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PubMed1992

Peptide-containing neurons in different regions of the submucous plexus of human sigmoid colon.

Specimens of the sigmoid colon were obtained from male and female patients (n = 11) with carcinoma of the colon or rectum and studied immunohistochemically for vasoactive intestinal polypeptide-, somatostatin-, substance P-, neuropeptide Y-, calcitonin gene-related peptide-, met- and leu-enkephalin-, 5-hydroxytryptamine-, and dopamine beta-hydroxylase-containing nerves. In the subdivisions of the submucous plexus (namely, Schabadasch's, Meissner's, and the intermediate plexuses), substance P- and vasoactive intestinal polypeptide-immunoreactive nerve fibers were the most numerous, and equal densities of these nerves were found in all three layers. In contrast, few neuropeptide Y-, met-enkephalin-, leu-enkephalin-, calcitonin gene-related peptide-, somatostatin-, 5-hydroxytryptamine-, and dopamine beta-hydroxylase-immunoreactive nerves were found in these regions. The nerve cell bodies of the submucous plexus contained vasoactive intestinal polypeptide, substance P, leu-enkephalin, somatostatin, and 5-hydroxytryptamine but not neuropeptide Y, met-enkephalin, calcitonin gene-related peptide, and dopamine beta-hydroxylase. Vasoactive intestinal polypeptide-containing nerve cell bodies were found in all three subdivisions. Substance P-, leu-enkephalin-, and somatostatin-immunoreactive nerve cell bodies were found in Schabadasch's plexus and the intermediate region of the submucous plexus, but they were absent from Meissner's plexus; 5-hydroxytryptamine-containing nerve cell bodies were only observed in Schabadasch's plexus. The possible function of the neuropeptide-, dopamine beta-hydroxylase-, and 5-hydroxytryptamine-containing neurons in the different layers of the submucous plexus is discussed.

Adult↗

PubMed1988

Electrophysiological characterization of functionally distinct 5-hydroxytryptamine receptors on guinea-pig submucous plexus.

Intracellular recordings were made from neurons of the guinea-pig submucous plexus and the actions of 5-hydroxytryptamine on the postsynaptic membrane and on evoked synaptic potentials were examined. 5-Hydroxytryptamine produced two types of direct postsynaptic responses: (1) A depolarization associated with a fall in input resistance was observed in all cells. Voltage-clamp and ion substitutions showed that this depolarization resulted primarily from an inward sodium current. This response could be as brief as 30 ms; it showed desensitization and was selectively abolished by 0.2-2 microM ICS 205-930. (2) A depolarization (or inward current) associated with a decreased conductance was observed in about 50% of neurons, usually after the first response was blocked by ICS 205-930. This response was due to a decreased potassium conductance; the minimum time course of this response was 8-10 s. It did not show desensitization and was not sensitive to blockade by currently available antagonists of 5-hydroxytryptamine, nicotinic and/or muscarinic receptors. Higher concentrations of 5-hydroxytryptamine were required to produce the sodium conductance increase than the potassium conductance decrease; 2-methyl-5-hydroxytryptamine was equally effective in producing these responses. 5-Hydroxytryptamine also caused a barrage of "spontaneous" nicotinic excitatory post-synaptic potentials which were sensitive to tetrodotoxin. This response desensitized, was blocked by ICS 205-930 and is presumed to reflect excitation of other cholinergic cell bodies in the plexus by the sodium conductance increase mechanism described. The evoked nicotinic excitatory postsynaptic potential and the adrenergic inhibitory postsynaptic potential were decreased by 5-hydroxytryptamine; a portion of this inhibition showed desensitization and was blocked by ICS 205-930 as well as by the muscarinic receptor antagonists, atropine and pirenzepine. The ICS 205-930-insensitive portion of this inhibition could not be attributed to activation of 5-hydroxytryptamine-1 or 5-hydroxytryptamine-2 receptors. Thus, the following conclusions are drawn: 5-hydroxytryptamine excites submucous plexus neurons by activating two distinct 5-hydroxytryptamine receptors. Activation of the 5-hydroxytryptamine-3 receptor (sensitive to ICS 205-930) produces a depolarization mediated by an increased sodium conductance. The same effect occurring in other cholinergic cell bodies initiates action potentials which are responsible for the 5-hydroxytryptamine-induced release of acetylcholine.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed2002

Colocalization of NO and VIP in neurons of the submucous plexus in the rat intestine.

Since very few previous studies have carried out the quantitative analysis for the colocalization of nitric oxide (NO) and vasoactive intestinal peptide (VIP) in the submucous neurons in the rat digestive tract, we applied in vivo treatment of colchicine to enhance the immunoreactivity and examined the colocalization of NO synthase (nNOS) and VIP in neurons of the submucous plexus throughout the rat digestive tract. The density of nNOS-containing neurons in the submucous plexus in the stomach corpus (103+/-25 cells/cm(2), n=3) and that in the antrum (157+/-9 cells/cm(2), n=3) were significantly lower than those in small and large intestine. However no difference was detected in the cell density among duodenum (1967+/-188 cells/cm(2), n=3), jejunum (2640+/-140 cells/cm(2), n=3), ileum (2070+/-42 cells/cm(2), n=3), proximal colon (2243+/-138 cells/cm(2), n=3) and distal colon (2633+/-376 cells/cm(2), n=3). The proportion of nNOS-immunoreactive (IR), nNOS/VIP-IR and VIP-IR neurons to the total number of submucous neurons was examined. nNOS/VIP-IR neurons comprised 45-55% of total number of submucous neurons from the duodenum to the proximal colon, however those comprised 66.4+/-5.1% in the distal colon. The results showed that the dense distribution of nNOS-containing neurons was found in the submucous plexus throughout the small and large intestine, and large population of submucous neurons co-stored nNOS and VIP.

Animals↗

PubMed1999

Functional group I metabotropic glutamate receptors in submucous plexus of guinea-pig ileum.

Intracellular recording methods and immunostaining revealed the existence of functional group I metabotropic glutamate receptors (mGluRs) in submucous plexus neurons of guinea-pig ileum. Selective group I, but not groups II or III metabotropic glutamate receptor agonists induced concentration-dependent, slowly-activating depolarizing responses. Group I metabotropic glutamate receptor antagonism observed with (S)-4-carboxyphenylglycine (S-4-CPG) (100 - 600 microM) was competitive as determined by Schild analysis (pA(2)=3.81+/-0.02). Neither the group II and III metabotropic nor ionotropic glutamate receptor antagonists altered responses evoked by group I receptor agonists. Immunoreactivities for metabotropic glutamate 1alpha and 5 receptors were found to locate exclusively in neurons in the submucous plexus of guinea-pig ileum with the highest density around the cell bodies. The results suggest that group I metabotropic glutamate receptors are functionally expressed in the submucous plexus of guinea-pig small intestine.

Animals↗

PubMed1989

Muscarinic excitation and inhibition of neurons in the submucous plexus of the guinea-pig caecum.

Intracellular recordings were made from neurons in the submucous plexus of the guinea-pig caecum. Muscarinic agonists (acetylcholine, bethanechol and muscarine) depolarized about 70%, and hyperpolarized about 30% of the submucous plexus neurons. Low concentrations of pirenzepine reversibly antagonized both responses. The measured dissociation constants (KD) of 10-30 nM for the depolarizations and 1-3 nM for the hyperpolarizations suggest that each response was mediated by muscarinic M1 cholinoceptors. The muscarinic depolarization and hyperpolarization were associated with a decreased and an increased conductance, respectively, and the reversal potential for the muscarinic responses varied as the potassium concentration varied, always being around the potassium equilibrium potential. In cells depolarized by muscarinic agonists these agents appeared to decrease a potassium conductance that could also be inactivated by substance P. In approximately 30% of the submucous neurons, the slow inhibitory postsynaptic potential, elicited in response to single or repetitive focal stimuli (1-10 pulses at 20-40 Hz), appeared to consist of a large component which was sensitive to the blocking action of idazoxan (100-300 nM) and a small component which was idazoxan-insensitive. The latter (muscarinic slow inhibitory postsynaptic potential) was completely abolished by pirenzepine. The concentrations of pirenzepine which caused a 50% depression ranged from 5 to 20 nM. The muscarinic slow inhibitory postsynaptic potential was increased in amplitude and duration by physostigmine (100-300 nM). The muscarinic slow inhibitory postsynaptic potential was accompanied by a decrease in membrane input resistance, and was reversed in polarity near the potassium equilibrium potential. When muscarine induced a hyperpolarization and/or focal stimulation elicited a muscarinic slow inhibitory postsynaptic potential in the presence of idazoxan (100-300 nM), the intracellular injection of guanosine 5'-O-(3-thiotriphosphate) produced a progressive membrane hyperpolarization during which the muscarinic hyperpolarizing responses were attenuated. It is concluded that the muscarine-induced reduction in potassium conductance is mediated through a muscarinic M1 receptor which has a relatively low affinity for pirenzepine. The muscarine-induced increase in potassium conductance is probably produced by the association of a guanine nucleotide-binding regulatory protein with another muscarinic M1 receptor that has a relatively high affinity for pirenzepine.

Acetylcholine↗

PubMed1986

Motoneurones of the submucous plexus regulate electrical activity of the circular muscle of canine proximal colon.

The hypothesis that the circular muscle of the canine proximal colon receives motor input from neurones in the submucous plexus was tested. Circular muscle cells were impaled with micro-electrodes and submucous plexus neurones were stimulated by electrical field stimulation and microejection of acetylcholine (ACh). In the presence of atropine to block the direct muscarinic effects, microejection of ACh onto the submucosa where intact submucous ganglia were suspended evoked: (i) an inhibitory junction potential (i.j.p.) that reduced the amplitude, duration and rate of rise of the subsequent slow wave; (ii) a slow wave of increased duration following the initial inhibitory response. These responses were enhanced by increasing the volume of ACh administered. Responses to ACh were blocked by hexamethonium, 10(-4) M; d-tubocurarine, 10(-4) M; or tetrodotoxin (TTX), 10(-6) M, suggesting they were neural in origin. Both inhibitory and excitatory responses were the result of non-cholinergic and non-adrenergic nerves. The transmitters mediating these effects are unknown. Removal of the longitudinal muscle, myenteric plexus, and the serosal portion of the circular muscle had no apparent effect on the responses to application of ACh to submucosal ganglia. In these preparations the responses to field stimulation were identical to those produced by ACh. The submucous plexus also provides cholinergic input to the circular muscle. When ACh was discretely applied to the submucosa cholinergic responses were elicited at the muscle cell which were significantly reduced by hexamethonium or TTX. These findings suggest that the cholinergic responses were the result of ACh release by neurones at the effector and not by overflow of the exogenous ACh. Cholinergic responses were also elicited in preparations in which the myenteric plexus had been removed. Slow waves in circular muscle of the proximal colon yield excitation-contraction coupling in the absence of Ca2+ action potentials. Therefore the influence of submucous neurones on electrical slow waves has direct consequences on motor activity. Reduction in the amplitude and duration of slow wave by i.j.p.s. results in reduction in the amplitude and duration of phasic contractions. Excitatory inputs enhance contractions. The data support a new concept: motoneurones emanating from submucous ganglia innervate the circular muscle and provide inhibitory and excitatory inputs to regulate slow wave activity.(ABSTRACT TRUNCATED AT 400 WORDS)

Acetylcholine↗

PubMed1984

Two types of neurones lacking synaptic input in the submucous plexus of guinea-pig small intestine.

Intracellular recordings were made from neurones in the submucous plexus of guinea-pig small intestine. Approximately 20% of all neurones studied appeared to lack synaptic input; two types of neurones without synaptic input were identified. The action potential recorded from one of these types of ganglion cells was followed by a prolonged after-hyperpolarization. The after-hyperpolarization was abolished by Co2+ or Mn2+. A Ca2+ component of the evoked action potential was demonstrated after the Na+ component had been abolished by tetrodotoxin (TTX). The other type of neurone was characterized by a high input resistance (220-320 M omega) and spontaneous action potentials which were unaffected by hexamethonium, curare, atropine or TTX. The ionic basis of the action potentials evoked from spontaneously active cells differed from other enteric neurones. These action potentials exhibited a TTX-insensitive Na+ component as well as a Ca2+ component. It is suggested the spontaneously active neurones may be responsible for the ongoing synaptic activity recorded from many submucous plexus neurones in the absence of stimulation.

Action Potentials↗

PubMed1985

Nerve cell density in submucous plexus throughout the gut of cat and opossum.

Quantitative differences in submucous plexus density were sought in cat and opossum gut by examining full-thickness whole mounts of the submucosa stained with silver, and counting ganglia per square centimeter and nerve cell bodies per ganglion in order to compute density of innervation (nerve cell bodies per square centimeter). In the cat, the nerve cell bodies per square centimeter in the 12 named regions were as follows: proximal esophagus, 0; mid-esophagus, 0; distal esophagus, 0; fundus, 84; gastric antrum, 18; duodenum, 5831; jejunum, 4632; ileum, 3191; proximal colon, 1275; mid-colon, 689; distal colon, 359; rectum, 144. In the opossum, values were as follows: proximal esophagus, 37; mid-esophagus, 52; distal esophagus, 84; duodenum, 1812; jejunum, 2234; ileum, 1488; proximal colon, 206; mid-colon, 197; distal colon, 121; rectum, 61. Adequate specimens could not be obtained from opossum stomach. Differences were due more to variations in distribution density of ganglia than in ganglionic size. The relatively dense submucous plexus of the intestine probably is related to the capacity of the intestinal mucosa for peptide secretion as well as to its absorptive function.

Animals↗

PubMed2004

Quantitative morphometric analysis of the submucous plexus in age-related control groups.

An increased number and density of the so-called "giant ganglia" (seven or greater ganglion cells per ganglion) serve as histopathological criteria for a bowel motility disorder called intestinal neuronal dysplasia of the submucous plexus (IND B). However, because these morphological criteria have been defined based upon observations in constipated patients, the diagnostic value of previous studies is open to controversy. Moreover, no age-related reference data from unaffected controls are available. This study reports on data from unaffected controls on the variability of size and distribution of ganglia in the submucous plexus during development. Therefore, for the first time, the normal status has been defined. Four age groups have been defined: (a) premature births, gestational age less than 35 weeks; (b) 1-365 days; (c) 1-14 years and (d) 15 years to greater than 70 years). All of these groups revealed giant ganglia in the submucous plexus. With advancing age, there was a decrease in the number of giant ganglia (from 32.7% in group a to 11.2% in group d) accompanied by an inverse increase in the mean distance between all ganglia (from 0.52 mm in group a to 1.17 mm in group d). The data presented permit the conclusion that the criteria mentioned above are not apt to define IND B as an entity, since they do not allow a sufficient demarcation from the age-correlated normal values presented here.

Adolescent↗

PubMed1986

Arterial submucous plexus of duodenum (1st 2.5 cm): vascular etiology in duodenal ulcers.

The arterial submucous plexus in 5 human duodena (1st 2.5 cm of duodenum) were studied by injecting India-ink. The submucous plexus was richly vascular, which excluded the possibility of the end arteries to be the causative factor in production of ulcers in this region. Perhaps, the ulcers in this region may be due to obstruction of blood flow caused by traction of an over filled stomach.

Arteries↗

PubMed1975

Observations on the submucous plexus and mucosal arteries of the dog's stomach and first part of the duodenum.

Arteriolar patterns of the submucous plexus were studied in all areas of the dog's stomach and in the first inch of the duodenum. There appeared to be no poverty of plexus, although in some cases the vessels were somewhat smaller in the pyloric part of the lesser curvature than elsewhere. Mucosal arteries arose from the plexus, and none appeared to have an extramural origin. In man, on the other hand, there is a poverty of the submucous plexus in the 'ulcer region', i.e. in the incisural region of the lesser curvature and in the first inch of the duodenum, associated in some cases with mucosal end arteries of extramural origin. The absence of these features in the dog, which does not suffer from spontaneous chronic ulceration, lends further support to the view that they play a role in the aetiology of the disease in man.

Angiography↗

PubMed2001

Outer submucous plexus: an intrinsic nerve network involved in both secretory and motility processes in the intestine of large mammals and humans.

The architecture of the enteric nerve networks in the gastrointestinal tract appears to be more complex in large mammals, including humans, than in small laboratory animals. At least two distinct ganglionic nerve plexuses could be identified in the submucous layer in the digestive tract of large mammals. While functionally and morphologically similar neuron populations are found in the intestinal wall of both small and large mammals, significant differences in their topographical organization and neurochemical features may be present. This short review clearly illustrates that the close and exclusive association, which has been assumed so far between the efferent pathways of the submucous plexus and regulation of intestinal secretion/absorption on the one hand and between the myenteric plexus and regulation of intestinal motility on the other hand, cannot be interpreted that strictly. An attempt has been made to give a briefoverview of the current status of the identification of distinct functional enteric neuronal classes in the gastrointestinal tract of large mammals using the pig and human intestine as references, and to compare these data with the more extensive information gathered from the guinea-pig intestine.

Animals↗

PubMed1990

Changes in adrenergic and peptidergic nerves in the submucous plexus of streptozocin-diabetic rat ileum.

The effect of streptozocin diabetes on the distribution of adrenergic and peptidergic nerves in the submucous plexus of rat ileum was investigated and compared with the changes in the myenteric plexus of the same region of ileum. There was an increase in the intensity of immunoreactivity in vasoactive intestinal polypeptide- and neuropeptide Y-like immunoreactive nerve fibers and neurons and a decrease in calcitonin gene-related peptide-like immunoreactivity but no change in substance P- and dopamine beta-hydroxylase-like immunoreactivity in the nerve fibers and neurons of the submucous plexus of both 8- and 16-wk streptozocin-diabetic rat ileum. However, in the myenteric plexus of the diabetic rat ileum, there was enlargement of varicosities and an increase followed by a slight decrease in the intensity of immunoreactivity of vasoactive intestinal polypeptide- and dopamine beta-hydroxylase-like immunoreactive nerve fibers and neurons, increased substance P-like immunoreactivity in diabetes at 16 wk, and an initial decrease (at 8 wk) followed by a recovery of calcitonin gene-related peptide-like immunoreactivity at 16 wk, but no change in neuropeptide Y-like immunoreactivity. The markedly different changes in peptidergic and adrenergic nerves between the two enteric plexuses show that diabetic neuropathy induced by streptozocin is not selective and involves factors other than neurotransmitter types.

Adrenergic Fibers↗

PubMed1999

Morphological relationship between serotonergic neurons and nitrergic neurons for electrolytes secretion in the submucous plexus of the guinea pig distal colon.

In the submucous plexus, double immunocytochemistry revealed that nitric oxide synthase (NOS)-immunoreactivity was found in both numerous nerve fibers and some nerve cell bodies, while 5-hydroxytryptamine (5-HT)-immunoreactivity was limited to many nerve fibers, but not any nerve cell bodies. About 30% of the total NOS positive neurons (978) had close or some contact with 5-HT positive nerve fiber, suggesting that NO may participate in the 5-HT-evoked chloride secretion.

Animals↗

PubMed1984

Slow excitatory synaptic potentials recorded from neurones of guinea-pig submucous plexus.

Intracellular recordings made from neurones of guinea-pig submucous plexus revealed three types of synaptic input: cholinergic excitatory synaptic potentials (fast e.p.s.p.s) of 50-80 ms duration, inhibitory synaptic potentials (i.p.s.p.s) of 1 s duration, and non-cholinergic, non-adrenergic slow e.p.s.p.s which lasted for 15-20 s. A single stimulus was sufficient to elicit the slow e.p.s.p. in all neurones in which this synaptic input was present. Slow e.p.s.p.s were recorded in those neurones which also displayed i.p.s.p.s. Both the i.p.s.p. and the slow e.p.s.p. appeared in an all-or-none fashion and were not affected by alterations in the stimulus strength. The inhibitory as well as the slow excitatory synaptic potentials reversed close to the K+ equilibrium potential, indicating that the i.p.s.p. is due to an activation of K+ conductance while the slow e.p.s.p. is due to its inactivation. Evidence is presented which suggests the slow e.p.s.p. may be generated at a synapse located some distance from the soma, presumably at a dendritic location. Only those cells which showed slow e.p.s.p.s responded to substance P with a depolarization which mimicked the slow e.p.s.p.

Action Potentials↗

PubMed1985

Slow postsynaptic potentials in neurones of submucous plexus of guinea-pig caecum and their mimicry by noradrenaline and various peptides.

Intracellular recordings of membrane potential and membrane currents were made from neurones in the submucous plexus of the guinea-pig caecum in vitro. Fast and slow excitatory postsynaptic potentials and slow inhibitory postsynaptic potentials were recorded from the majority of neurones following focal stimulation of presynaptic fibres in the plexus. The slow inhibitory postsynaptic potential was associated with an increase in membrane conductance and reversed its polarity at -90 mV; it was reversibly blocked by yohimbine. The slow excitatory postsynaptic potential and its underlying current was associated with a decrease in membrane conductance. Two kinds of voltage-dependence both of the slow excitatory postsynaptic potential and current were observed; in 80% of cells, the excitatory postsynaptic potential and current became smaller with membrane hyperpolarization and reversed polarity at -90 mV (reversing type) but in 20% of cells both the excitatory postsynaptic potential and current simply disappeared when the membrane potential reached -70 mV (non-reversing type). The effects of acetylcholine, adenosine 5'-triphosphate, bombesin, 5-hydroxytryptamine, neurotensin, noradrenaline, substance P and vasoactive intestinal polypeptide were examined. The only substance which mimicked the slow inhibitory postsynaptic potential was noradrenaline; brief applications of noradrenaline caused hyperpolarizations which had the same time-course, reversal potential and sensitivity to yohimbine as the slow inhibitory postsynaptic potential. The non-reversing type of slow excitatory postsynaptic potential was mimicked only by adenosine 5'-triphosphate. The reversing type of slow excitatory postsynaptic potential was mimicked by bombesin, neurotensin, substance P and vasoactive intestinal polypeptide. 5-Hydroxytryptamine and vasoactive intestinal polypeptide (in some neurones) caused a depolarization with an increase in membrane conductance. All three synaptic potentials were reversibly depressed by superfusion of noradrenaline but noradrenaline did not affect the potential changes evoked by brief application of exogenous acetylcholine or substance P. It is concluded that, in guinea-pig submucous plexus neurones, the slow inhibitory postsynaptic potential is mediated by noradrenaline and results from a potassium conductance increase.(ABSTRACT TRUNCATED AT 400 WORDS)

Adenosine Triphosphate↗

PubMed1998

Morphometric aspects of the submucous plexus in whole-mount preparations of normal human distal colon.

BACKGROUND/PURPOSE: The diagnosis of intestinal neuronal dysplasia (IND) has traditionally been based on the finding of hyperplasia of the submucous plexus and increased acetylcholinesterase activity in parasympathetic nerve fibers in the lamina propria. However, recently it has been suggested that proposed diagnostic criteria relating to nerve cell density may overlap with age-related changes and that the finding of giant ganglia (ganglia containing more than seven ganglion cells) is the most relevant diagnostic parameter of IND. Ganglion cell counting is usually performed on conventional histological sections, whereas the topology of whole ganglia has been poorly studied. The aim of this study was to define the number of ganglion cells per ganglion and the ganglion cell density (the number of ganglion cells per surface area) in submucous whole-mount preparations of normal human colon. METHODS: Specimens from distal colon were obtained during postmortem examination from 14 patients who died of nongastrointestinal disease. The submucous layer was prepared as a whole mount and stained for NADPH diaphorase (a nitrergic neurotransmitter marker) and cuprolinic blue (a general neuronal marker). Ganglion cell density was estimated by counting at least 10 mm2. The number of ganglion cells per ganglion was counted in at least 20 ganglia per case. RESULTS: Ganglion cell density (NADPH diaphorase) fell markedly during the first 5 to 6 years of life (r = -0.60, P < .05). Most ganglion cells formed ganglia of 3 to 64 cuprolinic blue staining cells. The mean number of ganglion cells per ganglion did not vary with age. CONCLUSIONS: The density of NADPH diaphorase-positive ganglion cells in the submucous plexus of human distal colon decreases markedly with age. However, the number of ganglion cells per ganglion remains constant. These findings indicate that the age of the patient has crucial importance for the histolopathologic evaluation of enteric nervous system disorders.

Adolescent↗

PubMed1979

Ultrastructural observations on the submucous plexus of the large intestine of the rainbow trout (Salmo gairdneri, Rich.).

Ultrastructural studies on the large intestine of the Rainbow trout revealed the hitherto unreported presence of neurons within the submucous plexus. The neurons were closely associated with axons which expanded intermittently along their length to form varicosities. According to the type and proportion of vesicles within the varicosities, three groups were distinguished. Group I varicosities contained a predominant population of small electronlucent vesicles measuring 30 nm in diameter. Group II varicosities contained a few medium-sized granular vesicles measuring 60 nm in diameter mixed with large numbers of small electronlucent vesicles. Most of the Group III varicosities contained large numbers of large granular vesicles, 90 nm in diameter. Varicosities of all groups formed axosomatic synapses. The significance of the presence of neurons in the submucous plexus of the trout large intestine is discussed. Group I and II varicosity profiles were interpreted as representing cholinergic and adrenergic innervation respectively. The identification of the transmitter substance in Group III varicosities, however, has still to be resolved.

Animals↗