Redefining dietary fibre: potentially a recipe for disaster.
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Biomedical subjects
Publications and source records attributed to H N Englyst.
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Prebiotics, as currently conceived of, are all carbohydrates of relatively short chain length. To be effective they must reach the cecum. Present evidence concerning the 2 most studied prebiotics, fructooligosaccharides and inulin, is consistent with their resisting digestion by gastric acid and pancreatic enzymes in vivo. However, the wide variety of new candidate prebiotics becoming available for human use requires that a manageable set of in vitro tests be agreed on so that their nondigestibility and fermentability can be established without recourse to human studies in every case. In the large intestine, prebiotics, in addition to their selective effects on bifidobacteria and lactobacilli, influence many aspects of bowel function through fermentation. Short-chain fatty acids are a major product of prebiotic breakdown, but as yet, no characteristic pattern of fermentation acids has been identified. Through stimulation of bacterial growth and fermentation, prebiotics affect bowel habit and are mildly laxative. Perhaps more importantly, some are a potent source of hydrogen in the gut. Mild flatulence is frequently observed by subjects being fed prebiotics; in a significant number of subjects it is severe enough to be unacceptable and to discourage consumption. Prebiotics are like other carbohydrates that reach the cecum, such as nonstarch polysaccharides, sugar alcohols, and resistant starch, in being substrates for fermentation. They are, however, distinctive in their selective effect on the microflora and their propensity to produce flatulence.
BACKGROUND: A chemically based classification of dietary carbohydrates that takes into account the likely site, rate, and extent of digestion is presented. The classification divides dietary carbohydrates into sugars, starch fractions, and nonstarch polysaccharides, and groups them into rapidly available glucose (RAG) and slowly available glucose (SAG) as to the amounts of glucose (from sugar and starch, including maltodextrins) likely to be available for rapid and slow absorption, respectively, in the human small intestine. OBJECTIVE: We hypothesize that RAG is an important food-related determinant of the glycemic response. DESIGN: The measurement of RAG, SAG, and starch fractions by an in vitro technique is described, based on the measurement by HPLC of the glucose released from a test food during timed incubation with digestive enzymes under standardized conditions. Eight healthy adult subjects consumed 8 separate test meals ranging in RAG content from 11 to 49 g. RESULTS: The correlation between glycemic response and RAG was highly significant (P < 0.0001) and a given percentage increase in RAG was associated with the same percentage increase in glycemic response. After subject variation was accounted for, RAG explained 70% of the remaining variance in glycemic response. CONCLUSIONS: We show the significance of in vitro measurements of RAG in relation to glycemic response in human studies. The simple in vitro measurement of RAG and SAG is of physiologic relevance and could serve as a tool for investigating the importance of the amount, type, and form of dietary carbohydrates for health.
We conducted a population-based case-control study among different ethnic groups in Hawaii to evaluate the role of various types and components of fiber, as well as micronutrients and foods of plant origin, on the risk of colorectal cancer. We administered personal interviews to 698 male and 494 female Japanese, Caucasian, Filipino, Hawaiian, and Chinese cases diagnosed during 1987-1991 with adenocarcinoma of the colon or rectum and to 1,192 population controls matched to cases by age, sex, and ethnicity. We used conditional logistic regression to estimate odds ratios, adjusted for caloric intake and other covariates. We found a strong, dose-dependent, inverse association in both sexes with fiber intake measured as crude fiber, dietary fiber, or nonstarch polysaccharides. We found inverse associations of similar magnitude for the soluble and insoluble fiber fractions and for cellulose and noncellulosic polysaccharides. This protective effect of fiber was limited to fiber from vegetable sources, with an odds ratio of 0.6 (95% confidence interval = 0.4-0.9) and 0.5 (95% confidence interval = 0.3-0.7) for the highest compared with the lowest quartile of intake for men and women, respectively. We found associations of the same magnitude for soluble and insoluble vegetable fiber, but no clear association with fiber from fruits or cereals. This pattern was consistent between sexes, across segments of the large bowel (right colon, left colon, and rectum), and among most ethnic groups. The effect of vegetable fiber may be independent of the effects of other phytochemicals, since the effect estimates remained unchanged after further adjustment for other nutrients. Intakes of carotenoids, light green vegetables, yellow-orange vegetables, broccoli, corn, carrots, bananas, garlic, and legumes (including soy products) were inversely associated with risk, even after adjustment for vegetable fiber. The data support a protective role of fiber from vegetables against colorectal cancer, which appears independent of its water solubility property and of the effects of other phytochemicals. The data also indicate that certain vegetables and fruits may be protective against this disease through mechanisms other than their fiber content.
The glycaemic index (GI) is an in vivo measurement based on the glycaemic response to carbohydrate-containing foods, and allows foods to be ranked on the basis of the rate of digestion and absorption of the carbohydrates that they contain. GI values are normalized to a reference amount of available carbohydrate and do no reflect the amounts of carbohydrate normally present in foods; for example, a food with a low content of carbohydrates will have a high GI value if that carbohydrate is digested and absorbed rapidly in the human small intestine. This is potentially confusing for a person wishing to control his or her blood glucose levels by the choice of foods. The rate and extent of starch digestion in vitro has been measured using a technique that classifies starch into three major fractions: rapidly digestible starch (RDS), slowly digestible starch (SDS) and resistant starch (RS). In addition, this technique gives a value for rapidly available glucose (RAG), which includes RDS, free glucose and the glucose moiety of sucrose. When the values for thirty-nine foods were expressed on the basis of the available carbohydrate content of these foods, highly significant (P < 0.001) positive correlations were observed between GI and both RDS and RAG. The measurement of RAG in vitro provides values for direct calculation of the amount of glucose likely to be rapidly absorbed in the human small intestine and, thus, to influence blood glucose and insulin levels. These values can be used to compare foods, as eaten, on an equal-weight basis. Food-table RAG values would allow simple calculation of the total amount of RAG provided by single foods, by whole meals and by whole diets. Studies are planned in which RAG and the glycaemic response in man will be measured for identical food products.
The digestion of four sources of resistant starch (RS) has been studied in twelve healthy volunteers who ate controlled diets for 15 d periods. RS from potato, banana, wheat and maize (17-30 g/d) was compared with a starch-free diet, a diet containing wheat starch that was fully digested in the small intestine, and with 18.4 g NSP from brand/d. RS increased stool wet weight by 1.6 g/d per g RS fed for potato, 1.7 for banana, 2.5 for wheat and 2.7 for maize, but this was significantly less than bran NSP at 4.9 g/g. RS was extensively digested in twenty-seven of thirty-four diet periods but five subjects were unable to break down one or two of the RS sources. Faecal N and energy excretion were increased. RS decreased NSP breakdown and RS2 (resistant starch granules) tended to prolong transit time. All forms of RS increased faecal total short-chain fatty acid excretion. RS2 (from potato and banana) gave greater proportions of acetate in faeces, and RS3 (retrograded starch from wheat and maize) more propionate. We have concluded that RS2 and RS3 are broken down in the human gut, probably in the colon although in 26% of cases this breakdown was impaired. RS exerts mild laxative properties, predominantly through stimulation of biomass excretion but also through some sparing of NSP breakdown.
The digestibility of the starch in plant foods is highly variable, and is dependent on a number of factors, including the physical structure of both the starch and the food matrix. An in vitro technique has been developed to categorize starch in plant foods according to its likely rate and extent of digestion in the human small intestine. The in vitro method provides values for rapidly digestible starch, slowly digestible starch and resistant starch (RS). In the present study values for the RS content of foods, as measured by the analytical technique, were compared with the recovery of starch from these foods when fed to healthy ileostomates. Nine ileostomy subjects were given a polysaccharide-free diet with a breakfast supplement, on eac of 2 d (two subjects) or 3 d (seven subjects), of biscuits made from wheat, potato or banana flours or from moist-heat-processed wheat or maize flours. RS intakes measured in vitro ranged from 8.5 to 15.0 g/d for the test biscuits, and mean starch recoveries in ileostomy effluent were 100.4 (n5, range 91-106)% of those values, but there was substantial variation between individuals. It is proposed that RS is defined as 'the sum of starch and starch-degradation products that, on average, reach the human large intestine'. The analytical method for the measurement of RS in vitro based on this definition is shown to provide an accurate prediction of the average amount of starch that is likely to escape complete digestion and absorption in the human small intestine.
Dietary intakes of starch and non-starch polysaccharides (NSP) have been estimated for a rural West African community. These people eat directly from shared bowls of cooked food, and so measurement of any individual's food intake is not possible. Recently developed methodology for estimating food intake under these circumstances and the analysis of samples for dietary polysaccharides are combined to yield estimates of intakes of about 375 g starch and 25 g NSP/d for adult males, with lower intakes for women and children, related to their smaller body weights. These intakes are not direct measurements but are extremely plausible when compared with values obtained for other societies where the complication of the shared food bowl does not impinge upon the measurements.
Dietary carbohydrate may be divided into monosaccharides and disaccharides (sugars), oligosaccharides [degree of polymerization (DP) 3-9], and polysaccharides (DP > 10). Their physiological properties and health benefits depend on the site, rate, and extent of their digestion or fermentation in the gut. Oligosaccharides are a diverse group of soluble carbohydrates, many of which are not digested by pancreatic enzymes. They are fermented in the colon and some have specific effects on bacteria. The major dietary polysaccharides are starch and nonstarch polysaccharides (NSPs). The digestion of starch depends on its physical form, the nature of the starch granule, and the effects of food processing. Starch may be rapidly digested, slowly digested, or resistant, the last passes into the colon for fermentation. The NSPs (cell wall polysaccharides) all resist digestion. They exert a physical effect in the upper gut, serving to moderate carbohydrate and possibly lipid absorption, whereas in the colon they affect bowel habit through fermentation, they affect epithelial cell metabolism, and, along with other fermented carbohydrates, they provide energy to humans.
Six subjects with ileostomies consumed five diets containing 61-164 g starch/d of which 0.4-34.8 g was resistant starch (RS). Ileal excretion of starch was 97% of that measured as dietary RS in vitro with no significant difference between RS fed and starch recovered on any of the test diets. Variation in starch excretion between subjects was partly due to differences in mouth-to-stoma transit time. In vitro fermentation of ileal effluent from RS-supplemented diets produced significantly more short-chain fatty acids, a higher molar proportion of butyrate (17% compared with 12%), and a lower concentration of ammonia compared with control subjects. These results indicate that the amount of starch that reaches the large intestine can be predicted from measurements in vitro for a wide range of RS intakes under normal eating conditions. They also support the hypothesis that RS, through fermentation, has distinctive influences on the colonic environment.
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Methods for the measurement of dietary fibre as non-starch polysaccharides (NSP) are described. A common enzymic removal of starch and acid hydrolysis of the NSP to their constituent sugars are followed by one of three alternative techniques, gas-liquid chromatography, high-performance liquid chromatography or spectrophotometry, for measurement of the released sugars. The results obtained by the three methods are in good agreement for a wide range of raw and processed foods. NSP compose approximately 90% of the plant cell-wall material and are therefore a good index of this material. Values for NSP therefore provide a good marker for a diet rich in fruit, vegetables and high-extraction cereal products associated with health and recommended in dietary guidelines. Values for total, soluble and insoluble NSP may be obtained with any of the end-point techniques, and the detailed information obtained from the chromatographic methods is useful in studies of the relationship between the intakes of various types of NSP and health. The causes of some potential interferences in the spectrophotometric assay, especially from processed foods, have been identified and eliminated. The rapid spectrophotometric version is suitable for food labelling purposes and for quality control, and the changes described have made it more robust.
A technique is described for measurement of the uronic acid constituents of non-starch polysaccharides (NSP) by high-performance liquid chromatography with pulsed amperometric detection using two sets of hydrolysis conditions. In one set of hydrolysis conditions the uronic acid-containing polymers are hydrolysed by prolonged treatment with sulfuric acid only; in the other, the hydrolysate obtained in the Englyst procedure for the measurement of dietary fibre as NSP is buffered to between pH 3.5 and 4 and the uronic acid-containing polymers present are depolymerized enzymically. The conditions for acid hydrolysis only or acid and enzymic hydrolysis of uronic acid-containing polymers optimized for a range of fruit, vegetable and cereal products. Values obtained for total uronic acids are very similar using either hydrolysis procedure, and are in good agreement with values obtained using the spectrophotometric assay as used in the Englyst procedure.
The plasma-cholesterol-lowering effects of some dietary legumes are now well established from animal and human studies, but the mechanism is not completely understood. The present study investigated the effect of baked beans (Phaseolus vulgaris) on steroid metabolism of hypercholesterolaemic pigs. Three groups of four pigs were studied: baseline (BL), normal pigs (NP) and those previously prepared with an ileo-rectal anastomosis to nullify the function of the large intestine (IR). All three groups were given a semi-purified control diet, with about 40% energy as fat (polyunsaturated:saturated fatty acid (P:S) ratio 0.3), supplemented with 10 g cholesterol/kg, for 14 d. Then IR and NP pigs were fed for 28 d on a diet supplemented with 10 g cholesterol/kg and 300 g baked beans/kg (dry-matter basis), so that the 40% contribution to energy from fat was maintained (P:S ratio 0.3). Group BL was fed on the control diet throughout. The intact pigs (NP) fed on baked beans showed considerable differences compared with the other groups, as follows: (a) reduced plasma cholesterol (NS); (b) higher concentration of cholesterol in bile (NS); (c) higher concentration of bile acids, especially secondary bile acids, in bile (P < 0.05); (d) reduced elimination of bile acids in faeces, especially secondary bile acids (P < 0.05); (e) higher excretion of coprostanol and lower elimination of cholesterol in faeces (P < 0.05). From these findings it is proposed that a baked-bean-enriched diet potentiates bacterial fermentation and steroid degradation in the large intestine and enhances conservation of bile acids and cholesterol within the enterohepatic circulation. The high concentration of bile acids and cholesterol in bile may thus promote feedback inhibition of hepatic cholesterol synthesis, and hence, reduce plasma cholesterol.
An improved method is described for the measurement of total, soluble and insoluble dietary fibre as non-starch polysaccharides (NSP). An established procedure is modified to allow more rapid removal of starch and hydrolysis of NSP. In its present form the procedure is simpler and more robust than those previously published. In the modified method starch is removed enzymically within 50 min and NSP is precipitated with ethanol and then hydrolysed by treatment with sulfuric acid for 2 h. The constituent sugars can in turn be measured by gas-liquid chromatography, high-performance liquid chromatography or more rapidly by colorimetry. The improved procedure described here for the removal of starch and hydrolysis of NSP applies to all three techniques, but only the method for measurement of sugars by gas-liquid chromatography is described here in full.
For nutritional purposes, starch in foods may be classified into rapidly digestible starch (RDS), slowly digestible starch (SDS) and resistant starch (RS). RS may be further divided into three categories according to the reason for resistance to digestion. A method is reported for the measurement of total starch, RDS, SDS, RS and three RS fractions in starchy foods, using controlled enzymic hydrolysis with pancreatin and amyloglucosidase. The released glucose is measured by colorimetry, using a glucose oxidase kit. Values for RDS and SDS in foods obtained by the method reflect the rate of starch digestion in vivo. Values for RS are similar to the amounts of starch escaping digestion in the small intestine of ileostomates, and are a guide to the amounts of starch likely to enter the colon for fermentation. Results are given for a number of starchy foods.
Starch, not fibre, is probably the major substrate for fermentation in the human colon. However, quantitating the amount of starch that resists pancreatic amylase and thus escapes digestion in the small bowel is difficult. A number of techniques have been employed in man and are reviewed here, including direct intubation of the ileum, the ileostomy model, and breath studies. The results of a series of studies of the digestion of starch from potato and banana are reported. When fed to ileostomy patients, 3% of hot potato starch and 12% of cold potato starch were resistant to digestion, as was 75% of banana starch. In feeding experiments with healthy volunteers none of the starch was recoverable in faces, indicating its complete fermentation in the colon. Breath H2 measurements after test meals of these starches indicated that only 2-5% of potato starch and 7-12% of banana starch was fermented. A single blood acetate measurement timed to coincide with peak breath H2 was not useful. However, a number of problems with breath H2 studies are discussed, and it is suggested that either ileal intubation or the ileostomy model are the most reliable techniques presently available, with serial blood acetate determinations also potentially valuable. Overall on Western diets, approximately 10% of all starch is probably resistant starch.
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