1 Conservatoire National des
Arts et Métiers, Stimulating water
absorption in the colon represents an important target to reduce stool
output in secretory diarrhea. Recently, a 153-amino-acid peptide was
isolated from porcine upper small intestine and purified, taking into
account the increase of water absorption in guinea pig gallbladder.
Accordingly, this peptide was named sorbin. The aim of the present
study was to determine if the COOH-terminal heptapeptide of sorbin
(C7-sorbin) participates in the
regulation of electrolyte transport in the colon. Different regions
(from duodenum to colon) of stripped intestinal mucosa from rats or
humans were mounted in Ussing chambers to measure the changes in
short-circuit current
(
sodium chloride absorption; ion transport; colon
THE GASTROINTESTINAL TRACT IS uniquely designed to
control water and electrolyte economy of the body (for recent review,
see Ref. 10). In one day, the gastrointestinal tract must deal with a
minimum of 8-9 liters of water from ingested meals and drinks and
digestive secretions. Much of this volume is absorbed in the small
intestine secondary to nutrient and electrolyte absorption, leaving
1-2 liters that enter the large intestine. Here, 90% of the
remaining water is absorbed, yielding an average daily stool volume of
100-200 ml.
In diarrheal diseases, the increase in stool volume is the consequence
of a situation in which the volumes presented to the colon exceed its
large reserve capacity. Thus, in secretory diarrhea such as cholera,
diarrhea occurs when the large volume produced by the secretory effect
of the toxins from the vibrio in the small intestine overwhelms the
absorptive capacity of the colon (for recent review, see Ref. 11). At a
constant secretion rate in the small intestine, the extent of water and
electrolyte losses in stools are directly related to the absorptive
capacity of the colon. Hence, the severity of dehydration in cholera
could be attributed in part to the decrease in colonic absorptive
capacity (36). In addition, the severity of viral infection in young piglets, before 3 days of age, appears to be the consequence of a lack
of stimulation of colonic absorption in response to the increased
volume presented to the colon from the small intestine (1). In contrast
to 3-day-old piglets, the colons of 3-wk-old infected piglets had
increased fluid absorption that was about six times over controls; this
compensatory response prevented diarrhea in these older animals.
The epithelial cells lining the gastrointestinal tract are the key
players in the absorptive and secretory events. Electrolytes are
actively transported from lumen to blood or in the opposite direction
as a result of the activity of the
Na+-K+-ATPase and
asymmetric distribution of specific transporters or channels, between
luminal and basolateral membranes of the enterocytes. Regulation of
intestinal electrolyte absorption or secretion occurs via intracellular
messengers that are modulated by various external stimuli, including
neuromediators. Several of them, including peptide YY (PYY) (9, 20),
somatostatin (5, 16, 28), ANG (7, 17), enkephalin (16, 24, 27), and
norepinephrine (28), reduce intestinal secretion in physiological and
pathological conditions. However, the role of these hormones in
stimulating colonic absorption as a response to increased secretion is
ill defined.
Sorbin is a peptide of 153 amino acids that was isolated and purified,
taking into account the increase in water absorption, from porcine
upper small intestine in guinea pig gallbladder (32, 37). In the rat,
the COOH-terminal heptapeptide of the natural molecule was found to be
the minimal biologically active fragment. The synthetic modified
fragment
[Pro-Val-Thr-Lys-Pro-Gln-(D-Ala)-NH2 (C7-sorbin)] was found to
increase water and electrolyte absorption in the rat duodenum and to
decrease intestinal secretion induced by vasoactive intestinal peptide
(VIP) perfusion (7, 23, 29). In addition, it reduced water secretion
induced by cholera toxin in the rat (J. Fioramonti and L. Bueno,
personal communication). Because sorbin has been selected and
identified based on its effect on water absorption in the gallbladder,
its main activity might be the stimulation of an NaCl neutral
absorptive process (18, 21). In addition, if sorbin is mainly active on
NaCl absorption, it may be predicted that its effect would be observed
in the ileum and the colon where the neutral NaCl transport is
primarily located (6).
Thus the aim of the present study was to determine if
C7-sorbin participates in the
regulation of electrolyte transport in the colon in basal and
stimulated secretion conditions. The results indicate that
C7-sorbin stimulated NaCl neutral
absorption and inhibited electrogenic
Cl Chemicals
ABSTRACT
Top
Abstract
Introduction
Materials and methods
Results
Discussion
References
Isc) and
net 22Na and
36Cl fluxes
(JNanet and
JClnet) after serosal exposure
of 10
7 to
10
3 M
C7-sorbin. In fasted rat
intestine, C7-sorbin
(10
4 M) induced an
immediate reduction in
Isc in the distal
ileum and proximal and distal colon but not in the duodenum and
jejunum. In the colon,
Isc reduction and
JNanet and JClnet stimulation were dose
dependent (EC50 = 2 × 10
5 M). At
10
3 M, maximal effect was
observed (
Isc =
1.14 ± 0.05,
JNanet = +4.97 ± 1.38, and
JClnet = +9.25 ± 1.44 µeq · h
1 · cm
2).
C7-sorbin
(10
3 M) inhibited the
increase in Isc
induced by a series of 10 secretory agents such as secretin, vasoactive
intestinal peptide, PGE2, and
serotonin. In HT-29-Cl19A cells,
C7-sorbin induced an increase in
Isc, with a
maximal effect at 10
3 M
(
Isc = 0.29 ± 0.10 µeq · h
1 · cm
2).
In human intestine, a dose-dependent decrease in
Isc was observed in right and sigmoid colons in basal and stimulated conditions (EC50
10
5 M; at
10
4 M,
Isc =
2.66 ± 0.17 µeq · h
1 · cm
2)
but not in the jejunum. The results indicate that
C7-sorbin stimulated NaCl neutral
absorption and inhibited electrogenic Cl
in rat and human
intestinal epithelia. In addition, the antisecretory effect was
essentially observed in the distal part of both rat and human intestine
and the magnitude of the proabsorptive effect was directly related to
the magnitude of the previously induced secretion.
INTRODUCTION
Top
Abstract
Introduction
Materials and methods
Results
Discussion
References
secretion in the rat
colon. In addition, the antisecretory effect was essentially observed
in the distal part of rat and human intestine. The magnitude of the
proabsorptive effect was directly related to the magnitude of the
previously induced secretion.
MATERIALS AND METHODS
Top
Abstract
Introduction
Materials and methods
Results
Discussion
References
-atrial
natriuretic factor-(1
28), neurotensin, ANG I, PYY, and somatostatin
(somatotropin release-inhibiting factor, SRIF), at least 95% pure,
were obtained from Neosystem (Strasbourg, France). H-sorbin
(C7-sorbin) was a kind gift from Institut Henri Beaufour (Les Ulis, France). Scintillation fluid (Aquasafe 300+) was purchased from BAI (Elancourt, France).
H36Cl (0.2 M, specific activity
0.34 MBq/mg Cl
) was from
Amersham (Les Ulis, France).
22NaCl (specific activity 23.38 MBq/mg Na+) was from DuPont NEN
(Les Ulis, France). All other chemicals were analytic grade reagents.
Peptide Solutions
Stock solutions (10Preparation of Tissues
Mature male Sprague-Dawley rats weighing 180-250 g were obtained from Iffa Credo (St. Germain s/Arbresle, France) and housed in individual cages and fed standard laboratory chow (UAR, Villemoisson s/Orge, France) until used in these studies. Food was withdrawn 18 h before the experiments, but animals had free access to drinking water.For electrophysiology studies, animals were killed by cervical dislocation, and segments of intestine from fasted animals were removed and rinsed free of intestinal content by flushing with ice-cold Ringer solution. The animals' stomachs were found to be empty. Tissues were stripped off the muscular layer, opened along the mesenteric border, and mounted as flat sheets between the two halves of acrylic Ussing chambers, as previously described (20, 35).
Human tissues were obtained at surgery from digestive cancer patients. Healthy pieces were stored in ice-cold Ringer medium until used (close to 1 h after the tissue was removed). Tissues were prepared as described above and mounted in Ussing chambers.
Short-Circuit Current Studies in Ussing Chambers
The isotonic Ringer solution used throughout the experiments contained (in mM) 115 NaCl, 25 NaHCO3, 1.2 MgCl2, 1.2 CaCl2, 2.4 K2HPO4, and 0.4 KH2PO4. The pH was 7.40 at 37°C when bubbled with the 95% O2-5% CO2 mixture used to circulate the chamber fluid. The ClThe spontaneous transmural electrical potential difference, reflecting
the asymmetry of electrical charges between the luminal and serosal
membranes, was measured via 3 M KCl solution in 4% (wt/vol) agar
bridges. These bridges were placed on both sides of the tissue and
adapted to calomel half-cells, linked to a high-impedance voltmeter.
Potential difference was short-circuited throughout the experiment by a
short-circuit current
(Isc) via 3 M
KCl in agar bridges placed in each reservoir, adapted to Ag-AgCl
electrodes in relation with an automatic voltage-clamp system (DVC
1000, World Precision Instruments, Sarasota, FL). Delivered
Isc, corrected for fluid resistance, was recorded continuously on a computer. Isc represents
the sum of the net ion fluxes transported across the epithelium in the
absence of an electrochemical gradient (mainly Na+,
Cl, and
HCO
3). Every 30 s, the tissue was
automatically clamped at +1 mV for 3 s to calculate transepithelial
electrical conductance, according to Ohm's law. Peptides were added to
the serosal medium after stabilization of
Isc.
Effect of C7-Sorbin on Isc
C7-Sorbin was used at both single and cumulative doses. In the single dose experiments, one piece of tissue per chamber was exposed to only one given concentration that ranged from 10Effect of C7-Sorbin on
Na+ and
Cl Fluxes
Effect of C7-Sorbin on Secretion Induced by Intestinal Peptides
In this experiment, one piece of tissue per chamber was exposed to 2 × 10Statistics
Results are reported as means ± SE. All determinations were performed in pieces of tissue (n) from at least seven rats or three humans. ANOVA was performed according to the general linear model procedure, and comparison of means was by the least-square difference test of the SAS package (SAS Institute, Cary, NC). ![]() |
RESULTS |
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Effect of C7-Sorbin in Rat Intestine
Effect of C7-sorbin on
Isc.
Typical recordings of the effect of
C7-sorbin on
Isc in rat colon
are presented in Fig. 1. In basal
condition, addition of 103
M C7-sorbin in the serosal
compartment was followed by an immediate and steady decrease in
Isc (Fig.
1A). A similar effect was observed (Fig. 1B) after stimulation of
Isc by
10
4 M 5-HT. Further
addition of 5 × 10
5 M
bumetanide in the serosal solution did not decrease the current, strongly suggesting that C7-sorbin
entirely inhibited electrogenic Cl
secretion. Previous
treatment with naloxone
(10
6 M) or TTX (2 × 10
6 M) did not alter the
response of colon to C7-sorbin.
|
Effect of C7-sorbin in different segments
of rat intestine.
A small antisecretory effect of
C7-sorbin was found in both
duodenum (Isc =
2.67 ± 0.67 µA/cm2) and jejunum
(
Isc =
5.27 ± 0.94 µA/cm2), as shown in Fig.
2. At
10
4 M, the effect of
C7-sorbin was more marked in the
ileum (
Isc =
8.16 ± 3.37 µA/cm2,
P < 0.05) and even more so in the
colon (
Isc =
17.69 ± 2.82 µA/cm2,
P < 0.001).
|
Dose response of C7-sorbin on
Isc.
In rat colon, C7-sorbin induced a
dose-dependent reduction in
Isc in basal
condition. The apparent threshold was at
106 M, the
EC50 value close to 2 × 10
5 M, and maximal response
was at 10
3 M (Fig.
3). Almost identical results were obtained
when the tissue was stimulated by
10
4 M 5-HT before
C7-sorbin addition (not shown).
|
Effect of C7-sorbin on
Cl and
Na+ fluxes in
rat colon.
The simplest explanation for the reduction in
Isc is an
inhibition of electrogenic
Cl
secretion linked to a
stimulation of neutral NaCl absorption that cannot be directly measured
by Isc.
Therefore, fluxes were measured by use of
22Na+
and
36Cl
.
Figure 4 shows that
C7-sorbin reduced
Isc and
stimulated Na+ and
Cl
absorption in a
dose-dependent manner. C7-sorbin
also induced a dose-dependent decrease in tissue conductance together
with an decrease in unidirectional fluxes from serosa to mucosa (Table 1). In jejunum, however, small but
significant effects of C7-sorbin on Isc and net
Cl
flux were only noticed
at 10
3 M (Table
2).
|
|
|
Effect of C7-sorbin on secretion induced
by secretory agents in rat colon.
Several secretory agents present in the intestinal mucosa can stimulate
Cl secretion, including
5-HT, PGE2, and VIP. Thus a series
of these compounds were first added to the serosal side of stripped
colonic mucosa, followed by serosal addition of
10
3 M
C7-sorbin. After a peak,
Isc returned to a
steady value greater than the basal value; further addition of
C7-sorbin elicited a decrease in
Isc until a new
steady state was reached. For most tested compounds, subsequent
addition of C7-sorbin was followed by a significant decrease in
Isc. The
magnitude of the
Isc decrease after C7-sorbin was inversely
related to the magnitude of
Isc stimulated by
secretory agents. The relationship between the steady-state Isc obtained
after stimulation by secretory agents and the decrease in
Isc elicited by
further addition of C7-sorbin is
presented in Fig. 5.
|
Comparison of different antisecretory agents in rat colon.
The dose responses of four antisecretory agents on
Isc were compared
in rat colon; EC50 of SRIF was
obtained at lower concentrations than that of other antisecretory
agents such as PYY, C7-sorbin, and
clonidine in the following sequence: SRIF > PYY > C7-sorbin > clonidine. When we
compared the maximal effect of these antisecretory agents, the effect
of C7-sorbin was similar to that
obtained with SRIF, whereas the lowest maximal effect was obtained with
clonidine (Fig. 6).
|
Effect of C7-Sorbin in HT-29 Cell Monolayers
The effect of C7-sorbin was assessed in vitro on human colonic cell line HT-29-Cl19A at concentrations ranging from 10
|
Effect of C7-Sorbin on Human Colon
Effect of C7-sorbin on different segments
of human intestine.
The effect of C7-sorbin was tested
on different segments of human intestine (Fig.
8).
C7-sorbin dose dependently reduced
basal Isc in the
colon (right colon and sigmoid), as shown in Table 3. Addition of bumetanide did not further
modify the Isc.
In the jejunum, C7-sorbin had no
effect on Isc.
|
|
Effect of C7-sorbin on human colon after
stimulation of Isc by VIP.
The effect of C7-sorbin was
assessed in human colon after stimulation of tissue by VIP (Fig.
9). After
Isc stimulation
by 2 × 107 M VIP,
C7-sorbin reduced secretion in
human colon. The threshold for the
C7-sorbin effect was
10
5 M, but significant
reduction in Isc
was observed at concentrations over
10
5 M.
|
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DISCUSSION |
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Our study clearly suggested that
C7-sorbin stimulated NaCl neutral
absorption and inhibited electrogenic
Cl secretion in the rat
colon. In addition, the antisecretory and proabsorptive effects were
essentially observed in the ileum and colon, in rat and human
intestine. The magnitude of the proabsorptive effect was directly
related to the magnitude of the previously induced secretion.
The proabsorptive effect is constitutive of the peptide sorbin, which
was selected from extracts from porcine mucosa on the basis of a
capacity to stimulate absorption of water in guinea pig gallbladder
(32, 37). The COOH-terminal heptapeptide has mainly been tested for its
effect on water and electrolyte absorption in vivo in intestinal loops
in the rat (7, 23, 29). It was found that
C7-sorbin induced absorption of
water and electrolytes in duodenum and ileum. In addition,
C7-sorbin decreased the
VIP-stimulated secretion of water (37%),
Na+ (30%),
Cl (63%), and
HCO
3 (25%) (29). These effects were
greater than those produced by equimolar doses of neuropeptide Y, SRIF,
and metenkephalinamide in rat ileum ligated loops (29). In our study
using isolated fragments of intestine mounted in Ussing chambers, we
have confirmed the antisecretory and proabsorptive effects of
C7-sorbin in the intestine of rats.
We also have extended the previous observations in terms of site and mechanism of action and relationship with the secretory status of the intestine. The present results clearly indicate that the C7-sorbin effect displays an aboral gradient: the decrease in Isc was hardly observed in the duodenum and jejunum, and in the ileum it was 46% lower than that observed in the colon. The present results might explain why large numbers of animals were necessary to demonstrate an effect of sorbin in the small intestine (7, 23, 29). The proabsorptive effect of sorbin in the colon is not unique, as several other agents have been found to display similar effects, including PYY (9, 20), somatostatin (5, 16, 28), ANG (7, 17), enkephalin (16, 24, 27), and norepinephrine (28). However, the present study clearly indicates that the main effect of sorbin is located in the distal part of the rat and human intestine.
The aboral gradient for the effect of sorbin on electrolyte transport
may be due to several factors, including the distribution for the
sorbin receptors and effectors. Very little is known about the
receptors. In recent experiments, no specific binding for sorbin was
found in basolateral membranes of intestinal epithelial cells (M. Laburthe and T. Voisin, personal communication). In terms of cellular
effectors, the effect of sorbin on stimulating water absorption in the
gallbladder pointed to the possibility of an effect on a neutral NaCl
absorptive process (18, 21, 37). We thus measured on the same pieces of
tissue the electrical parameters and the
Na+ and
Cl isotopic transepithelial
fluxes in absence of a transepithelial electrochemical gradient. Also,
because the coefficients of variation are commonly much greater for the
isotopic fluxes than for the electrical parameters, the determinations
were made at four different concentrations of
C7-sorbin, i.e., on 16 adjacent
pieces of colon or jejunum in the same experiment. The results
confirmed that the colon is the main target for sorbin, whereas
isotopic fluxes were not significantly modified in the jejunum. In the
colon, a dose-dependent stimulation of the neutral NaCl absorption
together with a smaller reduction in electrogenic
Cl
secretion was observed.
The results obtained after substitution of
Na+ or
Cl
indicate that
Na+ and
Cl
are required for the
effect of C7-sorbin. In addition,
comparison of the ionic unidirectional fluxes with the transepithelial
conductance further strengthened the possibility that sorbin stimulates
electrolyte absorption (transepithelial conductance expressed in
mS/cm2 is similar to the sum of
the diffusional ionic fluxes measured in the absence of an
electrochemical gradient and expressed in µeq · h
1 · cm
2)
(13). Thus the change in conductance could be compared with the change
in the unidirectional ionic fluxes. In the present experiments,
conductance decreased with increasing sorbin concentrations; concurrently, the unidirectional ionic fluxes from serosa to mucosa decreased, suggesting a reduction in transepithelial ionic diffusion, whereas the flux in mucosa to serosa was not significantly altered. This may indicate a decrease in the diffusional flux compensated for by
an increase in the nondiffusional flux from mucosa to serosa compatible
with a stimulation of a neutral NaCl absorptive flux. Although it was
beyond the scope of these experiments to identify the molecular
structures supporting the NaCl absorption and the Cl
secretion, the
literature suggests that
Na+/H+
exchanger (NHE3) (38) and cystic fibrosis transmembrane conductance regulator (2, 22) transporters are good candidates for membrane effectors that support the proabsorptive and antisecretory effects of sorbin.
Interestingly, we observed a similar aboral distribution in the human
intestine. This may indicate a similar distribution for the ionic
transporters involved in the effect of sorbin in the rat and human
intestine. Thus sorbin extracted from porcine intestine is also active
on rat and human intestine. However, these results do not indicate the
presence of sorbin or its receptors in the human intestine. When we
measured the effect of C7-sorbin on HT-29-Cl19A, an intestinal epithelial cell line derived from a human
colonic cancer patient, it came as a surprise that the Isc did not
decrease but increased at a relatively high
C7-sorbin concentration
(104 M). This may indicate
that sorbin does not act directly on the intestinal epithelial cells,
but, like the many mediators located in the lamina propria, sorbin may
interact with different cell types that control electrolyte transport
by the epithelial cells (3, 8, 10, 12). However, neither naloxone nor
TTX altered the C7-sorbin effect.
One of the most intriguing results is the effect of sorbin after secretion was stimulated: the intensity of the effect of sorbin appeared to be directly related to the intensity of the secretion previously induced by a variety of agents (Fig. 5). The physiological consequence of a regulatory link between absorption and secretion is obvious when considering the movement of water and electrolytes in the intestine after a meal (14, 34). During a meal, a large quantity of water enters the intestinal lumen as a result of the volume of the meal and the meal-induced secretion by the salivary glands, stomach, pancreas, biliary system, and intestine. Several hormones participate in the meal-induced secretion (30). Water and electrolytes thus secreted in the intestinal lumen are reabsorbed by the intestine, and the stimulation of intestinal absorption must be quantitatively adjusted to the amount secreted; the mechanism of meal-stimulated absorption involves the nutrients of the meal that stimulate electrolyte absorption by different mechanisms (4, 11, 14, 15, 25). In addition, several hormones display antisecretory and proabsorptive effects (5, 7, 9, 16, 17, 20, 24, 28). PYY acting in the upper part of the intestine is secreted in the blood from colonic cells at the beginning of the meal (19). However, it does not seem to display antisecretory effects until 1 h after the beginning of the meal (19). What is striking about sorbin is the link between the intensity of secretion and the intensity of the sorbin response. This suggests that if sorbin is a natural constituent of the human intestine it may play an important role in the economy of water and electrolyte handling. In addition, in diarrheal diseases, the symptoms may be reduced by stimulating colonic absorption in response to secretion. This has already been suggested for human cholera (36) and viral infection in piglets (1). In addition, sorbin has been found to display a beneficial effect in cholera toxin-induced diarrhea in the rat (J. Fioramonti and L. Bueno, personal communication).
At the cellular level, stimulated
Cl secretion has been found
to be associated with a stimulated glucose and
Na+ absorption (31). In
cAMP-induced Cl
secretion,
it has been suggested that the number of
Na+-glucose transporter (SGLT1)
molecules responsible for the
glucose-Na+ cotransport is
increased in the luminal membrane (26). In addition, it has been
reported that methylated casein, an antidiarrheal drug, exhibits
antisecretory activity only in cholera-treated rabbit ileum and has no
effect in control conditions (33). However, we have no direct
indication that a similar mechanism exists to explain the sorbin effect.
Finally, for the following reasons, the present results suggest that C7-sorbin is a good candidate for use in the reduction of water and electrolyte losses in diarrheal diseases: 1) it is mainly active in the distal part of the intestine; 2) its proabsorptive effect is apparent after intestinal secretion has been stimulated; 3) the intensity of its proabsorptive effect is grossly proportional to the intensity of previously induced secretion by a variety of mediators; 4) the size of the heptapeptide makes it feasible for synthesis in industrial environments; 5) there is no evidence of tachyphilaxie; the dose-response curves were similar when cumulative doses were given to the same tissue and when one dose was given to one tissue; and 6) the results of the dose-effect relationship may point to a limitation; apparently, in this in vitro system, the EC50 is one order of magnitude higher than other peptides, such as PYY and SRIF, that have antidiarrheal properties. However, the maximum effect appeared to be essentially the same as that of the most active peptides.
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ACKNOWLEDGEMENTS |
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This study was partly supported by the Institut Henri Beaufour, Avenue du Canada, Les Ulis, France.
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FOOTNOTES |
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The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. §1734 solely to indicate this fact.
Address for reprint requests: M. Boisset, Laboratoire de Biologie, Conservatoire National des Arts et Métiers, 2, rue Conté, 75141 Paris 03, France.
Received 25 February 1998; accepted in final form 14 September 1998.
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