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Planarian

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1168:. They discovered that one of the amputated fragments sent to space regenerated into a double-headed worm. The majority of such amputated worms (95%) did not do so, however. An amputated worm regenerated into a double-head creature after spending five weeks aboard the International Space Station (ISS) – though regeneration of amputated worms as double-headed heteromorphosis is not a rare phenomenon unique to a microgravity environment. In contrast, two-headed planaria regenerates can be induced by exposing amputated fragments to electrical fields. Such exposure with opposite polarity can induce a planarian with 2 tails. Two-headed planaria regenerates can be induced by treating amputated fragments with pharmacological agents that alter levels of calcium, cyclic AMP, and protein kinase C activity in cells, as well as by genetic expression blocks (interference RNA) to the canonical Wnt/β-Catenin signalling pathway. 1131:. Neoblasts are required for new cell production, and they therefore provide the cellular basis for planarian regeneration. Cell signaling mechanisms provide positional information that regulates the cell types and tissues that are produced from the neoblasts in regeneration. Many signaling molecules that provide positional information to neoblasts, in regeneration and homeostasis, are expressed in muscle cells. Following injury, muscle cells throughout the body can alter the expression of genes that encode molecules that provide positional information. Therefore, the activities of neoblasts and muscle cells following injuries are essential for successful regeneration. 1004:, adult stem cells, which proliferate and produce differentiated cells. Some researchers claim that the products derived from bisecting a planarian are similar to the products of planarian asexual reproduction; however, debates about the nature of asexual reproduction in planarians and its effect on the population are ongoing. Some species of planarian are exclusively asexual, whereas some can reproduce both sexually and asexually. In most of the cases the sexual reproduction involve two individuals; auto fecundation has been rarely reported (e.g. in 591: 982: 137: 110: 561: 885: 838: 576: 1190:
cut the worm in two, and allowed both worms to regenerate each half would develop the light-shock reaction. In 1963, McConnell repeated the experiment, but instead of cutting the trained flatworms in two he ground them into small pieces and fed them to other flatworms. He reported that the flatworms learned to associate the bright light with a shock much faster than flatworms who had not been fed trained worms.
43: 230: 780:. Planarians lack a circulatory system, and absorb oxygen through their body wall. They uptake food to their gut using a muscular pharynx, and nutrients diffuse to internal tissues. A three-branched intestine runs across almost the entire body, and includes a single anterior and two posterior branches. The planarian intestine is a 1000:. Eggs develop inside the body and are shed in capsules. Weeks later, the eggs hatch and grow into adults. In asexual reproduction, the planarian fissions and each fragment regenerates its missing tissues, generating complete anatomy and restoring functions. Asexual reproduction, similar to regeneration following injury, requires 1206:
has ever reproduced his results of planarians scrunching when exposed to light. Subsequent explanations of this scrunching behaviour associated with cannibalism of trained planarian worms were that the untrained flatworms were only following tracks left on the dirty glassware rather than absorbing
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This experiment intended to test whether memory could be transferred chemically. The experiment was repeated with mice, fish, and rats, but it always failed to produce the same results. The perceived explanation was that rather than memory being transferred to the other animals, it was the hormones
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planarian flatworms by pairing a bright light with an electric shock. After repeating this several times they took away the electric shock, and only exposed them to the bright light. The flatworms would react to the bright light as if they had been shocked. Thompson and McConnell found that if they
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The planarian has a soft, flat, wedge-shaped body that may be black, brown, blue, gray, or white. The blunt, triangular head has two ocelli (eyespots), pigmented areas that are sensitive to light. There are two auricles (earlike projections) at the base of the head, which are sensitive to touch and
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Vila-Farré, Miquel; Rozanski, Andrei; Ivanković, Mario; Cleland, James; Brand, Jeremias N.; Thalen, Felix; Grohme, Markus A.; von Kannen, Stephanie; Grosbusch, Alexandra L.; Vu, Hanh T.-K.; Prieto, Carlos E.; Carbayo, Fernando; Egger, Bernhard; Bleidorn, Christoph; Rasko, John E. J. (2023-10-19).
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that are found in the planarian parenchyma across the planarian body. They are small and round cells, 5 to 10 μm, and characterized by a large nucleus, which is surrounded by little cytoplasm. Neoblasts are required for regenerating missing tissues and organs, and they continuously replenish
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the presence of certain chemicals. The mouth is located in the middle of the underside of the body, which is covered with hairlike projections (cilia). There are no circulatory or respiratory systems; oxygen enters and carbon dioxide leaves the planarian's body by diffusing through the body wall.
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Historically, planarians have been considered "immortal under the edge of a knife." Very small pieces of the planarian, estimated to be as little as 1/279th of the organism it is cut from, can regenerate back into a complete organism over the course of a few weeks. New tissues can grow due to
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The organism itself does not have to be completely cut into separate pieces for the regeneration phenomenon to be witnessed. In fact, if the head of a planarian is cut in half down its center, and each side retained on the organism, it is possible for the planarian to regenerate two heads and
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Planarian regeneration combines new tissue production with reorganization to the existing anatomy, morphallaxis. The rate of tissue regrowth varies between species, but in frequently used lab species, functional regenerated tissues are available already 7–10 days following tissue amputation.
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any missing organ, which has made planarians a popular model in research of regeneration and stem cell biology. The genome sequences of several species are available, as are tools for molecular biology analysis.
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and excretory pores on them. Also, flame cells remove unwanted liquids from the body by passing them through ducts which lead to excretory pores, where waste is released on the dorsal surface of the planarian.
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Regeneration starts following an injury that require the growth of a new tissue. Neoblasts localized near the injury site proliferate to generate a structure of differentiating cells called
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of planarians make them a model system for investigating a number of biological processes, many of which may have implications for human health and disease. Advances in
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Rozanski, Andrei; Moon, HongKee; Brandl, Holger; Martín-Durán, José M; Grohme, Markus A; Hüttner, Katja; Bartscherer, Kerstin; Henry, Ian; Rink, Jochen C (2019-01-08).
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pp 3., "Planarians (the popular name for the group as a whole), or triclad flatworms (the more scientific designation of the same group), are acoelomate bilaterians".
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has emerged as the species of choice for modern molecular biology research, due to its diploid chromosomes and the availability of both asexual and sexual strains.
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Thus, one of their gametes will combine with the gamete of another planarian. Each planarian transports its secretion to the other planarian, giving and receiving
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Rilling M (1996). "The mystery of the vanished citations: James McConnell's forgotten 1960s quest for planarian learning, a biochemical engram, and celebrity".
799:, are usually found. Some species have auricles that protrude from the margins of the head. The auricles can contain chemical and mechanical sensory receptors. 2833:
Morokuma J, Durant F, Williams KB, Finkelstein JM, Blackiston DJ, Clements T, Reed DW, Roberts M, Jain M, Kimel K, Trauger SA, Wolfe BE, Levin M (April 2017).
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tissues by producing new cells. Neoblasts can self-renew and generate progenitors for different cell types. In contrast to adult vertebrate stem cells (e.g.,
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in the ingested ground animals that changed the behavior. McConnell believed that this was evidence of a chemical basis for memory, which he identified as
1037:(i.e., producing all somatic cell types). Moreover, they give rise to differentiating, post-mitotic, cells directly, and not by producing rapidly-dividing 1253: 818:). Sometimes, those species with two eyes may present smaller accessory or supernumerary eyes. The subterranean triclads are often eyeless or blind. 3319: 1210:
In 2012, Tal Shomrat and Michael Levin have shown that planarians exhibit evidence of long-term memory retrieval after regenerating a new head.
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Sluys, R.; Kawakatsu, M.; Riutort, M.; Baguñà, J. (2009). "A new higher classification of planarian flatworms (Platyhelminthes, Tricladida)".
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Saló E, Abril JF, Adell T, Cebrià F, Eckelt K, Fernandez-Taboada E, Handberg-Thorsager M, Iglesias M, Molina MD, Rodríguez-Esteban G (2009).
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in the U.S., sought to determine how microgravity and micro-geomagnetic fields would affect the growth and regeneration of planarian
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function possible in these animals and scientists are studying them worldwide. Like other invertebrate model organisms, for example
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Newmark PA, Sánchez Alvarado A (March 2002). "Not your father's planarian: a classic model enters the era of functional genomics".
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Triclads reproduce sexually and asexually, and different species may be able to reproduce by one or both modes. Planarians are
67: 51: 1713: 2138: 1459: 1384: 2010:"Vertically- and horizontally-transmitted memories - the fading boundaries between regeneration and inheritance in planaria" 3337: 1400: 1041:. Consequently, neoblasts divide frequently, and apparently lack a large sub-population of dormant or slow-cycling cells. 2835:"Planarian regeneration in space: Persistent anatomical, behavioral, and bacteriological changes induced by space travel" 1907:
Aoki R, Wake H, Sasaki H, Agata K (March 2009). "Recording and spectrum analysis of the planarian electroencephalogram".
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Cebrià, Francesc; Nakazawa, Masumi; Mineta, Katsuhiko; Ikeo, Kazuho; Gojobori, Takashi; Agata, Kiyokazu (2002-04-03).
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was responsible for some of the first systematic studies (that still underpin modern research) before the advent of
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is remodeled to restore symmetry and proportion of the new planaria that forms from a piece of a cut up organism.
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The number of eyes in the triclads is variable depending on the species. While many species have two eyes (e.g.
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which in addition to excellent regenerative abilities, are easy to culture in the lab. In recent decades,
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Observations on some interesting phenomena in animal physiology, exhibited by several species of planariae
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Triclads play an important role in watercourse ecosystems and are often very important as bio-indicators.
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Witchley, Jessica N.; Mayer, Mirjam; Wagner, Daniel E.; Owen, Jared H.; Reddien, Peter W. (2013-08-29).
2316:"Telomere maintenance and telomerase activity are differentially regulated in asexual and sexual worms" 902: 136: 17: 3399: 1245:
The most frequently used planarian in high school and first-year college laboratories is the brownish
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Several planarian species are commonly used for biological research. Popular experimental species are
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The order Tricladida is split into three suborders, according to their phylogenetic relationships:
2119:"Stem Cells, Patterning and Regeneration in Planarians: Self-Organization at the Organismal Scale" 3081: 2426:
Hayashi, Tetsutaro; Asami, Maki; Higuchi, Sayaka; Shibata, Norito; Agata, Kiyokazu (2006-07-13).
1272: 1072: 1038: 1030: 895: 3440: 3435: 3142: 2994:(1973). "Chapter 1: Behavioral studies of the molecular basis of memory". In Deutsch JA (ed.). 1259: 1220: 1186: 1096: 1066: 270: 219: 118: 2488:"Beta-catenin defines head versus tail identity during planarian regeneration and homeostasis" 1142:
that have the ability to create all the various cell types. These adult stem cells are called
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Chan JD, Agbedanu PN, Zamanian M, Gruba SM, Haynes CL, Day TA, Marchant JS (February 2014).
2161:"Clonogenic Neoblasts Are Pluripotent Adult Stem Cells That Underlie Planarian Regeneration" 1766:
Sluys, R. (1990). "A monograph of the Dimarcusidae (Platyhelminthes, Seriata, Tricladida)".
3241: 3189: 3180: 2938: 2499: 2379:"Planarian regeneration involves distinct stem cell responses to wounds and tissue absence" 2327: 1661:
Hallez P. (1892). Classification des Ticlades, Bulletin de la Société Zoologique de France.
1306: 595: 3057: 2428:"Isolation of planarian X-ray-sensitive stem cells by fluorescence-activated cell sorting" 1606: 853:. Longitudinal ventral nerve chords extend from the brain to the tail. Transverse nerves, 8: 3072: 2991: 2125:, Methods in Molecular Biology, vol. 1774, New York, NY: Springer, pp. 57–172, 1226: 637: 2884:"'Death and axes': unexpected Ca²⁺ entry phenologs predict new anti-schistosomal agents" 2503: 2331: 2314:
Tan TC, Rahman R, Jaber-Hijazi F, Felix DA, Chen C, Louis EJ, Aboobaker A (March 2012).
1526: 1491: 1310: 2910: 2883: 2859: 2834: 2718:"Planarian regeneration: achievements and future directions after 20 years of research" 2637: 2604: 2580: 2547: 2520: 2487: 2465: 2403: 2378: 2350: 2315: 2242: 2217: 2193: 2160: 2094: 2034: 2009: 1932: 1920: 1783: 1779: 1748: 1639: 1607:"PlanMine 3.0—improvements to a mineable resource of flatworm biology and biodiversity" 1587: 1539: 1446:, Methods in Molecular Biology, vol. 1774, New York, NY: Springer, pp. 1–56, 1322: 1183: 1100: 1080: 942: 590: 131: 2942: 2805: 1680: 3381: 3228: 3092:
A user-friendly visualization tool and database of planarian regeneration experiments
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Aboobaker AA (May 2011). "Planarian stem cells: a simple paradigm for regeneration".
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Riutort, Marta; Álvarez-Presas, Marta; Lázaro, Eva; Sol, Eduard; Paps, Jordi (2012).
1694: 1684: 1644: 1626: 1579: 1543: 1531: 1513: 1465: 1455: 1380: 1376: 1104: 607: 269:, which planarians use for regenerating missing body parts. Many species are able to 2469: 2098: 2059:"Not your father's planarian: a classic model enters the era of functional genomics" 1802: 1787: 1752: 1591: 1326: 3233: 3194: 3022: 2972: 2905: 2895: 2854: 2846: 2779: 2739: 2729: 2688: 2632: 2616: 2575: 2559: 2515: 2507: 2439: 2398: 2390: 2345: 2335: 2278: 2237: 2229: 2188: 2172: 2126: 2070: 2029: 2021: 1974: 1916: 1871: 1814: 1775: 1740: 1732: 1676: 1634: 1618: 1571: 1521: 1503: 1447: 1314: 1247: 1233: 1203: 1164: 1155: 1147: 1088: 1025: 63: 3207: 2900: 2563: 2267:"Bromodeoxyuridine Specifically Labels the Regenerative Stem Cells of Planarians" 1177: 1092: 825:. Between the epidermis and the gastrodermis there is a parenchymatous tissue or 262: 3350: 2394: 2130: 1451: 1114:. These animals have an apparently limitless regenerative capacity, and asexual 981: 3280: 3267: 2976: 2620: 2320:
Proceedings of the National Academy of Sciences of the United States of America
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The triclads have an anterior end or head where sense organs, such as eyes and
781: 725: 707: 447: 387: 286: 207: 3067: 3063: 2783: 2118: 1439: 1318: 3429: 3165: 2813: 2628: 2571: 2453: 2292: 2184: 2082: 1986: 1885: 1828: 1630: 1517: 1492:"Evolutionary dynamics of whole-body regeneration across planarian flatworms" 1368: 1199: 993:. In sexual reproduction, the mating generally involves mutual insemination. 990: 796: 677: 657: 649: 178: 2511: 2340: 2218:"Planarian stem cells specify fate yet retain potency during the cell cycle" 2176: 1736: 3036: 2919: 2868: 2791: 2753: 2702: 2646: 2589: 2529: 2461: 2412: 2359: 2300: 2283: 2266: 2251: 2202: 2090: 2043: 1994: 1928: 1893: 1698: 1648: 1583: 1535: 1469: 1136: 1079:
Planarians have a number of cell types, tissues and simple organs that are
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The body of the triclads is covered by a ciliated epidermis that contains
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Kenk, R., 1972. Freshwater planarians (Turbellarians) of North America.
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More information on the genetic screen to identify regeneration genes
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has been shown to maintain its telomere length through regeneration.
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Wagner, Daniel E.; Wang, Irving E.; Reddien, Peter W. (2011-05-13).
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The planarian nervous systems consists of a bilobed shaped cerebral
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Schmidtea mediterranea: a model system for analysis of motile cilia
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Link to an article discussing some work on planarian immortality
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Alejandro Sanchez-Alvarado's Seminar: Regeneration in Planarians
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Newmark, Phillip A.; Sánchez Alvarado, Alejandro (2000-04-15).
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Raz, Amelie A.; Wurtzel, Omri; Reddien, Peter W. (2021-04-20).
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Newmark, Phillip A.; Alvarado, Alejandro Sánchez (2002-03-01).
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Reddien, Peter W.; Alvarado, Alejandro Sánchez (2004-11-01).
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Sluys, Ronald; Riutort, Marta (2018), Rink, Jochen C. (ed.),
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More information on freshwater planarians and their biology
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Gurley KA, Rink JC, Sánchez Alvarado A (January 2008).
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As a model system in biological and biomedical research
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The excretory system is made of many tubes with many
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continue to live. Researchers, including those from
1906: 1110:Planarians are also an emerging model organism for 909:. Unsourced material may be challenged and removed. 2722:The International Journal of Developmental Biology 2675:Handberg-Thorsager M, Fernandez E, Salo E (2008). 2007: 1807:The International Journal of Developmental Biology 2215: 2158: 1214:Planarian species used for research and education 1171: 335:Phylogenetic supertree after Sluys et al., 2009: 3427: 2935:"Investigations of the cellular bases of memory" 2372: 2806:"Do it again. Round up of regenerating animals" 2117:Rink, Jochen C. (2018), Rink, Jochen C. (ed.), 2056: 2008:Neuhof M, Levin M, Rechavi O (September 2016). 1967:Annual Review of Cell and Developmental Biology 1960: 1275: – Satirical (and serious) science journal 2123:Planarian Regeneration: Methods and Protocols 1444:Planarian Regeneration: Methods and Protocols 1363: 1251:. Other common species used are the blackish 3118:Tricladida on the Encyclopedia of Life (EOL) 3008: 2668: 1437: 1198:. McConnell's results are now attributed to 315:, allowing them to glide along on a film of 3002: 2990: 2677:"Stem cells and regeneration in planarians" 1844: 1842: 1840: 1838: 1671:Rompolas P, Patel-King RS, King SM (2009). 1333: 108: 3096: 3026: 2909: 2899: 2858: 2769: 2743: 2733: 2692: 2636: 2579: 2519: 2443: 2432:Development, Growth & Differentiation 2402: 2349: 2339: 2282: 2241: 2192: 2033: 1875: 1864:Development, Growth & Differentiation 1818: 1638: 1525: 1507: 1292: 1290: 1288: 969:Learn how and when to remove this message 763: 325: 86:Learn how and when to remove this message 1979:10.1146/annurev.cellbio.20.010403.095114 1963:"Fundamentals of Planarian Regeneration" 1835: 980: 836: 589: 574: 559: 228: 2962: 2765: 2763: 2659: 2602: 1759: 1711: 1405:Integrated Taxonomic Information System 1357: 14: 3428: 1285: 3141: 3140: 2932: 2541: 2539: 2481: 2479: 2112: 2110: 2108: 1956: 1954: 1952: 1950: 1948: 1946: 1765: 849:that is referred to as the planarian 3400:ED5EC61A-9C78-8E2D-FF00-FF0F1072FC01 3374:dcd2f6bc-6b9f-4d0f-a7b6-392e887e5bf7 3286:8fadcfbd-6d55-473e-9fd8-aecf7ad4595f 3097:Aboobaker, Aziz (27 February 2008). 2760: 2116: 1557: 1555: 1553: 1433: 1431: 1429: 1427: 1425: 1423: 1421: 907:adding citations to reliable sources 878: 36: 3015:The Journal of Experimental Biology 3009:Shomrat T, Levin M (October 2013). 2373:Wenemoser, Danielle; Reddien, Peter 1440:"Planarian Diversity and Phylogeny" 1060:technologies has made the study of 24: 2536: 2476: 2105: 1943: 1921:10.1016/j.neuroscience.2008.11.011 1780:10.1111/j.1463-6409.1990.tb00237.x 1099:has attracted the most attention. 27:Flatworms of the Turbellaria class 25: 3462: 3046: 2996:The Physiological Basis of Memory 1550: 1418: 875:Reproductive system of planarians 832: 2603:Reddien, Peter W. (2018-10-04). 2445:10.1111/j.1440-169X.2006.00876.x 1877:10.1046/j.1440-169x.2002.00629.x 883: 135: 41: 2989:For a general review, see also 2983: 2956: 2926: 2875: 2826: 2798: 2709: 2653: 2596: 2419: 2366: 2307: 2258: 2209: 2152: 2050: 2001: 1900: 1851: 1794: 1705: 1664: 1121: 894:needs additional citations for 868: 59:it does not summarise the body. 3446:Negligibly senescent organisms 1655: 1598: 1496:Nature Ecology & Evolution 1482: 1393: 1172:Biochemical memory experiments 13: 1: 1681:10.1016/S0091-679X(08)93004-1 1345:Encyclopædia Britannica, Inc. 1279: 1182:In 1955, Robert Thompson and 985:Planarian reproductive system 861:) activity of other animals. 3064:Planaria eating worm segment 2901:10.1371/journal.ppat.1003942 2564:10.1016/j.celrep.2013.07.022 1207:the memory of their fodder. 1013: 330: 297:(freshwater planarian); and 7: 3133:Featured Creatures Web site 2395:10.1016/j.ydbio.2010.06.017 2131:10.1007/978-1-4939-7802-1_2 1725:Revue Écologie Terre et Vie 1452:10.1007/978-1-4939-7802-1_1 1266: 555: 304:Planarians move by beating 249:(triclads) are free-living 10: 3467: 2977:10.1037/0003-066X.51.6.589 2621:10.1016/j.cell.2018.09.021 2234:10.1016/j.stem.2021.03.021 1509:10.1038/s41559-023-02221-7 1299:Journal of Natural History 1175: 1017: 872: 29: 3451:Invertebrate common names 3149: 2784:10.1016/j.tcb.2011.01.005 1319:10.1080/00222930902741669 503: 488: 481: 445: 430: 423: 408: 401: 391: 373: 366: 351: 344: 203: 198: 132:Scientific classification 130: 116: 107: 102: 3111:University of Nottingham 1564:Nature Reviews. Genetics 1039:transit amplifying cells 774:fluid-filled body cavity 55:may need to be rewritten 30:Not to be confused with 3076:, facts, anatomy, image 2681:Frontiers in Bioscience 2512:10.1126/science.1150029 2341:10.1073/pnas.1118885109 2177:10.1126/science.1203983 2063:Nature Reviews Genetics 1737:10.3406/revec.2010.1533 1031:hematopoietic stem cell 1024:Neoblasts are abundant 841:Planaria nervous system 57:. The reason given is: 3074:Schmidtea mediterranea 2772:Trends in Cell Biology 2284:10.1006/dbio.2000.9645 1611:Nucleic Acids Research 1260:Girardia dorotocephala 1221:Schmidtea mediterranea 1116:Schmidtea mediterranea 986: 842: 772:flatworms that lack a 764:Anatomy and physiology 604: 587: 572: 326:Phylogeny and taxonomy 243: 242:Unidentified planarian 119:Dugesia subtentaculata 2965:American Psychologist 2735:10.1387/ijdb.072414es 2383:Developmental Biology 2271:Developmental Biology 1820:10.1387/ijdb.113441mr 984: 840: 593: 578: 563: 293:(marine planarians); 241: 3281:Fauna Europaea (new) 2939:University of Durham 1379:. pp. 1230 pp. 1305:(29–30): 1763–1777. 1273:Worm Runner's Digest 903:improve this article 596:Platydemus manokwari 3021:(Pt 20): 3799–810. 2660:Dalyell JG (1814). 2504:2008Sci...319..323G 2332:2012PNAS..109.4209T 2228:(7): 1307–1322.e5. 1623:10.1093/nar/gky1070 1311:2009JNatH..43.1763S 1227:Schmidtea polychroa 638:Centrovarioplanidae 301:(land planarians). 3028:10.1242/jeb.087809 2026:10.1242/bio.020149 1712:Manenti R (2010). 1204:blinded experiment 1184:James V. McConnell 1101:Thomas Hunt Morgan 987: 843: 610:ranks after Sluys 605: 588: 573: 565:Sabussowia ronaldi 244: 3423: 3422: 3382:Open Tree of Life 3143:Taxon identifiers 3078:at GeoChemBio.com 2728:(8–10): 1317–27. 2377:W. (2010-08-15). 2171:(6031): 811–816. 2140:978-1-4939-7802-1 1768:Zoologica Scripta 1617:(D1): D812–D820. 1502:(12): 2108–2124. 1461:978-1-4939-7802-1 1386:978-0-8053-7146-8 1377:Benjamin Cummings 1254:Planaria maculata 1107:as a discipline. 1105:molecular biology 1033:), neoblasts are 979: 978: 971: 953: 552: 551: 543: 542: 534: 533: 525: 524: 516: 515: 467: 466: 458: 457: 239: 227: 226: 194: 96: 95: 88: 68:lead layout guide 16:(Redirected from 3458: 3416: 3415: 3403: 3402: 3390: 3389: 3377: 3376: 3367: 3366: 3354: 3353: 3351:NHMSYS0000841064 3341: 3340: 3328: 3327: 3315: 3314: 3302: 3301: 3289: 3288: 3276: 3275: 3263: 3262: 3250: 3249: 3237: 3236: 3224: 3223: 3211: 3210: 3198: 3197: 3185: 3184: 3183: 3170: 3169: 3168: 3138: 3137: 3114: 3099:"Immortal Worms" 3062:YouTube videos: 3041: 3040: 3030: 3006: 3000: 2999: 2987: 2981: 2980: 2960: 2954: 2953: 2951: 2950: 2941:. Archived from 2930: 2924: 2923: 2913: 2903: 2879: 2873: 2872: 2862: 2830: 2824: 2823: 2821: 2820: 2802: 2796: 2795: 2767: 2758: 2757: 2747: 2737: 2713: 2707: 2706: 2696: 2672: 2666: 2665: 2657: 2651: 2650: 2640: 2600: 2594: 2593: 2583: 2543: 2534: 2533: 2523: 2483: 2474: 2473: 2447: 2423: 2417: 2416: 2406: 2376: 2370: 2364: 2363: 2353: 2343: 2311: 2305: 2304: 2286: 2262: 2256: 2255: 2245: 2213: 2207: 2206: 2196: 2156: 2150: 2149: 2148: 2147: 2114: 2103: 2102: 2054: 2048: 2047: 2037: 2005: 1999: 1998: 1958: 1941: 1940: 1904: 1898: 1897: 1879: 1855: 1849: 1846: 1833: 1832: 1822: 1798: 1792: 1791: 1763: 1757: 1756: 1722: 1716:Polycelis felina 1709: 1703: 1702: 1668: 1662: 1659: 1653: 1652: 1642: 1602: 1596: 1595: 1559: 1548: 1547: 1529: 1511: 1486: 1480: 1478: 1477: 1476: 1435: 1416: 1415: 1413: 1411: 1397: 1391: 1390: 1361: 1355: 1354: 1352: 1351: 1337: 1331: 1330: 1294: 1248:Girardia tigrina 1234:Dugesia japonica 1165:Dugesia japonica 1156:Tufts University 1026:adult stem cells 974: 967: 963: 960: 954: 952: 911: 887: 879: 580:Polycelis felina 484: 483: 426: 425: 404: 403: 394: 393: 369: 368: 347: 346: 339: 338: 263:adult stem cells 240: 192: 140: 139: 112: 100: 99: 91: 84: 80: 77: 71: 64:improve the lead 45: 44: 37: 21: 3466: 3465: 3461: 3460: 3459: 3457: 3456: 3455: 3426: 3425: 3424: 3419: 3411: 3406: 3398: 3393: 3385: 3380: 3372: 3370: 3362: 3357: 3349: 3344: 3336: 3331: 3323: 3318: 3310: 3305: 3297: 3292: 3284: 3279: 3271: 3266: 3258: 3253: 3245: 3240: 3232: 3227: 3219: 3214: 3206: 3201: 3193: 3188: 3179: 3178: 3173: 3164: 3163: 3158: 3145: 3123:Land planarians 3049: 3044: 3007: 3003: 2988: 2984: 2961: 2957: 2948: 2946: 2931: 2927: 2894:(2): e1003942. 2880: 2876: 2851:10.1002/reg2.79 2831: 2827: 2818: 2816: 2804: 2803: 2799: 2768: 2761: 2714: 2710: 2687:(13): 6374–94. 2673: 2669: 2658: 2654: 2601: 2597: 2544: 2537: 2498:(5861): 323–7. 2484: 2477: 2424: 2420: 2374: 2371: 2367: 2326:(11): 4209–14. 2312: 2308: 2263: 2259: 2214: 2210: 2157: 2153: 2145: 2143: 2141: 2115: 2106: 2055: 2051: 2006: 2002: 1959: 1944: 1905: 1901: 1856: 1852: 1847: 1836: 1813:(1–2–3): 5–17. 1799: 1795: 1764: 1760: 1720: 1710: 1706: 1691: 1669: 1665: 1660: 1656: 1603: 1599: 1560: 1551: 1487: 1483: 1474: 1472: 1462: 1436: 1419: 1409: 1407: 1399: 1398: 1394: 1387: 1362: 1358: 1349: 1347: 1339: 1338: 1334: 1295: 1286: 1282: 1269: 1240:S. mediterranea 1216: 1180: 1178:Memory transfer 1174: 1124: 1073:D. melanogaster 1047: 1022: 1016: 975: 964: 958: 955: 912: 910: 900: 888: 877: 871: 835: 768:Planarians are 766: 558: 553: 544: 535: 526: 517: 468: 459: 333: 328: 229: 191: 169:Platyhelminthes 134: 92: 81: 75: 72: 61: 46: 42: 35: 28: 23: 22: 15: 12: 11: 5: 3464: 3454: 3453: 3448: 3443: 3438: 3421: 3420: 3418: 3417: 3404: 3391: 3378: 3368: 3355: 3342: 3329: 3316: 3303: 3290: 3277: 3268:Fauna Europaea 3264: 3251: 3238: 3225: 3212: 3199: 3186: 3171: 3155: 3153: 3147: 3146: 3135: 3134: 3120: 3115: 3094: 3089: 3084: 3079: 3070: 3060: 3055: 3048: 3047:External links 3045: 3043: 3042: 3001: 2982: 2971:(6): 589–598. 2955: 2925: 2888:PLOS Pathogens 2874: 2825: 2797: 2759: 2708: 2667: 2652: 2615:(2): 327–345. 2595: 2558:(4): 633–641. 2535: 2475: 2438:(6): 371–380. 2418: 2389:(2): 979–991. 2365: 2306: 2277:(2): 142–153. 2257: 2222:Cell Stem Cell 2208: 2151: 2139: 2104: 2075:10.1038/nrg759 2069:(3): 210–219. 2049: 2020:(9): 1177–88. 2000: 1973:(1): 725–757. 1942: 1915:(2): 908–914. 1899: 1870:(2): 135–146. 1850: 1834: 1793: 1758: 1731:(2): 279–285. 1704: 1689: 1663: 1654: 1597: 1576:10.1038/nrg759 1549: 1481: 1460: 1417: 1392: 1385: 1356: 1332: 1283: 1281: 1278: 1277: 1276: 1268: 1265: 1215: 1212: 1176:Main article: 1173: 1170: 1123: 1120: 1112:aging research 1046: 1043: 1018:Main article: 1015: 1012: 1007:Cura foremanii 991:hermaphrodites 977: 976: 891: 889: 882: 873:Main article: 870: 867: 834: 833:Nervous system 831: 797:chemoreceptors 765: 762: 761: 760: 759: 758: 757: 756: 755: 754: 748: 736: 735: 734: 728: 726:Dendrocoelidae 722: 708:Continenticola 704: 703: 702: 690: 689: 688: 687: 686: 674: 673: 672: 666: 654: 653: 652: 646: 640: 557: 554: 550: 549: 546: 545: 541: 540: 537: 536: 532: 531: 528: 527: 523: 522: 519: 518: 514: 513: 510: 509: 502: 499: 498: 495: 494: 487: 482: 480: 474: 473: 470: 469: 465: 464: 461: 460: 456: 455: 452: 451: 448:Dendrocoelidae 444: 441: 440: 437: 436: 429: 424: 422: 419: 418: 415: 414: 407: 402: 400: 392: 390: 388:Continenticola 384: 383: 380: 379: 372: 367: 365: 362: 361: 358: 357: 350: 345: 343: 337: 332: 329: 327: 324: 287:Continenticola 225: 224: 223: 222: 216: 210: 208:Continenticola 201: 200: 196: 195: 186: 182: 181: 176: 172: 171: 166: 162: 161: 156: 152: 151: 146: 142: 141: 128: 127: 114: 113: 105: 104: 94: 93: 50:The article's 49: 47: 40: 26: 9: 6: 4: 3: 2: 3463: 3452: 3449: 3447: 3444: 3442: 3441:Animal models 3439: 3437: 3436:Rhabditophora 3434: 3433: 3431: 3414: 3409: 3405: 3401: 3396: 3392: 3388: 3383: 3379: 3375: 3369: 3365: 3360: 3356: 3352: 3347: 3343: 3339: 3334: 3330: 3326: 3321: 3317: 3313: 3308: 3304: 3300: 3295: 3291: 3287: 3282: 3278: 3274: 3269: 3265: 3261: 3256: 3252: 3248: 3243: 3239: 3235: 3230: 3226: 3222: 3217: 3213: 3209: 3204: 3200: 3196: 3191: 3187: 3182: 3176: 3172: 3167: 3161: 3157: 3156: 3154: 3152: 3148: 3144: 3139: 3132: 3128: 3124: 3121: 3119: 3116: 3112: 3108: 3104: 3100: 3095: 3093: 3090: 3088: 3085: 3083: 3080: 3077: 3075: 3071: 3069: 3065: 3061: 3059: 3056: 3054: 3051: 3050: 3038: 3034: 3029: 3024: 3020: 3016: 3012: 3005: 2997: 2993: 2992:Chapouthier G 2986: 2978: 2974: 2970: 2966: 2959: 2945:on 2012-10-15 2944: 2940: 2936: 2933:Kentridge B. 2929: 2921: 2917: 2912: 2907: 2902: 2897: 2893: 2889: 2885: 2878: 2870: 2866: 2861: 2856: 2852: 2848: 2845:(2): 85–102. 2844: 2840: 2836: 2829: 2815: 2814:New Scientist 2811: 2810:New Scientist 2807: 2801: 2793: 2789: 2785: 2781: 2778:(5): 304–11. 2777: 2773: 2766: 2764: 2755: 2751: 2746: 2741: 2736: 2731: 2727: 2723: 2719: 2712: 2704: 2700: 2695: 2690: 2686: 2682: 2678: 2671: 2663: 2656: 2648: 2644: 2639: 2634: 2630: 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1841: 1839: 1830: 1826: 1821: 1816: 1812: 1808: 1804: 1797: 1789: 1785: 1781: 1777: 1773: 1769: 1762: 1754: 1750: 1746: 1742: 1738: 1734: 1730: 1726: 1719: 1717: 1708: 1700: 1696: 1692: 1690:9780123813770 1686: 1682: 1678: 1674: 1667: 1658: 1650: 1646: 1641: 1636: 1632: 1628: 1624: 1620: 1616: 1612: 1608: 1601: 1593: 1589: 1585: 1581: 1577: 1573: 1569: 1565: 1558: 1556: 1554: 1545: 1541: 1537: 1533: 1528: 1523: 1519: 1515: 1510: 1505: 1501: 1497: 1493: 1485: 1471: 1467: 1463: 1457: 1453: 1449: 1445: 1441: 1434: 1432: 1430: 1428: 1426: 1424: 1422: 1406: 1402: 1396: 1388: 1382: 1378: 1374: 1370: 1366: 1360: 1346: 1342: 1336: 1328: 1324: 1320: 1316: 1312: 1308: 1304: 1300: 1293: 1291: 1289: 1284: 1274: 1271: 1270: 1264: 1262: 1261: 1256: 1255: 1250: 1249: 1243: 1241: 1237: 1235: 1230: 1228: 1223: 1222: 1211: 1208: 1205: 1201: 1200:observer bias 1197: 1191: 1188: 1185: 1179: 1169: 1167: 1166: 1161: 1157: 1151: 1149: 1145: 1141: 1138: 1132: 1130: 1119: 1117: 1113: 1108: 1106: 1102: 1098: 1094: 1090: 1086: 1082: 1077: 1075: 1074: 1069: 1068: 1063: 1059: 1056: 1052: 1042: 1040: 1036: 1032: 1027: 1021: 1011: 1009: 1008: 1003: 999: 994: 992: 983: 973: 970: 962: 959:November 2016 951: 948: 944: 941: 937: 934: 930: 927: 923: 920: –  919: 915: 914:Find sources: 908: 904: 898: 897: 892:This article 890: 886: 881: 880: 876: 866: 862: 860: 856: 852: 848: 839: 830: 828: 824: 819: 817: 813: 812: 807: 806: 800: 798: 793: 790: 785: 783: 779: 775: 771: 753: 749: 747: 743: 742: 741: 737: 733: 729: 727: 723: 721: 717: 716: 715: 711: 710: 709: 705: 701: 697: 696: 695: 691: 685: 681: 680: 679: 678:Procerodoidea 675: 671: 667: 665: 661: 660: 659: 658:Bdellouroidea 655: 651: 650:Meixnerididae 647: 645: 641: 639: 635: 634: 633: 629: 628: 627: 623: 622: 621: 617: 616: 615: 613: 609: 602: 598: 597: 592: 585: 581: 577: 570: 566: 562: 548: 547: 539: 538: 530: 529: 521: 520: 512: 511: 508: 507: 501: 500: 497: 496: 493: 492: 486: 485: 479: 476: 475: 472: 471: 463: 462: 454: 453: 450: 449: 443: 442: 439: 438: 435: 434: 428: 427: 421: 420: 417: 416: 413: 412: 406: 405: 399: 396: 395: 389: 386: 385: 382: 381: 378: 377: 371: 370: 364: 363: 360: 359: 356: 355: 349: 348: 341: 340: 336: 323: 320: 318: 314: 311: 307: 302: 300: 296: 292: 288: 284: 280: 275: 272: 268: 264: 260: 256: 253:of the class 252: 248: 221: 217: 215: 211: 209: 205: 204: 202: 197: 190: 187: 184: 183: 180: 179:Rhabditophora 177: 174: 173: 170: 167: 164: 163: 160: 157: 154: 153: 150: 147: 144: 143: 138: 133: 129: 125: 121: 120: 115: 111: 106: 101: 98: 90: 87: 79: 76:December 2023 69: 66:and read the 65: 60: 56: 53: 48: 39: 38: 33: 19: 3150: 3102: 3073: 3018: 3014: 3004: 2995: 2985: 2968: 2964: 2958: 2947:. Retrieved 2943:the original 2928: 2891: 2887: 2877: 2842: 2839:Regeneration 2838: 2828: 2817:. Retrieved 2809: 2800: 2775: 2771: 2725: 2721: 2711: 2694:10.2741/3160 2684: 2680: 2670: 2664:. Edinburgh. 2661: 2655: 2612: 2608: 2598: 2555: 2552:Cell Reports 2551: 2495: 2491: 2435: 2431: 2421: 2386: 2382: 2368: 2323: 2319: 2309: 2274: 2270: 2260: 2225: 2221: 2211: 2168: 2164: 2154: 2144:, retrieved 2122: 2066: 2062: 2052: 2017: 2014:Biology Open 2013: 2003: 1970: 1966: 1912: 1909:Neuroscience 1908: 1902: 1867: 1863: 1853: 1810: 1806: 1796: 1774:(1): 13–29. 1771: 1767: 1761: 1728: 1724: 1715: 1707: 1672: 1666: 1657: 1614: 1610: 1600: 1570:(3): 210–9. 1567: 1563: 1499: 1495: 1484: 1473:, retrieved 1443: 1408:. Retrieved 1401:"Tricladida" 1395: 1372: 1359: 1335: 1302: 1298: 1258: 1252: 1246: 1244: 1239: 1232: 1225: 1219: 1217: 1209: 1192: 1181: 1163: 1152: 1133: 1125: 1122:Regeneration 1115: 1109: 1097:regeneration 1078: 1071: 1065: 1051:life history 1048: 1023: 1005: 995: 988: 965: 956: 946: 939: 932: 925: 913: 901:Please help 896:verification 893: 869:Reproduction 863: 844: 820: 809: 803: 801: 794: 786: 767: 740:Geoplanoidea 738:Superfamily 714:Planarioidea 712:Superfamily 700:Dimarcusidae 684:Procerodidae 676:Superfamily 670:Bdellouridae 664:Uteriporidae 656:Superfamily 630:Superfamily 619: 611: 606: 594: 579: 564: 504: 489: 478:Geoplanoidea 446: 431: 409: 398:Planarioidea 374: 352: 334: 321: 303: 276: 258: 246: 245: 188: 117: 97: 82: 73: 62:Please help 58: 54: 52:lead section 3307:iNaturalist 3175:Wikispecies 3107:Brady Haran 2745:2445/192658 1745:2434/147984 1365:Campbell NA 1187:conditioned 1137:pluripotent 1095:. However, 1083:to our own 1035:pluripotent 918:"Planarian" 816:Geoplaninae 789:flame cells 752:Geoplanidae 720:Planariidae 694:Cavernicola 632:Cercyroidea 506:Geoplanidae 411:Planariidae 376:Cavernicola 342:Tricladida 283:Cavernicola 255:Turbellaria 220:Cavernicola 218:Infraorder 212:Infraorder 175:Subphylum: 3430:Categories 3208:Tricladida 3195:Tricladida 3181:Tricladida 3151:Tricladida 2949:2007-02-08 2819:2012-10-21 2146:2023-12-05 1475:2023-12-02 1350:2010-05-01 1348:Retrieved 1280:References 1196:memory RNA 1140:stem cells 1081:homologous 1067:C. elegans 929:newspapers 855:commissure 827:mesenchyme 811:Microplana 778:parenchyma 770:bilaterian 746:Dugesiidae 644:Cercyridae 620:Tricladida 491:Dugesiidae 295:Paludicola 271:regenerate 259:Tricladida 247:Planarians 193:Lang, 1884 189:Tricladida 103:Planarian 18:Planarians 3103:Test Tube 3068:Planarian 2629:0092-8674 2572:2211-1247 2454:0012-1592 2293:0012-1606 2185:0036-8075 2083:1471-0056 1987:1081-0706 1937:207244874 1886:0012-1592 1829:0214-6282 1631:0305-1048 1544:264347538 1518:2397-334X 1160:flatworms 1144:neoblasts 1055:molecular 1014:Neoblasts 1002:neoblasts 823:rhabdites 782:blind sac 732:Kenkiidae 706:Suborder 692:Suborder 624:Suborder 601:geoplanid 584:planariid 433:Kenkiidae 331:Phylogeny 299:Terricola 267:neoblasts 251:flatworms 206:Suborder 199:Subdivis 155:Kingdom: 149:Eukaryota 32:Planarium 3247:46481314 3160:Wikidata 3109:for the 3037:23821717 2920:24586156 2869:28616247 2792:21353778 2754:19247944 2703:18508666 2647:30290140 2590:23954785 2530:18063757 2470:10048289 2462:16872450 2413:20599901 2360:22371573 2301:10753506 2252:33882291 2203:21566185 2099:28379017 2091:11972158 2044:27565761 1995:15473858 1929:19063945 1894:11940100 1788:84915439 1753:54499235 1699:20409812 1649:30496475 1592:28379017 1584:11972158 1536:37857891 1527:10697840 1470:29916154 1410:July 23, 1371:(2019). 1369:Reece JB 1327:85174457 1267:See also 1129:blastema 1020:Neoblast 847:ganglion 626:Maricola 614:, 2009: 608:Linnaean 569:Maricola 556:Taxonomy 354:Maricola 291:Maricola 279:Maricola 257:, order 214:Maricola 165:Phylum: 159:Animalia 145:Domain: 124:dugesiid 3166:Q534535 3125:on the 2911:3930560 2860:5469732 2638:7706840 2581:4101538 2521:2755502 2500:Bibcode 2492:Science 2404:2950745 2351:3306686 2328:Bibcode 2243:8254784 2194:3338249 2165:Science 2035:5051648 1640:6324014 1373:Biology 1307:Bibcode 1089:tissues 1058:genetic 943:scholar 805:Dugesia 750:Family 744:Family 730:Family 724:Family 718:Family 698:Family 682:Family 668:Family 662:Family 648:Family 642:Family 636:Family 310:ventral 308:on the 265:called 185:Order: 3413:142028 3387:555387 3371:NZOR: 3260:1TRCDO 3221:165597 3035:  2918:  2908:  2867:  2857:  2790:  2752:  2701:  2645:  2635:  2627:  2588:  2578:  2570:  2528:  2518:  2468:  2460:  2452:  2411:  2401:  2375:  2358:  2348:  2299:  2291:  2250:  2240:  2201:  2191:  2183:  2137:  2097:  2089:  2081:  2042:  2032:  1993:  1985:  1935:  1927:  1892:  1884:  1827:  1786:  1751:  1697:  1687:  1647:  1637:  1629:  1590:  1582:  1542:  1534:  1524:  1516:  1468:  1458:  1383:  1325:  1148:tissue 1093:organs 945:  938:  931:  924:  916:  618:Order 612:et al. 313:dermis 3408:WoRMS 3395:Plazi 3338:54468 3325:12395 3320:IRMNG 3312:64222 3273:14248 3234:7NGC4 2466:S2CID 2095:S2CID 1933:S2CID 1784:S2CID 1749:S2CID 1721:(PDF) 1588:S2CID 1540:S2CID 1323:S2CID 1202:. 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Index

Planarians
Planarium
lead section
improve the lead
lead layout guide
Learn how and when to remove this message

Dugesia subtentaculata
dugesiid
Scientific classification
Edit this classification
Eukaryota
Animalia
Platyhelminthes
Rhabditophora
Tricladida
Continenticola
Maricola
Cavernicola
flatworms
Turbellaria
adult stem cells
neoblasts
regenerate
Maricola
Cavernicola
Continenticola
Maricola
Paludicola
Terricola

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