On the Rat Gastric Motility
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The Japanese Journal of Physiology 16, pp.497-508, 1966 ON THE RAT GASTRIC MOTILITY Takesi HUKUHARA AND Toshiaki NEYA Department of Physiology, Okayama University Medical School, Okayama From the results obtained in the experiments carried out on the automa- ticity of the motility of dogs small intestine, HUKUHARA, NAKAYAMA and FU- KUDA8) concluded that the origin of the intestinal motility was of neurogenic nature, that is, rhythmic contractions of the small intestine were maintained by acetylcholine which was spontaneously released from the intramural ganglion cells, including not only their cell bodies, but also their axons. This hypothesis is naturally expected to be applied to the gastric motility. Taking these facts and hypothesis into consideration, a series of experiments has been performed on the gastric motility. The experimental results here reported are concerned with the problems: the localization and specificity of the pacemaker, the difference of behavior of different regions of the stomach and the mechanism underlying these phenomena. As for the gastric peristalsis, the results obtained by investigators until 1924 were summarized by MCCREA et al.14) Since then there could be found only a few literatures4,6,10,11) related with the problems concerned. METHODS In order to observe the movement of the rat stomach in vivo, the well-fed animals weighing from 80 to 200 g were anesthetized with the intraperitoneal administration of 50 mg/kg pentobarbital sodium (Nembutal, ABBOT). It was characteristic that the movement of the rat stomach was not impaired despite administering such a large dose of the drug as described above. The animal was then set in supine position to the frames installed in the internal space of the double-walled trough, the lumen of the wall being irrigated with water appropriately warmed to keep the temperature of the space at about 37•Ž. After the abdominal cavity had been opened wide by the midline incision of the abdominal wall, the incised edges of the wall and the limb of the liver were connected at several spots to the frames by means of of cotton threads or serrefines in such a way that the ventral surface of the stomach was exposed to view. The abdominal cavity was then filled with Tyrode solution which was modified after TAKEMASA16)(1957) as follows: 0.221M-NaCl, 0.0028M-KCl, 0.0015M-CaCl2, 0.0024M-NaHCO3, 0.00032M-NaH2PO4 and 0.0101M-Glucose. For the purpose of observing the movements of the stomach in vitro the whole Received for publication February 8, 1966. *福 原 武 ,祢 屋 俊 昭 497 498 T. HUKUHARA AND T. NEYA stomach was isolated from the animal, which was not fed for 24 to 48 hours before the experiments, and immersed in the bath filled with Tyrode solution which was warmed to the constant temperature of 37•Ž and bubbled with oxgen gas . Leaving the small amount of original gastric content, the glass cannulae were inserted into the esophageal and duodenal lumens, respectively. The former was connected by means of a vinyl tube to a small, double-walled glass vessel which was filled with Tyrode solution and located as high as 5 cm H2O above the level of the stomach , while the latter was located at the level of the stomach. This Tyrode solution was kept at the temperature of about 37•Ž by irrigating the lumen of the wall of the vessel with water appropriately warmed. The gastric movement in vitro was observed sometimes on the dorsal side and sometimes on the ventral side of the stomach by means of a binocular dissecting microscope with low magnifications. Occasionally, the gastric motility was recorded by means of levers assorted with strain gauge transducers , the levers being set up at the two different points of the gastric surface, several millimeters apart from each other along its curved long axis. In order to study the features of the movements of various regions of the stomach strips consisting of the whole gastric layers, about 15 mm long and 1.5 mm wide, were made from the various regions of the stomach and immersed in Tyrode solution cooled to 3•‹ to 5•Ž (See FIG. 1): A) pacemaker strip which included the root of the left gastric artery and the spot at which the 2nd and the 3rd branches of artery bifurcated, B) body strip which was made from the region of the body distal to the pacemaker region, C) pyloric strip which was made from the pyloric part of the stomach distal to the angular incisure, D) sphincter strip which was made from the pyloric sphincter and E) fundus strip which was made from the pars proventricularis, to record their movements. The strips were suspended in the Tyrode bath, the temperature of which was kept constant at 37•Ž. Drugs used were physostigmine sulfate (Merck) and acetylcholine chloride (Daiichi Seiyaku). FIG. 1. Ventral view of rat stomach. In the diagram are shown the arteries providing the blood supply to the stomach. The pacemaker area is hatched. The areas surrounded by broken lines show those from which the strip preparations were made. ON THE RAT GASTRIC MOTILITY 499 RESULTS 1) Observations of the rat stomach in vivo. Rat stomach was supplied with the left and right gastric arteries as shown in FIG. 1: At the site just distal to the cardia, the left gastric artery bran- ched into two main branches, the ventral and dorsal ones, each of which then diverged into small branches spreading radially over the corresponding surfaces of the stomach. In the present experiments they were, for convenience sake, named V1, V2-V6 and D1, D2-D6, respectively, according as to whether they supplied the ventral or dorsal surfaces, and were arranged from the small curvature to the pars proventricularis. In addition, two isolated branches came from the left gastric artery on its way to the cardia to supply the pars proventricularis, and the right gastric artery supplied the pyloric part as well as the region of the corps ventriculi adjacent to the the greater curvature. Observing the ventral surface of the stomach, it was noted that the gas- tric peristalsis always started from the cross-sectional area a few millimeters wide in which the root of the left gastric artery and the point where branch V3 of the artery ramified from V2 were included (hatched area shown in FIG. 1). This area would be conveniently referred to as the pacemaker area. It was characteristic that every spot involved in this band of contraction almost simultaneously contracted. Every point of the band then caudally propagated with the same speed along the long axis of the stomach until it terminated at the pylorus, without, however, propagating beyond the pylorus to the duodenum. The peristaltic waves repeatedly appeared with a regular time interval of about 10 seconds. As the cardia was about 12 mm apart from the pylorus and the time elapsed in a whole swing was estimated at about 12 seconds, the velocity of the peristalsis was estimated at about 1 mm/sec. The peristaltic wave was considerably strong in the whole course along the lesser curvature, while along the greater curvature it was weak until it arrived at the angular incisure but then the nearer the wave approached the sphincter the stronger it became. On the other hand, as soon as the peristaltic wave nearly reached the middle of the pyloric part, the next one appeared at the pacemaker area, two bands of contractions being thus distinctly seen at the same time on the stomach. In FIG. 2 is shown the course of the gastric peristalsis. The pars proventricularis was apparently motionless. In addition, so far as the site of initiation of peristaltic waves was concerned, the bilateral severance of vagus and/or splanchnic nerves had no effect. 2) The movements of the stomach in vitro. A) On the peristalsis of the whole isolated stomach. Within the first 10 500 T. HUKUHARA AND T. NEYA . 2. Pictures indicating the course of the peristaltic waves of rat stomach in vivo. The stomach is view- ed from the ventral side. The pic- tures are taken with every 2 seconds intervals. Arrows show shallow de- pressions produced as the peristalsis sweeps down. Note that two depres- sions are seen at the same time in the pictures 1, 2, 5 and 6. minutes after the lumen of the isolated stomach had been parfused, the peris- taltic waves as usual appeared similarly to those observed in the stomach in vivo. And a minute amount of gastric content was expelled through the duodenal cannula, every time when the peristaltic wave reached the pylorus. On observing the stomach on its dorsal surface the gastric peristalsis showed the mirror image of that observed on the ventral surface. It was worthy to note that in this condition of the experiment the pacemaker shift occurred: FIG ON THE RAT GASTRIC MOTILITY 501 the transverse band of contraction so faint as one might overlook appeared sometimes at the midportion and sometimes at the top of the pars prov- entricularis, propagating distally to reach the pylorus. By utilizing the device described in Chapter 'Methods', the change of motility associated with the peristaltic wave was recorded at two different points which were several millimeters apart from each other along the curved long axis of the stomach. These mechanograms allowed to estimate accurately the velocity of the conduction as well as the period of the peristaltic contrac- tions. The example is shown in FIG. 3. In this case the period and the FIG. 3. Mechanograms of the peristalsis of the whole isolated rat stomach. Utilizing the levers assorted with strain gauge transducers the gastric peristalses were recorded at pacemaker area (A) and pyloric area (B) which were 11 millimeters apart from each other.