Cellular scaling rules for primate brains.
Open accessAbstract
Primates are usually found to have richer behavioral repertoires and better cognitive abilities than rodents of similar brain size. This finding raises the possibility that primate brains differ from rodent brains in their cellular composition. Here we examine the cellular scaling rules for primate brains and show that brain size increases approximately isometrically as a function of cell numbers, such that an 11x larger brain is built with 10x more neurons and approximately 12x more nonneuronal cells of relatively constant average size. This isometric function is in contrast to rodent brains, which increase faster in size than in numbers of neurons. As a consequence of the linear cellular scaling rules, primate brains have a larger number of neurons than rodent brains of similar size, presumably endowing them with greater computational power and cognitive abilities.
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Because EQ is sensitive to the species included and the human brain conforms to primate scaling rules, the authors suggested that great apes such as orangutans and gorillas have bodies much larger than expected for primates of their brain size, rather than humans having a larger brain than expected.
“we have suggested that, rather than humans having a larger brain than expected, it is the great apes such as orangutans and, more notably, gorillas that have bodies that are much larger than expected for primates of their brain size (Herculano-Houzel et al., 2007).”
From The human brain in numbers: a linearly scaled-up primate brain (2009) · cites this paper as Herculano-Houzel et al., 2007
Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measuredAbsolute brain mass and number of neurons must be considered because the exceeding number of neurons in a large brain should be larger than in a smaller brain of the same EQ.
“Absolute brain mass and number of neurons, left out of the encephalization equation, must clearly be taken into consideration, since the “exceeding number of neurons” in a large brain should necessarily be larger than that in a smaller brain of same EQ (Herculano-Houzel, 2007).”
From The human brain in numbers: a linearly scaled-up primate brain (2009) · cites this paper as Herculano-Houzel, 2007
The ‘Implications for Volume Comparisons Across Orders’ section says that comparisons of brain size should consider neuronal counts directly, not just brain volume. The paper also relates larger primate brain structures to greater numbers of neurons. It does not address encephalization quotient (EQ) or establish that a larger brain has more ‘exceeding’ neurons than a smaller brain with the same EQ; no page numbers are available in our copy. (LLM, conf=0.86)
Experimental — accuracy not yet measuredIn mammals, the pallium accounts for most of total brain volume, but the cerebellum houses a large majority of brain neurons.
“This markedly contrasts with the situation found in mammals, in which the pallium accounts for most of total brain volume, but the cerebellum houses a large majority of brain neurons (32–39)”
From Birds have primate-like numbers of neurons in the forebrain (2016) · cites this paper as [33]
The “Fractional Distribution of MBR and Neurons” section reports that the cerebral cortex makes up most brain mass while the cerebellum contains most brain neurons in the primates examined. The claim extends that finding to mammals generally, which this evidence does not establish. Page numbers are unavailable in our copy. (LLM, conf=0.86)
Experimental — accuracy not yet measuredPrimate brain size grows approximately isometrically with neuron number, with no systematic change in neuronal density or in the non-neuronal-to-neuronal ratio as brain size increases.
“primate brain size increases approximately isometrically as a function of neuron number, with no systematic change in neuronal density or in the non-neuronal/neuronal ratio with increasing brain size”
From The human brain in numbers: a linearly scaled-up primate brain (2009) · cites this paper as Herculano-Houzel et al., 2007
Like in mammals, the proportion of nonneuronal cells is very small in the cerebellum, varying between 12% and 19% in songbirds and between 14% and 19% in parrots, but, in contrast to mammals, nonneuronal cells also constitute a minor cellular fraction in the telencephalon, representing 21-40% of cells in songbirds and 31-43% of cells in parrots.
“Like in mammals (32–39, 46), the proportion of nonneuronal cells is very small in the cerebellum, varying between 12% and 19% in songbirds and between 14% and 19% in parrots, but, in contrast to mammals, nonneuronal cells also constitute a minor cellular fraction in the telencephalon, representing 21–40% of cells in songbirds and 31–43% of cells in parrots”
From Birds have primate-like numbers of neurons in the forebrain (2016) · cites this paper as [33]
Non-neuronal cell density is remarkably constant across these species.
“Given that non-neuronal cell density is remarkably constant across these species [24], [25]”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
There is no uniform scaling of neuronal density with brain size.
“However, it has now been demonstrated that there is no uniform scaling of neuronal density with brain size [24], [25], [41], which invalidates the previous estimate”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
Brain metabolism is not necessarily related to whole body metabolism in any determining way; any apparent relationship might be coincidental and dependent on the rate with which brain size scales as a function of its number of neurons.
“the scaling of brain metabolism as a function of total numbers of neurons opens the possibility that brain metabolism is not necessarily related to whole body metabolism in any determining way; any apparent relationship might be coincidental, and dependent on the rate with which brain size scales as a function of its number of neurons, which we have shown to vary across mammalian orders [24], [25]”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
Mammals whose brain scales with economical neuronal scaling rules, such as primates, have a large number of brain neurons for a given body size and would be expected to have a larger relative brain metabolic rate than other mammals, such as rodents, which have a smaller number of brain neurons for a same body or brain size.
“In this way, mammals whose brain scales with economical neuronal scaling rules, such as primates, have a large number of brain neurons for a given body size, and would accordingly be expected to have a larger relative brain metabolic rate than other mammals, such as rodents, which have a smaller number of brain neurons for a same body or brain size [24], [25].”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
The human brain's use of 20% of the whole body energy budget while representing only 2% of body mass can be explained by the large number of neurons in the human brain, about 10× larger than would be expected of a rodent brain of its size.
“According to the scaling of brain metabolism with its total number of neurons proposed above, the apparently remarkable use in humans of 20% of the whole body energy budget by a brain that represents only 2% of this mass can be explained quite simply by the large number of neurons in the human brain, about 10× larger than would be expected of a rodent brain of its size [44], given the different”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
The previous estimate that average energy use per neuron increases with brain size is invalidated.
“However, it has now been demonstrated that there is no uniform scaling of neuronal density with brain size [24], [25], [41], which invalidates the previous estimate”
From Scaling of Brain Metabolism with a Fixed Energy Budget per Neuron: Implications for Neuronal Activity, Plasticity and Evolution (2011) · cites this paper as [25]
The number of neurons in primate brains scales isometrically with brain size, indicating selection for constant neural density and neuron size, a scaling relationship that contrasts with other orders of animals.
“Second, the number of neurons in primate brains scales isometrically with brain size, indicating selection for constant neural density and neuron size, a scaling relationship that contrasts with other orders of animals (100).”
From The evolution of self-control. (2014) · cites this paper as [100]
The cortical reorganization hypothesis suggests that selection for absolutely larger brains, and concomitant cortical reorganization, was the predominant mechanism supporting cognitive evolution.
“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).”
From The evolution of self-control. (2014) · cites this paper as [100]
Brain (structure) size scales approximately isometrically as a function of cell number (and therefore computational power) in primates.
“Because brain (structure) size scales approximately isometrically as a function of cell number (and therefore computational power) in primates [24,25]”
From Brain reorganization, not relative brain size, primarily characterizes anthropoid brain evolution (2013) · cites this paper as [24]
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- Authors
- Herculano-Houzel, S., Collins, C. E., Wong, P., & Kaas, J. H.
- Affiliations
- *Departamento de Anatomia, Instituto de Ciências Biomédicas, Universidade Federal do Rio de Janeiro, 21941-590, Rio de Janeiro, Brazil
- Vanderbilt University (US)
- Corresponding author
- Kaas, Jon H.
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2007-02-27
- Volume
- 104
- Issue
- 9
- Pages
- 3562-3567
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC1805542
- OpenAlex
- W2033231119
- Cited by
- 422
- References
- 32
- Citations by year
- 2026: 17 · 2025: 19 · 2024: 15 · 2023: 24 · 2022: 18 · 2021: 22 · 2020: 57 · 2019: 17 · 2018: 15 · 2017: 28 · 2016: 30 · 2015: 19 · 2014: 33 · 2013: 18 · 2012: 23
- FWCI
- 21.97
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Eye Institute (R01EY002686, EY002686)
- National Institute of Neurological Disorders and Stroke (R01NS016446)
- Topics
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Life Sciences › Neuroscience › Developmental Neuroscience › Neurogenesis and neuroplasticity mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- primate brain, brain size scaling, neuron number, non-neuronal cells, isometric scaling, rodent brain, cognitive ability
- MeSH terms
- Animals, Brain, Indoles, Neurons, Organ Size, Primates, Species Specificity
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