Flax Stem Research Articles

Pectinmethylesterases (PME) and pectinmethylesterase inhibitors (PMEI) enriched during phloem fiber development in flax (Linum usitatissimum).

Flax phloem fibers achieve their length by intrusive-diffusive growth, which requires them to penetrate the extracellular matrix of adjacent cells. Fiber elongation therefore involves extensive remodelling of cell walls and middle lamellae, including modifying the degree and pattern of methylesterification of galacturonic acid (GalA) residues of pectin. Pectin methylesterases (PME) are important enzymes for fiber elongation as they mediate the demethylesterification of GalA in muro, in either a block-wise fashion or in a random fashion. Our objective was to identify PMEs and PMEIs that mediate phloem fiber elongation in flax. For this purpose, we measured transcript abundance of candidate genes at nine different stages of stem and fiber development and found sets of genes enriched during fiber elongation and maturation as well as during xylem development. We expressed one of the flax PMEIs in E. coli and demonstrated that it was able to inhibit most of the native PME activity in the upper portion of the flax stem. These results identify key genetic components of the intrusive growth process and define targets for fiber engineering and crop improvement.

Production of efficient enzymes for flax retting by solid state fermentation withAspergillus niger

Purpose– The purpose of this paper is to study the optimum culture condition of flax-retting enzyme, and applying the enzyme liquors to retting. In addition to research the structure of flax stem in enzyme retting using scanning electron microscopy (SEM).Design/methodology/approach– The influence of cultural parameters such as moisture level, temperature, incubation time and inoculum size were evaluated with respect to polygalacturonase and xylanase yield. The structure of flax stem was revealed by SEM.Findings– The flax was retted by dilute enzyme liquors (polygalacturonase/25 U/ml, xylanase/1 U/ml), and 24-h retting period was obtained. The SEM studies revealed the removal of a large amount of non-cellulosic gummy material of flax stem in enzyme retting.Originality/value– First, flax-retting enzyme containing higher-yield polygalacturonase, lower-yield xylanase and no cellulose was yielded by SSF. Second, the composition of enzyme could meet flax retting and no damage cellulosic fibre. Finally, only low-cost wheat bran, citrus peel powder and mineral salt were employed in substrate, which could reduce the overall cost of enzyme production and flax retting.

Microwave assisted retting – A novel method of processing of flax stems

A new method for retting of flax stem was introduced by using microwave energy. The combined effect of pre-soaking, microwave volumetric heating and the non-thermal effect of microwave energy were applied in the retting of flax stem. Effective retting was observed in various combinations of pre-soaking, Microwave treatment times and microwave energy. The samples soaked for 24 and 36 h with microwave treatments of 20 min showed better retting efficiency. Fried tests, Near Infrared analysis and fibre diameter analysis were done to find out the retting efficiency. The effect of non-thermal effect of microwave in flax retting was proved experimentally. Colourimetric tests and tensile strength tests were conducted to compare the quality of fibres after microwave retting and no significant changes in the physical qualities were observed. The water released after pre-soaking was not contaminated because of less microbial activity in short duration of the entire process. In industrial point of view, this method is acceptable because of the ease of recyclability of the pre-soaked water and comparably short duration of the process.

Application of Electro-Technologies in Processing of Flax Fiber

Flax fibers used for various applications are obtained from flax stems. Retting followed by drying and mechanical separation leads to the production of fibers. This review article discusses the application of electro-technologies in the production of bast fibers from the flax stem. In these technologies, flax stems harvested from the field are subjected to microwave assisted retting, followed by electro–osmotic dewatering which reduces the water content of the stems. Dewatered stems are transferred to a microwave chamber for further drying, thus retted stems are obtained for further processing.

Identification of cell wall proteins in the flax (Linum usitatissimum ) stem

Sequential salt (CaCl2 , LiCl) extractions were used to obtain fractions enriched in cell wall proteins (CWPs) from the stem of 60-day-old flax (Linum usitatissimum) plants. High-resolution FT-ICR MS analysis and the use of recently published genomic data allowed the identification of 11912 peptides corresponding to a total of 1418 different proteins. Subcellular localization using TargetP, Predotar, and WoLF PSORT led to the identification of 152 putative flax CWPs that were classified into nine different functional classes previously established for Arabidopsis thaliana. Examination of different functional classes revealed the presence of a number of proteins known to be involved in, or potentially involved in cell-wall metabolism in plants. The flax stem cell wall proteome was also compared with transcriptomic data previously obtained on comparable samples. This study represents a major contribution to the identification of CWPs in flax and will lead to a better understanding of cell wall biology in this species.

Effect of microwave and hot air drying on flax straw at controlled temperatures

Flax stems were subjected to microwave drying at controlled temperatures. The rate of drying was then compared with conventional hot air drying. The product temperature was maintained at 40°C, 60°C and 80°C for both microwave and hot air drying. The moisture content of flax stem was about 70% (wet basis). The microwave drying was conducted in a microwave apparatus which recorded mass, product temperature, incident microwave power, reflected microwave power and inlet/outlet air temperature. The final moisture content for experiment was set to 9% (wet basis). Microwave-convective drying ensured about 30% to 70% reduction of drying time for drying flax straw as compared to hot air drying. Curve fitting with different mathematical models were carried out and all the models were fitted well for both hot air and microwave drying. The tensile strength of flax straw, measured with an Instron apparatus, increased with an increase in the processing temperature of both processes. Hot air dried flax straw showed the greatest tensile strength and modulus of elasticity at processing temperatures of 60°C and 80°C with a significant difference.

Transfer of Near-Infrared Calibration Model for Determining Fiber Content in Flax: Effects of Transfer Samples and Standardization Procedure

The transfer of a calibration model for determining fiber content in flax stem was accomplished between two near-infrared spectrometers, which are the same brand but which require a standardization. In this paper, three factors, including transfer sample set, spectral type, and standardization method, were investigated to obtain the best standardization result. Twelve standardization files were produced from two sets of the transfer sample (sealed reference standards and a subset of the prediction set), two types of the transfer sample spectra (raw and preprocessed spectra), and three standardization methods (direct standardization (DS), piecewise direct standardization (PDS), and double window piecewise direct standardization (DWPDS)). The efficacy of the model transfer was evaluated based on the root mean square error of prediction, calculated using the independent prediction samples. Results indicated that the standardization using the sealed reference standards was unacceptable, but the standardization using the prediction subset was adequate. The use of the preprocessed spectra of the transfer samples led to the calibration transfers that were successful, especially for the PDS and the DWPDS correction. Finally, standardization using the prediction subset and their preprocessed spectra with DWPDS correction proved to be the best method for transferring the model.

Flax shive as a source of activated carbon for metals remediation

Flax shive constitutes about 70% of the flax stem and has limited use. Because shive is a lignocellulosic by-product, it can potentially be pyrolyzed and activated to produce an activated carbon. The objective of this study was to create an activated carbon from flax shive by chemical activation in order to achieve significant binding of selected divalent cations (cadmium, calcium, copper, magnesium, nickel, zinc). Shive carbons activated by exposure to phosphoric acid and com-pressed air showed greater binding of cadmium, copper, nickel or zinc than a sulfuric acid-activated flax shive carbon reported in the literature and a commercial, wood-based carbon. Uptake of calcium from a drinking water sample by the shive carbon was similar to commercial drinking water filters that contained cation exchange resins. Magnesium removal by the shive carbon was greater than a commercial drinking water filtration carbon but less than for filters containing cation exchange resins. The results indicate that chemically activated flax shive carbon shows considerable promise as a component in industrial and residential water filtration systems for removal of divalent cations.

Engineering of PHB Synthesis Causes Improved Elastic Properties of Flax Fibers

Flax stem is a source of fiber used by the textile industry. Flax fibers are separated from other parts of stems in the process called retting and are probably the first plant fibers used by man for textile purposes (1). Nowadays flax cultivation is often limited because of its lower elastic property compared to cotton fibers. Thus the goal of this study was to increase the flax fiber quality using a transgenic approach. Expression of three bacterial genes coding for beta-ketothiolase (phb A), acetoacetyl-CoA reductase (phb B), and PHB synthase (phb C) resulted in poly-beta-hydroxybutyrate (PHB) accumulation in the plant stem. PHB is known as a biodegradable thermoplastic displaying chemical and physical properties similar to those of conventional plastics (i.e., polypropylene). The fibers isolated from transgenic flax plants cultivated in the field and synthesizing PHB were then studied for biomechanical properties. All measured parameters, strength, Young's modulus, and energy for failure of flax fibers, were significantly increased. Thus the substantial improvement in elastic properties of fibers from the transgenic line has been achieved. Since the acetyl CoA, substrate for PHB synthesis, is involved not only for energy production but also for synthesis of many cellular constituents, the goal of this study was also the analysis of those metabolites, which interfere with plant physiology and thus fiber quality. The analyzed plants showed that reduction in lignin, pectin, and hemicellulose levels resulted in increased retting efficiency. A significant increase in phenolic acids was also detected, and this was the reason for improved plant resistance to pathogen infection. However, a slight decrease in crop production was detected.

MALDI-TOF MS evidence for the linking of flax bast fibre galactan to rhamnogalacturonan backbone

Fibre-specific (1 → 4)-β-galactan extracted from bast fibre peels of developing flax ( Linum usitatissimum L.) stem has been studied to elucidate its structural details. The polysaccharide was characterized by NMR and subjected to partial degradation protocols, including chemical and enzymatic approaches. The oligosaccharide fragments obtained were fractionated by gel permeation chromatography and analyzed for their molecular mass with MALDI-TOF MS. The obtained data show that this flax galactan is a complex RG-I polysaccharide with variable side chain structures. The backbone is composed of the common GalA-Rha repeats with a high degree of branching. These side chains are mainly composed of β-1,4-linked Gal oligomers: (1) short branches of only one or two Gal residue(s); (2) long (linear) branches of up to 26 Gal residues; (3) mixed branches of between 3 and 12 Gal residues (possibly derived from longer linear side chains), that are resistant to galactanase cleavage; (4) side chains of at least 17 Gal residues, decorated with single Ara moieties. The linkage between RG backbone and galactan side chains was confirmed by the presence of fragments with (Rha-GalA) n Hex m structure type. Neither chemical, nor enzymatic hydrolysis yielded oligomeric GalA residues, indicating that RG-I blocks are not interrupted by HGA regions. The polymer can be cleaved only partially by the rhamnogalacturonan hydrolase used, while the remaining part is resistant, probably due to peculiarities of side chain structure. Novel Rha-GalA oligomers were liberated by RG-hydrolase containing two or three Gal attached to Rha near the cleavage site. The native polymer is decorated by acetyl groups, with yet unknown distribution patterns. Treatment with purified and well-characterized galactanase does not change the hydrodynamic volume of flax galactan (despite considerable cleavage of Gal moieties), suggesting a complex “secondary” structure of the polymer.

How Does Light Intensity Affect the Elementary Fiber Length in Flax?

There are not enough antecedents about the effect of a decrease in photosynthesis on flax stem growth and elementary fiber length. Therefore, plots sowed with cultivar Diane were shaded with nets that intercepted 70% of photosynthetically active radiation. The nets were placed at three stages of the growth period: S1: 575°Cd (height: 25 cm), S2: 854°Cd (height: 54 cm), and S3: 1304°Cd (height: 84 cm). These applications resulted in stems shadowed only in the parts, which grew during the treatment. Additionally, control plots without shadows during the entire cycle (S0) were grown. In every treatment the production of fiber was quantified, and a harvest index (HI = fiber biomass obtained/stems biomass produced) was calculated. Fiber quality was also evaluated using elementary fiber length. The harvest index was bigger in control plants than shadow plants. The length of elementary fiber of the shadow treatments S1, S2, and S3 was 4.28 cm, 3.76 cm, and 3.02 cm, respectively, whereas the adjacent portions to the shaded one in each treatment had a length fiber increase of 21%, 34%, and 37%, respectively. In the same portions, S0 had fiber lengths of 4.94, 4.45, and 3.82 cm, respectively. The trend observed in all the shaded treatments showed a decrease of elementary fiber length. The fiber length in the adjacent portions to the shaded ones was not conditioned by the shadow treatment received. These results allow relating fiber quality with stresses generated by meteorological events that take place during the growth of the stem.

Lignification in the flax stem: evidence for an unusual lignin in bast fibers

In the context of our research on cell wall formation and maturation in flax (Linum usitatissimum L) bast fibers, we (1) confirmed the presence of lignin in bast fibers and (2) quantified and characterized the chemical nature of this lignin at two developmental stages. Histochemical methods (Weisner and Maüle reagents and KMnO(4)-staining) indicating the presence of lignin in bast fibers at the light and electron microscope levels were confirmed by chemical analyses (acetyl bromide). In general, the lignin content in flax bast fibers varied between 1.5% and 4.2% of the dry cell wall residues (CWRs) as compared to values varying between 23.7% and 31.4% in flax xylem tissues. Immunological and chemical analyses (thioacidolysis and nitrobenzene oxidation) indicated that both flax xylem- and bast fiber-lignins were rich in guaiacyl (G) units with S/G values inferior to 0.5. In bast fibers, the highly sensitive immunological probes allowed the detection of condensed guaiacyl-type (G) lignins in the middle lamella, cell wall junctions, and in the S1 layer of the secondary wall. In addition, lower quantities of mixed guaiacyl-syringyl (GS) lignins could be detected throughout the secondary cell wall. Chemical analyses suggested that flax bast-fiber lignin is more condensed than the corresponding xylem lignin. In addition, H units represented up to 25% of the monomers released from bast-fiber lignin as opposed to a value of 1% for the corresponding xylem tissue. Such an observation indicates that the structure of flax bast-fiber lignin is significantly different from that of the more typical 'woody plant lignin', thereby suggesting that flax bast fibers represent an interesting system for studying an unusual lignification process.

Occurrence of cell-specific galactan is coinciding with bast fiber developmental transition in flax

Bast fiber peels of developing flax ( Linum usitatissimum L.) stem contain tissue-specific soluble (1→4)-β-galactan. The occurrence and localization of this polymer have been investigated at various stages of bast fiber development. Tissue-specific galactan (molecular mass >2000 kDa) emerged right below the snap point—the manually identified spot on the stem, above which the fiber cells elongate, while below no more elongation occurs, but intensive secondary cell wall thickening takes place. In the course of plant development the amount of tissue-specific galactan gradually declined and in mature plants the polymer disappeared from the fraction, being either degraded or fixed within the cell wall. The immunolocalization of (1→4)-β-galactans with LM5 antibody revealed that in flax stem these polymers were present only in bast fiber cells. Two layers were well pronounced within the secondary cell wall, the density of gold particles in the inner layer was 5.2±1.5 times higher than in the outer layer. To distinguish soluble galactan from other (1→4)-β-galactans, the stem sections were treated with buffer, which elutes buffer-soluble galactan. The soluble high molecular mass (1→4)-β-galactan is a bast fiber cell-specific polymer, emerging at the transition from cell elongation to cell wall thickening. It is localized in the inner portion of the secondary cell wall and is modified or degraded in the course of further plant development. This polymer belongs to the rarely observed plant cell wall polysaccharides, which only occurs at one specific stage of cell development, suggesting that it could play a prominent role in cellulose deposition and secondary cell wall formation in flax fibers.

The snap point: a transition point in Linum usitatissimum bast fiber development

The developing stem of fibre flax ( Linum usitatissimum L.) contains a specific region called the ‘snap point’, where the fiber-enriched bast tissues considerably change their mechanical properties. The snap point was found to be present during a restricted period of plant development—the fast growth phase, and to disappear when stem growth was completed. To relate this snap point to bast fiber formation stages, the number of bast fiber cells and the thickness of their cell walls were followed on the stained cross-sections of the flax stem throughout plant development, using the progressing snap point as the reference. The snap point was shown to be the spot, above which the elongation of bast fiber cells is fully completed. This fast growth stage is the period when the maximum length of all bast fibers in the mature plant (a major characteristic of flax fiber quality) is fixed and would not be changed later. Autoradiography was used to visualize the mode of flax bast fiber elongation above the snap point. The even distribution of label was indicative for surface (diffusive) growth type. Elongation of individual fiber cells was estimated to take only 2–4 days with a rate of 1–2 cm per day, while cell wall thickening occurs mainly below snap point and lasts around 2 months. The special cell wall structural order, characteristic for mature bast fibers, first appeared at the snap point in the outer layer of the secondary cell wall. Schemes are included, illustrating the course of cell wall thickening and the localization of various stages of fiber formation on the stem of growing flax plant. The established exact localization and duration of flax bast fiber formation stages, and the existence of snap point as the manually identified morphological reference for the transition, permit to separate the bast fibers at different stages of development and make flax an attractive model system to study the functional genomics of fiber formation in technical crops.

The use of FT‐IR microspectroscopic mapping to study the effects of enzymatic retting of flax (Linum usitatissimum L) stems

AbstractFourier transform infrared (FT‐IR) microspectroscopic mapping was investigated as a tool to study the effects of enzymatic retting of flax stems. The FT‐IR technique permitted the elucidation of the relative loss or changes in the distribution of key chemical components after treatment with enzymes or enzyme/chelator mixtures in association with visible changes in structure. Cross‐sections of Ariane flax stems were treated with SP 249 (a pectinase‐rich enzyme mixture from Novo Nordisk) at 0.5, 0.7 or 1.0 ml l−1 concentration in pH 5 acetate buffer for 6 h at 40 °C. Flax stems treated with 0.5 or 0.7 ml l−1 SP 249 and 50 mM oxalic acid as a chelator were also investigated by the technique. The results indicated that treatment with 0.5 ml l−1 SP 249 alone was ineffective in releasing the fibre bundles from the surrounding tissue, but the release was increased by the addition of 50 mM oxalic acid as a likely chelator for the cations of pectate salts. However, the IR spectra of the bundles indicated that an insoluble oxalate salt remained on the tissue after this treatment. Increasing the concentration of SP 249 to 0.7 ml l−1 plus 50 mM oxalic acid was effective in releasing the fibre bundles and generating some ultimate fibres with no detectable oxalate expectate salt residues. Increasing the SP 249 concentration to 1.0 ml l−1 without using oxalic acid was effective in separating the fibre bundles into ultimate (individual) fibres, leaving no pectate salt residue and only a trace of pectic esters and/or acids. The use of infrared mapping, or so‐called chemical imaging, is shown to have advantages over visible imaging alone in that it can detect and locate the chemical species present after each treatment in relation to the anatomical features of the flax stem. This analytical tool shows promise as a technique by which to study the effects of enzymatic treatment of natural fibre materials.Published in 2002 for SCI by John Wiley & Sons, Ltd

Plant factors influencing enzyme retting of fiber and seed flax.

Retting, which is the microbial activity through which bast fibers are released from nonfiber tissues, is the limiting factor in flax processing. The objective of this work is to identify chemical and structural characteristics in a variety of fiber and seed flax types that influence enzyme retting in a recently developed method. Analyses of flax retted in a series of tests, including two enzyme rettings in some cases, indicated that lignin did not limit the separation of fibers from shive and showed that pectinases in enzyme-retting mixtures could ret fiber and seed flax. However, mature stems, such as that in flax produced for seed, had greater amounts of cutin and wax in the cleaned fiber product, suggesting that the cuticle could be a greater antiquality factor in seed versus fiber flax. With seed flax, the fraction of finer fibers produced during retting was significantly lower than with fiber flax. Results indicated that enzyme retting could be used to obtain flax fibers from seed flax stem residues and add value to this agricultural material.

Caffeoyl-coenzyme A 3-O-methyltransferase enzyme activity, protein and transcript accumulation in flax (Linum usitatissimum) stem during development.

Flax (Linum usitatissimum) is a commercially important fiber crop in Europe. Lignification of its phloem fibers, although weak, causes a decrease in their commercial quality. In flax, fiber lignin mainly consists of guaiacyl (G) units in contrast to the mixed guaiacyl-syringyl (G-S) lignin type occurring in xylem fibers. G lignins are reported as more condensed polymers due to a higher frequency of 5-5 linkages, whereas the deposition of syringyl end groups in lignins increases the proportion of alky-aryl ether linkages as beta-O-4-bonds. The type of linkages within a lignin polymer depends on the methylation of either the 3-hydroxyl groups or both 3-OH and 5-OH groups, which is controlled by two enzymes: caffeate 3-O-methyltransferase (COMT) and caffeoyl-coenzyme A 3-O-methyltransferase (CCoAOMT). First, we measured the in vitro activity of both OMTs in the flax stem tissues during stem development. CCoAOMT activity varied in the same way as COMT, i.e. increased gradually with stem maturity, from the top to the bottom of the stem, was maximum at the flowering stage and was lower, but still scorable, in the outer fiber-bearing tissues than in the xylem cells. In a second step, we focused our studies on the characterization of CCoAOMT in order to understand the implication of this enzyme in the lignification of flax fiber cells. CCoAOMT activity appeared to be associated with the accumulation of an acidic 33-kDa polypeptide identified as a CCoAOMT after immunological cross-reactivity with a poplar CCoAOMT and microsequencing. The differential accumulation of the CCoAOMT protein was confirmed by immunolocalization on tissue prints and correlated with that of the transcripts, suggesting a transcriptional regulation of CCoAOMT in the flax stem.

In vitro evaluation of dew‐retting of flax by fungi from southern Europe

SummaryFlax dew‐retting is widely adopted in most flax‐growing countries, but it does not represent a practical solution where dry weather conditions occur after harvest. A study of the local microbiological aspects was undertaken as a contribution to improve field‐retting of flax under southern European climates. Fungi were isolated from soil and dew‐retted flax in northern Italy, and 23 representative strains were chosen to test their ability to ret flax stems. Experiments were performed in vitro on flax stem pieces artificially inoculated with single fungal strains. Retting degree was assessed with a mechanical test, to evaluate the ease with which the bast was detached from the wood core, and by the analysis of the residual fibre pectins using uronic acid. Uronic acid dosage provided a better differentiation of the strains than the mechanical test. There was a large variability in retting ability among the species assayed and even among strains of the same species. The best results were obtained with all Aspergillus and Penicillium strains, while Mucor and Rhizopus strains showed a variable retting ability. Fusarium, Trichoderma strains and Epicoccum nigrum had the poorest retting abilities among all the fungal strains assayed.

Composition and Distribution of Cell Wall Phenolic Compounds in Flax (Linum usitatissimum L.) Stem Tissues

Cell wall phenolic compounds were analysed in xylem and bast fibre-rich peels of flax stems by biochemical, histochemical and ultrastructural approaches. Localization of cell wall phenolics by the enzyme-gold method using laccase revealed several gold particle distribution patterns. One of the major types (an even distribution of single gold particles) was present mainly in xylem, while the other (compact branched groups of ten–40 gold particles) was found both in xylem and fibre cells. The lignin content of the stem parts was estimated by the Klason procedure and by the thioglycolic acid assay, and the phenolic products recovered after alkaline cupric oxide oxidation of cell walls were analysed by GC. By combining chemical analysis data and the frequency of various gold particle types within the tissues, different patterns of gold particle distribution could be ascribed to certain cell wall phenolics; lignin was stained as evenly distributed single gold particles, while branched clusters represented hydroxycinnamic acids. The Klason procedure did not remove all the non-lignin components from flax fibres, known for their highly crystalline cellulose, and considerably overestimated the lignin content. The thioglycolic acid assay results were consistent with GC and microscopic observations.

Near-infrared–Fourier-transform–Raman microspectroscopic imaging of flax stems

Near-infrared–Fourier-transform–Raman (NIR–FT–Raman) microscopy was employed to image and subsequently produce in situ maps of the distribution of chemical components in flax ( Linum usitatissimum L.) stem tissue. Thick (∼80 μm) cross-sections of flax were cryotomed, thawed, equilibrated at 85% RH for >3 h and sealed under cover glasses on gold-mirrored microscope slides. Spectra for each image pixel were collected from selected tissue sections over the Raman shifted region 3600–300 cm −1 at 16 cm −1 with 256–512 scans and employing ∼185 mW of focused laser power at the sample. A computer controlled automated microscope stage was used to collect area maps of 50–150 μm in rectangular dimensions in 6–10 μm steps and register them to the visible images that were captured using a CCD camera. Data collection required 8–10 h per image. Chemical profiles were produced from area integrations of specific spectral regions. The chemical profiles showed the location of all of the major components of flax by anatomical cell type. A sharp shoulder at 2850 cm −1 in the CH stretch region provided evidence of waxes in the cuticular/epidermal tissue. The bands occurring around 1600 cm −1, due to aromatic ring stretching vibrations, gave evidence of lignin in core tissue and pigments in epidermal tissue. Bands occurring between 1175–1050 cm −1, due to COC heavy atom mixed mode vibrations, showed the greatest concentration of carbohydrate in fiber cells with lesser (but significant) amounts indicated in core tissue. In addition, bands occurring between 870–800, 515–476 and 400–360 cm −1 gave specific evidence for the presence of pectins, other non-cellulosic polysaccharides and cellulose, respectively, in parenchyma tissue.