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Cilia and flagella

Nine doublets around a central pair, with the motor on one side and the signal line running to the middle. The human body carries four kinds of it, not two.

15 pictures and 17 papers. Its organelle page: Cilia and flagella.

Gallery

Panel E, "iHTM Control" column: gamma-tubulin (red, basal body), ARL13B (green, ciliary axoneme) and DAPI (blue) in human iPSC-derived trabecular meshwork cells, showing a normal primary cilium. Part of a composite figure on OCRL mutations in Lowe syndrome.

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

Transmission electron microscope image of a thin section cut through the bronchiolar area of lung (mouse), showing cilia cross-sections. The inside of the cilium contain precisely arranged microtubules, the axoneme. The axoneme has a central unit containing two single microtubules and nine peripheral doublet microtubules (known as the "9+2"). Dynein sidearms project from the A tubule of each doublet.

Louisa Howard, Michael Binder (Dartmouth Electron Microscope Facility) · Public domain

High magnification of "Lung cilia#80311-50" image. This image is a thin x-section cut through cilia. The inside of the cilium contain precisely arranged microtubules, the axoneme. The axoneme has a central unit containing two single microtubules and nine peripheral doublet microtubules (known as the "9+2"). Dynein sidearms project from the A tubule of each doublet.

Louisa Howard, Michael Binder (Dartmouth Electron Microscope Facility) · Public domain

Performance of AAP purified gamma-Tubulin Antibody. ( A ) Immunofluorescence staining showing AAP purified gamma-Tubulin (#D-0006-100UG) performance in ciliated NIH-3T3 cells. Merged images including gamma-Tubulin (green), anti-acetylated tubulin (blue), and anti TMEM67 (red). The white square outlines a representative ciliary structure used for higher magnification in c-f. Scale bar represents 1 μm.

Karapetian, Gerhardt, Lemanska et al 2026, BMC Molecular and Cell Biology · CC BY 4.0

Decreased transition zone protein localization in ciliated trisomy 21 cells. ( A ) Representative confocal images of RPE1 D21 and T21 cells grown on coverslips and serum depleted for 24 hr. Cells were stained with GT335, PCNT, and the transition zone protein CEP290. Yellow arrows point to the CEP290 transition zone population and cyan arrows point to CEP290 satellites that colocalize with PCNT. (

Jewett, McCurdy, O'Toole et al 2023, eLife · CC BY 4.0

The GPCRs used in this study localize to the cilium. A, RPE1 cells transiently expressing indicated GFP- or Flag-tagged GPCR were fixed and processed for immunofluorescence. Cilia were identified by immunostaining endogenous ARL13B. In each image, the cilium of interest is enlarged and highlighted in a box at the upper right corner. Flag-HTR7 served as a nonciliary GPCR (negative control).

Mahajan, Madugula, Chia et al 2026, The Journal of Biological Chemistry · CC BY 4.0

Loss of IFT88 in Villin-Cre mice impairs ciliogenesis in the efferent ductules. A Gross anatomy of the reabsorptive mouse efferent ductules showing the proximal, conus, and distal segments. While the proximal segment, adjacent to the rete testis, consists of convoluted tubules, the distal segment converges into a single duct connecting to the epididymis. Its epithelium is composed of two main cell types: multiciliated cells (MCC) and reabsorptive primary ciliated cell (RPCC) that ensure sperm concentration through fluid reabsorption. C Immunofluorescent staining of efferent ductules from adult Ctl and Ift88 cKO mice showing IFT88 (gray), γ-tubulin (γ-tub, green) marking centrioles/basal bodies, and Acetylated α-tubulin (red) labeling axoneme in primary (arrow) and motile cilia. F Ultrastructural components of primary and motile cilia, including the basal body (Bb), axoneme (Ax), daughter centriole (Dc), transition zone (Tz), and basal foot (Bf), in the efferent ductules of control and Ift88 cKO mice by transmission electron microscopy (TEM).

Augiere, Hermo, Ferrier-Tarin et al 2026, Cellular and Molecular Life Sciences · CC BY 4.0

Cross-section of a normal tracheal axoneme by TEM, panel D of a figure comparing wild-type and Armc2-knockout ciliary axoneme structure. Scale bar 50 nm.

Giordani, Houdeline, Court et al 2026, Frontiers in Cell and Developmental Biology · CC BY 4.0

Motile cilia covering the apical surface of a bioengineered nasal epithelial construct, by scanning electron microscopy. One panel cropped from a larger figure on ciliary ultrastructure. Scale bar 10 um.

Bradbury, Dickson, Curtis et al 2026, Journal of Anatomy · CC BY 4.0

Transverse section of a fertile control sperm flagellum by TEM, labeled with the central pair (CP), outer dense fibers (ODF) and doublet microtubules (DMT), showing the canonical 9+2 axonemal organization. Scale bar 200 nm.

Li, Wan, Feng et al 2026, Frontiers in Cell and Developmental Biology · CC BY 4.0

Transmission electron microscope observation of the body surface cilia in trochophore larva of U. unicinctus .

Kong, Wei, Liu et al 2023, International Journal of Molecular Sciences · CC BY 4.0

The tracheal pseudostratified columnar ciliated epithelium reveals the columnar ciliated cells (C), the more electron-dense goblet cells (G), and basal cells (B). The columnar ciliated cells contain rounded to oval euchromatic nuclei (N) with small patches of peripheral heterochromatin. Regularly arranged cilia (arrows) are present. (rat (trachea); Fig. 9a.)

Sarhan, Elekhtiar, Elmowafy et al 2026, Histochemistry and Cell Biology · CC BY 4.0

TEM images of axonemal cross sections of human sperm flagella. Panel A, right: sperm from M1688, M1344, and M1670; arrows point to examples of outer dynein arms (ODAs). Panel B, right: sperm from M865, with causal DNAH17 variants, clearly show an absence of ODAs as well as axonemal disruption, including missing or supernumerary and misplaced microtubule doublets. Scale bar, 200 nm. (human (sperm flagellum); Figure 2, TEM panels A right and B right.)

Koser, Rieck, Aprea et al 2026, Human Reproduction Open · CC BY 4.0

Loss of zygotene cilia does not overtly affect meiotic chromosome pairing or DNA repair. (A) IF staining of seminiferous tubule squashes from control or cKO mice expressing EGFP-Centrin1 (centriole marker) with antibodies against A-tub and SYCP3 (meiotic chromosomal axis marker). Arrowheads indicate basal bodies extending primary cilia in zygotene cells. Scale bar, 5 um. Adapted: IF panel A only; quantification panel B and chromosome spread panels C-H excluded. (mouse (seminiferous tubule spermatocytes, CEP164-cKO); Figure 3, panel A only (IF of centrioles and primary cilia; quantification panel B and chromosome spread panels C-H excluded; rows 10-710 of full figure, 1600x700).)

Chen, Gong, Mak et al 2026, EMBO Reports · CC BY 4.0

Immunofluorescence staining for ARL13B marks primary cilia as slender projections extending from ARPE-19 human retinal pigment epithelial cells, with the ciliary axoneme clearly delineated against the nuclear DAPI counterstain. (Homo sapiens (ARPE-19 retinal pigment epithelial cells).)

Sui W 2025, Frontiers in cell and developmental biology · CC BY 4.0

Papers

  1. Ortiz-Álvarez G, Al Jord A. Repurposing Cell Cycle Oscillators for Multiciliated Cell Differentiation. Adv Exp Med Biol. 2026;1521:43-55. PMID: 42782440.
  2. Li Y, Yang Y, Yang S, Chang H, Li YP, Yang S. Ciliary Ift20 and Wwtr1 Coordinate Transforming Growth Factor-β Receptor II Signaling to Maintain Bone-Fat Balance in Skeletal Homeostasis. MedComm (2020). 2026;7:e70916. PMID: 42769548.
  3. Sarhan NR, Elekhtiar SA, Elmowafy R, Shrief AI. Empagliflozin mitigates lipopolysaccharide-induced tracheal injury in rats via downregulation of IL-6/JAK/STAT3 signaling pathway and stem cell preservation: histological and molecular study. Histochem Cell Biol. 2026;164:82. PMID: 42700260.
  4. Guo W, Yue J, Park JH, Valverde-Mendez D, Botting JM, Yu H, et al. Near-atomic in situ architecture and membrane-coupled dynamics of the Vibrio cholerae sheathed flagellum. Nat Commun. 2026;17:9199. PMID: 42660870.
  5. Koser SA, Rieck C, Aprea I, Krallmann C, Gaikwad AS, Wallmeier J, et al. HIPK4 is a novel gene associated with teratozoospermia and male infertility. Hum Reprod Open. 2026;2026:hoag066. PMID: 42597842.
  6. Hecquet A, Bonnefoy S, Bastin P. The trypanosome flagellum as model for parasitology, cell biology and ciliopathies. Semin Cell Dev Biol. 2026;184:103689. PMID: 42561750.
  7. Pistorio V, Vatier C, Capel , Beaupère C, Auclair M, Steunou V, et al. Centriolar Protein POC5 Regulates Human Adipogenesis and Cellular Senescence: Insights From a Novel Metabolic Ciliopathy. FASEB J. 2026;40:e72158. PMID: 42507085.
  8. Vijayakumaran A, Godbehere C, Abuammar A, Breusegem SY, Hurst LR, Morone N, et al. 3D nanoscale reconstruction of human airway multiciliogenesis reveals cellular architecture remodeling and cilia-mitochondria link via rootlets. Cell Rep. 2026;45:117714. PMID: 42485185.
  9. Chen JJ, Gong X, Mak M, Li FQ, Takemaru KI. Basal body docking failure triggers centriole clustering and elimination in mammalian spermatogenesis. EMBO Rep. 2026;27:5039-5061. PMID: 42471456.
  10. Boboltz A, Rathi V, Kumar S, Duncan GA. Myeloperoxidase impairs mucociliary transport on human airway epithelium. Dis Model Mech. 2026;19:dmm052764. PMID: 42063333.
  11. Wåhlander J, Schmidt T, Hansen CR, Kahn R, Påhlman LI. Sputum from Individuals with Primary Ciliary Dyskinesia Drives M2-like Macrophage Polarization. Lung. 2026;204:6. PMID: 41582098.
  12. Sui W, Ma X, Gong Y, Qiu K, Yuan M, Mao K, et al. Loss of a primary cilia protein ARL13B promotes TGFβ-1 induced EMT of RPE in proliferative vitreoretinopathy via increasing Smad3 expression. Front Cell Dev Biol. 2025;13:1661658. PMID: 41425088.
  13. Kovács P, Brazda P, Hajdú T, Harsányi B, Juhász K, Takács R, et al. Podocalyxin and ciliary neurotrophic factor receptor are novel components of the surfaceome of chondrogenic cells. Cell Commun Signal. 2025;24:8. PMID: 41327293.
  14. Liao L, Zhou R, Liu M. Primary cilia and their role in preeclampsia. Int J Med Sci. 2025;22:3260-3267. PMID: 40765552.
  15. Sun Y, Luo Z, Fu Y, Ngo T, Wang W, Wang Y, et al. Primary cilia and inflammatory response: unveiling new mechanisms in osteoarthritis progression. Exp Biol Med (Maywood). 2025;250:10490. PMID: 40357414.
  16. Wang X, Yin G, Yang Y, Tian X. Ciliary and Non-Ciliary Roles of IFT88 in Development and Diseases. Int J Mol Sci. 2025;26:2110. PMID: 40076734.
  17. Wu S, Zhou H, Ling H, Sun Y, Luo Z, Ngo T, et al. LIPUS regulates the progression of knee osteoarthritis in mice through primary cilia-mediated TRPV4 channels. Apoptosis. 2024;29:785-798. PMID: 38517601.

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