No abstract

Slow waves are known to originate orally in the stomach and to propagate toward the antrum, but the exact location of the pacemaker and the precise pattern of propagation have not yet been studied. Using assemblies of 240 extracellular electrodes, simultaneous recordings of electrical activity were made on the fundus, corpus, and antrum in open abdominal anesthetized dogs. The signals were analyzed off-line, pathways of slow wave propagation were reconstructed, and slow wave velocities and amplitudes were measured. The gastric pacemaker is located in the upper part of the fundus, along the greater curvature. Extracellularly recorded slow waves in the pacemaker area exhibited large amplitudes (1.8 +/- 1.0 mV) and rapid velocities (1.5 +/- 0.9 cm/s), whereas propagation in the remainder of the fundus and in the corpus was slow (0.5 +/- 0.2 cm/s) with low-amplitude waveforms (0.8 +/- 0.5 mV). In the antrum, slow wave propagation was fast (1.5 +/- 0.6 cm/s) with large amplitude deflections (2.0 +/- 1.3 mV). Two areas were identified where slow waves did not propagate, the first in the oral medial fundus and the second distal in the antrum. Finally, recordings from the entire ventral surface revealed the presence of three to five simultaneously propagating slow waves. High resolution mapping of the origin and propagation of the slow wave in the canine stomach revealed areas of high amplitude and rapid velocity, areas with fractionated low amplitude and low velocity, and areas with no propagation; all these components together constitute the elements of a gastric conduction system.

In an anesthetized, open-abdomen, canine model, the propagation pattern of the slow wave and its direction, velocity, amplitude, and frequency were investigated in the small intestine of 8 dogs. Electrical recordings were made using a 240-electrode array from 5 different sites, spanning the length of the small intestine. The majority of slow waves propagated uniformly and aborally (84%). In several cases, however, other patterns were found including propagation in the oral direction (11%) and propagation block (2%). In addition, in 69 cases (3%), a slow wave was initiated at a local site beneath the electrode array. Such peripheral pacemakers were found throughout the entire intestine. The frequency, velocity, and amplitude of slow waves were highest in the duodenum and gradually declined along the intestine reaching lowest values in the distal ileum (from 17.4+/-1.7 c/min to 12.2+/-0.7 c/min; 10.5+/-2.4 cm/s to 0.8+/-0.2 cm/s, and 1.20+/-0.35 mV to 0.31+/-0.10 mV, respectively; all p<0.001). Consequently, the wavelength of the slow wave was strongly reduced from 36.4+/-0.8 cm to 3.7 +/- 0.1 cm (p<0.001). We conclude that the patterns of slow wave propagation are usually, though not always, uniform in the canine small intestine and that the gradient in the wavelength will influence the patterns of local contractions.

The pattern of propagation of slow waves in the small intestine is not clear. Specifically, it is not known whether propagation is determined by a single dominant ICC-MP (Interstitial cells of Cajal located in the Myenteric Plexus) pacemaker unit or whether there are multiple active pacemakers. To determine this pattern of propagation, waveforms were recorded simultaneously from 240 electrodes distributed along the whole length of the intact isolated feline small intestine. After the experiments, the propagation patterns of successive individual slow waves were analysed. In the intact small intestine, there was only a single slow wave pacemaker unit active, and this was located at or 6-10 cm from the pyloric junction. From this site, slow waves propagated in the aboral direction at gradually decreasing velocities. The majority of slow waves (73%) reached the ileocaecal junction while the remaining waves were blocked. Ligation of the intestine at one to four locations led to: (a) decrease in the distal frequencies; (b) disappearance of distal propagation blocks; (c) increase in velocities; (d) emergence of multiple and unstable pacemaker sites; and (e) propagation from these sites in the aboral and oral directions. In conclusion, in the quiescent feline small intestine a single pacemaker unit dominates the organ, with occasional propagation blocks of the slow waves, thereby producing the well-known frequency gradient.

High resolution electrical mapping in the gastrointestinal system entails recording from a large number of extracellular electrodes simultaneously. It allows the collection of signals from 240 individual sites which are then amplified, filtered, digitized, multiplexed and stored on tape. After recording, periods of interest can be analysed and the original sequence of activity reconstructed. This technology, originally developed to study normal rhythms and abnormal dysrhythmias in the heart, has been modified to allow recordings from the gastrointestinal tract. In this report, initial results are presented describing the origin and propagation of the slow wave in the isolated stomach and the isolated duodenum in the cat. These results show that in both organs it not uncommon to have more than one focus active during a single cycle. The conduction of slow waves from such a multiple pacemaker environment can become quite complex, and this may play a role in determining the contractile pattern in these organs.

Previous studies have reported on propagation of individual spikes in isolated segments of the pregnant uterus, but there is no information on patterns of spike propagation in the intact organ. There is also no information on propagation of myometrial burst. The aim of this study was to record, at high resolution, patterns of propagation of electrical activities in the pregnant uterus. Sixteen timed-pregnant guinea pigs were euthanized at term, and their uteruses isolated. Fetuses were removed and replaced by an equal amount of Tyrode. A 240-electrode array was positioned at various locations along the organ, all signals were recorded simultaneously, and the electrical propagations were reconstructed. In the intact pregnant uterus at term, spikes propagated with high velocity in longitudinal (6.8 Ϯ 2.4 cm/s) and slower velocity in circular direction (2.8 Ϯ 1.0 cm/s; P Ͻ 0.01). Direction of propagation and frequency of activity were highly variable but showed similar patterns at the ovary or cervical end and along the anterior, posterior, and antimesometrial borders. Along mesometrium, spike propagation was sparse and fractionated. Migration of burst (0.6 Ϯ 0.4 cm/s) was significantly much slower than that of individual spikes (P Ͻ 0.001). Initial burst activity was located at variable locations along the ovarial end of the antimesometrial border, while the latest excitation occurred at the cervical end (1.2 Ϯ 0.9 min). In conclusion, high resolution electrical mapping of the intact pregnant uterus reveals fundamental properties in spatial and temporal patterns of spike and burst propagation that determine the contraction of the organ. spikes; myometrial burst migration ELECTRICAL ACTIVITY in the pregnant myometrium is characterized by phasic bursts of action potentials (spikes), which could be based on cyclic changes in transmembrane potential (24). Cyclic changes in potential resemble in some respects slow waves in the intestines and presumably propagate through the myometrium, similar to the propagation of the intestinal slow waves. In both cases, the depolarization induced by the (slow) waves initiate the opening of L-calcium channels leading to the occurrence of spikes. In the intestines, it is possible to record both slow waves and spikes with extracellular electrodes, and this made it possible to reconstruct the pattern of propagation of both signals and to study the complex interaction between these two electrical waveforms (22).In the myometrium, the basic electrical wave is too slow or its magnitude too small to be recorded extracellularly (9). The spikes, however, are visible in extracellular recordings and this led to several studies on their behavior. Miller et al. (25) analyzed spike propagation in pregnant uterine segments that were 3 ϫ 1 cm in size and showed that velocity in the longitudinal direction was much faster than in the circumferential direction. Lammers et al. (20) found similar results in equally small segments (8 ϫ 2 cm). Most of these studies were performed on isolated segments of...

Lammers WJ, Stephen B, Karam SM. Functional reentry and circus movement arrhythmias in the small intestine of normal and diabetic rats. Am J Physiol Gastrointest Liver Physiol 302: G684 -G689, 2012. First published December 29, 2011; doi:10.1152/ajpgi.00332.2011.-In a few recent studies, the presence of arrhythmias based on reentry and circus movement of the slow wave have been shown to occur in normal and diseased stomachs. To date, however, reentry has not been demonstrated before in any other part of the gastrointestinal system. No animals had to be killed for this study. Use was made of materials obtained during the course of another study in which 11 rats were treated with streptozotocin and housed with age-matched controls. After 3 and 7 mo, segments of duodenum, jejunum, and ileum were isolated and positioned in a tissue bath. Slow wave propagation was recorded with 121 extracellular electrodes. After the experiment, the propagation of the slow waves was reconstructed. In 10 of a total of 66 intestinal segments (15%), a circus movement of the slow wave was detected. These reentries were seen in control (n ϭ 2) as well as in 3-mo (n ϭ 2) and 7-mo (n ϭ 6) diabetic rats. Local conduction velocities and beat-to-beat intervals during the reentries were measured (0.42 Ϯ 0.15 and 3.03 Ϯ 0.67 cm/s, respectively) leading to a wavelength of 1.3 Ϯ 0.5 cm and a circuit diameter of 4.1 Ϯ 1.5 mm. This is the first demonstration of a reentrant arrhythmia in the small intestine of control and diabetic rats. Calculations of the size of the circuits indicate that they are small enough to fit inside the intestinal wall. Extrapolation based on measured velocities and rates indicate that reentrant arrhythmias are also possible in the distal small intestine of larger animals including humans. slow wave; diabetes; wavelength THE SMOOTH MUSCLE LAYER OF the gastrointestinal (GI) tract generates spontaneous electrical activities that are referred to as "slow waves." These slow waves are dependent on the interstitial cells of Cajal in the myenteric plexus (ICC-MY) and are important for maintaining normal GI motility (8, 9).In a recent study, we investigated the propagation of slow waves in a streptozotocin (STZ) model of diabetes in rats. The purpose of that study was to correlate the expected decrease in slow wave propagation with the reported decrease in the census of ICC-MY in the small intestine (8,10,20,30). To our initial surprise, we did not measure a significant decrease in slow wave propagation although there was a reduction in the number of ICC-MY by ϳ50% (10). We concluded from that study that the reduction of ICC-MY was not enough to block slow wave propagation.Further analysis of the propagation in both control and in 3-and 7-mo diabetic rats, however, revealed another unexpected result. In ϳ15% of the segments, slow wave propagation did not originate from one or a few foci (10, 11). Instead, the slow wave was seen to propagate in a circular fashion for relatively long periods of time, whereby the impulse could rotate ...

. Longitudinal and circumferential spike patches in the canine small intestine in vivo. Am J Physiol Gastrointest Liver Physiol 285: G1014-G1027, 2003. First published July 3, 2003 10.1152/ajpgi.00138. 2003In an open-abdominal anesthetized and fasted canine model of the intact small intestine, the presence, location, shape, and frequency of spike patches were investigated. Recordings were performed with a 240-electrode array (24 ϫ 10, 2-mm interelectrode distance) from several sites sequentially, spanning the whole length of the small intestine. All 240 electrograms were recorded simultaneously during periods of 5 min and were analyzed to reconstruct the origin and propagation of individual spikes. At every level in the small intestine, spikes propagated in all directions before stopping abruptly, thereby activating a circumscribed area termed a "patch." Two types of spikes were found: longitudinal spikes, which propagated predominantly in the longitudinal direction and occurred most often in the duodenum, and a second type, circumferential spikes, which propagated predominantly in the circular direction and occurred much more frequently in the jejunum and ileum. Circumferential spikes conducted faster than longitudinal spikes (17 Ϯ 6 and 7 Ϯ 2 cm/s, respectively; P Ͻ 0.001). Circumferential spikes originated in Ͼ90% of all cases from the antimesenteric border, whereas longitudinal spikes were initiated all around the circumference of the intestinal tube. Finally, the spatial sequence of spike patches after the slow wave was very irregular in the upper part of the intestine but much more regular in the lower part. In conclusion, spikes and spike patches occur throughout the small intestine, whereas their type, sites of origin, extent of propagation, and frequencies of occurrence differ along the length of the small intestine, suggesting differences in local patterns of motility. slow waves; spike patches; duodenum; jejunum; ileum INTESTINAL MOTILITY IS INITIATED by slow waves and by action potentials (spikes) that may or may not occur in the wake of the slow wave (6,18,19,23). Several studies have investigated the temporal relationship between slow waves and spikes and between spikes and motility (2,3,5,7,16,25,26). Not much attention, however, has been given to the spatial pattern of spike propagation (6, 17) or to possible regional variations along the small intestine. Recently, it was shown that spikes propagate for a limited extent in time and space before terminating spontaneously, thereby activating a relatively small area termed a spike "patch" (11). These patches were demonstrated in isolated tissues in vitro, in a single species, the cat, and in one part of the intestine, the duodenum. The question therefore arises as to whether spike patches also occur in another species, in the whole organism, and in other parts of the small intestine.We are now presenting an approach enabling us to record in vivo electrical signals from 240 extracellular sites simultaneously from the serosal surface of the intact c...