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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Background
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Method
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BackgroundMethod
Developmental changes of synaptic proteins expression within the hippocampal formation of the rat
Biranowska, J., Dziewíatkowski, J., Ludkiewicz, B., & Moryś, J. (2002) · Anatomy and Embryology · doi:10.1007/s00429-002-0277-2 · Cited by 14
Intact hippocampal gray matter in schizophrenia as revealed by automatized image analysis postmortem
Hurlemann, R., Tepest, R., Maier, W., Falkai, P., & Vogeley, K. T. (2005) · Anatomy and Embryology · doi:10.1007/s00429-005-0035-3 · Cited by 14
“According to cytoarchitectonic criteria ( West and Gundersen 1990; Arnold 2000), we manually delineated the outlines of (i) the pyramidal cell layer in each ammonic segment (CA1–4) and (ii) the granule cell layer (DG).”
Method
?-Amyloid precursor protein isoforms show correlations with neurones but not with glia of demented subjects
PROCTER, A. W., Francis, P. T., Holmes, C., Webster, M.-T., Qume, M., Stratmann, G. C., Doshi, R. N., Mann, D. M. A., Harrison, P. J., Pearson, R. C. A., et al. (1994) · Acta Neuropathologica · doi:10.1007/bf00296491 · Cited by 14
Stereological evaluation of treatment response in patients with non‐resectable hepatic alveolar echinococcosis using computed tomography via the Cavalieri method
Aydınlı, B., Kantarcı, M., Polat, K. Y., Ünal, B., Atamanalp, S. S., Durur, I., Ünal, D., & Akgün, M. (2006) · Liver International · doi:10.1111/j.1478-3231.2006.01363.x · Cited by 13
Prenatal Alcohol Exposure Affects Progenitor Cell Numbers in Olfactory Bulbs and Dentate Gyrus of Vervet Monkeys
Burke, M., Inyatkin, A., Ptito, M., Ervin, F. R., & Palmour, R. M. (2016) · Brain Sciences · doi:10.3390/brainsci6040052 · Cited by 13
“where ssf is section sampling fraction, asf is area sampling fraction, tsf is the thickness-sampling fraction and ∑Q− is the number of cells counted within the dissection [63].”
Background
The flavonoid 7,8-DHF fosters prenatal brain proliferation potency in a mouse model of Down syndrome
Stagni, F., Uguagliati, B., Emili, M., Giacomini, A., Bartesaghi, R., & Guidi, S. (2021) · Scientific Reports · doi:10.1038/s41598-021-85284-5 · Cited by 13
Cell to network computational model of the epileptic human hippocampus suggests specific roles of network and channel dysfunctions in the ictal and interictal oscillations
Aussel, A., Ranta, R., Aron, O., Colnat‐Coulbois, S., Maillard, L., & Buhry, L. (2022) · Journal of Computational Neuroscience · doi:10.1007/s10827-022-00829-5 · Cited by 13
“The number of neurons in each region and the proportion of excitatory and inhibitory cells were chosen in accordance with the literature ([Jinno and Kosaka, 2010], [West and Gundersen, 1990], [Patton and McNaughton, 1995], see [Aussel et al., 2018] for a detailed explanation).”
Memantine protects cholinergic and glutamatergic septal neurons from Aβ1–40-induced toxicity
Colom, L. V., Castaneda, M. T., Aleman, D. O., & Touhami, A. (2013) · Neuroscience Letters · doi:10.1016/j.neulet.2013.02.010 · Cited by 13
“Neuronal numbers were estimated using stereological approaches (StereoInvestigator software, MicroBrightField, Williston, VT, USA) [24].”
Growth patterns of human ovarian volume during intrauterine and postnatal organogenesis
Sforza, C., Ranzi, A., Ferrario, V. F., & Forabosco, A. (2004) · Early Human Development · doi:10.1016/j.earlhumdev.2004.05.005 · Cited by 13
“Indeed, both 3D US and the Cavalieri assessment measure the areas of multiple parallel sections at different distances along an axis of the organ [7,33,36].”
BackgroundMethod
4‐hydroxy tempo improves mitochondrial and neurobehavioral deficits in experimental model of H untington's disease
Sandhir, R., Mahajan, N., Mehrotra, A., Aggarwal, A., & Sunkaria, A. (2014) · Synapse · doi:10.1002/syn.21793 · Cited by 13
An isomorphic three‐dimensional cortical model of the pig rostrum
Ritter, C., Maier, E., Schneeweiß, U., Wölk, T., Simonnet, J., Malkawi, S., Eigen, L., Tunçkol, E., Purkart, L., & Brecht, M. (2020) · The Journal of Comparative Neurology · doi:10.1002/cne.25073 · Cited by 13
“Using the optical fractionator technique, we counted the nucleoli that come into focus and fall within the acceptance lines of the dissector, which were randomly placed on the series of sections (Howard & Reed, 2005; Naumann et al., 2012; West & Gundersen, 1990).”
Method
Immunocytochemical and stereological analysis of GABAB receptor subunit expression in the rat vestibular nucleus following unilateral vestibular deafferentation
Zhang, R., Ashton, J. C., Horii, A., Darlington, C. L., & Smith, P. F. (2005) · Brain Research · doi:10.1016/j.brainres.2005.01.018 · Cited by 13
The Weaver GIRK2 Mutation Leads to Decreased Levels of Serum Thyroid Hormone: Characterization of the Effect on Midbrain Dopaminergic Neuron Survival
Blum, M., Weickert, C. S., & Carrasco, E. (1999) · Experimental Neurology · doi:10.1006/exnr.1999.7231 · Cited by 13
Birthdates and Number of Neurons in the Serotonergic Raphe Nuclei in the Rat with Prenatal Protein Malnutrition
King, R. S., Kemper, T. L., DeBassio, W. A., Ramzan, M., Blatt, G. J., Rosene, D. L., & Galler, J. R. (2002) · Nutritional Neuroscience · doi:10.1080/1028415021000055934 · Cited by 13
“Cell counts were made using the optical fractionator method of West and Gundersen (1990) with a Bioquant TCW 3.12 (R and M Biometrics Inc.) image analysis system.”
Influential
Glial and neuronal cell numbers and cytochrome oxidase activity in CA1 and CA3 during postnatal development and aging of the rat
Cimadevilla, J. M., Moreno, L. M. G., González‐Pardo, H., Zahonero, M. C., & Árias, J. (1997) · Mechanisms of Ageing and Development · doi:10.1016/s0047-6374(97)00090-0 · Cited by 13
Maze learning and morphology of frontal cortex in adult and aged basal forebrain-lesioned rats.
Wellman, C. L., Logue, S. F., & Sengelaub, D. R. (1995) · Behavioral Neuroscience · doi:10.1037//0735-7044.109.5.837 · Cited by 13
“An unbiased stereological technique (e.g., Coggeshall, 1992; West, 1993; West & Gundersen, 1990 ) was used to obtain estimates of total neuron numbers in each lamina of frontal cortex.”
Age-related alterations of Nestin-immunoreactive neurons in rat basal forebrain with aged memory deficit
Li, D., Wang, J., Yew, D. T., Lucy Forster, E., & Yao, Z. (2008) · Neurochemistry International · doi:10.1016/j.neuint.2008.08.006 · Cited by 12
“The total number of Nestin-IR neurons within the MS, vDB and hDB were then estimated by combining the Nv with the Vref to give absolute numbers as described previously (West and Gundersen, 1990; Peterson et al., 1999).”
Cerebrocerebellar Loops in the Rodent Brain
Léna, C., & Popa, D. (2016) · Elsevier eBooks · doi:10.1016/b978-0-12-801386-1.00006-x · Cited by 12
The amygdala of the common shrew, guinea pig, rabbit, fox and pig: five flavours of the mammalian amygdala as a consequence of clade‐specific mosaic‐like evolution
Równiak, M., & Bogus‐Nowakowska, K. (2020) · Journal of Anatomy · doi:10.1111/joa.13148 · Cited by 11
“The numerical density of cells in a single amygdaloid nucleus was estimated using the optical dissector method described in detail by West and Gundersen (1990).”
MethodInfluential
Age-related differences in oligodendrogenesis across the dorsal-ventral axis of the mouse hippocampus
Yamada, J., & Jinno, S. (2014) · Hippocampus · doi:10.1002/hipo.22287 · Cited by 12
NMDA and kainate receptor expression, long-term potentiation, and neurogenesis in the hippocampus of long-lived Ames dwarf mice
Sharma, S., Darland, D. C., Lei, S., Rakoczy, S. G., & Brown‐Borg, H. M. (2011) · AGE · doi:10.1007/s11357-011-9253-1 · Cited by 12
“Non-biased quantification of DCX-positive nuclei was performed using the optical fractionator method as originally described by West and Gundersen (1990) and adapted from previous studies (Hattiangady et al.”
Method
Effects of Umbilical Cord Occlusion in Late Gestation on the Ovine Fetal Brain and Retina
Duncan, J. R., Camm, E. J., Loeliger, M. M., Cock, M. L., Harding, R., & Rees, S. M. (2004) · Journal of the Society for Gynecologic Investigation · doi:10.1016/j.jsgi.2004.03.004 · Cited by 12
Performance of semi-automated hippocampal subfield segmentation methods across ages in a pediatric sample
Schlichting, M. L., Mack, M. L., Guarino, K. F., & Preston, A. R. (2016) · NeuroImage · doi:10.1016/j.neuroimage.2019.01.051 · Cited by 12
“Importantly, these segmentations are all derived from a single tracing protocol; manual delineation was carried out by a single rater referencing histological work and printed atlases of human hippocampus (Duvernoy, 1998; Mai et al., 2007; West and Gundersen, 1990).”
Gender biased neuroprotective effect of Transferrin Receptor 2 deletion in multiple models of Parkinson’s disease
Milanese, C., Gabriels, S., Barnhoorn, S., Cerri, S., Ulusoy, A., Gornati, S. V., Wallace, D. F., Blandini, F., Monte, D. A. D., Subramaniam, V. N., et al. (2020) · Cell Death and Differentiation · doi:10.1038/s41418-020-00698-4 · Cited by 14
Memory and neurogenesis in aging and Alzheimer's disease.
Ávila, J., Insausti, R., & Rı́o, J. D. (2010) · Europe PMC (PubMed Central) · Cited by 12
Subchronic toluene exposure in low concentrations produces signs of reduced dysfunction in the 6-hydroxydopamine lesioned nigrostriatal dopaminergic system of the rat
Cintra, A., Aguirre, J. A., Andbjer, B., Finnman, U.-B., Hagman, M., Agnati, L. F., Höglund, C., Møller, A., & Fuxé, K. (1999) · Neuroscience Letters · doi:10.1016/s0304-3940(99)00112-3 · Cited by 12
Effect of hypothyroidism on vasoactive intestinal polypeptide-immunoreactive neurons in forebrain-neurohypophysial nuclei of the rat brain
Toni, R., Mosca, S., Ruggeri, F., Valmori, A., Orlandi, G., Toni, G., Lechan, R. M., & Vezzadini, P. (1995) · Brain Research · doi:10.1016/0006-8993(95)00340-v · Cited by 12
Efficient and unbiased estimation of volume and area of tissue components and cell number in gingival biopsies
Verwer, R. W. H., & Raber‐Durlacher, J. E. (1993) · Journal of Periodontal Research · doi:10.1111/j.1600-0765.1993.tb01075.x · Cited by 12
Changes in cell cycle parameters and cell number in the rat midbrain during organogenesis
Lewandowski, T. A., Ponce, R. A., Charleston, J. S., Hong, S., & Faustman, E. M. (2003) · Developmental Brain Research · doi:10.1016/s0165-3806(03)00004-x · Cited by 12
Clioquinol Protects Against Cell Death in Parkinson’s Disease Models In Vivo and In Vitro
Wilkins, S., Masters, C. L., Bush, A. I., Cherny, R. A., & Finkelstein, D. I. (2009) · Advances in behavioral biology · doi:10.1007/978-1-4419-0340-2_33 · Cited by 11
Selective brain neuronal and glial losses without changes in GFAP immunoreactivity: Young versus mature adult Wistar rats
Diene, L. D., Costa‐Ferro, Z. S. M., Barbosa, S., Milanesi, B. B., Lazzari, G. Z., Neves, L. T., Paz, L. V., Neves, P. F. R., Battisti, V., Martins, L. A., et al. (2019) · Mechanisms of Ageing and Development · doi:10.1016/j.mad.2019.111128 · Cited by 12
Quantified Assessment of Terminal Density and Innervation
Finkelstein, D. I., Stanić, D., Parish, C. L., Drago, J., & Horne, M. K. (2004) · Current Protocols in Neuroscience · doi:10.1002/0471142301.ns0113s27 · Cited by 11
Estudo estereológico das células de Purkinje cerebelares submetidas à intoxicação alcoólica em ratos Wistar
Apfel, M. Í. R., Esbérard, C. A., Rodrigues, F. K. P., Júnior, F. M. B., & Sillero, R. O. (2002) · Arquivos de Neuro-Psiquiatria · doi:10.1590/s0004-282x2002000200014 · Cited by 11
Identification of neuronal loci involved with displays of affective aggression in NC900 mice
Nehrenberg, D. L., Sheikh, A., & Ghashghaei, H. T. (2012) · Brain Structure and Function · doi:10.1007/s00429-012-0445-y · Cited by 12
“Counts were obtained in 50 9 50 lm counting frames (dissectors) with standard stereological inclusion and exclusion borders (West and Gundersen 1990).”
Method
APPLICATION OF STEREOLOGICAL METHODS TO STUDY THE WHITE MATTER AND MYELINATED FIBERS THEREIN OF RAT BRAIN
Shu, Y., Li, C., Zhang, W., Wang, W., Nyengaard, J. R., & Tang, Y. C. (2011) · Image Analysis & Stereology · doi:10.5566/ias.v27.p125-132 · Cited by 11
The evolutionary trajectories of the individual amygdala nuclei in the common shrew, guinea pig, rabbit, fox and pig: A consequence of embryological fate and mosaic‐like evolution
Równiak, M., Bogus‐Nowakowska, K., Kalinowski, D., & Kozłowska, A. (2021) · Journal of Anatomy · doi:10.1111/joa.13571 · Cited by 11
“2.3.3 | Estimation of neuron number The total number of neurons in a single nucleus was calculated by multiplying estimates of the nuclear volume by the numerical density in it (West & Gundersen, 1990).”
MethodInfluential
Pathological Nuclear Hallmarks in Dentate Granule Cells of Alzheimer’s Patients: A Biphasic Regulation of Neurogenesis
Gil, L., Chi-Ahumada, É., Niño, S. A., Capdeville, G., Méndez-Torres, A. M., Guerrero, C., Poves, A. B. R., Olazabal, I. M., & Jiménez-Capdeville, M. E. (2022) · International Journal of Molecular Sciences · doi:10.3390/ijms232112873 · Cited by 10
“This stabilization would compensate for the decrease in the constitutive heterochromatin marker H3K9me3 [139–141], which is fundamental for the structural organization of NADs and LADs [142] and genomic integrity [143].”
Background

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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