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Neuroepithelial circuit formed by innervation of sensory enteroendocrine cells
Open accessAbstract
Satiety and other core physiological functions are modulated by sensory signals arising from the surface of the gut. Luminal nutrients and bacteria stimulate epithelial biosensors called enteroendocrine cells. Despite being electrically excitable, enteroendocrine cells are generally thought to communicate indirectly with nerves through hormone secretion and not through direct cell-nerve contact. However, we recently uncovered in intestinal enteroendocrine cells a cytoplasmic process that we named neuropod. Here, we determined that neuropods provide a direct connection between enteroendocrine cells and neurons innervating the small intestine and colon. Using cell-specific transgenic mice to study neural circuits, we found that enteroendocrine cells have the necessary elements for neurotransmission, including expression of genes that encode pre-, post-, and transsynaptic proteins. This neuroepithelial circuit was reconstituted in vitro by coculturing single enteroendocrine cells with sensory neurons. We used a monosynaptic rabies virus to define the circuit's functional connectivity in vivo and determined that delivery of this neurotropic virus into the colon lumen resulted in the infection of mucosal nerves through enteroendocrine cells. This neuroepithelial circuit can serve as both a sensory conduit for food and gut microbes to interact with the nervous system and a portal for viruses to enter the enteric and central nervous systems.
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“A basolateral cytoplasmic process of the L cells -the neuropod -was shown to establish a functional synaptic contact with the enteric glia and vagal afferents.”
From The Microbiota-Gut-Brain Axis (2019) · cites this paper as Bohorquez et al., 2015
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“Enteroendocrine and enterochromaffin cells (ECCs) form close synaptic connections with certain vagal afferent fibers through cell extensions called neuropods [25, 26].”
From Role of diet and its effects on the gut microbiome in the pathophysiology of mental disorders (2022) · cites this paper as [26]
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Liu, W. W., & Bohórquez, D. V. (2022) · Nature reviews. Neuroscience · doi:10.1038/s41583-022-00613-5 · Cited by 65
Self-citationMicrobiome to Brain: Unravelling the Multidirectional Axes of Communication
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Gastrointestinal Immunity and Alpha-Synuclein
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Prox1-positive cells monitor and sustain the murine intestinal epithelial cholinergic niche
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Enteroendocrine Dynamics – New Tools Reveal Hormonal Plasticity in the Gut
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Gut mucosal cells transfer α-synuclein to the vagus nerve
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“Indeed, enteroendocrine cells have been reported to contain neuropods that are directly linked to neuronal cells and are able to transmit signals to the CNS (Bohórquez et al., 2015).”
BackgroundEnteroendocrine cell lineages that differentially control feeding and gut motility
Hayashi, M., Kaye, J. A., Douglas, E., Joshi, N. R., Gribble, F. M., Reimann, F. M., & Liberles, S. D. (2023) · eLife · doi:10.7554/elife.78512 · Cited by 63
“…cells are epithelial cells with neuron-like features, as they are electrically excitable, release vesicles upon elevation of intracellular calcium, and form synaptic connections with second-order neurons through specialized extrusions called neuropods (Bohorquez et al., 2015; Reimann et al., 2012).”
BackgroundRen, W., Chen, J., Wang, W., Li, Q., Yin, X., Zhuang, G., Zhou, H., & Zeng, W. (2024) · Neuron · doi:10.1016/j.neuron.2023.12.012 · Cited by 62
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“By using time-lapse photomicroscopy it was observed that fluorescent EECs interacted with sensory neurons labeled with the lipophilic dye, DiI.71 With an EEC and sensory nerve lying in close proximity, the nerve extended a small neurite toward the EEC.”
BackgroundMethodResultInfluentialSelf-citationOptogenetic and chemogenetic techniques for neurogastroenterology
Boesmans, W., Hao, M. M., & Berghe, P. V. (2017) · Nature Reviews Gastroenterology & Hepatology · doi:10.1038/nrgastro.2017.151 · Cited by 55
Enteroendocrine cells couple nutrient sensing to nutrient absorption by regulating ion transport
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Regulation of the Autonomic Nervous System on Intestine
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Gut Microbiome and Modulation of CNS Function
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“Moreover, vagal afferent terminals can form synapse-like connections with extensions of ECCs, so-called neuropods, providing the peripheral serotonin system a direct neural link to the brain (19).”
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A gut sense for a microbial pattern regulates feeding
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Sodium channel NaV1.3 is important for enterochromaffin cell excitability and serotonin release
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How dietary amino acids and high protein diets influence insulin secretion
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“Nutrient sensors are molecular and cellular machines responsible for detecting amino acids in addition to other nutrient components such as glucose, fatty acids, minerals, and anions (Bohorquez et al., 2015; Newsholme et al., 2010).”
BackgroundInfluentialEarly life interaction between the microbiota and the enteric nervous system
Foong, J. P. P., Hung, L. Y., Poon, S., Savidge, T., & Bornstein, J. C. (2020) · American Journal of Physiology-Gastrointestinal and Liver Physiology · doi:10.1152/ajpgi.00288.2020 · Cited by 50
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“Intriguingly, recent studies have uncovered that sensory neuropod cells in the gut [66] can synaptically signal with the juxtaposed vagal afferents using, among other possible mediators [67], the fast-acting neurotransmitter glutamate [10].”
BackgroundGut-brain axis in the pathogenesis of sepsis-associated encephalopathy
Wang, X., Wen, X., Yuan, S., & Zhang, J. (2024) · Neurobiology of Disease · doi:10.1016/j.nbd.2024.106499 · Cited by 47
“For example, enterochromaffin cells (ECCs) can release serotonin (5-HT) and peptide hormones that act on the vagal sensory neurons coupled with them, but also form synaptic connections with the vagus nerve (Bellono et al., 2017; Boh ´ orquez et al., 2015).”
BackgroundThe gut–brain axis and pain signalling mechanisms in the gastrointestinal tract
Meerschaert, K. A., & Chiu, I. M. (2024) · Nature Reviews Gastroenterology & Hepatology · doi:10.1038/s41575-024-01017-9 · Cited by 57
“Utilizing monosynaptic rabies virus tracing, it was shown that colonic enteroendocrine cells synapse with sensory neurons in mice 142 .”
Galanin inhibits GLP‐1 and GIP secretion via the GAL1 receptor in enteroendocrine L and K cells
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“Recent work (Bohorquez et al., 2014; Bohorquez et al., 2015) now suggests that enteroendocrine cells possess a prominent cytoplasmic process referred to as a “neuropod,” which is capable of direct contact with nerve fibres.”
BackgroundThe Brain–Gut–Bone Axis in Neurodegenerative Diseases: Insights, Challenges, and Future Prospects
Li, R., Miao, Z., Liu, Y., Chen, X., Wang, H., Su, J., & Chen, J. (2024) · Advanced Science · doi:10.1002/advs.202307971 · Cited by 54
Possible role of peptide YY (PYY) in the pathophysiology of irritable bowel syndrome (IBS)
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Enteroendocrine Regulation of Nutrient Absorption
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Layers of signals that regulate appetite
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“For this reason, there is the potential for considerable additional insight by using methods that record neural activity in real-time and during behavior.”
BackgroundRewiring the Brain Through the Gut: Insights into Microbiota–Nervous System Interactions
Săvulescu-Fiedler, I., Benea, S. N., Căruntu, C., Nancoff, A.-S., Homentcovschi, C., & Bucurica, S. (2025) · Current Issues in Molecular Biology · doi:10.3390/cimb47070489 · Cited by 51
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Neural control of gut homeostasis
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“The same study identified synaptically connected enteroendocrine cells (named neuropod cells) that synapse on afferent neurons and signal via the excitatory neurotransmitter glutamate to the brain stem, thus, rapidly informing the brain of gut luminal stimuli (17, 18, 87).”
BackgroundMelnik, B. C. (2021) · International Journal of Molecular Sciences · doi:10.3390/ijms22031059 · Cited by 43
Balasuriya, G. K., Hill‐Yardin, E. L., Gershon, M. D., & Bornstein, J. C. (2016) · The Journal of Physiology · doi:10.1113/jp272071 · Cited by 41
“2011) that were similar to the ‘neuropods’ of enteroendocrine cells (Bohórquez et al. 2015).”
ResultEnteroendocrine cells: the gatekeepers of microbiome-gut-brain communication
Grundeken, E., & Aidy, S. E. (2025) · npj Biofilms and Microbiomes · doi:10.1038/s41522-025-00810-x · Cited by 48
“EECs are capable of forming specialized structures known as neuropods, whichsynapsewiththenervesoftheENS,therebycreatinganeuroepithelial circuit 7,8 .”
Dodds, K. N., Travis, L., Kyloh, M. A., Jones, L. A., Keating, D. J., & Spencer, N. J. (2022) · American Journal of Physiology-Gastrointestinal and Liver Physiology · doi:10.1152/ajpgi.00019.2022 · Cited by 42
Enteroendocrine cells regulate intestinal homeostasis and epithelial function
Nwako, J. G., & McCauley, H. A. (2024) · Molecular and Cellular Endocrinology · doi:10.1016/j.mce.2024.112339 · Cited by 49
“Moreover, some EECs directly synapse with enteric neurons via a long cytoplasmic process termed the neuropod (Bohorquez et al., 2015), and can transmit the presence of sugar to the brain in milliseconds (Kaelberer et al., 2018).”
BackgroundEissa, N., Hussein, H. H., Hendy, G. N., Bernstein, C. N., & Ghia, J. (2018) · Biochemical Pharmacology · doi:10.1016/j.bcp.2018.04.009 · Cited by 42
Davis, E. A., Zhou, W., & Dailey, M. J. (2018) · Physiological Reports · doi:10.14814/phy2.13745 · Cited by 38
“SNS and PNS nerves come in close contact with the intestinal epithelium, which supports the idea of a direct autonomic effect on intestinal epithelial cell proliferation and tissue renewal (Gabella and Costa 1968; Bohorquez et al. 2015).”
BackgroundRodrigues, P. V., de Godoy, J. V. P., Bosque, B. P., Neto, D. P. A., Tostes, K., Palameta, S., Garcia-Rosa, S., Tonoli, C. C. C., Carvalho, H. F., & Fonseca, M. D. C. (2022) · Scientific Reports · doi:10.1038/s41598-022-08076-5 · Cited by 40
Vasoactive intestinal peptide regulates ileal goblet cell production in mice
Schwerdtfeger, L., & Tobet, S. A. (2020) · Physiological Reports · doi:10.14814/phy2.14363 · Cited by 38
“There have been demonstrations of neural – epithelial signaling in the intestine via enteroendocrine cells (Bohorquez et al., 2014, 2015), and cholinergic regulation of goblet cells in the eye conjunctiva (Garcia-Posadas et al., 2016).”
BackgroundMicrobiome–gut–brain dysfunction in prodromal and symptomatic Lewy body diseases
Ryman, S. G., Vakhtin, A. A., Richardson, S. P., & Lin, H. C. (2022) · Journal of Neurology · doi:10.1007/s00415-022-11461-9 · Cited by 39
“Historically, they were viewed as hormone-producing cells, but subsequent observations have demonstrated that they are electrically excitable, possess many neuronal features, and can communicate directly with the nervous system [145, 146].”
Cavin, J., Cuddihey, H., MacNaughton, W. K., & Sharkey, K. A. (2019) · American Journal of Physiology-Gastrointestinal and Liver Physiology · doi:10.1152/ajpgi.00186.2019 · Cited by 39
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The Gut‐Brain Axis in Parkinson disease: Emerging Concepts and Therapeutic Implications
Menozzi, E., Schapira, A. H. V., & Borghammer, P. (2025) · Movement Disorders Clinical Practice · doi:10.1002/mdc3.70029 · Cited by 42
“…1% of the total gut epithelial cells and mainly located in the small intestine and colon, 113 EECs attracted the interest of the PD fi eld as they not only display neuronal-like properties including synaptic features and electrical excitability, 108,114 but they also express α -synuclein.”
Zhang, S., Wu, Z., Zhang, S., Ru, Y., Wang, Q., Tong, H., Qin, Q., Yan, Q., Li, Z., & Wu, G. (2025) · Food & Function · doi:10.1039/d5fo03139g · Cited by 39
Gastrointestinal hormones and the gut connectome
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Self-citationAn intravital window to image the colon in real time
Rakhilin, N., Garrett, A., Eom, C., Chavez, K. R., Small, D. M., Daniel, A. R., Kaelberer, M. M., Mejooli, M. A., Huang, Q., Ding, S., et al. (2019) · Nature Communications · doi:10.1038/s41467-019-13699-w · Cited by 40
Self-citationNeuroendocrine gut–brain signaling in obesity
Gruber, T., Lechner, F., Krieger, J.-P., & García‐Cáceres, C. (2024) · Trends in Endocrinology and Metabolism · doi:10.1016/j.tem.2024.05.002 · Cited by 41
“Intriguingly, >50% of Cck + I-cells form pseudopod-like basal processes, which are neuropod specializations that facilitate direct synaptic transmission with underlying mucosal nerves [10,11].”
BackgroundProtease‐dependent excitation of nodose ganglion neurons by commensal gut bacteria
Pradhananga, S., Tashtush, A. A., Allen‐Vercoe, E., Petrof, E. O., & Lomax, A. E. (2020) · The Journal of Physiology · doi:10.1113/jp279075 · Cited by 34
“It is also likely that enteroendocrine cells/neuropod cells (Raybould et al. 2004; Bohorquez et al. 2015; Kaelberer et al. 2018) facilitate communication between luminal microbes and vagal afferent neurons.”
BackgroundSpencer, N. J., Kyloh, M. A., Travis, L., & Hibberd, T. J. (2024) · Cell and Tissue Research · doi:10.1007/s00441-024-03879-6 · Cited by 35
“The idea that EEC may communicate with sensory nerve endings was first conceived based on in vitro co-culturing studies of trigeminal neurons and enteroendocrine cells (Bohorquez et al. 2015).”
BackgroundBacterial modulation of visceral sensation: mediators and mechanisms
Lomax, A. E., Pradhananga, S., Sessenwein, J. L., & O’Malley, D. (2019) · American Journal of Physiology-Gastrointestinal and Liver Physiology · doi:10.1152/ajpgi.00052.2019 · Cited by 36
Zhang, W.-X., Zhang, Y., Qin, G., Li, K.-M., Wei, W., Li, S.-Y., & Yao, S.-K. (2019) · World Journal of Gastroenterology · doi:10.3748/wjg.v25.i43.6416 · Cited by 32
The gut-brain axis in Parkinson's disease
Bonaz, B. L. (2023) · Revue Neurologique · doi:10.1016/j.neurol.2023.11.004 · Cited by 34
IUPHAR review: Microbiota-gut-brain axis and its role in neuropsychiatric disorders
Lee, S.-H., Han, C., & Shin, C. (2025) · Pharmacological Research · doi:10.1016/j.phrs.2025.107749 · Cited by 38
The emerging role of microbiota derived SCFAs in neurodegenerative disorders
Wachamo, S., & Gaultier, A. (2025) · Brain Behavior & Immunity - Health · doi:10.1016/j.bbih.2025.101012 · Cited by 36
“Attributing to their locations and receptors, vagal afferents (VAs) can respond to mechanical, chemical, or hormonal signals in the gut, and transfer sensory information to the brain (Berthoud et al., 2004).”
InfluentialGanz, J., Melançon, E., & Eisen, J. S. (2016) · Methods in cell biology · doi:10.1016/bs.mcb.2016.02.003 · Cited by 32
Dorsal striatum dopamine oscillations: Setting the pace of food anticipatory activity
de Lartigue, G., & McDougle, M. J. (2018) · Acta Physiologica · doi:10.1111/apha.13152 · Cited by 33
“Paracrine signalling between enteroendocrine cells and vagal afferent nerves innervating the gut is thought to be the main mechanism of fat signalling to the brain, although there is also preliminary evidence for direct synaptic signalling between enteroendocrine cells and extrinsic afferent nerves.(100) In addition,…”
BackgroundBile acids, bioactive signalling molecules in interoceptive gut‐to‐brain communication
Joyce, S. A., & O'Malley, D. (2022) · The Journal of Physiology · doi:10.1113/jp281727 · Cited by 32
“, 2017), L-cells also form a neuroepithelial circuit, directly synapsing with sensory afferents (Bohorquez et al., 2015), which signal to the CNS (Buckley et al.”
BackgroundThe now and then of gut-brain signaling
Kaelberer, M. M., & Bohórquez, D. V. (2018) · Brain Research · doi:10.1016/j.brainres.2018.03.027 · Cited by 30
Self-citationEpithelial–Neuronal Communication in the Colon: Implications for Visceral Pain
Najjar, S. A., Davis, B. M., & Albers, K. M. (2020) · Trends in Neurosciences · doi:10.1016/j.tins.2019.12.007 · Cited by 32
“These epithelial cells are electrically excitable and can form synaptic structures with neurons [22, 23].”
BackgroundDistribution and Stimulus Secretion Coupling of Enteroendocrine Cells along the Intestinal Tract
Adriaenssens, A. E., Reimann, F. M., & Gribble, F. M. (2018) · Comprehensive physiology · doi:10.1002/cphy.c170047 · Cited by 39
Interaction between diet and gastrointestinal endocrine cells
El‐Salhy, M., Mazzawi, T. R. E., Hausken, T., & Hatlebakk, J. G. (2016) · Biomedical Reports · doi:10.3892/br.2016.649 · Cited by 30
“Based on these data, it was concluded that the gut endocrine cells have the necessary elements for afferent and efferent synaptic transmission (75).”
Dezfuli, B. S., Giari, L., & Bosi, G. (2021) · Advances in Parasitology/Advances in parasitology · doi:10.1016/bs.apar.2021.03.001 · Cited by 30
Optical Imaging of Pancreatic Innervation
Makhmutova, M., & Caicedo, A. (2021) · Frontiers in Endocrinology · doi:10.3389/fendo.2021.663022 · Cited by 27
“A beautiful example of synapse formation in vitro has been shown by culturing trigeminal sensory neurons with intestinal enteroendocrine cells (98).”
BackgroundGlucagon-like peptide-1 and insulin synergistically activate vagal afferent neurons
Iwasaki, Y., Goswami, C., & Yada, T. (2017) · Neuropeptides · doi:10.1016/j.npep.2017.05.003 · Cited by 31
Guardians of the gut: influence of the enteric nervous system on the intestinal epithelial barrier
Bubeck, M., Becker, C., & Patankar, J. V. (2023) · Frontiers in Medicine · doi:10.3389/fmed.2023.1228938 · Cited by 29
“has revealed the presence of neuropod extensions from the basolateral surfaces of intestinal and colonic enteroendocrine cells that allow them to directly synapse with mucosal efferent and afferent neuronal innervations (55, 56).”
BackgroundChen, Z., Maqbool, J., Sajid, F., Hussain, G., & Sun, T. (2020) · Microbial Pathogenesis · doi:10.1016/j.micpath.2020.104675 · Cited by 28
Gastrointestinal Vagal Afferents and Food Intake: Relevance of Circadian Rhythms
Page, A. J. (2021) · Nutrients · doi:10.3390/nu13030844 · Cited by 32
Boeri, L., Perottoni, S., Izzo, L., Giordano, C., & Albani, D. (2021) · Advanced Healthcare Materials · doi:10.1002/adhm.202002043 · Cited by 28
Li, X., Chen, L.-M., Kumar, G., Zhang, S.-J., Zhong, Q.-H., Zhang, H., Gui, G., Wu, L.-L., Fan, H., & Sheng, J.-W. (2022) · Frontiers in Neuroscience · doi:10.3389/fnins.2022.820106 · Cited by 29
Varied Composition and Underlying Mechanisms of Gut Microbiome in Neuroinflammation
Farooq, R. K., Alamoudi, W., Alhibshi, A., Rehman, S., Sharma, A. R., & Abdulla, F. A. (2022) · Microorganisms · doi:10.3390/microorganisms10040705 · Cited by 31
“Mucosal serotonin permits a pro-inflammatory reaction in the ENS, while neuronal serotonin delivers neuroprotection [52].”
BackgroundInfluentialEl‐Salhy, M., Hausken, T., Gilja, O. H., & Hatlebakk, J. G. (2016) · Expert Review of Gastroenterology & Hepatology · doi:10.1080/17474124.2017.1269601 · Cited by 33
Advances in VNS efficiency and mechanisms of action on cognitive functions
Wang, W., Li, R., Li, C., Liang, Q., & Gao, X. (2024) · Frontiers in Physiology · doi:10.3389/fphys.2024.1452490 · Cited by 31
“…permeability, VNS may also help reduce in fl ammation by controlling gut microbiota and improving cerebrospinal fl uid (CSF) exchange in the glymphatic system (Bohórquez et al., 2015; Bonaz et al., 2019; Cheng et al., 2020; Zhang et al., 2020), though the exact ways it does this need more research.”
BackgroundParkinson's Disease and the Gut: Models of an Emerging Relationship
Bindas, A. J., Kulkarni, S., Koppes, R. A., & Koppes, A. N. (2020) · Acta Biomaterialia · doi:10.1016/j.actbio.2021.03.071 · Cited by 27
The pathogenesis of proventricular dilatation disease
Tizard, I. R., Shivaprasad, H. L., Guo, J., Hameed, S. S., Ball, J. M., & Payne, S. (2016) · Animal Health Research Reviews · doi:10.1017/s1466252316000189 · Cited by 27
“These cells communicate with enteric nerve cells through a cytoplasmic process called a neuropod (Bohorquez et al., 2015).”
BackgroundEnterochromaffin Cells: Sentinels to Gut Microbiota in Hyperalgesia?
Xu, X., Chen, R., Zhan, G., Wang, D., Tan, X., & Xu, H. (2021) · Frontiers in Cellular and Infection Microbiology · doi:10.3389/fcimb.2021.760076 · Cited by 31
“Due to its autonomy, neurotransmitter diversity, and complex cell structure, ENS enjoys the “second brain” reputation (Fattahi et al., 2016).”
BackgroundInfluentialRecent advances in understanding the roles of the enteric nervous system
Chanpong, A., Borrelli, O., & Thapar, N. (2022) · Faculty Reviews · doi:10.12703/r/11-7 · Cited by 30
Neuroimmune pharmacological approaches
Holzer, P. D., Hassan, A. M., Jain, P., Reichmann, F., & Farzi, A. (2015) · Current Opinion in Pharmacology · doi:10.1016/j.coph.2015.09.003 · Cited by 27
“In addition, enteroendocrine cells are capable of direct neurotransmission through specialized cell-nerve contacts termed neuropods where basic elements of neurotransmission such as pre-synaptic, post-synaptic, and trans-synaptic proteins are expressed [22 ].”
The Bidirectional Signal Communication of Microbiota‐Gut‐Brain Axis in Hypertension
Wang, X., Chen, Z., Geng, B., & Cai, J. (2021) · International Journal of Hypertension · doi:10.1155/2021/8174789 · Cited by 29
Metadata
- Authors
- Bohórquez, D. V., Shahid, R. A., Erdmann, A., Kreger, A. M., Wang, Y., Calakos, N., Wang, F., & Liddle, R. A.
- Affiliations
- Duke Medical Center (US)
- Durham VA Medical Center (US)
- Venue
- Journal of Clinical Investigation
- Published
- 2015-01-02
- Volume
- 125
- Issue
- 2
- Pages
- 782-786
- ISSN
- 0021-9738 (ISSN-L)
- PMC
- PMC4319442
- OpenAlex
- W2119429325
- Cited by
- 484
- References
- 28
- Citations by year
- 2026: 42 · 2025: 51 · 2024: 45 · 2023: 45 · 2022: 47 · 2021: 60 · 2020: 47 · 2019: 38 · 2018: 41 · 2017: 22 · 2016: 29 · 2015: 17
- FWCI
- 38.12
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Journal of Clinical Investigation (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- DukeSpace (Duke University) (submittedVersion)
- DukeSpace (Duke University) (submittedVersion, cc-by-nc-nd)
- Funders
- U.S. Department of Veterans Affairs (I01 BX002230)
- National Institutes of Health
- National Institute of Diabetes and Digestive and Kidney Diseases (F32DK094704, R01DK098796, R01DK091946, K01DK103832)
- Topics
- Health Sciences › Medicine › Gastroenterology › Gastrointestinal motility and disorders
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Regulation of Appetite and Obesity
- Keywords
- enteroendocrine cells, sensory neurons, enteric nervous system, neurotransmission, gut-brain signaling, satiety, rabies virus, small intestine
- MeSH terms
- Animals, Cell Communication, Cells, Cultured, Colon, Intestine, Small, Coculture Techniques, Enteroendocrine Cells, Neuroepithelial Cells, Mice
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