of Cupressaceae exhibiting a broad range of cavitation resistances. Across species, cavitation resistance was most closely correlated to the ratio of the torus to pit aperture diameter but did not vary systematically with margo porosity. Furthermore, our data indicate that constraints on pit hydraulic efficiency are shared: the pit aperture limits pit conductivity in more drought-resistant taxa, while increased margo resistance is more likely to control pit conductivity in species that are more vulnerable to cavitation. These results are coupled with additional data concerning pit membrane structure and function and are discussed in the context of the evolutionary biogeography of the Cupressaceae.Water transport in conifers occurs through narrow, single-celled conduits (tracheids) that are organized in overlapping, longitudinal files. This simple, homoxylous arrangement represents an ancestral vascular design that has remained remarkably consistent since its first appearance in the progymnosperms of the Mid-Devonian (Taylor et al., 2009). However, the small size of tracheids can impose a high resistance to water transport as compared with the large, hydraulically efficient vessels present in many angiosperms (Hacke et al., 2004; Sperry et al., 2006). Despite this handicap, conifer tracheids can be just as hydraulically efficient as angiosperm xylem for a given conduit diameter, a result that can be wholly attributed to the distinctive structure of the conifer intertracheid pit membrane (Pittermann et al., 2005; Sperry et al., 2006).Because pit membranes also function to limit the spread of air from one conduit to another (cavitation), the physiological consequences of the transport efficiency versus cavitation safety tradeoffs in conifer and angiosperm pit membranes have received considerable attention at the pit and xylem levels, whereby cavitation resistance in north temperate woody plants appears to come at the cost of hydraulic efficiency (Pittermann et al., 2006a, 2006b; Sperry et al., 2006; Choat et al., 2008; Domec et al., 2008; Jansen et al., 2009; Schoonmaker et al., 2010). Previous work has shown that the integrated vascular performance of plants is key to understanding species distributions (Sperry et al., 1994; Brodribb and Hill, 1999; Pockman and Sperry, 2000; Choat et al., 2007), and within this framework, pit membranes have the potential to act as the nexus of the cavitation safety versus transport efficiency compromise. Yet, despite our progress, we are just starting to learn how these tradeoffs play out at the level of the pit membrane, particularly in one as complex as that of conifers. Hence, the goals of this study were to determine whether selection has acted to optimize conifer pit membrane performance in a manner that reflects species cavitation resistance and habitat distribution as well as to examine the role, if any, of evolutionary lineage.Unlike the homogenous pit membrane of angiosperm vessels, the conifer pit membrane is composed of two distinct regions: a thickened, centrally located torus and a porous margo region that surrounds it (Fig. 1; for study species, see Hacke et al., 2004; Choat et al., 2008; Choat and Pittermann, 2009). When tracheids are water filled, the pit membrane is centrally located in the pit chamber and water moves from tracheid to tracheid through the margo. Should an air-seeding event (cavitation) occur, causing a tracheid to become air filled, (i.e. embolized), the negative xylem pressure in the water-filled tracheid will act on the air-water interface in the margo pores by deflecting the pit membrane in the direction of the functional tracheid, thereby appressing the torus against the pit aperture border (Bailey, 1913; Liese, 1965; Liese and Bauch, 1967; Petty, 1972). This valve action of the membrane can create an effective seal that prevents further spread of air in the xylem. Cavitation is thought to occur when the water potential of the water-filled tracheid becomes negative enough to dislodge the torus from its sealing position, allowing air to enter the conduit. Overall, the structure of the torus-margo pit membrane must optimize what at first glance appear to be conflicting functional requirements: on the one hand, cavitation resistance selects for a combination of large tori and small apertures, but on the other hand, hydraulic efficiency favors porous margos, large apertures, and small tori.Open in a separate windowFigure 1.SEM images of intertracheid pit membranes belonging to nine Cupressaceae species (of 15) that represent the broad range of observed cavitation pressures. The opaque torus region of the membrane (T) is held centrally by the microfibrils of the margo (M). Visually, increased cavitation resistance appears to be associated with increased margo porosity, but quantitative estimates of margo resistance made on the most intact regions of the pit membranes (Fig. 8) revealed no differences among the species surveyed.

Table I.

Study species, figure abbreviations (Fig. Abbrevs.), locations (SFBG, San Francisco Botanical Garden, San Francisco; UCBG, University of California Botanical Garden, Berkeley, CA; UCSC, University of California, Santa Cruz, Arboretum, Santa Cruz, CA), and species natural history (Farjon, 2005)

Electrochemical Relations in the Transfer of Ions to the Xylem Sap of Maize Roots

The difference in electrical potential between the xylem exudateof isolated roots of 1-week-old maize seedlings and the culturesolution surrounding the roots has been determined togetherwith the concentrations of potassium, calcium, chloride, andsulphate in the xylem sap. The difference in electro-chemicalpotential (µ) for these ions has been calculated fromthe measurements. The effects on µ of varying the saltstatus of the roots, the composition of the culture solutionand of 2-4-dinitrophenol have been examined. µ for potassiumand chloride was always positive, implying that movement ofthese ions to the xylem sap was under metabolic constraint.However, pretreatment of the maize seedlings with potassiumchloride and increasing levels of dinitrophenol in the culturesolution over the exudation period caused little or no significantchange in µ for potassium although the rate of movementof ions to the xylem was substantially reduced. For calcium,µ was negative with roots in dilute culture solutionsin the absence of dinitrophenol, implying that calcium couldenter the xylem by passive diffusion. Addition of dinitrophenolchanged the sign of µ and thus brought about an apparentlyactive transport of calcium.Only in the absence of competinganions was the rate of entry of chloride significantly correlatedwith µ, and no such relationship was found for potassiumor calcium individually. These results encourage doubt as towhether measurements of µ provide a valid basis for decidingto what extent the movement of individual ions depends on specific‘active’ or ‘passive’ transport processes.Thedifference in electrical potential appears to be a characteristicof the living root and to depend on the total concentrationof ions in the external solution and on their rate of transferinto the xylem sap.     

几种生理因素对玉米木质部汁液中蛋白质含量的影响

受干旱胁迫的玉米叶和茎木质部汁液中蛋白质含量降低,根中蛋白质含量升高.偏酸性营养液中的玉米各营养器官木质部汁液中蛋白质含量降低,中性或碱性营养液中的则升高.以100mmol·L-1的ABA营养液处理后的玉米根、茎和叶片的木质部汁液中蛋白质含量都升高;而用2 mmol·L-1EGTA、80 mmol·L-1三氟啦嗪或100mmol·L-1异博定处理后的木质部汁液中蛋白质含量变化不明显.     

The Vascular System of Maize Stems Revisited: Implications for Water Transport and Xylem Safety

The plexus of vascular bundles in the nodes of grasses is notoriouslycomplex, where long axial bundles pass through a network oftransverse bundles. The xylem pathways for water in maize stemshave been investigated anatomically and with dye and particulatetracers, revealing some of the details of this complexity. Onlyapprox. 3% of axial vessels pass through nodes without beinginterrupted by end walls. Axial bundles at nodes differ fromthose in internodes in having the metaxylem and protoxylem vesselsconnected by small tracheary elements. So it is only at nodesthat exchange of sap occurs between the large vessels withina bundle. End walls, acting as filters for particles and gasbubbles, always separate axial vessels from vessels in transversebundles. The high redundancy of bundle connections in the nodalplexus is interpreted as providing alternative water pathwaysto bypass embolisms and damaged or diseased sections of thexylem. The pores in the filters at the base of nodes and betweenaxial and transverse vessels within nodes are <20 nm in diameter.Where axial vessels connect to transverse vessels, a varietyof unusual shapes of vessel elements mediate two- and three-wayconnections within the plexus.Copyright 2000 Annals of BotanyCompany Zea mays, cryoSEM, maize, node, pits, pit membranes, vessel ends, vessels, xylem embolism, xylem pathogens     

Molecular Basis of the Core Structure of Tight Junctions

The morphological feature of tight junctions (TJs) fits well with their functions. The core of TJs is a fibril-like proteinaceous structure within the lipid bilayer, the so-called TJ strands. TJ strands in apposing plasma membranes associate with each other to eliminate the intercellular space. A network of paired TJ strands generates a continuous belt that circumscribes each cell to establish the diffusion barrier to the solutes in the paracellular pathway throughout the cellular sheet. Identification and characterization of TJ-associated proteins during the last two decades has unveiled the nature of TJ strands and how they are spatially organized. The interplay between integral membrane proteins, claudins, and cytoplasmic plaque proteins, ZO-1/ZO-2, is critical for TJ formation and function.Tight junctions (TJs) are fascinating structures in terms of their function and morphology. In 1963, using ultrathin-section electron microscopy, Farquhar and Palade described the fine structure of TJs together with adherens junctions (AJs) and desmosomes at the most luminal side of the lateral membrane (Farquhar and Palade 1963). In addition, they demonstrated insightfully that TJs function as permeability seals for mass tracers. Indeed, the structure of TJs observed in electron microscopy indicates that TJs could physically restrict the leak of solutes through the intercellular space. However, physiological studies at the same time revealed that solute transport occurred via the intercellular space in a variety of epithelial cells. A resolution of these different views of TJ function comprises the current concept that the TJ regulates the diffusion of solutes with size and charge selectivity and that it is functionally different in physiologically diverse epithelial cell types (Powell 1981; Anderson and Cereijido 2001). To understand the molecular mechanism controlling TJ structure and function, it is important to determine their molecular composition and organization.Although purification of TJs is difficult, Stevenson and Goodenough developed an isolation method for a TJ-enriched plasma membrane fraction from rodent liver. They discovered the first TJ-associated protein, ZO-1, in 1986 by generating monoclonal antibodies against this fraction (Stevenson et al. 1986). Since then, many molecular components of TJs have been identified using immunological approaches or searches for binding proteins with known molecules, which have enabled detailed molecular cell biological analyses of TJs. Among the TJ-associated proteins, the claudin family of membrane proteins identified in 1998 by the Tsukita group are key molecules in the architecture and barrier function of TJs (Furuse et al. 1998a). Functional analyses of claudins have allowed remarkable progress in the development of a comprehensive understanding of the molecular basis of the ultrastructure and physiological characteristics of TJs (Van Itallie and Anderson 2006; Furuse and Tsukita 2006; Angelow et al. 2008). In addition, the cytoplasmic plaque proteins associated with TJs are important in regulating TJ architecture (Guillemot et al. 2008).In this article, we present the molecular basis for the core structure of TJs based on recent progress in functional analyses of TJ-associated proteins. The current molecular basis of TJ physiology is covered in detail in Anderson and Van Itallie (2009).     

Structure,Regulation and Function of Gap Junctions in Liver

Abstract

Gap junctions are a specialized group of cell-to-cell junctions that mediate direct intercellular communication between cells. They arise from the interaction of two hemichannels of adjacent cells, which in turn are composed of six connexin proteins. In liver, gap junctions are predominantly found in hepatocytes and play critical roles in virtually all phases of the hepatic life cycle, including cell growth, differentiation, liver-specific functionality and cell death. Liver gap junctions are directed through a broad variety of mechanisms ranging from epigenetic control of connexin expression to post-translational regulation of gap junction activity. This paper reviews established and novel aspects regarding the architecture, control and functional relevance of liver gap junctions.     

Origins of the Electrical Potential Difference between the Xylem Sap of Maize Roots and the External Solution

The effects of 2,4-dinitrophenol, of changes in the temperatureand concentration of the ambient solution and of variationsin salt status on the electrical potential difference betweenthe xylem exudate of maize roots and the ambient solution havebeen examined. The results are discussed in the light of someof the factors which could give rise to a potential differencebetween the sap and the solution. The rapid response of thepotential difference to dinitrophenol and to changes in temperaturesuggest that, at least in part, it arises directly from metabolicprocesses. Rapid changes in the potential difference broughtabout by addition of salts may be attributed to differentialrates of movement of anions and cations in the initial uptakeprocess. Over longer periods the potential difference appearsto be dependent on the concentration, but not the compositionof the ambient solution, and on the salt status of the roots.The salt status influences the relative rates at which anionsand cations are transported to the xylem sap, and a correlationhas been found between the potential difference and the ratioof the rates of movement of chloride or sulphate to potassiumto the sap. The implications of these findings on the elucidationof the pathways whereby ions are transported to the sap arediscussed.     

药用植物东北雷公藤茎次生木质部结构研究

对东北雷公藤苇次生木质部结构及其与卫矛科其它属进行比较研究,为东北雷公藤生药鉴定提供了理论依据。结果表明：东北雷公藤茎次生木质部早晚材导管分子大小差异极大．为环孔材;导管分子穿孔板均为单穿孔板;异型Ⅱ-b射线;纤维管胞具螺纹增厚;具分隔木纤维。     

The Supramolecular Architecture in Chloroplast Thylakoid Membranes of Warigal Wheat

Thylakoid membranes of chloroplast from first leaf and flag leaf of wheat Warigal were examined by freeze-fracture and rotary shadowing etectron microscopy. The shape, size, density and size distribution of freeze-fracture partieles of their four faces were measured and plotted as three-dimensional histograms by a Hewlett-packard 9874 A digitizer with a HP 9845 B Computer and HP 9872 C plotter. When comparisons were made among different fracture faces and between the corresponding faces of the first leaf and the flag leaf, we found that the supramolecular architecture on the four fracture faces of the flag leaf differs from that on the corresponding faces of the chloroplast thytakoid membranes of the first leaf. The most significant difference was that the EFs particles contain the photosystem Ⅱ reaction centres associated with LHCP and the PFs particles were mostly light-harvesting complex. There was a 15% increase in EFs particle density, a 22% increase in PFs particle density and a 28% increase in EFu particle density. The large PFu particles contained the photosystem Ⅰ reaction centre and the flag lcaves contained 5% more than the first leaves. In addition, the stacking of thylakoid membranes in the flag leaf was 5% more than those in the first leaf.Thus, it provides theoretical basis for the fact that the flag leaf has higher photosynthetic rate.     

Damage to Photosynthetic Membranes in Chilling-Sensitive Plants: Maize,a Case Study

Abstract

Biotechnology may soon take greater advantage of extremophiles — microorganisms that grow in high salt or heavy metal concentrations, or at extremes of temperature, pressure, or pH. These organisms and their cellular components are attractive because they permit process operation over a wider range of conditions than their traditional counterparts. However, extremophiles also present a number of challenges for the development of bioprocesses, such as slow growth, low cell yield, and high shear sensitivity. Difficulties inherent in designing equipment suitable for extreme conditions are also encountered. This review describes both the advantages and disadvantages of extremophiles, as well as the specialized equipment required for their study and application in biotechnology.     

Tritrichomonas foetus: Fine Structure of Freeze-Fractured Membranes1

ABSTRACT. Freeze-fracture techniques reveal differences in fine structure between the anterior three flagella of Tritrichomonas foetus and its recurrent flagellum. The anterior flagella have rosettes of 9–12 intramembranous particles on both the P and E faces. The recurrent flagellum lacks rosettes but has ribbon-like arrays of particles along the length of the flagellum, which may be involved in the flagellum's attachment to the cell body. This flagellum is attached to the membrane of the cell body along a distinct groove that contains few discernible particles. Some large intramembranous particles are visible on the P face of the cell body membrane at the point where the flagellum emerges from the cell body. The randomly distributed particles on the P and E faces of the plasma membrane have a particle density of 919/μm² and 468/μm² respectively, and there are areas on both faces that are devoid of particles. Freeze-fracture techniques also reveal numerous fenestrations in the membrane of the Golgi complex and about 24 pores per μm² in the nuclear. membrane.