The DOI matches its Crossref record; 138 of the 138 references it cites were found in a registry or an index.
A PHA-L analysis of ascending projections of the dorsal raphe nucleus in the rat
Abstract
Ascending projections from the dorsal raphe nucleus (DR) were examined in the rat by using the anterograde anatomical tracer, Phaseolus vulgaris leucoagglutinin (PHA-L). The majority of labeled fibers from the DR ascended through the forebrain within the medial forebrain bundle. DR fibers were found to terminate heavily in several subcortical as well as cortical sites. The following subcortical nuclei receive dense projections from the DR: ventral regions of the midbrain central gray including the 'supraoculomotor central gray' region, the ventral tegmental area, the substantia nigra-pars compacta, midline and intralaminar nuclei of the thalamus including the posterior paraventricular, the parafascicular, reuniens, rhomboid, intermediodorsal/mediodorsal, and central medial thalamic nuclei, the central, lateral and basolateral nuclei of the amygdala, posteromedial regions of the striatum, the bed nucleus of the stria terminalis, the lateral septal nucleus, the lateral preoptic area, the substantia innominata, the magnocellular preoptic nucleus, the endopiriform nucleus, and the ventral pallidum. The following subcortical nuclei receive moderately dense projections from the DR: the median raphe nucleus, the midbrain reticular formation, the cuneiform/pedunculopontine tegmental area, the retrorubral nucleus, the supramammillary nucleus, the lateral hypothalamus, the paracentral and central lateral intralaminar nuclei of the thalamus, the globus pallidus, the medial preoptic area, the vertical and horizontal limbs of the diagonal band nuclei, the claustrum, the nucleus accumbens, and the olfactory tubercle. The piriform, insular and frontal cortices receive dense projections from the DR; the occipital, entorhinal, perirhinal, frontal orbital, anterior cingulate, and infralimbic cortices, as well as the hippocampal formation, receive moderately dense projections from the DR. Some notable differences were observed in projections from the caudal DR and the rostral DR. For example, the hippocampal formation receives moderately dense projections from the caudal DR and essentially none from the rostral DR. On the other hand, virtually all neocortical regions receive significantly denser projections from the rostral than from the caudal DR. The present results demonstrate that dorsal raphe fibers project significantly throughout widespread regions of the midbrain and forebrain.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“The MR serotonin neurons project to structures near the midline, whereas the DR serotonin neurons target more lateral regions (Jacobs and Azmitia, 1992; Muzerelle et al., 2016; Vertes, 1991; Vertes et al., 1999).”
From Single-cell transcriptomes and whole-brain projections of serotonin neurons in the mouse dorsal and median raphe nuclei (2019) · cites this paper as Vertes, 1991
In the citing paper
“DRN 5-HT neurons send highly divergent projections that target many functionally distinct brain regions (Azmitia and Segal, 1978; Muzerelle et al., 2016; Vertes, 1991).”
From Molecular and anatomical organization of the dorsal raphe nucleus (2019) · cites this paper as Vertes, 1991
Citations
Cites
Steinbusch, H. W. M. (1981) · Neuroscience · doi:10.1016/0306-4522(81)90146-9 · Cited by 2,415
Central cholinergic pathways in the rat: An overview based on an alternative nomenclature (Ch1–Ch6)
Mesulam, M. M., Mufson, E. J., Wainer, B. H., & Levey, A. I. (1983) · Neuroscience · doi:10.1016/0306-4522(83)90108-2 · Cited by 2,329
Azmitia, E. C., & Segal, M. (1978) · The Journal of Comparative Neurology · doi:10.1002/cne.901790311 · Cited by 1,908
Emson, P. C., Heffher, R. R., & Pearson, J. (1984) · Journal of Neuropathology & Experimental Neurology · doi:10.1097/00005072-198409000-00010 · Cited by 1,651
Efferent connections of the substantia nigra and ventral tegmental area in the rat
Beckstead, R. M., Domesick, V. B., & Nauta, W. J. H. (1979) · Brain Research · doi:10.1016/0006-8993(79)91001-1 · Cited by 1,293
The cortical projections of the mediodorsal nucleus and adjacent thalamic nuclei in the rat
Krettek, J. E., & Price, J. L. (1977) · The Journal of Comparative Neurology · doi:10.1002/cne.901710204 · Cited by 1,277
Gerfen, C. R., & Sawchenko, P. E. (1984) · Brain Research · doi:10.1016/0006-8993(84)90940-5 · Cited by 1,205
Organization of motor and somatosensory neocortex in the albino rat
Hall, R. D., & Lindholm, E. P. (1974) · Brain Research · doi:10.1016/0006-8993(74)90076-6 · Cited by 897
Kelley, A. E., Domesick, V. B., & Nauta, W. J. H. (1982) · Neuroscience · doi:10.1016/0306-4522(82)90067-7 · Cited by 888
Leonard, C. M. (1969) · Brain Research · doi:10.1016/0006-8993(69)90003-1 · Cited by 850
Research works printed but not linked: 9
- Fraser H., 1976, Slow Virus Disease of Animals and Man, 267
- Irle E., 1986, Afferent connections of the subatantia innominata/basal nucleus of Meynert in carnivores and primates, J. Hirnforsch., 27, 343
- Klein J., 1985, The new drug they call “Ecstasy”. Is it too much to swallow?, 38
- Pasik T., 1982, Serontonergic afferents in the monkey neostriatum, Acta Biol. Sci. Hung., 33, 277
- Vertes R. P., 1989, An anterograde PHA study of the efferent projections of the dorsal raphe nucleus in the rat, Neurosci Abstr., 15, 450
- Vertes R. P., 1990, Brainstem Mechanisms of Behaviour, 33
- Vertes R. P., 1990, Brainstem Mechanisms of Behaviour, 535
- Zilles K., 1985, A stereotaxi atlas
- Zilles K., 1985, The Rat Nervous System, 375
Cited by
Sorted by citations
Items 801–843 of 843
Carrette, L. L. G., Sneddon, E. A., & George, O. (2023) · Elsevier eBooks · doi:10.1016/b978-0-12-823453-2.00003-5
Linking serotonergic median raphe input to dorsal CA1 with mnemonic functionsPreprint
Gerdey, J., & Masseck, O. A. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.09.04.556213
Rosário, B. D. A., de Lima, M. P., Vieira, M. G., de Aquino, P. É. A., Viana, G. S. D. B., Silveira, E. R., de Brito, D. H. A., Zampieri, D., Lemes, J. A., de Nazaré, M. D. F. S., et al. (2023) · Research Square · doi:10.21203/rs.3.rs-3705796/v1
(2024) · Elsevier eBooks · doi:10.1016/b978-0-443-15596-3.16001-1
Alosaimi, F., Temel, Y., & Jahanshahi, A. (2024) · Deep Brain Stimulation · doi:10.1016/j.jdbs.2024.12.002
Interplay Between Serotonin Transporter and Glucocorticoid Receptor on Anxiety and Depression
Padovan, C. M., Tirapelli, C. R., Campos‐Cardoso, R., & Souza, C. D. D. M. (2024) · Springer eBooks · doi:10.1007/978-3-031-32035-4_19-1
Serotonin neurons are necessary for tonic sodium intake inhibition
Gasparini, S., Buchanan, G. F., & Geerling, J. C. (2025) · American Journal of Physiology-Regulatory, Integrative and Comparative Physiology · doi:10.1152/ajpregu.00029.2025
Interplay Between Serotonin Transporter and Glucocorticoid Receptor on Anxiety and Depression
Padovan, C. M., Tirapelli, C. R., Campos‐Cardoso, R., & Souza, C. D. D. M. (2025) · Springer eBooks · doi:10.1007/978-3-031-73368-0_19
George, B. E., Vidrascu, E., Neira, S., Devine, M. P., & Kash, T. L. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.01.30.702832
“The ventral pallidum receives moderately dense input from the dorsal raphe nucleus 40–42 and possesses many of the 5-HT receptor subtypes, including from the 5-HT-1, -2, -3, and -4 classes 43–48 .”
Saidi, L., Rioux, V., Wallman, M., Rajan, S. S., Metzakopian, E., Lévesque, M., Proulx, C. D., & Parent, M. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.02.23.707446
Sapin, E. (2009) · theses.fr (ABES) · doi:10.70675/220c13b8z7dc6z47cazb09bz8dab989d204f
Contribution des récepteurs 5-HT4 à la motivation et la prise de décision de manger
Jean, A. (2010) · theses.fr (ABES) · doi:10.70675/804a4a80z3585z415fzba64z9a7f076dd88d
Rôle du noyau paragigantocellulaire latéral dans le réseau du sommeil paradoxal chez le rat
Sirieix, C. M. (2011) · theses.fr (ABES) · doi:10.70675/b976b36fzcb70z4aa8z88baze1ffe48eca8a
The role of serotoninergic neurons in amyotrophic lateral sclerosis
Oussini, H. E. (2016) · theses.fr (ABES) · doi:10.70675/2431e8ddzd4e4z4710z81dcz14100daa6294
Contribution de noyaux hypothalamiques et de leur interconnexion à la régulation du sommeil
Varin, C. (2016) · theses.fr (ABES) · doi:10.70675/72080df7z22e5z4258za973z0aeccd16aef5
Mansouri-Guilani, N. (2017) · theses.fr (ABES) · doi:10.70675/99ced3b5zeac6z4eb2z8e6cz01bcad7cf2d9
Rôle du transporteur plasmique des monoamines (PMAT) dans le système nerveux central
Amin, S. R. (2017) · theses.fr (ABES) · doi:10.70675/f562837bz937fz402eza824z24af264eaf90
Rôle de la protéine matricellulaire hevin dans la réponse à la cocaïne
Mongrédien, R. (2018) · theses.fr (ABES) · doi:10.70675/926d644czef3bz4ce3z9fd7z6080a7628079
Serotonin and emotional behaviours : developmental role of the 5-HT7 receptor
Olusakin, J. (2019) · theses.fr (ABES) · doi:10.70675/480ab91dz9252z4d95z8749z992be4697690
On the developmental origin of CA1 pyramidal neuron diversity in the adult mouse hippocampus
Cavalieri, D. (2021) · theses.fr (ABES) · doi:10.70675/af05b2d4zc031z48d7z9ebaz859f489a8450
All-optical interrogation of the role of interneurons in CA1 cell assemblies
Vorobyev, A. (2023) · theses.fr (ABES) · doi:10.70675/eb10a4e1za595z4f16z9e3bz115344f3d87a
Socio-communicative and nucleus accumbens core dysfunctions in fragile X syndrome
Giua, G. (2023) · theses.fr (ABES) · doi:10.70675/e45d048ez2538z4905zba74z1de16201406a
New functional insights from the monkey claustrum
Wulff, A. B., & Mathur, B. N. (2026) · National Science Review · doi:10.1093/nsr/nwag414
“These include the thalamic intralaminar regions, thalamic nucleus reuniens, basolateral amygdala, basal forebrain, supramammillary nucleus, dorsal raphe nucleus, and arguably the patch compartments of the striatum [79 ,80 ,89 –95 ].”
Contextualized phenovectors reveal a Drosophila serotonergic circuit controlling satietyPreprint
Xu, S., Zhang, X., Stewart, J. C., Ho, J., Choudhury, D., Wang, Z., & Claridge‐Chang, A. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.07.01.735755
Mutlu, A. K., Serneels, B., Wiest, C., Trinh, A., Bardenhewer, R., Palumbo, F., Frisvold, O. B., Aukrust, I. K. F., Ostenrath, A. M., & Yaksi, E. (2026) · Nature Communications · doi:10.1038/s41467-026-75490-y · Cited by 5
“Axonal projections of DRN neurons broadly innervate the vertebrate forebrain 35,54,90,91 .”
Ultrasonic potentiation of ketamine neuromodulationPreprint
Roy, K. S., Martinez, P. J., Ewbank, S. N., Shinozuka, K., Purohit, M. P., Xiang, Y., & Airan, R. D. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.07.29.741494
“…5-HT signal may reflect engagement of slower neuromodulatory loops, possibly via mPFC projections to the dorsal raphe and reciprocal serotonergic innervation of frontal and striatal targets, which integrate over longer timescales than the fast glutamatergic and GABAergic responses [58–60].”
Vertes, R. P., Linley, S. B., & Mitchell, A. S. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-24078-2.00014-0
Wheeler, D. W., Witter, M. P., & Ascoli, G. A. (2026) · Elsevier eBooks · doi:10.1016/b978-0-443-24078-2.00019-x
Yuan, Q., Wang, S., & Tang, Z. (2026) · European Journal of Neuroscience · doi:10.1111/ejn.70653
“The DRN sends dense projections to key limbic and cortical regions, including the prefrontal cortex, hippocampus, and amygdala (Ren et al. 2018; Vertes 1991), that are vital for emotional processing.”
Rouleau, R., Oubraim, S., Cirone, L., Shen, R., & Haj‐Dahmane, S. (2026) · Frontiers in Cellular Neuroscience · doi:10.3389/fncel.2026.1951375
“…studies have shown that the CLA receives strong inputs from serotonergic (5-HT) neurons of dorsal raphe nucleus (DRN) (Zingg et al., 2018; Vertes, 1991; Peyron et al., 1998), and expresses high levels of 5-HT receptors, including 5-HT1A, 5-HT1B, 5-HT1F, 5-HT2A and 5-HT2C receptors…”
Huang, K. W. (2019) · Harvard Dataverse · doi:10.7910/dvn/qb5cc8
The Serotoninergic System in Sleep and Narcolepsy
Alexandre, C., & Scammell, T. E. (2011) · Springer eBooks · doi:10.1007/978-1-4419-8390-9_7
“In particular, Vertes and colleagues demonstrated that both the DRN and the MRN project to the forebrain, but to separate regions with very little overlap [ 12 , 13].”
BackgroundDefelipe (1999) · doi:10.1002/(sici)1096-9861(19990927)412:3
Rinaman (1998) · doi:10.1002/(sici)1096-9861(19980914)399:1
Metadata
- Authors
- Vertes, R. P.
- Affiliations
- Florida Atlantic University (US)
- Corresponding author
- Vertes, Robert P.
- ORCID
- Vertes, Robert P.
- Venue
- Journal of Comparative Neurology
- Publisher
- Wiley
- Published
- 1991-11-22
- Volume
- 313
- Issue
- 4
- Pages
- 643-668
- ISSN
- 0021-9967 (ISSN-L)
- 1096-9861
- OpenAlex
- W1984112731
- Cited by
- 842
- References
- 138
- Citations by year
- 2026: 12 · 2025: 16 · 2024: 17 · 2023: 20 · 2022: 17 · 2021: 28 · 2020: 27 · 2019: 26 · 2018: 25 · 2017: 32 · 2016: 31 · 2015: 36 · 2014: 36 · 2013: 29 · 2012: 32
- FWCI
- 4.18
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- The Journal of Comparative Neurology (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institute of Mental Health
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- dorsal raphe nucleus, ascending projections, serotonergic system, medial forebrain bundle, thalamus, amygdala, striatum, hippocampal formation, anterograde tracing, Phaseolus vulgaris leucoagglutinin, periaqueductal gray, substantia nigra pars compacta, ventral tegmental area
- MeSH terms
- Animals, Cerebral Cortex, Hippocampus, Immunohistochemistry, Male, Neural Pathways, Phytohemagglutinins, Raphe Nuclei, Rats, Inbred Strains, Rats
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review