• Issigonis, M., Browder, K.L., Chen, R., Collins, J.J., and P.A. Newmark. 2023. A niche-derived nonribosomal peptide triggers planarian sexual development. Proc. Natl. Acad. Sci. 121: e2321349121 (cover)


  • Issigonis, M., Redkar, A.B., Rozario, T., Khan, U.W., Mejia-Sanchez, R., Lapan, S.W., Reddien, P.W., and P.A. Newmark. 2022. A Krüppel-like factor is required for development and regeneration of germline and yolk cells from somatic stem cells in planarians. PLOS Biology 20: e3001472.
  • Khan, U.W., and P.A. Newmark. 2022. Somatic regulation of female germ cell regeneration and development in planarians. Cell Reports 38: 110525.
  • P.A. Newmark and A. Sánchez Alvarado. 2022. Schmidtea happens: re-establishing the planarian as a model for studying the mechanisms of regeneration. Curr. Top. Dev. Biol. 147: 307-344.
  • Rozario, T., Collins, J.J., and P.A. Newmark. 2022. The good, the bad, and the ugly: from planarians to parasites. Curr. Top. Dev. Biol. 147: 345-373.
  • Rouhana, L., Chong, T, and P.A. Newmark. 2022. Analysis of morphogenesis and flagellar assembly during spermatogenesis in planarian flatworms. Methods Mol. Biol. 2364: 199-216.


  • Lee, J., Chong, T., and P.A. Newmark. 2020. The esophageal gland mediates host immune evasion by the human parasite Schistosoma mansoni. Proc. Natl. Acad. Sci. 117: 19299-19309. (cover)
  • Forsthoefel D.Jj, Cejda N.I., Khan U.W., and P.A. Newmark. 2020. Cell-type diversity and regionalized gene expression in the planarian intestine. eLife 9: e52613
  • Diaz Soria, C.L. *, Lee, J. *, Chong, T. , Coghlan, A., Tracey, A., Young, M.D., Andrews, T., Hall, C., Ling Ng, B., Rawlinson, K., Doyle, S.R., Leonard, S., Lu, Z., Bennett, H.M., Rinaldi, G., Newmark, P.A., and M. Berriman. 2020. Single-cell atlas of the first intra-mammalian developmental stage of the human parasite Schistosoma mansoni. Nature Communications 11: 6411.
    *equal contributions


  • Issigonis, M. and P.A. Newmark. 2019. From worm to germ: Germ cell development and regeneration in planarians. Curr. Top Dev. Biol. 135: 127-153.
  • Rozario, T., Quinn, E.B., Wang, J., Davis, R.E., and P.A. Newmark. 2019. Region-specific regulation of stem cell-driven regeneration in tapeworms. eLife 8: e48958
  • Gao, J., Yang, N., Lewis, F.A., Yau, P., Collins III, J.J., Sweedler, J.V., and P.A. Newmark. 2019. A rotifer-derived paralytic compound prevents transmission of schistosomiasis to a mammalian host. PLOS Biology 17(10): e3000485.


  • Saberi, A., Gulyaeva, A.A., Brubacher, J.L., Newmark, P.A., and A.E. Gorbalenya. 2018. A planarian nidovirus expands the limits of RNA genome size. PLOS Path. 14(11): e1007314.
  • Wang, B., Lee, J., Li, P., Saberi, A., Yang, H., Liu, C., Zhao, M., and P.A. Newmark. 2018. Stem cell heterogeneity drives the parasitic life cycle of Schistosoma mansoni. eLife 7: e35449.
  • Rozario, T., and P.A. Newmark. 2018. Prospecting for Planarian Pluripotency (Preview). Cell 173: 1566-1567.
  • King, R.S. and P.A. Newmark. 2018. Whole-mount in situ hybridization of planarians. Methods Mol. Biol. 1774: 379-392.
  • Forsthoefel, D.J., Ross K.G., Newmark P.A., Zayas R.M. Fixation, processing, and immunofluorescent labeling of whole-mount planarians. 2018. Methods Mol. Biol. 1774: 353-366.


  • Rouhana, L., Tasaki, J., Saberi, A., and P.A. Newmark. 2017. Genetic dissection of the planarian reproductive system through characterization of Schmidtea mediterranea CPEB homologs. Dev. Biol. 426: 43-55.


  • Roberts-Galbraith, R.H., Brubacher, J.L., and P.A. Newmark. 2016. A functional genomics screen in planarians reveals regulators of whole-brain regeneration. eLife 5: e17002.
  • Iyer, H., Issigonis, M., Sharma, P.P., Extavour, C.G., and P.A. Newmark. 2016. A pre-meiotic function for boule in the planarian Schmidtea mediterraneaProc. Natl. Acad. Sci. 113: E3509-3518.
  • Iyer. H., Collins III, J.J., and P.A. Newmark. 2016. NF-YB regulates spermatogonial stem cell self-renewal and proliferation in planarians. PLOS Genet. 12: e1006109.
  • Saberi, A., Jamal, A., Beets, I., Schoofs, L., and P.A. Newmark. 2016. GPCRs direct germline development and somatic gonad function in planarians. PLOS Biol. 14: e1002457 (cover).
  • Collins III, J.J., Wendt, G.R., Iyer. H., and P.A. Newmark. 2016. Stem cell progeny contribute to the schistosome host-parasite interface. eLife 5: e12473.
  • Ong, T.-H., Romanova, E.V., Roberts-Galbraith, R., Yang, N., Zimmerman, T.A., Collins, J.J., Lee, J.E., Kelleher, N.L., Newmark, P.A., and J.V. Sweedler. 2016. Mass spectrometry imaging and identification of peptides associated with cephalic ganglia regeneration in Schmidtea mediterraneaJ. Biol. Chem. 291: 8109-8120.


  • Issigonis, M. and P.A. Newmark. 2015. Heal thy cell(f): a single-cell view of regeneration. Dev. Cell. 35: 527-528.
  • Issigonis, M. and P.A. Newmark. 2015. Planarian “kidneys” go with the flow. eLife 4: e09353.
  • Rozario, T. and P.A. Newmark. 2015. A confocal microscopy-based atlas of tissue architecture in the tapeworm, Hymenolepis diminuta.  Exp. Parasitol. 158: 31-41.
  • Lambrus, B.G., Cochet-Escartin, O., Gao, J., Newmark, P.A., Collins, E-M. S., and J.J. Collins III. 2015. Tryptophan hydroxylase is required for eye melanogenesis in the planarian Schmidtea mediterraneaPLOS One 10(5): e0127074.
  • Roberts-Galbraith, R.H. and P.A. Newmark. 2015. On the organ trail: insights into organ regeneration in the planarian. Curr. Opin. Gen. Dev. 32: 37-46.


  • Forsthoefel, D.J., Waters, F.A., and P.A. Newmark. 2014. Generation of cell type-specific monoclonal antibodies for the planarian and optimization of sample processing for immunolabeling. BMC Dev. Biol. 14: 45.
  • Tharp, M.E., Collins, J.J., and P.A. Newmark. 2014. A lophotrochozoan-specific nuclear hormone receptor is required for reproductive system development in the planarian. Dev. Biol. 396: 150-157.
  • Rouhana, L., Weiss, J.A., King, R.S., and P.A. Newmark. 2014. PIWI homologs mediate Histone H4 mRNA localization to planarian chromatoid bodies.  Development 141: 2592-2601.
  • Brubacher, J.L., Vieira, A.P., and P.A. Newmark. 2014. Preparation of the planarian Schmidtea mediterranea for high-resolution histology and transmission electron microscopy. Nat. Protoc. 9: 661-673.


  • Wang, B., Collins III, J.J., and P.A. Newmark. 2013. Functional genomic characterization of neoblast-like stem cells in larval Schistosoma mansonieLife 2: e00768.
  • Sikes, J.M. and P.A. Newmark. 2013. Restoration of regeneration in a planarian with limited regenerative ability. Nature 500: 77-80.
  • Collins III, J.J. and P.A. Newmark. 2013. It’s no fluke: the planarian as a model for understanding schistosomes. PLOS Pathog. 9(7): e1003396.
  • Chong, T., Collins III, J.J., Brubacher, J.L., Zarkower, D., and P.A. Newmark. 2013. A sex-specific transcription factor controls male identity in a simultaneous hermaphrodite. Nat. Commun. 41814.
  • King, R.S. and P.A. Newmark. 2013. An in situ hybridization protocol for enhanced detection of gene expression in the planarian Schmidtea mediterraneaBMC Dev. Biol. 13:8.
  • Rouhana, L., Weiss, J.A., Forsthoefel, D.J., Lee, H., King, R.S., Inoue, T., Shibata, N., Agata, K., and P.A. Newmark. 2013. RNA interference by feeding in vitro synthesized double-stranded RNA to planarians: methodology and dynamics. Dev. Dynam. 242: 718-730 (cover).
  • Collins III, J.J., Wang, B., Lambrus, B.G., Tharp, M., Iyer, H., and P.A. Newmark. 2013. Adult somatic stem cells in the human parasite, Schistosoma mansoniNature 494: 476-479.
  • Roberts-Galbraith, R.H. and P.A. Newmark. 2013. Follistatin antagonizes Activin signaling and acts with Notum to direct planarian head regeneration. Proc. Natl. Acad. Sci. USA 110: 1363-1368.


  • Forsthoefel D.J., James, N.P., Escobar D.E., Stary, J.M., Vieira, A.P., Waters F.A., and P.A. Newmark. 2012. An RNAi screen reveals intestinal regulators of branching morphogenesis, differentiation, and stem cell proliferation in planarians. Dev. Cell 23: 691-704.
  • King, R.S. and P.A. Newmark. 2012. The cell biology of regeneration. J. Cell Biol. 196: 553-562.
  • Rouhana, L., Vieira, A.P., Roberts-Galbraith, R.H., and P.A. Newmark. 2012. PRMT5 and the role of symmetrical dimethylarginine in chromatoid bodies of planarian stem cells. Development 139: 1083-1094.
  • Miller, C.M. and P.A. Newmark. 2012. An insulin-like peptide regulates size and adult stem cells in planarians. Int. J. Dev. Biol. 56: 75-82.


  • Chong, T., Stary, J.M., Wang, Y., and P.A. Newmark. 2011. Molecular markers to characterize the hermaphroditic reproductive system of the planarian Schmidtea mediterraneaBMC Dev. Biol. 11: 69.
  • Forsthoefel, D.J., Park, A.E., and P.A. Newmark. 2011. Stem cell-based growth, regeneration, and remodeling of the planarian intestine. Dev. Biol. 356: 445-459.
  • Cogswell A.A., Collins J.J. III, Newmark, P.A., and D.L. Williams. 2011. Whole mount in situ hybridization methodology for Schistosoma mansoniMol. Biochem. Parasitol. 178: 46-50.
  • Collins J.J. III*, King R.S.*, Cogswell A., Williams D.L., and P.A. Newmark. 2011. An atlas for Schistosoma mansoni organs and life-cycle stages using cell type-specific markers and confocal microscopy. PLOS Negl. Trop. Dis. 5(3): e1009 (featured image).
    *authors contributed equally to the work


  • Collins, J.J., Hou, X., Romanova, E.V., Lambrus, B.G., Miller, C.M., Saberi, A., Sweedler, J.V., and P.A. Newmark. 2010. Genome-wide analyses reveal a role for peptide hormones in regulating planarian germline development. PLOS Biol. 8(10): e1000509.
  • Wang, Y., Stary, J.M., Wilhelm, J.E., and P.A. Newmark. 2010. A functional genomic screen in planarians identifies novel regulators of germ cell development. Genes Dev. 24: 2081-2092 (cover).
  • Zayas, R.M., Cebrià, F., Guo, T., Feng, J., and P.A. Newmark. 2010. The use of lectins as markers for differentiated secretory cells in planarians. Dev. Dynam. 239: 2888-2897.




  • Cebrià, F., Guo, T., Jopek, J., and P.A. Newmark. 2007. Regeneration and maintenance of the planarian midline is regulated by a slit orthologue. Dev. Biol. 307: 394-406 (cover).
  • Wang, Y., Zayas, R.M., Guo, T., and P.A. Newmark. 2007. nanos function is essential for development and regeneration of planarian germ cells. Proc. Natl Acad. Sci. 104: 5901-5906.
  • Cebrià, F. and P.A. Newmark. 2007. Morphogenesis defects are associated with abnormal nervous system regeneration following roboA RNAi in planarians. Development 134: 833-837.


  • Guo, T., Peters, A.H.F.M., and P.A. Newmark. 2006. A bruno-like gene is required for stem cell maintenance in planarians. Dev. Cell 11: 159-169 (cover).


  • Zayas, R.M., Hernández, A, Habermann, B., Wang, Y., Stary, J.M., and P.A. Newmark. 2005. The planarian Schmidtea mediterranea as a model for epigenetic germ cell specification: analysis of ESTs from the hermaphroditic strain. Proc. Natl. Acad. Sci. 102: 18491-18496.
  • Cebrià, F. and P.A. Newmark. 2005. Planarian homologues of netrin and netrin receptor are required for proper regeneration of the central nervous system and the maintenance of nervous system architecture. Development 132: 3691-3703.
  • Zayas, R.M., Bold, T.D., and P.A. Newmark. 2005. Spliced leader trans-splicing in freshwater planarians. Mol. Biol. Evol. 22: 2048-2054.
  • Newmark, P.A. 2005. Opening a new can of worms: a large-scale RNAi screen in planarians (Preview). Dev. Cell 8: 623-624.


  • Newmark, P.A., Reddien, P.W., Cebrià, F., and A. Sánchez Alvarado. 2003. Ingestion of bacterially expressed double-stranded RNA inhibits gene expression in planarians. Proc. Natl. Acad. Sci. 100 (suppl. 1): 11861-11865.
  • Oviedo, N., Newmark, P.A. and A. Sánchez Alvarado. 2003. Allometric scaling and proportion regulation in the freshwater planarian Schmidtea mediterraneaDev. Dyn. 226: 326-333.
  • Sánchez Alvarado, A.*, Newmark, P.A.*, Robb, S., and Juste, R. 2002. The Schmidtea mediterranea database as a molecular resource for studying platyhelminthes, stem cells and regeneration. Development 129: 5659-5665. *authors contributed equally to the work
  • Newmark, P.A. and A. Sánchez Alvarado. 2002. Not your father’s planarian: a classic model enters the era of functional genomics. Nat. Rev. Genet. 3: 210-219.
  • Newmark, P.A. and A. Sánchez Alvarado. 2001. Regeneration in planaria. In Encyclopedia of Life Sciences. Nature Publishing Group, London,
  • Newmark, P.A. and A. Sánchez Alvarado. 2000. Bromodeoxyuridine specifically labels the regenerative stem cells of planarians. Dev. Biol220: 142-153.
  • Sánchez Alvarado, A. and P.A. Newmark. 1999. Double-stranded RNA specifically disrupts gene expression during planarian regeneration. Proc. Natl. Acad. Sci. USA 96: 5049-5054.
  • Sánchez Alvarado, A. and P.A. Newmark. 1998. The use of planarians to dissect the molecular basis of metazoan regeneration. Wound Repair Regen6: 413-420.
  • Newmark, P.A. and A. Sánchez. 1997. Regeneration and scientific terminology. BioEssays 19: 535.