The appearance of tonic constrictions.—It has already been noted that as the food accumulates in the ascending colon it is at first confined to this region by antiperistaltic waves. With further accessions, however, the contents naturally must be pressed more and more into the transverse and descending colon. In the early stages of this accumulation, while the food lies chiefly in the ascending colon, the only activity of the muscular walls is the antiperistalsis. As the contents extend along the intestine a deep constriction appears near the advancing end and nearly separates a globular mass from the main body of the food (Fig. 6). The contents of the large intestine progress farther and farther from the cæcum; meanwhile new tonic constrictions appear which separate the contents into a series of globular masses. And as the number of these divisions increases they take a position farther from the cæcum, so that they are present chiefly in the descending colon (Fig. 7). Raiser has recorded a similar appearance in the terminal portion of the rabbit’s colon, in which deep circular constrictions separate the scybalous masses. He maintains that these masses are pushed onward by the constrictions. Comparing tracings made at rather long intervals (forty-five minutes), I found that the rings disappear from the transverse colon, and then are present with the waste material in the descending colon. Thus in the cat also these rings, which seem with short observation to be remaining in one position, are in reality moving slowly away from the cæcum, pushing the hardening contents before them. The contents at this stage are no longer fluid, and consequently they must offer considerable resistance to a force pushing them through the colon. It is an advantage to have this pultaceous substance propelled in divisions rather than in a uniformly cylindrical mass, since the fibres along the length of the mass are thereby rendered effective. Such are the functions of the persistent rings; they form the waste matter into globular masses at the end of the transverse colon and slowly push these masses onward.
Figure 7.—Radiograph showing the region of tonic constrictions (descending colon) and the region of antiperistalsis (transverse and ascending colon).
In the transverse colon, which is free from the slowly moving rings, the antiperistaltic waves have full sway. In the region of the tonic rings an infrequent or even a slowly periodic relaxation and contraction are often to be observed. These changes seem to take place in all the rings at about the same time. Once I saw antiperistaltic waves running over the uppermost of four segments, but since the rings on either side of the segment held tightly, the waves had merely the effect of churning the material of the segment and did not move it onward. Inasmuch as the material in these segments at first is soft, so that the segments are easily compressible, while the fæcal masses which are the final result are relatively hard and dry, it follows that even within the confines of these persistent rings some absorption is taking place.
Defecation
The process of clearing the colon is a process of repeated reduction of the amount of material present. Figure 8 (3.11) is a radiograph showing the food in the colon at 3.11 P.M. About 3.25, with a slow, sweeping movement, the gut swung around so that the ascending colon was lying in the position of the last half of the transverse colon, and the transverse colon had taken the position of the descending part (Fig. 8, 3.25). At the same time the tonic constrictions disappeared and were replaced by a strong, broad contraction of the circular muscle, tapering the contents off on either side in two cones. The region of strongest contraction was apparently drawn downward with the rest of the gut by a shortening of the descending colon. As the intestine swung around, more material was forced into the rectum, and when the swinging of the intestine stopped, the constriction which divided the lumen passed slowly downward, and with the aid of the muscles surrounding the abdominal cavity, pushed the separated mass out of the canal.[35] After the terminal mass had thus been pushed out, the colon with the remainder of its contents returned to nearly its former position (Fig. 8, 3.46). About two hours afterward this remnant had been spread throughout the length of the large intestine by means of the slowly moving rings. Figure 7 is a radiograph of the same colon pictured in Figure 8; the radiograph was taken at 11.50 A.M., and at 12.15 P.M. the material in the lower descending colon was forced out in the manner above described. Within three hours the remaining portion had been spread into the evacuated region, as shown in Figure 8, 3.11. The manner in which the material is spread from the region of the antiperistaltic waves into the region of the slowly advancing rings presents a problem. During normal living new food constantly arriving in the colon must force the old contents forward just as the later parts of a meal force forward the earlier parts; there is no doubt, however, that most of the contents of the cæcum and the ascending colon may be passed onward even during starvation. The emptying of these regions, according to my observations, is never complete; for after considerable time has elapsed and the large intestine is cleared and dilated with gas, some substance is still to be detected in the cæcum and clinging to the walls of the ascending colon. The only activities manifested here are the antiperistaltic waves and the strong tonic contraction of the whole circular musculature shown in Figure 6. It is clear that the latter activity would serve to press into the transverse colon a considerable portion of the contents of the ascending colon, and the remnant seen clinging to the walls would be the part not thus pressed forward.
Figure 8.—Two radiographs and a tracing showing the changes taking place in defecation. 3.11, material in the colon. 3.25, colon carried downward and terminal mass separated. 3.46, after defecation, when the colon returns to former position. Defecation occurred at 3.27.
Twice I have seen appearances which might account for the emptying of the first portion of the large intestine in a more thorough manner than that above described. At one time, without apparent stimulation, strong tonic contraction occurred along the entire length of the ascending colon, which forced the contents almost wholly into the transverse portion. This action seemed merely an exaggerated form of that observable after food passes the ileocæcal valve (see Fig. 6). At another time, after a mass of food had passed through the ileocæcal valve, after the ascending colon had contracted generally and the antiperistaltic waves had coursed over it in the usual manner, a deep constriction appeared at the valve and ran upward without relaxation nearly the length of the ascending colon, pushing the contents before it. For an instant the wave paused; then the constriction relaxed and the food returned towards the cæcum. These observations indicate that either a general contraction of the wall of the large intestine or a true peristalsis may be effective in pressing waste matter from the region where antiperistalsis is the usual activity into the region where the slowly advancing rings may carry it on to evacuation (see Fig. 7).