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The remarkable, yet not extraordinary, human brain as a scaled-up primate brain and its associated cost
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Neuroscientists have become used to a number of “facts” about the human brain: It has 100 billion neurons and 10- to 50-fold more glial cells; it is the largest-than-expected for its body among primates and mammals in general, and therefore the most cognitively able; it consumes an outstanding 20% of the total body energy budget despite representing only 2% of body mass because of an increased metabolic need of its neurons; and it is endowed with an overdeveloped cerebral cortex, the largest compared with brain size. These facts led to the widespread notion that the human brain is literally extraordinary: an outlier among mammalian brains, defying evolutionary rules that apply to other species, with a uniqueness seemingly necessary to justify the superior cognitive abilities of humans over mammals with even larger brains. These facts, with deep implications for neurophysiology and evolutionary biology, are not grounded on solid evidence or sound assumptions, however. Our recent development of a method that allows rapid and reliable quantification of the numbers of cells that compose the whole brain has provided a means to verify these facts. Here, I review this recent evidence and argue that, with 86 billion neurons and just as many nonneuronal cells, the human brain is a scaled-up primate brain in its cellular composition and metabolic cost, with a relatively enlarged cerebral cortex that does not have a relatively larger number of brain neurons yet is remarkable in its cognitive abilities and metabolism simply because of its extremely large number of neurons.
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“Brain evolution may often be driven by adaptive changes in cellular composition or molecular expression within conserved structures4–6.”
From Innovations present in the primate interneuron repertoire (2020) · cites this paper as [5]
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“Our neural network parameters represent a computational analog to biological neural capacity, directly reflecting the studies showing strong correlations between brain volume and total neuron count across species, humans, and mice brains (68–70).”
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“During much of evolutionary history, when food scarcity and caloric deprivation was a major evolutionary pressure, mechanisms evolved to minimize muscle size and thus decrease non-essential energy expenditure42,44,55.”
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“minimizing muscle mass during caloric restriction can substantially reduce energy expenditure, thus sparing energy for the maintenance of other more critical organ systems42–44.”
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“Beyond identifiable causes, a large part of phenotype variability across patients with the same ABCD1 genotype may “simply” be due to stochasticity, a factor hitherto considered “noise” by Cartesian researchers.”
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“The cortical reorganization hypothesis suggests that selection for absolutely larger brains-and concomitant cortical reorganization-was the predominant mechanism supporting cognitive evolution (77, 91, 100-106, 120).”
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Citations
Cites
Collective dynamics of ‘small-world’ networks
Watts, D. J., & Strogatz, S. H. (1998) · Nature · doi:10.1038/30918 · Cited by 43,060
(1982) · Trends in Neurosciences · doi:10.1016/0166-2236(82)90076-5 · Cited by 9,150
An Energy Budget for Signaling in the Grey Matter of the Brain
Attwell, D., & Laughlin, S. B. (2001) · Journal of Cerebral Blood Flow & Metabolism · doi:10.1097/00004647-200110000-00001 · Cited by 3,543
Henry, M. A. (1995) · The Journal of the American Dental Association · doi:10.14219/jada.archive.1995.0108 · Cited by 3,454
Specification of Cerebral Cortical Areas
Rakic, P. (1988) · Science · doi:10.1126/science.3291116 · Cited by 3,127 · cited 1×
“Cortical expansion is commonly envisioned as occurring laterally, through the increase of the number of progenitor cells in the subventricular zone and the consequent addition of radial columns containing a constant number of neurons across species (38).”
BackgroundScaling: Why is Animal Size so Important?
Schmidt-Nielsen, K. (1984) · doi:10.1017/cbo9781139167826 · Cited by 2,677
Scaling Why is animal size so important?
Schmidt‐Nielsen, K. (1984) · Cited by 2,654
Azevedo, F. A. C., Carvalho, L. R. B., Grinberg, L. T., Farfel, J. M., Ferretti, R. E. L., Leite, R. E. P., Filho, W. J., Lent, R., & Herculano‐Houzel, S. (2009) · Journal of Comparative Neurology · doi:10.1002/cne.21974 · Cited by 2,337
Self-citationThe expensive-tissue hypothesis: The brain and the digestive system in human and primate evolution
Aiello, L. C., & Wheeler, P. (1995) · Current Anthropology · doi:10.1086/204350 · Cited by 2,291
Kleiber, M. (1932) · Hilgardia · doi:10.3733/hilg.v06n11p315 · Cited by 2,222
Research works printed but not linked: 1
- GC Conroy Primate Evolution (WW Norton, New York, 1990).
Cited by
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Variation in cancer risk among tissues can be explained by the number of stem cell divisions
Tomasetti, C., & Vogelstein, B. (2015) · Science · doi:10.1126/science.1260825 · Cited by 1,927
Integrated information theory: from consciousness to its physical substrate
Tononi, G., Boly, M., Massimini, M., & Koch, C. (2016) · Nature Reviews Neuroscience · doi:10.1038/nrn.2016.44 · Cited by 1,727
Neural correlates of consciousness: progress and problems
Koch, C., Massimini, M., Boly, M., & Tononi, G. (2016) · Nature Reviews Neuroscience · doi:10.1038/nrn.2016.22 · Cited by 1,600
The cerebellum and cognitive function: 25 years of insight from anatomy and neuroimaging
Buckner, R. L. (2013) · Neuron · doi:10.1016/j.neuron.2013.10.044 · Cited by 1,173
von Bartheld, C. S., Bahney, J., & Herculano‐Houzel, S. (2016) · Journal of Comparative Neurology · doi:10.1002/cne.24040 · Cited by 1,063
Self-citationThe evolution of distributed association networks in the human brain
Buckner, R. L., & Krienen, F. M. (2013) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2013.09.017 · Cited by 836
BigBrain: An Ultrahigh-Resolution 3D Human Brain Model
Amunts, K., Lepage, C., Borgeat, L., Mohlberg, H., Dickscheid, T., Rousseau, M.-É., Bludau, S., Bazin, P., Lewis, L. B., Oros‐Peusquens, A., et al. (2013) · Science · doi:10.1126/science.1235381 · Cited by 884
The evolution of self-control.
MacLean, E. L., Hare, B., Nunn, C. L., Addessi, E., Amici, F., Anderson, R. C., Aureli, F., Baker, J. M., Bania, A. E., Barnard, A. M., et al. (2014) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1323533111 · Cited by 819
Consciousness: here, there and everywhere?
Tononi, G., & Koch, C. (2015) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2014.0167 · Cited by 811
Neural progenitors, neurogenesis and the evolution of the neocortex
Florio, M., & Huttner, W. Β. (2014) · Development · doi:10.1242/dev.090571 · Cited by 654
Establishing Cerebral Organoids as Models of Human-Specific Brain Evolution
Pollen, A. A., Bhaduri, A., Andrews, M. G., Nowakowski, T. J., Meyerson, O. S., Mostajo-Radji, M. A., Di Lullo, E., Alvarado, B., Bedolli, M., Dougherty, M. L., et al. (2019) · Cell · doi:10.1016/j.cell.2019.01.017 · Cited by 600
Microglial regional heterogeneity and its role in the brain
Tan, Y.-L., Yuan, Y., & Tian, L. (2020) · Molecular Psychiatry · doi:10.1038/s41380-019-0609-8 · Cited by 571
Survival of the Friendliest: Homo sapiens Evolved via Selection for Prosociality
Hare, B. (2017) · Annual Review of Psychology · doi:10.1146/annurev-psych-010416-044201 · Cited by 444
Innovations present in the primate interneuron repertoire
Krienen, F. M., Goldman, M., Zhang, Q., C. H. del Rosario, R., Florio, M., Machold, R., Saunders, A., Levandowski, K., Zaniewski, H., Schuman, B., et al. (2020) · Nature · doi:10.1038/s41586-020-2781-z · Cited by 381
Otani, T., Marchetto, M. C., Gage, F. H., Simons, B. D., & Livesey, F. J. (2016) · Cell Stem Cell · doi:10.1016/j.stem.2016.03.003 · Cited by 353
Innovations in Primate Interneuron Repertoire
Krienen, F. M., Goldman, M., Zhang, Q., del Rosario, R., Florio, M., Machold, R., Saunders, A., Levandowski, K., Zaniewski, H., Schuman, B., et al. (2019) · doi:10.1101/709501 · Cited by 372
Pietschnig, J., Penke, L., Wicherts, J. M., Zeiler, M., & Voracek, M. (2015) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2015.09.017 · Cited by 316
The cerebellum in Alzheimer’s disease: evaluating its role in cognitive decline
Jacobs, H. I. L., Hopkins, D. A., Mayrhofer, H. C., Bruner, E., van Leeuwen, F. W., Raaijmakers, W., & Schmahmann, J. D. (2017) · Brain · doi:10.1093/brain/awx194 · Cited by 349
Childhood as a solution to explore–exploit tensions
Alison Gopnik, C. (2020) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2019.0502 · Cited by 354
Emerging connections between cerebellar development, behaviour and complex brain disorders
Sathyanesan, A., Zhou, J., Scafidi, J., Heck, D. H., Sillitoe, R. V., & Gallo, V. (2019) · Nature reviews. Neuroscience · doi:10.1038/s41583-019-0152-2 · Cited by 341
Kong, X., Yu, F., Yao, W., Cai, S., Zhang, J., & Lin, H. (2023) · Neural Networks · doi:10.1016/j.neunet.2023.12.008 · Cited by 292
Boyd, J. L., Skove, S. L., Rouanet, J. P., Pilaz, L., Bepler, T., Gordân, R., Wray, G. A., & Silver, D. L. (2015) · Current Biology · doi:10.1016/j.cub.2015.01.041 · Cited by 289
Organoid intelligence (OI): the new frontier in biocomputing and intelligence-in-a-dish
Smirnova, L., Caffo, B. S., Gracias, D. H., Huang, Q., Pantoja, I. E. M., Tang, B., Zack, D. J., Berlinicke, C. A., Boyd, J. L., Harris, T. D., et al. (2023) · Frontiers in Science · doi:10.3389/fsci.2023.1017235 · Cited by 329
An early cell shape transition drives evolutionary expansion of the human forebrain
Benito-Kwiecinski, S. K., Giandomenico, S. L., Sutcliffe, M. A., Riis, E. S., Freire-Pritchett, P., Kelava, I., Wunderlich, S., Martin, U., Wray, G. A., McDole, K., et al. (2021) · Cell · doi:10.1016/j.cell.2021.02.050 · Cited by 272
Fedorenko, E., & Varley, R. (2016) · Annals of the New York Academy of Sciences · doi:10.1111/nyas.13046 · Cited by 247
Species-specific pace of development is associated with differences in protein stability
Rayón, T., Stamataki, D., Perez‐Carrasco, R., Garcia-Perez, L., Barrington, C., Melchionda, M., Exelby, K., Lázaro, J., Tybulewicz, V. L. J., Fisher, E., et al. (2020) · Science · doi:10.1126/science.aba7667 · Cited by 269
Glial Cells as Progenitors and Stem Cells: New Roles in the Healthy and Diseased Brain
Dimou, L., & Götz, M. (2014) · Physiological Reviews · doi:10.1152/physrev.00036.2013 · Cited by 285
Mechanisms of astrocyte development and their contributions to neurodevelopmental disorders
Sloan, S. A., & Barres, B. A. (2014) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2014.03.005 · Cited by 260
Human brain evolution: transcripts, metabolites and their regulators
Somel, M., Liu, X., & Khaitovich, P. (2013) · Nature reviews. Neuroscience · doi:10.1038/nrn3372 · Cited by 246
Human TKTL1 implies greater neurogenesis in frontal neocortex of modern humans than Neanderthals
Pinson, A., Xing, L., Namba, T., Kalebic, N., Peters, J., Oegema, C. E., Traikov, S., Reppe, K., Riesenberg, S., Maričić, T., et al. (2022) · Science · doi:10.1126/science.abl6422 · Cited by 236
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Cox, D., & Dean, T. (2014) · Current Biology · doi:10.1016/j.cub.2014.08.026 · Cited by 237
RNA-binding proteins in neurodegeneration: mechanisms in aggregatePreprint
Conlon, E. G., & Manley, J. L. (2017) · Genes & Development · doi:10.1101/gad.304055.117 · Cited by 229
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Filippi, M., van den Heuvel, M. P., Fornito, A., Yong He, A., Pol, H. E. H., Agosta, F., Comı, G., & Rocca, M. A. (2013) · The Lancet Neurology · doi:10.1016/s1474-4422(13)70144-3 · Cited by 219
Mouse and human share conserved transcriptional programs for interneuron development
Shi, Y., Wang, M., Mi, D., Lu, T., Wang, B., Dong, H., Zhong, S., Chen, Y., Sun, L., Zhou, X., et al. (2021) · Science · doi:10.1126/science.abj6641 · Cited by 196
A Survey of Robotics Control Based on Learning-Inspired Spiking Neural Networks
Bing, Z., Meschede, C., Röhrbein, F., Huang, K., & Knoll, A. (2018) · Frontiers in Neurorobotics · doi:10.3389/fnbot.2018.00035 · Cited by 197
Rossi, D. (2015) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2015.04.003 · Cited by 185
Brain Plasticity and Human Evolution
Sherwood, C. C., & Gómez-Robles, A. (2017) · Annual Review of Anthropology · doi:10.1146/annurev-anthro-102215-100009 · Cited by 170
Developmental mechanisms underlying the evolution of human cortical circuits
Vanderhaeghen, P., & Polleux, F. (2023) · Nature Reviews Neuroscience · doi:10.1038/s41583-023-00675-z · Cited by 173
Cognitive Ecology of Food Hoarding: The Evolution of Spatial Memory and the Hippocampus
Витальевич, П. В., & Roth, T. C. (2013) · Annual Review of Ecology Evolution and Systematics · doi:10.1146/annurev-ecolsys-110512-135904 · Cited by 181
Consciousness and complexity: a consilience of evidence
Sarasso, S., Casali, A. G., Casarotto, S., Rosanova, M., Sinigaglia, C., & Massimini, M. (2021) · Neuroscience of Consciousness · doi:10.1093/nc/niab023 · Cited by 171
The Neurology of Consciousness
Tononi, G., Boly, M., Gosseries, O., & Laureys, S. (2015) · Elsevier eBooks · doi:10.1016/b978-0-12-800948-2.00025-x · Cited by 178
Language is primarily a tool for communication rather than thought
Fedorenko, E., Piantadosi, S. T., & Gibson, E. A. F. (2024) · Nature · doi:10.1038/s41586-024-07522-w · Cited by 190
Towards a comparative science of emotion: Affect and consciousness in humans and animals
Paul, E. S., Sher, S., Tamietto, M., Winkielman, P., & Mendl, M. (2019) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2019.11.014 · Cited by 180
Herculano‐Houzel, S., WATSON, C. R. R., & Paxinos, G. (2013) · Frontiers in Neuroanatomy · doi:10.3389/fnana.2013.00035 · Cited by 164
Self-citationToxoplasma gondii: Entry, association, and physiological influence on the central nervous system
Mendez, O. A., & Koshy, A. A. (2017) · PLoS Pathogens · doi:10.1371/journal.ppat.1006351 · Cited by 169
Sexual Dimorphism in the Human Olfactory Bulb: Females Have More Neurons and Glial Cells than Males
de Oliveira-Pinto, A. V., Santos, R. M., Coutinho, R., Oliveira, L. M., dos Santos, G. B., Alho, A. T. D. L., Leite, R. E. P., Farfel, J. M., Suemoto, C. K., Grinberg, L. T., et al. (2014) · PLoS ONE · doi:10.1371/journal.pone.0111733 · Cited by 169
Glia Crosstalk in Neuroinflammatory Diseases
Bernaus, A., Blanco, S., & Sevilla, A. (2020) · Frontiers in Cellular Neuroscience · doi:10.3389/fncel.2020.00209 · Cited by 157
Herculano-Houzel, S., Avelino-de-Souza, K., Neves, K., PorfÃrio, J., Messeder, D., Mattos Feijó, L., Maldonado, J., & Manger, P. R. (2014) · Frontiers in Neuroanatomy · doi:10.3389/fnana.2014.00046 · Cited by 153
Self-citationGeneration of neuropeptidergic hypothalamic neurons from human pluripotent stem cells
Merkle, F. T., Maroof, A. M., Wataya, T., Sasai, Y., Studer, L., Eggan, K. C., & Schier, A. F. (2015) · Development · doi:10.1242/dev.117978 · Cited by 151
van Albada, S. J., Rowley, A. G. D., Senk, J., Hopkins, M., Schmidt, M., Stokes, A. B., Lester, D., Diesmann, M., & Furber, S. B. (2018) · Frontiers in Neuroscience · doi:10.3389/fnins.2018.00291 · Cited by 157
Gowlett, J. A. J., & Wrangham, R. W. (2013) · Azania Archaeological Research in Africa · doi:10.1080/0067270x.2012.756754 · Cited by 154
Is the brain really a small-world network?
Hilgetag, C. C., & Goulas, A. (2015) · Brain Structure and Function · doi:10.1007/s00429-015-1035-6 · Cited by 144
Abdelfattah, A. S., Farhi, S. L., Zhao, Y., Brinks, D., Zou, P., Ruangkittisakul, A., Platiša, J., Pieribone, V. A., Ballanyi, K., Cohen, A. E., et al. (2016) · Journal of Neuroscience · doi:10.1523/jneurosci.3484-15.2016 · Cited by 143
Kabadayi, C., Taylor, L. A., von Bayern, A. M. P., & Osvath, M. (2016) · Royal Society Open Science · doi:10.1098/rsos.160104 · Cited by 152
Cerebellar Neurodynamics Predict Decision Timing and Outcome on the Single-Trial Level
Lin, Q., Manley, J., Helmreich, M., Schlumm, F., Li, J. M., Robson, D. N., Engert, F., Schier, A., Nöbauer, T., & Vaziri, A. (2020) · Cell · doi:10.1016/j.cell.2019.12.018 · Cited by 122
Fast Approximate Quadratic Programming for Graph Matching
Vogelstein, J. T., Conroy, J. M., Lyzinski, V., Podrazik, L. J., Kratzer, S. G., Harley, E. T., Fishkind, D. E., Vogelstein, R. J., & Priebe, C. E. (2015) · PLoS ONE · doi:10.1371/journal.pone.0121002 · Cited by 167
Zeng, B., Liu, Z., Lu, Y., Zhong, S., Qin, S., Huang, L., Zeng, Y., Li, Z., Dong, H., Shi, Y., et al. (2023) · Cell stem cell · doi:10.1016/j.stem.2023.04.016 · Cited by 142
Genes, Cells and Brain Areas of Intelligence
Goriounova, N. A., & Mansvelder, H. D. (2019) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2019.00044 · Cited by 148
Unraveling the evolution of uniquely human cognition
MacLean, E. L. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1521270113 · Cited by 124
Beyond brain size: Uncovering the neural correlates of behavioral and cognitive specialization
Logan, C., Avin, S., Boogert, N. J., Buskell, A., Cross, F. R., Currie, A., Jelbert, S. A., Lukas, D., Marés, R., Navarrete, A. F., et al. (2018) · Comparative Cognition & Behavior Reviews · doi:10.3819/ccbr.2018.130008 · Cited by 125
The number of neurons in Drosophila and mosquito brains
Raji, J. I., & Potter, C. J. (2021) · PLoS ONE · doi:10.1371/journal.pone.0250381 · Cited by 123
The pathogenesis of cerebral small vessel disease and vascular cognitive impairment
Markus, H. S., & Joutel, A. (2025) · Physiological Reviews · doi:10.1152/physrev.00028.2024 · Cited by 140
Maiti, P., Manna, J., Ilavazhagan, G., Rossignol, J., & Dunbar, G. L. (2015) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2015.09.020 · Cited by 128
Evolution of cortical neurons supporting human cognition
Galakhova, A. A., Hunt, S., Wilbers, R., Heyer, D. B., de Kock, C. P. J., Mansvelder, H. D., & Goriounova, N. A. (2022) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2022.08.012 · Cited by 127
Probing human brain evolution and development in organoids
Giandomenico, S. L., & Lancaster, M. A. (2017) · Current Opinion in Cell Biology · doi:10.1016/j.ceb.2017.01.001 · Cited by 111
Mechanisms for the maintenance and regulation of axonal energy supply
Chamberlain, K. A., & Sheng, Z. (2019) · Journal of Neuroscience Research · doi:10.1002/jnr.24411 · Cited by 111
Fragility and volatility of structural hubs in the human connectome
Gollo, L. L., Roberts, J. A., Cropley, V., Biase, M. A. D., Pantelis, C., Zalesky, A., & Breakspear, M. J. (2018) · Nature Neuroscience · doi:10.1038/s41593-018-0188-z · Cited by 105
An energy costly architecture of neuromodulators for human brain evolution and cognition
Castrillón, G., Epp, S. M., Bose, A., Fraticelli, L., Hechler, A., Belenya, R., Ranft, A., Yakushev, I. Y., Utz, L., Sundar, L. K. S., et al. (2023) · Science Advances · doi:10.1126/sciadv.adi7632 · Cited by 102
Electronic and Ionic Materials for Neurointerfaces
Ferro, M. D., & Melosh, N. A. (2017) · Advanced Functional Materials · doi:10.1002/adfm.201704335 · Cited by 96
Frontoparietal Tracts Linked to Lateralized Hand Preference and Manual Specialization
Howells, H., de Schotten, M. T., Dell’Acqua, F., Beyh, A., Zappalà, G., Leslie, A., Simmons, A., Murphy, D., & Catani, M. (2018) · Cerebral Cortex · doi:10.1093/cercor/bhy040 · Cited by 96
Pandora's growing box: Inferring the evolution and development of hominin brains from endocasts
Zollikofer, C. P. E., & León, M. S. P. D. (2013) · Evolutionary Anthropology Issues News and Reviews · doi:10.1002/evan.21333 · Cited by 94
Bozek, K., Wei, Y., Yan, Z., Liu, X., Xiong, J., Sugimoto, M., Tomita, M., Pääbo, S., Pieszek, R., Sherwood, C. C., et al. (2014) · PLoS Biology · doi:10.1371/journal.pbio.1001871 · Cited by 93
Human evolution: the non-coding revolution
Franchini, L. F., & Pollard, K. S. (2017) · BMC Biology · doi:10.1186/s12915-017-0428-9 · Cited by 90
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Zhao, L., Zhang, L., Wu, Z., Chen, Y., Dai, H., Yu, X., Liu, Z., Zhang, T., Hu, X., Jiang, X., et al. (2023) · Meta-Radiology · doi:10.1016/j.metrad.2023.100005 · Cited by 116
Relating enhancer genetic variation across mammals to complex phenotypes using machine learning
Kaplow, I. M., Lawler, A. J., Schäffer, D. E., Srinivasan, C., Sestili, H. H., Wirthlin, M. E., Phan, B. N., Prasad, K., Brown, A. R., Zhang, X., et al. (2023) · Science · doi:10.1126/science.abm7993 · Cited by 97
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Krämer, P., & Bressan, P. (2017) · Perspectives on Psychological Science · doi:10.1177/1745691617718356 · Cited by 94
Ribeiro, P. F. M., Ventura-Antunes, L., Gabi, M., Mota, B., Grinberg, L. T., Farfel, J. M., Ferretti-Rebustini, R. E. L., Leite, R. E. P., Filho, W. J., & Herculano-Houzel, S. (2013) · Frontiers in Neuroanatomy · doi:10.3389/fnana.2013.00028 · Cited by 89
Self-citationMuench, N. A., Patel, S., Maes, M. E., Donahue, R., Ikeda, A., & Nickells, R. W. (2021) · Cells · doi:10.3390/cells10071593 · Cited by 93
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Jurcău, M. C., Jurcău, A., Cristian, A., Hogea, V. O., Diaconu, R. G., & Nunkoo, V. S. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms251910535 · Cited by 100
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Bailey, D. M. (2018) · The Journal of Physiology · doi:10.1113/jp276814 · Cited by 78
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Metadata
- Authors
- Herculano-Houzel, S.
- Affiliations
- Instituto de Ciências Biomédicas, Universidade Federal do Rio de Janeiro, 21941-902, Rio de Janeiro, Brazil; and Instituto Nacional de Neurociência Translacional, Instituto Nacional de Ciência e Tecnologia/Ministério de Ciência e Tecnologia, 04023-900, Sao Paulo, Brazil
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2012-06-22
- Volume
- 109
- Issue
- supplement_1
- Pages
- 10661-10668
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC3386878
- OpenAlex
- W1966084915
- Semantic Scholar
- 65f9d025887298541bab310b071635e4588d2b68
- Cited by
- 726
- References
- 100
- Citations by year
- 2026: 37 · 2025: 69 · 2024: 93 · 2023: 73 · 2022: 58 · 2021: 60 · 2020: 89 · 2019: 51 · 2018: 36 · 2017: 30 · 2016: 38 · 2015: 38 · 2014: 30 · 2013: 19 · 2012: 4
- FWCI
- 4.12
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Primate Behavior and Ecology
- Life Sciences › Neuroscience › Developmental Neuroscience › Neurogenesis and neuroplasticity mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Keywords
- human brain, primate brain, brain scaling, brain size, brain energy metabolism, brain energy consumption, cerebral cortex, neuron number, glial cells, brain evolution, cognitive ability
- MeSH terms
- Animals, Brain, Biological Evolution, Humans, Nerve Net, Neuroglia, Neurons, Primates
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