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*
Unité de Laboratoire pour l’Innovation dans les Céréales, ZAC "Les Portes de Riom," BP 173, F-63204 Riom, France and
Laboratoire Maladies Métaboliques et Micronutriments, Centre de Recherches en Nutrition Humaine Auvergne, INRA, Clermont-Ferrand/Theix, F-63122 St-Genès-Champanelle, France
2To whom correspondence should be addressed at U3M, INRA Theix, F-63122 Theix, France. E-mail: lopez{at}clermont.inra.fr
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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KEY WORDS: • calcium • copper • dietary fiber • iron • magnesium • zinc • rats
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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. This resistant
starch
(RS)3
may be an important substrate for the human colonic microflora, and its
fermentation may convey substantial benefits to the host
(2)
. Nevertheless, various factors, including physical
inaccessibility of the starch in the food, granular structure and the
degree of retrogradation, can limit ileal starch digestibility to
<100% (3)
. Classification of RS into four main types
[physically inaccessible starch (RS 1), resistant granules and high
amylose starches (RS 2), retrograded starches (RS 3) and chemically
modified starches (RS 4)] reflects these major differences
(4)
. Thus, raw native RS (RS 2) but not retrograded RS (RS
3) raises Mg and Ca absorption in rats (5)
.
Despite the classification, the same type of RS, such as RS 2,
refers to resistant granules and high amylose starches (HAS). Raw
potato starch (RPS), which contains a large proportion of resistant
granules of starch, is considered to be RS 2. Feeding RPS leads to very
active fermentations in the distal parts of the digestive tract
(6)
. As a result of the hypertrophy of the cecal wall and
of the cecal acidification, RPS fermentation increases intestinal
absorption of Ca, Mg, Fe, Zn and Cu (7
8
9)
. Furthermore,
RPS lowers plasma cholesterol and triglycerides (10
,11)
.
High amylose starches are also a source of RS 2 and have beneficial
nutritional effects. HAS acts as a prebiotic in promoting the fecal
excretion of probiotic organisms (12)
. Furthermore, HAS
prevents the development of nonreversible insulin resistance in rats
(13
,14)
and depresses plasma cholesterol and
triacylglycerol concentrations in rats (15)
. In humans,
HAS intake lowers plasma cholesterol and triglyceride concentrations
compared with a diet rich in amylopectin starch (16)
. In
contrast to RPS, only one study reported beneficial effects of HAS on
Ca and Fe absorption (17)
. The aim of the present work
thus was to compare the respective effects of RPS and HAS on intestinal
fermentation, mineral absorption (Ca, Mg, Fe, Zn and Cu) and lipid
metabolism in rats. |
MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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We used 64 male Wistar rats, weighing 
150 g, derived from the colony
of laboratory animals of the National Institute of Agronomic Research
(INRA, Clermont-Ferrand/Theix, France). The animals were housed in
wire-bottomed cages to limit coprophagy. The rats were fed a
fiber-free diet (control diet) for 6 d before the beginning of
the experiment (Table 1
). After this period, eight rats had blood samples taken (d 0 of the
experiment), and the others were divided into three groups:
1) 8 rats were fed the control semipurified diet (control
group or C), 2) 24 rats were fed a raw potato–containing
diet (RPS) (Louis François, St-Maur, France) and
3) 24 rats were fed a high amylose starch–containing diet
(HAS) (Hi-Maize; Starch Australasia Ltd., Lane Cove,
Australia).
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Blood collection.
Rats were killed at the end of the lights-out period (between 0800 and 0900 h) because cecal fermentation is still very active during this period. Before they were killed, the rats were anesthetized (40 mg/100 g sodium pentobarbital), and blood samples were taken from the aorta. Artery blood was placed in microfuge tubes containing heparin and centrifuged at 10,000 x g for 2 min. Plasma samples were stored at 4°C for lipid and lipoprotein analyses.
After blood sampling, the cecum (complete with contents) was removed and weighed. Duplicate samples of cecal contents were collected into 2-mL microfuge tubes that were immediately frozen and stored at -20°C. Then, the cecal wall was flushed clean with ice-cold saline, blotted onto filter paper and weighed (cecal wall weight). Cecal water was determined as the difference between wet weight and dry weight on aliquots of cecal contents that were dried to constant weight. Before short-chain fatty acid (SCFA) analysis, supernatants were obtained by centrifuging one of the microtubes at 20,000 x g for 10 min at 4°C.
Analytical procedures.
SCFA were measured in aliquots of cecal supernatants by gas-liquid
chromatography (18)
. Bile acids and neutral sterols were
extracted from feces by a two-step procedure. One volume of sample
was first dispersed in 10 volumes of ethanolic KOH (0.5 mol/L) using a
Polytron disintegrator (Lucerne, Switzerland) and extracted at 70°C
for 2 h. One volume of this suspension (typically 2.5 mL) was
redispersed in 4 volumes of ethanolic KOH and reextracted at 70°C for
2 h. Bile acids were quantified using the reaction catalyzed by 3
-hydroxysteroid dehydrogenase (EC 1.1.1.50; Sigma Chemical Co., St.
Louis, MO) (19)
. Cholesterol concentration was
enzymatically determined on the same extract and on plasma using a kit
purchased from BioMérieux (Charbonnières-les-Bains,
France). Triacylglycerols (Biotrol, Paris, France) were determined in
plasma via an enzymatic procedure. Liver lipids were extracted with
chloroform/methanol (2:1, v/v) (20)
.
Plasma lipoproteins were separated by density gradient
ultracentrifugation using pooled samples (21)
. After
centrifugation in a TST 41.14 swinging-bucket rotor at 100,000
x g for 24 h at 18°C, the gradient was divided
into 24 x 500-µL fractions and kept at 4°C for lipid
analysis. Due to the low level of LDL and the relative overlapping of
HDL1 and HDL2 fractions in
rat plasma, we decided to determine data on the d < 1.040 kg/L fraction (mainly triglyceride-rich lipoprotein, with a
minor contribution of LDL) and on the d > 1.040 kg/L
fraction (essentially HDL).
For mineral determinations, 0.25–0.5 g of dried samples (food and feces) was dry-ashed (10 h at 500°C) and then extracted at 130°C in HNO3/H2O2 (2:1) (Merck, Suprapur, Darmstadt, Germany) until discoloration. Final dilution was made in 1 g lanthanum chloride/L solution (for Ca and Mg determinations) or in 0.2 mL HNO3/L (for Fe, Zn and Cu determinations). Mineral concentrations were determined by atomic absorption spectrophotometry (560; Perkin-Elmer Cetus, Norwalk, CT) in an acetylene-air flame at the following wavelengths: 422 nm (Ca), 285 nm (Mg), 248 nm (Fe), 214 nm (Zn) and 325 (Cu). A nebulizer with high sensitivity was used for trace element determinations. Appropriate quality controls were analyzed with each set of measurements.
Statistical analysis.
Values are means ± SD. Results were compared by one-way ANOVA using the General Linear Models procedure of the SuperANOVA software (Abacus, Berkeley, CA). Post hoc comparisons were performed with Fisher’s least significant difference procedures. Differences between groups were considered significant at P < 0.05.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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The incorporation of 20% RPS or HAS did not markedly alter growth: the final body weights did not differ (control, 257 ± 21 g; RPS, 251 ± 12 g; HAS, 249 ± 8 g).
Cecum weights were low and almost constant in rats fed the control diet
(Fig. 1
). In rats fed the RS diets (RPS and HAS), a similar increase in cecal
weight was noted at 7 d after the beginning of the experiment. In
parallel with cecal contents, the cecal wall weight increased
progressively throughout the study. The cecal pH was close to 7.0 in
rats fed the fiber-free diet; in rats fed the RPS and HAS diets,
markedly acidic pH conditions (5–5.5) were found after 7 d of
treatment and remained steady.
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). However, a substantial level of SCFA (133 µmol) was also present in
the cecum of the rats fed the fiber-free control diet. The presence
of RS in the diet increased the acetate, propionate and butyrate
concentrations. The total cecal pool was 5-fold greater in rats fed the
RPS and HAS diets than in those fed the control diet (P
< 0.001). Furthermore, the fermentation of RPS and HAS resulted
in substantial lactate production.
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The Ca and Mg intakes were the same from the three experimental diets.
However, apparent Ca absorption (daily intake minus fecal excretion)
was significantly greater in rats fed RPS or HAS diets than in those
fed the control diet (P < 0.05) (Table 3
). In rats fed the control diet, half of the dietary Mg was apparently
absorbed. The incorporation of RPS or HAS into the diets significantly
enhanced apparent Mg absorption: 64 or 63% of ingested Mg was absorbed
when diets contained RPS or HAS. The dietary levels of Fe, Zn and Cu
were similar among the groups (Table 4
). Only 24% of Zn was absorbed in rats fed the fiber-free control
diet. This absorption was significantly enhanced by the presence of RPS
or HAS in the diets (27 and 24%, respectively, compared with the
fiber-free diet). Apparent Fe absorption was near 42% in the
controls, and this balance was significantly enhanced by dietary RPS
and HAS (P < 0.05). Regarding Zn and Fe, the apparent
absorption of Cu was increased by the introduction of RPS or HAS into
the diets: Cu absorption was doubled in rats fed the RS diets compared
with those fed the fiber-free control diet (P < 0.01)
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The daily cholesterol intake did not differ among groups (Table 5
). Neutral steroid excretion (fecal cholesterol plus fecal coprostanol)
in controls (61.5 µmol/d) corresponded to 53% of the daily food
supplied. This excretion was significantly higher in rats fed the RPS
or HAS diet and corresponded to 86% of the daily cholesterol intake.
Fecal bile acid excretion was significantly enhanced by dietary RPS or
HAS (Fig. 2
). Thus, unlike for the control diet, the total steroid balance was
negative in rats fed RPS or HAS.
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). Analysis of the plasma lipoprotein profile via gradient density
ultracentrifugation (Fig. 3
) showed that there was a reduction in cholesterol in the d
< 1.040 kg/L fractions (triglyceride-rich lipoprotein) in rats
fed the RPS and HAS diets. The triglyceride concentration of
triglyceride-rich lipoprotein fraction (d < 1.040
kg/L) was significantly depressed in the groups fed the RS diets. By
contrast, there was no significant change in cholesterol in the
d > 1.040 fraction (essentially HDL) in rats fed the
RPS or HAS diets. Moreover, when rats were fed RPS or HAS, the hepatic
cholesterol (-50 and -40%, respectively) and triglycerides (-53
and -47%, respectively) was reduced relative to controls
(P < 0.05).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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. In Western countries, RS constitutes one of the most
important sources of fermentable carbohydrates, and the estimated
average intake of RS in Europe is 5–9 g/d (23)
.
Intestinal fermentation of RS may have important implications for
health. In our experiment, two kinds of RS 2 (RPS and HAS) were
compared, and they elicited similar potent effects on cecal
fermentation and cholesterol and mineral balances.
RS increases cecal weight (total organ and cecal wall) in rats
(24)
, and this is accompanied by histological changes in
cecal mucosa, such as an elevated crypt height and cell number per
crypt (25)
. Thus, the diet containing RPS or HAS induced a
rapid increase in the weight of cecal wall from the first days of the
experiment, due to stimulation of cellular division. The pH fall and
the increase in the SCFA pool play essential roles in inducing cellular
division, because poorly fermentable fibers that accumulate in the
cecum have little hypertrophic effect (26)
. Furthermore,
the microbial breakdown of RS in the large intestine may promote
fermentation, producing a large amount of butyric acid, especially with
a human microflora (27)
. However, this depends on the type
of starch and associated microflora (28)
. Recently,
Ferguson et al. (29)
showed that potato starch, unlike
high amylose corn starch, enhances the proportion of butyrate,
suggesting that there are marked differences among different RS on SCFA
production, even though these were all classified as RS2. In the
present study, high propionic acid fermentation was observed in rats
fed diets containing 20% RPS or 20% HAS. With a higher percentage of
RS in the diet (40%), high butyric acid fermentations has been
observed (6)
.
Cecal hypertrophy under acidic fermentation conditions stimulated Ca
uptake in the distal absorption sites. The large augmentation of the Ca
solubility in the cecum allows this organ to play an important role in
total Ca absorption. In fact, the rat cecum presents the highest
density of Ca transport sites responsive to vitamin D metabolites
(30)
. The total number of these sites could increase, due
to cecal hypertrophy. It is also conceivable that SCFA may directly
influence Ca absorption by modifying electrolyte exchanges (Ca-H), and
Trinidad et al. (31)
proposed that Ca could pass through
the cell membrane more readily in the form of a less-charged
complex (Ca acetate)+ via a passive pathway.
Scharrer and Lutz (32)
also reported a stimulatory effect
of SCFA on Ca absorption in the rat large intestine. Moreover, Ohta et
al. (33)
reported that fermentable carbohydrates
such as fructooligosaccharides (FOS) change the concentration of
Calbindin-D9K differently in the mucosa of the small intestine and
large intestine of rats. Thus, some of the stimulatory effect of FOS
relates to the transcellular route of Ca absorption in the
large intestine of rats. However, the distribution of SCFA along the
large bowel of rats is different from that found in humans, and this
may influence the site of Ca absorption. In fact, the Ca absorption
rate is highly regulated. A high rate of Ca absorption in the large
intestine could trigger a feedback mechanism involving inhibition of Ca
proximal intestine absorption, because there is a control of the
digestive balance of Ca by endocrine factors (34)
. Despite
this feedback, an improvement in Ca assimilation was observed in the
presence of RS in rats or in pigs (5
,8
,9
,17)
. In humans, a
role for the colon in Ca absorption is supported by the observation
that the large intestine is able to maintain a near-normal rate of
Ca absorption in case of small intestine resection (35)
.
In contrast to Ca, the importance of the distal part of the digestive
tract for Mg absorption is well documented (36
,37)
, and it
was previously shown that RS stimulated Mg absorption in rats
(5
,7
8
9)
. Like for Ca, fermentable carbohydrates may also
raise the soluble Mg pool in the large intestine, as a consequence of
acidifying digestive tract contents. In addition, Mg solubility is
generally higher than that of Ca. Thus, the potent effects of RS on Mg
absorption result from large intestine hypertrophy, the increase in Mg
solubility and, possibly, a specific effect of SCFA on passive Mg
absorption (7
,38)
. SCFA are predominantly absorbed in an
undissociated form in the large intestine, although they mainly occur
as anions in the lumen (39)
. Protons needed for SCFA
absorption may be delivered by various ion exchangers (including Mg-H);
in return, SCFA absorption at acidic pH would supply more protons to
the exchangers, resulting in a higher transport rate (32)
.
Fe, Zn and Cu absorptions were significantly greater in animals fed the
RPS or HAS diet compared with the group fed the control diet. Several
explanations for this effect can be proposed. This absorption
improvement can result from an increase in the exchange area
(enlargement of cecum and longer transit time) and the elevation of the
cecal blood flow. Accordingly, Hara et al. (40)
showed
that the cecum and colon contribute to Zn absorption when absorption in
the small intestine is impaired. Moreover, Sakai et al.
(41)
reported that the cecum also plays an important role
in the mechanism by which fermentable carbohydrates prevent
postgastrectomy anemia. However, it is conceivable that the decrease in
cecal pH observed in the RPS and HAS diet groups was accompanied
through some improvement in the solubility of these minerals, because
for a given pH, their salts are generally more soluble than those of Ca
or Mg in the cecal contents. So far, few studies have documented the
specific effect of RS on the absorption and the balance of Fe, Zn and
Cu. Morais et al. (17)
reported in infant pigs that a meal
containing 16.4% HAS induces a greater apparent absorption of Ca and
Fe compared with a completely digestible starch meal. With the RPS
diet, our group found simulating on enhancing effects on Fe
absorption as well as on Zn and Cu absorptions (8
,9)
. The
present study clearly shows that RPS and HAS have similar potentials to
improve trace element absorption (Fe, Zn and Cu) in rats.
RPS and HAS feeding led to 31 and 27% lower plasma cholesterol,
respectively, than in controls. Some (15
,42)
have reported
a marginal cholesterol-lowering effect of amylomaize starch. The
cholesterol-lowering effect of fermentable carbohydrate may be
explained by various mechanisms: 1) inhibition of diffusion
of cholesterol and bile acids at the microvillous boundary layer,
2) steroid-binding capacities in the small intestine,
3) impairment of the passive reabsorption in the large
intestine by insolubilization of bile acids (by acidification of
luminal pH, microbial dehydroxylation to less polar metabolites, or
entrapping on various insoluble structures) and 4) metabolic
effects, especially on hepatic lipid metabolism. The first effect of
RPS and HAS is to reduce cholesterol absorption to a weak level
(14% in rats adapted to the RS diet versus 47% in control rats).
In parallel, a significant effect was observed of RPS and HAS on fecal
bile acid excretion. These effects of RS on the fecal excretion of bile
acids and neutral sterols have been described previously
(43
44
45)
. In humans, an accelerated transit rate was
reported after RS consumption, and this tends to limit the reabsorption
of bile acids (46)
. Thus, the total steroid balance was
negative when RPS or HAS was present in the diet. It must be noted that
the cholesterol-lowering effect of RS was more pronounced in the
triglyceride-rich lipoprotein fraction, which is in keeping with
the inhibition of cholesterol absorption. Moreover, the production of
SCFA from RS, particularly propionate, could potentiate the
consequences of enhanced fecal steroids (47
,48)
. A
putative role of SCFA to mediate the cholesterol-lowering effect of
fiber has been proposed, probably in relation to the inhibition of the
metabolism of the major lipogenic precursors, such as acetate and
lactate. Demigné et al. (49)
showed that propionate,
of which the production is high in animals fed the RPS and HAS diets,
inhibits cholesterogenesis and lipogenesis from acetate. A relatively
low insulinemia has been observed in rats fed dietary fiber. In
parallel to a high rate of SCFA absorption observed in rats fed fiber,
glucose absorption was lower than in fiber-free controls. Taken
together, these factors may account for depressed lipid synthesis
(50)
. Thus in the present study, RPS and HAS lowered
plasma lipids in rats. Nevertheless, some types of RS do not do so in
humans (51)
or in pigs (52)
. Clinical studies
showed no effect of RS on plasma lipids in humans (53)
.
Moreover, in contrast to rats, RS lowers fecal bile acid excretion in
humans (54)
. It appears that rats may differ fundamentally
from humans, because of the differences of intestinal microflora.
Indeed, the activity of colonic microflora appears to influence serum
lipid levels (55)
. However, as in rats, RS is fermented by
the human gut microflora into SCFA in the large intestine. These SCFA
generated by bacterial fermentation of fibers suppress cholesterol
synthesis in liver and intestine (56)
, thus reducing serum
and hepatic cholesterol concentrations (57)
In conclusion, although direct extrapolation to humans may be questionable due to differences in digestive tract structure and colonic microflora, the substitution of a portion of digestible starch by RPS or HAS leads to enhanced intestinal fermentation, improved mineral absorption and reduced cholesterol absorption in rats. It is of note that RPS or HAS may play a role by increasing mineral absorption in the large intestine, and this effect may be of particular interest when the overall process of digestive absorption is inefficient, such as in elderly subjects. This awaits further investigation in human subjects, particularly to assess whether low levels of RPS or HAS are effective as cholesterol-lowering and mineral-improving agents.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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3 Abbreviations used: C, control; FOS, fructooligosaccharides; HAS, high amylose starch; RPS, raw potato starch; RS, resistant starch; SCFA, short-chain fatty acids.
Manuscript received August 9, 2000. Initial review completed October 15, 2000. Revision accepted December 19, 2000.
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