Cheng LC, Alvarado AS. Whole-Mount BrdU Staining with Fluorescence In Situ Hybridization in Planarians. Methods Mol Biol. 2018;1774:423-434. doi: 10.1007/978-1-4939-7802-1_15.

Merryman MS, Alvarado AS, Jenkin JC. Culturing Planarians in the Laboratory. Methods Mol Biol. 2018;1774:241-258. doi: 10.1007/978-1-4939-7802-1_5.

Zhang S, Alvarado AS. Planarian High Molecular Weight DNA Isolation by Spooling. Methods Mol Biol. 2018;1774:277-284. doi: 10.1007/978-1-4939-7802-1_8.

Adler CE, Alvarado AS. Systemic RNA Interference in Planarians by Feeding of dsRNA Containing Bacteria. Methods Mol Biol. 2018;1774:445-454. doi: 10.1007/978-1-4939-7802-1_17.

Zeng A, Li H, Guo L, Gao X, McKinney S, Wang Y, Yu Z, Park J, Semerad C, Ross E, Cheng LC, Davies E, Lei K,Wang W, Perera A, Hall K, Peak A, Box A, Sánchez Alvarado A. Prospectively Isolated Tetraspanin Neoblasts Are Adult Pluripotent Stem Cells Underlying Planaria Regeneration. Cell. 2018 Jun 14;173(7):1593-1608.e20. doi: 10.1016/j.cell.2018.05.006.

Guo L, Accorsi A, He S, Guerrero-Hernández C, Sivagnanam S, McKinney S, Gibson M, Sánchez Alvarado A. An adaptable chromosome preparation methodology for use in invertebrate research organisms. BMC Biol. 2018 Feb 26;16(1):25. doi: 10.1186/s12915-018-0497-4.

Cheng LC, Tu KC, Seidel CW, Robb SMC, Guo F, Sánchez Alvarado A. Cellular, ultrastructural and molecular analyses of epidermal cell development in the planarian Schmidtea mediterranea. Dev Biol. 2018 Jan 15;433(2):357-373. doi: 10.1016/j.ydbio.2017.08.030. Epub 2017 Oct 31.

Sánchez Alvarado A. To solve old problems, study new research organisms. Dev Biol. 2018 Jan 15;433(2):111-114. doi: 10.1016/j.ydbio.2017.09.018. Epub 2017 Nov 24. No abstract available.


Sasidharan V, Marepally S, Elliott SA, Baid S, Lakshmanan V, Nayyar N, Bansal D, Sánchez Alvarado A, Vemula PK, Palakodeti D. The miR-124 family of microRNAs is crucial for regeneration of the brain and visual system in the planarian Schmidtea mediterranea. Development. 2017 Sep 15;144(18):3211-3223. doi: 10.1242/dev.144758. Epub 2017 Aug 14.

Adler CE, Sánchez Alvarado A. PHRED-1 is a divergent neurexin-1 homolog that organizes muscle fibers and patterns organs during regeneration. Dev Biol. 2017 Jul 1;427(1):165-175. doi: 10.1016/j.ydbio.2017.04.012. Epub 2017 Apr 29.

Davies EL, Lei K, Seidel CW, Kroesen AE, McKinney SA, Guo L, Robb SM, Ross EJ, Gotting K, Alvarado AS. Embryonic origin of adult stem cells required for tissue homeostasis and regeneration. Elife. 2017 Jan 10;6. pii: e21052. doi: 10.7554/eLife.21052.


Sánchez Alvarado, A. The joy of figuring things out: a story of worms, haplotypes & genetic ancestry. Dec 05, 2016

Stephanie H. Nowotarski and Alejandro Sánchez Alvarado (2016) Widening perspectives on regenerative processes through growth. npj Regenerative Medicine (2016) 1, 16015; doi:10.1038/npjregenmed.2016.15; published online 8 December 2016

Guo, L., Zhang, S., Rubinstein, B., Ross, E. & Sánchez Alvarado, A. Widespread maintenance of genome heterozygosity in Schmidtea mediterranea. Nat. Ecol. Evol. 1, 0019 (2016 Dec 5).

Luttrell SM, Gotting K, Ross E, Sánchez Alvarado A, Swalla BJ. Head regeneration in hemichordates is not a strict recapitulation of development. Dev Dyn. 2016 Dec;245(12):1159-1175. doi: 10.1002/dvdy.24457.

Lei K, Thi-Kim Vu H, Mohan RD, McKinney SA, Seidel CW, Alexander R, Gotting K, Workman JL, Sánchez Alvarado A. Egf Signaling Directs Neoblast Repopulation by Regulating Asymmetric Cell Division in Planarians. Dev Cell. 2016 Aug 22;38(4):413-29. doi: 10.1016/j.devcel.2016.07.012.

Arnold CP, Merryman MS, Harris-Arnold A, McKinney SA, Seidel CW, Loethen S, Proctor KN, Guo L, Sánchez Alvarado A. Pathogenic shifts in endogenous microbiota impede tissue regeneration via distinct activation of TAK1/MKK/p38. eLife. 2016 Jul 21;5. pii: e16793. doi: 10.7554/eLife.16793.

Eric Ross, David Blair, Carlos Guerrero Hernández and Alejandro Sánchez Alvarado (2016) Comparative and Transcriptome Analyses Uncover Key Aspects of Coding and Long Noncoding RNAs in Flatworm Mitochondrial Genomes. G3 (Bethesda) 2016 May 3;6(5): 1191-200. doi:10.1534/g3. 116.028175.


Elizabeth M. Duncan, Alex D. Chitsazan, Chris W. Seidel and Alejandro Sánchez Alvarado (2015) Set1 and MLL1/2 Target Distinct Sets of Functionally Different Genomic Loci In Vivo. Cell Reports 2015; doi:10.1016/j.celrep.2015.11.059

Tu KC, Cheng LC, T K Vu H, Lange JJ, McKinney SA, Seidel CW, Sánchez Alvarado A. Egr-5 is a post-mitotic regulator of planarian epidermal differentiation. elife. 2015 Oct 12;4:e10501. doi: 10.7554/eLife.10501.

Carolyn E. Adler and Alejandro Sánchez Alvarado (2015) Types or States? Cellular Dynamics and Regenerative Potential. Trends in Cell Biology 2015; doi: 10.1002/dvg.22872

Sofia M. C. Robb, Kirsten Gotting, Eric Ross and Alejandro Sánchez Alvarado (2015) SmedGD 2.0: The Schmidtea mediterranea genome database. genesis 2015; doi: 10.1002/dvg.22872

Hanh Thi-Kim Vu, Jochen C. Rink, Sean A. McKinney, Melainia McClain, Naharajan Lakshmanaperumal, Richard Alexander and Alejandro Sánchez Alvarado (2015) Stem cells and fluid flow drive cyst formation in an invertebrate excretory organ. eLife 2015; doi: 10.7554/eLife.07405

Sánchez Alvarado, A. Unravelling a can of worms. eLife. 2015; 4: e07431. Published online 2015 Apr 16. doi: 10.7554/eLife.07431


Youbin Xiang, Danny E. Miller, Eric J. Ross, Alejandro Sánchez Alvarado, and R. Scott Hawley (2014) Synaptonemal complex extension from clustered telomeres mediates full-length chromosome pairing in Schmidtea mediterranea. PNAS 2014; doi: 10.1073/pnas.1420287111

Carolyn E. Adler, Chris W. Seidel, Sean McKinney and Alejandro Sánchez Alvarado (2014) Selective amputation of the pharynx identifies a FoxA-dependent regeneration program in planaria. eLife 2014;3:e02238. (See editorial Insight by Dr. Janet Rossant)

Alejandro Sánchez Alvarado and Shinya Yamanaka (2014) Rethinking differentiation: stem cells, regeneration and plasticity. Cell, 2014; 157:110-119. doi:10.1016/j.cell.2014.02.041

Sofia MC Robb and Alejandro Sánchez Alvarado (2014) Histone Modifications and Regeneration in the Planarian Schmidtea mediterranea. Current Topics in Developmental Biology, 2014;108:71-93. doi: 10.1016/B978-0-12-391498-9.00004-8.


Alessandro Rossi, Eric J. Ross, Antonia Jack, and Alejandro Sánchez Alvarado (2013) Molecular cloning and characterization of SL3: A stem cell-specific SL RNA from the planarian Schmidtea mediterranea.Gene, 2013 Ocotober 8

Alejandro Sánchez Alvarado (2013) Learning about loss. eLife, 2013 Sep 10;2:e00533

Alejandro Sánchez Alvarado (2013) On the trail of a tropical disease. eLife, 2013 Jul 30;2:e01115.


Alejandro Sánchez Alvarado (2012) What is regeneration, and why look to planarians for answers? BMC Biology, 2012, 10:88 doi:10.1186/1741-7007-10-88

Alejandro Sánchez Alvarado (2012) Cellular hyperproliferation and cancer as evolutionary variables Current Biology, 2012 Septemeber 11, doi: 10.1016/j.cub.2012.08.008

Otto C. Guedelhoefer IV and Alejandro Sánchez Alvarado (2012) Amputation induces stem cell mobilization to sites of injury during planarian regeneration Development, 2012 August 16, doi: 10.1242/dev.082099

Otto C. Guedelhoefer IV and Alejandro Sánchez Alvarado (2012) Planarian immobilization, partial irradiation, and tissue transplantation Journal of Visualized Experiments (JoVE), 2012 August 5, doi: 10.3791/4015

Sarah A. Elliot C. and Alejandro Sánchez Alvarado (2012) The history and enduring contributions of planarians to the study of animal regeneration WIREs Developmental Biology, 2012 July 24, doi: 10.1002/wdev.82

Kimberly C. Tu, Bret J. Pearson, Alejandro Sánchez Alvarado (2012) TORC1 is required to balance cell proliferation and cell death in planarians Developmental Biology, 2012 March 16, doi: 10.1016/j.ydbio.2012.03.010

Cheryl E. Bender, Patrick Fitzgerald, Stephen W. Tait, Fabien Llambi, Gavin P. McStay, Douglas O. Tupper, Jason Pellettieri, Alejandro Sánchez Alvarado, Guy S. Salvesena, and Douglas R. Green (2012) Mitochondrial pathway of apoptosis is ancestral in metazoans PNAS, 2012 March 13.

Juliette Azimzadeh, Mei Lie Wong, Diane Miller Downhour, Alejandro Sánchez Alvarado and Wallace F. Marshall (2012) Centrosome loss in the evolution of planarians reveals the molecular architecture of the human centrosome Science, 2012 January 6, doi: 10.1126/science.1214457


Jochen C. Rink, Hanh Thi-Kim Vu and Alejandro Sánchez Alvarado (2011) The maintenance and regeneration of the planarian excretory system are regulated by EGFR signaling Development, 2011 August 8, doi: 10.1242/dev.066852

Catherine Adamidi, Yongbo Wang, Dominic Gruen, Guido Mastrobuoni, Xintian You, Dominic Tolle, Matthias Dodt, Sebastian D. Mackowiak, Andreas Gogol-Doering, Pinar Oenal, Agnieszka Rybak, Eric Ross, Alejandro Sánchez Alvarado, Stefan Kempa, Christoph Dieterich, Nikolaus Rajewsky, and Wei Chen (2011) De novo assembly and validation of planaria transcriptome by massive parallel sequencing and shotgun proteomics Genome Research, 2011 May 2, doi:10.1101/gr.113779.110


Kyle A. Gurley, Sarah A. Elliott, Oleg Simakov, Heiko A. Schmidt, Thomas W. Holstein and Alejandro Sánchez Alvarado (2010) Expression of secreted Wnt pathway components reveals unexpected complexity of the planarian amputation response Developmental Biology, 2010 August 10, doi:10.1016/j.ydbio.2010.08.007


Bret J. Pearson and Alejandro Sánchez Alvarado (2009) A Planarian p53 Homolog Regulates Proliferation and Self-Renewal in Adult Stem Cell Lineages Development, 2009 December 29, doi: 10.1242/dev.044297.

Jochen C. Rink, Kyle A. Gurley, Sarah A. Elliott, and Alejandro Sánchez Alvarado (2009) Planarian Hh signaling regulates regeneration polarity and links Hh pathway evolution to cilia Science, 2009 October 22

Jason Pellettieri, Patrick Fitzgerald, Shigeki Watanabe, Joel Mancuso, Douglas R. Green, Alejandro Sánchez Alvarado (2009) Cell Death and Tissue Remodeling in Planarian Regeneration Developmental Biology, 2009 September 11, doi:10.1016/j.ydbio.2009.09.015

Alejandro Sánchez Alvarado (2009) A Cellular View of Regeneration Nature, 460:39-40

Marc R. Friedländer, Catherine Adamidi, Ting Han, Svetlana Lebedeva, Thomas A. Isenbarger, Martin Hirst, Marco Marra, Chad Nusbaum, William L. Lee, James C. Jenkin, Alejandro Sánchez Alvarado, John K. Kim, and Nikolaus Rajewsky (2009) High-resolution profiling and discovery of planarian small RNAs PNAS, 2009 June 29, doi:10.1073/pnas.0905222106

Hara Kang and Alejandro Sánchez Alvarado (2009) Flow Cytometry Methods for the Study of Cell-cycle Parameters of Planarian Stem Cells Developmental Dynamics, 2009 Mar 25

Bret J. Pearson, George T. Eisenhoffer, Kyle A. Gurley, Jochen C. Rink, Diane E. Miller, and Alejandro Sánchez Alvarado (2009) A Formaldehyde-based Whole-Mount In Situ Hybridization Method for Planarians Developmental Dynamics, 238:443-450, 2009.

Bret J. Pearson and Alejandro Sánchez Alvarado (2009) Regeneration, Stem Cells, and the Evolution of Tumor Suppression Cold Spring Harb Symp Quant Biol. 2009 Jan 15.


Alejandro Sánchez Alvarado (2008) The Shredding of a Caricature Cell, 135:991-992, December 12, 2008

Kyle A. Gurley and Alejandro Sánchez Alvarado (2008) Stem Cells in Animal Models of Regeneration StemBook, ed. The Stem Cell Research Community, StemBook, doi/10.3824/stembook.1.32.1 

Néstor J. Oviedo, Bret J. Pearson, Michael Levin and Alejandro Sánchez Alvarado (2008) Planarian PTEN Homologs Regulate Stem Cells and Regeneration Through TOR Signaling Disease Models and Mechanisms, 1:131-143, Published online: September 18, 2008

George T. Eisenhoffer, Hara Kang, and Alejandro Sánchez Alvarado (2008) Molecular Analysis of Stem Cells and Their Descendants during Cell Turnover and Regeneration in the Planarian Schmidtea mediterranea Cell Stem Cell, 3:327-339

Alejandro Sánchez Alvarado (2008) Stem Cells: Time to Check Our Premises Cell Stem Cell, 3:25-9

Reddien, P.W., Newmark, P.A., and Alejandro Sánchez Alvarado (2008) Gene Nomenclature Guidelines for the Planarian Schmidtea mediterranea Developmental Dynamics, DOI:10.1002/dvdy.21623, Published online: July 15, 2008. 

Kenneth D. Birnbaum and Alejandro Sánchez Alvarado (2008) Slicing Across Kingdoms: Regeneration in Plants and Animals Cell, 132:697-710


Brandi Cantarel, Ian Korf, Sofia M. C. Robb, Genis Parra, Eric Ross, Barry Moore, Carson Holt, Alejandro Sánchez Alvarado, and Mark Yandell (2008) MAKER: An Easy-to-use Annotation Pipeline Designed for Emerging Model Organism Genomes Genome Research, 18:188-196 (e-publication date: November 19, 2007)

Jason Pellettieri and Alejandro Sánchez Alvarado (2007) Cell Turnover and Adult Tissue Homeostasis: From Humans to Planarians Annual Review of Genetics. 41:83-105.

Sofia M.C. Robb, Eric Ross and Alejandro Sánchez Alvarado (2008) SmedGD: the Schmidtea mediterranea Genome Database Nucleic Acids Research, 36:D599-D606, doi:10.1093/nar/gkm684  (e-publication date: September 18, 2007)

Peter W. Reddien, Adam L. Bermange, Adrienne M. Kicza and Alejandro Sánchez Alvarado (2007) BMP signaling regulates the dorsal planarian midline and is needed for asymmetric regeneration Development. 134:4043-4051 ( See Journal’s Preview )

Kyle A. Gurley, Jochen C. Rink and Alejandro Sánchez Alvarado (2007) ß-Catenin Defines Head Versus Tail Identity During Planarian Regeneration and Homeostasis Science, 319:323-327   (Editor’s Choice)


Alejandro Sánchez Alvarado and Panagiotis A. Tsonis (2006) Bridging the Regeneration Gap: Genetic Insights from Diverse Animal Models Nature Reviews | Genetics, 7:873:884.

Allan Spradling, Barry Ganetsky, Phil Hieter, Mark Johnston, Maynard Olson, Terry Orr-Weaver, Janet Rossant, Alejandro Sánchez Alvarado and Robert Waterston (2006) New Roles for Model Genetic Organisms in Understanding and Treating Human Disease: Report From The 2006 Genetics Society of America Meeting Genetics 172(4): 2025�2032.

Anita M. Orendt, B.H., David Richardson, Sofia Robb, Alejandro Sánchez Alvarado and Julio C. Facelli (2006) Design, Implementation and Deployment of a Commodity Cluster for Periodic Comparisons of Gene Sequences, in High-Performance Computing Paradigm and Infrastructure M.G. Laurence, T. Yang, Editor. 733-744.

Alejandro Sánchez Alvarado (2006) Planarian Regeneration: Its End is Its Beginning Cell. 124(2):241-5.


Peter W. Reddien, Néstor J. Oviedo, Joya R. Jennings, James C. Jenkin, and Alejandro Sánchez Alvarado (2005) SMEDWI-2 Is a PIWI-Like Protein That Regulates Planarian Stem Cells Science. 310:1327-1330.

Alejandro Sánchez Alvarado and Hara Kang. (2005) Multicellularity, stem cells, and the neoblasts of the planarian Schmidtea mediterranea Exp. Cell Res.. 306:299-308. 

Peter W. Reddien, Adam L. Bermange, Kenneth J. Murfitt, Joya R. Jennings and Alejandro Sánchez Alvarado (2005) Identification of Genes Needed for Regeneration, Stem Cell Function, and Tissue Homeostasis by Systematic Perturbation in Planarians Developmental Cell. 8:635-649.  ( See Journal’s Preview )


Peter W. Reddien and Alejandro Sánchez Alvarado (2004) Fundamentals of Planarian Regeneration Annual Review of Cell and Developmental Biology. 20:725-757. 

Alejandro Sánchez Alvarado (2004) Planarians Current Biology. 14:R737-8. 

Alejandro Sánchez Alvarado (2004) Regeneration and the need for simpler model organisms Philosophical Transactions of the Royal Society of London. Series B 359: 759-763.

Takeshi Inoue, Hiroshi Kimamoto, Keiji Okamoto, Yoshihiko Umesono, Hiroshi Kimamoto, Masaki Sakai, Alejandro Sánchez Alvarado and Kiyokazu Agata (2004) Morphological and Functional Recovery of the Planarian Photosensing System during Head Regeneration Zoological Science. 21: 275-283.  


Phillip A. Newmark, Peter W. Reddien, Francesc Cebrià and Alejandro Sánchez Alvarado (2003) Ingestion of bacterially expressed double-stranded RNA inhibits gene expression in planarians Proceedings of the National Academy of Sciences USA. 100 (Suppl 1): 11861-5.  

Alejandro Sánchez Alvarado (2003) The freshwater planarian Schmidtea mediterranea: embryogenesis, stem cells and regeneration Current Opinion in Genetics and Development. 13:438-444. 

Néstor J. Oviedo, Phillip A. Newmark, and Alejandro Sánchez Alvarado (2003) Allometric Scaling and Proportion Regulation in the Freshwater Planarian Schmidtea mediterranea Developmental Dynamics. 226:326-333.  


Alejandro Sánchez Alvarado, Phillip A. Newmark, Sofia M. C. Robb and Réjeanne Juste (2002) The Schmidtea mediterranea database as a molecular resource for studying platyhelminthes, stem cells and regeneration Development. 129:5659-5665.  

Francesc Cebrià, Chiyoko Kobayashi, Yoshihiko Umesono, Masumi Nakazawa, Katsuhiko Mineta, Kazuho Ikeo, Takashi Gojobori, Mari Itohk, Masanori Tairak, Alejandro Sánchez Alvarado, Kiyokazu Agata (2002) FGFR-related gene noudarake restricts brain tissues to the head region of planarians Nature. 419:620-624.  

Phillip A. Newmark and Alejandro Sánchez Alvarado (2002) Not Your Father’s Planarian: A Classic Model Enters The Era of Functional Genomics Nature Reviews | Genetics. 3:210-220.  

Sofia M.C. Robb and Alejandro Sánchez Alvarado (2002) Identification of immunological reagents for use in the study of freshwater planarians by means of whole-mount immunofluorescence and confocal microscopy Genesis:Journal of Genetics and Development. 34(4):293-298.  



Alejandro Sánchez Alvarado (2000) Regeneration in the Metazoans: Why Does it Happen? BioEssays. 22:578-590.  

Phillip A. Newmark and Alejandro Sánchez Alvarado (2000) Bromodeoxyuridine Specifically Labels the Regenerative Stem Cells of Planarians Developmental Biology. 220:142-153.  

Alejandro Sánchez Alvarado (2000) The Case for Comparative Regeneration: Learning from Simpler Organisms How to Make New Parts from Old e-biomed: The Journal of Regenerative Medicine. 1: 31-36.  


Alejandro Sánchez Alvarado and Phillip A. Newmark (1998) The Use of Planarians to Dissect the Molecular Basis of Metazoan Regeneration Wound Repair and Regeneration. 6: 413-420.  


Alejandro Sánchez Alvarado and Phillip A. Newmark (1998) The Use of Planarians to Dissect the Molecular Basis of Metazoan Regeneration Wound Repair and Regeneration. 6: 413-420.