MOTOR FUNCTIONS OF THE LARGE INTESTINE IN SHEEP VERSUS CATTLE

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1 MOTOR FUNCTIONS OF THE LARGE INTESTINE IN SHEEP VERSUS CATTLE J. Fioramonti, Marie-Françoise Hubert To cite this version: J. Fioramonti, Marie-Françoise Hubert. MOTOR FUNCTIONS OF THE LARGE INTESTINE IN SHEEP VERSUS CATTLE. Annales de Recherches Vétérinaires, INRA Editions, 1980, 11 (1), pp <hal > HAL Id: hal Submitted on 1 Jan 1980 HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific research documents, whether they are published or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers. L archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

2 MOTOR FUNCTIONS OF THE LARGE INTESTINE IN SHEEP VERSUS CATTLE J. FIORAMONTI Marie-Françoise HUBERT!.a&ofafo/re (/e P/7/x/o/06f/e, Fco/e /Var/ona/e!ëf!/v!a/re, Laboratoire 23, chemin des Physiologie, Capelles, Ecole Toulouse Nationale Cedex, Vétérinaire, France Résumé ACTIVITÉ MOTRICE COMPARÉE DU GROS INTESTIN CHEZ LE MOUTON ET LA VACHE. - La motricité des différents segments du gros intestin (caecum, côlon proximal, spiral et distal) a été comparée chez 3 brebis, 2 vaches et 3 veaux avant et après sevrage à partir des électromyogrammes. Ceux-ci sont caractérisés en l absence d un rythme électrique de base par la présence de 2 types de salves de potentiels : des salves de longue durée (5 à 13 s), localisées au caecum et au côlon proximal, et des salves de courte durée (3 à 4 s), surtout présentes au niveau du côlon spiral. Le côlon distal présente chez la brebis mais non chez la vache les deux types d activité. Une autre différence entre les deux espèces est une activité permanente du type salves de courte durée au niveau du côlon spiral chez le mouton (95% au lieu de 25% chez la vache du temps d enregistrement). La vache et le veau après sevrage présentent, toutes les 2,5 à 4 heures, des périodes d activité continue durant 6 min et propagées lentement depuis le côlon proximal jusqu au côlon distal. Le rôle de ces profils moteurs spécifiques du côlon est envisagé par rapport à la formation des fèces. The large intestine of ruminants comprised the following regions : caecum, proximal colon, centripetal and centrifugal.coils of the spiral colon, distal colon and rectum in both sheep and cattle (Ulyatt et al., Information on its motor functions is restricted to recent studies of the motility of the caecum and the proximal colon in sheep (Mac Rae et a/., 1973 ; Fioramonti and Ruckebusch, 1978). However, an important role of the spiral colon seemed involved in the formation of pellets in sheep for which the colonic retention time averaged 20 hours instead of only 9 hours in cattle (Hecker and Grovum, 1975). That cattle excrete moister faeces than sheep despite an identical length of the large intestine and a greater rate of water absorption in the cow might thus be linked to peculiar patterns of motility of the spiral colon. The aim of this study was to compare the colonic motility in sheep and cows and in calves before and after weaning in order to emphasize the factors involved in the different aspects of faeces and in the rate of passage of digesta. Material and Methods Three adult Lacaune ewes, two adult dry Francaise Frisonne Pie Noire cows and three calves of the same breed at 10 days of age at

3 the beginning of the experiments were used. Ewes and cows were fed ad libitum on hay diet. The calves received reconstituted full cream cow s milk (60 g dry matter/kg live weight, daily) in two meals. At five weeks of age, the amount of milk was progressively reduced over a period of 10 days and hay was given ad libitum. In all animals, eight pairs of intestinal electrodes made of insulated nichrome wire were implanted under halothane anaesthesia. The pairs of electrodes, 2 mm apart, were positionned on the ileum at 50 cm from the ileo-caecal valve (ICV), on the caecum near the ICV, on the proximal colon at about half distance between the ICV and the spiral colon, on the 1st, 3rd and 5th coil of the spiral colon, on the distal colon at about 50 cm from the rectum. In the two cows four additional groups of electrodes were implanted at 15 cm intervals on the 3rd coil of the spiral colon. After surgery animals were placed in individual cages. Elec-

4 tromyograms were recorded for 3 days per week during 8-12 weeks on an e.e.g. machine (Reega VIII, Alvar, Paris). In addition, the electrical activity of four chosen sites was automatically plotted each 20 by means of a four-line integrator on a potentiometric recorder (Latour, 1973). Results Sheep The electrical activity of the caecum and of the proximal colon already described (Fioramonti and Ruckebusch, 1979) consisted of

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6 long spike burst (LSB) potentials lasting about 6 s, propagated in either direction and recurring at a frequency of 0.7/min at the blind pole of the caecum, 1.1/min near the ICV and 1.5/min on the proximal colon. Periods of hyperactivity characterized by 4-12 spike bursts in series, lasting 1-3 min and followed by an inhibition for min occurred at intervals of 2-3 hours (fig. 11. These periods were detected when the phases of regular spiking activity (RSA) of the migrating myoelectric complexes (MMC) propagated along the terminal ileum (fig. 2). On the spiral colon, short spike burst (SSB) potentials lasting about 4 s occurred at a rate of about 15 per min during nearly 95% of the recording time so that the activity seemed continuous (table 11. This basal activity was disrupted on the 1st coil by strong spike bursts (8.5/h) propagated (about 3 cm/s) from the proximal colon and followed by quiescence during 10 to 30 s. Only 40% of these bursts reached the last centripetal coil. The electrical activity of the distal colon exhibited LSB potentials lasting about 7 s at the rate of 5/min and SSB potentials lasting 3 s and grouped in periods of 30 to 60 s in duration. The occurrence of these spiking activities was always very irregular so that no profile could be defined. Cow The electrical activity of the caecum and the proximal colon was similar to that observed in sheep. The major differences consisted of an increased duration of the spike bursts of the caecum ( s vs s) and of the periods of hyperactivity of the caecum ( min vs min) linked to the occurrence of a phase of regular spiking activity of an MMC on the terminal ileum (table 2). The electromyogram of the spiral colon consisted during 25% of the recording time of SSB potentials lasting about 4 s at a mean rate of 7.0 per min. On the distal colon only LSB potentials, 8.1 ± 1.7 s in duration at a mean rate of 6 per min, were recorded. A striking feature of the spiral and distal colon shown in figure 3 is the propagation of 8-9 phases per day of hyperactivity which lasted 5 min and were followed by a period of quiescence of nearly 30 min. The velocity of propagation averaged 2.1 ± 0.4 cm/min from the proximal colon to the distal colon. Calf Electrical activity of the large intestine in the calf before weaning differed from that after weaning. The mean frequency of spike bursts was very low (0.6 ± 0.1 /min) on the caecum and the proximal colon (table 11. The number of phases of hyperactivity of the caecum and proximal colon linked to the presence of ileal RSA phases was reduced. The periods of migrating activity from the spiral to the distal colon were irregular as well as their velocity of propagation which varied from 1 to 10 cm per min. During weaning, the motor profile of the large intestine became similar to that observed in cow within 10 days, although only 2/3 of the RSA phases of the MMCs were accompanied by motor responses of the caecum. Discussion The motor functions of the colon in ruminants are characterized by two patterns of colonic spiking activity, long and short spike

7 burst potentials, the SSB potentials being predominant in the spiral colon of sheep (fig. 4). The presence in cattle and in weaned calves of periods of migrating spiking activity along the spiral and distal colon are also of importance. The duality of spiking activity of the colon seems to be a general phenomenon already described in the rabbit (Ruckebusch and Fioramonti, 1976), the pig (Fioramonti and Bueno, 1980), the dog (Garcia-Villar, 1979) and human (Bueno et al., 1980). Long spike burst potentials are characterized by their duration and propagation. Short spike burst potentials lasting less than 5 s occurred frequently in series and remained localized. By contrast to the «mixed» pattern of SSB and LSB potentials observed in the species previously described, each segment of the ruminant large intestine exhibited its own characteristics of spiking activity. The caecum, the proximal colon and the distal colon presented mainly LSB potential bursts. SSB potentials largely predominated on the spiral colon. The motor function of the caecum previously studied by means of intraluminal balloons in cattle was described as contractions at a rate of 1.4 per min (Svendsen and Kristensen, A lower rate of contraction (1.2 per min) was found here. Nevertheless, the motility of bovine large intestine is specially characterized by the aboral propagation of periods of continuous spiking activity along the colon. Such an activity has never been reported for the colon of other species. The periods resemble the phases of RSA of the myoelectric complexes of the small bowel, especially because they are followed by a period of quiescence. However, their velocity of propagation is nearly 10 times lower than that seen in the bovine jejunum (Ruckebusch and Bueno, 1973). That in calf the frequency of caecal contractions suddenly increased after weaning, in association with an increased volume of nutrients is in accordance with a control of caecal motility by the volume of its contents (Fioramonti and Ruckebusch, 1979). Moreover, an increased residue diets seems to be important for the coordination of caecal motility since the frequency of periods of hyperactivity increased after weaning. Finally, differences in the motility patterns between sheep and cattle are in accordance with the differences in colonic transit time between these species. The relative low level of spiking activity observed in the bovine spiral 95% in colon (25% of recording time vs. sheep) could explain a shorter colonic transit time and lower storage function in cattle. Conversely, the nearly permanent spiking activity of the ovine spiral colon may be related to the formation of pellets. The specific migrating spiking activity of the bovine colon may be related to the propulsion towards the rectum of large amounts of residue diets. Accepted for publication September 28 th Summary The electrical activity of the caecum and of the different parts of the colon was recorded from chronically implanted electrodes in the ovine and bovine species, the latter including cattle and calves before and after weaning. In the both species, colonic spiking activity consisted of spike bursts lasting from 5 to 13 s localized to the caecum and proximal colon, and of spike bursts las-

8 ting 3 to 8 s, localized to the spiral colon. An intermingled pattern of long and short spike bursts was recorded on the distal colon in sheep but not in cattle. Another major difference between the two species was a permanent short spike burst activity in the ovine spiral colon (95 versus 25% of the recording time in cattle). Periods of continuous spiking activity lasting 6 min, migrated from the proximal colon to the distal colon in the cows and calves but not in sheep. The role of the different patterns of colonic activity is discussed in terms of colonic motor functions involved in the formation of faeces. References BUENO L., FIORAMONTI J., RUCKEBUSH Y., FREXINOS J., COULOM P., An evaluation of colonic myoelectrical activity in health and functional diseases. Gut, in press. FIORAMONTI J., BUENO L., Motor activity in the large intestine of the pig related to dietary fibre and retention time. Br. J. Nutr., 43, FIORAMONTI J., RUCKEBUSCH Y., On the control of caecal motility in sheep. Ann. Rech. Vét., 9, FIORAMONTI J., RUCKEBUSCH Y., Diet and caecal motility in sheep. Ann. Rech. Vét., 10, GARCIA-VILLAR R., Recherches sur la motricité colique chez le chien. These doct. 3e cycle, Toulouse, n 45, 126 p. HECKER J.F., GROVUM W.L., Rates of passage of digesta and water absorption along the large intestines of sheep, cows and pigs. Aust. J. Biol. Sci., 28, LATOUR A., Un dispositif simple d analyse quantitative de I electromyogramme intestinal chronique. Ann. Rech. V!t., 4, MAC RAE J.C., REID C.S.W., DELLOW D.W., WYBURN R.S., Caecal cannulation in the sheep. Res. Vet. Sci., 14, RUCKEBUSCH Y., BUENO L., The effect of weaning on the motility of the small intestine in the calf. Br. J. Nutr., 30, RUCKEBUSCH Y., FIORAMONTI J., The Fusus coli of the rabbit as a pacemaker area. Experientia, 32, ULYATT M.J., DELLOW D.W., REID C.S.W., BAUCHOP T., Structure and function of the large intestine of ruminants. /n Mc Donald I.W., Warner A.C.K., Digestion and metabolism in the ruminant, pp , Armidale, Australia. SVENDSEN P., KRISTENSEN B., Cecal dilatation in cattle. Nord. Vet. Med., 22,