Model for Cell Wall Growth of Streptococcus faecalis

In exponentially growing and dividing cells of Streptococcus faecalis, it is proposed that the leading edge of the annularly closing cross wall is the point of extension for both cross wall and peripheral wall. Peripheral wall extension is thought to be produced by the separation or splitting of the cross wall at its junction with peripheral wall. This results in the pushing of the equatorial wall bands, found on S. faecalis walls, to subsequatorial positions. These bands therefore mark the separation of old wall from new wall. Mesosomal formation was observed usually to precede cross wall initiation.     

Wall Relaxation and the Driving Forces for Cell Expansive Growth

When water uptake by growing cells is prevented, the turgor pressure and the tensile stress in the cell wall are reduced by continued wall loosening. This process, termed in vivo stress relaxation, provides a new way to study the dynamics of wall loosening and to measure the wall yield threshold and the physiological wall extensibility. Stress relaxation experiments indicate that wall stress supplies the mechanical driving force for wall yielding. Cell expansion also requires water absorption. The driving force for water uptake during growth is created by wall relaxation, which lowers the water potential of the expanding cells. New techniques for measuring this driving force show that it is smaller than believed previously; in elongating stems it is only 0.3 to 0.5 bar. This means that the hydraulic resistance of the water transport pathway is small and that rate of cell expansion is controlled primarily by wall loosening and yielding.     

Pulse wave velocity as a diagnostic index: the pitfalls of tethering versus stiffening of the arterial wall

Pulse wave velocity (PWV) is often used as a clinical index of aging, vascular disease, or age related hypertension. This practice is based on the assumption that a higher wave speed indicates vascular stiffening. This assumption is well grounded in the physics of pulsatile flow of an incompressible fluid where it is fully established that a pulse wave travels faster in a tube of stiffer wall, the wave speed becoming infinite in the mathematical limit of a rigid wall. However, in this paper we point out that the physical principal of higher pulse wave velocity in a stiffer tube is strictly valid only when the wall is free from outside constraints, which in the physiological setting is present in the form of tethering of the vessel wall. The use of PWV as an index of arterial stiffening may thus lose its validity if tethering is involved. A solution of the problem of vessel wall mechanics as they arise from the physiological pulsatile flow problem is presented for the purpose of resolving this issue. The vessel wall is considered to have finite thickness with or without tethering and with a range of mechanical properties ranging from viscoelastic to stiff. The results show that, indeed, while the wave speed becomes infinite in the mathematical limit of a rigid free wall, the opposite actually happens if the vessel wall is tethered. Here the wave speed actually diminishes as the degree of tethering increases. This dichotomy in the effects of tethering versus stiffening of the arterial wall may clearly lead to error in the interpretation of PWV as an index of vessel wall stiffness. In particular, a normal value of PWV may lead to the conclusion that vessel wall stiffening is absent while this value may in fact have been lowered by tethering. In other words, the diagnostic test may lead to a false negative diagnosis. Our results indicate that the reason for which PWV is lower in a tethered wall compared with that in a free wall of the same stiffness is that the radial movements of the wall are greatly reduced by tethering. More precisely, the results show that PWV depends strongly on the ratio of radial to axial displacements and that this ratio is much lower in a tethered wall than it is in a free wall of the same stiffness.     

Differential histone modification and protein expression associated with cell wall removal and regeneration in rice (Oryza sativa)

The cell wall is a critical extracellular structure that provides protection and structural support in plant cells. To study the biological function of the cell wall and the regulation of cell wall resynthesis, we examined cellular responses to enzymatic removal of the cell wall in rice (Oryza sativa) suspension cells using proteomic approaches. We find that removal of cell wall stimulates cell wall synthesis from multiple sites in protoplasts instead of from a single site as in cytokinesis. Nucleus DAPI stain and MNase digestion further show that removal of the cell wall is concomitant with substantial chromatin reorganization. Histone post-translational modification studies using both Western blots and isotope labeling assisted quantitative mass spectrometry analyses reveal that substantial histone modification changes, particularly H3K18(AC) and H3K23(AC), are associated with the removal and regeneration of the cell wall. Label-free quantitative proteome analyses further reveal that chromatin associated proteins undergo dramatic changes upon removal of the cell wall, along with cytoskeleton, cell wall metabolism, and stress-response proteins. This study demonstrates that cell wall removal is associated with substantial chromatin change and may lead to stimulation of cell wall synthesis using a novel mechanism.     

Estimating wall guidance and attraction in mouse free locomotor behavior

In this study, we estimate the influence exerted by the wall of the Open Field on the trajectory of the mouse. The wall exerts two types of influence on the mouse's path: one of guidance and one of attraction. The guiding influence is expressed by the tendency of mice to progress in parallel to the wall. This tendency wanes with increasing distance from the wall but is observed at large distances from it. The more parallel the mouse is to the wall the higher is its speed, even when distant from the wall. This association between heading direction and speed shows that the mouse controls its heading in reference to the wall. It is also observed in some blind strains, revealing that wall-guidance is not based exclusively on vision. The attraction influence is reflected by movement along the wall and by the asymmetry between speed during movement toward, and during movement away from the wall: sighted mice move faster toward the wall, whereas blind mice use similar speeds in both directions. Measures characterizing these influences are presented for five inbred strains, revealing heritable components that are replicable across laboratories. The revealed structure can lead to the identification of distinct groups of genes that mediate the distinct influences of guidance and attraction exerted by the wall. It can also serve as a framework for the decoding of electrophysiological data recorded in free moving rodents in the Open Field.     

On Vessel Member Differentiation in the Bean (Phaseolus vulgaris L.)

Certain ultrastructural features of vessel member differentiationwere examined in the primary xylem of petiole of bean (Phaseolusvulgaris L.). The cells used had helical secondary wall thickeningsand simple perforation plates. The primary cell wall increasesin thickness before the helices of secondary wall develop. Ina common wall between two vessel members of different ages,theprimary thickening occurs first in the older cell so thatfor a time the middle lamella is located closer to the youngercell rather than medianly. Apparently the helix is depositedafter the primary wall of a given cell reaches maximum thickness.The perforation of the end wall is preceded by primary thickeningof the part of the wall that is later removed. The marginalregion remains relatively thin and becomes covered with a rimof secondary wall. Vesicles with fibrous content appear nearthe surface within the end wall shortly before the perforationoccurs. A highly vacuolated protoplast with a much enlargednucleus and numerous organdIes is present during cell wall differentiation.After that process is completed, the protoplast disintegratesand the primary wall bearing the helix is hydrolysed where itis exposed to the cell lumen and, under certain conditions,also under the secondary wall.     

Regulation of cell wall biogenesis in Saccharomyces cerevisiae: the cell wall integrity signaling pathway

The yeast cell wall is a strong, but elastic, structure that is essential not only for the maintenance of cell shape and integrity, but also for progression through the cell cycle. During growth and morphogenesis, and in response to environmental challenges, the cell wall is remodeled in a highly regulated and polarized manner, a process that is principally under the control of the cell wall integrity (CWI) signaling pathway. This pathway transmits wall stress signals from the cell surface to the Rho1 GTPase, which mobilizes a physiologic response through a variety of effectors. Activation of CWI signaling regulates the production of various carbohydrate polymers of the cell wall, as well as their polarized delivery to the site of cell wall remodeling. This review article centers on CWI signaling in Saccharomyces cerevisiae through the cell cycle and in response to cell wall stress. The interface of this signaling pathway with other pathways that contribute to the maintenance of cell wall integrity is also discussed.     

Cell wall analysis

The cell wall is a rigid structure essential for survival of the fungal cell. Because of its absence in mammalian cells, the cell wall is an attractive target for antifungal agents. Thus, for different reasons, it is important to know how the cell wall is synthesized and how different molecules regulate that synthesis. The Schizosaccharomyces pombe cell wall is mainly formed by glucose polysaccharides and some galactomannoproteins. Here, we describe a fast and reliable method to analyze changes in S. pombe cell wall composition by using specific enzymatic degradation and chemical treatment of purified cell walls. This approach provides a powerful means to analyze changes in (1,3)beta-glucan and (1,3)alpha-glucan, two main polysaccharides present in fungal cell walls. Analysis of cell wall polymers will be useful to search for new antifungal drugs that may inhibit cell wall biosynthesis and/or alter cell wall structure.     

The molecular mechanism of stipe cell wall extension for mushroom stipe elongation growth

Stipe elongation growth is one of the remarkable characteristics of the growth and development of basidiomycete fruiting bodies. Stipe elongation is resulting from the lateral extension of stipe cells. The stipe cell is enclosed within a thin cell wall which must be loosened to expand the wall surface area for accommodation of the enlarged protoplast as the stipe cell elongates. In fungal cell walls, chitin molecules associate with each other by interchain hydrogen bonds to form chitin microfibrils which are cross-linked covalently to matrix polysaccharides. Early, some scientists proposed that stipe elongation was the result of enzymatic degradation of wall polysaccharides, whereas other researchers suggested that stipe elongation resulted from nonhydrolytic disruption of the hydrogen bonds by turgor pressure between wall polysaccharides. Recently, an extensometer was used to determine stipe wall extension for elucidation of the molecular mechanism of stipe elongation. In Coprinopsis cinerea, the native stipe cell wall is induced to extend by acidic buffers and the acid-induced native wall extension activity is located in the growing apical stipe region. A series of current experiments indicate that chitinases play a key role in the stipe wall extension, and β-glucanases mainly function in the wall remodeling for regulation of stipe wall expansibility to cooperate with chitinase to induce stipe wall extension. In addition, fungal expansin-like proteins can bind to chitin to enhance chitin hydrolysis, and their expression pattern is consistent with the stipe elongation growth, which is suggested to play an auxiliary role in the stipe wall extension.     

Cellulose synthesis in two secondary cell wall processes in a single cell type

Plant cells have a rigid cell wall that constrains internal turgor pressure yet extends in a regulated and organized manner to allow the cell to acquire shape. The primary load-bearing macromolecule of a plant cell wall is cellulose, which forms crystalline microfibrils that are organized with respect to a cell''s function and shape requirements. A primary cell wall is deposited during expansion whereas secondary cell wall is synthesized post expansion during differentiation. A complex form of asymmetrical cellular differentiation occurs in Arabidopsis seed coat epidermal cells, where we have recently shown that two secondary cell wall processes occur that utilize different cellulose synthase (CESA) proteins. One process is to produce pectinaceous mucilage that expands upon hydration and the other is a radial wall thickening that reinforced the epidermal cell structure. Our data illustrate polarized specialization of CESA5 in facilitating mucilage attachment to the parent seed and CESA2, CESA5 and CESA9 in radial cell wall thickening and formation of the columella. Herein, we present a model for the complexity of cellulose biosynthesis in this highly differentiated cell type with further evidence supporting each cellulosic secondary cell wall process.     

Feedback from the wall

The ability of cells to perceive changes in the composition and mechanical properties of their cell wall is crucial for plants to achieve coordinated growth and development. Evidence is accumulating to show that the plant cell wall, like its yeast counterpart, is capable of triggering multiple signalling pathways. The components of the cell wall that are responsible for initiating these signal responses remain unknown; however, recent technological advances in cell wall analysis may now facilitate the identification of these components and accelerate the characterisation of changes that occur in cell wall mutants.     

ULTRASTRUCTURE AND INITIATION OF WALL PATTERN IN PEDIASTRUM BORYANUM

The reticulate pattern in the wall of Pediastrum boryanum emerges rapidly during wall formation following aggregation of the swarming zoospores to form the coenobium. Electron micrographs during colony formation show that microtubules, present during the motile phase and aggregation, are gone prior to wall formation and probably do not participate in wall pattern regulation. A single dictyosome lies adjacent to the nucleus and from blebs of the nuclear membrane receives vesicles at its forming face. Vesicles formed at the maturing face have not been observed to contribute to the cell wall. Electron-lucent patches occur in the plasma membrane prior to wall formation. The first indication of a reticulate pattern in wall development is the deposition on the plasma membrane of interconnected plaques of outer wall material at the corners of hexagons. The sites of the plaques may correspond to clusters of ribosomes on endoplasmic reticulum underlying the plasmalemma. Following completion of the outer wall the thicker inner wall layer is deposited and within it the reticulate pattern of ridges is soon evident in tangential sections as strips of greater electron density. It is suggested that the pattern of the wall is templated by the plasma membrane.     

Wall stress and flow dynamics in abdominal aortic aneurysms: finite element analysis vs. fluid-structure interaction

Abdominal aortic aneurysm (AAA) rupture is the clinical manifestation of an induced force exceeding the resistance provided by the strength of the arterial wall. This force is most frequently assumed to be the product of a uniform luminal pressure acting along the diseased wall. However fluid dynamics is a known contributor to the pathogenesis of AAAs, and the dynamic interaction of blood flow and the arterial wall represents the in vivo environment at the macro-scale. The primary objective of this investigation is to assess the significance of assuming an arbitrary estimated peak fluid pressure inside the aneurysm sac for the evaluation of AAA wall mechanics, as compared with the non-uniform pressure resulting from a coupled fluid-structure interaction (FSI) analysis. In addition, a finite element approach is utilised to estimate the effects of asymmetry and wall thickness on the wall stress and fluid dynamics of ten idealised AAA models and one non-aneurysmal control. Five degrees of asymmetry with uniform and variable wall thickness are used. Each was modelled under a static pressure-deformation analysis, as well as a transient FSI. The results show that the inclusion of fluid flow yields a maximum AAA wall stress up to 20% higher compared to that obtained with a static wall stress analysis with an assumed peak luminal pressure of 117 mmHg. The variable wall models have a maximum wall stress nearly four times that of a uniform wall thickness, and also increasing with asymmetry in both instances. The inclusion of an axial stretch and external pressure to the computational domain decreases the wall stress by 17%.     

Wall stress and flow dynamics in abdominal aortic aneurysms: finite element analysis vs. fluid–structure interaction

Abdominal aortic aneurysm (AAA) rupture is the clinical manifestation of an induced force exceeding the resistance provided by the strength of the arterial wall. This force is most frequently assumed to be the product of a uniform luminal pressure acting along the diseased wall. However fluid dynamics is a known contributor to the pathogenesis of AAAs, and the dynamic interaction of blood flow and the arterial wall represents the in vivo environment at the macro-scale. The primary objective of this investigation is to assess the significance of assuming an arbitrary estimated peak fluid pressure inside the aneurysm sac for the evaluation of AAA wall mechanics, as compared with the non-uniform pressure resulting from a coupled fluid–structure interaction (FSI) analysis. In addition, a finite element approach is utilised to estimate the effects of asymmetry and wall thickness on the wall stress and fluid dynamics of ten idealised AAA models and one non-aneurysmal control. Five degrees of asymmetry with uniform and variable wall thickness are used. Each was modelled under a static pressure-deformation analysis, as well as a transient FSI. The results show that the inclusion of fluid flow yields a maximum AAA wall stress up to 20% higher compared to that obtained with a static wall stress analysis with an assumed peak luminal pressure of 117 mmHg. The variable wall models have a maximum wall stress nearly four times that of a uniform wall thickness, and also increasing with asymmetry in both instances. The inclusion of an axial stretch and external pressure to the computational domain decreases the wall stress by 17%.     

Synthesis of the inner cell wall layer of the chlamydomonad flagellate,Gloeomonas kupfferi

Summary An antibody to the inner wall layer ofGloeomonas kupfferi was isolated and used in a developmental analysis of cell wall processing, secretion and extracellular assembly. The focus of the processing of this matrix layer is the endomembrane system, in particular the Golgi apparatus (GA) and contractile vacuole (CV). During interphase, inner wall materials are processed in the GA, packaged in trans face vesicles and transported to the CV, the final internal depository of wall precursors until release to the cell surface. During cell division, significant changes occur in the inner wall layer processing. Early on in cytokinesis, the GA does not label with our antibody, suggesting that other wall layers are being processed. In later stages of cytokinesis, the GA changes in morphology and begins to produce inner wall layer materials. These wall precursors are shuttled to the CV where they are released around the daughter cell protoplasts. The first wall layer that is formed around daughter cells is the crystalline median wall layer. Once assembled, the inner wall layer condenses upon the crystalline layer and grows in size.     

Dynamics of cell wall structure in Saccharomyces cerevisiae

The cell wall of Saccharomyces cerevisiae is an elastic structure that provides osmotic and physical protection and determines the shape of the cell. The inner layer of the wall is largely responsible for the mechanical strength of the wall and also provides the attachment sites for the proteins that form the outer layer of the wall. Here we find among others the sexual agglutinins and the flocculins. The outer protein layer also limits the permeability of the cell wall, thus shielding the plasma membrane from attack by foreign enzymes and membrane-perturbing compounds. The main features of the molecular organization of the yeast cell wall are now known. Importantly, the molecular composition and organization of the cell wall may vary considerably. For example, the incorporation of many cell wall proteins is temporally and spatially controlled and depends strongly on environmental conditions. Similarly, the formation of specific cell wall protein-polysaccharide complexes is strongly affected by external conditions. This points to a tight regulation of cell wall construction. Indeed, all five mitogen-activated protein kinase pathways in bakers' yeast affect the cell wall, and additional cell wall-related signaling routes have been identified. Finally, some potential targets for new antifungal compounds related to cell wall construction are discussed.     

Cell wall sorting of lipoproteins in Staphylococcus aureus.

Many surface proteins are thought to be anchored to the cell wall of gram-positive organisms via their C termini, while the N-terminal domains of these molecules are displayed on the bacterial surface. Cell wall anchoring of surface proteins in Staphylococcus aureus requires both an N-terminal leader peptide and a C-terminal cell wall sorting signal. By fusing the cell wall sorting of protein A to the C terminus of staphylococcal beta-lactamase, we demonstrate here that lipoproteins can also be anchored to the cell wall of S. aureus. The topology of cell wall-anchored beta-lactamase is reminiscent of that described for Braun's murein lipoprotein in that the N terminus of the polypeptide chain is membrane anchored whereas the C-terminal end is tethered to the bacterial cell wall.     

The glycoprotein NOWA and minicollagens are part of a disulfide-linked polymer that forms the cnidarian nematocyst wall

The nematocyst is a unique extrusive organelle involved in the defense and capture of prey in cnidarians. Minicollagens and the glycoprotein NOWA are major components of the nematocyst capsule wall, which resists osmotic pressure of 15 MPa. Here we present the recombinant expression of NOWA, which spontaneously assembles to globular macromolecular particles that are sensitive to reduction as the native wall structure. Ultra-structural analysis showed that the Hydra nematocyst wall is composed of several layers of globular particles, which are interconnected via radiating rodlike protrusions. Evidence is presented that native wall particles contain NOWA and minicollagen, supposed to be linked via disulfide bonds between their homologous cysteine-rich domains. Our data suggest a continuous suprastructure of the nematocyst wall, assembled from wall proteins that share a common oligomerization motif.     

Cell Wall Modifying Proteins Mediate Plant Acclimatization to Biotic and Abiotic Stresses

Cell wall modification is an important aspect of plant acclimation to environmental stresses. Structural changes of the existing cell wall mediated by various cell wall modifying proteins help a plant adjust to environmental changes by regulating growth and policing the entry of biotic agents. For example, accelerated shoot growth during submergence and shading allows some plants to escape these unfavorable conditions. This is mediated by the regulation of wall modifying proteins that alter cell wall structure and allow it to yield to turgor, thus fueling cellular expansion. Regulation of cell wall protein activity results in growth modulation during drought, where maintenance of root growth through changes in wall extensibility is an important adaptation to water deficit. Freeze-tolerant plants adjust their cell wall properties to prevent freezing-induced dehydration and also use the cell wall as a barrier against ice crystal propagation. Cell wall architecture is an important determinant of plant resistance to biotic stresses. A rigid cell wall can fend off pathogen attack by forming an impenetrable, physical barrier. When breached, products released during wall modification can trigger plant defense signaling. This review documents and discusses studies demonstrating the importance of timely cell wall modification during plant stress responses by focusing on a well-researched subset of wall modifying proteins.