Dietary Energy Source and Supplements in Broiler Diets Containing Defatted Rice Bran

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1 2002 Poultry Science Association, Inc. Dietary Energy Source and Supplements in Broiler Diets Containing Defatted Rice Bran Adrizal,* S. Ohtani,,1 and M. Yayota *The United Graduate School of Agricultural Science, Gifu University, Gifu , Japan; and Faculty of Agriculture, Gifu University, Gifu , Japan Primary Audience: Nutritionists, Researchers, Feed Manufacturers SUMMARY Increasing the concentration of defatted rice bran (DRB) in the diet will increase the concentration of energy sources such as fat (oil) and carbohydrate because DRB contains relatively lower energy compared with the most common energy source, grain (corn). An experiment with broiler chicks was conducted to determine the influence of a high level of fat versus a high level of carbohydrate on the nutritional value of a diet containing 30% DRB. In order to improve the digestibility of fat and to anticipate the negative effect of increasing the concentration of nonstarch polysaccharide in the DRB during the early age of broiler chicks, two supplements [a bile salt (sodium taurocholate) or a fiber-degrading mixture of enzymes (Grindazyme GP 5000)] were included. The protein and fat digestibilities and ME n value were greater with high-fat diets compared with high-carbohydrate diets. These effects of the high-fat diet were reflected in superior growth and feed intake of chicks from 1 to 14 d of age. The supplementation of the DRB diets with the bile salt was of greater benefit for chick performance than the inclusion of a mixture of fiber-degrading enzymes, especially with high-fat diets. Key words: energy source, supplement, defatted rice bran, broiler chick 2002 J. Appl. Poult. Res. 11: DESCRIPTION OF PROBLEM The use of cereals or their by-products in broiler diets has been drawing much attention recently because these ingredients contain considerable amounts of nonstarch polysaccharides (NSP), substances that may impair nutrient utilization. A study performed by Annison et al. [1], however, showed that a diet containing 6% rice bran-soluble NSP contributed positively to dietary energy values in broiler chickens fed the diet from 28 to 34 d of age. A subsequent study by Farrell and Martin [2] indicated that feeding broiler chickens up to 23 d of age a diet containing 20% of rice bran did not depress gain and feed utilization. But, when the dietary concentration of rice bran was increased to 40%, gain and feed conversion ratio was adversely affected and no beneficial effect was observed after fiber-degrading enzyme supplementation [2]. Up to now, the optimum level of defatted rice bran (DRB) in broiler chick diets relating to the potential negative effect of NSP of DRB has not been clearly elucidated. Because information on the effect of NSP isolated from rice bran on the early growth of chicks is lacking, three preliminary studies 1 To whom correspondence should be addressed: sohtani@cc.gifu-u.ac.jp.

2 ADRIZAL ET AL: ENERGY SOURCE IN RICE BRAN DIET 411 were conducted [3, 4, 5]. In the first study [3], depressed pancreatic lipase activity was noted in chicks fed 7% total NSP extracted from DRB. This effect of total NSP was ameliorated by xylanase supplementation of the diet. In the second study [4], the negative effect of NSP on lipase activity disappeared when a diet containing 15% of total NSP, supplied by intact DRB, was supplemented with a bile salt. This result was confirmed by the results of the third study [5]. The use of fiber-degrading enzymes in diets containing NSP has been shown to decrease the viscosity of digesta and increase the ME value of the diet, evidently by increasing nutrient digestibility [6, 7, 8, 9]. Mateos and Sell [10] reported that nutrient digestibilities and digestive enzyme activities were altered when chicks were fed a high-carbohydrate diet in comparison with a high-fat energy diet. This finding indicates that providing an NSP diet with energy of fat or carbohydrate origin may result in different efficiency of energy utilization. The present study was designed to determine the influence of energy source (high fat versus high carbohydrate) in diets containing 30% DRB on nutrient digestibility and performance of broiler chicks. In addition, the influence of bile salt or fiber-degrading enzyme supplementation of the 30% DRB diets was also evaluated because these two supplements were noted to influence nutrient digestibility of the NSP diets [3, 4, 5]. MATERIALS AND METHODS One hundred twenty 1-d-old male broiler chicks (Chunky) were obtained from a local commercial hatchery and distributed into 12 pens with 10 chicks per pen. Two kinds of supplements (sodium taurocholate [11] and a fiber-degrading enzyme mixture [12]) were combined with two major sources of dietary energy (oil and carbohydrate) in the diets containing 30% DRB. Bile salt (sodium taurocholate) was used at 0.5 % of the diet, whereas fiber-degrading enzyme (Grindazyme GP 5000; contained 12,000 FXU of xylanase, 5,000 BGU of β-glucanase, and 10 FPU of pectinase/g of granules) derived from a selected strain of fungus was included at 0.05% of the diet. The high-fat diets that were supplemented with bile salt or enzyme contained and 10.43% blended oils [soy oil + palm oil (1:1), respectively]. The counterpart high-carbohydrate diets contained and 39.32% soluble starch [13] and 4.77 and 4.50% blended oil, respectively. Diets were formulated to meet the nutrient requirements of broiler chicks [14] (Table 1). Feed and water were provided to chicks for consumption ad libitum from 1 to 14 d of age. Feed intake, body weight gain (BWG), and feed:gain (F:G) ratio were recorded from 1 to 7 d of age and 8 to 14 d of age. After recording BW and collecting excreta on Days 7 and 14, three birds per pen were killed by decapitation [15], and samples of pancreatic tissue for lipase analysis [16] were taken. Ileal digesta (from yolk stalk to ileocecal junction) was collected by flushing 10 ml of deionized water from a syringe through the ileum portion into a plastic tube. The tubes were capped and stored at 30 C for later analyses of DM [17], fat, and protein [18]. Celite (acid-insoluble ash, AIA) [13] was used at 2% of the diet as a marker to determine the digestibility coefficients [19] of fat and protein. Left tibias were removed for ash [20] determination. Determinations of gross energy of feed and excreta by a bomb calorimeter [21], and nitrogen contents [18] of feed, digesta, and excreta were measured for calculating the ME n values [22] of the diets. Fat contents in feed, digesta, and excreta were also performed [18]. Data were analyzed by two-way analysis of variance [23]. A factorial experimental design was used with two energy sources two supplements three replications; levels of significance were based on P RESULTS Data presented in Table 2 show that, although main sources of dietary energy did not affect feed intake from 1 to 7 d of age, both BWG and F:G ratio were superior (P < 0.001) for chicks fed the high-fat diets. Similarly, chicks fed the high-fat diets from 8 to 14 d of age performed better than those fed the highcarbohydrate diets. There was also a main effect of supplement (P < 0.05) showing greater feed intake in the enzyme-supplemented dietary

3 412 JAPR: Research Report TABLE 1. Ingredient and nutrient composition of diets containing 30% defatted rice bran with high-fat or highcarbohydrate concentrations supplemented with bile salt (B) or fiber-degrading enzyme (E) High-fat High-carbohydrate Ingredients B E B E (%, air-dry basis) Soy protein (CP: 84.1%) Yellow corn Soybean meal Defatted rice bran Soy oil:palm oil (1:1) Starch (soluble) Sodium taurocholate Grindazym GP 5000 A DL-Methionine L-Cystine Dicalcium phosphate CaCO Vitamin mix B Mineral mix C NaCl Celite Calculated values ME n, kcal/kg 3,000 3,000 3,000 3,000 Ether extract, % CP, % Lys, % Met + Cys, % Ca, % Nonphytate P, % A Supplied per kilogram of diet: 6,000 U of xylanase, 2,500 U of β-glucanase, and 5Uofpectinase. B Provided per kilogram of diet: vitamin A (palmitate), 10,000 IU; cholecalciferol, 3,000 IU, vitamin E (DL-α-tocopherol), 30 IU; vitamin K 3 (menadione), 2 mg; vitamin B 12, 0.01 mg; biotin, 0.20 mg; choline, 550 mg; folacin, 1.10 mg; niacin, 35 mg; pantothenic acid, 10 mg; pyridoxine, 3.50 mg; riboflavin, 4 mg; thiamin, 1.50 mg; and ethoxyquin, 40 mg. C Provided per kilogram of diet: copper, 6 mg; iodine, 0.50 mg; iron, 50 mg; manganese, 60 mg; selenium, 0.15; and zinc, 40 mg. groups compared with those of the bile saltsupplemented dietary groups from 1 to 7dof age. However, the enzyme-supplemented groups had poorer F:G ratio than the bile saltsupplemented groups in this period of age. No interaction between the dietary energy sources and supplements on chick performance were observed. Fat and protein digestibilities by chicks fed the high-fat diets were consistently superior compared with chicks fed the high-carbohydrate diets, irrespective of age or whether ileal digesta or excreta collection was used (Table 3). The data also show that supplementing diets with bile salt increased 14-d fat digestibility as measured with ileal digesta (P < 0.001) or excreta (P < 0.01) but only increased 7-d ileal protein digestibility (P < 0.001). Interaction effects between energy source and supplement were observed for ileal fat digestibility. Ileal fat digestibilities at Days 7 (P < 0.05) and 14 (P < 0.01) were greatly enhanced by bile salt supplementation compared with enzyme supplementation. The interaction effect was not observed with excreta for fat digestibility. Significant interactions between energy source and supplement on protein digestibilities demonstrated inconsistent results. Ileal protein digestibility was improved (P < 0.01) more by bile salt supplementation compared with enzyme supplementation, apparently, in the highfat diets at Day 7. In contrast, the greater increase (P < 0.05) in excreta protein digestibility of the high-fat diet, clearly observed at Day 7, occurred when diet was supplemented with enzyme, whereas in the high-carbohydrate diet,

4 ADRIZAL ET AL: ENERGY SOURCE IN RICE BRAN DIET 413 TABLE 2. Feed intake, BWG, and F:G ratio of broiler chicks fed diets containing 30% defatted rice bran with highfat or high-carbohydrate concentrations supplemented with bile salt (B) or fiber-degrading enzyme (E) from 1 to 14dofage Feed intake BWG F:G ratio Diet 1 to 7 d 8 to 14 d 1 to 7 d 8 to 14 d 1 to 7 d 8 to 14 d (g/bird) (g:g) High-fat + B High-fat + E High-carbohydrate + B High-carbohydrate + E Energy source High-fat High-carbohydrate Supplement Bile Enzyme SEM A P Source of variance Energy source (ES) NS * *** *** *** *** Supplement (SPL) * NS NS NS * NS ES SPL NS NS NS NS NS NS A Means of three pens per treatment with 10 (1 to 7 d) or seven chicks (8 to 14 d) per pen. *P < 0.05; ***P < TABLE 3. Excreta and ileal digestibilities of fat and protein of diets containing 30% defatted rice bran with highfat or high-carbohydrate concentrations supplemented with bile salt (B) or fiber-degrading enzyme (E) fed to broiler chicks at 1 and 14 d of age Protein digestibility (%) A Fat digestibility (%) A Ileal Excreta Ileal Excreta Diet Day 7 Day 14 Day 7 Day 14 Day 7 Day 14 Day 7 Day 14 High-fat + B High-fat + E High-carbohydrate + B High-carbohydrate + E Energy sources High-fat High-carbohydrate Supplement Bile Enzyme SEM B (P) Sources of variance Energy sources (ES) *** ** *** *** * *** *** *** Supplements (SPL) *** NS NS NS NS *** NS ** ES SPL ** NS * * * *** NS NS A Dry matter basis. B Means of three pens per treatment with 10 (Day 7) or seven chicks (Day 14) for excreta except for ileal digestibility (n = 3; Days 7 and 14). *P < 0.05; **P < 0.01; ***P <

5 414 JAPR: Research Report TABLE 4. Dietary nitrogen-corrected energy (ME n ) values, pancreatic lipase activities, and tibia ash percentages of broiler chicks fed diets containing 30% defatted rice bran with high-fat or high-carbohydrate concentrations supplemented with bile salt (B) or fiber-degrading enzyme (E) at 1 and 14 d of age Lipase activity Tibia ash A ME n (kcal/kg) B (U/g of pancreas) C (%, fat-free) Diet Day 7 Day 14 Day 7 Day 14 Day 7 Day 14 High-fat + B 3,077 3, High-fat + E 3,053 3, High-carbohydrate + B 2,855 2, High-carbohydrate + E 2,687 2, Energy source High-fat 3,065 3, High-carbohydrate 2,771 2, Supplement Bile 2,966 3, Enzyme 2,870 2, SEM D (P) Sources of variance Energy source (ES) *** ** NS * NS Supplement (SPL) * NS NS NS NS ES SPL * NS * NS NS A Data were missed due to a technical error, so only data of Day 14 are presented. B Air-dry basis. C One unit represents milligrams of naphthol released in 10 min at 37 C. D Means of three pens per treatment with 10 (Day 7) or seven chicks (Day 14) per pen except for lipase activity and tibia ash (n = 3; Days 7 and 14). *P < 0.05; **P < 0.01; ***P < such increased excreta protein values were more pronounced when the diets were supplemented with bile salt (Days 7 and 14). Data presented in Table 4 show that the ME n values of the high-fat diets were greater than the values of the high-carbohydrate diets at Days 7 (P < 0.001) and 14 (P < 0.01). In addition to the greater effect of bile salt vs. enzyme supplementation on the ME n values of the diets at Day 7 (P < 0.05), bile salt supplementation also enhanced (P < 0.05) ME n values of diets with high levels of fat (interaction; P < 0.05) at this age. Similarly, this energy source also resulted in higher energy values for the enzyme-supplemented diet with a high level of fat compared to those in the diet with a high level of carbohydrate. No significant differences between major dietary energy sources or supplements were detected in pancreatic lipase activity at Day 7 (Table 4). An interaction effect (P < 0.05), however, was observed whereby bile salt supplementation of the high-carbohydrate diet increased lipase activity compared with the highfat diets. Enzyme supplementation of the highcarbohydrate diet decreased lipase activity compared with the same supplement in the high-fat diets. At Day 14, pancreatic lipase activity was greater (P < 0.05) for chicks fed the high-fat diets. Moreover, dietary treatments had no effect on tibia ash measurements (Table 4). DISCUSSION The superior performance of high-fat over high-carbohydrate (Table 2) in the diet indicated not only that chicks could tolerate high level of dietary fat but also chicks benefited from high level of fat compared to high level of carbohydrate. The beneficial effect of supplemental fat is because it interacts and promotes the digestibilities of inherent-fats and certain nonlipids in other dietary components for better energy utilization [10]. The age-related responses of chicks on fat and protein digestibilities (Table 3) are difficult to explain. However, these results illustrate that

6 ADRIZAL ET AL: ENERGY SOURCE IN RICE BRAN DIET 415 supplemental bile salt directly influenced the target nutrients, particularly as an emulsifier [24] for fat and as a stabilizer [25] for digestive enzymes associating with protein. Greater concentrations of fat emulsification provide greater possibilities of triglyceride hydrolysis by lipase. In addition, the capability of bile salt in enhancing trypsin and chymotrysin stabilities might also be an explanation for improved protein digestibility. On the other hand, the main target of supplemental fiber-enzyme mixtures is on the depolymerization of NSP followed with the ease of penetration of digestive enzymes to nutrients [26], but its indirect, favorable effect on nutrients was not as great as that of supplemental bile salt. Ileal fat digestibilities (Table 3), which were greatly enhanced by bile salt supplementation compared with enzyme supplementation at Days 7 and 14, indicated the synergetic effect between supplemental bile salt and dietary fat. Additionally, blended oils of soy and palm were used as supplemental fat in the current experiments (Table 1). Palm oil is highly saturated, and so it is very dependent on the availability of bile salt for absorption [24]. The interaction effect was not observed with excreta for fat digestibility. It can be speculated that the amount of bile salt that could be adsorbed in the lower intestine and returned to the liver were reduced due to the deconjugating action of lower gastrointestinal microfloras [27]. Hence, the secretion rate of bile salt into the intestine for facilitating fat emulsification would possibly be impaired, which will occur if the diet contains appreciable amounts of NSP [27] as likely in the DRB diet of the present experiment. Although significant interaction effects between energy source and supplement on protein digestibilities demonstrated inconsistent results, excellent effects of supplemental bile salt on protein digestibility when measured in ileal digesta were apparent (Table 3). This finding is likely related to the increases of stabilities of trypsin and chymotrypsin as the results of the effects of exogenous and intestinal bile salt [25]. The significant interaction between energy sources and supplements also indicated an effect of supplemental enzyme on protein digestibility of the high-fat diet when measured in excreta. This interaction possibly took place because the depolymerization of the dietary NSP due to supplemental enzyme provided some carbon-source energy to the particular hindgut microflora for the synthesis of amino acids that were utilized by the host. This phenomenon has been suggested by some authors [28, 29] although it is still debatable. The greater ME n values of the high-fat diets (Table 4) compared to the high-carbohydrate diets were reflected by nutrient digestibility and growth performance of chicks, which demonstrated an extracaloric effect of supplemental fat [10, 30]. The beneficial effect of supplemental fat on dietary energy value is sometimes greater than the predicted energy value of fat on the basis of calculation [10, 30]. A parallel correlation between the energy value of the diet and amino acids [31] or fat [32] digestibility has been reported. The energy source by supplement interaction effect also resulted in a greater effect of enzyme in the diet with a high level of fat than in the diet with a high level of carbohydrate. It is likely that the diet with a high level of fat with bile salt or enzyme supplementation resulted in a positive synergistic effect. This finding suggests that the beneficial effect of additional bile salt for fat emulsification would consequently favor the digestibility of fat concomitantly with energy utilization. The pronounced effect of supplemental enzyme in the high-fat diet also implies that the depolymerizing action of enzyme on NSP leads to greater access of not only digestive enzymes on nutrients but also favors compounds (bile salts) on fat emulsification and digestibility. This is because NSP is capable of binding with bile salt (7, 29). The reason for inconsistent energy source and supplement interaction effects on pancreatic lipase activity is unclear. A greater pancreatic lipase activity at Day 14 for chicks fed the high-fat diets, however, indicated a correlation with a greater ileal or excreta fat digestibility (Table 3) caused by the high-fat level in the diet. The greater fat digestibility could indicate greater micelle formation, which also implies greater intestinal bile salt concentration. Brand and Morgan [33] reported that an increased bile salt concentration in the intestine could be a factor contributing to increased cholecystoki-

7 416 nin, which subsequently stimulated pancreatic lipase secretion and activities. Dietary treatments, which had no effects on tibia ash, indicated that none of the dietary treatments, particularly the two kinds of supplemental components, had a direct effect on the releases of minerals from their binding [34] with NSP. In general, the results reported here show that when DRB, which is relatively low in energy and high in NSP, was used at 30% of a JAPR: Research Report broiler chick diet, protein and fat digestion and dietary ME n content were greater with high-fat diets compared with high-carbohydrate diets. These effects of high fat were reflected in superior growth and feed efficiency of young chicks. The data also showed that supplementation of the 30% DRB diet with the bile salt was of greater benefit than inclusion of a mixture of fiber-degrading enzymes, especially with highfat diets. CONCLUSIONS AND APPLICATIONS 1. Fat and protein digestion and dietary ME n content was greater with high-fat diets compared with high-carbohydrate diets. The data also showed that supplementation of the 30% DRB diet with bile salt was of greater benefit than inclusion of a mixture of fiber-degrading enzymes, especially with high-fat diets. 2. Diets of up to 30% defatted rice bran could be given to broiler chicks if supplemented with bile salt. REFERENCES AND NOTES 1. Annison, G., P. J. Moughan, and D. V. Thomas Nutritive activity of soluble rice bran arabinoxylans in broiler diets. Br. Poult. Sci. 36: Farrell, D. J., and E. A. Martin Strategies to improve the nutritive value of rice bran in poultry diets. I. The addition of food enzymes to target the non-starch polysaccharide fractions in diets of chickens and ducks gave no response. Br. Poult. Sci. 39: Adrizal, and S. Ohtani Effects of rice bran-nonstarch polysaccharides and fiber-degrading enzymes on performance and nutrient digestibility in broiler chicks. J. Poult. Sci. 39: Adrizal, and S. Ohtani. Gifu University, 1-1 Yanagido, Gifu , Japan. Unpublished data. 5. Adrizal, and S. Ohtani Defatted rice bran nonstarch polysaccharides in broiler diets: Effects of supplements on nutrient digestibilities. J. Poult. Sci. 39: Pettersson, D., H. Graham, and P. Aman Enzyme supplementation of low or high crude protein concentration diets for broiler chickens. Anim. Prod. 51: Dänicke, S., O. Simon, H. Jeroch, and M. R. Bedford Interactions between dietary fat type and xylanase supplementation when rye-based diets are fed to broiler chickens. 2. Performance, nutrient digestibility and the fat-soluble vitamin status of livers. Br. Poult. Sci. 38: Choct, M., R. J. Hughes, and M. R. Bedford Effects of a xylanase on individual bird variation, starch digestion, throughout intestine, and ileal and caecal volatile fatty acid production in chickens fed wheat. Br. Poult. Sci. 40: Whitehead, C. C Nutrition: The integrative science. Br. Poult. Sci. 41: Mateos, G. G., and J. L. Sell Influence of carbohydrate and supplemental fat source on the metabolizable energy of the diet. Poult. Sci. 59: Becton Dickinson and Company, Sparks, MD. 12. Danisco Cultor, Brabrand, Denmark. 13. Wako Pure Chemical Industries, Ltd., Osaka, Japan. 14. National Research Council Nutrient Requirements of Poultry. 9th rev. ed. Natl. Acad. Press, Washington, DC. 15. Federation of Animal Science Societies Guide for the Care and Use of Agricultural Animals in Agricultural Research and Teaching. 1st rev. ed. Fed. Anim. Sci. Soc., Savoy, IL. 16. O Sullivan, N. P., E. A. Dunnington, A. S. Larsen, and P. B. Siegel Correlated responses in lines of chickens divergently selected for fifty-six-day body weight. 3. Digestive enzymes. Poult. Sci. 71: Association of Official Analytical Chemists International Official Methods of Analysis. 16th ed. Assoc. Off. Anal. Chem. Int., Arlington, VA. 18. Association of Official Analytical Chemists Official Methods of Analysis. 13th ed. Assoc. Off. Anal. Chem., Washington, DC. 19. Vogtmann, H., H. P. Pfirter, and A. L. Prabucki A new method of determining metabolizability of energy and digestibility of fatty acids in broiler diets. Br. Poult. Sci. 16: Dried tibias were ashed in a furnace at 600 C for 24 h and cooled in dessicator; ash weights were then recorded. 21. Model P-202 CA-4PJ, Shimadzu Co., Kyoto, Japan. 22. Hill, F. W., and D. L. Anderson Comparison of metabolizable and productive energy determinations with growing chicks. J. Nutr. 64: Campbell, R. C Statistics for Biologists. 3rd ed. Cambridge Univ. Press, Cambridge, UK. 24. Krogdahl, A Digestion and absorption of lipids in poultry. J. Nutr. 115: Classen, H. L., and Bedford, M. R The use of enzymes to improve the nutritive value of poultry feeds. Pages in Recent Advances in Animal Nutrition. W. Haresign and D. J. A. Cole, ed. Butterworth Heinemann, Nottingham, UK.

8 ADRIZAL ET AL: ENERGY SOURCE IN RICE BRAN DIET Malathi, V., and G. Devegowda In vitro evaluation of nonstarch polysaccharide digestibility of feed ingredients by enzymes. Poult. Sci. 80: Smits, C. H. M., A. Veldman, H. J. Verkade, and A. C. Beynen The inhibitory effect of carboxymethylcellulose with high viscosity on lipid absorption in broiler chickens coincides with reduced bile salt concentration and raised microbial numbers in the small intestine. Poult. Sci. 77: Karasawa, Y Effect of colostomy on nitrogen nutrition in the chicken fed a low protein diet plus urea. J. Nutr. 119: Karasawa, Y., and M. Maeda A role of caecum in nitrogen nutrition of the chicken fed moderate protein diet and low protein diet plus urea. Br. Poult. Sci. 35: Nitsan, Z., A. Dvorin, Z. Zoref, and S. Mokady Effect of added soyabean oil and dietary energy on metabolizable and net energy of broiler diets. Br. Poult. Sci. 38: Ravindran, V., L. I. Hew, G. Ravindran, and W. L. Bryden A comparison of ileal digesta and excreta analysis for the determination of amino acid digestibility in food ingredients for poultry. Br. Poult. Sci. 40: Dvorin, A., Z. Zoref, S. Mokady, and Z. Nitsan Nutritional aspects of hydrogenated and regular soybean oil added to diets of broiler chickens. Poult. Sci. 77: Brand, S. J., and R. G. H. Morgan Stimulation of pancreatic secretion and growth in the rat after feeding cholestyramine. Gastroenterology 83: Frolich, W Chelating properties of dietary fiber and phytate: The role for mineral availability. Pages in New Developments in Dietary Fiber. I. Furda and C. J. Brine, ed. Plenum Press, New York, NY. Acknowledgment The supply of Grindazyme GP 5000 by Danisco Cultor Brabrand, Denmark, is acknowledged.

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