Skip to main content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Review Article
  • Published:

Not your father's planarian: a classic model enters the era of functional genomics

Key Points

  • The planarian was established as a system for the study of regenerative development more than 200 years ago. As attention shifted towards organisms with more rigid patterns of development, and molecular and genetic techniques gained popularity, planarians fell out of favour.

  • Planarians show remarkable developmental plasticity. For example, a planarian can regenerate from a piece of tissue that represents less than 1/279 of the adult organism.

  • They also de-grow in nutritionally limiting conditions and can shrink to a size that is smaller than their size at hatching. De-growth is accompanied by loss of reproductive structures, which re-form when nutrition becomes plentiful again.

  • Developmental plasticity is dependent on a population of pluripotent cells called neoblasts. Neoblasts are thought to represent stem cells and are stimulated to migrate, grow and divide by discontinuities in the adult structure. Planarians therefore have great potential as a model system for studying stem-cell biology.

  • Genetic markers have recently become available that will help to study neoblasts, and to determine, for example, whether the cells are a homogeneous population or consist of a collection of lineage-restricted cells.

  • Further technological improvements in this system have been the development of methods for in situ hybridization, the successful application of RNA interference and an accumulation of genomic resources.

  • The current fascination with stem-cell biology, along with the technological advances in planarians, has set the stage for a resurgence of interest in these organisms. Planarians could provide important insights into the mechanisms that underlie regeneration and development.

Abstract

Freshwater planarians were a classic model for studying the problems of development and regeneration. However, as attention shifted towards animals with more rigid developmental processes, the planarians, with their notoriously plastic ontogeny, declined in significance as a model system. This trend was exacerbated with the introduction of genetic and molecular approaches, which did not work well in planarians. More recently, the heightened interest in stem-cell biology, along with the successful application of molecular, cellular and genomic approaches in planarians, is re-establishing these fascinating organisms as models for studying regeneration and developmental plasticity.

This is a preview of subscription content, access via your institution

Access options

Buy this article

Prices may be subject to local taxes which are calculated during checkout

Figure 1: Diagrams of the major organ systems in freshwater planarians.
Figure 2: The planarian's regenerative and remodelling abilities.
Figure 3: Growth and de-growth in response to food availability.
Figure 4: Regeneration in Schmidtea mediterranea.
Figure 5: Distribution of proliferating cells in the intact planarian.
Figure 6: The planarian Schmidtea mediterranea.
Figure 7: Representative whole-mount in situ hybridizations.
Figure 8: Scheme for using RNA interference to identify genes that are required for regenerative processes.

Similar content being viewed by others

References

  1. Lenhoff, S. G. & Lenhoff, H. M. Hydra and the Birth of Experimental Biology, 1744: Abraham Trembley's Memoirs Concerning the Natural History of a Type of Freshwater Polyp with Arms Shaped like Horns (Boxwood, Pacific Grove, California, 1986).

    Google Scholar 

  2. Agata, K. & Watanabe, K. Molecular and cellular aspects of planarian regeneration. Sem. Cell Dev. Biol. 10, 377–383 (1999).

    Article  CAS  Google Scholar 

  3. Lender, T. in Advances in Morphogenesis Vol. 2 (eds Abercrombie, M. & Brachet, J.) 305–331 (Academic, Inc., New York, 1962).

    Google Scholar 

  4. Wolff, E. in Regeneration: 20th Growth Symp. (ed. Rudnick, D.) 53–84 (The Ronald Press Co., New York, 1962).

    Google Scholar 

  5. Brøndsted, H. V. Planarian Regeneration 276 (Pergamon, London, 1969).An excellent compendium of experiments carried out on planarians with an extensive bibliography of the classic literature.

    Book  Google Scholar 

  6. Baguñà, J. et al. in Experimental Embryology in Aquatic Plants and Animals (ed. Marthy, H.-J.) 129–162 (Plenum, New York, 1990).

    Book  Google Scholar 

  7. Baguñà, J. in Cellular and Molecular Basis of Regeneration: From Invertebrates to Humans (eds Ferretti, P. & Géraudie, J.) 135–165 (John Wiley & Sons Ltd, Chichester, UK, 1998).

    Google Scholar 

  8. Hyman, L. H. The Invertebrates: Protozoa through Ctenophora 726 (McGraw–Hill Book Co., Inc., New York, 1940).

    Google Scholar 

  9. Willmer, P. Invertebrate Relationships (Cambridge Univ. Press, New York, 1990).

    Book  Google Scholar 

  10. Brusca, R. C. & Brusca, G. J. Invertebrates (Sinauer Associates, Sunderland, Massachusetts, 1990).

    Google Scholar 

  11. Barnes, R. S. K., Calow, P. & Olive, P. J. W. The Invertebrates: a New Synthesis 488 (Blackwell Science, Oxford, 1993).

    Google Scholar 

  12. Hyman, L. H. The Invertebrates: Platyhelminthes and Rhynchocoela. The Acoelomate Bilateria 550 (McGraw–Hill, New York, 1951).A detailed and comprehensive survey of the Platyhelminthes.

    Google Scholar 

  13. Giribet, G., Distel, D. L., Polz, M., Sterrer, W. & Wheeler, W. C. Triploblastic relationships with emphasis on the Acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: a combined approach of 18S rDNA sequences and morphology. Syst. Biol. 49, 539–562 (2000).

    Article  CAS  PubMed  Google Scholar 

  14. Tyler, S. in Interrelationships of the Platyhelminthes (eds Littlewood, D. T. J. & Bray, R. A.) 3–12 (Taylor & Francis, London, 2001).

    Google Scholar 

  15. Zrzavy, J., Mihulka, S., Kepka, P., Bezd, A. & Tietz, D. Phylogeny of the Metazoa based on morphological and 18S ribosomal DNA evidence. Cladistics 14, 249–285 (1998).

    Article  PubMed  Google Scholar 

  16. Adoutte, A. et al. The new animal phylogeny: reliability and implications. Proc. Natl Acad. Sci. USA 97, 4453–4456 (2000).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  17. Aguinaldo, A. et al. Evidence for a clade of nematodes, arthropods and other moulting animals. Nature 387, 489–493 (1997).

    Article  CAS  PubMed  Google Scholar 

  18. Baguñà, J., Carranza, S., Paps, J., Ruiz-Trillo, I. & Riutort, M. in Interrelationships of the Platyhelminthes (eds Littlewood, D. T. J. & Bray, R. A.) 49–56 (Taylor & Francis, London, 2001).

    Google Scholar 

  19. Ishii, S. The ultrastructure of the protonephridial flame cell of the freshwater planarian Bdellocephala brunnea. Cell Tissue Res. 206, 441–449 (1980).

    CAS  PubMed  Google Scholar 

  20. Ishii, S. The ultrastructure of the protonephridial tubules of the freshwater planarian Bdellocephala brunnea. Cell Tissue Res. 206, 451–458 (1980).

    CAS  PubMed  Google Scholar 

  21. Agata, K. et al. Structure of the planarian central nervous system (CNS) revealed by neuronal cell markers. Zool. Sci. 15, 433–440 (1998).

    Article  CAS  Google Scholar 

  22. Carpenter, K., Morita, M. & Best, J. Ultrastructure of the photoreceptor of the planarian Dugesia dorotocephala. I. Normal eye. Cell Tissue Res. 148, 143–158 (1974).

    Article  CAS  PubMed  Google Scholar 

  23. MacRae, E. The fine structure of sensory receptor processes in the auricular epithelium of the planarian, Dugesia tigrina. Z. Zellf. 82, 479–494 (1967).

    Article  CAS  Google Scholar 

  24. Pedersen, K. J. Some features of the fine structure and histochemistry of planarian subepidermal gland cells. Z. Zellf. 50, 121–142 (1959).

    Article  Google Scholar 

  25. Pedersen, K. J. Slime-secreting cells of planarians. Ann. NY Acad. Sci. 106, 424–443 (1963).

    Article  CAS  PubMed  Google Scholar 

  26. Rieger, R. M., Tyler, S., Smith, J. P. S. I. & Rieger, G. M. in Microscopic Anatomy of Invertebrates Vol. 3 (eds Harrison, F. W. & Bogitsch, B. J.) 7–140 (Wiley–Liss, New York, 1991).

    Google Scholar 

  27. Pallas, P. S. Miscellanea zoologica, quibus novae imprimis atque obscurae animalium species (Hagae Comitum, apud Pterum van Cleef, Holland, 1766).

    Google Scholar 

  28. Randolph, H. Observations and experiments on regeneration in planarians. Arch. Entwm. Org. 5, 352–372 (1897).

    Google Scholar 

  29. Morgan, T. H. Experimental studies of the regeneration of Planaria maculata. Arch. Entwm. Org. 7, 364–397 (1898).

    Google Scholar 

  30. Allen, G. E. Thomas Hunt Morgan 447 (Princeton Univ. Press, New Jersey, 1978).

    Google Scholar 

  31. Morgan, T. H. Regeneration in planarians. Arch. Entwm. Org. 10, 58–119 (1900).

    Google Scholar 

  32. Morgan, T. H. Growth and regeneration in Planaria lugubris. Arch. Entwm. Org. 13, 179–212 (1902).

    Google Scholar 

  33. Morgan, T. H. Regeneration of heteromorphic tails in posterior pieces of Planaria simplicissima. J. Exp. Zool. 1, 385–393 (1904).

    Article  Google Scholar 

  34. Morgan, T. H. Regeneration in Allolobophora foetida. Arch. Entwm. Org. 5, 570–586 (1897).

    Google Scholar 

  35. Morgan, T. H. 'Polarity' considered as a phenomenon of gradation of materials. J. Exp. Zool. 2, 495–506 (1905).

    Article  Google Scholar 

  36. Lawrence, P. A. Background to bicoid. Cell 54, 1–2 (1988).

    Article  CAS  PubMed  Google Scholar 

  37. Wolpert, L. in A History of Regeneration Research: Milestones in the Evolution of a Science (ed. Dinsmore, C. E.) 201–217 (Cambridge Univ. Press, Cambridge, UK, 1991).

    Google Scholar 

  38. Morgan, T. H. Regeneration 316 (The Macmillan Co., New York, 1901).

    Book  Google Scholar 

  39. Lillie, F. R. Some notes on regeneration and regulation in planarians. I. The source of material of new parts and limits of size. Am. Nat. 34, 173–177 (1900).

    Article  Google Scholar 

  40. Schultz, E. Über Reduktionen. I. Über Hungerserscheinungen bei Planaria lactea. Arch. Entwm. Org. 18, 555–577 (1904).

    Google Scholar 

  41. Berninger, J. Über die Einwirkung des Hungers auf Planarien. Zool. Jahrb. 30, 181–216 (1911).

    Google Scholar 

  42. Abeloos, M. Recherches expérimentales sur la croissance et la régénération chez les planaires. Bull. Biol. 1, 1–140 (1930).

    Google Scholar 

  43. Baguñà, J. & Romero, R. Quantitative analysis of cell types during growth, degrowth and regeneration in the planarians Dugesia mediterranea and Dugesia tigrina. Hydrobiologia 84, 181–194 (1981).

    Article  Google Scholar 

  44. Child, C. M. Starvation, rejuvenescence and acclimation in Planaria dorotocephala. Arch. Entwm. Org. 38, 418–446 (1914).

    CAS  Google Scholar 

  45. Calow, P. Life Cycles: an Evolutionary Approach to the Physiology of Reproduction, Development and Ageing 164 (Chapman & Hall, London, 1978).

    Book  Google Scholar 

  46. Romero, R. & Baguñà, J. Quantitative cellular analysis of growth and reproduction in freshwater planarians (Turbellaria; Tricladida). I. A cellular description of the intact organism. Invert. Reprod. Dev. 19, 157–165 (1991).

    Article  Google Scholar 

  47. Baguñà, J. Mitosis in the intact and regenerating planarian Dugesia mediterranea n. sp. I. Mitotic studies during growth, feeding and starvation. J. Exp. Zool. 195, 53–64 (1976).

    Article  Google Scholar 

  48. Kobayashi, K., Koyanagi, R., Matsumoto, M., Padilla Cabrera, J. & Hoshi, M. Switching from asexual to sexual reproduction in the planarian Dugesia ryukyuensis: bioassay system and basic description of sexualizing process. Zool. Sci. 16, 291–298 (1999).

    Article  Google Scholar 

  49. Best, J. B., Goodman, A. B. & Pigon, A. Fissioning in planarians: control by the brain. Science 164, 565–566 (1969).

    Article  CAS  PubMed  Google Scholar 

  50. Pigon, A., Morita, M. & Best, J. B. Cephalic mechanism for social control of fissioning in planarians. II. Localization and identification of the receptors by electron micrographic and ablation studies. J. Neurobiol. 5, 443–462 (1974).

    Article  CAS  PubMed  Google Scholar 

  51. Morita, M. & Best, J. B. Effects of photoperiods and melatonin on planarian asexual reproduction. J. Exp. Zool. 231, 273–282 (1984).

    Article  CAS  Google Scholar 

  52. Child, C. M. Physiological isolation of parts and fission in Planaria. Arch. Entwm. Org. 30, 159–205 (1910).

    Google Scholar 

  53. Sonneborn, T. M. Genetic studies on Stenostomum incaudatum (nov. spec.). I. The nature and origin of differences among individuals formed during vegetative reproduction. J. Exp. Zool. 57, 57–108 (1930).

    Article  Google Scholar 

  54. Nuttycombe, J. W. & Waters, A. J. The American species of the genus Stenostomum. Proc. Am. Phil. Soc. 79, 213–300 (1938).

    Google Scholar 

  55. Le Moigne, A. Etude du développement embryonnaire et recherches sur les cellules de régénération chez l'embryon de la Planaire Polycelis nigra (Turbellarié, Triclade). J. Embryol. Exp. Morphol. 15, 39–60 (1966).

    CAS  PubMed  Google Scholar 

  56. Curtis, W. C. The life history, the normal fission, and the reproductive organs of Planaria maculata. Proc. Boston Soc. Nat. Hist. 30, 515–559 (1902).

    Google Scholar 

  57. Ogawa, K. et al. Identification of a receptor tyrosine kinase involved in germ cell differentiation in planarians. Biochem. Biophys. Res. Commun. 248, 204–209 (1998).

    Article  CAS  PubMed  Google Scholar 

  58. Shibata, N. et al. Expression of vasa(vas)-related genes in germline cells and totipotent somatic stem cells of planarians. Dev. Biol. 206, 73–87 (1999).

    Article  CAS  PubMed  Google Scholar 

  59. Schüpbach, T. & Wieschaus, E. Maternal-effect mutations altering the anterior–posterior pattern of the Drosophila embryo. Roux' Arch. Dev. Biol. 195, 302–317 (1986).

    Article  Google Scholar 

  60. Hay, B., Jan, L. Y. & Jan, Y. N. A protein component of Drosophila polar granules is encoded by vasa and has extensive sequence similarity to ATP-dependent helicases. Cell 55, 577–587 (1988).

    Article  CAS  PubMed  Google Scholar 

  61. Fujiwara, Y. et al. Isolation of a DEAD-family protein gene that encodes a murine homolog of Drosophila Vasa and its specific expression in germ cell lineage. Proc. Natl Acad. Sci. USA 91, 12258–12262 (1994).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  62. Olsen, L. C., Aasland, R. & Fjose, A. A vasa-like gene in zebrafish identifies putative primordial germ cells. Mech. Dev. 66, 95–105 (1997).

    Article  CAS  PubMed  Google Scholar 

  63. Tsunekawa, N., Naito, M., Sakai, Y., Nishida, T. & Noce, T. Isolation of chicken vasa homolog gene and tracing the origin of primordial germ cells. Development 127, 2741–2750 (2000).

    Article  CAS  PubMed  Google Scholar 

  64. Yoon, C., Kawakami, K. & Hopkins, N. Zebrafish vasa homologue RNA is localized to the cleavage planes of 2- and 4-cell-stage embryos and is expressed in the primordial germ cells. Development 124, 3157–3165 (1997).

    Article  CAS  PubMed  Google Scholar 

  65. Gruidl, M. E. et al. Multiple potential germ-line helicases are components of the germ-line-specific P granules of Caenorhabditis elegans. Proc. Natl Acad. Sci. USA 93, 13837–13842 (1996).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  66. Mochizuki, K., Nishimiya-Fujisawa, C. & Fujisawa, T. Universal occurrence of the vasa-related genes among metazoans and their germline expression in Hydra. Dev. Genes Evol. 211, 299–308 (2001).

    Article  CAS  PubMed  Google Scholar 

  67. Randolph, H. The regeneration of the tail in Lumbriculus. Zool. Anz. 14, 154–156 (1891).

    Google Scholar 

  68. Baguñà, J. Mitosis in the intact and regenerating planarian Dugesia mediterranea n. sp. II. Mitotic studies during regeneration, and a possible mechanism of blastema formation. J. Exp. Zool. 195, 65–80 (1976).

    Article  Google Scholar 

  69. Wolff, E. & Dubois, F. Sur la migration des cellules de régénération chez les Planaires. Revue Suisse Zool. 55, 218–227 (1948).

    Article  Google Scholar 

  70. Dubois, F. Contribution à l'étude de la migration des cellules de régénération chez les Planaires dulcicoles. Bull. Biol. Fr. Belg. 83, 213–283 (1949).

    Google Scholar 

  71. Fedecka-Bruner, B. in Regeneration in Animals and Related Problems (eds Kiortsis, V. & Trampusch, H. A. L.) 185–192 (North-Holland Publishing Co., Amsterdam, 1965).

    Google Scholar 

  72. Baguñà, J., Saló, E. & Auladell, C. Regeneration and pattern formation in planarians. III. Evidence that neoblasts are totipotent stem cells and the source of blastema cells. Development 107, 77–86 (1989).An elegant demonstration of the pluripotency of planarian neoblasts.

    Article  Google Scholar 

  73. Bardeen, C. R. & Baetjer, F. H. The inhibitive action of the Roentgen rays on regeneration in planarians. J. Exp. Zool. 1, 191–195 (1904).

    Article  Google Scholar 

  74. Miller, J. A. Studies on heteroplastic transplantation in triclads. I. Cephalic grafts between Euplanaria dorotocephala and E. tigrina. Phys. Zool. 11, 214–247 (1938).

    Article  Google Scholar 

  75. Santos, F. V. Studies on transplantation in planaria. Phys. Zool. 4, 111–164 (1931).

    Article  Google Scholar 

  76. Okada, Y. K. & Sugino, H. Transplantation experiments in Planaria gonocephala. Jpn. J. Zool. 7, 373–439 (1937).

    Google Scholar 

  77. Kato, K., Orii, H., Watanabe, K. & Agata, K. The role of dorsoventral interaction in the onset of planarian regeneration. Development 126, 1031–1040 (1999).Demonstrates the use of molecular tools to re-examine classical axial determination experiments in planarians.

    Article  CAS  PubMed  Google Scholar 

  78. Kato, K., Orii, H., Watanabe, K. & Agata, K. Dorsal and ventral positional cues required for the onset of planarian regeneration may reside in differentiated cells. Dev. Biol. 233, 109–121 (2001).

    Article  CAS  PubMed  Google Scholar 

  79. Salvetti, A., Rossi, L., Deri, P. & Batistoni, R. An MCM2-related gene is expressed in proliferating cells of intact and regenerating planarians. Dev. Dyn. 218, 603–614 (2000).

    Article  CAS  PubMed  Google Scholar 

  80. Ito, H., Saito, Y., Watanabe, K. & Orii, H. Epimorphic regeneration of the distal part of the planarian pharynx. Dev. Genes Evol. 211, 2–9 (2001).

    Article  CAS  PubMed  Google Scholar 

  81. Newmark, P. & Sánchez Alvarado, A. Bromodeoxyuridine specifically labels the regenerative stem cells of planarians. Dev. Biol. 220, 142–153 (2000).This paper debunks the dogma that planarian cells could not be labelled with exogenous thymidine analogues.

    Article  CAS  PubMed  Google Scholar 

  82. Gratzner, H. G. A new reagent for detection of DNA replication. Science 218, 474–475 (1982).

    Article  CAS  PubMed  Google Scholar 

  83. Benazzi, M. & Benazzi Lentati, G. Animal Cytogenetics Vol. 1 Platyhelminthes (ed. Bernard, J.) 182 (Gebrüder Borntraeger, Berlin, 1976).

    Google Scholar 

  84. Bayascas, J. R., Castillo, E., Muñoz-Mármol, A. M. & Saló, E. Planarian hox genes: novel patterns of expression during regeneration. Development 124, 141–148 (1997).

    Article  CAS  PubMed  Google Scholar 

  85. Orii, H. et al. The planarian HOM/HOX homeobox genes (Plox) expressed along the anteroposterior axis. Dev. Biol. 210, 456–468 (1999).A comprehensive characterization of HOM/HOX homeobox genes in planarians with spatial expression data for Plox5 that indicates its possible involvement in anteroposterior patterning.

    Article  CAS  PubMed  Google Scholar 

  86. Saló, E. & Baguñà, J. Regeneration and pattern formation in planarians. I. The pattern of mitosis in anterior and posterior regeneration in Dugesia (G) tigrina, and a new proposal for blastema formation. J. Embryol. Exp. Morphol. 83, 63–80 (1984).

    PubMed  Google Scholar 

  87. Garcia-Fernandez, J. et al. High copy number of highly similar mariner-like transposons in planarian (Platyhelminthe): evidence for a trans-phyla horizontal transfer. Mol. Biol. Evol. 12, 421–431 (1995).

    CAS  PubMed  Google Scholar 

  88. Baguñà, J. et al. From morphology and karyology to molecules. New methods for taxonomical identification of asexual populations of freshwater planarians. A tribute to Professor Mario Benazzi. Ital. J. Zool. 66, 207–214 (1999).

    Article  Google Scholar 

  89. Prats, J. M. An`alisi del contingut de DNA a planària d'aigües dolces per citometria de flux i citoespectrofotmetria. Universitat de Barcelona (1991).

    Google Scholar 

  90. Umesono, Y., Watanabe, K. & Agata, K. A planarian orthopedia homolog is specifically expressed in the branch region of both the mature and regenerating brain. Dev. Growth Differ. 39, 723–727 (1997).The first description of whole-mount in situ hybridization in planarians.

    Article  CAS  PubMed  Google Scholar 

  91. Pedersen, K. J. Cytological studies on the planarian neoblast. Z. Zellf. 50, 799–817 (1959).

    Article  CAS  Google Scholar 

  92. Nogi, T. & Watanabe, K. Position-specific and non-colinear expression of the planarian posterior (Abdominal-B-like) gene. Dev. Growth Differ. 43, 177–184 (2001).

    Article  CAS  PubMed  Google Scholar 

  93. Orii, H., Kato, K., Agata, K. & Watanabe, K. Molecular cloning of bone morphogenetic protein (BMP) gene from the planarian Dugesia japonica. Zool. Sci. 15, 871–877 (1998).

    Article  CAS  Google Scholar 

  94. Kobayashi, C., Nogi, T., Watanbe, K. & Agata, K. Ectopic pharynxes arise by regional reorganization after anterior/posterior chimera in planarians. Mech. Dev. 89, 25–34 (1999).The elegant use of molecular markers to address the problems of axial rearrangement described in classic transplantation studies.

    Article  CAS  PubMed  Google Scholar 

  95. Umesono, Y., Watanabe, K. & Agata, K. Distinct structural domains in the planarian brain defined by the expression of evolutionarily conserved homeobox genes. Dev. Genes Evol. 209, 31–39 (1999).

    Article  CAS  PubMed  Google Scholar 

  96. Cebrià, F. et al. Dissecting planarian CNS regeneration by the expression of neural-specific genes. Dev. Growth Differ. (in the press).

  97. Fire, A. et al. Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans. Nature 391, 806–811 (1998).

    Article  CAS  PubMed  Google Scholar 

  98. Sánchez Alvarado, A. & Newmark, P. A. Double-stranded RNA specifically disrupts gene expression during planarian regeneration. Proc. Natl Acad. Sci. USA 96, 5049–5054 (1999).Introduction of loss-of-gene-function assays to the study of metazoan regeneration using dsRNA in planarians.

    Article  PubMed  PubMed Central  Google Scholar 

  99. Cheyette, B. N. et al. The Drosophila sine oculis locus encodes a homeodomain-containing protein required for the development of the entire visual system. Neuron 12, 977–996 (1994).

    Article  CAS  PubMed  Google Scholar 

  100. Serikaku, M. A. & O'Tousa, J. E. sine oculis is a homeobox gene required for Drosophila visual system development. Genetics 138, 1137–1150 (1994).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  101. Pineda, D. et al. Searching for the prototypic eye genetic network: sine oculis is essential for eye regeneration in planarians. Proc. Natl Acad. Sci. USA 97, 4525–4529 (2000).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  102. Timmons, L. & Fire, A. Specific interference by ingested dsRNA. Nature 395, 854 (1998).

    Article  CAS  PubMed  Google Scholar 

  103. Timmons, L., Court, D. L. & Fire, A. Ingestion of bacterially expressed dsRNAs can produce specific and potent genetic interference in Caenorhabditis elegans. Gene 263, 103–112 (2001).

    Article  CAS  PubMed  Google Scholar 

  104. Fraser, A. G. et al. Functional genomic analysis of C. elegans chromosome I by systematic RNA interference. Nature 408, 325–330 (2000).

    Article  CAS  PubMed  Google Scholar 

  105. Betchaku, T. Isolation of planarian neoblasts and their behavior in vitro with some aspects of the mechanism of the formation of regeneration blastema. J. Exp. Zool. 164, 407–433 (1967).One of the first systematic examinations of neoblast behaviour in culture.

    Article  CAS  PubMed  Google Scholar 

Download references

Acknowledgements

We thank S. Robb and R. Juste for their outstanding contributions to the S. mediterranea EST project; M. Sepanski for electron microscopy; M. Pala for her generous gift of the sexual strain of S. mediterranea; K. Agata and the members of his lab for sharing unpublished results; N. Oviedo for sharing in situ results before publication; anonymous referees for their helpful suggestions; and the National Institutes of Health, National Institute of General Medical Science for supporting our work on planarians.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Phillip A. Newmark.

Related links

Related links

DATABASES

LocusLink 

MCM2

PCNA

sine oculis

vasa 

FURTHER INFORMATION

Alejandro Sánchez Alvarado's lab

Phillip Newmark's lab

Planarian papers published in the 18th–19th centuries in Europe

Planarian resources on the Web

Glossary

PLURIPOTENCY

The ability of a cell to contribute to multiple tissues in a developing organism. If a cell is able to contribute to all tissues, it is said to be totipotent.

COELOM

A fluid-filled body cavity that is lined by mesodermal cells.

BILATERIA

Animals in which the right and left halves are mirror images. Bilaterians include most animals: chordates, arthropods, worms, molluscs and others.

PROTOSTOME

Bilaterian animals, including arthropods, molluscs and worms, whose mouth develops before the anus during embryogenesis.

DEUTEROSTOME

Animals, including chordates and echinoderms, whose mouth develops after the anus during embryogenesis.

FLAME CELL

A cell that is distinguished by a tuft of beating cilia (resembling a flame) and that filters waste materials into the excretory system.

NERVOUS PLEXUS

A bundle or collection of nerves.

POLYP

A sessile form of an animal, such as a Hydra, that is attached to a substrate.

MORPHOGENETIC GRADIENT

A progressive increase or decrease in the concentration of molecules that cause cells to adopt different developmental fates at different concentrations.

POLAR GRANULE

A cytoplasmic organelle that is associated with the germ plasm (germ-line material) in Drosophila.

ANNELID

The phylum of segmented worms.

PARENCHYMA

Mesodermal tissue that fills the space between the epidermis and the gut in acoelomates (animals that lack a coelom).

BLASTEMA

A specialized structure that is composed of an epithelial layer and mesodermally derived, undifferentiated cells.

MIXOPLOID

Containing cells that are of different ploidy — for example, diploid and polyploid.

MARINER

A transposable element that was originally discovered in Drosophila and has since been shown to be present in the genomes of diverse species.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Newmark, P., Alvarado, A. Not your father's planarian: a classic model enters the era of functional genomics. Nat Rev Genet 3, 210–219 (2002). https://doi.org/10.1038/nrg759

Download citation

  • Issue Date:

  • DOI: https://doi.org/10.1038/nrg759

This article is cited by

Search

Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing