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Article: Circadian rhythms in colonic function.

Hibberd, Timothy J / Ramsay, Stewart / Spencer-Merris, Phaedra / Dinning, Phil G / Zagorodnyuk, Vladimir P / Spencer, Nick J

Frontiers in physiology

2023 Volume 14, Page(s) 1239278

Abstract: A rhythmic expression of clock genes occurs within the cells of multiple organs and tissues throughout the body, termed "peripheral clocks." Peripheral clocks are subject to entrainment by a multitude of factors, many of which are directly or indirectly ... ...

Abstract	A rhythmic expression of clock genes occurs within the cells of multiple organs and tissues throughout the body, termed "peripheral clocks." Peripheral clocks are subject to entrainment by a multitude of factors, many of which are directly or indirectly controlled by the light-entrainable clock located in the suprachiasmatic nucleus of the hypothalamus. Peripheral clocks occur in the gastrointestinal tract, notably the epithelia whose functions include regulation of absorption, permeability, and secretion of hormones; and in the myenteric plexus, which is the intrinsic neural network principally responsible for the coordination of muscular activity in the gut. This review focuses on the physiological circadian variation of major colonic functions and their entraining mechanisms, including colonic motility, absorption, hormone secretion, permeability, and pain signalling. Pathophysiological states such as irritable bowel syndrome and ulcerative colitis and their interactions with circadian rhythmicity are also described. Finally, the classic circadian hormone melatonin is discussed, which is expressed in the gut in greater quantities than the pineal gland, and whose exogenous use has been of therapeutic interest in treating colonic pathophysiological states, including those exacerbated by chronic circadian disruption.
Language	English
Publishing date	2023-08-30
Publishing country	Switzerland
Document type	Journal Article ; Review
ZDB-ID	2564217-0
ISSN	1664-042X
ISSN	1664-042X
DOI	10.3389/fphys.2023.1239278
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Article ; Online: Morphological and neurochemical characterisation of anterogradely labelled spinal sensory and autonomic nerve endings in the mouse bladder.

Sharma, Harman / Kyloh, Melinda / Brookes, Simon J H / Costa, Marcello / Spencer, Nick J / Zagorodnyuk, Vladimir P

Autonomic neuroscience : basic & clinical

2020 Volume 227, Page(s) 102697

Abstract: The bladder is innervated by axons of sympathetic and parasympathetic efferent nerves, and by spinal afferent neurons. The objective was to characterise anatomically and immunohistochemically the terminal endings of sensory and autonomic motor nerve ... ...

Abstract	The bladder is innervated by axons of sympathetic and parasympathetic efferent nerves, and by spinal afferent neurons. The objective was to characterise anatomically and immunohistochemically the terminal endings of sensory and autonomic motor nerve endings in wholemount preparations of the mouse bladder. We used both anterograde labelling of pelvic and hypogastric nerves ex vivo and anterograde labelling from lumbosacral dorsal root ganglia (DRG) in vivo in male and female mice. These were combined with immunohistochemistry for major markers of sensory, sympathetic and parasympathetic nerves. Selective labelling of spinal afferent endings following dextran biotin-labelling from DRGs in vivo showed no co-localisation of VAChT or TH in sensory terminals in the detrusor and suburothelial plexus. Biotinamide was applied ex vivo to nerve trunks arising in the pelvic ganglion and running towards the bladder. Among the filled axons, 38% of detrusor fibres and 47% of suburothelial axons were immunoreactive for calcitonin-gene related peptide (CGRP). Vesicular acetylcholine transporter (VAChT) immunoreactivity was present in 26% of both detrusor and suburothelial axons. For tyrosine hydroxylase (TH), the proportions were 15% and 17%, respectively. Three major morphological types of CGRP-immunoreactive nerve endings were distinguished in the bladder wall: simple, branching and complex. VAChT-immunoreactive parasympathetic axons had simple and branching endings; TH immunoreactive axons all had simple morphologies. Our findings revealed that different subtypes of sensory and autonomic nerve endings can be reliably identified by combining anterograde labelling ex vivo with specific immunohistochemical markers, although morphologically some of these types of endings were indistinguishable.
MeSH term(s)	Animals ; Axons/chemistry ; Female ; Immunohistochemistry ; Male ; Mice ; Mice, Inbred C57BL ; Nerve Endings/chemistry ; Neuroanatomical Tract-Tracing Techniques ; Parasympathetic Nervous System/anatomy & histology ; Sympathetic Nervous System/anatomy & histology ; Urinary Bladder/innervation
Language	English
Publishing date	2020-07-01
Publishing country	Netherlands
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2020105-9
ISSN	1872-7484 ; 1566-0702
ISSN (online)	1872-7484
ISSN	1566-0702
DOI	10.1016/j.autneu.2020.102697
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Article ; Online: Disengaging spinal afferent nerve communication with the brain in live mice.

Kyloh, Melinda A / Hibberd, Timothy J / Castro, Joel / Harrington, Andrea M / Travis, Lee / Dodds, Kelsi N / Wiklendt, Lukasz / Brierley, Stuart M / Zagorodnyuk, Vladimir P / Spencer, Nick J

Communications biology

2022 Volume 5, Issue 1, Page(s) 915

Abstract: Our understanding of how abdominal organs (like the gut) communicate with the brain, via sensory nerves, has been limited by a lack of techniques to selectively activate or inhibit populations of spinal primary afferent neurons within dorsal root ganglia ...

Abstract	Our understanding of how abdominal organs (like the gut) communicate with the brain, via sensory nerves, has been limited by a lack of techniques to selectively activate or inhibit populations of spinal primary afferent neurons within dorsal root ganglia (DRG), of live animals. We report a survival surgery technique in mice, where select DRG are surgically removed (unilaterally or bilaterally), without interfering with other sensory or motor nerves. Using this approach, pain responses evoked by rectal distension were abolished by bilateral lumbosacral L5-S1 DRG removal, but not thoracolumbar T13-L1 DRG removal. However, animals lacking T13-L1 or L5-S1 DRG both showed reduced pain sensitivity to distal colonic distension. Removal of DRG led to selective loss of peripheral CGRP-expressing spinal afferent axons innervating visceral organs, arising from discrete spinal segments. This method thus allows spinal segment-specific determination of sensory pathway functions in conscious, free-to-move animals, without genetic modification.
MeSH term(s)	Animals ; Brain ; Colon ; Ganglia, Spinal/metabolism ; Mice ; Pain
Language	English
Publishing date	2022-09-14
Publishing country	England
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ISSN	2399-3642
ISSN (online)	2399-3642
DOI	10.1038/s42003-022-03876-x
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Article ; Online: Localization of the sensory neurons and mechanoreceptors required for stretch-evoked colonic migrating motor complexes in mouse colon.

Zagorodnyuk, Vladimir P / Spencer, Nick J

Frontiers in physiology

2011 Volume 2, Page(s) 98

Abstract: The pacemaker and pattern generator that underlies the cyclical generation of spontaneous colonic migrating motor complexes (CMMCs) has recently been identified to lie within the myenteric plexus and/or muscularis externa. Neither the mucosa, nor the ... ...

Abstract	The pacemaker and pattern generator that underlies the cyclical generation of spontaneous colonic migrating motor complexes (CMMCs) has recently been identified to lie within the myenteric plexus and/or muscularis externa. Neither the mucosa, nor the release of substances from the mucosa were found to be required for the spontaneous generation of CMMCs. However, it is known that stretch applied to the colonic wall can also evoke CMMCs and since stretch of the gut wall is known to stimulate the mucosa, it is not clear whether release of substances from the mucosa and/or submucosal plexus are required for stretch-evoked CMMCs. Therefore, the aim of this study was to determine whether circumferential stretch-evoked CMMCs require the presence of the mucosa and/or submucosal plexus in isolated mouse colon. Spontaneous CMMCs were recorded from full length sheet preparations of colon in vitro. Graded circumferential stretch (at a rate of 100 μm/s) applied to a 15-mm segment of mid-distal colon reliably evoked a CMMC, which propagated to the oral recording site. Sharp dissection to remove the mucosa and submucosal plexus from the entire colon did not prevent spontaneous CMMCs and circumferential stretch-evoked CMMCs were still reliably evoked by circumferential stretch, even at significantly lower thresholds. In contrast, in intact preparations, direct stimulation of the mucosa (without accompanying stretch) proved highly inconsistent and rarely evoked a CMMC. These observations lead to the inescapable conclusion that the sensory neurons activated by colonic stretch to initiate CMMCs lie in the myenteric plexus, while the mechanoreceptors activated by stretch, lie in the myenteric ganglia and/or muscularis externa. Stretch activation of these mechanoreceptors does not require release of any substance(s) from the mucosa, or neural inputs arising from submucosal ganglia.
Language	English
Publishing date	2011-12-21
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	2564217-0
ISSN	1664-042X ; 1664-042X
ISSN (online)	1664-042X
ISSN	1664-042X
DOI	10.3389/fphys.2011.00098
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Article ; Online: Characterization of putative interneurons in the myenteric plexus of human colon.

Humenick, Adam / Chen, Bao Nan / Wattchow, David A / Zagorodnyuk, Vladimir P / Dinning, Phil G / Spencer, Nick J / Costa, Marcello / Brookes, Simon J H

Neurogastroenterology and motility : the official journal of the European Gastrointestinal Motility Society

2020 Volume 33, Issue 1, Page(s) e13964

Abstract: Background: The enteric nervous system contains multiple classes of neurons, distinguishable by morphology, immunohistochemical markers, and projections; however, specific combinations differ between species. Here, types of enteric neurons in human ... ...

Abstract	Background: The enteric nervous system contains multiple classes of neurons, distinguishable by morphology, immunohistochemical markers, and projections; however, specific combinations differ between species. Here, types of enteric neurons in human colon were characterized immunohistochemically, using retrograde tracing combined with multiple labeling immunohistochemistry, focussing on non-motor neurons. Methods: The fluorescent carbocyanine tracer, DiI, was applied to the myenteric plexus in ex vivo preparations, filling neurons projecting within the plexus. Limits of projection lengths of motor neurons were established, allowing them to be excluded from the analysis. Long ascending and descending interneurons were then distinguished by labeling for discriminating immunohistochemical markers: calbindin, calretinin, enkephalin, 5-hydroxytryptamine, nitric oxide synthase, and substance P. These results were combined with a previous published study in which nitric oxide synthase and choline acetyltransferase immunoreactivities were established. Key results: Long ascending neurons (with projections longer than 8 mm, which excludes more than 95% motor neurons) formed four types, in descending order of abundance, defined by immunoreactivity for: (a) ChAT+/ENK+, (b) ChAT+/ENK+/SP+, (c) ChAT+/Calb+, and (d) ChAT+/ENK+/Calb+. Long descending neurons, up to 70 mm long also formed at least four types, distinguished by immunoreactivity for (a) NOS + cells (without ChAT), (b) ChAT+/NOS+, (c) ChAT+/Calret+, and (d) ChAT+/5HT + cells (with or without NOS). Conclusions and inferences: Long interneurons, which do not innervate muscularis externa, are likely to coordinate neural activity over distances of many centimeters along the colon. Characterizing their neurochemical coding provides a basis for understanding their roles, investigating their connectivity, and building a comprehensive account of human colonic enteric neurons.
MeSH term(s)	Aged ; Calbindin 2/metabolism ; Calbindins/metabolism ; Choline O-Acetyltransferase/metabolism ; Colon/innervation ; Enkephalins/metabolism ; Female ; Humans ; Interneurons/metabolism ; Male ; Middle Aged ; Motor Neurons/metabolism ; Myenteric Plexus/cytology ; Myenteric Plexus/metabolism ; Neurons, Afferent/metabolism ; Neurons, Efferent/metabolism ; Nitric Oxide Synthase/metabolism ; Serotonin/metabolism ; Substance P/metabolism
Chemical Substances	Calbindin 2 ; Calbindins ; Enkephalins ; Serotonin (333DO1RDJY) ; Substance P (33507-63-0) ; Nitric Oxide Synthase (EC 1.14.13.39) ; Choline O-Acetyltransferase (EC 2.3.1.6)
Language	English
Publishing date	2020-08-24
Publishing country	England
Document type	Journal Article ; Research Support, N.I.H., Extramural
ZDB-ID	1186328-6
ISSN	1365-2982 ; 1350-1925
ISSN (online)	1365-2982
ISSN	1350-1925
DOI	10.1111/nmo.13964
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Article ; Online: Functional changes in low- and high-threshold afferents in obstruction-induced bladder overactivity.

Zagorodnyuk, Vladimir P / Keightley, Lauren J / Brookes, Simon J H / Spencer, Nick J / Costa, Marcello / Nicholas, Sarah J

American journal of physiology. Renal physiology

2019 Volume 316, Issue 6, Page(s) F1103–F1113

Abstract: Neural mechanisms of lower urinary tract symptoms in obstruction-induced bladder overactivity remain unclear. We made the first single unit recordings from different types of spinal afferents to determine the effects of bladder outlet obstruction in ... ...

Abstract	Neural mechanisms of lower urinary tract symptoms in obstruction-induced bladder overactivity remain unclear. We made the first single unit recordings from different types of spinal afferents to determine the effects of bladder outlet obstruction in guinea pigs. A model of gradual bladder outlet obstruction in male guinea pigs was used to produce overactive bladder. Conscious voiding was assessed in metabolic cages, and micturition was recorded in anesthetized guinea pigs in vivo. Single unit extracellular recordings were made ex vivo from spinal afferent nerves in flat sheet preparations of the bladder. Guinea pigs with partially obstructed bladders showed a significant increase in conscious voiding frequency compared with sham-operated guinea pigs. Also, nonvoiding contractions increased significantly in both frequency and amplitude. Although spontaneous firing of low-threshold bladder afferents was increased, their stretch-induced firing was reduced. The proportion of capsaicin-sensitive low-threshold afferents increased in obstructed bladders. Interestingly, spontaneous and stretch-induced firing were both significantly increased in high-threshold afferents after obstruction. In summary, sensory signaling increased in the obstructed bladder during the filling phase. This is largely mediated by low-threshold stretch-sensitive afferents that are activated by increased local nonvoiding contractions. Increased spontaneous firing by high-threshold afferents also contributes. Our findings revealed a complex effect of bladder outlet obstruction on different types of bladder afferents that needs consideration for potential therapeutic targeting of lower urinary tract symptoms in obstruction-induced bladder overactivity.
MeSH term(s)	Action Potentials ; Afferent Pathways/metabolism ; Afferent Pathways/physiopathology ; Animals ; Disease Models, Animal ; Guinea Pigs ; Male ; Mechanoreceptors/metabolism ; Sensory Thresholds ; Spinal Nerves/metabolism ; Spinal Nerves/physiopathology ; Urinary Bladder/innervation ; Urinary Bladder Neck Obstruction/complications ; Urinary Bladder Neck Obstruction/metabolism ; Urinary Bladder Neck Obstruction/physiopathology ; Urinary Bladder, Overactive/etiology ; Urinary Bladder, Overactive/metabolism ; Urinary Bladder, Overactive/physiopathology ; Urination ; Urodynamics
Language	English
Publishing date	2019-03-25
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	603837-2
ISSN	1522-1466 ; 0363-6127
ISSN (online)	1522-1466
ISSN	0363-6127
DOI	10.1152/ajprenal.00058.2019
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Article ; Online: Extrinsic primary afferent signalling in the gut.

Brookes, Simon J H / Spencer, Nick J / Costa, Marcello / Zagorodnyuk, Vladimir P

Nature reviews. Gastroenterology & hepatology

2013 Volume 10, Issue 5, Page(s) 286–296

Abstract: Visceral sensory neurons activate reflex pathways that control gut function and also give rise to important sensations, such as fullness, bloating, nausea, discomfort, urgency and pain. Sensory neurons are organised into three distinct anatomical ... ...

Abstract	Visceral sensory neurons activate reflex pathways that control gut function and also give rise to important sensations, such as fullness, bloating, nausea, discomfort, urgency and pain. Sensory neurons are organised into three distinct anatomical pathways to the central nervous system (vagal, thoracolumbar and lumbosacral). Although remarkable progress has been made in characterizing the roles of many ion channels, receptors and second messengers in visceral sensory neurons, the basic aim of understanding how many classes there are, and how they differ, has proven difficult to achieve. We suggest that just five structurally distinct types of sensory endings are present in the gut wall that account for essentially all of the primary afferent neurons in the three pathways. Each of these five major structural types of endings seems to show distinctive combinations of physiological responses. These types are: 'intraganglionic laminar' endings in myenteric ganglia; 'mucosal' endings located in the subepithelial layer; 'muscular-mucosal' afferents, with mechanosensitive endings close to the muscularis mucosae; 'intramuscular' endings, with endings within the smooth muscle layers; and 'vascular' afferents, with sensitive endings primarily on blood vessels. 'Silent' afferents might be a subset of inexcitable 'vascular' afferents, which can be switched on by inflammatory mediators. Extrinsic sensory neurons comprise an attractive focus for targeted therapeutic intervention in a range of gastrointestinal disorders.
MeSH term(s)	Gastrointestinal Diseases/physiopathology ; Gastrointestinal Tract/innervation ; Gastrointestinal Tract/physiology ; Humans ; Muscle, Smooth/innervation ; Muscle, Smooth/physiology ; Visceral Afferents/physiology
Language	English
Publishing date	2013-02-26
Publishing country	England
Document type	Journal Article ; Review
ZDB-ID	2493722-8
ISSN	1759-5053 ; 1759-5045
ISSN (online)	1759-5053
ISSN	1759-5045
DOI	10.1038/nrgastro.2013.29
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Article ; Online: Structure-function relationship of sensory endings in the gut and bladder.

Zagorodnyuk, Vladimir P / Brookes, Simon J H / Spencer, Nick J

Autonomic neuroscience : basic & clinical

2010 Volume 153, Issue 1-2, Page(s) 3–11

Abstract: Visceral afferents play a key role in neural circuits underlying the physiological function of visceral organs. They are responsible for the detection and transmission of a variety of visceral sensations (e.g. satiety, urge, discomfort and pain) from the ...

Abstract	Visceral afferents play a key role in neural circuits underlying the physiological function of visceral organs. They are responsible for the detection and transmission of a variety of visceral sensations (e.g. satiety, urge, discomfort and pain) from the viscera to the central nervous system. A comprehensive account of the different functional types of visceral sensory neurons would be invaluable in understanding how sensory dysfunction occurs and how it might be diagnosed and treated. Our aim was to explore the morphology of different nerve endings of visceral afferents within the gastrointestinal tract and urinary bladder and how the morphology of these nerve endings may relate to their functional properties. Morphological studies of mechanosensitive endings of visceral afferents to the gut and bladder correlated with physiological recordings have added a new dimension to our ability to distinguish different functional classes of visceral afferents.
MeSH term(s)	Action Potentials/physiology ; Animals ; Gastrointestinal Tract/anatomy & histology ; Gastrointestinal Tract/innervation ; Gastrointestinal Tract/physiology ; Humans ; Mechanoreceptors/physiology ; Nerve Endings/physiology ; Receptors, Purinergic P2/metabolism ; Receptors, Purinergic P2X2 ; Sensory Receptor Cells/cytology ; Urinary Bladder/anatomy & histology ; Urinary Bladder/innervation ; Urinary Bladder/physiology ; Vesicular Acetylcholine Transport Proteins/metabolism ; Visceral Afferents/physiology
Chemical Substances	P2RX2 protein, human ; Receptors, Purinergic P2 ; Receptors, Purinergic P2X2 ; Vesicular Acetylcholine Transport Proteins
Language	English
Publishing date	2010-02-16
Publishing country	Netherlands
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2020105-9
ISSN	1872-7484 ; 1566-0702
ISSN (online)	1872-7484
ISSN	1566-0702
DOI	10.1016/j.autneu.2009.07.018
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Article ; Online: Translating peripheral bladder afferent mechanosensitivity to neuronal activation within the lumbosacral spinal cord of mice.

Grundy, Luke / Harrington, Andrea M / Caldwell, Ashlee / Castro, Joel / Staikopoulos, Vasiliki / Zagorodnyuk, Vladimir P / Brookes, Simon J H / Spencer, Nick J / Brierley, Stuart M

Pain

2019 Volume 160, Issue 4, Page(s) 793–804

Abstract: Primary afferent neurons transduce distension of the bladder wall into action potentials that are relayed into the spinal cord and brain, where autonomic reflexes necessary for maintaining continence are coordinated with pathways involved in sensation. ... ...

Abstract	Primary afferent neurons transduce distension of the bladder wall into action potentials that are relayed into the spinal cord and brain, where autonomic reflexes necessary for maintaining continence are coordinated with pathways involved in sensation. However, the relationship between spinal circuits involved with physiological and nociceptive signalling from the bladder has only been partially characterised. We used ex vivo bladder afferent recordings to characterise mechanosensitive afferent responses to graded distension (0-60 mm Hg) and retrograde tracing from the bladder wall to identify central axon projections within the dorsal horn of the lumbosacral (LS) spinal cord. Labelling of dorsal horn neurons with phosphorylated-MAP-kinase (pERK), combined with labelling for neurochemical markers (calbindin, calretinin, gamma aminobutyric acid, and parvalbumin) after in vivo bladder distension (20-60 mm Hg), was used to identify spinal cord circuits processing bladder afferent input. Ex vivo bladder distension evoked an increase in primary afferent output, and the recruitment of both low- and high-threshold mechanosensitive afferents. Retrograde tracing revealed bladder afferent projections that localised with pERK-immunoreactive dorsal horn neurons within the superficial laminae (superficial dorsal horn), dorsal gray commissure, and lateral collateral tracts of the LS spinal cord. Populations of pERK-immunoreactive neurons colabelled with calbindin, calretinin, or gamma aminobutyric acid, but not parvalbumin. Noxious bladder distension increased the percentage of pERK-immunoreactive neurons colabelled with calretinin. We identified LS spinal circuits supporting autonomic and nociceptive reflexes responsible for maintaining continence and bladder sensations. Our findings show for the first time that low- and high-threshold bladder afferents relay into similar dorsal horn circuits, with nociceptive signalling recruiting a larger number of neurons.
MeSH term(s)	Afferent Pathways/physiology ; Animals ; Calbindin 2/metabolism ; Cholera Toxin/metabolism ; Female ; Ganglia, Spinal/cytology ; Lumbosacral Region ; MAP Kinase Signaling System/physiology ; Mechanoreceptors/physiology ; Mice ; Mice, Inbred C57BL ; Neurons, Afferent/physiology ; Physical Stimulation/adverse effects ; Spinal Cord/cytology ; Statistics, Nonparametric ; Urinary Bladder/innervation ; gamma-Aminobutyric Acid/metabolism
Chemical Substances	Calbindin 2 ; gamma-Aminobutyric Acid (56-12-2) ; Cholera Toxin (9012-63-9)
Language	English
Publishing date	2019-02-04
Publishing country	United States
Document type	Journal Article
ZDB-ID	193153-2
ISSN	1872-6623 ; 0304-3959
ISSN (online)	1872-6623
ISSN	0304-3959
DOI	10.1097/j.pain.0000000000001453
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Article ; Online: Characterization of projections of longitudinal muscle motor neurons in human colon.

Humenick, Adam / Chen, Bao Nan / Lauder, Chris I W / Wattchow, David A / Zagorodnyuk, Vladimir P / Dinning, Phil G / Spencer, Nick J / Costa, Marcello / Brookes, Simon J H

Neurogastroenterology and motility : the official journal of the European Gastrointestinal Motility Society

2019 Volume 31, Issue 10, Page(s) e13685

Abstract: Background: The enteric nervous system contains inhibitory and excitatory motor neurons which modulate smooth muscle contractility. Cell bodies of longitudinal muscle motor neurons have not been identified in human intestine.: Methods: We used ... ...

Abstract	Background: The enteric nervous system contains inhibitory and excitatory motor neurons which modulate smooth muscle contractility. Cell bodies of longitudinal muscle motor neurons have not been identified in human intestine. Methods: We used retrograde tracing ex vivo with DiI, with multiple labeling immunohistochemistry, to characterize motor neurons innervating tenial and inter-tenial longitudinal muscle of human colon. Key results: The most abundant immunohistochemical markers in the tertiary plexus were vesicular acetylcholine transporter, nitric oxide synthase (NOS), and vasoactive intestinal polypeptide (VIP). Of retrogradely traced motor neurons innervating inter-tenial longitudinal muscle, 95% were located within 6mm oral or anal to the DiI application site. Excitatory motor neuron cell bodies, immunoreactive for choline acetyltransferase (ChAT), were clustered aborally, whereas NOS-immunoreactive cell bodies were distributed either side of the DiI application site. Motor neurons had small cell bodies, averaging 438 + 18µm Conclusions and inferences: Tenial and inter-tenial motor neurons innervating the longitudinal muscle have short projections. Inhibitory motor neurons have less polarized projections than cholinergic excitatory motor neurons. Longitudinal and circular muscle layers are innervated by distinct local populations of excitatory and inhibitory motor neurons. A population of human enteric neurons that contribute significantly to colonic motility has been characterized.
MeSH term(s)	Aged ; Cell Size ; Choline O-Acetyltransferase/metabolism ; Colon/innervation ; Colon/metabolism ; Colon/pathology ; Enteric Nervous System/cytology ; Enteric Nervous System/metabolism ; Female ; Fluorescent Dyes ; Gastrointestinal Motility ; Humans ; Immunohistochemistry ; Male ; Middle Aged ; Motor Neurons/cytology ; Motor Neurons/metabolism ; Muscle, Smooth/innervation ; Muscle, Smooth/metabolism ; Muscle, Smooth/pathology ; Neuroanatomical Tract-Tracing Techniques ; Nitric Oxide Synthase/metabolism
Chemical Substances	Fluorescent Dyes ; Nitric Oxide Synthase (EC 1.14.13.39) ; Choline O-Acetyltransferase (EC 2.3.1.6)
Language	English
Publishing date	2019-07-29
Publishing country	England
Document type	Journal Article ; Research Support, N.I.H., Extramural
ZDB-ID	1186328-6
ISSN	1365-2982 ; 1350-1925
ISSN (online)	1365-2982
ISSN	1350-1925
DOI	10.1111/nmo.13685
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