The orexigenic hormone acyl-ghrelin increases adult hippocampal neurogenesis and enhances pattern separation
Open accessAbstract
An important link exists between intact metabolic processes and normal cognitive functioning; however, the underlying mechanisms remain unknown. There is accumulating evidence that the gut hormone ghrelin, an orexigenic peptide that is elevated during calorie restriction (CR) and known primarily for stimulating growth hormone release, has important extra-hypothalamic functions, such as enhancing synaptic plasticity and hippocampal neurogenesis. The present study was designed to evaluate the long-term effects of elevating acyl-ghrelin levels, albeit within the physiological range, on the number of new adult born neurons in the dentate gyrus (DG) and performance on the Spontaneous Location Recognition (SLR) task, previously shown to be DG-dependent and sensitive to manipulations of plasticity mechanisms and cell proliferation. The results revealed that peripheral treatment of rats with acyl-ghrelin enhanced both adult hippocampal neurogenesis and performance on SLR when measured 8-10 days after the end of acyl-ghrelin treatment. Our data show that systemic administration of physiological levels of acyl-ghrelin can produce long-lasting improvements in spatial memory that persist following the end of treatment. As ghrelin is potentially involved in regulating the relationship between metabolic and cognitive dysfunction in ageing and neurodegenerative disease, elucidating the underlying mechanisms holds promise for identifying novel therapeutic targets and modifiable lifestyle factors that may have beneficial effects on the brain.
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Ghrelin protects neurons from β-amyloid-induced toxicity.
“It binds to growth hormone secretagogue receptor (GHSR), and it has been shown to protect neurons from β-amyloid-induced toxicity, reduce tau phosphorylation, and enhance synaptic plasticity [20, 44–46].”
From Protective PLCG2 variants associate with a delayed onset of Alzheimer’s disease among heterozygous APOE ε4 carriers (2026) · cites this paper as [45]
The Introduction describes ghrelin as having neuroprotective properties, supporting the general idea of neuronal protection. It does not identify β-amyloid-induced toxicity as a condition against which ghrelin protects neurons; searches for amyloid-related toxicity found no such passage in our copy. (LLM, conf=0.78)
Experimental — accuracy not yet measuredGhrelin enhances synaptic plasticity.
“It binds to growth hormone secretagogue receptor (GHSR), and it has been shown to protect neurons from β-amyloid-induced toxicity, reduce tau phosphorylation, and enhance synaptic plasticity [20, 44–46].”
From Protective PLCG2 variants associate with a delayed onset of Alzheimer’s disease among heterozygous APOE ε4 carriers (2026) · cites this paper as [45]
The Introduction says ghrelin treatments have been shown to affect measures of hippocampal synaptic plasticity, but does not say whether those measures increased. The Results report enhanced neurogenesis in adult rats, which the Introduction describes as a form of plasticity; that does not establish enhanced synaptic plasticity specifically. (LLM, conf=0.82)
Experimental — accuracy not yet measuredElevated ghrelin levels exert anti-inflammatory effects, metabolic regulation, and promote cognitive enhancement.
“This is an interesting finding, as elevated ghrelin levels are known to exert anti-inflammatory effects, metabolic regulation, and promote cognitive enhancement [20, 45, 47].”
From Protective PLCG2 variants associate with a delayed onset of Alzheimer’s disease among heterozygous APOE ε4 carriers (2026) · cites this paper as [45]
The Introduction says ghrelin regulates energy homeostasis, appetite, body weight, and adiposity, and describes pro-cognitive effects. The Discussion reports enhanced pattern-separation memory after elevated systemic acyl-ghrelin treatment. Neither section supports the attributed anti-inflammatory effect; a search of our copy for inflammation found no such passage. Page numbers were unavailable. (LLM, conf=0.87)
Experimental — accuracy not yet measuredGhrelin binds to growth hormone secretagogue receptor (GHSR).
“It binds to growth hormone secretagogue receptor (GHSR), and it has been shown to protect neurons from β-amyloid-induced toxicity, reduce tau phosphorylation, and enhance synaptic plasticity [20, 44–46].”
From Protective PLCG2 variants associate with a delayed onset of Alzheimer’s disease among heterozygous APOE ε4 carriers (2026) · cites this paper as [45]
Ghrelin reduces tau phosphorylation.
“It binds to growth hormone secretagogue receptor (GHSR), and it has been shown to protect neurons from β-amyloid-induced toxicity, reduce tau phosphorylation, and enhance synaptic plasticity [20, 44–46].”
From Protective PLCG2 variants associate with a delayed onset of Alzheimer’s disease among heterozygous APOE ε4 carriers (2026) · cites this paper as [45]
The Introduction discusses ghrelin’s effects on cognition and hippocampal plasticity, while the Results report increased neurogenesis in treated rats. Neither those passages nor searches for “tau phosphorylation” and phosphorylated tau address whether ghrelin reduces tau phosphorylation. (LLM, conf=0.76)
Experimental — accuracy not yet measured
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“demands on behavioral pattern separation processes than when the objects are further apart (Bekinschtein et al. 2013, 2014; Kent et al. 2015).”
BackgroundMethodInfluentialMetabolic hormones mediate cognition
Ghosh‐Swaby, O. R., Reichelt, A. C., Sheppard, P. A. S., Davies, J. S., Bussey, T. J., & Saksida, L. M. (2022) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2022.101009 · Cited by 25 · see citing works
Self-citationPeptides and aging: Their role in anorexia and memory.
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Advances in the Development of Nonpeptide Small Molecules Targeting Ghrelin Receptor
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Ghrelin-Mediated Regeneration and Plasticity After Nervous System Injury
Stoyanova, I., & Lutz, D. (2021) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2021.595914 · Cited by 23 · see citing works
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Less is more: Caloric regulation of neurogenesis and adult brain function
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Self-citationAcyl Ghrelin Improves Synapse Recovery in an In Vitro Model of Postanoxic Encephalopathy
Stoyanova, I., Hofmeijer, J., van Putten, M., & le Feber, J. (2015) · Molecular Neurobiology · doi:10.1007/s12035-015-9502-x · Cited by 19 · see citing works
“synaptic plasticity [26], memory consolidation [32], and adult neurogenesis [33, 34].”
BackgroundBeyond Appetite: Acylated Ghrelin As A Learning, Memory and Fear Behavior-modulating Hormone.
Reich, N., & Hölscher, C. (2022) · Neuroscience and Biobehavioral Reviews · doi:10.1016/j.neubiorev.2022.104952 · Cited by 18 · see citing works
“In opposition to the latter study, (Kent et al., 2015) observed no improvements in DG neuroblast differentiation following the 14 day i.p. administration of AG.”
InfluentialAcylation, a Conductor of Ghrelin Function in Brain Health and Disease
Thomas, A. S., Sassi, M., Angelini, R., Morgan, A. H., & Davies, J. S. (2022) · Frontiers in Physiology · doi:10.3389/fphys.2022.831641 · Cited by 17 · see citing works
Self-citationRatcliff, M., Rees, D., McGrady, S., Buntwal, L., Hornsby, A., Bayliss, J., Kent, B. A., Bussey, T., Saksida, L., Beynon, A. L., et al. (2019) · Journal of neuroendocrinology · doi:10.1111/jne.12755 · Cited by 17 · see citing works
Self-citationPreclinical evidence of ghrelin as a therapeutic target in epilepsy
Ge, T., Yang, W., Fan, J., & Li, B.-J. (2017) · OncoTarget · doi:10.18632/oncotarget.18349 · Cited by 17
“demonstrated that systemic administration of physiological levels of acylated ghrelin for 8–10 days enhances hippocampal neurogenesis and spatial memory in rats [116].”
BackgroundHippocampal neurogenesis and memory in adolescence following intrauterine growth restriction
Gilchrist, C. P., Cumberland, A. L., Kondos‐Devcic, D., Hill, R., Khore, M., Quezada, S., Reichelt, A. C., & Tolcos, M. (2020) · Hippocampus · doi:10.1002/hipo.23291 · Cited by 18 · see citing works
“…studies could use a smaller separation (i.e., xs-SLR) which is shown to require increased hippocampal BDNF (Bekinschtein et al., 2013) and neuroproliferation (Kent et al., 2015) to determine whether subtle cognitive changes in IUGR rats may be parsed with more taxing pattern separation processes.”
MethodRusso, C., Mannino, G., Patanè, M., Parrinello, N., Pellitteri, R., Stanzani, S., Giuffrida, R., Furno, D. L., & Russo, A. (2021) · Histochemistry and Cell Biology · doi:10.1007/s00418-021-01980-3 · Cited by 12 · see citing works
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“To help elucidate the cognitive enhancing effects of ghrelin, we administered a treatment of acyl-ghrelin, peripherally, to adult rats [27 ].”
BackgroundMethodInfluentialSelf-citationGhrelin mediated hippocampal neurogenesis.
Davies, J. (2022) · Vitamins and hormones · doi:10.1016/bs.vh.2021.12.003 · Cited by 12 · see citing works
Self-citationThe 5:2 diet does not increase adult hippocampal neurogenesis or enhance spatial memory in mice
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“These studies provide strong evidence in support of the pro-neurogenic role of acyl-ghrelin signalling (Kent et al., 2015).”
BackgroundInfluentialSelf-citationGhrelin Acylation—A Post-Translational Tuning Mechanism Regulating Adult Hippocampal Neurogenesis
Sassi, M., Morgan, A., & Davies, J. (2022) · Cells · doi:10.3390/cells11050765 · Cited by 10 · see citing works
Self-citationSoldier Health Habits and the Metabolically Optimized Brain
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Glucose availability regulates ghrelin‐induced food intake in the ventral tegmental area
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Self-citationTherapeutic potential of ghrelin/GOAT/GHSR system in gastrointestinal disorders
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Ghrelin promotes neurologic recovery and neurogenesis in the chronic phase after experimental stroke
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“Ghrelin has also been shown to have beneficial effects on learning and memory formation, which may be directly related to its neurogenic properties [9, 21, 22].”
Yun, S., Soler, I., Tran, F. H., Haas, H. A., Shi, R., Bancroft, G. L., Suarez, M., de Santis, C. R., Reynolds, R. P., & Eisch, A. (2023) · bioRxiv · doi:10.1101/2023.01.26.525756 · Cited by 5
“…DG neurogenesis is linked to improved behavioral pattern separation and cognitive flexibility (Bekinschtein et al., 2011; Burghardt et al., 2012; Kent et al., 2015c; McAvoy and Sahay, 2017), we hypothesized that LEC TRIP8b shRNA mice would have more DG neurogenesis compared to SCR shRNA mice.”
BackgroundEffects of Ghrelin on Olfactory Ensheathing Cell Viability and Neural Marker Expression
Russo, C., Patanè, M., Russo, A., Stanzani, S., & Pellitteri, R. (2020) · Journal of Molecular Neuroscience · doi:10.1007/s12031-020-01716-3 · Cited by 5 · see citing works
Factors Regulating Neurogenesis in the Adult Dentate Gyrus
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Neuronal migration depends on blood flow in the adult mammalian brain
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“…Chopin et al., 2012 ; Soleyman-Jahi et al., 2019 ; Stoyanova and Lutz, 2021 ), ghrelin influences neurogenesis in the V-SVZ and subgranular zone of the hippocampal dentate gyrus ( Hornsby et al., 2016 ; Kent et al., 2015 ; Kim et al., 2015 ; Li et al., 2014 ; Li et al., 2013 ; Moon et al., 2009 ).”
Abdulazeez, Y., Utami, R., Al-Jamal, K. T., & Mok, Z.-H. (2025) · ACS Chemical Neuroscience · doi:10.1021/acschemneuro.5c00683 · Cited by 3 · see citing works
Neuronal migration depends on blood flow in the adult brainPreprint
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“…et al., 2017; Chopin et al., 2012; Soleyman-Jahi et al., 2019; Stoyanova and Lutz, 2021) , ghrelin influences neurogenesis in the V-SVZ and subgranular zone of the hippocampal dentate gyrus (Hornsby et al., 2016; Kent et al., 2015; Kim et al., 2015; Li et al., 2014, 2013; Moon et al., 2009) .”
BackgroundSong, W., Lee, H., Lim, J. M., Yoon, S. H., Kim, Y. S., Kim, H., & Kim, M.-H. (2025) · Neuropsychopharmacology · doi:10.1038/s41386-025-02268-z · Cited by 2 · see citing works
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“This is an interesting finding, as elevated ghrelin levels are known to exert anti-inflammatory effects, metabolic regulation, and promote cognitive enhancement [20, 45, 47].”
Reichelt, A., Hashad, A., Ghosh-Swaby, O. R., Skirzewski, M., Scott, G. A., Machado, M. M. F., Kramar, C. P., Princz-Lebel, O., Inoue, W., Bussey, T., et al. (2026) · Neuropsychopharmacology · doi:10.1038/s41386-026-02343-z · Cited by 1 · see citing works
“…(B6N;129-Tg(CAG-CHRM3*,- mCitrine)1Ute/J) mice or R26-hM4Di/mCitrine (B6N.129-Tg(CAG-CHRM4*,- mCitrine)Ute/J) mice expressing a CAG promotor-driven HA-tagged hM3Dq or hM4Di with an upstream fl oxed STOP cassette [39], to create PV hM4Di(+/ − ) or PV hM3Dq(+/ − ) offspring were used.”
Self-citationHornsby, A. K. E., Haywood-Sheldrake, B., Gryksa, K., Havasi, A., Roberts, L. D., Humby, T., Neumann, I., Davies, J. S., & Wells, T. (2025) · Journal of neuroendocrinology · doi:10.1111/jne.70106 · Cited by 1 · see citing works
“22 Indeed, deletion of GHSR, the receptor for ghrelin, inhibits CR-induced augmentation of AHN and associated fear-memory performance.”
Self-citationStark, R., Zigman, J., & Andrews, Z. (2025) · Journal of neuroendocrinology · doi:10.1111/jne.70096 · Cited by 1 · see citing works
Effect of unacylated ghrelin on peripheral nerve regeneration
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Self-citationExtracellular vesicles: Multi-source regulatory mechanisms in hippocampal homeostasis
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“…in hypothalamus and pituitary (20, 23), fat accumulation (17, 24), via GHSR in white (25), and brown (26) adipocytes, including marrow adipocytes (25), suppression of gonadotrophin secretion (27), via hypothalamic GHSR and promotion of adult hippocampal neurogenesis via hippocampal GHSR (28, 29).”
Self-citationThe protective PLCG2 variants delay Alzheimer's disease onset age in APOE ϵ4 carriersPreprint
Jeskanen, H., Heikkinen, S., Kervinen, I., Lehtisalo, J., Ngandu, T., Willman, R.-M., Rosa, J., Hoffmann, D., FinnGen, Leinonen, V., et al. (2025) · medRxiv · doi:10.1101/2025.09.22.25336304
“This is an interesting finding, as elevated ghrelin levels are known to exert anti-inflammatory effects, metabolic regulation, and promote cognitive enhancement (40,42,44).”
Altered protein lipidation in cancer: A hidden link to sensory neuronal dysfunction
Ibrahim, I., Abdellatif, A., & Eassa, H. A. (2025) · Comput. Biol. Chem. · doi:10.1016/j.compbiolchem.2025.108656
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“[14] found that ghrelin is important for significant”
BackgroundKent, B. A. (2015) · Apollo (University of Cambridge) · doi:10.17863/cam.16474
“…(Lee & Sollivan, 2010), touchscreen Location Discrimination (LD) (McTighe et al., 2009; Clelland et al., 2009; Creer et al., 2010), and Spontaneous Location Recognition (SLR) (Hunsaker, Rosenberg, & Kesner, 2008; Bekinschtein et 18 Chapter 1 al., 2013; 2014; Kent et al., 2015).”
InfluentialSelf-citation
- Authors
- Kent, B. A., Beynon, A. L., Hornsby, A. K. E., Bekinschtein, P., Bussey, T. J., Davies, J. S., & Saksida, L. M.
- Affiliations
- University of Cambridge (GB)
- Swansea University (GB)
- Corresponding author
- Davies, Jeffrey S.
- Saksida, Lisa M.
- ORCID
- Venue
- Psychoneuroendocrinology
- Publisher
- Elsevier BV
- Published
- 2015-01
- Volume
- 51
- Pages
- 431-439
- ISSN
- 0306-4530 (ISSN-L)
- PMC
- PMC4275579
- OpenAlex
- W2101777754
- Cited by
- 94
- References
- 36
- Citations by year
- 2026: 7 · 2025: 10 · 2024: 8 · 2023: 4 · 2022: 9 · 2021: 7 · 2020: 6 · 2019: 9 · 2018: 9 · 2017: 12 · 2016: 6 · 2015: 7
- FWCI
- 2.90
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Psychoneuroendocrinology (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- CONICET Digital (CONICET) (publishedVersion, cc-by-nc-sa)
- Americanae (AECID Library) (submittedVersion, cc-by)
- Scholarship@Western (Western University) (submittedVersion)
- LA Referencia (Red Federada de Repositorios Institucionales de Publicaciones Científicas) (submittedVersion, cc-by)
- Apollo (University of Cambridge) (submittedVersion, cc-by)
- Funders
- Royal Society
- Medical Research Council (G0902250/94306)
- Biotechnology and Biological Sciences Research Council (BB/G019002/1)
- LMS
- Gates Cambridge
- Topics
- Life Sciences › Neuroscience › Developmental Neuroscience › Neurogenesis and neuroplasticity mechanisms
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Regulation of Appetite and Obesity
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- acylated ghrelin, ghrelin, adult hippocampal neurogenesis, dentate gyrus, pattern separation, spatial memory, caloric restriction, gastrointestinal hormones, synaptic plasticity, neurodegeneration
- MeSH terms
- Animals, Hippocampus, Male, Rats, Ghrelin, Neurogenesis, Spatial Memory
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