Individual Cell Wall Research Articles

Multiscale stress-strain characterization of onion outer epidermal tissue in wet and dry states.

• Quantitative measurements of water's effects on the tension response of plant tissue will assist in understanding the regulatory mechanism underlying expansive growth. Such measurements should be multiscale in nature to account for plants' hierarchical structure.• Outer onion epidermal tissues were cut and bonded to uniaxial displacement-controlled mechanical loading devices to apply and measure the force on the sample. Fluorescent polystyrene beads (500 nm in diameter) were dispersed on the sample surface under various levels of tensile load conditions to obtain displacement maps with a confocal fluorescent microscope. The resulting strain was measured using a digital image correlation technique by tracking individual bead displacements. The applied forces were obtained by measuring the displacement of the calibrated force-sensing device. Tissue- and cell-scale mechanical properties were quantified by calculating the applied stress and the corresponding global and local strains.• The Young's modulus values of individual cell walls of dehydrated and rehydrated samples were 3.0 ± 1.0 GPa and 0.4 ± 0.2 GPa, respectively, and are different from the Young's modulus values of the global tissue-scale dehydrated and rehydrated samples, which were 1.9 ± 0.3 GPa and 0.08 ± 0.02 GPa, respectively. Poisson's ratio increased more than 3-fold due to hydration.• The results on global, cell-to-cell, and point-to-point mechanical property variations suggest the importance of the mechanical contribution of extracellular features including the middle lamella, cell shape, and dimension. This study shows that a multiscale investigation is essential for fundamental insights into the hierarchical deformation of biological systems.

Strain-rate sensitivity of foam materials: A numerical study using 3D image-based finite element model

Realistic simulations are increasingly demanded to clarify the dynamic behaviour of foam materials, because, on one hand, the significant variability (e.g. 20% scatter band) of foam properties and the lack of reliable dynamic test methods for foams bring particular difficulty to accurately evaluate the strain-rate sensitivity in experiments; while on the other hand numerical models based on idealised cell structures (e.g. Kelvin and Voronoi) may not be sufficiently representative to capture the actual structural effect. To overcome these limitations, the strain-rate sensitivity of the compressive and tensile properties of closed-cell aluminium Alporas foam is investigated in this study by means of meso-scale realistic finite element (FE) simulations. The FE modelling method based on X-ray computed tomography (CT) image is introduced first, as well as its applications to foam materials. Then the compression and tension of Alporas foam at a wide variety of applied nominal strain-rates are simulated using FE model constructed from the actual cell geometry obtained from the CT image. The stain-rate sensitivity of compressive strength (collapse stress) and tensile strength (0.2% offset yield point) are evaluated when considering different cell-wall material properties. The numerical results show that the rate dependence of cell-wall material is the main cause of the strain-rate hardening of the compressive and tensile strengths at low and intermediate strain-rates. When the strain-rate is sufficiently high, shock compression is initiated, which significantly enhances the stress at the loading end and has complicated effect on the stress at the supporting end. The plastic tensile wave effect is evident at high strain-rates, but shock tension cannot develop in Alporas foam due to the softening associated with single fracture process zone occurring in tensile response. In all cases the micro inertia of individual cell walls subjected to localised deformation is found to have negligible effect on the macro strain-rate sensitivity of Alporas foam.

Bacterial cell wall research in Tübingen: A brief historical account

Research in Tübingen on bacterial cell walls began in 1951 and continues to this day. The studies over the decades reflect the development in the field, which was strongly influenced by the design of suitable biochemical and genetic methods used to unravel the highly complex envelope structure. At the beginning of this period, improper crude extraction and solubilization methods were employed in an attempt to isolate pure components. Nevertheless, progress was steady and culminated in major insights into the structure and function of individual cell wall components and the cell wall as a whole. The “cell wall” has various definitions. In this short overview, the term includes the cell wall of gram-positive bacteria in the strict sense, and also the outer membrane, the murein (peptidoglycan) and the outer membrane of gram-negative bacteria and the cytoplasmic membranes.

Morphology and ultrastructure of Interfilum and Klebsormidium (Klebsormidiales, Streptophyta) with special reference to cell division and thallus formation

Representatives of the closely related genera, Interfilum and Klebsormidium, are characterized by unicells, dyads or packets in Interfilum and contrasting uniseriate filaments in Klebsormidium. According to the literature, these distinct thallus forms originate by different types of cell division, sporulation (cytogony) versus vegetative cell division (cytotomy), but investigations of their morphology and ultrastructure show a high degree of similarity. Cell walls of both genera are characterized by triangular spaces between cell walls of neighbouring cells and the parental wall or central space among the walls of a cell packet, exfoliations and projections of the parental wall and cap-like and H-like fragments of the cell wall. In both genera, each cell has its individual cell wall and it also has part of the common parental wall or its remnants. Therefore, vegetative cells of Interfilum and Klebsormidium probably divide by the same type of cell division (sporulation-like). Various strains representing different species of the two genera are characterized by differences in cell wall ultrastructure, particularly the level of preservation, rupture or gelatinization of the parental wall surrounding the daughter cells. The differing morphologies of representatives of various lineages result from features of the parental wall during cell separation and detachment. Cell division in three planes (usual in Interfilum and a rare event in Klebsormidium) takes place in spherical or short cylindrical cells, with the chloroplast positioned perpendicularly or obliquely to the filament (dyad) axis. The morphological differences are mainly a consequence of differing fates of the parental wall after cell division and detachment. The development of different morphologies within the two genera mostly depends on characters such as the shape of cells, texture of cell walls, mechanical interactions between cells and the influence of environmental conditions.

Plant and algal cell walls: diversity and functionality.

Although plants and many algae (e.g. the Phaeophyceae, brown, and Rhodophyceae, red) are only very distantly related they are united in their possession of carbohydrate-rich cell walls, which are of integral importance being involved in many physiological processes. Furthermore,wall components have applications within food, fuel, pharmaceuticals, fibres (e.g. for textiles and paper) and building materials and have long been an active topic of research. As shown in the 27 papers in this Special Issue, as the major deposit of photosynthetically fixed carbon, and therefore energy investment, cell walls are of undisputed importance to the organisms that possess them, the photosynthetic eukaryotes ( plants and algae). The complexities of cell wall components along with their interactions with the biotic and abiotic environment are becoming increasingly revealed. The importance of plant and algal cell walls and their individual components to the function and survival of the organism, and for a number of industrial applications, are illustrated by the breadth of topics covered in this issue, which includes papers concentrating on various plants and algae, developmental stages, organs, cell wall components, and techniques. Although we acknowledge that there are many alternative ways in which the papers could be categorized (and many would fit within several topics), we have organized them as follows: (1) cell wall biosynthesis and remodelling, (2) cell wall diversity, and (3) application of new technologies to cell walls. Finally, we will consider future directions within plant cell wall research. Expansion of the industrial uses of cell walls and potentially novel uses of cell wall components are both avenues likely to direct future research activities. Fundamentally, it is the continued progression from characterization (structure, metabolism, properties and localization) of individual cell wall components through to defining their roles in almost every aspect of plant and algal physiology that will present many of the major challenges in future cell wall research.

A rapid live-cell ELISA for characterizing antibodies against cell surface antigens of Chlamydomonas reinhardtii and its use in isolating algae from natural environments with related cell wall components.

BackgroundCell walls are essential for most bacteria, archaea, fungi, algae and land plants to provide shape, structural integrity and protection from numerous biotic and abiotic environmental factors. In the case of eukaryotic algae, relatively little is known of the composition, structure or mechanisms of assembly of cell walls in individual species or between species and how these differences enable algae to inhabit a great diversity of environments. In this paper we describe the use of camelid antibody fragments (VHHs) and a streamlined ELISA assay as powerful new tools for obtaining mono-specific reagents for detecting individual algal cell wall components and for isolating algae that share a particular cell surface component.ResultsTo develop new microalgal bioprospecting tools to aid in the search of environmental samples for algae that share similar cell wall and cell surface components, we have produced single-chain camelid antibodies raised against cell surface components of the single-cell alga, Chlamydomonas reinhardtii. We have cloned the variable-region domains (VHHs) from the camelid heavy-chain-only antibodies and overproduced tagged versions of these monoclonal-like antibodies in E. coli. Using these VHHs, we have developed an accurate, facile, low cost ELISA that uses live cells as a source of antigens in their native conformation and that requires less than 90 minutes to perform. This ELISA technique was demonstrated to be as accurate as standard ELISAs that employ proteins from cell lysates and that generally require >24 hours to complete. Among the cloned VHHs, VHH B11, exhibited the highest affinity (EC50 < 1 nM) for the C. reinhardtii cell surface. The live-cell ELISA procedure was employed to detect algae sharing cell surface components with C. reinhardtii in water samples from natural environments. In addition, mCherry-tagged VHH B11 was used along with fluorescence activated cell sorting (FACS) to select individual axenic isolates of presumed wild relatives of C. reinhardtii and other Chlorphyceae from the same environmental samples.ConclusionsCamelid antibody VHH domains provide a highly specific tool for detection of individual cell wall components of algae and for allowing the selection of algae that share a particular cell surface molecule from diverse ecosystems.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-014-0244-0) contains supplementary material, which is available to authorized users.

Mechanical modelling quantifies the functional importance of outer tissue layers during root elongation and bending.

Root elongation and bending require the coordinated expansion of multiple cells of different types. These processes are regulated by the action of hormones that can target distinct cell layers. We use a mathematical model to characterise the influence of the biomechanical properties of individual cell walls on the properties of the whole tissue.Taking a simple constitutive model at the cell scale which characterises cell walls via yield and extensibility parameters, we derive the analogous tissue-level model to describe elongation and bending. To accurately parameterise the model, we take detailed measurements of cell turgor, cell geometries and wall thicknesses.The model demonstrates how cell properties and shapes contribute to tissue-level extensibility and yield. Exploiting the highly organised structure of the elongation zone (EZ) of the Arabidopsis root, we quantify the contributions of different cell layers, using the measured parameters. We show how distributions of material and geometric properties across the root cross-section contribute to the generation of curvature, and relate the angle of a gravitropic bend to the magnitude and duration of asymmetric wall softening.We quantify the geometric factors which lead to the predominant contribution of the outer cell files in driving root elongation and bending.

The PGRS domain is responsible for translocation of PE_PGRS30 to cell poles while the PE and the C‐terminal domains localize it to the cell wall

PE_PGRS proteins localize in the mycobacterial cell wall and the cell wall localization of PE_PGRS33 has been shown to be attributed to its PE domain. In this study, we expressed deletion mutants of PE_PGRS30 in Mycobacterium smegmatis to characterize the role of its domains in protein localization. It was revealed that, apart from the PE domain, the C-terminal domain present in few PE_PGRS proteins carries individual cell wall localization signals. Proteinase K sensitivity assay showed that PE_PGRS30 is exposed on the mycobacterial surface through its PGRS domain. PGRS domain was also shown to be responsible for polar localization of PE_PGRS30.

Effects of biodegradation by brown-rot decay on selected wood properties in eastern white cedar (Thuja occidentalis L.)

The effects of decay on wood density, wood structure, chemical composition, and mechanical properties in eastern white cedar (Thuja occidentalis L.) were investigated. Decay stage was determined by X-ray densitometry and scanning electron microscopy (SEM). Chemical and mechanical properties of decayed and sound wood samples were also determined. SEM results showed that decay colonization varied among wood cell types and within individual cell wall layers. Growth of fungi causing brown rot decay was limited and slower in latewood than in earlywood due to the narrow cell lumen, thicker wall, and higher density of latewood. Decay-related changes in wood density were more prominent in earlywood than latewood tracheids. Brown-rot decay selectively removes structural carbohydrate components, increasing the higher lignin/carbohydrate ratio as decay progresses. The relationships between chemical composition, mechanical properties, and weight loss were highly significant, with 15% weight loss leading to 40% loss in modulus of rupture (MOR) and 30% loss in bending modulus of elasticity (MOE). The strength loss (MOR) was attributable to arabinan and galactan loss. The loss in wood stiffness was associated with cellulose (glucan) loss, and weight loss was associated with mannan and xylans losses.

Induction of innate immunity by Aspergillus fumigatus cell wall polysaccharides is enhanced by the composite presentation of chitin and beta-glucan

Chitin and β-glucan are conserved throughout evolution in the fungal cell wall and are the most common polysaccharides in fungal species. Together, these two polysaccharides form a structural scaffold that is essential for the survival of the fungus. In the present study, we demonstrated that Aspergillus fumigatus alkali-insoluble cell wall fragments (AIF), composed of chitin linked covalently to β-glucan, induced enhanced immune responses when compared with individual cell wall polysaccharides. Intranasal administration of AIF induced eosinophil and neutrophil recruitment, chitinase activity, TNF-α and TSLP production in mice lungs. Selective destruction of chitin or β-glucan from AIF significantly reduced eosinophil and neutrophil recruitment as well as chitinase activity and cytokine expression by macrophages, indicating the synergistic effect of the cell wall polysaccharides when presented together as a composite PAMP. We also showed that these cell wall polysaccharides induced chitin-specific IgM in mouse serum. Our in vivo and in vitro data indicate that chitin and β-glucan play important roles in activating innate immunity when presented as composite cell wall PAMPs.

Nanoindentation of wood cell walls: effects of sample preparation and indentation protocol

Nanoindentation has become a valuable tool in wood science. It enables to examine the mechanical properties of the wood cell walls, which are polymeric, multi-layered structures with typical thicknesses of a few micrometers. Despite the intensive use of the method for the characterization of wood cell walls, it is not entirely clear yet how the measurement results may be affected by the way the sample is prepared and the indentation is carried out. This manuscript contributes to clarify these issues, by presenting indentation data for a variety of sample preparation techniques and indentation protocols, and by critically evaluating the observed differences of the obtained indentation moduli and hardnesses. Investigations covered the effect of different embedding materials, including testing of non-embedded cell walls, and of repeated exposure to high temperatures during harsh drying before the indentation test. Moreover, potential edge effects were studied when the indentation size approaches the width of the individual cell wall layers. Using different embedding materials as well as testing non-embedded cell walls did not lead to significant changes in the measured properties. Due to damage during the sample preparation, non-embedded cell walls tend to show substantially higher experimental scatter. Repeated drying prior to embedding had no significant effect on the resulting moduli and hardnesses. Finally, it was found that reasonable mechanical properties can be extracted from the cell corner middle lamella (CCML), even when the size of the indent approaches the diameter of the CCML.

Characterisation of cell wall polysaccharides from rapeseed (Brassica napus) meal

To enable structural characteristics of individual cell wall polysaccharides from rapeseed (Brassica napus) meal (RSM) to be studied, polysaccharide fractions were sequentially extracted. Fractions were analysed for their carbohydrate (linkage) composition and polysaccharide structures were also studied by enzymatic fingerprinting.The RSM fractions analysed contained pectic polysaccharides: homogalacturonan in which 60% of the galacturonic acid residues are methyl-esterified, arabinan branched at the O-2 position and arabinogalactan mainly type II. This differs from characteristics previously reported for Brassica campestris meal, another rapeseed cultivar. Also, in the alkali extracts hemicelluloses were analysed as xyloglucan both of the XXGG- and XXXG-type decorated with galactosyl, fucosyl and arabinosyl residues, and as xylan with O-methyl-uronic acid attached. The final residue after extraction still contained xyloglucan and remaining (pectic) polysaccharides next to cellulose, showing that the cell wall matrix of RSM is very strongly interconnected.

X‐ray imaging of leaf venation networks

Leaf venation networks mediate many plant resource fluxes and are therefore of broad interest to research questions in plant physiology, systematics, paleoecology, and physics. However, the study of these networks is limited by slow and destructive imaging methods. X-ray imaging of leaf veins is potentially rapid, of high resolution, and nondestructive. Here, we have developed theory for absorption- and phase-contrast X-ray imaging. We then experimentally test these approaches using a synchrotron light source and two commercially available X-ray instruments. Using synchrotron light, we found that major veins could be consistently visualized using absorption-contrast imaging with X-ray energies <10keV, while both major and minor veins could be consistently visualized with the use of an iodine contrast agent at an X-ray energy of 33.269keV. Phase-contrast imaging at a range of energies provided high resolution but highlighted individual cell walls more than veins. Both approaches allowed several hundred samples to be processed per d. Commercial X-ray instruments were able to resolve major veins and some minor veins using absorption contrast. These results show that both commercial and synchrotron X-ray imaging can be successfully applied to leaf venation networks, facilitating research in multiple fields.

Determining the polysaccharide composition of plant cell walls

The plant cell wall is a chemically complex structure composed mostly of polysaccharides. Detailed analyses of these cell wall polysaccharides are essential for our understanding of plant development and for our use of plant biomass (largely wall material) in the food, agriculture, fabric, timber, biofuel and biocomposite industries. We present analytical techniques not only to define the fine chemical structures of individual cell wall polysaccharides but also to estimate the overall polysaccharide composition of cell wall preparations. The procedure covers the preparation of cell walls, together with gas chromatography-mass spectrometry (GC-MS)-based methods, for both the analysis of monosaccharides as their volatile alditol acetate derivatives and for methylation analysis to determine linkage positions between monosaccharide residues as their volatile partially methylated alditol acetate derivatives. Analysis time will vary depending on both the method used and the tissue type, and ranges from 2 d for a simple neutral sugar composition to 2 weeks for a carboxyl reduction/methylation linkage analysis.

Nitrosative stress and combinatorial stresses in the pathogen Candida albicans

Candida albicans is a major fungal pathogen that causes frequent mucosal infections in otherwise healthy individuals, and life-threatening systemic infections in immune-compromised patients. The first line of defence against Candida albicans is the release of reactive nitrogen species (RNS), reactive oxygen species (ROS) and cationic fluxes by macrophages and neutrophils. As a result robust responses to RNS, ROS and cationic stresses play major roles in the virulence of C. albicans . This fungus is exposed to all of these stresses simultaneously following phagocytosis. Consequently in this study we examined the responses of this pathogen to combinations of nitrosative, oxidative and cationic stress. The roles of the Cta4-Yhb1, Cap1, Hog1 and Mkc1 signalling pathways, which activate responses to individual nitrosative, oxidative, cationic and cell wall stresses, respectively, were examined. The sensitivity of null mutants to individual and combinatorial stresses was examined, and the activation of these pathways was confirmed by assaying the phosphorylation levels of the key protein kinases. Further evidence of the pathway activation was provided by transcription factor localisation and the expression of downstream target genes. We show that combinatorial nitrosative, oxidative and/or cationic stresses exert unexpected effects upon the classical stress signalling pathways. The relevance to C. albicans infections will be discussed.

Topochemical and transmission electron microscopic studies of bacterial decay in pine (Pinus sylvestris L.) harbour foundation piles

Bacterial degradation of Pinus sylvestris harbour foundation piles was studied topochemically by scanning UV-microspectrophotometry. This analytical technique enables direct imaging of lignin distribution within individual cell wall layers. Additionally, light and transmission electron microscopy (TEM) was used to characterise structural changes of the cell walls. Various decay stages were found in the samples. TEM revealed that the bacterial degradation occurred mainly in the S2, leaving granular remnants in degraded wall portions with lower as well as higher electron density than the surrounding unmodified wall. In the initial stages, topochemical investigations revealed that lignin modification starts in the innermost parts of the secondary wall, most clearly observed in latewood tracheids. During advanced degradation, lignin modification occurs more or less severe in walls of all cell types. However, even in cell portions with intensive decay, the compound middle lamellae and ray tracheids were undegraded. The knowledge about lignin modification at initial stages of wood degradation by bacteria is of fundamental importance to provide more information on the process of cell wall decay.

On the dynamic mechanical properties of open-cell metal foams – A re-assessment of the ‘simple-shock theory’

Metal foams are increasingly used for energy absorption especially in lightweight structures and to resist blast and impact loads. This requires an understanding of the dynamic response of these materials for modelling purposes. As a supplement to Tan et al. (2005a,b), hereinafter referred to as T–L for brevity, this paper provides experimental data for the dynamic mechanical properties of open-cell Duocel® foams having a three-dimensional (3D) distribution of cells. These confirm significant enhancement of the foam’s compressive strength, accompanied by changes in their deformation pattern in certain loading régimes, particularly what has come to be described as the ‘shock’ régime by Zheng et al. (2012). This paper examines experimentally, in a similar fashion as T–L, how the structural response of the individual cell walls is affected by cell-shape anisotropy at the cell (meso)-scale and how this, in turn, alters the pattern of cell crushing and the dynamic, mechanical properties. The distinctive role of cell microinertia and ‘shock’ formation are discussed in relation to the mechanical properties measured for these 3D cylindrical specimens. For consistency the same procedures described in T–L are used. The features identified are shown to be consistent with those observed in finite-element simulations of two-dimensional (2D) honeycombs as estimated by the one-dimensional (1D) steady-shock theory summarised in T–L. The different deformation patterns that develop in the various loading régimes are categorised according to the compression rate/impact speed. Critical values of impact velocity, corresponding to the transition from one pattern to the other, are quantified and predictive formulae for the compressive uniaxial strengths in the directions of two of the principal axes of the material in each loading régime are derived and discussed. The accuracy of the predictive formula in T–L is shown to critically depend on the ‘densification strain’ of the foam specimens. This parameter and the discussion that follows could assist the formulation and validation of alternative theoretical/computational models on the dynamic deformation of such materials.

Validating classical line profile analyses using microbeam diffraction from individual dislocation cell walls and cell interiors

Dislocation structures in deformed metals produce broad asymmetric diffraction line profiles. During analysis, these profiles are generally separated into two nearly symmetric subprofiles corresponding to diffraction by dislocation cell walls and cell interiors. These subprofiles are then interpreted using complex models of dislocation-based line broadening. Until now, it has not been possible to test the many assumptions that are made in such an analysis. Here, depth-resolved microbeam diffraction was used to measure diffraction line profiles from numerous individual dislocation cell walls and cell interiors in a heavily deformed Cu single crystal. Summing these profiles directly constructed the cell-interior and cell-wall subprofiles that have been approximated in the line profile analysis literature for the past 30 years. Direct comparison between the reconstructed subprofiles and the macroscopic asymmetric line profile from the same sample allows the first direct tests of many of the assumptions that have been used for interpreting these X-ray measurements.

Ultrastructural topochemistry of cell wall polymers in Populus nigra by transmission electron microscopy and Raman imaging

The topochemical distribution of lignin and cellulose in individual cell wall layers of Populus nigra stem was determined by transmission electron microscopy (TEM) and confocal Raman microscopy. TEM images exhibited the fiber wall as being typically differentiated into three layers: middle lamella (ML), primary wall (P), and secondary wall (S1, S2, and S3). Higher magnification views showed the S2 layer to be differentiated into electron lucent and dense areas in the radial direction. In situ Raman images calculated by integrating over the intensity of characteristic spectral bands enabled visualization of the spatial variation in lignin and cellulose. Raman images acquired by integrating over the spectral band at 1605 cm-1 suggested that higher lignin content was visualized in the cell corner (CC), the compound middle lamella (CML), and the secondary wall of ray parenchyma. Cellulose distribution followed by taking the band regions around 2897 cm-1 into account showed the opposite pattern, with the highest content in fiber secondary wall. The SEM-EDXA provided semi-quantitative results, showing that the lignin content ratio in various cell wall layers was 1.4 (CC):1.1(CML):1(S2).

An extensin-rich matrix lines the carinal canals in Equisetum ramosissimum, which may function as water-conducting channels.

The anatomy of Equisetum stems is characterized by the occurrence of vallecular and carinal canals. Previous studies on the carinal canals in several Equisetum species suggest that they convey water from one node to another. Cell wall composition and ultrastructure have been studied using immunocytochemistry and electron microscopy, respectively. Serial sectioning and X-ray computed tomography were employed to examine the internode-node-internode transition of Equisetum ramosissimum. The distribution of the LM1 and JIM20 extensin epitopes is restricted to the lining of carinal canals. The monoclonal antibodies JIM5 and LM19 directed against homogalacturonan with a low degree of methyl esterification and the CBM3a probe recognizing crystalline cellulose also bound to this lining. The xyloglucan epitopes recognized by LM15 and CCRC-M1 were only detected in this lining after pectate lyase treatment. The carinal canals, connecting consecutive rings of nodal xylem, are formed by the disruption and dissolution of protoxylem elements during elongation of the internodes. Their inner surface appears smooth compared with that of vallecular canals. The carinal canals in E. ramosissimum have a distinctive lining containing pectic homogalacturonan, cellulose, xyloglucan and extensin. These canals might function as water-conducting channels which would be especially important during the elongation of the internodes when protoxylem is disrupted and the metaxylem is not yet differentiated. How the molecularly distinct lining relates to the proposed water-conducting function of the carinal canals requires further study. Efforts to elucidate the spatial and temporal distribution of cell wall polymers in a taxonomically broad range of plants will probably provide more insight into the structural-functional relationships of individual cell wall components or of specific configurations of cell wall polymers.