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PubMed1975

Use of L-rhamnose to study irreversible adsorption of bacteriophage PL-1 to a strain of Lactobacillus casei.

L-rhamnose has been found to be useful to investigate the process of irreversible adsorption of PL-I phage to its host bacterium, Lactobacillus casei ATCC27092. L-rhamnose inhibited phage adsorption to cells without inactivating free phages. Adsorption inhibition was correlated with the concentrations of L-rhamnose. The inhibitory effect of L-rhamnose on phage adsorption was of a competitive nature against host cells. Among other saccharides tested, L-fucose, L-mannose and D-ribose showed a slight degree of adsorption-inhibiting activity. In early stages of phage adsorption in a tris-maleate buffer, where the binding of phages to cells was still reversible, addition of L-rhamnose resulted in the partial desorption of phages from the cells to which they had adsorbed. However, the number of infective phages desorbed by L-rhamnose treatment gradually fell off as incubation continued, showing that the phages became firmly bound to the cells. Therefore, it is possible to determine the number of phages irreversibly adsorbed to cells by using this desorption technique with L-rhamnose. The process of irreversible phage adsorption, that is, the formation of phage-cell complexes from which no more infective phages could be desorbed, was dependent on temperature and strongly inhibited at 0 degrees C.

Adsorption↗
PubMed1986

Differentiation of Streptococcus sanguis and S. mitior by whole-cell rhamnose content and possession of arginine dihydrolase.

Whole-cell rhamnose concentrations were measured in 48 strains of streptococci resembling Streptococcus sanguis and S. mitior. Physiological characteristics were tested by the API-20/Strep system, and it was found that "typical" S. sanguis (arginine positive, aesculin positive) contained significant amounts of rhamnose, while "typical" S. mitior (arginine negative, aesculin negative) contained very low or undetectable amounts of rhamnose. Both groups contained dextran-positive and dextran-negative strains. Organisms that were more difficult to speciate (those giving positive results in the arginine or the aesculin test, but not in both) could also be divided into a rhamnose-positive and a rhamnose-negative group; with one exception, all of the rhamnose-positive strains gave a positive result with arginine in the API-20/Strep test. There were several discrepancies between the results of conventional tests for arginine and aesculin hydrolysis and those of the corresponding API test. The results of conventional tests for arginine hydrolysis did not correlate closely with rhamnose content, and conventional tests for aesculin hydrolysis were less sensitive than API tests. With the API-20/Strep system, S. sanguis can almost always be distinguished from S. mitior by its ability to hydrolyse arginine.

Arginine↗
PubMed1984

L-Rhamnose utilisation in Salmonella typhimurium.

L-Rhamnose is degraded by strains of Salmonella typhimurium by isomerisation to L- rhamnulose , phosphorylation to L- rhamnulose -1-phosphate and cleavage to lactaldehyde and dihydroxyacetone phosphate. The enzymes involved are, respectively, rhamnose isomerase ( RhaI ), rhamnulokinase ( RhuK ) and an aldolase (Ald). Strains able to grow rapidly on L-rhamnose contained a high-affinity uptake system for 3H-L-rhamnose that was induced by L-rhamnose and repressed by D-glucose. The synthesis of RhaI and RhuK was also induced by L-rhamnose but was not repressed by D-glucose. The synthesis of Ald was constitutive. Data are presented on some strains which grow very slowly on L-rhamnose and on others which do not utilise it.

Mutation↗
PubMed1990

The bacteriophage kh receptor of Lactococcus lactis subsp. cremoris KH is the rhamnose of the extracellular wall polysaccharide.

A receptor for bacteriophages of lactic acid bacteria, including Lactococcus lactis subsp. cremoris KH, was found on the cell wall and not on the cell membrane, as determined by a phage-binding assay of sodium dodecyl sulfate- and mutanolysin-treated cell walls. The cell wall carbohydrates of L. lactis subsp. cremoris KH were analyzed by gas chromatography and mass spectrometry and found to contain rhamnose, galactose, glucose and N-acetylglucosamine. Similar analysis of mutants that were reduced in the ability to bind phages kh, 643, c2, ml3, and 1 indicated that galactose was essential for binding all phages. In addition, rhamnose was required for binding phages kh and ml3. Inhibition studies of phage binding by using two different lectins with a specificity for galactose indicated that phage kh may not bind directly to galactose. Rather, galactose may be an essential structural component located in the vicinity of the receptor. Incubation of any of the five phages with rhamnose or of phage kh with purified cell walls inactivated the phages. Inactivation required divalent cations and was irreversible. Inactivation of phages was stereospecific for rhamnose, as neither L-(+)- nor D-(-)-fucose (the stereoisomers of rhamnose) inhibited the phage. Furthermore, phage infection of a culture was completely inhibited by the addition of rhamnose to the medium. Therefore, the receptor for phage kh appears to be a rhamnose component of the extracellular wall polysaccharide.

Bacteriophages↗
PubMed2004

A genetic locus necessary for rhamnose uptake and catabolism in Rhizobium leguminosarum bv. trifolii.

Rhizobium leguminosarum bv. trifolii mutants unable to catabolize the methyl-pentose rhamnose are unable to compete effectively for nodule occupancy. In this work we show that the locus responsible for the transport and catabolism of rhamnose spans 10,959 bp. Mutations in this region were generated by transposon mutagenesis, and representative mutants were characterized. The locus contains genes coding for an ABC-type transporter, a putative dehydrogenase, a probable isomerase, and a sugar kinase necessary for the transport and subsequent catabolism of rhamnose. The regulation of these genes, which are inducible by rhamnose, is carried out in part by a DeoR-type negative regulator (RhaR) that is encoded within the same transcript as the ABC-type transporter but is separated from the structural genes encoding the transporter by a terminator-like sequence. RNA dot blot analysis demonstrated that this terminator-like sequence is correlated with transcript attenuation only under noninducing conditions. Transport assays utilizing tritiated rhamnose demonstrated that uptake of rhamnose was inducible and dependent upon the presence of the ABC transporter at this locus. Phenotypic analyses of representative mutants from this locus provide genetic evidence that the catabolism of rhamnose differs from previously described methyl-pentose catabolic pathways.

Bacterial Proteins↗
PubMed2004

Identification and functional characterization of the Lactococcus lactis rfb operon, required for dTDP-rhamnose Biosynthesis.

dTDP-rhamnose is an important precursor of cell wall polysaccharides and rhamnose-containing exopolysaccharides (EPS) in Lactococcus lactis. We cloned the rfbACBD operon from L. lactis MG1363, which comprises four genes involved in dTDP-rhamnose biosynthesis. When expressed in Escherichia coli, the lactococcal rfbACBD genes could sustain heterologous production of the Shigella flexneri O antigen, providing evidence of their functionality. Overproduction of the RfbAC proteins in L. lactis resulted in doubled dTDP-rhamnose levels, indicating that the endogenous RfbAC activities control the intracellular dTDP-rhamnose biosynthesis rate. However, RfbAC overproduction did not affect rhamnose-containing B40-EPS production levels. A nisin-controlled conditional RfbBD mutant was unable to grow in media lacking the inducer nisin, indicating that the rfb genes have an essential role in L. lactis. Limitation of RfbBD activities resulted in the production of altered EPS. The monomeric sugar of the altered EPS consisted of glucose, galactose, and rhamnose at a molar ratio of 1:0.3:0.2, which is clearly different from the ratio in the native sugar. Biophysical analysis revealed a fourfold-greater molecular mass and a twofold-smaller radius of gyration for the altered EPS, indicating that these EPS are more flexible polymers with changed viscosifying properties. This is the first indication that enzyme activity at the level of central carbohydrate metabolism affects EPS composition.

Bacterial Proteins↗
PubMed1986

Lactulose, 51Cr-labelled ethylenediaminetetra-acetate, L-rhamnose and polyethyleneglycol 400 [corrected] as probe markers for assessment in vivo of human intestinal permeability.

The urinary excretion of lactulose, 51Cr-labelled ethylenediaminetetra-acetate (51Cr-EDTA), L-rhamnose and polyethyleneglycol 400 (PEG-400) has been measured after intravenous and oral administration in healthy volunteers. Intestinal permeation of the probes was compared after their ingestion in iso-osmolar, hyperosmolar and cetrimide-containing test solutions. Urinary recovery of lactulose and 51Cr-EDTA after intravenous administration reached 75% by 5 h, and exceeded 90% at 24 h, and these values were 62 and 72%, respectively, for L-rhamnose. Recovery of PEG-400, however, varied with the relative molecular mass (Mr) of each polymer from 25.9 to 68.5% in 24 h. Intestinal permeation of ingested lactulose and 51Cr-EDTA was low, but that of L-rhamnose was 45-fold, and that of PEG-400 100-fold, greater. Permeation of lactulose and 51Cr-EDTA was markedly increased by cetrimide and hyperosmolar stress, whereas that of L-rhamnose showed little change. PEG-400 permeation was not affected by cetrimide, but was slightly increased by hyperosmolar stress. The 5 h permeation of lactulose, but not of L-rhamnose or PEG-400, correlated with that of 51Cr-EDTA (r = 0.98, P less than 0.001). These findings are compatible with three distinct pathways of unmediated mucosal permeation, L-rhamnose (radius less than 0.4 nm) passing mainly through small aqueous 'pores' of high incidence, lactulose and 51Cr-EDTA (radius greater than 0.5 nm) through larger aqueous 'channels' of low incidence susceptible to cetrimide and hyperosmolar stress, and PEG-400, which has appreciable lipid solubility, by partition through cell membrane lipid as well as the aqueous 'pores'.

Cell Membrane Permeability↗
PubMedDate not supplied

The absence of rhamnose in human urine following the ingestion of gum karaya (Sterculia).

Urinary rhamnose estimations following ingestion of gum karaya were requested by the Scientific Committee for Food (EEC) in July 1983. Five male volunteers have therefore made 24-h urine collections prior to, and following, the ingestion of 10 g gum karaya for 15 days, an intake ten-fold greater than that approved in terms of the present temporary ADI (0-12 X 5 mg/kg b.w.). Paper chromatographic separations, with two solvent systems, were made on the fresh urine specimens and also after ten-fold enrichments of all urinary constituents. Standard aqueous solutions of rhamnose, and urine to which rhamnose had been added, showed the detection limit to be 0.2 microgram rhamnose. Independent examinations in two laboratories failed to detect rhamnose at this level in any of the urine specimens, Had 1% of the rhamnose present in 10 g gum karaya appeared in the 24-h urine specimens, it would have been detected. This confirms previous evidence that dietary gum karaya is neither digested nor degraded by enteric bacteria and is not absorbed to any significant extent in Man.

Adult↗
PubMed2004

L-Rhamnose increases serum propionate in humans.

BACKGROUND: Acetic and propionic acids are produced by colonic bacterial fermentation of unabsorbed carbohydrates and are absorbed into the portal circulation. From there, they travel to the liver, where acetate is a lipogenic substrate and propionate can inhibit lipogenesis. The extent to which peripheral blood short-chain fatty acid concentrations reflect differences in colonic fermentation is uncertain. The unabsorbed sugar lactulose produces mainly acetate when fermented in vitro, whereas L-rhamnose yields propionate. OBJECTIVE: The objective of the study was to ascertain whether ingestion of L-rhamnose and lactulose would have different acute effects on peripheral acetate and propionate concentrations and on breath hydrogen and methane concentrations. DESIGN: Twenty-two subjects were fed 25 g L-rhamnose, lactulose, or glucose on 3 separate occasions in a randomized crossover design. Blood and breath samples were collected hourly for 12 h. RESULTS: Serum propionate was significantly higher with ingestion of L-rhamnose than with that of lactulose or glucose (P < 0.001). The area under the curve for serum acetate was significantly higher with ingestion of lactulose than with that of glucose (P < 0.03). The ratio of serum acetate to propionate was significantly higher with ingestion of lactulose than with that of glucose or L-rhamnose (P < 0.01). Breath hydrogen was significantly higher with ingestion of lactulose than with that of L-rhamnose or glucose (P < 0.0001). CONCLUSIONS: The selective increases in serum acetate and propionate concentrations in humans were obtained by feeding specific fermentable substrates. Presumably, these changes in serum concentrations reflect changes in colonic production. Selective alteration of colonic fermentation products could yield a new mechanism for modifying blood lipids.

Acetates↗
PubMed2006

L-rhamnose and lactulose decrease serum triacylglycerols and their rates of synthesis, but do not affect serum cholesterol concentrations in men.

Colonic short-chain fatty acids (SCFA) may affect hepatic lipid metabolism. Lactulose increases colonic acetate production, whereas L-rhamnose increases propionate. To test the effects of oral L-rhamnose and lactulose for 28 d on fasting concentrations and hepatic synthesis of lipids in humans, 18 men were administered 25 g/d of L-rhamnose, lactulose, or d-glucose for 4 wk in a partially randomized crossover design, with blood collected from fasting subjects on the first and last day of each period. Cholesterol and triacylglycerol (TG) synthesis rates were determined using deuterated water uptake rate over the last 24 h of each period. Postprandial blood lipids, and glucose and insulin were assessed in 11 subjects on d 28. Fasting serum cholesterol was unchanged; however, when expressed as a percentage change, TG were decreased, relative to baseline (P < 0.04), by L-rhamnose (-10%) and lactulose (-10%), compared with D-glucose, which increased serum TG (+11%). Net TG-fatty acid (TGFA) synthesis on d 28 was lower with L-rhamnose (2.42 +/- 0.38 g/d) and lactulose (2.62 +/- 0.35 g/d) than with D-glucose (2.96 +/- 0.31 g/d, P < 0.01). We conclude that these results do not support a primary role for propionate in the cholesterol-lowering effect of soluble fiber. However, both lactulose and L-rhamnose lowered serum TG (expressed as a percentage change) and TGFA synthesis, compared with d-glucose, which increased them. Although these data are consistent with inhibition of TGFA synthesis by SCFA, other aspects of the metabolism of these sugars cannot be ruled out as putative agents of their TG-lowering effects.

Adult↗
PubMedDate not supplied

Lactulose-rhamnose intestinal permeability in children with cystic fibrosis.

The lactulose-rhamnose intestinal permeability test was performed in 10 cystic fibrosis (CF) children. Urine was collected for 5 h after oral intake of the test solution. A single thin-layer chromatography followed by densitometry was used for the measurement of lactulose and rhamnose concentrations in the urine specimens. The excretion of each molecule was expressed as the percentage of the orally administered dose excreted and the lactulose-rhamnose ratio was the ratio of the percentage quantities of each probe molecule excreted. The mean lactulose-rhamnose excretion ratio in CF patients and controls was 0.16 and 0.038, respectively (p less than 0.001). The mean 5-h lactulose excretion was 2% and 0.28% in CF patients and control subjects, respectively (p less than 0.001). The mean 5-h rhamnose excretion was 12.8% and 7.6% in CF patients and control subjects (p less than 0.05). We conclude that CF patients have an increased intestinal permeability to lactulose and rhamnose. Further studies are needed in order to elucidate the mechanisms involved as well as the eventual nutritional implications.

Adolescent↗
PubMed1987

Cross-induction of the L-fucose system by L-rhamnose in Escherichia coli.

Dissimilation of L-fucose as a carbon and energy source by Escherichia coli involves a permease, an isomerase, a kinase, and an aldolase encoded by the fuc regulon at minute 60.2. Utilization of L-rhamnose involves a similar set of proteins encoded by the rha operon at minute 87.7. Both pathways lead to the formation of L-lactaldehyde and dihydroxyacetone phosphate. A common NAD-linked oxidoreductase encoded by fucO serves to reduce L-lactaldehyde to L-1,2-propanediol under anaerobic growth conditions, irrespective of whether the aldehyde is derived from fucose or rhamnose. In this study it was shown that anaerobic growth on rhamnose induces expression of not only the fucO gene but also the entire fuc regulon. Rhamnose is unable to induce the fuc genes in mutants defective in rhaA (encoding L-rhamnose isomerase), rhaB (encoding L-rhamnulose kinase), rhaD (encoding L-rhamnulose 1-phosphate aldolase), rhaR (encoding the positive regulator for the rha structural genes), or fucR (encoding the positive for the fuc regulon). Thus, cross-induction of the L-fucose enzymes by rhamnose requires formation of L-lactaldehyde; either the aldehyde itself or the L-fuculose 1-phosphate (known to be an effector) formed from it then interacts with the fucR-encoded protein to induce the fuc regulon.

Alcohol Oxidoreductases↗
PubMed2006

Preparative synthesis of dTDP-L-rhamnose through combined enzymatic pathways.

dTDP-L-rhamnose, an important precursor of O-antigen, was prepared on a large scale from dTMP by executing an one-pot reaction in which six enzymes are involved. Two enzymes, dTDP-4-keto-6-deoxy-D-glucose 3,5-epimerase and dTDP-4-keto-rhamnose reductase, responsible for the conversion of dTDP-4-keto-6-deoxy-D-glucose to dTDP-L-rhamnose, were isolated from their putative sequences in the genome of Mesorhizobium loti, functionally expressed in Escherichia coli, and their enzymatic activities were identified. The two enzymes were combined with an enzymatic process for dTDP-4-keto-6-deoxy-D-glucose involving TMP kinase, acetate kinase, dTDP-glucose synthase, and dTDP-glucose 4,6-dehydratase, which allowed us to achieve a preparative scale synthesis of dTDP-L-rhamnose using dTMP and glucose-1-phosphate as starting materials. About 82% yield of dTDP-L-rhamnose was obtained based on initial dTMP concentration at 20 mM dTMP, 1 mM ATP, 10 mM NADH, 60 mM acetyl phosphate, and 80 mM glucose-1-phosphate. From the reaction with 20 ml volume, approximately 180 mg of dTDP-L-rhamnose was obtained in an overall yield of 60% after two-step purification, that is, anion exchange chromatography and gel filtration for desalting. The purified product was identified by HPLC, ESI-MS, and NMR, showing about 95% purity.

Bacterial Proteins↗
PubMed2001

Lactulose-L-rhamnose intestinal permeability test in patients with liver cirrhosis.

A lactulose-L-rhamnose intestinal permeability test was conducted on 35 patients with liver cirrhosis and six normal controls. Gas chromatography was used to measure lactulose and L-rhamnose concentrations in blood and urine specimens. The excretion of each molecule was expressed as the percentage of the orally administrated dose and the lactulose-L-rhamnose ratio as the ratio of the percentage of each probe molecule excreted. The mean 8-h lactulose excretion ratios were 0.56 and 0.16% in patients with liver cirrhosis and the control subjects, respectively (P<0.05), whereas the corresponding excretion ratios for L-rhamnose were 4.40 and 3.49%. The mean lactulose-L-rhamnose excretion ratios in patients with liver cirrhosis and the control subjects were 0.124 and 0.049, respectively (P<0.05). The lactulose-L-rhamnose excretion ratio increased in patients with liver cirrhosis complicated by large intestinal vascular ectasia of the large intestine or rectal varices, which were used as parameters for evaluating the effects of portal hypertension on the lower digestive tract. These results suggest that an increase in lactulose intestinal permeability in patients with liver cirrhosis proves the effects of portal hypertension extending to the lower digestive tract.

Journal Article↗
PubMed1997

Induction by (alpha)-L-Arabinose and (alpha)-L-Rhamnose of Endopolygalacturonase Gene Expression in Colletotrichum lindemuthianum.

The production of endopolygalacturonase (endoPG) by Colletotrichum lindemuthianum, a fungal pathogen causing anthracnose on bean seedlings, was enhanced when the fungus was grown in liquid medium with L-arabinose or L-rhamnose as the sole carbon source. These two neutral sugars are present in plant cell wall pectic polysaccharides. The endolytic nature of the enzyme was demonstrated by its specific interaction with the polygalacturonase-inhibiting protein of the host plant as well as by sugar analysis of the products released from its action on oligogalacturonides. Additional characterization of the protein was achieved with an antiserum raised against the pure endoPG of the fungus. Induction by arabinose and rhamnose was more prolonged and led to a level of enzyme activity at least five times higher than that on pectin. Northern blot experiments showed that this effect was correlated to the induction of a 1.6-kb transcript. A dose-response study indicated that the endoPG transcript level was already increased at a concentration of each sugar as low as 2.75 mM in the medium and was maximum at 55 mM arabinose and 28 mM rhamnose. Glucose, the main plant cell wall sugar residue which is also present in the apoplast, prevented endoPG gene expression, partially when added to pectin at concentrations ranging from 5 to 110 mM and totally when added at 55 mM to arabinose. Inhibition by glucose of the rhamnose-induced endoPG was correlated to nonuptake of rhamnose. This is the first report that arabinose and rhamnose stimulate endoPG gene expression in a fungus. The possible involvement of these various sugars on endoPG gene expression during pathogenesis is discussed.

Journal Article↗
PubMed1997

Identification of a fourth gene involved in dTDP-rhamnose synthesis in Streptococcus mutans.

We had isolated three genes (rmlA, rmlB, and rmlC) involved in dTDP-rhamnose synthesis in Streptococcus mutans and found that three genes were insufficient for dTDP-rhamnose synthesis (Y. Tsukioka, Y. Yamashita, T. Oho, Y. Nakano, and T. Koga, J. Bacteriol. 179:1126-1134, 1997). The rmlD gene of S. mutans, encoding the enzyme which catalyzes the last step of dTDP-rhamnose synthesis, has been cloned and sequenced. The cell extract of Escherichia coli expressing the rmlD gene of S. mutans exhibited enzymatic activity corresponding to its counterpart in Shigella flexneri, a gram-negative bacterium. Rhamnose was not detected in the cell wall preparation purified from the mutant in which the cloned gene was insertionally inactivated. Rabbit antiserum against S. mutans serotype c-specific antigen did not react with autoclaved extracts from the mutant. The rmlD gene product of S. mutans compensated for the incompleteness of dTDP-rhamnose synthesis by the three previously isolated genes. These results indicate that the rmlD gene product is indispensable for the dTDP-rhamnose pathway and subsequently for the synthesis of serotype-specific antigen in S. mutans. Furthermore, conservation of the rmlD gene in Streptococcus species was demonstrated by Southern blot analysis.

Amino Acid Sequence↗
PubMed2003

Simultaneous gas-chromatographic measurement of rhamnose, lactulose and sucrose and their application in the testing gastrointestinal permeability.

BACKGROUND: As it is important to test gastric and intestinal permeability simultaneously in gastrointestinal disorders such as Celiac disease, we developed a gas-chromatographic (GC) method to estimate rhamnose (L-rh), lactulose (Lacl) and sucrose (Suc) in urine. METHODS: The method is based on the use of alditol acetate derivatives giving a lower number of GC peaks than reducing sugars do. Acetate derivatives are more stable and less expensive than GC silylates and keep the flame-detector cleaner. We checked the chemical stability of alditol acetates by verifying the reproducibility of the standard curve of a sugar derivative sample which had been stored for 2 months at -20 degrees C. RESULTS: The calibration proved linear over the range 0.1-1 microg of sugar injected. Analytical sugar recovery was 88%+/-19.4% (mean+/-S.D.) for rhamnose, 105%+/-7.4% for sucrose and 102%+/-2.4% for lactulose. Mean within-day precision (CV) was 7.7% for rhamnose, 5.7% for sucrose and 1.9% for lactulose, and between-day (CV) was 6.7% for rhamnose, 3.9% for sucrose and 1.6% for lactulose. The rhamnose, lactulose and sucrose as the lactulose/rhamnose ratio clearly differentiated 25 healthy controls from 36 patients with active gluten-sensitive enteropathy. CONCLUSIONS: A fast, reliable and cheap gas-chromatographic method is presented here to evaluate gastric and intestinal permeability.

Celiac Disease↗
PubMed1998

Scarcity or complete lack of single rhamnose residues interspersed within the homogalacturonan regions of citrus pectin.

Commercial citrus pectin containing galacturonic acid and rhamnose in a ratio of approximately 40:1 was saponified and then exhaustively digested with endopolygalacturonase (EPG). The products were separated by ultrafiltration into low-molecular-weight (LMW) and high-molecular-weight (HMW) fractions. The LMW fraction accounted for 80% of the starting material, but for only 10% of the total rhamnose. The molar ratio of galacturonic acid to rhamnose of the LMW fraction was 236, suggesting that very few small Rha-containing oligomers were generated by the EPG digestion. No distinct Rha-containing oligomers were found by various chromatographic analyses of the LMW fraction. The HMW fraction, which only accounted for 10% by weight of the starting pectin, contained more than 85% of the rhamnose. The ratio of GalA to Rha in the HMW fraction was 1.7:1 and partial acid hydrolysis of this fraction produced a series of oligomers consisting of GalA-Rha repeating units, suggesting that it contained rhamnogalacturonan, which has a backbone composed of GalA-Rha disaccharide repeating units. The HMW fraction also contained large amounts of arabinose and galactose, which probably originated from side chains linked to some of the rhamnose residues. We propose that commercial citrus pectin is composed of two regions: the predominant region consists of chains of uninterrupted 1,4-linked alpha-D-GalA residues with between 60-70% of the residues methyl esterified; and the other region consists of rhamnogalacturonan with a backbone composed of GalA-Rha disaccharide repeating units and neutral sugar side chains.

Carbohydrate Sequence↗