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PubMed1971

Biosynthesis of levan and a new method for the assay of levansucrase activity.

The polysaccharide levan was synthesized in a solidified agar medium containing sucrose as a source of fructose. The biosynthesis was achieved by the enzyme levansucrase (2,6-fructan-d-glucose 6-fructosyltransferase, EC 2.4.1.10), a small quantity of which was placed in circular wells cut in the agar gel. The enzyme slowly diffused through the agar-sucrose medium and the synthesis of levan was observed as circular white areas, the size of which was dependent on the time of incubation and the concentration of enzyme used.

Agar↗

PubMed2006

Crystal structure of inactivated Thermotoga maritima invertase in complex with the trisaccharide substrate raffinose.

Thermotoga maritima invertase (beta-fructosidase), a member of the glycoside hydrolase family GH-32, readily releases beta-D-fructose from sucrose, raffinose and fructan polymers such as inulin. These carbohydrates represent major carbon and energy sources for prokaryotes and eukaryotes. The invertase cleaves beta-fructopyranosidic linkages by a double-displacement mechanism, which involves a nucleophilic aspartate and a catalytic glutamic acid acting as a general acid/base. The three-dimensional structure of invertase shows a bimodular enzyme with a five bladed beta-propeller catalytic domain linked to a beta-sandwich of unknown function. In the present study we report the crystal structure of the inactivated invertase in interaction with the natural substrate molecule alpha-D-galactopyranosyl-(1,6)-alpha-D-glucopyranosyl-beta-D-fructofuranoside (raffinose) at 1.87 A (1 A=0.1 nm) resolution. The structural analysis of the complex reveals the presence of three binding-subsites, which explains why T. maritima invertase exhibits a higher affinity for raffinose than sucrose, but a lower catalytic efficiency with raffinose as substrate than with sucrose.

Aspergillus↗

PubMed2003

The role of invertases and hexose transporters in controlling sugar ratios in maternal and filial tissues of barley caryopses during early development.

To analyse carbohydrate metabolism and its role during early seed development of barley we characterised genes encoding two cell wall-bound invertases (HvCWINV1 and HvCWINV2) and two putative hexose transporter-like genes (HvSTP1 and HvSTP2). No typical vacuolar invertase gene could be identified. Instead, a gene encoding sucrose:fructan 6-fructosyltransferase (HvSF6FT1), an enzyme with soluble acid invertase activity, was isolated and characterised. Furthermore, enzyme activities and sugar levels were measured. HvSF6FT1-mRNA levels and acid soluble invertase activity are highest in the maternal pericarp 1-2 days after flowering (DAF). HvSF6FT1 is strongly expressed in regions flanking the main vascular bundle and to a lower extent in filial endospermal transfer cells, which persist until maturity and never accumulate starch. In contrast, cell wall-bound invertase HvCWINV2 is expressed early in development mainly in the style region and later on in pericarp areas which transiently accumulate starch and undergo degradation later in development. The hexose transporter HvSTP2 shows a temporal and spatial expression pattern similar to HvCWINV2. Transcripts of HvCWINV1 have been localised within the first row of endospermal cells and in the outermost area of the nucellar projection as well as in endospermal transfer cells before starch filling; the same regions of the endosperm are labelled with a hexose transporter HvSTP1-probe. HvSTP1 is expressed at very low levels within the pericarp but much higher in the syncytial endosperm at 3 DAF and in endospermal transfer cells 7 DAF. The temporal and spatial association of HvCWINV1 and HvSTP1 expression indicates that hexoses liberated by the invertase within the endospermal cavity are taken up by the transporter to be delivered into the central uncellularised space of the endosperm to supply mitotically active endosperm cells with hexoses. The results are discussed and compared with published data on the role of soluble sugars as signal molecules in seed developmental processes.

Carbohydrate Metabolism↗

PubMedDate not supplied

Non-digestible oligosaccharides used as prebiotic agents: mode of production and beneficial effects on animal and human health.

Prebiotic agents are food ingredients that are potentially beneficial to the health of consumers. The main commercial prebiotic agents consist of oligosaccharides and dietary fibres (mainly inulin). They are essentially obtained by one of three processes: 1) the direct extraction of natural polysaccharides from plants; 2) the controlled hydrolysis of such natural polysaccharides; 3) enzymatic synthesis, using hydrolases and/or glycosyl transferases. Both of these enzyme types catalyse transglycosylation reactions, allowing synthesis of small molecular weight synthetic oligosaccharides from mono- and disaccharides. Presently, in Europe, inulin-type fructans, characterised by the presence of fructosyl units bound to the beta-2,1 position of sucrose, are considered as one of the carbohydrate prebiotic references. Prebiotics escape enzymatic digestion in the upper gastrointestinal tract and enter the caecum without change to their structure. None are excreted in the stools, indicating that they are fermented by colonic flora so as to give a mixture of short-chain fatty acids (acetate, propionate and butyrate), L-lactate, carbon dioxide and hydrogen. By stimulating bifidobacteria, they may have the following implications for health: 1) potential protective effects against colorectal cancer and infectious bowel diseases by inhibiting putrefactive bacteria (Clostridium perfringens ) and pathogen bacteria (Escherichia coli, Salmonella, Listeria and Shigella ), respectively; 2) improvement of glucid and lipid metabolisms; 3) fibre-like properties by decreasing the renal nitrogen excretion; 4) improvement in the bioavailability of essential minerals; and 5) low cariogenic factor. These potential beneficial effects have been largely studied in animals but have not really been proven in humans. The development of a second generation of oligosaccharides and the putative implication of a complex bacterial trophic chain in the intestinal prebiotic fermentation process are also discussed.

Animals↗

PubMed2006

Structural and functional insights into intramolecular fructosyl transfer by inulin fructotransferase.

Inulin fructotransferase (IFTase), a member of glycoside hydrolase family 91, catalyzes depolymerization of beta-2,1-fructans inulin by successively removing the terminal difructosaccharide units as cyclic anhydrides via intramolecular fructosyl transfer. The crystal structures of IFTase and its substrate-bound complex reveal that IFTase is a trimeric enzyme, and each monomer folds into a right-handed parallel beta-helix. Despite variation in the number and conformation of its beta-strands, the IFTase beta-helix has a structure that is largely reminiscent of other beta-helix structures but is unprecedented in that trimerization is a prerequisite for catalytic activity, and the active site is located at the monomer-monomer interface. Results from crystallographic studies and site-directed mutagenesis provide a structural basis for the exolytic-type activity of IFTase and a functional resemblance to inverting-type glycosyltransferases.

Amino Acid Sequence↗

PubMed2003

Isolation and characterisation of an invertase cDNA from perennial ryegrass (Lolium perenne).

An invertase (LpFT2) cDNA from perennial ryegrass was isolated and sequenced. Nucleotide sequence analysis revealed an ORF of 2016 bp encoding a protein of 671 amino acids. LpFT2 is 76% identical to sugarcane soluble acid invertase, and contains invertase and fructosyltransferase functional domains. LpFT2 is present as a single copy gene and maps to the distal region of LG6 in perennial ryegrass. The expression pattern analysis of LpFT2 revealed transcript accumulation in seedlings and in mature leaf sheaths. The LpFT2 recombinant protein expressed in yeast showed invertase and fructan exohydrolase-like activities with complete breakdown of sucrose, 1-kestose (DP3), 1,1-kestotetraose (DP4) and 1,1,1-kestopentaose (DP5) into glucose and fructose.

Amino Acid Sequence↗

PubMed1995

Recovery of inulin from Jerusalem artichoke (Helianthus tuberosus L.) in the small intestine of man.

The recovery of inulin, a naturally occurring beta (2-->1)-fructan isolated from Jerusalem artichoke (Helianthus tuberosus L.), in the small intestine of man was studied in ileostomy subjects. The ileostomists were given a low-dietary-fibre diet based on white wheat bread and virtually free of inulin, and the same diet with the addition of 10 g and 30 g inulin product respectively, and the recovery and mean transit time (MTT) of inulin were estimated by tracking inulin in ileal effluent. The recovery of inulin was approximately 87% at both ingestion levels. MTT was 4.9 (SE 0.6) h at an intake of 10 g inulin product decreasing to 3.4 (SE 0.3) h at an intake of 30 g inulin product. A significant change in the fructose:glucose ratio of inulin from ingestion (4.1) to recovery in ileal effluent (4.5-4.7) and a lower recovery of the glucose residue than of the fructose residue of inulin indicate that the low-molecular-weight inulins are more sensitive to hydrolysis than the high-molecular-weight fragments. The loss of inulin during passage through the small intestine is presumably due to hydrolysis by either acids or enzymes and to microbial degradation by the microflora permanently colonizing the distal small intestine. The concentrations of lactic acid (LA) and short-chain fatty acids (SCFA) in frequently collected ileal effluents on the control day were approximately 6 nmol/l and approximately 55 mmol/l respectively. During periods with inulin ingestion the concentration of LA increased to 18-26 mmol/l (P < 0.052), while the concentration of SCFA ran converse and decreased to 18-32 mmol/l (P < 0.023). The osmotic loads (68 and 204 mosmol/l) associated with the ingestion of inulin product caused minor malabsorption of low-molecular-weight sugars.

Adult↗

PubMed2005

Inulin and oligofructose: impact on intestinal diseases and disorders.

A large and diverse variety of bacteria have evolved and adapted to live in the human intestinal habitat in a symbiotic arrangement that influences both physiology and pathology in the host. Symbiosis between host and flora can be optimised by prebiotics. Inulin-type fructans have been shown to improve the metabolic functions of the commensal flora. Clinical and experimental data suggest that they also improve the gut mucosal barrier. Furthermore, modulation of the trophic functions of the flora by these prebiotics could help in the prevention of inflammatory bowel diseases. The anti-inflammatory effects of inulin or oligofructose have been assessed in the rat model of distal colitis induced by dextran sodium sulphate, which histologically resembles human ulcerative colitis, and in the trinitrobenzene sulphonic acid model that resembles human Crohn's disease. Both inulin and oligofructose stimulate colonic production of SCFA and favour the growth of indigenous lactobacilli and/or bifidobacteria. These effects are associated with reduced mucosal inflammation and decreased mucosal lesion scores. Inulin has also been tested in a placebo-controlled clinical trial in patients with relapsing pouchitis. Treatment reduced endoscopic and histological parameters of inflammation of the pouch mucosa. Inulin and oligofructose may offer an opportunity to prevent chronic inflammatory intestinal disorders, and this potential should be tested in further clinical studies.

Animals↗

PubMed2002

Functional foods: concepts and application to inulin and oligofructose.

A food can be regarded as functional if it is satisfactorily demonstrated to affect beneficially one or more target functions in the body, beyond adequate nutritional effects, in a way which is relevant to either the state of well-being and health or the reduction of the risk of a disease. Health claims are expected to be authorized for functional foods based either on enhanced function (type A claim) or disease risk reduction (type B claim). Their development is a unique opportunity to contribute to the improvement of the quality of the food offered to consumer's choice for the benefit of his well-being and health. But only a rigorous scientific approach producing sound data will guarantee its success. The functional food components that are discussed in the proceedings of the 3rd ORAFTI Research Conference are the inulin-type fructans, natural food components found in miscellaneous edible plants. They are non-digestible oligosaccharides that are classified as dietary fiber. The targets for their functional effects are the colonic microflora that use them as selective 'fertilizers'; the gastrointestinal physiology; the immune functions; the bioavailability of minerals; and the metabolism of lipids. Potential health benefits may also concern reduction of the risk of some diseases like intestinal infections, constipation, non-insulin dependent diabetes, obesity, osteoporosis or colon cancer. The present proceedings review the scientific data available and, by reference to the concepts in functional food science, they assess the scientific evidence which will be used to substantiate health claims.

Bacteria↗

PubMed2002

Gastrointestinal effects of prebiotics.

The defining effect of prebiotics is to stimulate selectively the growth of bifidobacteria and lactobacilli in the gut and, thereby, increase the body's natural resistance to invading pathogens. Prebiotic carbohydrates may also have additional, less specific, benefits because they are fermented in the large intestine. The prebiotic carbohydrates that have been evaluated in humans at the present time largely consist of fructans or galactans. There is consistent evidence from in vitro and in vivo studies that these are not digested by normal human enzymes, but are readily fermented by anaerobic bacteria in the large intestine. There are no reports of faecal recovery of measurable quantities of prebiotic carbohydrates. Through fermentation in the large intestine, prebiotic carbohydrates yield short-chain fatty acids, stimulate the growth of many bacterial species in addition to the selective effects on lactobacilli and bifidobacteria, they can also produce gas. Along with other fermented carbohydrates, prebiotics have mild laxative effects, although this has proved difficult to demonstrate in human studies because the magnitude of laxation is small. Potentially, the most important effect of prebiotic carbohydrates is to strengthen the body's resistance to invading pathogens and, thereby, prevent episodes of diarrhoea. At the present time, this effect has not been convincingly demonstrated in either adults or children, although there have been attempts to ameliorate the diarrhoea associated with antibiotics and travel, but without success. However, prebiotic carbohydrates clearly have significant and distinctive physiological effects in the human large intestine, and on the basis of this it is likely that they will ultimately be shown to be beneficial to health.

Bifidobacterium↗

PubMed2003

On defining dietary fibre.

Establishing a definition for dietary fibre has historically been a balance between nutrition knowledge and analytical method capabilities. While the most widely accepted physiologically-based definitions have generally been accurate in defining the dietary fibre in foods, scientists and regulators have tended, in practice, to rely on analytical procedures as the definitional basis in fact. As a result, incongruities between theory and practice have resulted in confusion regarding the components that make up dietary fibre. In November 1998 the president of the American Association of Cereal Chemists (AACC) appointed an expert scientific review committee and charged it with the task of reviewing and, if necessary, updating the definition of dietary fibre. The committee was further charged with assessing the state of analytical methodology and making recommendations relevant to the updated definition. After due deliberation, an updated definition of dietary fibre was delivered to the AACC Board of Directors for consideration and adoption (Anon, 2000; Jones 2000b). The updated definition includes the same food components as the historical working definition used for approximately 30 years (a very important point, considering that the majority of the research of the past 30 years delineating the positive health effects of dietary fibre is based on that working definition). However, the updated definition more clearly delineates the make-up of dietary fibre and its physiological functionality. As a result, relatively few changes will be necessary in analytical methodology. Current methodologies, in particular AACC-approved method of analysis 32-05 (Grami, 2000), Association of Official Analytical Chemists' official method of analysis 985.29 (Horwitz, 2000a) or AACC 32-07 (Grami, 2000) Association of Official Analytical Chemists 991.43 (Horwitz, 2000a) will continue to be sufficient and used for most foods. A small number of additional methods will be necessary to quantify the dietary fibre levels in foods containing fibres such as fructans (polymers and oligomers of fructose, inulin), modified dextrins, and/or synthetic dietary fibre analogues.

Chemistry Techniques, Analytical↗

PubMed1993

Dietary fiber, inulin, and oligofructose: a review comparing their physiological effects.

Dietary fiber is a general term. It covers a wide variety of substances that belong to the family of carbohydrates that resist hydrolysis by human alimentary enzymes but are fermented by colonic microflora. The main physiological effects of dietary fiber are primarily on gastric emptying and small intestinal transit time, resulting in an improved glucose tolerance and a decreased digestion of starch: second, on colonic transit time and large bowel functions due to fermentation by ceco-colonic microbial flora or bulking action. The so-called soluble dietary fibers are fermented to a large extent by a wide variety of anaerobic bacteria that result in an increase in bacterial biomass, an increase in fecal mass, a change in intracolonic pH, and production of short chain fatty acids and various gases as metabolic end products. The insoluble fibers are only marginally fermented: they serve almost exclusively as bulking agents that result in shorter transit time and increased fecal mass. The short chain fatty acids resulting from the colonic fermentation of dietary fiber are largely absorbed via the portal blood and reach both the liver and the peripheral tissues. They induce changes in glucose and fat metabolism leading to post-prandial hypoglycemia and long-term hypolipidemia. Inulin and oligofructose are fructans with a degree of polymerization of 2 to 60 and 2 to 20, respectively. Due to the structural conformation of their osidic bridge (beta 2-1), they both resist the hydrolysis by human alimentary enzymes. Moreover, when reaching the colon, both inulin and oligofructose are almost quantitatively fermented almost exclusively by colonic bifidobacteria and bacteroides. Such an extensive fermentation causes an increase in fecal bacterial biomass, a decrease in ceco-colonic pH, and produces a large amount of fermentation products among which the short chain fatty acids that exert systemic effects on lipid metabolism. Thus, both inulin and oligofructose have most of the characteristics of a dietary fiber and the proposal is made to classify them as such. Moreover, they are bifidogenic factors, because, due to still unknown reasons, they are primarily fermented by bifidobacteria. It is concluded from this review that "nondigestible fructo-oligosaccharides," even though they are not included in the carbohydrate fraction that is quantified as dietary fiber by classic analytical methods, have most of the physiological effects of a dietary fiber.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1995

On the presence of inulin and oligofructose as natural ingredients in the western diet.

The classic definitions of inulin and oligofructose are constructively criticized. It is observed that inulin cannot unequivocally be described as a polydisperse 1-kestose-based (GFn) beta (2-->1) linear fructan chain, but that inulin always contains small amounts of Fm and branched molecules. This review article describes the presence of inulin and oligofructose in common foodstuffs. Historical data on human consumption add an extra dimension. Modern analytical techniques (HPLC, LGC, HPAEC-PAD) are used to check the variety of data mentioned in the literature throughout the past century. Methods to determine inulin and oligofructose in natural foodstuffs (cereals, fruit, and vegetables) are optimized and used to determine the loss of inulin during storage and during preparation of the food. These findings allow quantification of the amount of inulin and oligofructose in the average daily western diet. The daily per capita intake is estimated to range from 1 to 10 g, depending on geographic, demographic, and other related parameters (age, sex, season, etc.). Inulin and oligofructose are not measured by classic methods of dietary fiber analysis and consequently are often not mentioned in food tables. Their significant contribution (1 to 10 g/d/per capita) to the dietary fiber fraction (recommended at 25 g/d/per capita) is not taken into account in any nutritional recommendations. In view of this, inulin and oligofructose deserve more attention, both in food composition tables and in diet or nutrition studies.

Diet↗

PubMed2003

Nutritional responses to the presence of inulin and oligofructose in the diets of domesticated animals: a review.

Inulin and oligofructose are prebiotic oligosaccharides fermented in the large intestine. This article provides an extensive review of the effects of these oligosaccharides on gastrointestinal characteristics (microflora, pathogen control, epithelial cell proliferation, putrefactive compound production, fecal characteristics, and nutrient digestibility) and systemic metabolism of carbohydrates, nitrogen, lipids, and minerals in dogs, cats, horses, calves, pigs, poultry, and rabbits. In addition, intake of inulin and oligofructose and considerations in their supplementation to animal diets are discussed. Growth performance and meat production in livestock in response to inulin and oligofructose supplementation are addressed. Finally, the possible substitution of antibiotics with fructans in animal diets and directions for future research are presented.

Animal Feed↗

PubMed2006

Direct haplotype-specific DNA sequencing.

Determining haplotype-specific DNA sequence information is very important in a wide range of research fields. However, no simple and robust approaches are currently available for determining haplotype-specific sequence information. We have addressed this problem by developing a very simple and robust haplotype-specific sequencing approach. We utilise the fact that DNA sequencing polymerases are sensitive to 3'end mismatches in the sequencing primer. By using two sequencing primers with 3'end corresponding to the two alleles in a given SNP locus, we are able to obtain allele-specific DNA sequences from both alleles. We evaluated this direct haplotype-specific approach by determining haplotypes within the intron 2 sequence of the fructan-6-fructosyltransferase (6-ft) gene in Lolium perenne L. We obtained reliable haplotype-specific sequences for all primers and genotypes evaluated. We conclude that the haplotype-specific sequencing is robust, and that the approach has a potentially very wide application range for any diploid organism.

Base Sequence↗

PubMed1988

Production of acids from inulin by a mixed culture of rumen microorganisms.

The production of volatile fatty acids (VFA) and lactic acid from inulin (plant fructan with 2-1 bonds) in in vitro rumen fermentations was followed. The experiments were performed with inocula from wethers receiving two rations, with two pH regimes and using as inocula either whole rumen contents or the corresponding rumen fluid. The following results were obtained: The VFA production was higher when using inocula from hay and concentrate fed wethers. In contrast to this, the lactic acid production was higher when using inocula from hay fed wethers. The VFA production tends to be higher in a weakly acid medium than in a neutral medium. Acetate-to-propionate molar ratio was lower at lower pH. The amounts and composition of the VFA were not different when using both types of inocula. Thus, we believe that micro-organisms colonising plant fibres obviously do not play an important role in inulin splitting.

Animal Feed↗

PubMed2001

Inulin and oligofructose as dietary fiber: a review of the evidence.

This critical review article examines the composition and source of inulin and oligofructose, the physiological effects of their consumption, and how these materials relate to the concept of dietary fiber. Inulin and oligofructose are fructans extracted on a commercial basis from the chicory root. Inulin has been defined as a polydisperse carbohydrate material consisting mainly, if not exclusively, of beta (2-1) fructosyl-fructose links ranging from 2 to 60 units long. Native chicory inulin has an average degree of polymerization (DP) of 10 to 20, whereas oligofructose contains chains of DP 2 to 10, with an average DP of 4. While a universally accepted definition for dietary fiber does not exist, it is generally agreed that this term includes saccharides (+ lignin) that are not hydrolyzed or absorbed in the upper part of the gastrointestinal tract. These materials reach the colon, where they may be totally fermented, partially fermented, or remain unfermented. In addition, fibers contribute to fecal bulking. Inulin and oligofructose are not digested in the upper part of the gastrointestinal tract or are they absorbed and metabolized in the glycolytic pathway, or directly stored as glycogen like 'sugars' or starches. None of the molecules of fructose and glucose that form inulin and oligofructose appear in the portal blood. These materials are quantitatively fermented by the microflora of the colon; further, it has been demonstrated that this fermentation leads to the selective stimulation of the growth of the bifidobacteria population. After reviewing their chemistry, origin, and physiological effects, it is the opinion of the authors that inulin and oligofructose are dietary fiber. They share the basic common characteristics of dietary fibers, that is, saccharides of plant origin, resistance to digestion and absorption in the small intestine, and fermentation in the colon to produce short-chain fatty acids that are absorbed and metabolized in various parts of the body. Moreover, this fermentation induces a bulking effect.

Animals↗

PubMed2000

Prebiotics and probiotics: are they functional foods?

A probiotic is a viable microbial dietary supplement that beneficially affects the host through its effects in the intestinal tract. Probiotics are widely used to prepare fermented dairy products such as yogurt or freeze-dried cultures. In the future, they may also be found in fermented vegetables and meats. Several health-related effects associated with the intake of probiotics, including alleviation of lactose intolerance and immune enhancement, have been reported in human studies. Some evidence suggests a role for probiotics in reducing the risk of rotavirus-induced diarrhea and colon cancer. Prebiotics are nondigestible food ingredients that benefit the host by selectively stimulating the growth or activity of one or a limited number of bacteria in the colon. Work with prebiotics has been limited, and only studies involving the inulin-type fructans have generated sufficient data for thorough evaluation regarding their possible use as functional food ingredients. At present, claims about reduction of disease risk are only tentative and further research is needed. Among the claims are constipation relief, suppression of diarrhea, and reduction of the risks of osteoporosis, atherosclerotic cardiovascular disease associated with dyslipidemia and insulin resistance, obesity, and possibly type 2 diabetes. The combination of probiotics and prebiotics in a synbiotic has not been studied. This combination might improve the survival of the bacteria crossing the upper part of the gastrointestinal tract, thereby enhancing their effects in the large bowel. In addition, their effects might be additive or even synergistic.

Bifidobacterium↗