Comparison of cell wall compositions of a desert xerophyte and a related mesophyte

A comparison was made of the cell wall compositions of stem internode tissues from two members of the Chenopodiaceae. Cell walls from Anabasis syriaca (a desert xerophyte) contained non-cellulosic polysaccharides rich in arabinose, xylose and galacturonic acid. The non-cellulosic polysaccharides from cell walls of Spinacia oleracea (a mesophyte) were rich in glucose. Anabasis syriaca cell walls contained relatively more cellulose and lignin than those of Spinacia oleracea.     

Quantitative and qualitative changes in cell wall polysaccharides in relation to growth and cell wall loosening in Lactuca sativa hypocotyls

The correlation between hypocotyl elongation, cell wall loosening and changes in cell wall polysaccharides was studied using intact lettuce seedlings grown in the dark or in light together with gibberellic acid (GA) and/or 5-fluorodeoxyuridine (FUDR). The following results were obtained:
1) The production of pectic, hemicellulosic and cellulosic polysaccharides look place in parallel with hypocotyl elongation, which was substantially affected by different growth conditions.
2) The mole percentage sugar composition of pectic and hemicellulosic polysaccharides changed in response to dark, light, GA, or FUDR treatments.
3) The amounts of xylose and glucose in hemicellulosic polysaccharides and those of galactosc, rhumnose and uronic acid in pectic polysaccharides increased in parallel with hypocotyl elongation.
4) Statistical analysis of the quantitative relationship between sugars composing polysaccharides revealed that the uronic acid content changed in parallel with those of rhamnose and galactose in pectic polysaccharides, and the content of xylose varied in parallel with those of fucose and glucose.
5) The content of hemicellulosic polysaccharides was correlated with cell wall loosening represented by a decrease in the minimum stress-relaxation time. Changes in the stress-relaxation time value were correlated with those in the content of araltinose and galactose in hemicellulosic polysaccharides.
Based on these results, the relationship between hypocotyl elongation, changes in cell wall polysaccharides, and cell wall loosening is discussed with respect to the effect of GA and FUDR on hypocotyl elongation.     

Aluminum-Induced Rapid Root Inhibition and Changes in Cell-Wall Components of Squash Seedlings

Growth of squash (Cucurbita maxima Duch.) roots was significantly inhibited by 1 mM AlCl3 as early as 1 h after the treatment. The growth inhibition was confined to the elongating zone (1-6 mm from the root tip). Chemical analysis of cell-wall polysaccharides from roots revealed that aluminum increased pectin, hemi-cellulose, and cellulose contents after 3 h of treatment. The effect of aluminum on pectin content was found in the elongating zone including the root tip, whereas change in cellulose content was confined to only nonelongating zones. Hemicellulose content increased in all of the regions along the root axis. The increase in the pectin fraction was due to the increases in uronic acids, galactose, and arabinose constituents, whereas hemicellulose content changed due to increases in glucose, xylose, galactose, and arabinose. The results clearly indicate that aluminum rapidly reduced squash root growth by inhibiting cell elongation and altering metabolism of cell-wall polysaccharides in the nonelongating zone as well as in the elongating zone.     

Changes in the composition of cotton fibre cell walls during development

Purified cell walls, prepared from cotton fibres (Gossypium arboreum L.) at different growth stages, were subjected to successive extractions to give pectic, hemicellulosic, and -cellulosic fractions. The protein content and sugars obtained after hydrolysis of the total cell walls and of the various fractions were quantitatively estimated. The amount of protein in the fibre cell walls from one ovule reached a maximum value at the end of the elongation growth, decreased, and then reached a second maximum at the end of the secondary wall deposition. The absolute amounts of fucose, galactose, mannose, rhamnose, arabinose, uronic acid, and non-cellulosic glucose residues all reached a maximum at the end of the primary wall formation or at the beginning of the secondary wall formation. Only the absolute amounts of xylose and of the cellulosic glucose residues increased until the end of the fibre development. Most conspicuous was the decrease in the absolute amounts of non-cellulosic glucose and of arabinose residues during the secondary wall formation, possibly indicating a turnover of at least some of the hemicellulosic wall material.Abbreviations DPA days post anthesis - TLC thin layer chromatography - SDS sodium dodecyl sulphate     

Changes in cell wall neutral sugar composition during fruit ripening: a species survey

Non-cellulosic neutral sugar composition of cell walls from seventeen fruit types were analysed during ripening. Galactose was the major non-cellulosic neutral sugar in cell walls of cucurbit and solanaceous fruit, xylose was the predominant non-cellulosic neutral component of berries, and arabinose was the major non-cellulosic component of pome fruits. The major non-cellulosic neutral sugar residue in cell walls of stone fruits varied. In nectarine and peach, plum, and apricot, the major sugar was arabinose, galactose, and xylose, respectively. In 15 of the 17 types of fruit, a net loss of non-cellulosic neutral sugar residues occurred during ripening. No net loss occurred in plums and cucumbers. A net loss of cell wall galactose and/or arabinose occurred in 14 of the types of fruit. Xylose was the major neutral sugar residue lost from walls of apricot during ripening. In general, berry cell walls were comparatively low in galactose and arabinose content.     

Rumen digestion of rice straw structural polysaccharides: effect of ammonia treatment and lucerne extract supplementation in vitro

The combined effects of lucerne (Medicago sativa L.) extract supplementation and ammonia treatment of rice straw (Oryza sativa, variety Thaibonnet) on the ruminal digestion of cell wall components were investigated in six continuous culture systems using a randomised complete block design. Data were fitted to second-order polynomial models. Untreated rice straw had higher contents of ash-free cell wall residues (CWR; 763 v. 687 g/kg dry matter (DM)) and non-cellulosic sugars (191 v. 166 g/kg DM) than treated rice straw. Ammoniation preferentially removed xylose, which resulted in a lower xylose-to-arabinose ratio (5.1 v. 5.8). In absence of lucerne supplementation and ammoniation, degradability coefficients were 0.54, 0.46, 0.58, 0.54, 0.42 and 0.60 for cellulose–glucose, xylose, arabinose, galactose, mannose and uronic acids, respectively. Both factors had significant effects on the microbial degradation of structural polysaccharides. With lucerne extract at an optimal level, ammonia treatment increased ash-free cell wall degradation by more than 10%. The degradability coefficients were increased by ammoniation without any significant interaction with lucerne extract, except for glucose, whose degradability was mostly influenced by lucerne extract in a curvilinear way. The comparison of regression coefficients in cell wall and CWR models suggested that ammoniation improved the degradabilities of xylose, galactose and mannose by partly solubilising the corresponding hemicelluloses and by improving the susceptibility of the remaining fraction to microbial attack, whereas it increased the degradability of arabinose only by favouring microbial attack.     

Comparison between maize root cells and their respective regenerating protoplasts: wall polysaccharides

The cell-wall polysaccharides from different parts of maize roots have been analysed. The arabinose, galactose and mannose contents are influenced by cell differentiation, whereas xylose, rhamnose and uronic-acid contents are not. In cap cells, the pectin content is low but rhamnose and fucose are present in larger quantities. The cell-wall polysaccharides from cells of the elongation zone and their respective regenerating protoplasts were also analysed. The walls of the protoplasts contained higher xylose and mannose levels and a much lower level of cellulose than the cells from which they were derived.     

Glucuronoarabinoxylan structure in the walls of Aechmea leaf chlorenchyma cells is related to wall strength

In CAM-plants rising levels of malic acid in the early morning cause elevated turgor pressures in leaf chlorenchyma cells. Under specific conditions this process is lethal for sensitive plants resulting in chlorenchyma cell burst while other species can cope with these high pressures and do not show cell burst under comparable conditions. The non-cellulosic polysaccharide composition of chlorenchyma cell walls was investigated and compared in three cultivars of Aechmea with high sensitivity for chlorenchyma cell burst and three cultivars with low sensitivity. Chlorenchyma layers were cut from the leaf and the non-cellulosic carbohydrate fraction of the cell wall fraction was analyzed by gas-liquid chromatography. Glucuronoarabinoxylans (GAXs) were the major non-cellulosic polysaccharides in Aechmea. The fine structure of these GAXs was strongly related to chlorenchyma wall strength. Chlorenchyma cell walls from cultivars with low sensitivity to cell burst were characterized by an A/X ratio of ca. 0.13 while those from cultivars with high sensitivity showed an A/X ratio of ca. 0.23. Xylose chains from cultivars with high cell burst sensitivity were ca. 40% more substituted with arabinose compared to cultivars with low sensitivity for cell burst. The results indicate a relationship in vivo between glucuronoarabinoxylan fine structure and chlorenchyma cell wall strength in Aechmea. The evidence obtained supports the hypothesis that GAXs with low degrees of substitution cross-link cellulose microfibrils, while GAXs with high degrees of substitution do not. A lower degree of arabinose substitution on the xylose backbone implies stronger cell walls and the possibility of withstanding higher internal turgor pressures without cell bursting.     

Soluble Cell Wall Polysaccharides Released from Pea Stems by Centrifugation : II. EFFECT OF ETHYLENE

The effect of ethylene on cell wall metabolism in sections excised from etiolated pea stems was studied. Ethylene causes an inhibition of elongation and a pronounced radial expansion of pea internodes as shown by an increase in the fresh weight of excised, 1-cm sections. Cell wall metabolism was studied using centrifugation to remove the cell wall solution from sections. The principal neutral sugars in the cell wall solution extracted with H₂O are arabinose, xylose, galactose, and glucose. Both xylose and glucose decline relative to controls in air within 1 hour of exposure to ethylene. Arabinose and galactose levels are not altered by ethylene until 8 hours of treatment, whereupon they decline in controls in air relative to ethylene treatment. When alcohol-insoluble polymers are fractionated into neutral and acidic polysaccharides, xylose and glucose predominate in the neutral fraction and arabinose and galactose in the acidic fraction. Ethylene depresses the levels of xylose and glucose in the neutral fraction and elevates arabinose and galactose in the acidic fraction. Ethylene treatment does not affect the level of uronic acids extracted with H₂O; however, the level of hydroxyproline-rich proteins in this water-extracted cell wall solution is increased by ethylene. Extraction of sections with CaCl₂ results in an increase in the levels of neutral sugars particularly arabinose. Ethylene depresses the yield of arabinose in calcium-extracted solution relative to controls in air. Similarly, extraction with CaCl₂ increases the yield of extracted hydroxyproline in ethanol-insoluble polymers and ethylene depresses its level relative to controls. Metabolism of uronic acids and neutral sugars and growth in response to ethylene treatment contrast markedly with auxin-induced polysaccharide metabolism and growth. With auxin, sections increase mostly in length not radius, and this growth form is associated with an increase in the levels of xylose, glucose, and uronic acids. With ethylene, on the other hand, stem elongation is suppressed and expansion is promoted, and this growth pattern is associated with a decrease in xylose and glucose in the ethanol-insoluble polysaccharides.     

Arabidopsis dynamin-like protein DRP1A: a null mutant with widespread defects in endocytosis, cellulose synthesis, cytokinesis, and cell expansion

Dynamin-related proteins are large GTPases that deform and cause fission of membranes. The DRP1 family of Arabidopsis thaliana has five members of which DRP1A, DRP1C, and DRP1E are widely expressed. Likely functions of DRP1A were identified by studying rsw9, a null mutant of the Columbia ecotype that grows continuously but with altered morphology. Mutant roots and hypocotyls are short and swollen, features plausibly originating in their cellulose-deficient walls. The reduction in cellulose is specific since non-cellulosic polysaccharides in rsw9 have more arabinose, xylose, and galactose than those in wild type. Cell plates in rsw9 roots lack DRP1A but still retain DRP1E. Abnormally placed and often incomplete cell walls are preceded by abnormally curved cell plates. Notwithstanding these division abnormalities, roots and stems add new cells at wild-type rates and organ elongation slows because rsw9 cells do not grow as long as wild-type cells. Absence of DRP1A reduces endocytotic uptake of FM4-64 into the cytoplasm of root cells and the hypersensitivity of elongation and radial swelling in rsw9 to the trafficking inhibitor monensin suggests that impaired endocytosis may contribute to the development of shorter fatter roots, probably by reducing cellulose synthesis.     

Galactose inhibition of auxin-induced cell elongation in oat coleoptile segments

Auxin-induced cell elongation in oat coleoptile segments was inhibited by galactose; removal of galactose restored growth. Galactose did not appear to affect the following factors which modify cell elongation: auxin uptake, auxin metabolism, osmotic concentration of cell sap, uptake of tritium-labeled water, auxin-induced wall loosening as measured by a decrease in the minimum stress-relaxation time and auxininduced glucan degradation. Galactose markedly prevented incorporation of [¹⁴C]-glucose into cellulosic and non-cellulosic fractions of the cell wall. It was concluded that galactose inhibited auxin-induced long-term elongation of oat coleoptile segments by interfering with cell wall synthesis.     

Cell wall composition of leaves of an inherently fast- and an inherently slow-growing grass species

Differences in the relative growth rules of the inherently slow-growing Deschampsia flexuosa L. and the inherently fast-growing Holcus lanatus L. were reflected in cell wall synthesis in the elongation zone of the leaves. Leaf elongation rates depended on the size of the plant and ranged from 6 to 14 mm d^?1 in Deschampsia and from 12 to 42 mm d^?1 in Holcus. Anatomical data showed that the epidermis and vascular tissue are the important tissues controlling leaf extension. The cell wall polysaccharides of fully expanded leaves of the two species were identical in sugar composition. Enzymatic hydrolysis of polymeric sugars in the cell walls of the sheath and the lamina gave glucose (85%), arabinose (3.5%), fucose (0.5%), xylose (5.0%), mannose (0.5%), galaclose (0.8%) and galacturonic acid (3–4%). This composition applied throughout the blade and the sheath and did not change with ageing. Polysaccharides in the meristems of the two species showed identical sugar compositions with 51–55% glucose, 13–15% galactoronic acid and 13–14% arabinose as the main components. The extension zone was marked by a gradual increase of driselase-digestable polymers (per mm tissue) and a concurrent shift in sugar composition. The massive increase of glucose in the cell wall polymers of the elongation zone is probably caused by cellulose synthesis. The rate of synthesis of cell wall polysaccharides in Holcus was twice as high as that in Deschampsia. The slower-growing Deschampsia has more ferulic acid esterified with cell walls, which might contribute to the slowing of leaf growth. Lignin is not significantly deposited until growth has essentially ceased and is not responsible for the difference in growth rate.     

Growth and cell wall properties of two wheat cultivars differing in their sensitivity to aluminum stress

The present study was conducted to investigate the cell wall properties in two wheat (Triticum aestivum L.) cultivars differing in their sensitivity to Al stress. Seedlings of Al-resistant, Inia66 and Al-sensitive, Kalyansona cultivars were grown in complete nutrient solutions for 4 days and then subjected to treatment solutions containing Al (0, 50 microM) in a 0.5 mM CaCl(2) solution at pH 4.5 for 24 h. Root elongation was inhibited greatly by the Al treatment in the Al-sensitive cultivar compared to the Al-resistant cultivar. The Al-resistant cultivar accumulated less amount of Al in the root apex than in the Al-sensitive cultivar. The contents of pectin and hemicellulose in roots were increased with Al stress, and this increase was more conspicuous in the Al-sensitive cultivar. The molecular mass of hemicellulosic polysaccharides was increased by the Al treatment in the Al-sensitive cultivar. The increase in the content of hemicellulose was attributed to increase in the contents of glucose, arabinose and xylose in neutral sugars. Aluminum treatment increased the contents of ferulic acid and p-coumaric acid especially in the Al-sensitive cultivar by increasing the activity of phenylalanine ammonia lyase (PAL, EC 4.3.1.5). Aluminum treatment markedly decreased the beta-glucanase activity in the Al-sensitive cultivar, but did not exert any effect in the Al-resistant cultivar. These results suggest that the modulation of the activity of beta-glucanase with Al stress may be involved in part in the alteration of the molecular mass of hemicellulosic polysaccharides in the Al-sensitive cultivar. The increase in the molecular mass of hemicellulosic polysaccharides and ferulic acid synthesis in the Al-sensitive cultivar with Al stress may induce the mechanical rigidity of the cell wall and inhibit the elongation of wheat roots.     

Hypocotyl growth of Pinus pinaster seedlings. Changes in osmotic potential and cell wall composition

The changes in osmotic potential and cell wall composition of hypocotyl cell walls from different hypocotyl regions were investigated during growth of etiolated seedlings of Pinus pinaster Aiton. The osmotic potential in the subapical 5 mm part was minimum when hypocotyl growth rate was low, and increased when the fast growth phase began. The main non-cellulosic sugars of the cell wall from pine hypocotyl were arabinose, galactose, xylose, glucose and uronic acids, although their relative proportions were different from those found for angiosperm cell walls. Non-cellulosic glucose was the sugar showing the most important changes during hypocotyl growth as well as along the hypocotyl, suggesting that a glucose-rich polysaccharide is involved in a very active turnover during growth. A partial degradation of a xyloglucan during growth is suggested.     

Oxaziclomefone, a new herbicide, inhibits wall expansion in maize cell-cultures without affecting polysaccharide biosynthesis, xyloglucan transglycosylation, peroxidase action or apoplastic ascorbate oxidation

BACKGROUND AND AIMS: Oxaziclomefone (OAC), a new herbicide, inhibits cell expansion, especially in roots and cell-cultures of gramineous monocots. OAC does not affect turgor in cultured maize cells, and must therefore inhibit wall-loosening or promote wall-tightening. METHODS: The effects of OAC in living cultured maize cells on various biochemical processes thought to influence wall extension were studied. KEY RESULTS: OAC did not affect 14C-incorporation from D-[U-14C]glucose into the major sugar residues of the cell wall (cellulosic glucose, non-cellulosic glucose, arabinose, xylose, galactose, mannose or uronic acids). OAC had no effect on 14C-incorporation from trans-[U-14C]cinnamate into wall-bound ferulate or its oxidative coupling-products. OAC did not influence the secretion or in-vivo action of peroxidase or xyloglucan endotransglucosylase activities-proposed wall-tightening and -loosening activities, respectively. The herbicide did not affect the consumption of extracellular L-ascorbate, an apoplastic solute proposed to act as an antioxidant and/or to generate wall-loosening hydroxyl radicals. CONCLUSIONS: OAC decreased wall extensibility without influencing the synthesis or post-synthetic modification of major architectural wall components, or the redox environment of the apoplast. The possible value of OAC as a probe to explore aspects of primary cell wall physiology is discussed.     

Selected cell wall proteins from Zea mays: Assessment of their role in wall hydrolysis

Coleoptile cell wall proteins from Zea mays L. hybrid B 37 × Mo 17 were extracted and fractionated. Three enzymes identified in that extract were examined to determine their role in cell wall hydrolysis with a goal of evaluating the extent to which they participated in autohydrolytic reactions. Two separate proteins were identified as endo- and exo-glucanases. Incubation of these enzymes with heat inactivated cell walls, liberates products derived from the constitutive (1→3), (1→4)-β- d -glucan. The release of sugars from walls resembles that of cell wall autolysis. A third cell wall protein degraded polysaccharides in a more general manner, releasing carbohydrates containing xylose, arabinose, galactose and glucose. Polyclonal antibodies raised against the exoglucanase protein suppressed autolytic reactions of isolated cell wall.     

Incorporation of 14C-l-arabinose into polysaccharides of maize root-tips

Summary [1-¹⁴C]-l-arabinose was supplied to maize roots over a range of concentrations extending from 0.1 M to 0.04 mM. In each case, only xylose and arabinose units in the cell wall polysaccharides became labelled. However, although uptake increased with concentration, the conversion of l-arabinose to these cell wall units was not greatly influenced by raising the external sugar concentration, and there was no marked accumulation of UDP-pentose under any of the experimental conditions tested. Furthermore, specific activity of the arabinose isolated from the cell wall hydrolysates was always higher than that of the xylose. Because the labelling was so specific, patterns of pentose deposition could be followed by preparing autoradiographs of sections from roots fed with ¹⁴C-l-arabinose. In the pith and cortex, which are typically parenchymatous tissues, the maximum rate of incorporation was observed in cell walls at around 2 mm from the cap-stele junction. These cells had just reached their full width and were about to undergo a phase of rapid elongation. Results are in essential aggrement with those obtained earlier with d-glucuronate in similar experiments.     

Auxin-induced changes in the cell wall structure: Changes in the sugar compositions, intrinsic viscosity and molecular weight distributions of matrix polysaccharides of the epicotyl cell wall of Vigna angularis

Rapid effects of indole-3-acetic acid (IAA) on the mechanical properties of cell wall, and sugar compositions, intrinsic viscosity and molecular weight distribution of cell wall polysaccharides were investigated with excised epicotyl segments of Vigna angularis Ohwi et Ohashi cv. Takara.

• 1 IAA caused cell wall loosening as studied by stress-relaxation analysis within 15 min after the IAA application.
• 2 IAA stimulated the decrease in the content of arabinose and galactose in the hemicellulose 1 h after its application. The amounts of other component sugars in the cell wall polysaccharides remained constant during the IAA-induced segment growth.
• 3 The intrinsic viscocity of the pectin increased as early as 30 min after the IAA application. This effect was not prevented when elongation growth of the segment was osmotically suppressed by 0.15 M mannitol.
• 4 Gel permeation chromatography of the pectin on a Sepharose 4 B column demonstrated that IAA caused increase in the mass-average molecular weight of the pectin. Analysis of the sugar compositions of the pectin eluted from the Sepharose 4 B column indicated that IAA increased the molecular weight of the polysaccharides composed of uronic acid, galactose, rhamnose and arabinose. This effect became apparent within 30 min after the IAA application. Furthermore, IAA increased the molecular weight of the pectin when elongation growth of the epicotyl segments was osmotically suppressed by 0.15 M mannitol.
• 5 Hemicellulose of the cell wall chromatographed on a Sepharose CL-4 B column. Analysis of the neutral sugar compositions and the iodine staining property (specific for xyloglucans) of the polysaccharide solution eluted from the column indicated that the hemicellulose consisted of xyloglucans, arabinogalactans and polysaccharides composed of xylose and/or mannose. IAA caused a decrease in the arabinogalactan content and depolymerization of xyloglucans. These IAA effects became apparent within 30 min after the IAA application. These changes occurred even when elongation growth of the epicotyl segments was osmotically suppressed by 0.15 M mannitol.

Polymerization of the pectin, degradation of arabinogalactans and depolymerization of xyloglucans appear to be involved in the mechanism by which IAA induces cell wall loosening and therefore extension growth of cells.     

Changes in cell wall polysaccharides and monosaccharides during development and ripening of Japanese pear fruit

1. Changes in polysaccharide and monosaccharide components in thecell wall were studied during cell division, cell enlargmementand softening in Japanese pear fruit. Wall polysaccharides werefractionated into water soluble carbohydrate, NaClO₂ solublecarbohydrate, EDTA soluble carbohydrate, acid soluble hemicellulose,alkali soluble hemicellulose and cellulose. These polysaccharideswere composed of glucose, uronic acid, xylose, arabinose, galactose,rhamnose, mannose and fucose.
2. The total polysaccharide contentof the cell wall per cell (DNAcontent basis) remained constantduring the cell division period(S1). But during the pre-enlargementperiod (S2) it began toincrease rapidly in spite of the slightnessof cell enlargement.Thereafter, during the enlargement period(S3) the polysaccharidesremained almost constant although thefruits enlarged dramatically,and the polysaccharides increasedsomewhat with ripening. Thequality of the polysaccharides,however, seemed to change activelyat each stage. This suggestedthat the extensive fruit enlargementdid not require an increasein polysaccharide content, and wasrather accompanied by thepartial breakdown or partial interconversionof polysaccharidecomponents already present.
3. The loss of arabinose and galactosein acid soluble hemicellulosewas prominant in fruit softeningoccurring in the ripening stage.The cellulose component decreasedwith overripening. Water solublepectin increased parallel tothe increase in total pectin withripening. On the other hand,xylose and non-cellulosic glucoseresidues did not alter withripening or overripening. Non-cellulosicglucose continued toaccumulate during cell enlargement.

¹ This paper is Contribution A-88, Fruit Tree Research Station. (Received August 4, 1978; )     

Cell wall formation in zoospores of Allomyces arbuscula

Carbohydrate composition was determined in isolated cell walls of meiospores of Allomyces arbuscula after incubation for 15 min (encysted meiospores: cysts), 150 min (germlings: cysts + rhizoids) and 24 h (cysts + rhizoids + hyphae). The principal constituent in all cell wall samples is chitin, accounting for about 75% of the recovered carbohydrates. In addition, cell walls of all stages examined contain polysaccharides which release galactose, glucose, mannose, arabinose, xylose, fucose, and rhamnose on acid hydrolysis. While different developmental stages show minor quantitative changes in chitin, the ratio of galactose to glucose decreases sharply during differentiation of ungerminated cysts into germlings with rhizoids and hyphae. The increase in glucose is accompanied by a decrease in the amount of xylose and/or fucose and of galactose.List of Abbreviation TFA trifluoroacetic acid