Affinage

WAPL

Wings apart-like protein homolog · UniProt Q7Z5K2

Round 2 corrected
Length
1190 aa
Mass
132.9 kDa
Annotated
2026-04-28
130 papers in source corpus 14 papers cited in narrative 14 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

WAPL is the principal cohesin release factor that drives dynamic turnover of the cohesin ring on chromatin throughout the cell cycle, governing sister chromatid resolution, chromosome loop architecture, and cell-type-specific gene regulation. WAPL uses conserved FGF and YSR motifs to bind PDS5 and the cohesin regulatory subunits SA1/SA2, promoting opening of the Smc3–Scc1 DNA exit gate to release cohesin from DNA; this activity is counteracted in S/G2 by sororin (which displaces WAPL from PDS5 after cohesin acetylation), at centromeres by SGO1-PP2A (which dephosphorylates sororin to exclude WAPL), and by Haspin kinase (which phosphorylates WAPL's YSR motif to block PDS5B binding) (PMID:17112726, PMID:21111234, PMID:23242214, PMID:29138236). WAPL-mediated cohesin turnover limits the processivity of cohesin-driven loop extrusion, thereby setting chromatin loop size and enforcing CTCF boundary directionality genome-wide (PMID:28475897, PMID:29217591). Paradoxically, the free cohesin pool generated by WAPL-driven release and reload is required for cohesin loading at cell-type-specific enhancers and promoters, such that WAPL ablation stabilizes cohesin globally yet depletes it from regulatory elements, impairing gene expression and differentiation (PMID:33318687).

Mechanistic history

Synthesis pass · year-by-year structured walk · 7 steps
  1. 2006 High

    Identification of WAPL as a stoichiometric cohesin-binding protein that promotes cohesin release from chromosomes resolved the longstanding question of how cohesin is removed from chromosome arms during mitotic prophase.

    Evidence Reciprocal Co-IP from HeLa nuclear extracts, in vitro reconstitution of WAPL–SA/PDS5 ternary complex, siRNA depletion and overexpression phenotypes; replicated simultaneously by two independent labs using FRAP to show increased cohesin residence time upon WAPL depletion

    PMID:17112726 PMID:17113138

    Open questions at the time
    • Mechanism of ring opening by WAPL not determined
    • Whether WAPL acts catalytically or stoichiometrically on individual cohesin rings unknown
    • Chromatin context (loop architecture) not yet explored
  2. 2009 High

    Discovery that cohesin acetylation (by ESCO1/ESCO2) renders cohesin resistant to WAPL, and that WAPL's FGF motifs mediate its functional interactions with PDS5, established the molecular logic of how cohesion is stabilized after DNA replication.

    Evidence FGF motif mutagenesis and Xenopus egg extract reconstitution; single-molecule DNA fiber analysis showing WAPL/PDS5A removal rescues replication defects of acetylation-deficient cells; genetic epistasis with Sgo1

    PMID:19696148 PMID:19907496

    Open questions at the time
    • Structural basis of FGF motif–PDS5 interaction not resolved
    • Direct demonstration of ring opening at the Smc3–Scc1 interface not yet achieved
  3. 2010 High

    The finding that sororin displaces WAPL from PDS5 to maintain cohesion, and that sororin becomes dispensable when WAPL is absent, established WAPL as the direct target of the cohesion-maintenance pathway downstream of cohesin acetylation.

    Evidence Co-IP competition assays and siRNA double-depletion epistasis (WAPL/sororin) in vertebrate cells

    PMID:21111234

    Open questions at the time
    • Structural detail of sororin–PDS5 interface displacing WAPL not determined
    • Whether sororin fully occludes WAPL binding or acts allosterically unclear
  4. 2012 High

    Two studies revealed how centromeric cohesin is specifically protected from WAPL: SGO1-PP2A dephosphorylates sororin at centromeres to exclude WAPL, placing WAPL exclusion as the key mechanism; separately, yeast genetics showed WAPL also restricts interphase chromosome condensation independent of cohesion.

    Evidence Phosphomimetic and non-phosphorylatable sororin mutants with forced SGO1 targeting; yeast wapl-null condensation and cohesion assays

    PMID:23219725 PMID:23242214

    Open questions at the time
    • Whether condensation role in yeast is conserved in vertebrates not established
    • Full spectrum of WAPL post-translational regulation at centromeres unknown
  5. 2017 High

    Genome-wide chromatin conformation studies established that WAPL limits cohesin-mediated loop extrusion: WAPL depletion leads to extended loops, loss of CTCF boundary directionality, and 'vermicelli' chromosomes, while Haspin kinase was shown to directly phosphorylate WAPL's YSR motif to inhibit its PDS5B binding at centromeres.

    Evidence Auxin-inducible degron depletion with Hi-C, 4C-seq, and ChIP-seq in human cells; in vitro Haspin kinase assays and phospho-mimetic WAPL mutants; super-resolution microscopy

    PMID:28475897 PMID:29138236 PMID:29217591

    Open questions at the time
    • Direct visualization of loop extrusion processivity change upon WAPL removal not achieved
    • Whether WAPL release occurs during active extrusion or only at stalled complexes unclear
    • Full set of kinases regulating WAPL activity beyond Haspin and CDK not mapped
  6. 2020 High

    WAPL-driven cohesin turnover was shown to be paradoxically required for cell-type-specific cohesin loading at enhancers and promoters, linking the cohesin release factor to gene regulation and differentiation through a reload mechanism dependent on pioneer transcription factors.

    Evidence WAPL knockout in mouse embryonic stem cells with ChIP-seq, Hi-C, and RNA-seq; OCT4/SOX2 co-occupancy analysis

    PMID:33318687

    Open questions at the time
    • Whether the reload mechanism operates through the same NIPBL/MAU2 loader at all cell-type-specific sites unresolved
    • Kinetics of cohesin reload after WAPL-mediated release not measured in real time
  7. 2022 High

    Acute (3-hour) WAPL depletion was found to perturb TAD structure without significantly affecting enhancer–promoter contacts or transcription, suggesting the transcriptional effects of WAPL loss arise from chronic redistribution of cohesin rather than immediate loop changes.

    Evidence Auxin-inducible degron acute depletion with Micro-C, TT-seq nascent transcription, and live-cell single-molecule imaging in mESCs

    PMID:36471071

    Open questions at the time
    • Time course between acute and chronic WAPL depletion effects not fully characterized
    • Whether transcription factor target-search facilitation by cohesin depends on WAPL-driven dynamics specifically or total cohesin levels remains ambiguous

Open questions

Synthesis pass · forward-looking unresolved questions
  • A complete structural and kinetic understanding of how WAPL opens the cohesin ring in real time — including whether WAPL acts during active loop extrusion or only at stalled/paused complexes — remains unresolved, as does the full regulatory code of WAPL post-translational modifications beyond Haspin phosphorylation.
  • No high-resolution cryo-EM or crystal structure of WAPL engaged with an intact cohesin ring
  • In vivo single-molecule measurement of WAPL-mediated release kinetics during loop extrusion not performed
  • Comprehensive phosphoproteomics of WAPL across the cell cycle lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 6
Localization
GO:0005694 chromosome 6 GO:0005654 nucleoplasm 3
Pathway
R-HSA-1640170 Cell Cycle 6 R-HSA-4839726 Chromatin organization 4 R-HSA-74160 Gene expression (Transcription) 2
Complex memberships
cohesin–WAPL–PDS5 release complex

Evidence

Reading pass · 14 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 Human WAPL is a cohesin-binding protein that directly associates with the regulatory subunits of cohesin (SA1/SA2 and Pds5) and is required for timely release of cohesin from chromosome arms during mitotic prophase; Wapl depletion causes retention of cohesin on chromosomes and poorly resolved sister chromatids, while overexpression causes premature sister chromatid separation. In vitro reconstitution showed Wapl forms a stoichiometric ternary complex with cohesin regulatory subunits, inhibiting cohesin's ability to interact with chromatin. Co-immunoprecipitation from HeLa nuclear extracts, siRNA depletion with mitotic phenotype analysis, in vitro reconstitution of ternary complex, overexpression studies Current biology : CB High 17112726
2006 WAPL controls the dynamic association of cohesin with chromatin throughout the cell cycle; Wapl depletion blocks cohesin dissociation from chromosomes during early mitosis, prevents sister chromatid resolution until anaphase, and increases cohesin residence time on chromatin in interphase. WAPL is associated with cohesin throughout the cell cycle. siRNA depletion in HeLa cells, live-cell imaging, FRAP to measure cohesin residence time, immunofluorescence Cell High 17113138
2009 WAPL and PDS5 directly modulate conformational changes of cohesin to promote its dissociation from chromatin during prophase; FGF motifs in Wapl coordinate its physical and functional interactions with Pds5 and cohesin subunits. Wapl depletion from Xenopus egg extracts severely impairs sister chromatid resolution, and human Wapl rescues these defects in a defined early-mitosis time window. Sgo1 antagonizes Wapl-Pds5 to stabilize cohesin on chromosome arms. Xenopus egg extract in vitro system, immunodepletion and rescue, FGF motif mutagenesis, co-immunoprecipitation Genes & development High 19696148
2009 Cohesin acetylation of SMC3 (by ESCO1/ESCO2) counteracts WAPL and PDS5A's destabilizing activity on cohesin; unacetylated cohesin engages WAPL and PDS5A in a hyperstable interaction that impedes replication fork progression, and removal of either WAPL or PDS5A rescues slow replication forks in cells lacking ESCO1, ESCO2, or CTF18-RFC. Single-molecule DNA fiber analysis, siRNA depletion, SMC3 acetylation-deficient mutants, double-depletion epistasis Nature High 19907496
2010 Sororin maintains sister chromatid cohesion by inhibiting WAPL's ability to dissociate cohesin from DNA; DNA replication and cohesin acetylation promote Sororin binding to cohesin, whereupon Sororin displaces WAPL from its binding partner PDS5. In the absence of WAPL, Sororin becomes dispensable for cohesion. Co-immunoprecipitation, siRNA double-depletion epistasis (Wapl/Sororin), binding competition assays in vertebrate cells Cell High 21111234
2007 Sororin is required for the stable chromatin-bound population of cohesin in G2 phase; it interacts with chromatin-bound cohesin and functions to establish or maintain cohesion, operating in the same pathway as WAPL regulation. siRNA depletion, chromatin fractionation, cell-cycle-staged analysis of cohesin stability Current biology : CB Medium 17349791
2012 Budding yeast Wapl acts as a negative regulator of cohesion maintenance in G2 (not during replication fork progression); cohesin acetylation renders cohesin Wapl-resistant from S phase onward. In the absence of Wapl, chromosome condensation is increased in both interphase and mitosis, revealing a role for Wapl in adjusting chromosome condensation status independent of sister chromatid cohesion. Yeast genetics, cohesion assays, condensation measurements, cell-cycle-staged experiments in wapl-null and eco1 mutants Current biology : CB High 23219725
2012 SGO1-PP2A protects centromeric cohesin from WAPL during mitosis; CDK-mediated phosphorylation of SGO1 enables its direct binding to cohesin, recruiting PP2A which dephosphorylates sororin on PDS5-bound cohesin, thereby excluding WAPL from the centromeric cohesin complex. Expression of non-phosphorylatable sororin bypasses the requirement for SGO1-PP2A, placing WAPL's exclusion as the key mechanism. Co-immunoprecipitation, phosphomimetic and non-phosphorylatable sororin mutants, CDK inhibition, epistasis experiments Nature cell biology High 23242214
2017 WAPL is a cohesin release factor that restricts chromatin loop extension; the duration with which cohesin embraces DNA (controlled by WAPL) determines loop size. WAPL also prevents looping between incorrectly oriented CTCF sites. The SCC2/SCC4 loading complex promotes loop extension, and balanced activity of SCC2/SCC4 and WAPL is required for correct chromosome structure. WAPL depletion by auxin-inducible degron and siRNA in human cells, Hi-C, 4C-seq, CTCF ChIP-seq, live-cell cohesin dynamics Cell High 28475897
2017 WAPL and its PDS5 binding partners control the length of chromatin loops genome-wide; in the absence of WAPL and PDS5 proteins, cohesin forms extended loops (passing CTCF sites), accumulates in axial chromosomal positions called 'vermicelli', and condenses chromosomes. PDS5 proteins are also required for CTCF boundary function. These results support the loop extrusion model where WAPL releases cohesin to limit loop size. Auxin-inducible degron depletion of WAPL and PDS5 in HCT116 cells, Hi-C, ChIP-seq, immunofluorescence, super-resolution microscopy The EMBO journal High 29217591
2017 Haspin kinase antagonizes WAPL at centromeres through a kinase-dependent mechanism: the C-terminal kinase domain of Haspin binds and phosphorylates the YSR motif of WAPL, directly inhibiting the YSR motif-dependent interaction of WAPL with PDS5B, thereby blocking cohesin release at centromeres. Phospho-mimetic mutation in WAPL-YSR prevents WAPL from binding PDS5B and releasing cohesin. In vitro kinase assays, co-immunoprecipitation, phospho-mimetic and binding-deficient mutants, forced centromere targeting of Haspin kinase domain, Haspin inhibitor treatment EMBO reports High 29138236
2017 Pds5 and WAPL open the Smc3-Scc1 interface (DNA exit gate) of the cohesin ring to release DNA; a model proposes that Pds5, WAPL, and SA1/2 form a rigid scaffold docking on Scc1, anchoring Scc1-N to the Smc1 ATPase head, with ATP-driven relative movements between Smc1-3 ATPase heads disrupting the Smc3-Scc1 interface. WAPL's FGF motifs are critical for this interaction. Structural modeling, review of biochemical evidence including ATP hydrolysis requirements, FGF motif function, and DNA exit gate studies BioEssays : news and reviews in molecular, cellular and developmental biology Medium 28220956
2020 WAPL creates a pool of free cohesin through cohesin turnover (release and reload cycle), which is required for cohesin to bind cell-type-specific sites enriched at enhancers and promoters. Paradoxically, WAPL ablation stabilizes cohesin binding genome-wide but depletes it from cell-type-specific regions, causing loss of promoter-enhancer loops, gene expression, and differentiation. Cohesin loading at cell-type-specific sites depends on pioneer transcription factors OCT4 and SOX2. WAPL knockout in mouse embryonic stem cells, ChIP-seq, Hi-C, chromosome conformation capture, RNA-seq, OCT4/SOX2 ChIP-seq Nature genetics High 33318687
2022 Acute depletion of WAPL (3 hours) does not significantly affect enhancer-promoter (E-P) interactions or transcription, despite substantially perturbing TAD structure; live-cell single-molecule imaging revealed that cohesin depletion (but not WAPL depletion per se) reduces transcription factor binding to chromatin, suggesting cohesin facilitates TF target search efficiency rather than directly maintaining E-P loops. Auxin-inducible degron acute depletion (3h) of WAPL, CTCF, cohesin in mESCs; high-resolution Micro-C; nascent transcript profiling (TT-seq); live-cell single-molecule imaging of TF binding Nature genetics High 36471071

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Global, in vivo, and site-specific phosphorylation dynamics in signaling networks. Cell 2861 17081983
2005 Myeloperoxidase: friend and foe. Journal of leukocyte biology 1654 15689384
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2004 Large-scale characterization of HeLa cell nuclear phosphoproteins. Proceedings of the National Academy of Sciences of the United States of America 1159 15302935
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2019 Neuroinflammation: friend and foe for ischemic stroke. Journal of neuroinflammation 1109 31291966
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2012 A census of human soluble protein complexes. Cell 689 22939629
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2017 Topologically associating domains and chromatin loops depend on cohesin and are regulated by CTCF, WAPL, and PDS5 proteins. The EMBO journal 612 29217591
2017 The Cohesin Release Factor WAPL Restricts Chromatin Loop Extension. Cell 589 28475897
2006 Wapl controls the dynamic association of cohesin with chromatin. Cell 498 17113138
2020 Exercise-induced oxidative stress: Friend or foe? Journal of sport and health science 438 32380253
2015 A Dynamic Protein Interaction Landscape of the Human Centrosome-Cilium Interface. Cell 433 26638075
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
2016 Inflammatory Response in the CNS: Friend or Foe? Molecular neurobiology 421 27889895
2010 Systematic analysis of human protein complexes identifies chromosome segregation proteins. Science (New York, N.Y.) 421 20360068
2006 Human Wapl is a cohesin-binding protein that promotes sister-chromatid resolution in mitotic prophase. Current biology : CB 356 17112726
2021 A proximity-dependent biotinylation map of a human cell. Nature 339 34079125
2006 Cardiac fibroblasts: friend or foe? American journal of physiology. Heart and circulatory physiology 337 16617141
2008 The osteoclast: friend or foe? Annual review of pathology 323 18039135
2013 Genome-wide meta-analysis identifies new susceptibility loci for migraine. Nature genetics 314 23793025
2010 Sororin mediates sister chromatid cohesion by antagonizing Wapl. Cell 305 21111234
2022 Enhancer-promoter interactions and transcription are largely maintained upon acute loss of CTCF, cohesin, WAPL or YY1. Nature genetics 302 36471071
2019 More friend than foe: the emerging role of neutrophils in tissue repair. The Journal of clinical investigation 296 31205028
2004 Phosphoproteomic analysis of the developing mouse brain. Molecular & cellular proteomics : MCP 291 15345747
2018 Nutrients and Oxidative Stress: Friend or Foe? Oxidative medicine and cellular longevity 244 29643982
2016 ICOS Co-Stimulation: Friend or Foe? Frontiers in immunology 234 27559335
2018 Ubiquitination: Friend and foe in cancer. The international journal of biochemistry & cell biology 223 29864543
2017 Optimized fragmentation schemes and data analysis strategies for proteome-wide cross-link identification. Nature communications 221 28524877
2000 Lentiviral vectors: turning a deadly foe into a therapeutic agent. Gene therapy 211 10680011
2009 Cohesin acetylation speeds the replication fork. Nature 208 19907496
2013 PRP19 transforms into a sensor of RPA-ssDNA after DNA damage and drives ATR activation via a ubiquitin-mediated circuitry. Molecular cell 204 24332808
2018 An AP-MS- and BioID-compatible MAC-tag enables comprehensive mapping of protein interactions and subcellular localizations. Nature communications 201 29568061
2009 The transcriptional corepressor CtBP: a foe of multiple tumor suppressors. Cancer research 194 19155295
2007 Sororin is required for stable binding of cohesin to chromatin and for sister chromatid cohesion in interphase. Current biology : CB 191 17349791
2013 Autophagy: cancer's friend or foe? Advances in cancer research 183 23768510
1996 Prediction of the coding sequences of unidentified human genes. VI. The coding sequences of 80 new genes (KIAA0201-KIAA0280) deduced by analysis of cDNA clones from cell line KG-1 and brain. DNA research : an international journal for rapid publication of reports on genes and genomes 178 9039502
2009 Releasing cohesin from chromosome arms in early mitosis: opposing actions of Wapl-Pds5 and Sgo1. Genes & development 174 19696148
2009 Luminal sulfide and large intestine mucosa: friend or foe? Amino acids 168 20020161
2006 Immunomodulatory effects of caffeine: friend or foe? Pharmacology & therapeutics 165 16540173
2015 Hydrogen sulfide in cancer: Friend or foe? Nitric oxide : biology and chemistry 163 26297862
2000 Dinucleoside polyphosphates-friend or foe? Pharmacology & therapeutics 163 11007992
2012 Phosphorylation-enabled binding of SGO1-PP2A to cohesin protects sororin and centromeric cohesion during mitosis. Nature cell biology 162 23242214
2020 Synthetic Lethal and Resistance Interactions with BET Bromodomain Inhibitors in Triple-Negative Breast Cancer. Molecular cell 159 32416067
2017 AGEs, RAGEs and s-RAGE; friend or foe for cancer. Seminars in cancer biology 155 28712719
2015 De novo lipogenesis in metabolic homeostasis: More friend than foe? Molecular metabolism 145 25973385
2009 Ubiquitin-mediated proteolysis of HuR by heat shock. The EMBO journal 142 19322201
2005 A scan of chromosome 10 identifies a novel locus showing strong association with late-onset Alzheimer disease. American journal of human genetics 137 16385451
2020 G-Quadruplexes at Telomeres: Friend or Foe? Molecules (Basel, Switzerland) 135 32823549
2020 WAPL maintains a cohesin loading cycle to preserve cell-type-specific distal gene regulation. Nature genetics 135 33318687
2016 Gamma-glutamyltransferase-friend or foe within? Liver international : official journal of the International Association for the Study of the Liver 132 27512925
2009 Friend or foe? Resolving the impact of glial responses in glaucoma. Journal of glaucoma 126 19525723
2017 STAT1 in cancer: friend or foe? Discovery medicine 124 28950072
2014 Protein mistranslation: friend or foe? Trends in biochemical sciences 123 25023410
2018 Crosstalk between phosphorylation and O-GlcNAcylation: friend or foe. The FEBS journal 118 29717537
2012 Adipose tissue: friend or foe? Nature reviews. Cardiology 104 23149834
2018 NFKB1 and Cancer: Friend or Foe? Cells 100 30205516
2017 Immune System, Friend or Foe of Oncolytic Virotherapy? Frontiers in oncology 99 28589085
2002 Nitric oxide in liver diseases: friend, foe, or just passerby? Annals of the New York Academy of Sciences 99 12076981
2012 Budding yeast Wapl controls sister chromatid cohesion maintenance and chromosome condensation. Current biology : CB 98 23219725
2018 Oxygen in the critically ill: friend or foe? Current opinion in anaesthesiology 97 29334496
2020 Zinc in the Brain: Friend or Foe? International journal of molecular sciences 85 33255662
2016 Emerging role of HMGB1 in lung diseases: friend or foe. Journal of cellular and molecular medicine 75 28039939
2013 IL-17 in lung disease: friend or foe? Thorax 74 23604380
2018 PGC1α: Friend or Foe in Cancer? Genes 73 29361779
2020 Platelets in Neurodegenerative Conditions-Friend or Foe? Frontiers in immunology 72 32431701
2008 Amyloids: friend or foe? Journal of Alzheimer's disease : JAD 72 18487849
2019 SOX11: friend or foe in tumor prevention and carcinogenesis? Therapeutic advances in medical oncology 69 31210798
2021 Pyroptosis in Cancer: Friend or Foe? Cancers 68 34298833
2022 The Role of SLC7A11 in Cancer: Friend or Foe? Cancers 64 35804831
2016 Cancer and necroptosis: friend or foe? Cellular and molecular life sciences : CMLS 63 27048810
2019 Friend or foe, the role of EGR-1 in cancer. Medical oncology (Northwood, London, England) 62 31748910
2018 Microbe Profile: Mycobacterium tuberculosis: Humanity's deadly microbial foe. Microbiology (Reading, England) 62 29465344
2015 Lithium in the Kidney: Friend and Foe? Journal of the American Society of Nephrology : JASN 61 26577775
2021 NF-κB in Cancer Immunity: Friend or Foe? Cells 60 33572260
2022 The Role of NRF2 in Bone Metabolism - Friend or Foe? Frontiers in endocrinology 57 35282459
2010 Cannabinoids and bone: friend or foe? Calcified tissue international 57 20532878
2018 MDR1 in immunity: friend or foe? Oncoimmunology 55 30524890
2015 Phosphoinositide 3-kinase: friend and foe in cardiovascular disease. Frontiers in pharmacology 54 26321955
2009 Magnesium and tumors: ally or foe? Cancer treatment reviews 54 19203841
2016 MTH1 as a nucleotide pool sanitizing enzyme: Friend or foe? Free radical biology & medicine 52 27833032
2022 Microglia: Friend and foe in tauopathy. Progress in neurobiology 47 35714860
2017 A kinase-dependent role for Haspin in antagonizing Wapl and protecting mitotic centromere cohesion. EMBO reports 47 29138236
2001 NF-kappaB in transplantation: friend or foe? Transplant infectious disease : an official journal of the Transplantation Society 47 11844153
2016 Alternative Polyadenylation: Another Foe in Cancer. Molecular cancer research : MCR 46 27075335
2013 Wnt signaling in remyelination in multiple sclerosis: friend or foe? Molecular neurobiology 46 24243343
2018 Role of Autophagy in Proteostasis: Friend and Foe in Cardiac Diseases. Cells 43 30572675
2011 Calcineurin inhibitors in kidney transplantation: friend or foe? Current opinion in nephrology and hypertension 43 21885969
2014 Stat3: friend or foe in colitis and colitis-associated cancer? Inflammatory bowel diseases 42 25185686
2020 Tim-4 in Health and Disease: Friend or Foe? Frontiers in immunology 41 32300343
2020 p62: Friend or Foe? Evidences for OncoJanus and NeuroJanus Roles. International journal of molecular sciences 41 32708719
2020 Malignancy and IFITM3: Friend or Foe? Frontiers in oncology 41 33364194
2014 Mitochondrial stress: balancing friend and foe. Experimental gerontology 41 24603155
2023 ARID1A in cancer: Friend or foe? Frontiers in oncology 40 36890819
2022 TLR9: A friend or a foe. Life sciences 40 35963302
2022 Browning Epicardial Adipose Tissue: Friend or Foe? Cells 39 35326442
2021 Post-stroke Neurogenesis: Friend or Foe? Frontiers in cell and developmental biology 39 33834025
2020 The Aryl Hydrocarbon Receptor in Asthma: Friend or Foe? International journal of molecular sciences 39 33233810
2018 A Friend or Foe: Calcineurin across the Gamut of Neurological Disorders. ACS central science 39 30062109
2014 Aldehyde dehydrogenase 1A1: friend or foe to female metabolism? Nutrients 39 24594504
2021 Natural Killer Cells: Friend or Foe in Metabolic Diseases? Frontiers in immunology 38 33717101
2021 Autophagy: A Friend or Foe in Allergic Asthma? International journal of molecular sciences 38 34204710
2021 Neuronal Mitophagy: Friend or Foe? Frontiers in cell and developmental biology 37 33537304
2018 KISS1/KISS1R in Cancer: Friend or Foe? Frontiers in endocrinology 37 30123188
2024 Friend or foe: The role of stress granule in neurodegenerative disease. Neuron 36 38744273
2023 TP53 Mutation in Acute Myeloid Leukemia: An Old Foe Revisited. Cancers 36 37835510
2021 TNFα and Immune Checkpoint Inhibition: Friend or Foe for Lung Cancer? International journal of molecular sciences 36 34445397
2023 The intratumoral microbiota: friend or foe? Trends in cancer 34 37061408
2022 The chaperone Clusterin in neurodegeneration-friend or foe? BioEssays : news and reviews in molecular, cellular and developmental biology 34 35521968
2019 LRRK2 in Infection: Friend or Foe? ACS infectious diseases 34 30915830
2007 COX-2: friend or foe? Current pharmaceutical design 34 17584101
2017 The Top1 paradox: Friend and foe of the eukaryotic genome. DNA repair 33 28641942
2021 PERK signaling pathway in bone metabolism: Friend or foe? Cell proliferation 32 33615575
2018 Neutrophil extracellular traps in vasculitis, friend or foe? Current opinion in rheumatology 31 28957962
2017 Releasing the cohesin ring: A rigid scaffold model for opening the DNA exit gate by Pds5 and Wapl. BioEssays : news and reviews in molecular, cellular and developmental biology 29 28220956
2016 Potassium: friend or foe? Pediatric nephrology (Berlin, Germany) 29 27194424
2022 Nutritional Sensor REDD1 in Cancer and Inflammation: Friend or Foe? International journal of molecular sciences 28 36077083
2020 The proteasome: friend and foe of mitochondrial biogenesis. FEBS letters 28 33249599
2024 Friend or foe: Lactate in neurodegenerative diseases. Ageing research reviews 27 39127445
2022 APOBEC3: Friend or Foe in Human Papillomavirus Infection and Oncogenesis? Annual review of virology 27 35671565
2021 Inflammasomes in dendritic cells: Friend or foe? Immunology letters 27 33848562
2018 Molecular Regulation of Carcinogenesis: Friend and Foe. Toxicological sciences : an official journal of the Society of Toxicology 27 30053205
2014 NK cells after transplantation: friend or foe. Immunologic research 27 24522700
2024 Folic acid: friend or foe in cancer therapy. The Journal of international medical research 26 38229460
2017 The spoilage yeast Zygosaccharomyces bailii: Foe or friend? Yeast (Chichester, England) 26 28556381
2016 Human endogenous retroviruses: friend or foe? APMIS : acta pathologica, microbiologica, et immunologica Scandinavica 26 26818257
2006 Macrophage and dengue virus: friend or foe? The Indian journal of medical research 26 16926454