Gupta S., Heinrichs, E., Novitch, B.G. and S. J. Butler (2023). Investigating the basis of lineage decisions and developmental trajectories in the dorsal spinal cord through pseudotime analyses, Development, in revision  BioRxiv,

Alvarez, S., Gupta S., Honeychurch. K., Mercado-Ayon Y., Kawaguchi R. and S. J. Butler (2023).  Netrin1 patterns the dorsal spinal cord through modulation of Bmp signaling, Cell Reports, in revision  BioRxiv,


Gupta S., Kawaguchi R., Heinrichs, E., Gallardo, S., Castellanos, S., Mandric I., Novitch, B.G. and Butler S.J. (2022). In vitro atlas of dorsal spinal interneurons reveals Wnt signaling as a critical regulator of progenitor expansion, Cell Reports, 40:111119  doi: 10.1016/j.celrep.2022.111119


S.J. Butler (2021). The confidence to question. Science (Working Life) 374:230  doi: 10.1126/science.acx9307

Alvarez, S., Varadarajan, S.V. and S.J. Butler (2021). Dorsal commissural axon guidance in the developing spinal cord.  Current Topics in Developmental Biology: Neural Development and Disease.  142: 197-231

Gupta S., Yamauchi, K., Novitch, B.G. and S. J. Butler (2021). Derivation of dorsal spinal sensory interneurons from human pluripotent stem cells.  STAR Protocols.  2: 100319 10.1016/j.xpro.2021.100319

Gupta S. and S.J. Butler (2021).  Getting in touch with your senses: mechanisms specifying sensory interneurons in the dorsal spinal cord.  WIREs Mechanisms of Disease. e1520


Vonica, A., Bhat, N., Phan K.D., Lancu, L., O’Donell, A., Christiano, A.*, Riley B.*, Butler, S.J.* and A.V. Luria*  (2020).  APCDD1 is a dual BMP/WNT inhibitor in the developing nervous system and skin. Developmental Biology, 464:71-87
* co-corresponding authors


Frendo, M.E., da Silva, A., Phan, K.D., Riche, S. and S.J. Butler (2019).  The cofilin/Limk1 pathway controls the growth rate of both developing and regenerating motor axons.  J Neuroscience, DOI:

Comer, J., Alvarez, S., Butler, S.J. and J. Kaltschmidt.  (2019).  Commissural neurons in the developing spinal cord: from Cajal to the present day.  Neural Development, 14:9, DOI:

Andrews, M.G., Kong, J., Novitch, B.G, and S.J. Butler.  (2019). New perspectives on the mechanisms establishing the dorsal-ventral axis of the spinal cord. Current Topics in Developmental Biology: Organ Development 132: 417-450


Butler, S. J.  (2018).  The feel good factor. Laboratory News.

Gupta. S., Sivaligam, D., Hain, S., Makkar, C., Sosa, E., Clark. A. and S.J. Butler.  (2018). Deriving dorsal spinal interneurons from human pluripotent stem cells. Stem Cell Reports 10:1-16


Andrews, M.G., del Castillo, L.  M., Ochoa-Bolton, E., Yamauchi, K., Smogorzewski, J. and S.J.  Butler.  (2017).  BMPs direct sensory interneuron identity in the developing spinal cord using signal-specific not morphogenic activities. eLife 6:e30647 DOI: 10.7554/eLife.30647

Varadarajan, S.G and S. J. Butler.  (2017). Netrin1 establishes multiple boundaries for axon growth in the developing spinal cord.  Developmental Biology, 430: 177-187

Butler, S.J. (2017). The evolution of an axon guidance model: from chemotaxis to haptotaxis.  The Node (Development)

Varadarajan, S.G, Kong, J., Phan, K.D.,  Kao, T.-J., Panaitof, S.C., Cardin, J., Eltzschig, H.,, Kania, A., Novitch, B.G. and S. J. Butler.  (2017). Netrin1 produced by neural progenitors, not floor plate cells, is required for axon guidance in the spinal cord.  Neuron, 94:790-799


Butler, S.J. and M. E. Bronner. (2015). From classical to current: analyzing peripheral nervous system and spinal cord lineage and fate.  Developmental Biology 398:135-46.


Pearson, C.A, Butler, S.J and B. G. Novitch (2014). Neuronal organization: unsticking the cadherin code. Current Biology 24: R1127-9.

Kandyba, E., Hazen, V.M., Kobielak, A., Butler, S.J. and K. Kobielak. (2014). Smad1 and 5 but not Smad8 establish stem cell quiescence which is critical to transform the premature hair follicle during morphogenesis toward the postnatal state.  Stem Cells, 32: 534-547


Phan, K.D. and S.J. Butler  (2013). Bilaterally symmetric populations of chicken dI1 (commissural) axons cross the floor plate independently of each other.  PLoS One 8: e62977

Yamauchi, K., Varadarajan, S.G., Li. J., and S.J. Butler.  (2013).  Type Ib BMP receptors mediate the rate of commissural axon extension through inhibition of cofilin activity.  Development, 140:333-42

Kong, J., Butler S.J. and B.G. Novitch (2013). The brain told me to do it. Developmental Cell. 25:436-8

Gaber, Z.B., Butler, S.J. and B. G. Novitch (2013). PLZF regulates Fibroblast Growth Factor responsiveness and maintenance of neural progenitors. PLoS Biology 11::e1001676


Hazen, V.M., Andrews, M.G., Umans L., Crenshaw III, E.B, Zwijsen A., and Butler S.J.  (2012).  BMP receptor-activated Smads direct diverse functions during the development of the dorsal spinal cord.  Developmental Biology, 367: 216-227.


Hazen, V.M., Phan, K.D., Hudiburgh S. and S.J. Butler  (2011). Inhibitory Smads can differentially regulate cell fate specification and axon dynamics in the dorsal spinal cord.  Developmental Biology 336: 566-575.

Phan, K.D., Croteau, L.-P., Kam, J.W.K., Kania A., Cloutier, J.-F. and Butler S.J.  (2011).  Neogenin may functionally substitute for Dcc in chicken.  PLoS One 6: e22072


Phan, K.D., Hazen, V.M., Frendo, M.E., Jia, Z. and S.J. Butler.  (2010).  The bone morphogenetic protein roof plate chemorepellent regulates the rate of commissural axonal growth.  J. Neuroscience 30: 15430-40

Hazen, V.M., Phan K.D., Yamauchi K. and S. J. Butler (2010).  Assaying the ability of diffusible signaling molecules to reorient embryonic spinal commissural axons.  J Vis Exp. 37:1853


Novitch, B.G., and S.J. Butler. (2009).  Reducing the mystery of neuronal differentiation.  Cell 138: 1062-1064


Yamauchi, K., Phan, K.D. and S.J. Butler.  (2008).  BMP type I receptors have distinct activities in mediating cell fate and axon guidance. Development 135:1119-28.


Butler, S.J. and G. Tear.  (2007).  Getting axons onto the right path: the role of transcription factors in axon guidance.  Development 134: 439-448.

before 2007

Butler, S.J. and J. Dodd.  (2003).  A role for BMP heterodimers in roof plate-mediated repulsion of commissural axons.  Neuron 38: 389-401

Augsburger, A*., Schuchardt, A., Hoskins, S., Dodd, J. and S. J. Butler*,   (1999).  BMPs as mediators of roof plate repulsion of commissural neurons.  Neuron 24: 127-141.
* joint first authors

Butler, S.J., Ray, S. and Y. Hiromi.  (1997).  klingon, a novel member of the Drosophila immunoglobulin superfamily, is required for the development of the R7 neuron.  Development 124: 781-792.