The Journal of Nutrition 1964 Volume.82 No.2
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Influences of Dietary Carbohydrate-Fat Combinations on Various Functions Associated with Glycolysis and Lipogenesis in Rats II. GLUCOSE VS. SUCROSE WITH CORN OIL AND TWO HYDROGENATED OILS* 1 CATHERINE CARROLL Department of Home Economics, Agricultural Experiment Station, University of Arkansas, Fayetteville, Arkansas ABSTRACT Weanling rats were fed diets differing only in source of carbohydrate and fat for 2 to 4 weeks. Livers were assayed for glucose-6-phosphatase and fructose diphosphatase activities, and for content of glycogen and lipids. Effects on enzyme activities of substituting fructose for glucose were similar to those observed on sub stituting sucrose for rice starch (previous report). Feeding either hydrogenated coconut oil (HCO) or hydrogenated peanut oil (HPO) in place of corn oil (CO) modified the enzymatic responses to dietary fructose. Results with HPO were some what different than those with HCO. Labile phosphorus values were highest in groups fed sucrose or fructose with CO, and lowest in those fed rice starch or glucose with HPO. Effects of dietary carbohydrate on accumulation of lipid in liver appeared to be a function of the type of fat fed, namely, substitution of a fructose source for a direct glucose source resulted in the accumulation of less fat in livers of rats fed CO, but of more fat in livers of rats fed a hydrogenated oil. Proportions of phospholipid and cholesterol in liver lipid, and concentration of cholesterol in serum also varied with the combination of carbohydrate and fat fed. The type of carbohydrate fed to rats starch as compared with that in livers influences the activity of several enzyme from rats fed corn oil with rice starch. systems involved in carbohydrate metabo Lipid content of the liver and cholesterol lism (1-4). Freedland and Harper (1) concentration of the serum also varied noted that the stimulation of glucose-6- with the combination of carbohydrate and phosphatase activity resulting from the fat in the diet. substitution of an indirect source of glu The experiments reported in the present cose (e.g., fructose) for a direct source paper were designed to gain more informa (dextrin) appeared to be an adaptation of tion about the above interrelationships by the enzyme system due to increased gluco- the following means: 1) Determining neogenesis. Recent studies from this whether substitution of fructose for glu laboratory demonstrated that metabolic cose with corn oil and with saturated fat responses to the dietary source of carbo would produce the same results as had the hydrate are partially dependent on the substitution of sucrose for rice starch; 2) type of fat in the diet (5). For example, comparing effects of feeding a hydrogen adaptation in the activities of the glucose- ated fat containing long-chain fatty acids 6-phosphatase and fructose diphosphatase with those observed on feeding coconut oil; enzyme systems, and changes in glycogen and 3) following the course of liver lipid content of the liver resulting from the sub variations for an additional 2 weeks. stitution of sucrose for rice starch were Criteria for evaluating differences in modified by the type of fat fed (com oil carbohydrate and lipid metabolism were vs. hydrogenated coconut oil). Further evi the same as used in the previous study dence for an influence of the dietary (5) except that in the second part of the source of fat on carbohydrate metabolism current study livers were also assayed for was the significantly greater glucose-6- labile phosphorus from ADP and ATP. phosphatase activity in livers from rats Received for publication August 12, 1963. fed hydrogenated coconut oil with rice 1 Supported in part by NIH grant A-4854-C1. J . N u t r it io n , 8 2 : *64 163 164 CATHERINE CARROLL EXPERIMENTAL were decapitated. Methods used for serum Experimental animals, cholesterol determinations and for liver diets, and groups assays for activities of the glucose-6-phos- phatase (G-6-Pase) and fructose diphos Male, weanling rats2 of the Sprague- phatase (FDPase) enzyme systems, for Dawley strain, weighing approximately glycogen, and for total lipid, phospholipid, 50 g each at the beginning of the study, and cholesterol are described in a previous were fed different carbohydrate-fat com paper (5). An additional assay carried out binations. Rations for all groups consisted in part 2 was the determination of labile of the following: (in per cent) protein, phosphorus from ADP + ATP. The adeno 20 (casein supplemented with 2% of d l - sine phosphates from samples of liver ho methionine); fat, 15; carbohydrate, 60.55; mogenates were adsorbed onto charcoal, choline chloride, 0.2; salts (6),3 4; and and labile phosphorus from ADP + ATP vitamin mix,4 0.25. Variables were the was hydrolyzed off by boiling in HC1 (7). type of carbohydrate and the type of fat. The inorganic phosphorus released was Part 1. A total of 52 rats was divided determined by the method of Fiske and into 8 groups (6 or 7 rats /group). Each Subbarow (8). group was fed a diet containing one of the following carbohydrate - fat combinations RESULTS for 2 weeks: corn oil5 (CO) with glucose, CO with fructose, hydrogenated coconut Growth rates and relative oil6 (HCO) with glucose, HCO with fruc liver weights tose, and each of these 4 combinations The feeding of hydrogenated oil in place with added cholesterol.7 of com oil resulted in a depression of the Part 2. Since some differences were growth rate, more severe with HPO than observed between effects of monosaccha with HCO. However, the average weekly rides in part 1, and those of rice starch and weight gain for each group for the entire sucrose in the previous study (5), part 2 experimental period was at least 30 g, and was designed to investigate these differ did not fall below 26 g for any group in ences by feeding the monosaccharides and any single week (table 1). the more complex carbohydrates simulta Relative liver weights (liver weight/ neously to different groups of rats. Also, 100 g of body weight) were significantly hydrogenated peanut oil8 (HPO) was used greater in all groups fed a source of fruc in place of HCO, to observe effects of pro tose (sucrose or fructose) as compared viding more long-chain fatty acids. The with those of corresponding groups fed a approximate fatty acid content of the direct source of glucose (rice starch or 2 fats, as supplied by the manufacturers, glucose) (table 1). Differences were sig was as follows: for HPO (in per cent) nificant at the 1 or 2% level10 in all in palmitic, 12; stearic, 79; arachidic and stances except one, sucrose vs. rice starch longer saturated, 6; and mono-unsatu- with CO at 4 weeks. This observation is rated, 3; as compared with HCO (in per consistent with previous reports from other cent) caprylic, 6; capric, 6; lauric, 44; laboratories that relative liver weight is myristic, 18; palmitic, 11; and stearic, 15. increased by feeding fructose in place of The iodine number of the HPO was ap glucose (9) or dextrin (1). proximately 4, and that of the HCO was 2 Obtained from Hormone Assay Laboratories, Inc., 1 to 3. A total of 96 lats was divided into C hicago. 3 Salt Mixture-W, obtained from Nutritional Bio 8 groups of 12 rats each. Four rats from chemicals Corporation, Cleveland. 4 The vitamin mix provided the following: (in m g/ each group were fed the experimental diet 100 g ration) thiamine-HCl, 0.08; riboflavin, 0.6; for 2 weeks, four for 3 weeks, and four pyridoxine, 0.4; Ca pantothenate, 4.0; niacin, 5.0; inositol, 20.0; folic acid, 0.04; vitamin B i2, 0.004; for 4 weeks. The 8 carbohydrate-fat com biotin, 0.02; vitamin A powder, 10.0 (200 units); calciferol, 0.18 (150 units); Dn-a-tocopherol powder, binations were: CO and HPO each with 30.0 (7.5 units); and menadione, 0.38. 3 Mazola, Com Products Company, Argo, Illinois. rice starch,9 sucrose, glucose, and fructose. 6 Hydrol, Durkee Famous Foods, Chicago. 7 Cholesterol added at level of 1% of total ration at PROCEDURES expense of dietary fat. 8 Obtained from Procter and Gamble Company, After animals had been fed experimen Cincinnati. 9 Obtained from Morningstar-Paisley, Inc., New York. tal diets for the times specified above, they 10 S tu d e n t’s t test. DIETARY CARBOHYDRATE-FAT COMBINATIONS 165 TA BLE 1 Groiuth rates and relative liver weights of rats fed different combinations of carbohydrates and fats Direct Direct Fructose Weeks glucose Fructose glucose source fed diet source source, with source with CO saturated saturated with CO i fat fat 9 g g g Weight gain/week G and CO F and CO G and HCO F and HCO 1 (7 )2 44 ± 1 3 4 1 * 2 4 1 * 1 3 7 * 1 2 (7 ) 4 7 ± 2 4 9 * 3 4 1 * 2 4 1 * 2 G and CO + C F and CO + C G and HCO + C F and HCO + C 1 (6 ) 44 ± 2 4 5 * 1 4 1 * 1 4 1 * 2 2 ( 6 ) 4 7 * 3 4 2 * 3 3 9 * 3 3 2 * 3 RS and CO S and CO RS and HPO S and HPO 1 (1 2 ) 45 ± 1 4 6 * 1 3 3 * 2 2 6 * 2 2 ( 1 2 ) 53 ± 1 5 3 * 1 4 0 * 2 4 5 * 1 3 (8 ) 4 9 * 1 4 4 * 2 4 1 * 1 4 4 * 1 4 (4 ) 52 ± 3 5 3 * 1 4 6 * 1 4 2 * 4 G and CO F and CO G and HPO F and HPO 1 (1 2 ) 43 ± 1 4 0 * 1 3 3 * 1 2 7 * 2 2 (1 2 ) 48 ± 1 4 8 * 1 3 7 * 1 3 3 * 1 3 (8 ) 46 ± 2 4 3 * 1 3 4 * 2 3 0 * 2 4 (4 ) 50 ± 4 4 5 * 6 3 3 * 2 2 9 * 2 Relative liver weights 1 G and CO F and CO G andHCO F and HCO 2 (7 ) 4.81 ± 0 .1 0 6.22 * 0.22 5 .0 5 * 0 .1 2 6 .2 6 * 0 .1 5 G and CO + C F and CO + C G and HCO + C F and HCO + C 2 (6 ) 4 .9 2 ± 0.14 6 .4 4 * 0 .2 2 4 .8 5 * 0 .2 3 5 .9 2 * 0 .2 3 RS and CO S and CO RS and HPO S and HPO 2 (4 ) 4 .9 2 ± 0.06 5 .7 2 * 0 .1 0 4 .4 0 * 0 .1 3 5 .4 7 * 0 .1 9 3 (4 ) 4 .7 7 * 0 .1 0 5 .4 3 * 0 .1 3 4 .5 4 * 0 .1 2 5 .2 8 * 0 .0 7 4 (4 ) 4 .5 4 * 0 .0 8 4 .8 6 * 0 .1 8 4 .0 4 * 0 .1 6 5 .4 6 * 0 .3 3 G and CO F and CO G and HPO F and HPO 2 (4 ) 5 .0 1 * 0 .0 7 6 .3 3 * 0 .1 1 4 .5 7 * 0 .2 2 5 .5 0 * 0 .1 9 3 (4 ) 4 .7 9 * 0 .0 5 5 .8 5 * 0 .1 2 4 .6 2 * 0 .1 2 5 .7 0 * 0 .1 7 4 (4 ) 4 .4 6 * 0 .0 9 5 .9 9 * 0 .4 1 4 .2 4 * 0 .0 8 5 .5 2 * 0 .1 1 1 Abbreviations: CO = corn oil, HCO = hydrogenated coconut oil, HPO = hydrogenated peanut oil, C = cho lesterol (1% of ration), G = glucose, F = fructose, RS = rice starch, S = sucrose.