Tension Receptors Research Articles

Morphological relationships of choleragenoid horseradish peroxidase‐labeled spinal primary afferents with myenteric ganglia and mucosal associated lymphoid tissue in the cat esophagogastric junction

The goal of the present study was to gain insight into the environmental factors influencing the activity of primary spinal afferent fibers in the different layers of the esophagogastric junction of the cat and, thus, to analyze the relationships of these afferents with various cellular components. Spinal primary afferent fibers were selectively labeled by anterogradely transported choleragenoid horseradish peroxidase conjugate (B-HRP). B-HRP was injected into the thoracic dorsal root ganglion at the T8-T13 levels. 6-Hydroxydopamine-induced sympathectomy was performed prior to B-HRP injection in order to prevent otherwise unavoidable labeling of sympathetic fibers in the gut wall. Numerous labeled fibers ran between, around, and within the myenteric ganglia. Others crossed the muscle layers directly and entered the mucosa, where some ran near granulocytes and around or through solitary lymphoid follicles. Labeled fibers were observed in the squamous esophageal epithelium but not in the fundic glandular epithelium. The fibers in the myenteric area are probably connected to the muscular tension receptors that have been detected by electrophysiologic techniques. This assumption is based on the observation that only a few fibers appear to terminate in muscle layers and on the fact that the myenteric area is very narrow and subject to powerful forces. Fibers in the myenteric ganglia could be involved in local efferent functions. Fibers in the mucosa could act as nociceptors and might be involved in local immunological responses.

Vagal reflex inhibition of motility in the abomasal body of sheep by antral and duodenal tension receptors.

Vagally-mediated regulation of motility in the abomasal body by duodenal and abomasal antral motility was demonstrated in acutely prepared anaesthetized sheep. The enteric plexuses between the abomasal body, antrum and duodenum were interrupted by transection. Antral contractions were more effective than duodenal contractions at causing inhibition of the abomasal body, and antral isometric conditions were more effective than antral isotonic conditions. Inhibition of motility in the abomasal body was reduced by unilateral cervical vagotomy, was abolished by bilateral cervical vagotomy, and was reversibly inhibited by cervical vagus cold block. The demonstration of vagal pathways in abomaso-abomasal reflexes confirms a functional homology of the mechanisms in the ruminant and animals with simple forms of stomach.

Electrophysiological characteristics of tension receptors in the abomasal antrum of sheep.

Unitary impulse activity from tension receptors in the abomasal antrum in sheep anaesthetized with chloralose was studied with concurrent measurements of isometric tension in the antral wall, intraluminal pressures and electromyography (EMG). Rhythmical background activity had a close correlation with intraluminal pressure changes and usually a close correlation with local increases in EMG activity and isometric tension. Impulse frequencies increased, but were not sustained, following changes from isotonic to isometric conditions. Tension receptors were excited by transmural compression, by antral distension and by local intra-arterial injections of acetylcholine (2.5-25 micrograms), pentagastrin (10-20 micrograms), 5-hydroxytryptamine (1-20 micrograms), adrenaline (10 micrograms) cholecystokinin-8 (1-5 micrograms) or bradykinin (0.5-20 micrograms), and by systemic injections of bethanecol (5-1000 micrograms). Local intra-arterial perfusions of atropine (50 micrograms) and hexamethonium (100 micrograms) did not abolish spontaneous antral movements or affect the enhanced impulse activity evoked by acetylcholine or 5-hydroxytryptamine. Most antral tension receptors were excited by increases in antral tension or passive distension; they therefore functioned 'in series' with the muscularis externa. The discharges of three tension receptors were 'off-loaded' early during the contraction evoked by local injections of pentagastrin (10-20 micrograms).

Bladder wall tension and mechanoreceptor discharge

Bladder wall mechanoreceptors are essential elements in micturition and continence reflexes. While they have been described as tension receptors, their response to bladder wall deformation has always been characterised in terms of pressure. The firing patterns of 10 bladder wall mechanoreceptors were determined during bladder distensions at a natural and a much faster rate. In all units firing rate was higher at any given pressure at the slower bladder distension rate. This inverse rate dependence was reduced when firing rate was related to a derived measure of bladder wall tension and abolished when multi-fibre recordings were used. We conclude that it is important to incorporate volume effects in studies of continence control systems.

Sensory innervation of the viscera: peripheral basis of visceral pain.

Sensory innervation of the viscera: peripheral basis of visceral pain.

Discrimination of transiently applied mechanical loads: breathing vs. pulling

Two groups of 24 subjects each attempted to discriminate between large elastic and resistive loads during 50 randomized presentations of each load. Breathers inspired from the loads through a J valve, whereas pullers reciprocally stroked the plunger of a 2-liter syringe connected to the J valve. A range of load durations was obtained in each subject by prematurely unloading approximately one-half of the trials at graded times from their onset. Breathers produced random discrimination scores [50.8 +/- 2.5% (SE) correct] when loaded inspirations were shorter than unloaded inspirations [trials in which both loads induced equal airway pressure (and probably respiratory muscle tension) waveforms] and nonrandom discrimination scores (65.7 +/- 2.8% correct) when loaded inspirations were longer than unloaded inspirations (trials in which both loads induced different waveforms). In contrast, pullers produced nonrandom discrimination scores (62.2 +/- 2.9% correct) when loaded airflow durations were shorter than unloaded inspirations [trials in which both loads induced equal line pressure (and therefore limb muscle tension) waveforms]. Supplemental audio feedback related to instantaneous airflow (an expression of movement) improved load perception in breathers (to 64.2 +/- 3.0% correct; P < 0.01), indicating that airflow feedback introduced load-specific information that was lacking during breathing but redundant during pulling. In support of this hypothesis, airflow feedback by itself enabled a third group of listeners to identify load type with equal accuracy as pullers but with greater accuracy than breathers. These findings suggest that 1) uniformed subjects rely heavily on feedback from airway pressure and/or muscle tension receptors to perceive added loads to breathing and 2) limb mechanoreceptors provide a more sensitive appreciation of movement than do respiratory mechanoreceptors.

Effects of 5-hydroxytryptamine on discharge of vagal mucosal afferent fibres from the upper gastrointestinal tract of the ferret

This study was undertaken to determine the effects of 5-hydroxytryptamine (5-HT) on vagal mucosal afferent endings and how this may relate to their sensitivity to other stimuli. Single afferent fibres with receptive fields in the mucosa of the upper gastrointestinal tract were recorded from the cervical vagus of Urethane anaesthetized ferrets. The selection criteria included failure to respond to luminal distension (i.e., vagal tension receptors were excluded). All fibres tested responded to mucosal stroking. The majority of these ( 28 32 ) also responded to close-intrarterially applied 5-HT (10 μg) with a brief burst (usually < 15 s) of action potentials, which in 6 6 cases was reduced or abolished by the 5-HT 3 receptor antagonist granisetron (0.1–0.5 mg/kg i.v.), as were four responses to cholecystokinin-octapeptide (100–400 pmol close I.A.). The response to 5-HT was shown to be dose-dependent over the dose range 2–75 μg on six occasions. Responses to luminal stimuli, which included 150 mM HC1, 1 M NaCl, and mucosal stroking, were not blocked by granisetron although in three fibres, the resting discharge was reduced by the antagonist, suggesting that resting discharge in vagal mucosal afferents may be influenced by endogenously released 5-HT.

Sensory innervation of the hairy skin (light- and electronmicroscopic study.

The sense of touch develops early in phylogeny and is one of the most important senses for the survival of the animal. Touch organs of hairy skin in mammals include the so-called "Haarscheiben" (also Pinkus corpuscles) and all types of hair follicles with their nerve endings. The touch organs of the skin consist of a mechanical transducing component and the sensory component. The epithelium and its derivatives like hair follicles and sebaceous glands are the mechanical transducing component transmitting the mechanical forces like pressure or touch to the second component--the sensory nerve endings. In mammalian hairy skin all sinus and guard hairs and many vellus hairs are touch organs. The sinus hair is a typical example of a touch organ. All mammals except humans are equipped with these highly differentiated touch organs. The hair follicle is almost completely embedded in a blood sinus and equipped with more than 2,000 sensory nerve endings. All sinus and guard hairs are equipped with free nerve endings (nociceptors), Merkel nerve endings (slowly adapting [SA I] mechanoreceptor units-pressure detectors), palisades of lanceolate nerve endings (velocity detectors), and pilo-Ruffini corpuscles (tension receptors). In most of the sinus hairs lamellated corpuscles of Pacini type could be found (rapidly adapting receptors-acceleration detectors). Most vellus hairs are equipped with free and lanceolate nerve endings. Some of the vellus hairs of the upper portion of the body (head, upper extremity) are innervated by Merkel nerve endings. The presence of pilo-Ruffini nerve endings in vellus hairs is very unusual.

Sensory innervation of the hairy skin (light-and electronmicroscopic study)

The sense of touch develops early in phylogeny and is one of the most important senses for the survival of the animal. Touch organs of hairy skin in mammals include the so-called “Haarscheiben” (also Pinkus corpuscles) and all types of hair follicles with their nerve endings. The touch organs of the skin consist of a mechanical transducing component and the sensory component. The epithelium and its derivatives like hair follicles and sebaceous glands are the mechanical transducing component transmitting the mechanical forces like pressure or touch to the second component—the sensory nerve endings. In mammalian hairy skin all sinus and guard hairs and many vellus hairs are touch organs.The sinus hair is a typical example of a touch organ. All mammals except humans are equipped with these highly differentiated touch organs. The hair follicle is almost completely embedded in a blood sinus and equipped with more than 2,000 sensory nerve endings. All sinus and guard hairs are equipped with free nerve endings (nociceptors), Merkel nerve endings (slowly adapting [SAI] mechanoreceptor units-pressure detectors), palisades of lanceolate nerve endings (velocity detectors), and pilo-Ruffini corpuscles (tension receptors). In most of the sinus hairs lamellated corpuscles of Pacini type could be found (rapidly adapting receptors-acceleration detectors).Most vellus hairs are equipped with free and lanceolate nerve endings. Some of the vellus hairs of the upper portion of the body (head, upper extremity) are innervated by Merkel nerve endings. The presence of pilo-Ruffini nerve endings in vellus hairs is very unusual.

Mechanoreceptor afferent activity compared with receptor field dimensions and pressure changes in feline urinary bladder.

The relationship between vesical mechanoreceptor field dimensions and afferent nerve activity recorded in pelvic plexus nerve filaments was examined in chloralose-anesthetized cats. Orthogonal receptor field dimensions were monitored with piezoelectric ultrasonic crystals. Reflexly generated bladder contractile activity made measurements difficult, therefore data were collected from cats subjected to actual sacral rhizotomy. Afferent activity was episodic and was initiated at different pressure and receptor field dimension thresholds. Maximum afferent activity did not correlate with maximum volume or pressure. Furthermore, activity was not linearly related to intravesical pressure, receptor field dimensions, or calculated wall tension. Pressure-length hysteresis of the receptor fields occurred. The responses of identified afferent units and their associated receptor field dimensions to brief contractions elicited by the ganglion stimulant 1,1-dimethyl-4-phenylpiperazinium iodide (2.5-20 micrograms i.a.), studied under constant volume or constant pressure conditions, are compatible with bladder mechanoreceptors behaving as tension receptors. Because activity generated by bladder mechanoreceptors did not correlate in a simple fashion with intravesical pressure or receptor field dimensions, it is concluded that such receptors are influenced by the viscoelastic properties of the bladder wall. Furthermore, as a result of the heterogeneity of the bladder wall, receptor field tension appears to offer a more precise relationship with the activity of bladder wall mechanoreceptors than does intravesical pressure.

Treatment of lower urinary tract dysfunction in patients with multiple sclerosis. Committee of the European Study Group of SUDIMS (Sexual and Urological Disorders in Multiple Sclerosis)

Bladder symptoms in patients with multiple sclerosis (MS) are common and usually arise as a result of spinal lesions which interrupt the neural pathways connecting the pontine micturition centre to the sacral spinal cord. Thus these symptoms are particularly likely to occur in those with lower limb neurological deficits. Fortunately bladder dysfunction in MS is rarely associated with serious upper tract disease so that the problem is usually one of symptomatic management. Lower urinary tract symptoms may be both "irritative" or "obstructive" in nature and can be explained in terms of underlying detrusor hyperreflexia and incomplete bladder emptying. Treatment is aimed at minimising both these effects. Oral anticholinergic medication can be effective in reducing detrusor hyperreflexia and intermittent catheterisation is used to reduce abnormally high post micturition residual volumes. With this simple treatment, often used in combination, many less severely affected patients with MS can gain considerable improvement in controlling urinary continence.

An excitatory body-antral reflex in the sheep abomasum.

This paper describes an abomasal body-antral excitatory reflex in acute experiments in sheep anaesthetized with chloralose. The abomasum was either surgically intact or acutely transected to form separate compartments of the abomasal body and antrum. Inflation of a balloon situated in the abomasal body and active contraction of the abomasal body increased the amplitude of antral contraction. In the intact preparation, inflation of the abomasal body increased the amplitude of antral contraction before and after extrinsic denervation. The antral response was reduced by 75% after extrinsic denervation. In the acutely transected preparation, inflation of the abomasal body and electrically induced contraction of the abomasal body also increased antral contraction amplitude. The antral response was abolished by bilateral section of the cervical vagus nerves in five of six experiments. Selective extrinsic neurotomy of either compartment abolished all responses. It was concluded that the neural mechanisms controlling abomasal motility in the sheep resemble those found in species with simple stomachs: excitation of in-series tension receptors of the abomasal body causes an increase in antral contraction amplitude by vago-vagal reflex pathways. Antral motility is also modulated by changes of tension in the abomasal body by intrinsic reflex pathways and reflexes involving abdominal preganglia may also be present. In a restricted number of experiments the splanchnic nerve was shown to play an important role.

Vagal afferent innervation of the rat fundic stomach: morphological characterization of the gastric tension receptor.

Although the gastric tension receptor has been characterized behaviorally and electrophysiologically quite well, its location and structure remains elusive. Therefore, the vagal afferents to the rat fundus (forestomach or nonglandular stomach) were anterogradely labeled in vivo with injections of the carbocyanine dye Dil into the nodose ganglia, and the nerves and ganglia of the enteric nervous system were labeled in toto with intraperitoneal Fluorogold injection. Dissected layers and cryostat cross sections of the fundic wall were mounted in glycerin and analyzed by means of conventional and laser scanning confocal microscopy. Particularly in the longitudinal, and to a lesser extent in the circular, smooth muscle layers, Dil-labeled fibers and terminals were abundant. These processes, which originated from fibers coursing through the myenteric ganglia and connectives, entered either muscle coat and then ran parallel to the respective muscle fibers, often for several millimeters. They ran in close association with the Fluorogold-labeled network of interstitial cells of Cajal, upon which they appeared to form multiple spiny appositions or varicosities. In the myenteric plexus, two different types of afferent vagal structures were observed. Up to 300 highly arborizing endings forming dense accumulations of small puncta similar to the esophageal intraganglionic laminar endings (Rodrigo et al., '75 Acta Anat. 92:79-100) were found in the fundic wall ipsilateral to the injected nodose ganglion. They often covered small clusters of myenteric neurons or even single isolated ganglion cells (mean = 5.8 neurons) and tended to extend throughout the neuropil of the ganglia. In a second pattern, fine varicose fibers with less profuse arborizations innervated mainly the central regions of myenteric ganglia. Camera lucida analyses established that single vagal afferent fibers had separate collaterals in both a smooth muscle layer and the myenteric ganglia. Finally, Dil-labeled afferent vagal fibers were also found in the submucosa and mucosa. Control experiments in rats with supranodose vagotomy as well as rats with Dil injections directly in the distal cervical vagus ruled out the possibility of colabeling of afferent fibers of passage. In triple labeling experiments, in conjunction with Dil labeling of afferents and Fluorogold labeling of enteric neurons, the carbocyanine dye DiA was injected into the dorsal motor nucleus of the vagus to anterogradely label the efferent vagal fibers and terminals. The different distributions and morphological characteristics of the vagal afferents and efferents could be simultaneously compared. In some instances the same myenteric ganglion was apparently innervated by an afferent laminar ending and an efferent terminal.(ABSTRACT TRUNCATED AT 400 WORDS)

Abdominal chemo‐ and mechanosensitivity in ruminants and its role in the control of food intake

Ruminant animals have evolved a large and complex set of stomachs which allow fermentation of fibrous food by symbiotic micro-organisms. These stomachs are well innervated and generate signals which are thought to be important in the control of voluntary food intake. Tension receptors in the muscular wall of the rumen and reticulum are slowly adapting and provide a measure of distension while epithelial receptors are rapidly adapting and provide information on the fibrousness of the digesta; they are involved in the control of stomach motility and voluntary food intake in order to prevent excessive distension. The epithelial receptors are also sensitive to the chemical nature of the digesta, particularly acidity. There are mechano- and chemoreceptors in the abomasum (true stomach) and duodenum and chemoreceptors in the liver, all of which have been implicated in the control of intake. It is relatively easy to prepare and maintain ruminants with a rumen fistula and many studies have shown the effects of such manipulations as distension of balloons in the rumen on voluntary intake. With fibrous, slowly digested feeds intake is primarily limited by rumen distension. With more rapidly digested feeds, however, the products of digestion play an important role in controlling intake. Short-chain fatty acids are the main products of fermentation and infusion of their salts into the rumen depresses food intake to a much greater extent than infusion into the general circulation. Acetate or propionate given into the rumen are more effective, mole for mole, than butyrate but must be given at rates exceeding the natural rate of production in order to have a significant effect. It has been suggested that much of the effect of sodium acetate is via the increase in the osmolality of rumen fluid but there is considerable uncertainty as to the physiological significance of osmotic effects, especially when animals have free access to water and can prevent excessive increases in tonicity by increasing their water intake. Other constituents of rumen fluid have been implicated in the control of food intake, particularly lactic acid and nitrogenous compounds, especially as these can be found in fermented feeds such as silage. The omasum controls the flow of digesta to the abomasum; it is therefore well placed to control rumen fill and thus intake but there has been little study in this area and this is also true for the abomasum.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of cholecystokinin (CCK-8) on two classes of gastroduodenal vagal afferent fibre

In order to investigate the vagal afferent pathway responsible for the previously reported effects of cholecystokinin (CCK) on gastric emptying and food intake, single afferent fibres were recorded from the cervical vagus of urethane-anaesthetized ferrets. Sixty tension receptor afferents with receptive fields in the corpus, antrum, duodenum, jejunum and ileum all showed a resting level of discharge which was augmented powerfully by distension of the segment containing the ending. Close intraarterial injection of CCK-8 (100-200 pmol) caused relaxation in proximal regions, but enhanced contractile activity in more distal regions. Mechanoreceptor discharge closely followed intraluminal pressure at all times, indicating a sensitivity primarily to tension and no direct sensitivity to CCK. Only duodenal tension receptors were significantly excited by CCK (due to increased contractile activity), whereas those in the stomach showed a net decrease. Thirty-seven mucosal receptors from the corpus, antrum, duodenum and jejunum showed responses to luminal stimuli: predominantly light stroking, acidity and hypertonicity as has been previously described. No responses to glucose or amino-acid infusions could be evoked. However, mucosal fibres showed a strong sensitivity to close-intraarterially injected CCK-8 (3-200 pmol) in 19/26 fibres tested. These responses were unaffected by cholinergic blockade when tested. The data strongly suggest that in the ferret only vagal mucosal receptors are directly sensitive to CCK-8. These fibres are therefore likely candidates for mediating some of the reflex and behavioural effects of CCK when it is released from the gastrointestinal tract and acts directly on vagal sensory endings.

Reflex co‐ordination of corporal and antral contractions in the conscious dog

This study characterizes the role of extrinsic nerves in the co-ordination of corporal and antral contractions in the dog. Fasting motor activity was recorded in conscious dogs with stomachs previously divided into separate corporal and antral pouches. Both corpus and antrum showed synchronized phases of activity and quiescence recognizable as migrating motor complexes (MMCs, duration 81.2 +/- 9.6 min, n = 4). Moreover, individual contractions were temporally linked such that corpus contractions, occurring at 76 +/- 4 s intervals, were each followed by a burst of one to three antral contractions at a frequency of 4-5 min-1. The mean latency between the onset of individual contractions in the corpus and antrum was 10.9 +/- 2.6 s (n = 4). Denervation of the antral pouch in two additional dogs did not affect the MMC cycle (mean durations 106.6 and 82.1 min) and the onset of activity in the corpus and antrum was generally co-ordinated but less precise. However, individual antral contractions were no longer linked to corporal contractions, occurring randomly throughout the corpus contraction cycle. This was associated with a lower contraction frequency in the denervated antral pouches than in the corpus (0.3 +/- 0.1 min-1 compared to 0.6 +/- 0.08 min-1). It is proposed that a vagal reflex, excited by corporal tension receptors, provides phasic excitation facilitating the generation of antral contractions. Such a reflex is likely to reinforce the myogenic mechanisms which occur in the intact stomach and thus plays a role in co-ordinating gastric peristalsis.

Colonic mechanosensory afferents projecting to the mouse superior mesenteric ganglion are in parallel tension receptors located in the circular muscle layer

Colonic mechanosensory afferents projecting to the mouse superior mesenteric ganglion are in parallel tension receptors located in the circular muscle layer

Central neural control of esophageal motility: A review

We review recent studies on the central neural control of esophageal motility, emphasizing the anatomy and chemical coding of esophageal pathways in the spinal cord and medulla. Sympathetic innervation of the proximal esophagus is derived primarily from cervical and upper thoracic paravertebral ganglia, whereas that of the lower esophageal sphincter and proximal stomach is derived from the celiac ganglion. In addition to noradrenaline, many sympathetic fibers in the esophagus contain neuropeptide Y (NPY), and both noradrenaline and NPY appear to decrease blood flow and motility. Preganglionic neurons innervating the cervical and upper thoracic ganglia are located at lower cervical and upper thoracic spinal levels. The preganglionic innervation of the celiac ganglion arises from lower thoracic spinal levels. Both acetylcholine (ACh) and enkephalin (ENK) have been localized in sympathetic preganglionic neurons, and it has been suggested that ENK acts to pre-synaptically inhibit ganglionic transmission. Spinal afferents from the esophagus are few, but have been described in lower cervical and thoracic dorsal root ganglia. A significant percentage contain calcitonin gene-related peptide (CGRP) and substance P (SP). The central distribution of spinal afferents, as well as their subsequent processing within the spinal cord, have not been addressed. Medullary afferents arise from the nodose ganglion and terminate peripherally both in myenteric ganglia, where they have been postulated to act as tension receptors, and, to a lesser extent, in more superficial layers. Centrally, these afferents appear to end in a discrete part of the nucleus of the solitary tract (NTS) termed the central subnucleus. The transmitter specificity of the majority of these afferents remains unknown. The central subnucleus, in turn, sends a dense and topographically discrete projection to esophageal motor neurons in the rostral portion of the nucleus ambiguous (NA). Both somatostatin-(SS) and ENK-related peptides have been localized in this pathway. Finally, motor neurons from the rostral NA innervate striated portions of the esophagus. In addition to ACh, these esophageal motor neurons contain CGRP, galanin (GAL), N-acetylaspartylglutamate (NAAG), and brain natriuretic peptide (BNP). The physiological effect of these peptides on esophageal motility remains unclear. Medullary control of smooth muscle portions of the esophagus have not been thoroughly investigated.

Muscarinic cholinergic receptors on the endothelium of human cerebral arteries.

To characterize the muscarinic cholinergic receptors on the endothelium of human cerebral arteries, isometric tension measurement and receptor autoradiographic studies were performed. Acetylcholine (ACh) induced dose-dependent relaxation of human cerebral arteries precontracted by 10(-5) M serotonin, with an EC50 of 1.9 +/- 0.6 X 10(-6) M (n = 7). The relaxation was abolished by 10(-5) M hemoglobin. Autoradiography, using the muscarinic antagonist [3H]propylbenzilycholine mustard, demonstrated the high density of muscarinic cholinergic receptors on the endothelial cells of human cerebral arteries, especially on the luminal surface of the endothelium. These findings suggest that ACh-induced relaxation mediated by muscarinic cholinergic receptors on the endothelium has a physiological function in human cerebral arteries.

Responses of vagal efferent fibres to stimulation of gastric mechano- and chemoreceptors in the anaesthetized ferret

The involvement of gastric tension receptors and mucosal chemoreceptors in vagal reflexes was assessed directly by recording single efferent fibres from the cervical vagus nerve in the urethane anaesthetised ferret, 32/39 fibres responded to distension of the gastric corpus, mainly with excitation of firing, 24/39 fibres responded to antral distension, with a higher proportion showing a reduction in firing. Fifty-seven percent of fibres showed a convergent input from corpus and antrum. The magnitude of these responses was large (usually > 50%) and of short latency (< 1 s). Responses to chemical perfusions in the antrum were seen in 14/31 efferent fibres, most of which showed excitation. Latency of response was variable (5–120 s) and may have been influenced by diffusion through the antral mucous barrier. These data indicate a complex arrangement of vagal reflexes involved in pre-pyloric regulation of gastric emptying by mechanical and chemical stimuli in the lumen.