DOI: 10.1126/science.aal3321. A subcellular map of the human proteome

10.1126/science.aal3321

Crossref journal-article

American Association for the Advancement of Science (AAAS)

Science (221)

Abstract

Mapping the proteome Proteins function in the context of their environment, so an understanding of cellular processes requires a knowledge of protein localization. Thul et al. used immunofluorescence microscopy to map 12,003 human proteins at a single-cell level into 30 cellular compartments and substructures (see the Perspective by Horwitz and Johnson). They validated their results by mass spectroscopy and used them to model and refine protein-protein interaction networks. The cellular proteome is highly spatiotemporally regulated. Many proteins localize to multiple compartments, and many show cell-to-cell variation in their expression patterns. Presented as an interactive database called the Cell Atlas, this work provides an important resource for ongoing efforts to understand human biology. Science , this issue p. eaal3321 ; see also p. 806

Authors 43

Peter J. Thul (first)
Lovisa Åkesson (additional)
Mikaela Wiking (additional)
Diana Mahdessian (additional)
Aikaterini Geladaki (additional)
Hammou Ait Blal (additional)
Tove Alm (additional)
Anna Asplund (additional)
Lars Björk (additional)
Lisa M. Breckels (additional)
Anna Bäckström (additional)
Frida Danielsson (additional)
Linn Fagerberg (additional)
Jenny Fall (additional)
Laurent Gatto (additional)
Christian Gnann (additional)
Sophia Hober (additional)
Martin Hjelmare (additional)
Fredric Johansson (additional)
Sunjae Lee (additional)
Cecilia Lindskog (additional)
Jan Mulder (additional)
Claire M. Mulvey (additional)
Peter Nilsson (additional)
Per Oksvold (additional)
Johan Rockberg (additional)
Rutger Schutten (additional)
Jochen M. Schwenk (additional)
Åsa Sivertsson (additional)
Evelina Sjöstedt (additional)
Marie Skogs (additional)
Charlotte Stadler (additional)
Devin P. Sullivan (additional)
Hanna Tegel (additional)
Casper Winsnes (additional)
Cheng Zhang (additional)
Martin Zwahlen (additional)
Adil Mardinoglu (additional)
Fredrik Pontén (additional)
Kalle von Feilitzen (additional)
Kathryn S. Lilley (additional)
Mathias Uhlén (additional)
Emma Lundberg (additional)

References 66 Referenced 2,252

Dates

Type	When
Created	8 years, 3 months ago (May 11, 2017, 9 p.m.)
Deposited	1 year, 7 months ago (Jan. 15, 2024, 11:20 a.m.)
Indexed	3 hours, 33 minutes ago (Aug. 31, 2025, 7:24 p.m.)
Issued	8 years, 3 months ago (May 26, 2017)
Published	8 years, 3 months ago (May 26, 2017)
Published Print	8 years, 3 months ago (May 26, 2017)

Funders 2

Familjen Erling-Perssons Stiftelse 10.13039/100007436 Erling-Perssons Stiftelse
Region: Europe
pri (Trusts, charities, foundations (both public and private))
Labels2
1. Erling-Persson Family Foundation
2. Familjen Erling-Perssons Stiftelse
Awards1
1. award303545
Knut och Alice Wallenbergs Stiftelse 10.13039/501100004063
Region: Europe
pri (Trusts, charities, foundations (both public and private))
Labels1
1. Knut and Alice Wallenberg Foundation
Awards1
1. award303544

BibTeX

@article{Thul_2017, title={A subcellular map of the human proteome}, volume={356}, ISSN={1095-9203}, url={http://dx.doi.org/10.1126/science.aal3321}, DOI={10.1126/science.aal3321}, number={6340}, journal={Science}, publisher={American Association for the Advancement of Science (AAAS)}, author={Thul, Peter J. and Åkesson, Lovisa and Wiking, Mikaela and Mahdessian, Diana and Geladaki, Aikaterini and Ait Blal, Hammou and Alm, Tove and Asplund, Anna and Björk, Lars and Breckels, Lisa M. and Bäckström, Anna and Danielsson, Frida and Fagerberg, Linn and Fall, Jenny and Gatto, Laurent and Gnann, Christian and Hober, Sophia and Hjelmare, Martin and Johansson, Fredric and Lee, Sunjae and Lindskog, Cecilia and Mulder, Jan and Mulvey, Claire M. and Nilsson, Peter and Oksvold, Per and Rockberg, Johan and Schutten, Rutger and Schwenk, Jochen M. and Sivertsson, Åsa and Sjöstedt, Evelina and Skogs, Marie and Stadler, Charlotte and Sullivan, Devin P. and Tegel, Hanna and Winsnes, Casper and Zhang, Cheng and Zwahlen, Martin and Mardinoglu, Adil and Pontén, Fredrik and von Feilitzen, Kalle and Lilley, Kathryn S. and Uhlén, Mathias and Lundberg, Emma}, year={2017}, month=may }

JSON

{
  "indexed": {
    "date-parts": [
      [
        2025,
        8,
        31
      ]
    ],
    "date-time": "2025-08-31T23:24:18Z",
    "timestamp": 1756682658140,
    "version": "3.41.2"
  },
  "reference-count": 66,
  "publisher": "American Association for the Advancement of Science (AAAS)",
  "issue": "6340",
  "license": [
    {
      "start": {
        "date-parts": [
          [
            2018,
            5,
            26
          ]
        ],
        "date-time": "2018-05-26T00:00:00Z",
        "timestamp": 1527292800000
      },
      "content-version": "vor",
      "delay-in-days": 365,
      "URL": "http://www.sciencemag.org/about/science-licenses-journal-article-reuse"
    }
  ],
  "funder": [
    {
      "DOI": "10.13039/100007436",
      "name": "Familjen Erling-Perssons Stiftelse",
      "doi-asserted-by": "publisher",
      "award": [
        "award303545"
      ],
      "id": [
        {
          "id": "10.13039/100007436",
          "id-type": "DOI",
          "asserted-by": "publisher"
        }
      ]
    },
    {
      "DOI": "10.13039/501100004063",
      "name": "Knut och Alice Wallenbergs Stiftelse",
      "doi-asserted-by": "publisher",
      "award": [
        "award303544"
      ],
      "id": [
        {
          "id": "10.13039/501100004063",
          "id-type": "DOI",
          "asserted-by": "publisher"
        }
      ]
    }
  ],
  "content-domain": {
    "domain": [],
    "crossmark-restriction": false
  },
  "published-print": {
    "date-parts": [
      [
        2017,
        5,
        26
      ]
    ]
  },
  "abstract": "<jats:title>Mapping the proteome</jats:title>\n          <jats:p>\n            Proteins function in the context of their environment, so an understanding of cellular processes requires a knowledge of protein localization. Thul\n            <jats:italic>et al.</jats:italic>\n            used immunofluorescence microscopy to map 12,003 human proteins at a single-cell level into 30 cellular compartments and substructures (see the Perspective by Horwitz and Johnson). They validated their results by mass spectroscopy and used them to model and refine protein-protein interaction networks. The cellular proteome is highly spatiotemporally regulated. Many proteins localize to multiple compartments, and many show cell-to-cell variation in their expression patterns. Presented as an interactive database called the Cell Atlas, this work provides an important resource for ongoing efforts to understand human biology.\n          </jats:p>\n          <jats:p>\n            <jats:italic>Science</jats:italic>\n            , this issue p.\n            <jats:related-article xmlns:xlink=\"http://www.w3.org/1999/xlink\" ext-link-type=\"doi\" issue=\"6340\" page=\"eaal3321\" related-article-type=\"in-this-issue\" vol=\"356\" xlink:href=\"10.1126/science.aal3321\">eaal3321</jats:related-article>\n            ; see also p.\n            <jats:related-article xmlns:xlink=\"http://www.w3.org/1999/xlink\" ext-link-type=\"doi\" issue=\"6340\" page=\"806\" related-article-type=\"in-this-issue\" vol=\"356\" xlink:href=\"10.1126/science.aan5955\">806</jats:related-article>\n          </jats:p>",
  "DOI": "10.1126/science.aal3321",
  "type": "journal-article",
  "created": {
    "date-parts": [
      [
        2017,
        5,
        12
      ]
    ],
    "date-time": "2017-05-12T01:00:39Z",
    "timestamp": 1494550839000
  },
  "source": "Crossref",
  "is-referenced-by-count": 2252,
  "title": "A subcellular map of the human proteome",
  "prefix": "10.1126",
  "volume": "356",
  "author": [
    {
      "ORCID": "https://orcid.org/0000-0002-6107-1465",
      "authenticated-orcid": false,
      "given": "Peter J.",
      "family": "Thul",
      "sequence": "first",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-2387-3491",
      "authenticated-orcid": false,
      "given": "Lovisa",
      "family": "\u00c5kesson",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-6368-6690",
      "authenticated-orcid": false,
      "given": "Mikaela",
      "family": "Wiking",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0003-0750-1070",
      "authenticated-orcid": false,
      "given": "Diana",
      "family": "Mahdessian",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-0530-4252",
      "authenticated-orcid": false,
      "given": "Aikaterini",
      "family": "Geladaki",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Cambridge Centre for Proteomics, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        },
        {
          "name": "Department of Genetics, University of Cambridge, Downing Street, Cambridge CB2 3EH, UK."
        }
      ]
    },
    {
      "given": "Hammou",
      "family": "Ait Blal",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-2643-8241",
      "authenticated-orcid": false,
      "given": "Tove",
      "family": "Alm",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Anna",
      "family": "Asplund",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, SE-751 85 Uppsala, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-8116-882X",
      "authenticated-orcid": false,
      "given": "Lars",
      "family": "Bj\u00f6rk",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-8918-7171",
      "authenticated-orcid": false,
      "given": "Lisa M.",
      "family": "Breckels",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Cambridge Centre for Proteomics, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        },
        {
          "name": "Computational Proteomics Unit, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        }
      ]
    },
    {
      "given": "Anna",
      "family": "B\u00e4ckstr\u00f6m",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Frida",
      "family": "Danielsson",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0003-0198-7137",
      "authenticated-orcid": false,
      "given": "Linn",
      "family": "Fagerberg",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Jenny",
      "family": "Fall",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-1520-2268",
      "authenticated-orcid": false,
      "given": "Laurent",
      "family": "Gatto",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Cambridge Centre for Proteomics, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        },
        {
          "name": "Computational Proteomics Unit, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        }
      ]
    },
    {
      "given": "Christian",
      "family": "Gnann",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Sophia",
      "family": "Hober",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Proteomics, School of Biotechnology, KTH Royal Institute of Technology, SE-106 91 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-7375-9681",
      "authenticated-orcid": false,
      "given": "Martin",
      "family": "Hjelmare",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-5160-9543",
      "authenticated-orcid": false,
      "given": "Fredric",
      "family": "Johansson",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-6428-5936",
      "authenticated-orcid": false,
      "given": "Sunjae",
      "family": "Lee",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Cecilia",
      "family": "Lindskog",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, SE-751 85 Uppsala, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0003-3717-5018",
      "authenticated-orcid": false,
      "given": "Jan",
      "family": "Mulder",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, Department of Neuroscience, Karolinska Institute, SE-171 77 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-2989-2052",
      "authenticated-orcid": false,
      "given": "Claire M.",
      "family": "Mulvey",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Cambridge Centre for Proteomics, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-4657-8532",
      "authenticated-orcid": false,
      "given": "Peter",
      "family": "Nilsson",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0003-3014-5502",
      "authenticated-orcid": false,
      "given": "Per",
      "family": "Oksvold",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-9977-5724",
      "authenticated-orcid": false,
      "given": "Johan",
      "family": "Rockberg",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Proteomics, School of Biotechnology, KTH Royal Institute of Technology, SE-106 91 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-8787-8868",
      "authenticated-orcid": false,
      "given": "Rutger",
      "family": "Schutten",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-8141-8449",
      "authenticated-orcid": false,
      "given": "Jochen M.",
      "family": "Schwenk",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-8800-8469",
      "authenticated-orcid": false,
      "given": "\u00c5sa",
      "family": "Sivertsson",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-0327-7377",
      "authenticated-orcid": false,
      "given": "Evelina",
      "family": "Sj\u00f6stedt",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, SE-751 85 Uppsala, Sweden."
        }
      ]
    },
    {
      "given": "Marie",
      "family": "Skogs",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-6781-1938",
      "authenticated-orcid": false,
      "given": "Charlotte",
      "family": "Stadler",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-6176-108X",
      "authenticated-orcid": false,
      "given": "Devin P.",
      "family": "Sullivan",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-7067-9173",
      "authenticated-orcid": false,
      "given": "Hanna",
      "family": "Tegel",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Proteomics, School of Biotechnology, KTH Royal Institute of Technology, SE-106 91 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-0028-5865",
      "authenticated-orcid": false,
      "given": "Casper",
      "family": "Winsnes",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-3721-8586",
      "authenticated-orcid": false,
      "given": "Cheng",
      "family": "Zhang",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-0064-4776",
      "authenticated-orcid": false,
      "given": "Martin",
      "family": "Zwahlen",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-4254-6090",
      "authenticated-orcid": false,
      "given": "Adil",
      "family": "Mardinoglu",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0003-0703-3940",
      "authenticated-orcid": false,
      "given": "Fredrik",
      "family": "Pont\u00e9n",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, SE-751 85 Uppsala, Sweden."
        }
      ]
    },
    {
      "given": "Kalle",
      "family": "von Feilitzen",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "given": "Kathryn S.",
      "family": "Lilley",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Cambridge Centre for Proteomics, Department of Biochemistry, University of Cambridge, Tennis Court Road, Cambridge CB2 1QR, UK."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0002-4858-8056",
      "authenticated-orcid": false,
      "given": "Mathias",
      "family": "Uhl\u00e9n",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    },
    {
      "ORCID": "https://orcid.org/0000-0001-7034-0850",
      "authenticated-orcid": false,
      "given": "Emma",
      "family": "Lundberg",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Science for Life Laboratory, School of Biotechnology, KTH Royal Institute of Technology, SE-171 21 Stockholm, Sweden."
        }
      ]
    }
  ],
  "member": "221",
  "reference": [
    {
      "key": "e_1_3_2_2_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1186/1471-2164-10-122"
    },
    {
      "key": "e_1_3_2_3_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/msb.2011.29"
    },
    {
      "key": "e_1_3_2_4_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1074/mcp.T400009-MCP200"
    },
    {
      "key": "e_1_3_2_5_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.cell.2006.03.022"
    },
    {
      "key": "e_1_3_2_6_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/emboj.2011.63"
    },
    {
      "key": "e_1_3_2_7_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/ncomms9992"
    },
    {
      "key": "e_1_3_2_8_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.7554/eLife.16950"
    },
    {
      "key": "e_1_3_2_9_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1002/0471140864.ps1923s74"
    },
    {
      "key": "e_1_3_2_10_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1126/science.1230593"
    },
    {
      "key": "e_1_3_2_11_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.molcel.2014.06.003"
    },
    {
      "key": "e_1_3_2_12_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.celrep.2016.04.064"
    },
    {
      "key": "e_1_3_2_13_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/embo-reports/kvd058"
    },
    {
      "key": "e_1_3_2_14_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nature02026"
    },
    {
      "key": "e_1_3_2_15_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.cell.2015.04.051"
    },
    {
      "key": "e_1_3_2_16_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1074/mcp.M700325-MCP200"
    },
    {
      "key": "e_1_3_2_17_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.jprot.2009.10.012"
    },
    {
      "key": "e_1_3_2_18_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkm259"
    },
    {
      "key": "e_1_3_2_19_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1039/C1MB05420A"
    },
    {
      "key": "e_1_3_2_20_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gku989"
    },
    {
      "key": "e_1_3_2_21_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1021/pr200379a"
    },
    {
      "key": "e_1_3_2_22_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nmeth.2377"
    },
    {
      "key": "e_1_3_2_23_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1074/mcp.M116.064527"
    },
    {
      "key": "e_1_3_2_24_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nbt1210-1248"
    },
    {
      "key": "e_1_3_2_25_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1126/science.1260419"
    },
    {
      "key": "e_1_3_2_26_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkr1079"
    },
    {
      "key": "e_1_3_2_27_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkv1157"
    },
    {
      "key": "e_1_3_2_28_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkq1112"
    },
    {
      "key": "e_1_3_2_29_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1126/science.1262110"
    },
    {
      "key": "e_1_3_2_30_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gku1178"
    },
    {
      "key": "e_1_3_2_31_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/msb.2011.82"
    },
    {
      "key": "e_1_3_2_32_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1074/mcp.M111.014050"
    },
    {
      "key": "e_1_3_2_33_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1002/pmic.200500072"
    },
    {
      "key": "e_1_3_2_34_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nbt0516-452c"
    },
    {
      "key": "e_1_3_2_35_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.2144/000113382"
    },
    {
      "key": "e_1_3_2_36_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/521274a"
    },
    {
      "key": "e_1_3_2_37_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nmeth.3995"
    },
    {
      "key": "e_1_3_2_38_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.jprot.2012.01.030"
    },
    {
      "key": "e_1_3_2_39_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1021/acs.jproteome.6b00821"
    },
    {
      "key": "e_1_3_2_40_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.3851/IMP1993"
    },
    {
      "key": "e_1_3_2_41_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkq1009"
    },
    {
      "key": "e_1_3_2_42_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1210/me.2010-0216"
    },
    {
      "key": "e_1_3_2_43_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkp914"
    },
    {
      "key": "e_1_3_2_44_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/bioinformatics/btu013"
    },
    {
      "key": "e_1_3_2_45_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1371/journal.pcbi.1004920"
    },
    {
      "key": "e_1_3_2_46_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gkv1351"
    },
    {
      "key": "e_1_3_2_47_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1126/science.1257601"
    },
    {
      "key": "e_1_3_2_48_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.cell.2007.12.033"
    },
    {
      "key": "e_1_3_2_49_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/nar/gku1179"
    },
    {
      "key": "e_1_3_2_50_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.tibtech.2012.02.002"
    },
    {
      "key": "e_1_3_2_51_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/ncomms8412"
    },
    {
      "key": "e_1_3_2_52_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.canlet.2015.09.022"
    },
    {
      "key": "e_1_3_2_53_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.2144/05385ST02"
    },
    {
      "key": "e_1_3_2_54_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nmeth.1701"
    },
    {
      "key": "e_1_3_2_55_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1016/j.jmb.2004.03.016"
    },
    {
      "key": "e_1_3_2_56_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/bioinformatics/btn550"
    },
    {
      "key": "e_1_3_2_57_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1002/pmic.200900258"
    },
    {
      "key": "e_1_3_2_58_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1021/pr3009308"
    },
    {
      "key": "e_1_3_2_59_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nbt.3519"
    },
    {
      "key": "e_1_3_2_60_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1101/gr.092759.109"
    },
    {
      "key": "e_1_3_2_61_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/bioinformatics/btp536"
    },
    {
      "key": "e_1_3_2_62_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1038/nprot.2008.211"
    },
    {
      "key": "e_1_3_2_63_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1101/gr.1239303"
    },
    {
      "key": "e_1_3_2_64_2",
      "unstructured": "R. Core Team R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing 2013); http://www.R-project.org/."
    },
    {
      "key": "e_1_3_2_65_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1186/gb-2004-5-10-r80"
    },
    {
      "key": "e_1_3_2_66_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1093/bioinformatics/btr645"
    },
    {
      "key": "e_1_3_2_67_2",
      "doi-asserted-by": "publisher",
      "DOI": "10.1002/pmic.201000359"
    }
  ],
  "container-title": "Science",
  "original-title": [],
  "language": "en",
  "link": [
    {
      "URL": "https://www.science.org/doi/pdf/10.1126/science.aal3321",
      "content-type": "unspecified",
      "content-version": "vor",
      "intended-application": "similarity-checking"
    }
  ],
  "deposited": {
    "date-parts": [
      [
        2024,
        1,
        15
      ]
    ],
    "date-time": "2024-01-15T16:20:10Z",
    "timestamp": 1705335610000
  },
  "score": 1,
  "resource": {
    "primary": {
      "URL": "https://www.science.org/doi/10.1126/science.aal3321"
    }
  },
  "subtitle": [],
  "short-title": [],
  "issued": {
    "date-parts": [
      [
        2017,
        5,
        26
      ]
    ]
  },
  "references-count": 66,
  "journal-issue": {
    "issue": "6340",
    "published-print": {
      "date-parts": [
        [
          2017,
          5,
          26
        ]
      ]
    }
  },
  "alternative-id": [
    "10.1126/science.aal3321"
  ],
  "URL": "http://dx.doi.org/10.1126/science.aal3321",
  "relation": {
    "has-review": [
      {
        "id-type": "doi",
        "id": "10.3410/f.727603588.793532452",
        "asserted-by": "object"
      },
      {
        "id-type": "doi",
        "id": "10.3410/f.727603588.793532933",
        "asserted-by": "object"
      },
      {
        "id-type": "doi",
        "id": "10.3410/f.727603588.793533693",
        "asserted-by": "object"
      },
      {
        "id-type": "doi",
        "id": "10.3410/f.727603588.793538041",
        "asserted-by": "object"
      },
      {
        "id-type": "doi",
        "id": "10.3410/f.727603588.793532595",
        "asserted-by": "object"
      }
    ]
  },
  "ISSN": [
    "0036-8075",
    "1095-9203"
  ],
  "subject": [],
  "container-title-short": "Science",
  "published": {
    "date-parts": [
      [
        2017,
        5,
        26
      ]
    ]
  },
  "article-number": "eaal3321"
}