Colonie Function and Fermentation Consuming High Fiber Diets1

in Men

S. E. FLEMING, D. MARTHINSEN ANDH. KUHNLEIN2 Department of Nutritional Sciences, University of California, Berkeley, CA 94720 ABSTRACT The relationships between fermentation in the gut and colonie func tion were studied by using data derived from a human metabolic study. Five healthy men were fed diets that were fiber-free or contained cellulose, xylan, pectin or corn bran. Fermentation was assessed by measuring the excretion of flatus gas, volatile fat ty acids (VFA) in feces and fecal pH. Colonie function was assessed by measuring tran sit time, fecal frequency, fecal output and fecal composition. Fibers that were only marginally fermented included cellulose and corn bran, and these fibers caused large fecal output, frequent defecations and prolonged the residence time of digesta in the gut; feces contained high levels of neutral detergent fiber (NDF); neither diet influ enced fecal pH nor flatus gas excretion; and only corn bran increased VFA excretion in feces. Fibers that were mostly fermented in the gut included xylan and pectin, and these fibers did not influence fecal frequency or fecal output, but they did decrease transit time; both diets caused high levels of flatus gas to be excreted; pectin caused higher VFA excretion in feces and lower fecal pH. The excretion of VFA in feces was highly correlated with total fecal output, and high levels of VFA were associated with low flatus gas excretion. Mutagenic activity in feces was negligible for all subjects on all diets as measured by the fluctuation test. J. Nutr. 113: 2535-2544, 1983. INDEXING KEY WORDS flatus short-chain fatty acids . volatile fatty acids pectin xylan corn bran cellulose fecal pH mutagens neutral detergent fiber human fecal output

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The incorporation of fibers into diets is currently being encouraged because of reported desirable effects on health. Dietary fibers are widely distributed in foods and are highly variable in both chemical and physical characteristics (1, 2). Some dietary fibers such as cellulose and cereal brans contain insoluble, complex carbohydrates, They have been shown to increase fecal output and, in some cases, to normalize intestinal transit time (3-5). These effects are thought to alleviate constipation and diarrhea, and to reduce intracolonic pressure so that the risk of diverticular disease is lessened (6, 7). Other dietary fibers such as
Pectin and plant gUmS are Water Soluble and L 1-X1.1 n ' ' r IV. -ihave little mtlUenCe On teca! Characteristics , ,

a glucose challenge test (10) so may be effective in treating some diabetic patients. One of the major limitations to including fibers in diets is the intestinal discomfort which often results. Anecdotal evidence indicates that distension and flatulence are undesirable side effects of consuming high levels of dietary fiber. This has been confirmed by a few studies. In particular, wheat bran was shown to increase the excretion of fermentation-produced gases such as hydrogen in breath (11). Oat gum was
Ap, g American Inst-tute ofNutri"on' Receiwd torpub"cation 12
'Supported in part by National Institutes of Health grant HOI AM 10202, U.S. Department of Agriculture grant 12-14-5001-2,a donation by the QuakerOats Co., Chicago, IL, and by Natural Science and EngineerPresent address: Division of Human Nutntion, University of British

(8, 9). However,

i t

'_

they appear
* rr

to reduce the

"g Research council gram A-JUS.

blood glucose and serum insulin responses to

Columbia, Vancouver, B.C., Canada.

2535

2536

FLEMING, MARTHINSEN AND KUHNLEIN

also shown to increase flatus excretion (12) and, recently, we reported that pectin and xylan also caused high levels of flatus gases to be excreted (13). In addition to these gases, the microorganisms also produce volatile fatty acids (VFA) including acetate, propionate, butyrate and valerate. Some dietary fibers have been reported to increase the quantity of VFA that are excreted in feces (14, 15). These compounds provide utilizable energy for the host and also may contribute to fecal output and to diarrheal diseases since they are an important factor governing the rates of sodium and water absorption within the colon (16). In this study, we investigated the mecha nism by which dietary fibers influence coIonic function. We hypothesized that since dietary fibers are fermented to various ex tents by intestinal microbes, it is the degree of fermentation and the products of this fermentation that largely determine the influence a fiber will have on colonie func tion. To investigate this hypothesis, three purified fibers and one cereal fiber were fed to healthy men. Gastrointestinal parameters such as transit time, fecal frequency, fecal output and fecal composition were mea sured. Additionally, the excretion of fer mentation products such as hydrogen, car bon dioxide and methane were measured in breath and flatus, and VFA were measured in feces. The excretion of fermentation products was correlated to the measures of gastrointestinal function in order to investi gate their interrelationships.

a sequence of six test diets. Two diets were basal, fiber-free formulations providing either 100% (basal) or 85% of the energy (low E) requirement for each subject. The basal formulation provided 0.8 g protein per kilogram body weight daily from egg albu men (Seymour Foods, Inc., San Francisco, CA). Fat (butterfat and cottonseed oil, local suppliers) provided 30% of caloric require ment, and the remainder was provided by cornstarch (CPC International, Englewood Cliffs, NJ), dextramaltse [American Maize Products Co. (mfg), Monarch Foods, Bris bane, CA (supplier)] and sucrose in a ratio of 5:5:1. The fiber components were cellu lose (Alphacel, ICN Pharmaceuticals, Inc., Cleveland, OH), pectin (Sigma Chemical Co., St. Louis, MO), xylan (ICN Pharma ceuticals, Inc.) and raw corn bran (Quaker Oats, Chicago, IL). Cellulose, xylan and pectin were each fed at 0.5 g per kilogram body weight, and corn bran was fed at 0.6 g per kilogram, daily. The fibers were sub stituted for the other dietary carbohydrate constituents. These levels represent a reason able daily intake of dietary fiber, yet an individual would not consume this quantity of fiber from a single source as is fed here In addition, daily supplements of vitamins, minerals and choline were taken. Total fecal collections were made through out the study. Their handling has been previ ously described in some detail (15). Daily fecal excretion was determined, and 3-day collections were pooled and homogenized with a known quantity of water. At this time, the pH of the homogenate was determined (Beckman Expandometer Model 76A, combi MATERIALS AND METHODS nation electrode; Beckman Instruments, Palo Five normal-weight males (coded 01, 02, Alto, CA). 03, 05, 06) aged 21-32 years were selected Aliquots of the homogenate prepared from from the community.3 The subjects were fecal specimens produced during the last 3 assessed to be free of gastrointestinal dis days of each metabolic period were analyzed orders and intestinal parasites, and to have for solids. The weight of excreted solids was normal small bowel absorption as previously calculated by determining the moisture con described (13). The experimental design and tent (25 mm Hg, 60,18 hours). Nitrogen diet formulations were described in detail in content was determined by the microthis earlier publication, so these will be only Kjeldahl procedure (17). Fecal volatile fatty acids were determined by steam distillation briefly outlined. The 63-day metabolic study was divided and gas-liquid chromatography as previously into seven metabolic periods each of 9 day's described (15). Neutral detergent fiber (NDF) duration. All subjects received a basal, fiber-free diet during the first metabolic 3This research was evaluated by the Committee for the Protection of period. Subsequently, each subject received Humans in Research, University of California, Berkeley, CA.

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COLONIC FUNCTION AND FERMENTATION WITH FIBERS

2537

analysis was conducted by using the standard AACC (18) procedure. Breath and flatus gases were analyzed on days 2, 5 and 9 of each metabolic period. A detailed description of the procedures has been reported (13). Data collected on days 5 and 9 are reported here since the data from day 2 often reflected the gas excretion from the previous metabolic period. Intestinal transit time was determined by administering a nonabsorbable dye (50 mg FDC Blue No. 2) to the subjects and record ing the time of appearance and disappear ance of the dye in the feces. The fluctuation test for fecal mutagens was carried out as previously described (19-21). Salmonella typhimurium TA 98 was used to analyze for mutagenic activity. In preparation for anal ysis, the fecal homogenates were centriand were sterilized by filtration through 0.4and 0.2-fim filters (Acrodisc, American Sci entific Products, McGaw Park, IL). Several blanks, a standard sample and duplicate ex tracts at either 15 mg dry weight or 30 mg

dry weight were tested in each run. Throughout the study, comparisons be tween treatments were generally conducted by using a factorial design and analysis of variance. Differences among means were determined by significance of Tukey's Studentized range test and Duncan's multiplerange tests. Correlation coefficients were determined and regression analyses were performed when appropriate. The mutagenesis data were compared by using the chi-square test for unrelated samples (22).
RESULTSAND DISCUSSION Transit time and fecal output

The time required for appearance and then disappearance of the fecal dye marker was strikingly variable among subjects (fig. 1) but the fiber-containing diets tended to hasten appearance of the dye and this was particularly noticeable for the pectin-, xylan- and corn bran-containing diets. The time required for the dye to disappear was

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HOURS FROM INGESTION OF DYE Fig. 1 Transit time of a fecal dye marker in men fed fiber-free diets or diets containing various fibers. Appear ance time ()and disappearance time (O) are shown for each subject (codes 01, 02, etc.) and means (bars) are provided for each diet. The standard deviations for appearance and disappearance, respectively, of the dye are as follows: basal, 28, 36; low E, 55, 69; cellulose, 19, 30; xylan, 12, 55; pectin, 12, 31; corn bran, 14, 33.

2538

FLEMING, MARTHINSEN AND KUHNLEIN

more difficult to assess, but, with the excep tion of corn bran, the fiber-containing diets tended to extend the time necessary for total disappearance. Other researchers have re ported that cellulose and wheat bran, or products containing wheat bran, tend to reduce or to normalize transit time (4, 5, 14, 23, 24). However, cellulose did not reduce transit time in this study, and these differ ences may be due to differences in the tech niques of measurement. A combination of transit-time measuring techniques using, for example, a dye that marks the liquid phase and radioopaque pellets that mark the solid phase may have provided a better overall view of the influence that these fibers have on gut transit. Fecal output and fecal frequency ap peared to be closely associated. For in stance, both parameters were highest for corn bran, and showed decreasing values for cellulose and pectin (fig. 2). Both fecal frequency and fecal output for the corn bran diet were significantly greater than for the fiber-free or xylan diet. However, diet influenced fecal output to a greater extent than fecal frequency. The average weights per defecation were calculated to be 87.0 and 74.7 g for the fiber-free diets, whereas the values for the fiber-containing diets in creased in the order of xylan (108.3 g), pectin (118.6 g), cellulose (138.3 g) and corn bran (140.0 g). Fecal output and fecal frequency were also related when considered among the five subjects (fig. 3). For example, subjects 05 and 06 had significantly fewer defecations than subjects 01, 02, and 03, and subject 03 had, statistically, the greatest fecal output. The average weight per defecation was cal culated to be 181.1 g for subject 05, whereas the values for the other four subjects ranged from 73 to 128 g. Fecal composition Fecal dry matter excretion was closely re lated to total fecal weight (fig. 2). The fecal output was increased for corn bran and cel lulose over the fiber-free diets, and these re sults confirm those of other workers (3, 4, 22, 24, 25). Pectin caused a small increase in total fecal output and in fecal solids excre-

120 100 80 60 40 20
BASAL LOW E CELLULOSE XYLAN PECTIN CORN BRAN

2 |

LO I
0.8 I
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6 g
0.4 p

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DIET

Fig. 2 Effect of dietary fibers on the frequency of defecation, fecal output and fecal composition. The values are means of five subjects. These values were taken from specimens provided during days 7-9 of each metabolic period. Symbols: fecal frequency (solid), fecal output of water (open), fecal output of neutral detergent fiber, dry basis (dots), fecal output of solids other than NDF (hatched).

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tion. This effect has been reported in another study when 30 g pectin were fed daily (8), but other workers showed no ef fect when quantities of only 6 or 9 g pectin were administered (14, 26). The composition of the fecal solids was influenced by diet (fig. 2). In particular, negligible quantities of NDF were excreted for the fiber-free diets or for the xylan- and pectin-containing diets. This material must have been of endogenous origin since NDF was not measurable in these diets. For the cellulose- and corn bran-containing diets, larger quantities of NDF were excreted which indicates that much of this material is of dietary origin and survived passage through the intestinal tract. The NDF con tent of cellulose and corn bran were de termined to be 100 and 70%, respectively. Based on the average excretion of NDF in feces, 20 and 2% of ingested fiber from cel lulose and corn bran, respectively, were hydrolyzed during passage along the gut. This hydrolysis would have occurred via fermen tation. However, these values also include any endogeneous material that is measur able by the NDF assay. Therefore, these values will underestimate dietary NDF dis appearance along the gut.

COLONIC FUNCTION AND FERMENTATION WITH FIBERS

2539

excretion, but xylan and glucose (two of the components of the constituent polymers) re 1.2 o o leased by TFA hydrolysis appeared to repre sent from 10-40% of intake, so that 60-90% i.o of ingested xylan must have been fermented. E Analyses for fecal content of glucoronic acid 0.8 I were not conducted, but other researchers reported complete fermentation of 30 g O 0.6 g pectin per day (8). Thus, the 28-42 g pectin 3 eaten, daily, was probably fermented.
0.4 e _l Ul

Fecal output and fermentation

in the colon

0.2

01

02

03

05

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SUBJECT CODE

Fig. 3 Variability among subjects in frequency of defecation, fecal output and fecal composition. The values are means of all six diets, and these values were taken from specimens provided during days 7-9 of each metabolic period. Symbols: fecal frequency (solid), fecal output of water (open), fecal output of neutral detergent fiber, dry basis (dots), fecal output of solids other than NDF (hatched).

The average weight of non-NDF solids that were excreted ranged from 14 to 20 g, daily, for four fiber-containing diets (fig. 2). This material constituted only 12-13% of fecal weight for the cellulose- and corn bran-containing diets and 20-23% of fecal weight for all other diets. The subjects showed a fourfold difference in the amount of non-NDF solids excreted (fig. 3) and this material constituted 12-23% of fecal weight by subject. This material will be composed of sloughed intestinal cells, bacterial mass, undigested dietary constituents and endogeneous secretions. Earlier, Olson et al. (27) reported on the quantities and types of carbohydrates that were excreted in feces following direct hy drolysis with trifluoroacetic acid (TFA). From the fecal carbohydrate and NDF data presented here, great variability in fermen tation of fibers in the gastrointestinal tract is evident. Cellulose and corn bran were large ly excreted in the feces, and the carbohy drate profile of feces after TFA hydrolysis indicated the carbohydrate profile of the ingested fiber source (27). Xylan disap pearance could not be monitored by NDF

We previously reported that some of these dietary fibers influenced the quantities of VFA that were excreted in feces (15). These VFA are produced via microbial fermenta tion in the colon and have been thought to increase fecal output by increasing fecal water excretion due to their hyperosmolarity. From our data, it appears that VFA ex cretion was more closely related to total fe cal output (fig. 4, T = 0.83) than to fecal output of dry matter (r = 0.68). Much of the increase in VFA that occurred when corn bran and pectin were fed was due to higher excretion of acetic acid (15). The

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Fig. 4 Relationship of fecal output and excretion of volatile fatty acids (VFA) in feces. Values for each subject are provided (A, 01; D, 02; O, 03; , 05; T, 06) for all diets and these values were taken from speci mens provided during days 7-9 of each metabolic period. The regression equation for the best-fit curve was calculated by using x = fecal output (grams/day), y - VFA excretion in feces (grams/day), m = slope; b = /-intercept.

2540

FLEMING, MARTHINSEN AND KUHNLEIN

quantity of acetic acid excreted in feces is highly correlated with total fecal output (r = 0.81) and fecal output of dry matter (r = 0.70). Thus, our results show that VFA excretion is more closely related to the out put of fecal water than of fecal solids. However, when expressed as a percentage of fecal water, the concentration of VFA was not constant among subjects and diets (data not shown), which indicates that factors such as the diffusibility of VFA within the lumen or rate of absorption from the colon are likely influenced by lumenal contents and may vary with subject. The pH of the fecal homogenates were generally neutral or slightly alkaline (fig. 5). Only feces produced during pectin con sumption had a lower pH than during con sumption of the fiber-free diets, and the diet means were 6.8 and 7.3, respectively. Other workers were able to reduce fecal pH by feeding five to seven oranges daily to chil dren (28). However, dietary fiber in the form of wheat bran did not influence fecal pH in either of two studies (28, 29). Simi larly, we showed that corn bran had no ef fect on fecal pH (fig. 5). Our data show no association between fecal pH and VFA excretion. In particular, nonsignificant correlation coefficients were determined between fecal pH and daily VFA excretion (r = -0.38), concentration of VFA in feces (r = - 0.13) or concentra tion of VFA in fecal water (r = - 0.18). As suming that fecal pH reflects the pH of the lumenal contents of the colon, our data show that VFA can be accumulated in the lumen without causing simultaneous acidi fication. Other workers reported a lack of correlation between intestinal pH and VFA concentration in the pig (30). In fact these workers reported that the contents of the cranial half of the stomach showed a slight increase in pH when the highest levels of VFA and lactic acid were also measured. Similarly, Ruppin et al. (31) reported an in crease in pH with increasing propionate concentration in the intestinal lumen of humans. Bicarbonate is secreted during VFA absorption, and there is HCO3" ac cumulated in the contents as a result of nonionic VFA transport across the mucosa (31-33). These changes are probably

8.0 7.5 2 7.0 U 6.5 6.0

BASAL LOW CELLE ULOSE XYLAN PECTIN CORN BRAN

DIET

Fig. 5 Effect of dietary fibers on the pH of fecal homogenates. Values represent the mean pH calcu lated for the five subjects, and the range of values is represented by the bar.

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responsible for the constant or slight increase in pH that is observed in the presence of an increase in VFA absorption. If VFA absorption is accompanied by an increase in pH, then it is possible that a lowering of pH, as occurred with pectin in this study, may be associated with an increase in VFA concentration without a concomitant increase in absorption. Further studies are necessary to determine such an effect. The colonie environment and gas excretion High levels of VFA in feces (15) were as sociated with low levels of excretion of fer mentation gases in flatus (13). For example, when the subjects excreted high levels of VFA while consuming corn bran, they ex creted negligible levels of hydrogen, carbon dioxide and methane in flatus (fig. 6). From fecal analysis, we determined that corn bran, xylan and pectin were fermented in the gut, and these three diets showed an in verse relationship between VFA excretion and flatus gas excretion. Low levels of VFA and gases were excreted while consuming the fiber-free or cellulose-containing diets, and these diets likely provided little fer mentable substrate. From these data we can speculate that high VFA output may be inhibitory to the production of flatus gases. Alternatively, there may be competition among the microbial populations for fer-

COLONIC FUNCTION AND FERMENTATION WITH FIBERS

2541

4.0 cc CM 3.5

3.0
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(P > 0.05). None of the fibers fed in the previous study or in this study enhanced fecal mutagenesis activity as measured by the fluctuation test. Low levels of mutagens were previously reported in feces of persons consuming semipurified diets (20). Interrelationships and fermentation of colonie function

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EXCRETED

IN FECES (g/DAY)

Fig. 6 Relationship of the excretion of volatile fatty acids (VFA) in feces and of fermentation gases (H2, CH4, COj) in flatus. The values are means of all five subjects for each metabolic period. These values were determined on fecal specimens provided during days 7-9 of each metabolic period and on flatus gases col lected and analyzed during 1200-1800 hours on days 5 and 9 of each metabolic period.

mentable substrate, and VFA may be pro duced in preference to gases. As another explanation, the fast rate of intestinal transit noted for the corn bran diet may have dis couraged the utilization of VFA for gas production if the residence time was of in sufficient duration for this secondary fer mentation step. Fecal mutagenic activity The fluctuation test, used to quantify fecal mutagens, showed very low levels of activity for all subjects on all diets. This is evidenced from the raw data, which are presented (table 1) so that mutagenesis researchers can use them for alternate statis tical treatment (34). The ratio of the total number of revertant to nonrevertant tubes was calculated in order to compare the dif ferent diet periods (table 2). These data showed that the diet periods did not differ

Two of the dietary fibers, xylan and pec tin, were largely fermented in the gut since neither total fecal output nor output of solids was increased when these fibers were incorporated into the diet. Additionally, these fibers caused high levels of gas to be excreted. That VFA excretion was not in creased in accordance with the greater de gree of fermentation may be due to the ability of the microbes to utilize VFA to produce gases as secondary products. The other two fibers, cellulose and corn bran, were largely excreted in the feces and, therefore, were fermented to only a small degree by the intestinal microbes. As a con sequence, these fibers increased fecal output and fecal frequency, but did not cause gases to be produced as a result of fermentation. Corn bran stimulated VFA excretion and also caused the largest fecal output. This was accounted for by an increase in the ex cretion of moisture. Excretion of VFA has been previously associated with a hyperosmolar condition in the gut. Therefore, in addition to corn bran increasing the output of fecal solids due to its unfermentable constituents, the VFA that are produced caused a further increase in the moisture content of the feces. From these results, we see that the effect that dietary fibers have on colonie function is largely determined by their fermentability in the gut. If fermentable, the fibers will have a negligible influence on defecation, and may cause intestinal gas production. On the other hand, unfermented fiber increases fecal output. Corn bran represented a unique fiber source since it influenced defe cation in a manner that was representative of an unfermentable fiber, yet it was the only fiber to stimulate VFA excretion and fecal water output, but did not cause intesti nal gas to be increased. These effects make

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2542

FLEMING, MARTHINSEN AND KUHNLEIN TABLE l Number of revertant tubes' determined for fecal extracts Subject

Diet

01 29.5, 32.5" 32.5 26.5 28.5 32.5 29.0, 32.04

02

03

05

06 25.0, 18.5' 26.0 21.5 21.0 19.0 28.5

Basal Low E Cellulose Xylan Pectin Corn Bran

Mean values using 15 mg fecal extract 28.5, 28.02 22.03 21.0, 23.52 19.5, 25.5* 29.5 25.5 25.5, 22.04 18.5 20.5 30.5, 21.04 NE3 21.0 NE 21.5 21.0 17.0 32.5 16.0, 23.54

Mean values using 30 mg fecal extract 29.0229.031.032.024.040.5, 28.0223.535.0, 29.5233.027.5NE28.020.5, BasalLow 25.0431.020.5, ECelluloseXylanPectinCorn 5422.5NE33.519.0324.0, 33. 30.536.025.521.5, bran31.5, 37.0432.0,

02
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24.5427.5,30.533.024.522.025.028.

'Mean of two determinations, number of revertant tubes out of 50. 2The basal diet was repeated so values are given for both periods. 3No extract was available for determination. 4Repeated assays.

it a very desirable source of dietary fiber since it causes a large fecal output which is associated with reduced incidence of colon disease, and the VFA that are produced may provide a continuous source of energy to the host following the initial postprandial ab sorption.

CONCLUSIONS

Dietary fibers vary in their effects on colonie function and in their fermentability by intestinal microbes. To evaluate the influ ence of this fermentation on colonie func tion, we fed large quantities (0.5 g/kg body weight) of several sources of fiber and simul taneously measured parameters of gastro intestinal function and the excretion of fer mentation products. The purified fiber, cellulose, and the natural fiber, corn bran, which are com posed largely of insoluble dietary fibers, caused similar effects on gastrointestinal function and fermentation. When these dietary fibers were consumed, the subjects generally showed large fecal output, fre quent defecations, and a prolonging of the residence time of digesta within the GI tract

when determined by using a dye marker. Neither cellulose nor corn bran significantly influenced fecal pH nor flatus gas excretion. Fecal excretion of NDF and TFA hydrolyzable carbohydrates was high and the con centration of VFA in feces was low for the two diets. However, total VFA excretion over a given time period was high during corn bran consumption and the fecal components had a dilution effect on the fecal VFA con centration. These data indicate that cellu lose and corn bran remained largely unfermented in the gut and, thus, increased fecal output. The purified dietary fibers, xylan and pectin, which are composed of water soluble
TABLE 2 Number of revertant tubes and nonrevertant summed for each diet period Diets Revertant tubes (RT) Nonrevertant tubes (NRT) tubes

Ratio (RT:NRT)

BasalLow ECelluloseXylanPectinCorn

bran10676436515044307489335575494963706521.141.151.191.021.161.15

COLONIC FUNCTION AND FERMENTATION WITH FIBERS

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constituents, did not influence fecal output ble, and to the many staff members who or frequency of defecation, but they did were involved at each step. increase the rate of transit along the GI tract and increased excretion of intestinal gases. LITERATURE CITED Fecal NDF excretion was low, but TFA hydrolyzable carbohydrates were excreted in 1. Eastwood, M. A. & Kay, R. M. (1979) An hy feces during xylan consumption. Neither pothesis for the action of dietary fiber along the gastrointestinal tract. Am. J. Clin. Nutr. 32, fiber caused a large quantity of VFA to be 364-367. excreted in fces,but fecal pH was lowered 2. Stevens, B. J. H. & Selvendhan, R. R. (1981) A during pectin feeding. These data indicate comparison of the compositions of dietary fiber that xylan and pectin were mostly fer from some cereal and vegetable products in rela tion to observed effects in faecal weights. mented in the gut and, thus, the fermenta Lebensm. Wiss. Technol. 14, 301-305. tion products, gases and VFA, were respon 3. Eastwood, M. A., Kirkpatrick, J. R., Mitchell, sible for changes in colonie function. W. D., Bone, A. & Hamilton, T. (1973) Ef The excretion of VFA in feces was highly fects of dietary supplements of wheat bran and correlated with total fecal output and excre cellulose on faeces and bowel function. Br. J. Nutr. 4, 392-394. tion of dry matter. Neither total fecal VFA 4. Stephen, A. M. & Cummings, J. H. (1980) excretion nor VFA concentration was re Mechanism of action of dietary fibre in the hu lated to fecal pH. man colon. Nature (London) 284, 283-284. High VFA excretion was associated with 5. Harrison, R. J., Leeds, A. R., Bolster, N. R. & Judd, P. A. (1980) Exercise and wheat bran: the excretion of low levels of flatus gases. effect on whole gut transit. Proc. Nutr. Soc. 39(1), However, when only low levels of VFA were 22A. excreted, flatus gas excretion was variable 6. Findlay, J. M., Smith, A. N., Mitchell, N. D., This suggests that the gas-producing organ Anderson, A. J. B. & Eastwood, M. A. (1974) isms are active under these conditions and Effects of unprocessed bran on colon function in normal subjects and in diverticular disease. can produce gases via fermentation provid Lancet 1, 146-149. ing that suitable substrates are available 7. Brodribb, A. M. J. & Humphreys, D. M. (1976) Corn bran caused low levels of flatus gas but Diverticular disease: three studies. Br. Med. J. 1, 424-430. high levels of VFA to be excreted, whereas 8. Cummings, J. H., Southgate, D. A. T., Branch, pectin caused VFA to be preferentially ex W. J., Wiggins, H. S., Houston, H., Jenkins, creted by most subjects. D. J. A., Jivraj, T. & Hill, M. J. (1979) The Mutagenic activity was negligible for all digestion of pectin in the human gut and its effect subjects on all diets. Consequently, it ap on calcium absorption and large bowel function. Br. J. Nutr. 41, 477-485. pears that these dietary fibers did not stimu 9. Cummings, J. H. & Stephen, A. M. (1980) late the excretion of mutagenic compounds The role of dietary fiber in the human colon. in feces. Can. Med. Assoc. J. 123, 1109-1114. The effect that dietary fibers have on 10. Jenkins, D. J. A., Leeds, A. R., Gassull, M. A., colonie function appears to be largely deter Cochet, B. & Alberti, G. M. M. (1977) De mined by their fermentability. The fibers crease in postprandial insulin and glucose concen trations by guar and pectin. Ann. Intern. Med. that were fermentable caused intestinal gas 86, 20-23. to be excreted and had no effect on defeca 11. Bond, J. H. & Levitt, M. D. (1978) Effect of tion or intestinal transit time. The fibers dietary fiber on intestinal gas production and that were largely unfermented increased small bowel transit time in man. Am. J. Clin Nutr. 31, S169-S174. fecal output but did not increase gas excre tion. One fiber, corn bran, was unique by 12. Meyers, S. & Calloway, D. H. (1977) Gastro intestinal response to oat and wheat milling func increasing fecal output while simultaneous tion in older women. Cereal Chem. 54, 110-119. ly stimulating VFA to be produced as a re 13. Marthinsen, D. & Fleming, S. E. (1982) Ex sult of fermentation. cretion of breath and flatus gases by humans con
ACKNOWLEDGMENTS suming high fiber diets. J. Nutr. 112, 1133-1143. 14. Spiller, G. A., Chernoff, M. C., Hill, R. A., Gates, J. E., Nassar, J. J., Shipley, E. A. (1980) Effect of purified cellulose, pectin, and a low-residue diet on fecal volatile fatty acids, transit time, and fecal weight in humans. Am. J. Clin. Nutr. 33 754-759.

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The authors are indebted to the men who sacrificed much to make such a study possi

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FLEMING, MARTHINSEN AND KUHNLEIN 26. Stasse-Wolthuis, M., Albers, H. F. F., van Jeversen, J. G. C., de Jong, J. W., Hautvast, J. G. A. J., Hermus, R. J. J., Katan, M. B., Brydon, G. W. & Eastwood, M. A. (1980) Influence of dietary fiber from vegetables and fruits, bran or citrus pectin on serum lipids, fecal lipids and colonie function. Am. J. Clin. Nutr. 33, 1745-1756. 27. Olson, A. C., Gray, C. M., Chi,M.-C & Fleming, S. E. (1983) Cellulose, xylan, corn bran, and pectin in the human digestive process. In: Uncon ventional Sources of Dietary Fiber, ACS Symposium Series, No. 214 (Furda, L, ed.), p. 221, American Chemical Society, Washington, DC. 28. Walker, A. R. P., Walker, B. F. & Segal, I. (1979) Faecal pH value and its modification by dietary means in South African black and white schoolchildren. S. Afr. Med. J. 55, 495-498. 29. Stephen, A. M. & Cummings, J. H. (1981) The effect of wheat fiber on faecal pH in man. Gastroenterology 80, 1294 (abs.). 30. Argenzio, R. A., & Southworth, M. (1975) Sites of organic acid production and absorption in gastrointestinal tract of the pig. Am. J. Physiol. 228, 454-460. 31. Ruppin, H., Bar-Meir, S., Soergel, K. H., Wood, C. M. & Schmitt, M. G., Jr. (1980) Absorption of short chain fatty acids by the colon. Gastroenterology 78, 1500-1507. 32. McNeil, N. I., Cummings, J. H. & James, W. P. T. (1978) Short chain fatty acid absorp tion by the human large intestine. Gut 19, 819-822. 33. Roediger, W. E. W. & Moore, A. (1981) Effect of short-chain fatty acids on sodium absorption in isolated human colon perfused through the vascu lar bed. Dig. Dis. Sci. 26, 100-106. 34. Venitt, S. (1982) Mutagens in human feces: are they relevant to cancer of the large bowel? Mut. Res. 98, 265-286.

15. Fleming, S. E. & Rodriguez, M. A. (1983) In fluence of dietary fiber on fecal excretion of vola tile fatty acids by human adults. J. Nutr. 113, 1613-1625. 16. Argenzio, R. A. (1981) Short-chain fatty acids and the colon. Dig. Dis. Sci. 26(2), 97-99. 17. Association of Official Analytical Chemists (1975) Official Methods of Analysis, 12th ed., AOAC, Washington, DC. 18. American Association of Cereal Chemists (1970) Approved Methods of the AACC, AACC, St. Paul, MN. 19. Kuhnlein, U., Bergstrom, D. & Kuhnlein, H. V. (1981) Mutagens in feces from vegetarians and nonvegetarians. Mut. Res. 85, 1-12. 20. Kuhnlein, H. V. & Kuhnlein, U. (1980) Muta gens in feces from subjects on controlled formula diets. Nutr. Cancer 2, 119-124. 21. Kuhnlein, H. V., Levander, O. A., King, J. C., Sutherland, B. & Riskie, L. (1983) Dietary selenium and fecal mutagenicity in young men. Nutr. Res. 3, 203-209. 22. Siegel, S. (1956) Nonparametric Statistics for the Behavioral Sciences, McGraw-Hill, New York. 23. Cummings, J. H., Hill, M. J., Jenkins, D. J. A., Pearson, J. R., & Wiggins, H. S. (1976) Changes in fecal composition and colonie func tion due to cereal fiber. Am. J. Clin. Nutr. 29, 1468-1473. 24. Spiller, G. A., Shipley, E. A., Chernoff, M. C. & Cooper, W. C. (1979) Bulk laxative efficacy of a Psyllium seed hydrocolloid and of a mixture of cellulose and pectin. J. Clin. Pharmacol. 19, 313-320. 25. Southgate, D. A. T., Branch, W. }., Hill, M. J., Drasar, B. S., Walters, R. L., Davies, P. S. & McLean, B. I. (1976) Metabolic responses to dietary supplements of bran. Metabolism 25, 1129-1135.

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