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Protein Domains

Molecular Biology · NLM MeSH · updated 2026-10-29 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 3 min read

Discrete protein structural units that may fold independently of the rest of the protein and have their own functions.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (643 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

Discrete protein structural units that may fold independently of the rest of the protein and have their own functions.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Protein Domain; Domain, Protein; Domains, Protein; Peptide Domain; Domain, Peptide; Domains, Peptide; Peptide Domains.

Where it fits in the MeSH classification

Broader topics
Protein Structural Elements; Protein Structure, Tertiary
Narrower topics
C2 Domains; Carbohydrate Binding Modules; Catalytic Domain; Discoidin Domain; HMG-Box Domains; Immunoglobulin Domains; Kringles; Methyl CpG Binding Domain; Pleckstrin Homology Domains; PR-SET Domains; Proline-Rich Protein Domains; Protein Interaction Domains and Motifs

Research landscape

PubMed indexes 82,533 records tagged with the MeSH heading “Protein Domains” (counts retrieved 2026-10-29). 21,871 of them (26.5%) were published from 2020 onward, and the busiest period so far is 2010-2019 with 44,065 records.

Reviews account for 5,649 records (6.8%) and meta-analyses for 23. Randomized controlled trials number 17 (0.0%) as a share of all records on the topic. Randomized trials make up only a small share of this literature, so much of what is known comes from laboratory, observational or descriptive work.

PubMed records by publication periodthrough 1999: 2,346 records; 2000-2009: 14,251 records; 2010-2019: 44,065 records; 2020 onward: 21,871 records2,346through 199914,2512000-200944,0652010-201921,8712020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 19992,346
2000-200914,251
2010-201944,065
2020 onward21,871
Share of records by publication typeReviews: 6.8 percent; Meta-analyses: 0.0 percent; Randomized controlled trials: 0.0 percentReviews6.8% (5,649)Meta-analyses0.0% (23)Randomized controlled trials0.0% (17)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 82,533 PubMed records
TypeRecordsShare
Reviews5,6496.8%
Meta-analyses230.0%
Randomized controlled trials170.0%

Method: PubMed query "Protein Domains"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-29.

Reading the evidence in molecular biology

Molecular biology establishes mechanisms through controlled experiments, often in simplified systems such as purified proteins, cultured cells or model organisms. Such work can show what is possible in a defined system, but whether a pathway matters in a whole organism or in human disease needs further evidence. Check the experimental system, whether findings were reproduced by independent groups, and whether the authors tested the mechanism directly or inferred it from correlation.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Site-saturation mutagenesis of 500 human protein domains
Beltran A, Jiang X, Shen Y, Lehner B
2025Research article
Design not recorded in structured metadata
125cc by
PMID 39779847 · DOI · PMC11754108
actifpTM: a refined confidence metric of AlphaFold2 predictions involving flexible regions
Varga JK, Ovchinnikov S, Schueler-Furman O
2025Research article
Design not recorded in structured metadata
67cc by
PMID 40080667 · DOI · PMC11925850
Origin and functional diversification of PAS domain, a ubiquitous intracellular sensor
Xing J, Gumerov VM, Zhulin IB
2023Research article
Design not recorded in structured metadata
51cc by
PMID 37647406 · DOI · PMC10468136
Structural ontogeny of protein-protein interactions
Yang A, Jiang H, Jude KM, Akpinaroglu D, Allenspach S, Li AJ, et al.
2026Research article
Design not recorded in structured metadata
5cc by
PMID 41678610 · DOI · PMC12904254

Site-saturation mutagenesis of 500 human protein domains [1]

Research article Beltran A, Jiang X, Shen Y, Lehner B · · 2025 · cited 125 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Missense variants that change the amino acid sequences of proteins cause one-third of human genetic diseases1.
Closing excerpt
Our data demonstrate the feasibility of assaying human protein variants at scale and provides a large consistent reference dataset for clinical variant interpretation and training and benchmarking of computational methods.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (175 words)

Missense variants that change the amino acid sequences of proteins cause one-third of human genetic diseases1. Tens of millions of missense variants exist in the current human population, and the vast majority of these have unknown functional consequences. Here we present a large-scale experimental analysis of human missense variants across many different proteins. Using DNA synthesis and cellular selection experiments we quantify the effect of more than 500,000 variants on the abundance of more than 500 human protein domains. This dataset reveals that 60% of pathogenic missense variants reduce protein stability. The contribution of stability to protein fitness varies across proteins and diseases and is particularly important in recessive disorders. We combine stability measurements with protein language models to annotate functional sites across proteins. Mutational effects on stability are largely conserved in homologous domains, enabling accurate stability prediction across entire protein families using energy models. Our data demonstrate the feasibility of assaying human protein variants at scale and provides a large consistent reference dataset for clinical variant interpretation and training and benchmarking of computational methods.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 39779847 · DOI · PMC11754108

actifpTM: a refined confidence metric of AlphaFold2 predictions involving flexible regions [2]

Research article Varga JK, Ovchinnikov S, Schueler-Furman O · · 2025 · cited 67 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
SummaryOne of the main advantages of deep learning models of protein structure, such as Alphafold2, is their ability to accurately estimate the confidence of a generated structural model, which allows us to focus on highly confident predictions.
Closing excerpt
com/sokrypton/ColabFold) repository, installable both locally or usable with Colab notebook.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (191 words)

SummaryOne of the main advantages of deep learning models of protein structure, such as Alphafold2, is their ability to accurately estimate the confidence of a generated structural model, which allows us to focus on highly confident predictions. The ipTM score provides a confidence estimate of interchain contacts in protein-protein interactions. However, interactions, in particular motif-mediated interactions, often also contain regions that remain flexible upon binding. These noninteracting flanking regions are assigned low confidence values and will affect ipTM, as it considers all interchain residue-residue pairs, and two models of the same motif-domain interaction, but differing in the length of their flanking regions, would be assigned very different values. Here, we propose actual interface pTM (actifpTM), a modified ipTM measure, that focuses on the residues participating in the interaction, resulting in a more robust measure of interaction confidence. Besides, actifpTM is calculated both for the full complex as well as for each pair of chains, making it well-suited for evaluating multi-chain complexes with a particularly critical binding interface, such as antibody-antigen interactions.Availability and implementationThe method is available as part of the ColabFold (https://github.com/sokrypton/ColabFold) repository, installable both locally or usable with Colab notebook.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 40080667 · DOI · PMC11925850

Origin and functional diversification of PAS domain, a ubiquitous intracellular sensor [3]

Research article Xing J, Gumerov VM, Zhulin IB · · 2023 · cited 51 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Signal perception is a key function in regulating biological activities and adapting to changing environments.
Closing excerpt
The close relatedness of human PAS domains to those in bacteria provides an opportunity for drug design by exploring potential natural ligands and cofactors for bacterial homologs.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (153 words)

Signal perception is a key function in regulating biological activities and adapting to changing environments. Per-Arnt-Sim (PAS) domains are ubiquitous sensors found in diverse receptors in bacteria, archaea, and eukaryotes, but their origins, distribution across the tree of life, and extent of their functional diversity are not fully characterized. Here, we show that using sequence conservation and structural information, it is possible to propose specific and potential functions for a large portion of nearly 3 million PAS domains. Our analysis suggests that PAS domains originated in bacteria and were horizontally transferred to archaea and eukaryotes. We reveal that gas sensing via a heme cofactor evolved independently in several lineages, whereas redox and light sensing via flavin adenine dinucleotide and flavin mononucleotide cofactors have the same origin. The close relatedness of human PAS domains to those in bacteria provides an opportunity for drug design by exploring potential natural ligands and cofactors for bacterial homologs.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37647406 · DOI · PMC10468136

Structural ontogeny of protein-protein interactions [4]

Research article Yang A, Jiang H, Jude KM, Akpinaroglu D, Allenspach S, Li AJ, et al. · · 2026 · cited 5 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Understanding how protein binding sites evolve interactions with other proteins could hold clues to targeting "undruggable" surfaces.
Closing excerpt
Our results suggest that "silent" surfaces have a shallower energy landscape than natural binding sites, disfavoring tight binding, likely owing to evolutionary counterselection.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (124 words)

Understanding how protein binding sites evolve interactions with other proteins could hold clues to targeting "undruggable" surfaces. We used synthetic coevolution to engineer new interactions between naïve surfaces, simulating the de novo formation of protein complexes. We isolated seven distinct structural families of protein Z-domain complexes and found that synthetic complexes explore multiple shallow energy wells through ratchet-like docking modes, whereas complexes formed by natural binding sites converged in a deep energy well with a relatively fixed geometry. Epistasis analysis of a machine learning-estimated fitness landscape revealed "seed" contacts between binding partners that anchored the earliest stages of encounter complex formation. Our results suggest that "silent" surfaces have a shallower energy landscape than natural binding sites, disfavoring tight binding, likely owing to evolutionary counterselection.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 41678610 · DOI · PMC12904254

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article Small molecules, big targets: drug discovery faces the protein-protein interaction challenge [5]
    Scott DE, Bayly AR, Abell C, Skidmore J · Nat Rev Drug Discov · 2016 · cited 868 times (Europe PMC)
  • Research article The pyruvate dehydrogenase complexes: structure-based function and regulation [6]
    Patel MS, Nemeria NS, Furey W, Jordan F · J Biol Chem · 2014 · cited 530 times (Europe PMC)
  • Research article Structure-Based Design of Inhibitors of Protein-Protein Interactions: Mimicking Peptide Binding Epitopes [7]
    Pelay-Gimeno M, Glas A, Koch O, Grossmann TN · Angew Chem Int Ed Engl · 2015 · cited 508 times (Europe PMC)
  • Research article Designed ankyrin repeat proteins (DARPins): binding proteins for research, diagnostics, and therapy [8]
    Plückthun A · Annu Rev Pharmacol Toxicol · 2015 · cited 478 times (Europe PMC)
  • Research article The cullin-RING ubiquitin-protein ligases [9]
    Hua Z, Vierstra RD · Annu Rev Plant Biol · 2011 · cited 400 times (Europe PMC)
  • Research article WD40 repeat domain proteins: a novel target class? [10]
    Schapira M, Tyers M, Torrent M, Arrowsmith CH · Nat Rev Drug Discov · 2017 · cited 287 times (Europe PMC)

References

  1. Beltran A, Jiang X, Shen Y, Lehner B. Site-saturation mutagenesis of 500 human protein domains. . 2025. PMID 39779847 · DOI · PMC11754108
  2. Varga JK, Ovchinnikov S, Schueler-Furman O. actifpTM: a refined confidence metric of AlphaFold2 predictions involving flexible regions. . 2025. PMID 40080667 · DOI · PMC11925850
  3. Xing J, Gumerov VM, Zhulin IB. Origin and functional diversification of PAS domain, a ubiquitous intracellular sensor. . 2023. PMID 37647406 · DOI · PMC10468136
  4. Yang A, Jiang H, Jude KM, Akpinaroglu D, Allenspach S, Li AJ, et al.. Structural ontogeny of protein-protein interactions. . 2026. PMID 41678610 · DOI · PMC12904254
  5. Scott DE, Bayly AR, Abell C, Skidmore J. Small molecules, big targets: drug discovery faces the protein-protein interaction challenge. Nat Rev Drug Discov. 2016. PMID 27050677 · DOI
  6. Patel MS, Nemeria NS, Furey W, Jordan F. The pyruvate dehydrogenase complexes: structure-based function and regulation. J Biol Chem. 2014. PMID 24798336 · DOI · PMC4059105
  7. Pelay-Gimeno M, Glas A, Koch O, Grossmann TN. Structure-Based Design of Inhibitors of Protein-Protein Interactions: Mimicking Peptide Binding Epitopes. Angew Chem Int Ed Engl. 2015. PMID 26119925 · DOI · PMC4557054
  8. Plückthun A. Designed ankyrin repeat proteins (DARPins): binding proteins for research, diagnostics, and therapy. Annu Rev Pharmacol Toxicol. 2015. PMID 25562645 · DOI
  9. Hua Z, Vierstra RD. The cullin-RING ubiquitin-protein ligases. Annu Rev Plant Biol. 2011. PMID 21370976 · DOI
  10. Schapira M, Tyers M, Torrent M, Arrowsmith CH. WD40 repeat domain proteins: a novel target class?. Nat Rev Drug Discov. 2017. PMID 29026209 · DOI · PMC5975957

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-29.

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Related research topics

  • Catalytic DomainMolecular Biology
  • Protein ConformationMolecular Biology
  • Amino Acid SequenceMolecular Biology
  • Binding SitesMolecular Biology
  • Clustered Regularly Interspaced Short Palindromic RepeatsMolecular Biology
  • Consensus SequenceMolecular Biology

All 100 articles in Molecular Biology →

At a glance

82,533
PubMed records on this heading
26%
published since 2020
5,649
review articles
17
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-29; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • Catalytic DomainMolecular Biology
  • Protein ConformationMolecular Biology
  • Amino Acid SequenceMolecular Biology
  • Binding SitesMolecular Biology
  • Clustered Regularly Interspaced Short Palindromic RepeatsMolecular Biology

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