Where are Primary Cilia Found?

This list is maintained in collaboration with Prof. Denys Wheatley (Department of Cell Pathology, University of Aberdeen, MacRobert Building, 581 King Street, Aberdeen AB24 5UA, UK). We invite others to help us maintain an accurate and up-to-date database.

NOTICE: Any errors or omissions in this compilation are Bowser's responsibility. Apologies are extended to Prof. Lynn Margulis for our stubborn use of traditional cilium terminology. Please feel free to e-mail comments or suggestions to bowser@global2000.net .

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Kidney tubules [Mammals, Birds, Lizards, Turtle, Marine fish]

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Adrenal Glands [Rat, Squirrel, Hamster, chicken, frog]

Thyroid Gland [Man, Dog, Pig, Chicken]

Parathyroid Glands [Man, Monkey, Dog, Deer, Mouse]

Female Reproductive System

Ovary: follicular cells of various developing follicles, germinal epithelium, stromal cells, theca interna [Man, Cat, Rat, Rabbit, Guinea Pig, Mouse]

Fallopian tube ampullar mucosa

Vagina--stratified epithelium [Man]

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Uterus--Endometrium [Man, Rabbit, Rat, Mouse, Hamster]

Male Reproductive system--Rete testes, prostate, ductuli efferentes, peritubular myoid cells & interstitial cells of testes, seminal vesicles [Man, Rat, Mouse, Rabbit, Dog, Mouse, Locust].

Pancreas--exocrine, endocrine, ductular cells [Man, Rat, Mouse, Rabbit, Dog, Chicken, Guinea Pig, Hamster, Gerbil, Toad].


fat storing cells [Man, Monkey, Rat, Bat, Guinea Pig, birds]

Perisinusoidal cells [Man]

Intrahepatic/cystic bile duct [Man, Monkey, Cat, Pig, Rat, Hamster, Bat, Birds]

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Mesothelial and serosal cells [Cat]

Myoepithelial cells--breast, salivary gland [man]

Mammary gland [goat]

Thymus [Mouse]

Pituitary [Mouse, Guinea Pig, Rat, Rabbit, Dog, Man]

Neuroepithelial cells

Ventral aspect of third ventricle/infundibulum [Man, Rat]

Ependymal cells [frog]

NOTE: In most cases, ependymal cells have tufts of motile 9+2 cilia; they are readily cultured from fetal and neonate rat/mouse brain. Cells containing bona fide primary cilia are also found among the kinetociliated cells. (SSB)

Choroid Plexus [Rabbit, Chick]

Central & peripheral nervous system [Man, Rabbit, Rat, Mouse, Bird, Fish, Hydra]

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Epidermis--basal cells, keratinocytes, melanocytes [Man, Ox]

Oral Tissues-gingiva, oral mucosa, ameloblasts, odontoblast [Man, Cat, Rat, Guinea Pig]

Spleen--reticular cells [Rat, Rabbit]

Vascular endothelium--heart myocardium, aorta, arteries, capillaries [Man, Rabbits]

Chondrocytes [Man, Dog, Cat, Rabbit, Mouse, Guinea Pig]

Synovial cells [Man, Rabbit, Guinea Pig]

Osteocytes [Mice]

Heart Myocardium [Man, Rabbit, Mice, Chicks, Lizards]

Smooth muscle [Rat, Frog, Chick]

Invertebrate Muscle [Brittle-star tube feet, Polychaete body wall]

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Fibroblasts--adult tissues: Tendon [Man, Mouse] Fibrous connective tissue [Rabbit, Rat]

Respiratory epithelium [Mouse, Guinea Pig] PROBABLY ALL DEVELOPMENTAL SYSTEMS

CELLS OF SENSORY ORGANS: In many cases, the axonemal symmetry of the primary cilia listed here is not 9+0; they may be found as single, nonmotile 9+2 cilia (e.g., kinocilium of hair cells), tufts of long, nonmotile(?) 9+2 cilia (olfactory neurons), or highly modified 9+0 cilia (e.g., rods and cones). Recent papers are no longer included in this database unless they contain novel information.

Vertebrate eye [Man, Monkey, Porpoise, Ox, Cat, Rabbit, Mouse, Guinea Pig]


Pericanalicular mesothelium (patients w/glaucoma or cataracts)

Trabecular endothelium



Ciliary epithelium [rabbit] and Schwann cells of ciliary process stroma

Other eye bits:

Pineal photoreceptors

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Lower vertebrate & invertebrate eyes/photoreceptors

Middle and Inner ear [Monkey, Rat, Guinea Pig, Newt, Fish] Lateral line organ and macula utriculi [Fish]

Taste Buds [Rabbit, Mouse]

Olfactory epithelium [Dog, Frog] These are tufts of cilia with 9+2 symmetry proximally, 9+0 (or fewer doublets) peripherally. Motility unknown(?).

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Other sensory cells

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Note: Carefully compare the occurrence of primary cilia in normal cells (above) with the following reports of "defective" or "aberrant" cilia as an indication of a pathological state. You will see that in certain cases, the cells of pathological tissues normally express primary cilia.

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Atherosclerotic lesions (de-differentiation of endothelial cells?)

Other pathologic conditions

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Cells with tufts of 9+2 cilia found in "unusual" places:


There is a growing literature regarding the putative role of "motile" primary cilia in determining left/right symmetry in early embryonic development. See: Nonaka et al. (1998) Randomization of left­right asymmetry due to loss of nodal cilia generating leftward flow of extraembryonic fluid in mice lacking KIF3B motor protein. Cell 95:829­837.

The following citations support the notion that if a fully-differentiated cell type lacks a primary cilium, it most likely has a precursor that is ciliated.

  • Collin, 1971 [review]
  • [neonatal rat, hamster] Clabough, 1973
  • [Neonatal rat] Zimmerman & Tso, 1975; [Fetus, man] Møller, 1974
  • Insect (Coccinella) photoreceptor Home, 1975

    Insect (Lepidoptera) gland morphogenesis Barbier,1975

  • Insect (Silk moth) gland morphogenesis Berry & Johnson, 1975
  • Insect (Cockroach) gland morphogenesis Sreng & Quennedey, 1976
  • Kidney, fetal [Man] DeMartino & Zamboni, 1966
  • Zimmermann, 1971
  • Arey, 1974
  • Gudima et al., 1986
  • Postnatal renal collecting duct [Rabbit] Evan et al., 1991
  • Olfactory epithelium [Rat] Menco & Farbman, 1985 (a,b)
  • Menco & Farbman, 1987
  • Menco, 1988 (a,b)
  • Spleen, fetal--mesenchyme [Chick] De Harven & Bernhard, 1956
    • [Man] Zamboni & Westin, 1964
  • Parathyroid, fetus [Man] Nakagami et al., 1968
  • Thymus [Man] Sebuwufu, 1968
  • Endothelium heart [Man] Bystrevskaya et al., 1988
  • pineal [Man] Yamamoto & Fujimoto, 1980
  • Primordial erythroblasts [Chick, Rat] Breton-Gorius & Stralin, 1967
  • Tunica propria of bursa [Chick embryo] Quesada et al., 1985
  • Limb bud--mesenchyme [Chick, Mouse] Sorokin, 1962
    • --epithelium Jurand, 1965
      • Trelstad et al., 1967
      • Fonte et al., 1971
      • Synovial cells [Rabbit; regeneration] Campbell & Callahan, 1971
      • Tooth [Rat molar amelogenesis] Reith, 1967
        • [Rat incisor ameloblasts] Sasano, 1986
      • Bone [Rat] Federman & Nichols, 1974
      • Pituitary, fetal Ellis, 1966 (see Curry & Wheatley)
      • Brain [Tadpole] Kemali & Gioffre, 1978
    • Nervous tissue [Mouse] Cohen et al., 1988
      • Neural tube [Chick] Duncan, 1957
      • Sotelo & Trujillo-Cenóz
      • [Mouse] Herman & Kauffman, 1966
        • Supraoptic nucleus, neonate [Rat] Lafarga et al., 1980
        • Hippocampal neurons, embryo [Rat] Deitch, unpublished
        • Cerebral cortex, fetal [Man] Currie & Wheatley, 1966
    • Ependymal cells
      • 3rd ventricle [Rat] Ugrumov & Mitskevich, 1980
      • cerebellum [tadpole] Gona & Hauser, 1982

    Pulmonary epithelial precursors [Mice] Rehm et al., 1988

    • Ciliated epithelium
    • Lung [Rat] Sorokin, 1968
    • Trachea [Rat, Guinea Pig] Friedmann & Bird, 1971
    • Kober & Herbst, 1975
    • Dalen, 1981
    • Gaillard et al., 1989
    • Chondrogenesis [Man, Dog] Stockwell, 1971
      • Kelley, 1975
      • Cox & Peacock, 1977
      • Wilsman & Fletcher, 1978

    Muscle --skeletal/cardiac [Chick, Mouse] Przybylski & Blumberg, 1966

    • Rash et al., 1969
    • Przybylski, 1971
    • Viragh & Challice, 1973
    • Zummo et al., 1984

    --Regeneration [Tadpole tail] Warren, 1974

    • Myklebust et al., 1977

    Oviduct [Rabbit, Mouse]

    • Dirksen, 1971 (check this)
    • Komatsu & Fujita, 1978 (a,b)
    • Odor & Blandau, 1985

    Go to bibliography


    Primary culture:

    • Fibroblasts
      • [Man] Wheatley, 1969; Strugnell et al., 1996
      • [Mouse] Wheatley, 1969
      • [Rat] Wheatley, 1969
      • [Chick] Wheatley, 1969; Bowser, unpub.
      • [Prostate; Man] Bayne et al., 1998
    • Keratinocytes [Man] Strugnell et al., 1996
    • Gingiva [Man] Rose, 1970
    • Osteoblast [Fetal rat leg bone] Rose, 1970
    • Cornea [Frog] Hayden, unpub.
    • Corpus luteum epithelial cells [cow] Herrman et al., 1996
    • Retinal pigment epithelium [Man] Topp et al., 1996
    • Kidney epithelia
      • [Rat] Curthoys & Bellemann, 1979
      • [Rabbit] Minuth & Kriz, 1982
      • [Human] Trifillis & Kahng, 1990
      • [Bird] Sutterlin & Laverty, 1998
    • Anterior pituitary [Rat] Fazekas et al., 1982

  • Bile duct epithelium [Pig] Talbot & Caperna, 1998

    Continuous cell lines:

    • 3T3 Fibroblast
    • Goldberg & Green, 1964
    • Osborn & Weber, 1976
    • Albrecht-Buehler, 1977
    • Tucker et al., 1979a,b
    • Albrecht-Buehler & Bushnell, 1979
    • Albrecht-Buehler & Bushnell, 1980
    • Ho & Tucker, 1989
  • 3T6 Fibroblast Wheatley, 1971; 1972
  • 3Y1 Fibroblast Katsumoto et al., 1994
  • DonC Fibroblast Wheatley, 1969
  • BHK21/C13 Fibroblast (hamster)
    • Wheatley, 1969
    • Archer & Wheatley, 1971
  • Nil/2e Fibroblast Wheatley, 1969
  • CH3 Fibroblast (hamster) Wheatley, 1982
  • WI-38 Fibroblast (human) Wheatley, 1982
  • Vero Fibroblast (monkey) Wheatley, 1972
  • PE Mesenchyme (pig embryo) Vorobjev & Chentsov, 1982
  • PtK1, PtK2 Kidney epithelium (rat kangaroo)
    • Jensen et al., 1979
    • Rieder et al., 1979
    • Roth et al., 1988
  • LLC-PK1 Kidney epithelium (pig) Roth et al., 1988
  • MDCK Kidney epithelium (dog) Roth et al., 1988
  • BSC-40 Kidney epithelium Roth et al., 1988
  • GH3 PC 9+0 Pituitary tumor Bowser, unpub.
  • HFH-18 Melanoma (mouse) Rose, 1970
  • BPAE Pulmonary aortic endothel. (cow) Bowser, unpubl.
  • Li10 Liver epithelium Mori et al., 1979
  • CO25 Myoblast (mouse) Zeytinoglu et al., 1997

    Go to bibliography