Cited by 1,153Point processes and geometric inequalitiesMemory and Neural MechanismsClassical Anatomy and Histology

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Unbiased stereological estimation of the number of neurons in the human hippocampus

West, Mark J., Gundersen, H. J. G.·1990·Journal of Comparative Neurology

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Abstract

The total numbers of neurons in five subdivisions of human hippocampi were estimated using unbiased Stereological principles and systematic sampling techniques. The method addresses the problems associated with the results and conclusions of previous quantitative studies, virtually all of which have been based on biased estimates of neuron densities. For each subdivision, the total number of neurons was calculated as the product of the estimate of the volume of the neuron‐containing layers and the estimate of the numerical density of neurons in the layers. Each hippocampus was cut into 3‐mm‐thick slabs, transverse to the rostrocaudal axis. One 70‐μUm‐thick section from each slab was used in the analysis. The volumes of the layers containing neurons in five major subdivisions of the hippocampus (granule cell layer, hilus, CA3–2, CA1, and subiculum) were estimated with point‐counting techniques after delineation of the layers on each section. The numerical densities of neurons in each subdivision were estimated on the same sections with optical directors. The sampling used in both estimates was performed systematically in all three dimensions. In an example of five hippocampi, the mean numbers of neurons (CV = SD/mean) in the different subdivisions were as follows: granule cells 15 × 10 6 (0.28), hilus 2.0 × 10 6 (0.16), CA3‐2 2.7 × 10 6 (0.22), CA1 16 × 10 6 (0.32), subiculum 4.5 × 10 6 (0.19). The stereological measurements contributed approximately 25% of the observed variance. Among the five subjects there was a significant inverse relationship between age (which ranged from 47 to 85 years) and the total number of neurons in CA1 (which ranged from 24 to 11 × 10 6). An optimized sampling scheme for studies of the number of neurons in the human hippocampus has been designed on the basis of an analysis of variance of the estimates at different levels of the sampling scheme. Counting neurons in the five subdivisions of the human hippocampus with the optimized sampling scheme takes less than 4 hours.

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“Estimates Developmental Neurobiology of the total number of labeled SNB motoneurons were obtained using the optical dissector method outlined by Coggeshall (1992) and a procedure similar to that of West and Gunderson (1990).”
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“The procedure for acquiring images, and unbiased stereological analysis were the same as we previously reported or reported by others (West and Gundersen, 1990; Bose et al., 2005; Bernstein et al., 2011; Bernstein and O’Malley, 2013; Hou et al., 2014; Mustafa et al., 2016).”
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Background
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Method
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Self-citation
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“The two-stage (Nv × Vref) approach using the optical dissector and Cavalieri method (West and Gundersen, 1990) was used to calculate the total number of TH-immunoreactive cells in animals subjected to saline or MPTP-treatment regimen.”
Method
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“Because the reflected blade can be difficult to separate from the hilar polymorphic cell layer, it is usually considered to be part of a hilar cell population that comprises both CA3 reflected blade cells and hilar polymorphic cells (West and Gundersen, 1990; Holm and West, 1994; Harding et al., 1998; Keuker et al., 2…”
Background
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“The composition of excitatory (90%) and inhibitory (10%) neurons in the hippocampus (West and Gundersen 1990; Walker et al. 2002; Booker and Vida 2018) makes it more vulnerable to the effects of excitation–inhibition imbalance than other brain regions (Freund and Buzsáki 1996).”
Background
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“Gundersen, H.J.G. Stereology of arbitrary particles: A review of unbiased number and size estimators and the presentation of some new ones, in memory of William, R. Thompson.”
BackgroundInfluential
A Morphometric Comparative Study of the Medial Geniculate Body of the Rabbit and the Fox
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“The numerical density of the cells in each of the MGB nuclei was estimated using the optical dissector method described in detail by West and Gundersen (1990). ª 2011 Blackwell Verlag GmbH • Anat.”
MethodInfluential
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“The MetaMorph software package (Molecular Devices, Sunnyvale, USA) with an unbiased stereological protocol as previously described was used to count [27, 28].”
Method
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“Subsequently, based on unbiased stereological techniques to estimate neuron number, the overall results suggested that there was no widespread hippocampal cell loss in both human and rodent models during aging [12-18].”
Method
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“Details of stereological counting techniques have been described (West and Gundersen, 1990; West, 2002).”
Method
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Cell density quantification of high resolution Nissl images of the juvenile rat brain
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“The conventional stereological approach to prevent cell overcounting during consecutive slice density estimation involves excluding cells on half of the boundaries of the sampled areas (West, 1999; West and Gundersen, 1990).”
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“Hippocampal formation areas (GD), hilus, CA2/3, CA1, SUB, and white matter were identified according to the method described by West and Gundersen [65].”
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Neuronal volume of the hippocampal regions in ageing
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“…of the hippocampal neurons is affected and this may be detected by precise quantitative analyses that have been widely used in the neurosciences (West and Gundersen, 1990; Rapp et al., 1996; West and Slomianka, 1998; Slomianka and West, 2005) in the successful assessment of different…”
Validation and use of a computer-assisted counting procedure to quantify BrdU-labeled proliferating cells in the early postnatal mouse hippocampus
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“First, although typical sterology utilizes a top and bottom guard zone of the section to avoid double counting of cells present on two adjacent sections (West and Gundersen, 1990), this is difficult to implement for automated counting.”
Effect of corticosterone on developing hippocampus: Short‐term and long‐term outcomes
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Thalamic neuropathology in the chronic pilocarpine and picrotoxin model of epilepsy
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Compensatory paracrine mechanisms that define the urothelial response to injury in partial bladder outlet obstruction
Lendvay, T. S., Sweet, R. M., Han, C. H., Soygür, T., Cheng, J., Plaire, J. C., Charleston, J. S., Charleston, L. B., Bagai, S., Cochrane, K., et al. (2007) · American Journal of Physiology-Renal Physiology · doi:10.1152/ajprenal.00006.2007 · Cited by 7
Persistent neuronal density changes related to the establishment of a motor memory
Morales, P., Pinto‐Hamuy, T., Fernández, V., & Dı́az, E. (1999) · Behavioural Brain Research · doi:10.1016/s0166-4328(98)00099-0 · Cited by 7
“Neuronal density was quantified using the optical disector method [17,42,43].”
Method
Stereological Estimation of Neuronal Number: The Optical Disector/Cavalieri Combination
Nurcombe, V., Wreford, N. G., & Bertram, J. F. (2003) · Humana Press eBooks · doi:10.1385/1-59259-060-8:251 · Cited by 7
Pattern of retinoid-induced teratogenic effects: Possible relationship with relative selectivity for nuclear retinoid receptors RARα, RARβ, and RARγ
Elmazar, M. M., Reichert, U., Shroot, B., & Nau, H. (1996) · Teratology · doi:10.1002/(sici)1096-9926(199603)53:3 · Cited by 45
Adult neurogenesis—a reality check
Frielingsdorf, H., & Kuhn, H. G. (2007) · Debates in Neuroscience · doi:10.1007/s11559-007-9006-6 · Cited by 7
“Furthermore, the number of new cells born each day make up about 1% of the total number of granule cells in an adult rat [49, 85].”
Background
Inhibition of reactive oxygen species production accompanying alternatively activated microglia by risperidone in a mouse ketamine model of schizophrenia
Fujikawa, R., Yamada, J., Maeda, S., Iinuma, K., Moriyama, G., & Jinno, S. (2024) · Journal of Neurochemistry · doi:10.1111/jnc.16133 · Cited by 6
“A total of 18 mice (control, n = 6; KET-treated, n = 6; KET/RIS-treated, n = 6) were used for counting TMEM119 + /Iba1 + microglia and TMEM119 − /Iba1 + macrophages with an optical disector (West & Gundersen, 1990 ).”
Distribution and Morphology of Cortical Terminals in the Cat Thalamus from the Anterior Ectosylvian Sulcus
Huppé‐Gourgues, F., Farishta, R. A., Boire, D., Ptito, M., & Casanova, C. (2019) · Scientific Reports · doi:10.1038/s41598-019-39327-7 · Cited by 6
Dose-Related Reduction in Hippocampal Neuronal Populations in Fetal Alcohol Exposed Vervet Monkeys
Burke, M., Slimani, H., Ptito, M., Ervin, F. R., & Palmour, R. M. (2022) · Brain Sciences · doi:10.3390/brainsci12091117 · Cited by 6
“Stereology Design-based stereological estimation of neuronal numbers was achieved by using the optical fractionator method [32] on cresyl violet-stained sections.”
Method
Natural and Pathological Aging Distinctively Impacts the Pheromone Detection System and Social Behavior
Portalés, A., Chamero, P., & Jurado, S. (2023) · Molecular Neurobiology · doi:10.1007/s12035-023-03362-3 · Cited by 9
“unbiased quantification method: a 3D optical dissector for cell counting and a fractionator, based on a systematic, uniform, and random sampling over a known area [28, 29].”
Orthotopic transplantation of immortalized mesencephalic progenitors (CSM14.1 cells) into the substantia nigra of hemiparkinsonian rats induces neuronal differentiation and motoric improvement
Haas, S. J.-P., Petrov, S., Kronenberg, G., Schmitt, O., & Wree, A. (2007) · Journal of Anatomy · doi:10.1111/j.1469-7580.2007.00834.x · Cited by 6
“The coefficient of error (West & Gundersen, 1990) was smaller than 0.1.”
BackgroundInfluential
Conditional deletion of Neurog1 in the cerebellum of postnatal mice delays inhibitory interneuron maturation
Obana, E. A., Zhou, Q., Furmanski, O., & Doughty, M. L. (2018) · Journal of Neuroscience Research · doi:10.1002/jnr.24247 · Cited by 6
“All stereological measures yielded a Gundersen coefficient error of <.16 (Geinisman, Gundersen, van der Zee, & West, 1996; West & Gundersen, 1990).”
Background
Ectopic expression of aPKC-mediated phosphorylation in p300 modulates hippocampal neurogenesis, CREB binding and fear memory differently with age
Syal, C., Seegobin, M., Sarma, S. N., Gouveia, A., Hsu, K., Niibori, Y., He, L., Wondisford, F. E., Frankland, P. W., & Wang, J. (2018) · Scientific Reports · doi:10.1038/s41598-018-31657-2 · Cited by 6
Large time step discrete-time modeling of sharp wave activity in hippocampal area CA3
Malerba, P., Rulkov, N. F., & Bazhenov, M. (2018) · Communications in Nonlinear Science and Numerical Simulation · doi:10.1016/j.cnsns.2018.12.009 · Cited by 6
Structural features of neurons in whole grafts of the rat inferior colliculus
Zrull, M. C., & Coleman, J. R. (1991) · Hearing Research · doi:10.1016/0378-5955(91)90098-t · Cited by 6
Temporal adaptation in the testes of rat administered single dose Momordica charantia for three interrupted spermatogenic cycles: Cytometric quantification
Yama, O. E., & Duru, F. I. (2011) · Middle East Fertility Society Journal · doi:10.1016/j.mefs.2011.07.001 · Cited by 6
Matrix Metalloproteinases 2 and 9 Fail to Influence Drug-Induced Neuroapoptosis in Developing Rat Brain
Uckermann, O., Luksch, H., Stefovska, V. G., Hoehna, Y., Marzahn, J., Theil, M., Pesic, M., Górkiewicz, T., Gawlak, M., Wilczynski, G. M., et al. (2010) · Neurotoxicity Research · doi:10.1007/s12640-010-9211-1 · Cited by 6
“The numerical densities of degenerating cells in each brain region were estimated using the stereological dissector method (West and Gundersen 1990).”
Method
Age‐dependent changes in the glial cell nests of the canine rhinencephalic allocortex. A morphometric study
Reihlen, A., Weis, S., Obermaier, G., & Dahme, E. (1994) · The Anatomical Record · doi:10.1002/ar.1092380316 · Cited by 6
Absolute number of parvicellular and magnocellular neurons in the red nucleus of the rat midbrain: a stereological study
Aghoghovwia, B. E., & Oorschot, D. E. (2016) · Journal of Anatomy · doi:10.1111/joa.12495 · Cited by 6
“…once more, highlights that the major factor contributing to the overall variability was the biological dif- ferences (biological variation) in absolute neuronal number and not the precision of the estimates made with the stere- ological techniques applied (West & Gundersen, 1990; Oorschot, 1996).”
BackgroundMethodInfluential
Effect of Prenatal Protein Malnutrition on Birthdates and Number of Neurons in the Rat Locus Coeruleus
King, R. S., Kemper, T. L., DeBassio, W. A., Blatt, G. J., Ramzan, M., Rosene, D. L., & Galler, J. R. (1999) · Nutritional Neuroscience · doi:10.1080/1028415x.1999.11747283 · Cited by 6
Towards obtaining unbiased estimates of the total number of synapses in a brain region: problems of primary and secondary importance
Geinisman, Y., Gundersen, H. J. G., van der Zee, E., & West, M. J. (1997) · Journal of Neurocytology · doi:10.1023/a:1018558130094 · Cited by 6
Self-citation
Absolute Number of Three Populations of Interneurons and All GABAergic Synapses in the Human Hippocampus
Takács, V. T., Papp, P., Orosz, Á., Bardóczi, Z., Zsoldos, T., Zichó, K., Watanabe, M., Maglóczky, Z., Gombás, P., Freund, T. F., et al. (2025) · Journal of Neuroscience · doi:10.1523/jneurosci.0372-24.2024 · Cited by 6
Neuronal deletion of phosphatase and tensin homolog in mice results in spatial dysregulation of adult hippocampal neurogenesis
Latchney, S. E., Lopez, B. R. R., Womble, P. D., Blandin, K. J., & Lugo, J. N. (2023) · Frontiers in Molecular Neuroscience · doi:10.3389/fnmol.2023.1308066 · Cited by 5
“Moreover, when the number of cells is reported as per unit volume of tissue, there is no information about the volume of the structure (West and Gundersen, 1990).”
BackgroundInfluential
Decreased neuronal and increased endothelial fractalkine expression are associated with neuroinflammation in Parkinson’s disease and related disorders
Chu, Y., Harms, A. S., Boehringer, A. V., & Kordower, J. H. (2025) · Frontiers in Cellular Neuroscience · doi:10.3389/fncel.2025.1557645 · Cited by 5
A simplified application of systematic field sampling and low‐cost video recording set‐up for viewing disector pairs – exemplified in the rat cochlear nucleus
Korkmaz, A., Çiftçi, N., Boşnak, M., & Agˇar, E. (2000) · Journal of Microscopy · doi:10.1046/j.1365-2818.2000.00749.x · Cited by 5
“The literature dealing with the issue of systematic random section field sampling shows that the use of an automated stage is almost an integral part of the procedure (e.g. West & Gundersen, 1990; West et al., 1991; Skoglund et al., 1996; Howard & Reed, 1998; Hyman et al., 1998).”
BackgroundMethod
Quantitative analysis of odontoblast cells in fluorotic and nonfluorotic primary tooth pulp
Şentut, T. K., Kırzıoğlu, Z., Gökçımen, A., Aslan, H., & Erdoğan, Y. (2012) · TURKISH JOURNAL OF MEDICAL SCIENCES · doi:10.3906/sag-1007-964 · Cited by 5
Stereological Methods to Quantify Cell Loss in the Huntington’s Disease Human Brain
Mehrabi, N. F., Singh‐Bains, M. K., Waldvogel, H. J., & Faull, R. L. M. (2018) · Methods in molecular biology · doi:10.1007/978-1-4939-7825-0_1 · Cited by 5
Society of Agents
Walczak, S. M. (2018) · International Journal of Intelligent Information Technologies · doi:10.4018/ijiit.2018100101 · Cited by 5
Chronic administration of quetiapine stimulates dorsal hippocampal proliferation and immature neurons of male rats, but does not reverse psychosocial stress-induced hyponeophagic behavior
Murata, Y., Matsuda, H., Mikami, Y., Hirose, S., Mori, M., Ohe, K., Mine, K., & Enjoji, M. (2018) · Psychiatry Research · doi:10.1016/j.psychres.2018.12.137 · Cited by 5
Classes and gradients of prefrontal cortical organization in the primate
Hilgetag, C. C., Dombrowski, S. M., & Barbas, H. (2002) · Neurocomputing · doi:10.1016/s0925-2312(02)00479-4 · Cited by 5
Pyramidal neurons in the septal and temporal CA1 field of the human and hedgehog tenrec hippocampus
Liagkouras, I., Michaloudi, H. C., Batzios, C., Psaroulis, D., Georgiadis, M., Künzle, H., & Papadopoulos, G. C. (2008) · Brain Research · doi:10.1016/j.brainres.2008.04.037 · Cited by 5
Aortic coarctation hypertension induces fibroblast growth factor-2 immunoreactivity in the stimulated nucleus tractus solitarii
Fior-Chadi, D. R., Varella, T. C. N., Maximino, J. R., & Chadi, G. (2007) · Journal of Molecular Histology · doi:10.1007/s10735-007-9101-x · Cited by 5
“The optical fractionator ( West and Gundersen 1990; Chadi et al. 1993a; Janson and Moller 1993) was used to estimate the number of FGF-2, GFAP and OX42 immunoreactive profiles in the sampled volume of the NTS, bilaterally.”
Method
About regeneration of the brain (Lecture II)
Paltsyn, A. A., & Свиридкина, Н. Б. (2018) · Nauchno-prakticheskii zhurnal «Patogenez» · doi:10.25557/2310-0435.2018.01.83-91 · Cited by 6
Factors Influencing Neuronal Density on Sections: Quantitative Data Obtained by Computer Simulation
Duyckaerts, C. J., Delaère, P., Costa, C., & Hauw, J. (1992) · Methods in neurosciences · doi:10.1016/b978-0-12-185269-6.50037-0 · Cited by 5
Research uses of neuropathological data.
Coleman, P. D. (1997) · Neurobiology of Aging · doi:10.1016/s0197-4580(97)00059-6 · Cited by 4
What to Report: Information to Be Included in the Publication of a Stereological StudyPreprint
West, M. J. (2013) · Cold Spring Harbor Protocols · doi:10.1101/pdb.top071894 · Cited by 5
Self-citation
Untitled work
(2001) · doi:10.1002/1096-8644(200103)114:3
Volume and number of neurons of the human hippocampal formation in normal aging and Alzheimer’s disease
?imi?, G., Kostovi?, I., Winblad, B., & Bogdanovi?, N. (1997) · The Journal of Comparative Neurology · doi:10.1002/(sici)1096-9861(19970324)379:4
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Wang (1996) · doi:10.1002/(sici)1096-9861(19960318)366:4
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(1997) · doi:10.1002/(sici)1098-1063(1997)7:1
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Herman (2000) · doi:10.1002/1096-9861(20000703)422:3
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Kritzer (1997) · doi:10.1002/(sici)1096-9861(19970310)379:2
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(1999) · doi:10.1002/(sici)1098-1063(1999)9:5

Authors: West, M. J., & Gundersen, H. J. G.
Affiliations: Aarhus University (DK)
Corresponding author: West, Mark J.
ORCID: West, Mark J.
Venue: Journal of Comparative Neurology
Publisher: Wiley
Published: 1990-06
Volume: 296
Issue: 1
Pages: 1-22
ISSN: 0021-9967 (ISSN-L)
1096-9861
DOI: 10.1002/cne.902960102
OpenAlex: W2014892312
Cited by: 1,153
References: 47
Citations by year: 2026: 7 · 2025: 13 · 2024: 10 · 2023: 17 · 2022: 24 · 2021: 22 · 2020: 24 · 2019: 19 · 2018: 40 · 2017: 27 · 2016: 32 · 2015: 30 · 2014: 38 · 2013: 40 · 2012: 41
FWCI: 46.01
Type: journal article
Language: en
Open access: closed
Licence: onlinelibrary.wiley.com/termsAndConditions#vor (vor, 5244 d)
Locations: The Journal of Comparative Neurology (publishedVersion)
PubMed (publishedVersion)
Funders: Sundhed og Sygdom, Det Frie Forskningsråd
Topics: Physical Sciences › Mathematics › Applied Mathematics › Point processes and geometric inequalities
Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
Health Sciences › Medicine › Anatomy › Classical Anatomy and Histology
Keywords: human hippocampus, neuron number estimation, neuronal density, unbiased stereology, optical disector, point counts, CA1, CA3, subiculum, granular layer, hilus, age-related neuronal loss
MeSH terms: Aged, Aged, 80 and over, Cell Count, Hippocampus, Humans, Middle Aged

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Unbiased stereological estimation of the number of neurons in the human hippocampus

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