Publications from the Freeman Lab


Ma, Z., Stork, T., Bergles, DE. And M.R. Freeman. (2016) Neuromodulators signal through astrocytes to alter neural circuit activity and behavior. Nature. 2016Nov9,539(7629):428-432. doi: 10.11038/nature20145. PMC5161596

He, J., Zhou, R., Wu, Z., Carrasco, M.A., Kurshan, P.T., Farley, J.E., Simon, D.J., Wang, G.,Han, B., Hao, J., Heller, E., M.R. Freeman, Shen, K., Maniatis, T., Tessier-Lavigne, M., Zhuang, X. (2016) Prevalent presence of periodic actin-spectrin-based membrance skeleton in a broad range of neuronal cell types and animal specific.  Proc Natl Acad Sci USA. 2016 May 24: 113(21): 6029-34. doi: 10.01073/pnas.1605707113.  Epub 2016 May 9. PMC4889411

Francis, M.M. and M.R.Freeman. (2016) Dendrites actively restrain axon outgrowth and regeneration. Proc Natl Acad Sci USA. 2016 May 17; 113(2): 5465-6. doi: 10.1073/pnas.1605215113.  PMC4878526

Henninger, N., Bouley, J., Sikoglu, E.M., An, J., Moore, C.M., King, J.A., Bowser, R., M.R.Freeman, Brown, R.H.Jr. (2016) Attenuated traumatic axonal injury and improved functional outcome after traumatic brain injury in mice lacking Sarm1. Brain. 2016Feb11. [epub ahead of print]


Freeman, M.R. (2015) Drosphila Central Nervous System Glia.  Cold Spring Harbor Perspect Biol. Feb 26. pii:a020552 doi: 10.1101/cshperspect.a080552.  [Epub ahead of print]

Farely, J.E.and M.R. Freeman. (2015) Letting go of JuNK by disassembly of adhesive complexes. Neuron. 2015Dec2:88(5):848-50. doi:10.1016/j.neuron.2015.11.015.

Sreedharan, J., Neukomm, L.J., Brown, R.H. Jr., and M.R. Freeman. (2015) Age-dependent TRDP-43-medicated motor neuron degeneration requires GSK3, hat-trick, and xmas-2. Curr Bio. 2015Aug17:25(16):2130-6. doi: 10.1016/j.cub.2015.06.045. epub 2015Jul30.  PMC4546534.


Doherty, J., Sheehan, A.E., Bradshaw, R., Fox, A.N., Lu, T.Y. and M.R. Freeman. (2014) PI3K signaling and stat92E converge to modulate glial responsiveness to axonal injury. PLoS Biol. 2014 Nov4;12(11): e1001985.  doi:10.1371/journal.pbio.1001985. eCollection2014Nov.  PMC4219656.

Muthumukar, A.K., Stork, T. and M.R. Freeman. (2014) Activity-dependent regulation of astrocyte GAT levels during synaptogenesis. Nat Neurosci. Aug 24. doi:10.1038/nn.3791. [Epub ahead of print]

Lu,T.Y., Doherty, J. and M.R. Freeman. (2014)  DRK/DOS/SOS converge with Crk/Mbc/dCed-12 to activate Rac1 during glial engulfment of aconal debris.  Proc Natil Adac Sci.  

Burdett, T.C. and M.R. Freeman.(2014)  Astrocytes eyeball axonal mitochondria. Science. Jul 25; 345 (6195): 385-6. doi: 10.1126/science.1258295. 

Freeman, M.R. (2014) Signaling mechanism regulating Wallerian degeneration. Curr Opin Neurobiol.  Aug;27C:224-231. doi:10.1016/j.conb.2014.05.001. PMC4122608

Neukomm, L.J. and M.R. Freeman.  (2014)  Diverse cellular and molecular modes of axon degeneration. Trends in Cell Biology  doi:10.1015/j.tcb.2014.04.003 [Epub ahead of print]

Stork, T., Sheehan, A., Tasdemir-Yilmaz, O.E., and M.R. Freeman. (2014) Neuron-Glia Interactions through the Heartless FGF Receptor Signaling Pathway Mediate Morphogenesis of Drosophila Astrocytes.  Neuron.  Jul 16;83(2):388-40. doi: 10.1016/j.neuron.2014.06.026. 

Rooney, T.M. and M.R. Freeman. (2014) Drosophila models of neuronal injury.  ILAR J.  54(3):  291-5.  doi: 10.1093/ilar/ilt057.  PMC3962260.

Neukomm, L.J., Burdett, T.C, Gonzalez, M.A., Zuchner, S., Freeman, M.R. (2014) Rapid in vivo forward genetic approach for identifying axon death genes in Drosophila. Proc Natl Acad Sci U S A. 2014 Jun 23. pii: 201406230. [Epub ahead of print]

Kerr, K.S., Fuentes-Medel, Y., Brewer, C., Barria, R., Ashley J., Abruzzi, K.C., Sheehan, A., Tasdemir-Yilmaz, O.E., Freeman, M.R.*, and V. Budnik* (2014), Glial wingless/Wnt regulates glutamate receptor clustering and synaptic physiology at the Drosophila neuromuscular junction.  J Neurosci. 34:  2910-20.  doi:  10.1523/JNEUROSCI.3414-13.2014. *corresponding authors PMC3931504.

Tasdemir-Yilmaz, O.E., and M.R. Freeman (2014) Astrocytes engage unique molecular programs to engulf pruned neuronal debris from distinct subsets of neurons.  Genes Dev. 28: 20-33., doi:10.1101/gad.229518.113.  PMC3894410.


Corty, M.M. and M.R. Freeman. (2013) Cell biology in neuroscience:  Architects in neural circuit design: glia control neuron numbers and connectivity.  J. Cell Biology.  203(3): 395-405. doi: 10.1083.  PMC3824021.

Milde, S., Fox, A.N., Freeman, M.R., and Coleman, M.P. (2013) Deletions within its subcellular targeting domain enhance the axon protective capacity of Nmnat2 in vivo.  Sci Rep. 3: 2567.  doi: 10.1038/srep02567.  PMC3759051.

Coutinho-Budd, J. and M.R. Freeman.  (2013) Probing the enigma: Unraveling glial cell biology in invertebrates.  Current Opinion in Neurobiology. 23(6): 1073-9. doi: 10.1016/j.conb.2013.07.002.  Epub 2013 Jul 26.  PMC3830651.

Freeman, M.R. and D.H. Rowitch.  (2013)  Evolving concepts in gliogenesis: a look way back and ahead to the next 25 years.  Neuron, 80(3): 613-23. doi: 10.1016/j.neuron.2013.10.034.


Osterloh, J.M., Wang, J., Rooney, T.M., Adalbert, R., Powell, E.H.,  Avery, M.A., Logan, M.A., MacDonald, J.M., Ziegenfuss, J., Sheehan, A.E., Fox, A.N., Ding, A., Zuchner, S. Coleman, M.P., Tessier-Lavigne, M., and M.R. Freeman.  (2012) dSarm/Sarm1 is required for activation of an injury-induced axon death pathway.  Science, 337: 481-4.

MacDonald, J.M., Doherty, J., Hackett, R., and Freeman, M.R. (2013)  The c-Jun kinase signaling cascade promotes glial engulfment activity through activation of draper and phagocytic function.  Cell Death and Differentiation, 20: 1140-8. doi: 10.1038

Gonzalez, M.A., Van Booven, D., Hulme, W., Ulloa, R.H., Acosta-Lebrigio R., Osterloh, J., Logan, M., Freeman, M., and S. Zuchner.  (2012) Whole Genome Sequencing and a New Bioinformatics Platform Allow for Rapid Gene Identification in D. melanogaster EMS Screens. Biology, 1: 766-777; doi:10.3390/biology1030766

Stork, T., Bernardos, R. and M.R. Freeman.  (2012) Analysis of glial cell development and function in Drosophila.  Cole Spring Harbor Protoc.  2012: 1-17  doi:10.1101/pdb.top067587.

Fuentes-Medel, Y., Ashley, J., Barria, R., Maloney, R., Freeman, M.R.*, Budnik V* (2012)  Integration of a retrograde signal during synapse formation by glia-secreted TGF-ß ligand.  Current Biology,  22: 1831-8.  *corresponding/senior authors.  PMC3065899.

Ziegenfuss, J., Doherty, J., and M.R. Freeman. (2012) Distinct molecular pathways mediate glial activation and engulfment of axonal debris after axotomy.  Nature Neuroscience, 15: 979-87 doi:10.1038/nn.3135.

Avery, M., Rooney, T.M., Wishart, T.M., Gillingwater, Pandya, J.D., Geddes, J.W., Sullivan, P., and M.R. Freeman. (2012)  WldS protects axons through increased mitochondrial flux and enhanced mitochondrial Ca2+ buffering.  Current Biology, 22: 596-600.

Logan, M.A., Hackett, R., Doherty, J., Sheehan, A., and  M.R. Freeman.  (2012) Negative regulation of glial engulfment activity by Draper and Corkscrew terminates glial responses to axon injury. Nature Neuroscience 15: 722-30. PMC3337949.

Wishart T.M., Rooney T.M., Lamont D.J., Wright A.K., Morton A.J., Jackson M., Freeman M.R., Gillingwater T.H. (2012) Combining comparative proteomics and molecular genetics uncovers regulators of synaptic and axonal stability and degeneration in vivo. PLoS Genetics, 8(8):e1002936. PMC3431337.


McPhee, C.K., Logan, M.A., Freeman, M.R., and E.H. Baehrecke (2010) Activation of autophagy during cell death requires the engulfment receptor Draper.  Nature, 465: 1093-6.  PMC2892814.

Freeman, M.R.  (2010)  Specification and morphogenesis of astrocytes.  Science, 330: 774-778.

Coleman M.P. and M.R. Freeman (2010) Wallerian degeneration, WldS, and Nmnat.  Annual Reviews of Neuroscience, 33: 245-67.


Fuentes-Medel, Y., Logan, M.A., Ashley, J., Ataman, B., *Budnik, V., and *M.R. Freeman (2009) Glia and muscle sculpt neuromuscular arbors by engulfing destablilized synaptic boutons and shed presynaptic debris. PLoS-Biology, 7:e1000184. *corresponding/senior authors.  PMC2724735.

 Doherty, J., Logan, M.A., Tasdemir, O., and M.R. Freeman (2009)  Ensheathing glia function as phagocytes in the adult Drosophila brain.  J. Neuroscience, 29: 4768-81.  PMC2674269.

Osterloh, J.M., and M.R. Freeman (2009)  Neuronal death and dismemberment mediated by Sox14.  Nature Neuroscience 12, 1479-80.

Avery, M.A., Tasdemir, O., Sheehan, A., Wang, J., and M.R. Freeman (2009) Wlds requires Nmnat1-dependent NAD biosynthetic activity and N16-VCP interactions to suppress Wallerian degeneration.  J. Cell Biology 184: 501-513.  PMC2654119.


Ziegenfuss, J.S., Biswas, R., Avery, M.A., Hong, K., Sheehan, A.E., Yeung, Y.G., Stanley, E.R., and M.R. Freeman  (2008)  Draper-dependent glial phagocytic activity is mediated by Src family kinase signaling cascade.  Nature, 453: 935-9.   PMC2493287.


Emery, P. and M.R. Freeman (2007)  Glia got rhythm.  Neuron 55, 337-339.

Logan, M.A. and M.R. Freeman (2007)  The scoop on the fly brain:  Glial engulfment functions in DrosophilaNeuron Glia Biology 3: 63-74.  PMC2171361.


Freeman, M.R.  (2006)  Sculpting the nervous system:  Glial control of neuronal development.  Current Opinion in Neurobiology 16: 119-125.

Freeman, M.R. and J. Doherty (2006)  Glial cell biology in Drosophila and mammals.  Trends in Neurosciences 29: 82-90.

MacDonald, J.M., Beach M.G., Popiglia, E., Sheehan, A.E., Watts, R.J., and M.R.Freeman.  (2006) The Drosophila cell corpse engulfment receptor Draper mediates glial clearance of severed axons.  Neuron, 50: 869-881.


Freeman, M.R.  (2005)  Glial control of synaptogenesis.  Cell  120, 292-293.

Freeman, M.R. (2005)  Glial (and neuronal) cells missing.  Neuron 48, 163-165.


Freeman, M.R., Delrow, J., Kim, J., Johnson, E., and C.Q. Doe  (2003)  Unwrapping glial biology:  Gcm target genes regulating glial development, diversification, and function.  Neuron  38: 567-580.


Freeman, M.R., and C.Q. Doe (2001) Asymmetric Prospero localization is required to generate mixed neuronal/glial lineages in the Drosophila CNS.  Development 128: 4103-12.

Freeman, M.R., and C.Q. Doe (2001)  Moving muscle: Jeb signaling in DrosophilaDevelopmental Cell 1, 587-8.


Freeman, M.R., A. Dobritsa, P. Gaines, W.A. Segraves, and J.R. Carlson (1999)  The dare gene: Steroid hormone production, olfactory behavior, and neural degeneration in DrosophilaDevelopment 126: 4591-602.

Buszczak, M., M.R. Freeman, J.R. Carlson, M. Bender, L. Cooley, and W.A. Segraves (1999)  Ecdysone response genes govern egg chamber development during mid-oogenesis in DrosophilaDevelopment 126: 4581-4589.

Clyne, P.J., C.G. Warr, M.R. Freeman, D. Lessing, J. Kim, and J.R. Carlson (1999) A novel family of divergent seven-transmembrane proteins: candidate odorant receptors in DrosophilaNeuron 22: 327-338.