Top spots: The lists surface several important trends, including a focus on translational work, the rise of artificial intelligence and recent technical advances including Neuropixels and brain-computer interfaces.

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Highly cited papers reflect the surge in artificial-intelligence research in the field and other technical advances, plus prizewinning work on analgesics, the fusiform face area and ion channels.

Citations are an imperfect measure of scientific value, but they can provide a snapshot of papers that have proved impactful over time. Some highly cited papers put forth useful techniques or scintillating new results. Others provide fodder for debate. They all reflect research areas that scientists have been excited about at some point in time.

*The Transmitter *gathered citation metrics for the top neuroscience papers published over the past 30 years and assembled a list of the top 20 to try to capture a different view of the field’s changing landscape. To incorporate recent work that hasn’t had time to amass top-20-level counts, we also created lists of the top-five cited papers for each of the past five years.

Together, the lists surface a number of pivotal advances. As our surveys make clear, work in artificial intelligence, and computational neuroscience in general, has exploded over a relatively short span of time: Here, too, the top two most-cited neuroscience papers are both less than a decade old and concern an AI’s ability to master the game Go, outperforming a human professional player and earlier versions of itself. The last author on both papers, Demis Hassabis, co-founder and CEO of GoogleDeepMind, won the Nobel Prize in Chemistry in 2024 for his work on AlphaFold, applying AI to predict protein structure.

Work with translational potential also collected high citation numbers—a clinical bias that is also apparent in our interactive trends map. The third paper on the list, describing efforts that earned last author David Julius the Nobel Prize in Physiology or Medicine in 2021, helped define the mechanism of action and new targets for analgesic drugs. The paper at No. 14 first identified TDP-43 as a protein of interest in amyotrophic lateral sclerosis; No. 6 revealed how the serotonin transporter gene moderates the impact of stress in depression; and two papers from the past five years examine the effects of SARS-CoV-2.

Research detailing brain structure and brain networks features prominently on these top citation lists, too. No. 10, which describes findings that earned last author Roderick MacKinnon the Nobel Prize in Chemistry in 2003, offers a deep dive into the structure of the potassium channel. The paper at No. 7 shows that the fusiform gyrus area of the brain activates strongly when participants view a human face versus control—work that earned first author Nancy Kanwisher last year’s Kavli Prize. And the No. 9 paper demonstrates that salience and executive function networks, though often coactivated, are distinct; first author William Seeley received a MacArthur “Genius” grant in 2011. Still other papers mark important technical leaps, including Neuropixels 2.0, brain-computer interfaces and the Allen Brain Mouse Brain Common Coordinate Framework.

To generate the lists, we used a dataset—created by Dennis Vasquez Montes, a research data analyst at the Simons Foundation—of papers published in 12 leading journals between 1994 and 2024 indexed in PubMed. (The journal names are included below, beneath the final citation list; The Transmitter is editorially independent of but supported by the Simons Foundation.) Vasquez Montes obtained the papers’ metadata through the Entrez Programming Utilities provided by the National Center for Biotechnology Information. He also excluded papers that could not be linked to Semantic Scholar via DOI, PMID or PMCID, and used Google Gemini to exclude papers unrelated to neuroscience based on the title and abstract. The Transmitter’s editorial team removed additional outliers by hand. He obtained citation data for each paper through Semantic Scholar’s application programming interface.

We plan to draw on this dataset again for future coverage, but in the meantime, read on to see how many of the top highly cited papers you are already familiar with.

1. **17,264 citations: **Mastering the game of Go with deep neural networks and tree search. Silver D., Huang A., Maddison C.J., Guez A., Sifre L., van den Driessche G., Schrittwieser J., Antonoglou I., Panneershelvam V. … Hassabis D. *Nature *(2016)

2. 9,520 citations: Mastering the game of Go without human knowledge. Silver D., Schrittwieser J., Simonyan K., Antonoglou I., Huang A., Guez A., Hubert T., Baker L., Lai M. … Hassabis D. Nature (2017)

3. 8,681 citations: The capsaicin receptor: A heat-activated ion channel in the pain pathway. Caterina M.J., Schumacher M.A., Tominaga M., Rosen T.A., Levine J.D., Julius D. *Nature *(1997)

4. 8,260 citations: Whole brain segmentation: Automated labeling of neuroanatomical structures in the human brain. Fischl B., Salat D.H., Busa E., Albert M., Dieterich M., Haselgrove C., van der Kouwe A., Killiany R., Kennedy D. … Dale A.M. *Neuron *(2002)

5. 8,211 citations: The human brain is intrinsically organized into dynamic, anticorrelated functional networks. Fox M.D., Snyder A.Z., Vincent J.L., Corbetta M., Van Essen D.C., Raichle M.E. *Proceedings of the National Academy of Sciences *(2005)

6. 7,847 citations: Influence of life stress on depression: Moderation by a polymorphism in the 5-HTT gene. Caspi A., Sugden K., Moffitt T.E., Taylor A., Craig I.W., Harrington H., McClay J., Mill J., Martin J., Braithwaite A., Poulton R. *Science *(2003)

7. 7,680 citations: The fusiform face area: A module in human extrastriate cortex specialized for face perception. Kanwisher N., McDermott J., Chun M.M. *The Journal of Neuroscience *(1997)

8. 6,892 citations: Memory, visual discrimination performance, and the human hippocampus. Kim S., Jeneson A., van der Horst A.S., Frascino J.C., Hopkins R.O., Squire L.R. *The Journal of Neuroscience *(2011)

9. 6,592 citations: Dissociable intrinsic connectivity networks for salience processing and executive control. Seeley W.W., Menon V., Schatzberg A.F., Keller J., Glover G.H., Kenna H., Reiss A.L., Greicius M.D. *The Journal of Neuroscience *(2007)

10. 6,496 citations: The structure of the potassium channel: Molecular basis of K+ conduction and selectivity. Doyle D.A., Cabral J.M., Pfuetzner R.A., Kuo A., Gulbis J.M., Cohen S.L., Chait B.T., MacKinnon R. *Science *(1998)

11. 6,436 citations: Neurophysiological investigation of the basis of the fMRI signal. Logothetis N.K., Pauls J., Augath M., Trinath T., Oeltermann A. *Nature *(2001)

12. **6,313 citations: **Functional connectivity in the resting brain: A network analysis of the default mode hypothesis. Greicius M.D., Krasnow B., Reiss A.L., Menon V. *Proceedings of the National Academy of Sciences *(2002)

13. 6,223 citations: Emergence of simple-cell receptive field properties by learning a sparse code for natural images. Olshausen B.A., Field D.J. *Nature *(1996)

14. **5,973 citations: **Ubiquitinated TDP-43 in frontotemporal lobar degeneration and amyotrophic lateral sclerosis. Neumann M., Sampathu D.M., Kwong L.K., Truax A.C., Micsenyi M.C., Chou T.T., Bruce J., Schuck T., Grossman M. … Lee V.M.Y. *Science *(2006)

15. **5,961 citations: **A robust and high-throughput Cre reporting and characterization system for the whole mouse brain. Madisen L., Zwingman T.A., Sunkin S.M., Oh S.W., Zariwala H.A., Gu H., Ng L.L., Palmiter R.D., Hawrylycz M.J., Jones A.R., Lein E.S., Zeng H. *Nature Neuroscience *(2009)

16. 5,797 citations: Epigenetic programming by maternal behavior. Weaver I.C., Cervoni N., Champagne F.A., D’Alessio A.C., Sharma S., Seckl J.R., Dymov S., Szyf M., Meaney M.J. *Nature Neuroscience *(2004)

17. 5,674 citations: Measuring the thickness of the human cerebral cortex from magnetic resonance images. Fischl B., Dale A.M. *Proceedings of the National Academy of Sciences *(2000)

18. 5,598 citations: Opposing effects of ERK and JNK-p38 MAP kinases on apoptosis. Xia Z., Dickens M., Raingeaud J., Davis R.J., Greenberg M.E. *Science *(1995)

19. 5,356 citations: Resting microglial cells are highly dynamic surveillants of brain parenchyma in vivo. Nimmerjahn A., Kirchhoff F., Helmchen F. *Science *(2005)

20. **5,316 citations: **Dynamic mapping of human cortical development during childhood through early adulthood. Gogtay N., Giedd J.N., Lusk L., Hayashi K.M., Greenstein D., Vaituzis A.C., Nugent T.F., Herman D.H., Clasen L.S., Toga A.W., Rapoport J.L., Thompson P.M. *Proceedings of the National Academy of Sciences *(2004)

21. **216 citations: **Synergizing habits and goals with variational Bayes. Han D., Doya K., Li D., Tani J. *Nature Communications *(2024)

22. 160 citations: APOE4/4 is linked to damaging lipid droplets in Alzheimer’s disease microglia. Haney M.S., Pálovics R., Munson C.N., Long C., Johansson P.K., Yip O., Dong W., Rawat E., West E. … Wyss-Coray T. *Nature *(2024)

23. 126 citations: A petavoxel fragment of human cerebral cortex reconstructed at nanoscale resolution. Shapson-Coe A., Januszewski M., Berger D.R., Pope A., Wu Y., Blakely T., Schalek R.L., Li P.H. … Jain V., Lichtman J.W. *Science *(2024)

24. **126 citations: **A body-brain circuit that regulates body inflammatory responses. Jin H., Li M., Jeong E., Castro-Martinez F., Zuker C.S. *Nature *(2024)

25. 115 citations: Neurotransmitter classification from electron microscopy images at synaptic sites in Drosophila melanogaster. Eckstein N., Bates A.S., Champion A., Du M., Yin Y., Schlegel P., Lu A.K.Y., Rymer T. … Jefferis G.S.X.E., Funke J. *Cell *(2024)

26. 400 citations: Fast and sensitive GCaMP calcium indicators for imaging neural populations. Zhang Y., Rózsa M., Liang Y., Bushey D., Wei Z. … Turner G.C., Wang S.S.-H., Svoboda K., Hasseman J.P., Looger L.L. *Nature *(2023)

27. 312 citations: Transcriptomic diversity of cell types across the adult human brain. Siletti K., Hodge R., Mossi Albiach A., Lee K.W., Ding S.L., Hu L., Lönnerberg P., Bakken T., Casper T. … Linnarsson S. *Science *(2023)

28. 309 citations: Microglia-mediated T cell infiltration drives neurodegeneration in tauopathy. Chen X., Firulyova M., Manis M., Herz J., Smirnov I., Aladyeva E., Wang C., Bao X., Finn M.B. … Holtzman D.M. *Nature *(2023)

29. **294 citations: **Walking naturally after spinal cord injury using a brain-spine interface. Lorach H., Galvez A., Spagnolo V., Martel F., Karakas S., Intering N., Vat M. … Charvet G., Bloch J., Courtine G. *Nature *(2023)

30. 267 citations: Semantic reconstruction of continuous language from non-invasive brain recordings. Tang J., LeBel A., Jain S., Huth A.G. *Nature Neuroscience *(2023)

31. 1,612 citations: Mapping genomic loci implicates genes and synaptic biology in schizophrenia. Trubetskoy V., Pardiñas A.F., Qi T., Panagiotaropoulou G., Awasthi S., Bigdeli T.B., Bryois J. … Ripke S., Walters J.T.R., O’Donovan M.C. *Nature *(2022)

32. **1,258 citations: **Reproducible brain-wide association studies require thousands of individuals. Marek S., Tervo-Clemmens B., Calabro F.J., Montez D.F., Kay B.P., Hatoum A.S., Donohue M.R., Foran W. … Fair D.A., Dosenbach N.U. *Nature *(2022)

33. 1,107 citations: SARS-CoV-2 is associated with changes in brain structure in UK Biobank. Douaud G., Lee S., Alfaro-Almagro F., Arthofer C., Wang C., McCarthy P., Lange F., Andersson J.L., Griffanti L. … Smith S.M. *Nature *(2022)

34. 519 citations: Rare coding variants in ten genes confer substantial risk for schizophrenia. Singh T., Poterba T., Curtis D., Akil H., Al Eissa M., Barchas J.D., Bass N., Bigdeli T.B. … Neale B.M., Daly M.J. *Nature *(2022)

35. 473 citations: A human brain vascular atlas reveals diverse mediators of Alzheimer’s risk. Yang A.C., Vest R.T., Kern F., Lee D.P., Agam M., Maat C.A., Losada P.M., Chen M.B., Schaum N. … Wyss-Coray T. Nature (2022)

36. 785 citations: Neuropixels 2.0: A miniaturized high-density probe for stable, long-term brain recordings. Steinmetz N.A., Aydin C., Lebedeva A., Okun M., Pachitariu M., Bauza M., Beau M., Bhagat J. … Carandini M., Harris T.D. *Science *(2021)

37. 705 citations: Transcriptome-scale spatial gene expression in the human dorsolateral prefrontal cortex. Maynard K.R., Collado-Torres L., Weber L.M., Uytingco C., Barry B.K., Williams S.R., Catallini J.L., Tran M.N. … Martinowich K., Jaffe A.E. *Nature Neuroscience *(2021)

38. 678 citations: A taxonomy of transcriptomic cell types across the isocortex and hippocampal formation. Yao Z., van Velthoven C.T.J., Nguyen T.N., Goldy J., Sedeno-Cortes A.E., Baftizadeh F., Bertagnolli D., Casper T., Chiang M. … Zeng H. *Cell *(2021)

39. **626 citations: **High-performance brain-to-text communication via handwriting. Willett F.R., Avansino D.T., Hochberg L.R., Henderson J.M., Shenoy K.V. *Nature *(2021)

40. 539 citations: Structure-based classification of tauopathies. Shi Y., Zhang W., Yang Y., Murzin A.G., Falcon B., Kotecha A., van Beers M., Tarutani A. … Goedert M., Scheres S.H. *Nature *(2021)

41. **1,670 citations: **Large-scale exome sequencing study implicates both developmental and functional changes in the neurobiology of autism. Satterstrom F.K., Kosmicki J.A., Wang J., Breen M.S., De Rubeis S., An J.Y. … Sanders S.J., Roeder K., Daly M.J., Buxbaum J.D. *Cell *(2020)

42. 1,220 citations: Parameterizing neural power spectra into periodic and aperiodic components. Donoghue T., Haller M., Peterson E.J., Varma P., Sebastian P., Gao R., Noto T., Lara A.H., Wallis J.D. … Voytek B. *Nature Neuroscience *(2020)

43. 1,110 citations: Olfactory transmucosal SARS-CoV-2 invasion as a port of central nervous system entry in individuals with COVID-19. Meinhardt J., Radke J., Dittmayer C., Franz J., Thomas C., Mothes R., Laue M., Schneider J. … Radbruch H., Heppner F.L. *Nature Neuroscience *(2020)

44. **1,010 citations: **The Allen Mouse Brain Common Coordinate Framework: A 3D reference atlas. Wang Q., Ding S.L., Li Y., Royall J., Feng D., Lesnar P., Graddis N., Naeemi M. … Harris J.A., Ng L. *Cell *(2020)

45. **856 citations: **APOE4 leads to blood-brain barrier dysfunction predicting cognitive decline. Montagne A., Nation D.A., Sagare A.P., Barisano G., Sweeney M.D., Chakhoyan A., Pachicano M., Joe E., Nelson A.R. … Zlokovic B.V. *Nature *(2020)

Journals included in dataset: Cell, *Cell Reports, *Current Biology, eLife, The Journal of Neuroscience, Nature, Nature Communications, Nature Neuroscience, Neuron, Proceedings of the National Academy of Sciences, *Science *and Science Advances.

tags:

Publishing, Academia, Community, Depression, Face processing, fMRI, Hippocampus, Ion channels, Neural networks, Neurodegeneration, neuroimaging