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Nucleus

The compartment that holds the genome, wrapped in a double envelope whose pores are the only way in or out.

15 pictures and 65 papers. Its organelle page: Nucleus.

Gallery

Panel of a six-part figure, cropped to this panel and otherwise unaltered, which CC BY allows with attribution. Human astrocyte differentiated from induced pluripotent stem cells of a healthy donor, 25 days in vitro. The authors describe it as a typical cell nucleus with an eccentrically located nucleolus, marked by the yellow arrow, and shallow invaginations of the envelope, marked by the red arrow. Scale bar 2 micrometres. The invaginations are worth noticing: the envelope shown as a smooth sphere on most diagrams is not smooth here.

Kutukova, Ivanov, Novosadova et al 2026, Modern Technologies in Medicine · CC BY 4.0

HeLa cells stained with Hoechst 33258 stain.

TenOfAllTrades at English Wikipedia · Public domain

Bridging scales of resolution by combining variety of imaging modalities.

Kechagia, Medalia, Uversky et al 2026, Nucleus · CC BY 4.0

(A) Double fluorescent labeling of the 20S proteasome and propidium iodide (PI) in mechanically dissociated supraoptic neuron (SON) cell bodies from a control rat. Note the high nuclear concentration of the 20S proteasome. Structures, enriched in rRNAs, namely nucleoli and polyribosome-rich cytoplasmic areas (Nissl substance), appear intensely contrasted with PI. (B,C) Immunolabeling illustrating the nuclear and cytoplasmic distribution of the regulatory 19S and catalytic 20S proteasomes in rat trigeminal ganglion neurons (TGN). Note the higher concentration of proteasomes within the nucleus, the existence of nucleoplasmic domains with reduced proteasome signals (white asterisks), and the absence of both 19S and 20S proteasomes within the nucleolus (No). (D) Immunogold electron microscopy detecting the 20S proteasome and showing its localization in euchromatin regions (red arrows). The nucleolus (No) lacks proteasome signal.

Lafarga, Berciano, Rueda et al 2026, Frontiers in Neuroanatomy. Figure 1 · CC BY 4.0

NPM1 colocalizes and interacts with the PRC2.1 subcomplex. (A) Left panels: Subcellular localization of H3K27me3 or H3K9me3 (green), NPM1 (red), and DAPI (blue) in HeLa cells is shown in the representative immunofluorescence (IF) images. Right panels: intensity profiles of NPM1, H3K27me3 and H3K9me3 correspond to the zoomed regions (white rectangle) indicated by white lines crossing the nucleolus in the left panels.

Zhu, Wang, Chen et al 2026, Advanced Science · CC BY 4.0

HP1a condensate properties in resting conditions in IMR90 and HeLa. A Airyscan super resolution images of nuclei IMR90 transiently transfected with HP1a-GFP (green), fixed and stained with anti-H3K9me3 (red) antibodies. Merged image shown in the right panel. Scale bar = 2 um. B HeLa cells stably transfected with HP1a-GFP (green), fixed and stained with antibody against H3K9me3 (red). Merged image showing HP1a condensate distribution.

Hovet, Nahali, Halaburkova et al 2025, Communications Biology · CC BY 4.0

H3K9me3-stained heterochromatin is detached from the nuclear periphery of human cells upon depletion of eight INM proteins and the nuclear lamins. a , The selected set of 12 NE factors and their corresponding categories (lamins: lamin A, lamin C, lamin B1 and lamin B2; INM proteins: LBR, LAP2, emerin, LEMD2, MAN1, SUN1, SUN2 and LAP1) targeted by a combination of siPOOLs to assess heterochromatin tethering at the nuclear periphery by immunofluorescence.

Lewis, Sinigiani, Maziak et al 2026, Nature Cell Biology · CC BY 4.0

The lobulated nuclear phenotype associated with HGPS is dependent on cell division, with arrested cells displaying a distinct nuclear phenotype. (A) Time-lapse imaging of U2OS cells with doxycycline (dox)-inducible GFP-progerin expression. Cells were incubated with 500 ng/ml dox at t=0. Images were acquired every 30 min for 7 h using a spinning-disc confocal microscope. DNA was visualized using NucBlue.

Turkmen, Wang, Frost et al 2026, Journal of Cell Science · CC BY 4.0

The lobulated nuclear phenotype associated with HGPS is dependent on cell division, with arrested cells displaying a distinct nuclear phenotype. (A) Time-lapse imaging of U2OS cells with doxycycline (dox)-inducible GFP-progerin expression. Cells were incubated with 500 ng/ml dox at t=0. Images were acquired every 30 min for 7 h using a spinning-disc confocal microscope. DNA was visualized using NucBlue.

Turkmen, Wang, Frost et al 2026, Journal of Cell Science · CC BY 4.0

Whole-cell TEM overview of a cluster-stage mouse oocyte, its germinal vesicle (nucleus) visible as the large pale circle with a dense nucleolus, boxed regions marking areas magnified elsewhere in the source figure. Scale bar 10 um.

Zollo, Zaffagnini, Haidar et al 2026, Science Advances · CC BY 4.0

Transmission electron micrographs (TEM) of two daughter regenerative cells (rc1 and rc2) after symmetrical division: nu (nucleus), glg (glycogen granules), ld (lipid droplet), m (mitochondria), dc (digestive cells), bm (basal membrane).

Puljas, Bocina, Kevic et al 2026, Insects · CC BY 4.0

A control Ishikawa spheroid cell by TEM, showing an intact plasma membrane, continuous nuclear envelope, and mitochondria with intact cristae. Scale bar visible at lower right.

Bilir, Yildirim, Karalok et al 2026, International Journal of Molecular Sciences · CC BY 4.0

TEM image of a crescent-shaped nucleus in Chara sp

Böhm, Peters, Herburger et al 2026, Planta · CC BY 4.0

Structure of a camel liver hepatocyte. H--hepatocyte; N--nucleus; LD--lipid droplets; M--mitochondria; GER--granular endoplasmic reticulum channels; sinusoidal capillaries (↑). Electron microphotography. (Bactrian camel (Camelus bactrianus, liver hepatocyte); Figure 8single.)

Musina, Lebedeva, Drozdova et al 2025, Animals · CC BY 4.0

Representative confocal images of immunofluorescence-stained sections of human myocardium: lamin B1 (green), DAPI (blue), and sarcomeric alpha-actinin (red). A) cardiomyocyte nuclei. (human; Fig. 2, panel A (cardiomyocyte nuclei IF; rows 142-1193 of figure).)

Ventimiglia, Zelezniak, dos Remedios et al 2026, Journal of Molecular and Cellular Cardiology Plus · CC BY 4.0

Papers

  1. Wang X, Mu X, Fu Y, Dong W, Lu W, Li X, et al. Spatial, single-nucleus and pathological profiling of the invasive front in early hepatocellular carcinoma for characterizing specific leading-edge cell niche and improving recurrence modeling. Int J Biol Sci. 2026;22:8090-8118. PMID: 42807944.
  2. Thompson C, Meier-Stephenson V. Targeting cccDNA for Degradation: Endogenous- and Exogenous-Based Methods of Nuclear-Localized DNA Destruction in the Context of Chronic Hepatitis B Infection. Viruses. 2026;18:929. PMID: 42797762.
  3. Zhao S, Sui S, Guo X, Ji X, Liu T, Yang H. Integrated Single-Nucleus and Epigenomic Profiling Reveals Coordinated Neuro-Glial and Chromatin Remodeling in the Type 2 Diabetic Hypothalamus. Cells. 2026;15:1672. PMID: 42782771.
  4. Qiao Q, Katsumata Y, Lai HY, Lin KZ, Claas SA, Gabitto MI, et al. Distinct single-nucleus RNA-seq changes among non-neuronal cells in ADNC, LATE-NC, and mixed pathologies. Alzheimers Dement. 2026;22:e71810. PMID: 42779047.
  5. Du Q, Qin T, Wang J, Jiang Y, Xu Q, Li X, et al. XPO1-mediated TRIM21 nuclear export reprograms TREM2+ macrophage polarization by targeting IRF3 to augment anti-PD-1 efficacy in small cell lung cancer. Nat Commun. 2026;17:10050. PMID: 42768009.
  6. Gao X, Zhang M, Li L, Lyu H, Liu Z, Wei X, et al. Dynamic Changes of Neutrophil Nuclear Membrane CD63 as a Potential Biomarker for Early Adjunctive Diagnosis and Prognostic Evaluation of Sepsis in Critically Ill Patients: A Prospective Cohort Study. MedComm (2020). 2026;7:e71007. PMID: 42756472.
  7. Ojetunde M, Ma L, Goodney HG, Stevens AM, Mizener A, Pistilli EE, et al. Exploratory single-nucleus multiomics analysis of myeloid cell states associated with neoadjuvant chemotherapy response in pancreatic ductal adenocarcinoma. Clin Transl Med. 2026;16:e70810. PMID: 42735906.
  8. Liao XR, Zheng H, Luo JY, Zheng BX. Single-cell and single-nucleus transcriptomics of the dorsal root ganglion in neuropathic pain: cell-state remodeling and translational prospects. J Transl Med. 2026;24:1126. PMID: 42687187.
  9. Ventimiglia LN, Zelezniak A, Dos Remedios CG, Lal S, Ehler E, De Nicola GF. Lamin B1 loss in human cardiomyocytes is associated with nuclear shape deformation in dilated cardiomyopathy. J Mol Cell Cardiol Plus. 2026;17:100865. PMID: 42668459.
  10. Cui B, Lei M, Liu L, Shu M, Yu F, Yang K, et al. Single-cell chromatin profiling reveals relapse-related priming in pediatric acute myeloid leukemia. Life Sci Alliance. 2026;9:e202603746. PMID: 42637517.
  11. Kokki K, Wowro SJ, Robciuc M, Hallasaari A, Porri L, Kinnunen M, et al. BHLHE40 and ChREBP associate with hepatic enhancer clusters containing PPARα, RXRα, and HNF4 nuclear receptors. Endocrinology. 2026;167:bqag095. PMID: 42626966.
  12. Li Y, Wang K, Shi M, Zhang MQ, Gao J. Sequential-GAM constructs the single-cell geometric 3D genome structure. Nucleic Acids Res. 2026;54:gkag794. PMID: 42598856.
  13. Kong L, Iwabuchi S, Li YY, Murai K, Korai R, Kawaguchi K, et al. Chromatin Accessibility-Guided Targeting Identifies Structurally Constrained Regions in HBV cccDNA and Suppresses Viral Replication. Int J Mol Sci. 2026;27:6980. PMID: 42589635.
  14. Newman AG, Singh PB. HUSH, NEXT PROMPT: Epigenetics and the Nuclear RNA Exosome in Human Aging and Disease. Cells. 2026;15:1378. PMID: 42587787.
  15. Mu C, Ding Y, Weiss A, Rosenfeld S, Li F, Geng X. Analytical Methods and Application of Single-Cell and Single-Nucleus Transcriptomics in the Study of Ischemic Stroke. Biomolecules. 2026;16:1054. PMID: 42509846.
  16. Kunkanjanawan H, Kunkanjanawan T, Khemarangsan V, Suvithayasiri S, Suriyo T, Satayavivad J, et al. Comparative Study of Mesenchymal Stem Cell Identity and Chondrogenic Differentiation Capacity of Human Nucleus Pulposus, Adipose and Bone Marrow Tissues Derived Mesenchymal Stem Cells. Tissue Eng Regen Med. 2026;23:1075-1088. PMID: 42484960.
  17. Oda H, Yonezawa N, Yamagata K. Nuclear Dynamics and Its Timing Regulation Revealed by Live-Cell Imaging. Adv Exp Med Biol. 2026;1517:177-194. PMID: 42455444.
  18. Kreysing JP, Cruz-León S, Betz J, Penzo C, Majtner T, Schreiber M, et al. Molecular architecture of heterochromatin at the nuclear periphery of primary human cells. Nat Commun. 2026;17:5844. PMID: 42399220.
  19. Gigan JP, Garcia-Gonzalez P, Pilotti A, Galliot L, Reynaud A, Ghaffar Y, et al. Single-nuclei UPR profiling by flow cytometry reveals bortezomib resistance mechanisms in multiple myeloma. EMBO Mol Med. 2026;18:3057-3085. PMID: 42373888.
  20. Duan J, Fu Q, Huo Y, Wang C, Shen Y, Zhong M, et al. Critical changes in whole-brain gene networks in response to small-cell lung cancer as revealed by single-nucleus RNA sequencing. Front Immunol. 2026;17:1860628. PMID: 42311671.
  21. Lambertini E, Penolazzi L, Chierici A, Nadalini R, Sief C, Begnozzi F, et al. Participation of the purinergic P2X7 receptor in molecular complexes in the nucleus of human chondrocytes. Cell Mol Life Sci. 2026;83:319. PMID: 42307773.
  22. Cavallo M, Diaf A, Milanesi G, Biggiogera M, Casali C. Resolving Sub-Nuclear Architecture from Compartments to Functional Domains. Int J Mol Sci. 2026;27:4680. PMID: 42278215.
  23. Wang M, Wu X, Wang T, Liu W, Zhang Y, Zhang L, et al. SUMOylation-driven nuclear translocation of HSF2BP alleviates MASLD via COX6A1-dependent mitochondrial reprogramming. Cell Death Dis. 2026;17:718. PMID: 42259788.
  24. Tahasildar J, Shekhar S, Srinath S, Anjana K, Shiragur S, Annegowda VM. Cytomorphometric Study on Micronuclei and Nuclear-Cytoplasmic Variations of Exfoliated Buccal Cells in Tobacco Habituates Using Phase Contrast Microscope. Indian J Dent Res. 2026;37:61-65. PMID: 42241137.
  25. Kechagia Z, Medalia O. High-resolution nuclear cell biology by cryo-electron tomography. Nucleus. 2026;17:2675754. PMID: 42226577.
  26. Okada M, Subramaniam B, Geng B, Jung J, Yu B, Lin F, et al. Single-nucleus profiling of postmortem diffuse midline gliomas identifies mitochondrial biogenesis as a resistance mechanism to imipridone therapy. Neuro Oncol. 2026;28:2248-2262. PMID: 42178382.
  27. Vergara J, Perez-Gallardo C, Velasco R, Martinez D, Badilla D, Freire-Navarrete D, et al. NucVerse3D: generalizable 3D nuclear instance segmentation across heterogeneous microscopy modalities. Sci Rep. 2026;16:22197. PMID: 42141034.
  28. Liu Y, Yun X, Ding W, Li S, Liu H. Targeting the nuclear export receptor exportin-1 in acute myeloid leukaemia: From biology to clinical translation. Clin Transl Med. 2026;16:e70676. PMID: 42071259.
  29. Shirude MB, Devarajan A, Sahu SA, Mukherjee A, Dutta D. HIRA-SETDB1-H3K9me3 axis regulates chromatin architecture in leukemia cells. J Biol Chem. 2026;302:113081. PMID: 42055326.
  30. Wang Y, Hu O, Wu J, Huang S, Lin P, Yan P, et al. Harnessing Metabolic Priming to Engineer Human Nucleus Pulposus Macromass Overcoming Scalability-Phenotype Tradeoff. Cell Prolif. 2026;59:e70215. PMID: 42049200.
  31. Kabbe M, Agirre E, Carlström KE, Dumral , Lor YK, Pohl FB, et al. Single-nucleus epigenomic profiling of the adult human central nervous system unveils epigenetic memory of developmental programs. Nat Neurosci. 2026;29:992-1006. PMID: 41857393.
  32. Casali C, Cavallo M, Diaf A, Milanesi G, Biggiogera M. Now You See Me: Visualizing nuclear complexity by selective staining. Methods. 2026;250:46-54. PMID: 41831750.
  33. Li X, Yao X, Guo C, Liu Y, Liu M, Li J, et al. Nuclear Lipin1 recruits HDAC2 to epigenetically repress SREBP1-dependent lipid synthesis and myelination in hypoxic preterm white matter injury. Stem Cell Reports. 2026;21:102853. PMID: 41825445.
  34. Schol J, Ambrosio L, Ruiz-Fernandez C, Schlagenhof L, Voskamp C, Laagland LT, et al. TIE2-positive cells in the nucleus pulposus with a purpose: the who, what and why. J Biomed Sci. 2026;33:24. PMID: 41772594.
  35. Sun R, Gao C, Lu RO. Tumor-intrinsic chromatin programs enforce immune evasion in glioblastoma. J Clin Invest. 2026;136:e203222. PMID: 41766661.
  36. Khalaji S, Venit T, Lukáčová Z, Fambri V, Shrestha R, Kaluarachchi S, et al. Nuclear Myosin 1 links genomic architecture to adipose tissue remodeling, metabolic inflammation and obesity in mice. Cell Death Dis. 2026;17:270. PMID: 41748556.
  37. Pastinen T, Grundberg E, Bradley T, Honkanen J, Cheung WA, Vuorela A, et al. Evolving epigenomics of immune cells at single-nucleus resolution in children en route to type 1 diabetes. Nat Commun. 2026;17:3168. PMID: 41741442.
  38. Bouwman BA, Zhao L, de Luca KL, Bienko M. Emerging oligo-based imaging methods to study chromatin organization in cells and tissue. Curr Opin Genet Dev. 2026;97:102447. PMID: 41707606.
  39. Natale CF, Messina L, Panzetta V, Saporito S, Menna C, Ventre M, et al. Material-Induced Nuclear Deformation Controls Chromatin Architecture in Adipose Stem Cells. Adv Sci (Weinh). 2026;13:e14458. PMID: 41615150.
  40. Chiou LW, Jhuang YL, Hsu CL, Yuan RH, Cheng YH, Lee CH, et al. Pbrm1 Loss Induces a Permissive Chromatin State for Cholangiocytic Differentiation and Cholangiocarcinoma Formation. Cell Mol Gastroenterol Hepatol. 2026;20:101720. PMID: 41513002.
  41. Renkawitz J, Yesin A, Kroll J, Cabral AT, D'Annunzio S, Thiam HR. Nuclear mechanobiology rules immune cells' functions: from differentiation to cell trafficking and pathogen killing. Nucleus. 2026;17:2590843. PMID: 41495604.
  42. Takeuchi E, Yasumizu Y, Morita J, Ishikawa M, Ogawa K, Motooka D, et al. Single-nucleus multiome shows motor neuron glutamate overactivation in amyotrophic lateral sclerosis. Brain. 2026;149:2480-2494. PMID: 41218062.
  43. Zhu W, Wang W, Wang Y, Tong X, Liu X, Zhang L, et al. Hepatocyte nuclear factor 1 in renal lipid metabolism: molecular mechanisms and therapeutic potentials. Cell Biol Toxicol. 2025;41:157. PMID: 41291157.
  44. Klein K, Burk O, Tremmel R, Buettner FA, Kaestle L, Schroth W, et al. Transcription factors of the Nuclear Factor I (NFI) family control hepatocyte differentiation and cytochrome P450 activity in human liver. Pharmacol Res. 2025;221:107998. PMID: 41106572.
  45. Musina L, Lebedeva A, Drozdova L, Prusakov A, Ponamarev V. Features of the Liver Microstructural Organization of the Bactrian Camel. Animals (Basel). 2025;15:2921. PMID: 41096516.
  46. Zhang L, Xiu C, You H, Chen J. Dual-Lineage Chondrocyte-Like Cells in the Nucleus Pulposus of Aging Intervertebral Discs Are Accelerated by Hedgehog Signaling Inactivation. Aging Cell. 2025;24:e70248. PMID: 40977276.
  47. Abdel-Rasol MA, El-Sayed WM. Nuclear receptors in metabolic, inflammatory, and oncologic diseases: mechanisms, therapeutic advances, and future directions. Eur J Med Res. 2025;30:843. PMID: 40926269.
  48. Sánchez-Román EJ, Villa-Villegas L, Leal-Ortega R, Lira-Jaime LG, Rivas-Ruvalcaba F, Díaz-Ramírez K, et al. "Deep Brain Stimulation of the Ventral Intermediate Nucleus of the Thalamus for Tremor in Polr3a-Related Tremor-ataxia Syndrome: A Two-case Report". Tremor Other Hyperkinet Mov (N Y). 2025;15:42. PMID: 40917817.
  49. Liu Y, Yang H, Ren M, Yu Q, Xu Q, Fu X. Orphan Nuclear Receptors TR2 and TR4 in Erythropoiesis: From Mechanisms to Therapies. Biomolecules. 2025;15:798. PMID: 40563438.
  50. Nair A, Khanna J, Kler J, Ragesh R, Sengupta K. Nuclear envelope and chromatin choreography direct cellular differentiation. Nucleus. 2025;16:2449520. PMID: 39943681.
  51. Kastani IA, Soltani PK, Baltogiannis GG, Christou GA, Bairaktari ET, Kostara CE. Nuclear Magnetic Resonance (NMR)-Based Lipidomics Reveal the Association of Altered Red Blood Cell (RBC) Membrane Lipidome with the Presence and the Severity of Coronary Artery Stenosis. Molecules. 2024;30:36. PMID: 39795094.
  52. Atteberry B, Berman BP, Kelly TK, Cayford J. Understanding the complex chromatin dynamics in primary human neutrophils during PMA-induced NET formation. Front Immunol. 2024;15:1445638. PMID: 39524441.
  53. Rayl ML, Nemetchek MD, Voss AH, Hughes TS. Agonists of the Nuclear Receptor PPARγ Can Produce Biased Signaling. Mol Pharmacol. 2024;106:309-318. PMID: 39443155.
  54. Sladek FM, Apte U, Deol P. Editorial: Hepatocyte nuclear factor 4 alpha - new insights into an old receptor. Front Endocrinol (Lausanne). 2024;15:1491965. PMID: 39387053.
  55. Limone F, Mordes DA, Couto A, Joseph BJ, Mitchell JM, Therrien M, et al. Single-nucleus sequencing reveals enriched expression of genetic risk factors in extratelencephalic neurons sensitive to degeneration in ALS. Nat Aging. 2024;4:984-997. PMID: 38907103.
  56. Puvogel S, Alsema A, North HF, Webster MJ, Weickert CS, Eggen BJL. Single-Nucleus RNA-Seq Characterizes the Cell Types Along the Neuronal Lineage in the Adult Human Subependymal Zone and Reveals Reduced Oligodendrocyte Progenitor Abundance with Age. eNeuro. 2024;11:ENEURO.0246-23.2024. PMID: 38351133.
  57. Luo Z, Wei Z, Zhang G, Chen H, Li L, Kang X. Achilles' Heel-The Significance of Maintaining Microenvironmental Homeostasis in the Nucleus Pulposus for Intervertebral Discs. Int J Mol Sci. 2023;24:16592. PMID: 38068915.
  58. Chaand M, Fiore C, Johnston B, D'Ippolito A, Moon DH, Carulli JP, et al. Erythroid lineage chromatin accessibility maps facilitate identification and validation of NFIX as a fetal hemoglobin repressor. Commun Biol. 2023;6:640. PMID: 37316562.
  59. Lim RG, Al-Dalahmah O, Wu J, Gold MP, Reidling JC, Tang G, et al. Huntington disease oligodendrocyte maturation deficits revealed by single-nucleus RNAseq are rescued by thiamine-biotin supplementation. Nat Commun. 2022;13:7791. PMID: 36543778.
  60. Nofi CP, Wang P, Aziz M. Chromatin-Associated Molecular Patterns (CAMPs) in sepsis. Cell Death Dis. 2022;13:700. PMID: 35961978.
  61. Dreischer P, Duszenko M, Stein J, Wieder T. Eryptosis: Programmed Death of Nucleus-Free, Iron-Filled Blood Cells. Cells. 2022;11:503. PMID: 35159312.
  62. Gauthier BR, Comaills V. Nuclear Envelope Integrity in Health and Disease: Consequences on Genome Instability and Inflammation. Int J Mol Sci. 2021;22:7281. PMID: 34298904.
  63. Garcia M, Thirouard L, Sedès L, Monrose M, Holota H, Caira F, et al. Nuclear Receptor Metabolism of Bile Acids and Xenobiotics: A Coordinated Detoxification System with Impact on Health and Diseases. Int J Mol Sci. 2018;19:3630. PMID: 30453651.
  64. Skinner BM, Johnson EE. Nuclear morphologies: their diversity and functional relevance. Chromosoma. 2017;126:195-212. PMID: 27631793.
  65. Gorth DJ, Mauck RL, Chiaro JA, Mohanraj B, Hebela NM, Dodge GR, et al. IL-1ra delivered from poly(lactic-co-glycolic acid) microspheres attenuates IL-1β-mediated degradation of nucleus pulposus in vitro. Arthritis Res Ther. 2012;14:R179. PMID: 22863285.

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