The DOI matches its Crossref record; 27 of the 28 references it cites were found in a registry or an index.
Brain power
Open accessAbstract
The human brain is just 2% of the body’s weight, but 20% of its metabolic load (1⇓–3), and 10 times more expensive per gram than muscle. On the other hand, the brain manages to produce poetry, design spacecraft, and create art on an energy budget of ∼20 W, a paltry sum given that the computer on which this article is being typed requires 80 W. So where in the brain is power consumed, what is it used for, why is it so expensive relative to other costs of living, and how does it achieve its power efficiency relative to engineered silicon? Many classic papers have studied these questions. Attwell and Laughlin (4) developed detailed biophysical estimates suggesting that neural signaling and the postsynaptic effects of neurotransmitter release combined to account for 80% of the brain’s adenosine triphosphate (ATP) consumption, conclusions that are also supported by the overall physiology and anatomy of neural circuits (5, 6). Numerous studies explored the structural and functional consequences of this expenditure for limiting brain size (7) and scaling (8), efficient wiring patterns (9), analog (graded potential) vs. digital (spiking) signaling (10), distributed neural codes (11⇓–13), the distribution of information traffic along nerve tracts and their size distribution (14⇓–16), and computational heterogeneity and efficiency (17). Many of these ideas have been synthesized by Sterling and Laughlin (18) into a set of principles governing the design of brains. Now, in PNAS, Levy and Calvert (19) propose a functional accounting of the power budget of the mammalian brain, suggesting that communication is vastly more expensive than computation, and exploring the functional consequences for neural circuit organization. Levy and Calvert (19) build on the earlier literature by focusing primarily on the relative power committed to different modes of information … [↵][1]1Email: vijay{at}physics.upenn.edu. [1]: #xref-corresp-1-1
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“The mechanism of its emergence is very energy efficient.”
From There is no such thing as conscious artificial intelligence (2025) · cites this paper as Balasubramanian, 2021
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“The brain itself uses approximately 400 kCal per day, which is less than 0.5 kWh”
From There is no such thing as conscious artificial intelligence (2025) · cites this paper as Balasubramanian, 2021
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Cites
Irreversibility and heat generation in the computing process
Landauer, R. (1961) · IBM Journal of Research and Development · doi:10.1147/rd.53.0183 · Cited by 4,980
Interplay between Cortical Spreading Depolarization and Seizures
Kramer, D. R., Fujii, T., Ohiorhenuan, I. E., & Liu, C. Y. (2017) · Stereotactic and Functional Neurosurgery · doi:10.1159/000452841 · Cited by 3,721
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
Irreversibility and heat generation in the computing process
Landauer, R. W. (2000) · IBM Journal of Research and Development · doi:10.1147/rd.441.0261 · Cited by 2,665
The 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
Cellular energy utilization and molecular origin of standard metabolic rate in mammals
Rolfe, D. F., & Brown, G. C. (1997) · Physiological Reviews · doi:10.1152/physrev.1997.77.3.731 · Cited by 2,021
Siesjö, B. K. (1978) · Medical Entomology and Zoology · Cited by 1,426
The cost of cortical computation.
Lennie, P. (2003) · Current Biology · doi:10.1016/s0960-9822(03)00135-0 · Cited by 1,134
CNS energy metabolism as related to function
Ames, A. (2000) · Brain Research Reviews · doi:10.1016/s0165-0173(00)00038-2 · Cited by 663
Analog Versus Digital: Extrapolating from Electronics to Neurobiology
Sarpeshkar, R. (1998) · Neural Computation · doi:10.1162/089976698300017052 · Cited by 496
Flagged first, then by citations
1 of the 28 references couldn’t be matched to a known paper.
Research works printed but not linked: 1
- L. Sokoloff, “The metabolism of the central nervous system in vivo” in Neurophysiology, J. Field, H. W. Magoun, V. E. Hall, Eds. (Handbook of Physiology, American Physiological Society, 1960), sec. 1, vol. 3, pp. 1843–1864.
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Items 101–113 of 113
Spermidine protects mitochondria against amyloid-β oligomers
Mosca, D., Parfenova, E., Bie, Y., Mutsaers, A. J., & Spagnuolo, P. A. (2026) · Journal of Alzheimer s Disease Reports · doi:10.1177/25424823261449500
System-level FPGA validation of a trainable and robust multiplier-free spiking neural network
Li, Q., & (38117), L. Z. (2026) · iScience · doi:10.1016/j.isci.2026.115985
Metabolic Remodeling of the Parkinson’s Disease Frontal Cortex Revealed by LC-MS/MS Metabolomics
Daramola, O. E., Nwaiwu, J. I., Oluokun, O. O., Fowowe, M., Lux, A., Lopez, I., Bennett, A. I., & Mechref, Y. S. (2026) · Biomolecules · doi:10.3390/biom16060866
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Goldhaber-Gordon, S., Akwaboah, A. D., Sampson, A. L., Etienne‐Cummings, R., Andreou, A. G., Mihalaş, Ş., & Niebur, E. (2026) · International Conference on Neuromorphic Systems (ICONS) · doi:10.1145/3822454.3822479
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Steinkühler, J., & Mussel, M. (2026) · Advances in biomembranes and lipid self-assembly · doi:10.1016/bs.abl.2026.06.001
Fully in-memory photonic computing
Zhou, T., Wu, W., & Fang, L. (2026) · Nature Sensors · doi:10.1038/s44460-026-00123-2
Neto, M. C. (2026) · Revista de Geopolítica · doi:10.56238/revgeov17n9-016
Transcending natural evolution in the age of artificial intelligence
Guo, P., Jia, X., Tao, L.-Q., Yang, Y., & Ren, T.-L. (2026) · Science China Information Sciences · doi:10.1007/s11432-026-5041-7
Nutritional Modulation of Cognitive Function and Brain Networks: A Comprehensive Review
Krishna, D., Mishra, A., J, S. K., & Dhevi, B. (2026) · Brain Science Advances · doi:10.26599/bsa.2026.905024
OscVision: Oscillator Networks for Low Power Vision Inference
Cai, W., Li, Z., Keshavarzi, A., & Lee, T. H. (2026) · International Journal of Computer Vision · doi:10.1007/s11263-026-03044-2
Energetic dysregulations in psychotic disorders: Targets for new therapeutics
Anderson, J., Sonntag, K., Le, T. D. V., Shannon, K., Wang, C., Du, F., Chen, X., McPhie, D. L., Öngür, D., Cohen, B. M., et al. (2026) · FEBS Journal · doi:10.1111/febs.70719
Spatio-temporal plasticity in coupled stochastic relaxation-type oscillators
Noll, M., Madurawala, R., Birkoben, T., Kaps, S., Adelung, R., Kreuz, T., & Kohlstedt, H. (2026) · Engineering Research Express · doi:10.1088/2631-8695/aeb108
Neurochemical Pathways in Energy Metabolism and Mitochondrial Function
Abubakar, M., Dahiya, R., Ambawatiya, A., Maharana, K. C., Alom, S., Rawat, A., Rai, A., Yadav, D., Adhikary, K., & Kumar, N. (2026) · Royal Society of Chemistry eBooks · doi:10.1039/9781837071395-00102
Metadata
- Authors
- Balasubramanian, V.
- Affiliations
- University of Pennsylvania (US)
- Corresponding author
- Balasubramanian, Vijay
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2021-08-02
- Volume
- 118
- Issue
- 32
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC8364152
- OpenAlex
- W3196841063
- Cited by
- 113
- References
- 28
- Citations by year
- 2026: 28 · 2025: 38 · 2024: 25 · 2023: 19 · 2022: 3
- FWCI
- 6.18
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- Funders
- Simons Foundation (400425)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Physical Sciences › Computer Science › Artificial Intelligence › Neural Networks and Applications
- Keywords
- brain energy consumption, neural communication, neural computation, energy efficiency, ATP consumption, neural circuit organization, mammalian brain
- MeSH terms
- Brain, Head
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