David M. Miller, III, Ph.D.



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Faculty Appointments
Professor of Cell and Developmental Biology
Ph.D., Biochemistry, Rice University, Houston, TexasB.S., Biology, University of Southern Mississippi, Hattiesburg, Mississippi
Office Address
Cell and Developmental Biology
465 21st Ave South
Nashville, TN 8240
Research Description
The human brain embodies the most complex and functionally remarkable tissue in biology. These attributes are defined by elaborate, highly connected networks in which myriad types of neurons are linked together in circuits with discrete physiological roles. I am interested in fundamental mechanisms that drive the creation and maintenance of this intricate structure. To obviate the need to study the brain directly, we are using the model organism, C. elegans, to reveal the underlying programs that specify neural architecture. With its simple, well-defined nervous system and facile genetics, C. elegans, is especially well-suited to this approach. For example, the phenomenon of synaptic specificity is readily evident in the wiring diagram of the C. elegans nervous system which catalogs synaptic partners for all 302 neurons in the circuit. We have exploited this resource to identify genetic mutants that alter connectivity and thus define pathways that are normally required for directing the creation of synapses between specific neurons. The execution of these developmental programs depends on expression of unique combinations of genes in different types of neurons. With the goal of identifying these genetic signatures, the Miller lab has pioneered the development of robust methods for generating neuron-specific gene expression profiles. Reverse genetic strategies (e.g., RNAi) are then employed to test candidate genes from these lists for key roles in circuit architecture. Perturbations are readily detected in this small, transparent organism with high-resolution light microscopy of neurons and synapses marked with fluorescent proteins (e.g., GFP). In addition to uncovering mechanisms that direct synaptic specificity, the Miller lab has also used these methods to address other fundamental processes in neural development including synaptic plasticity, dendrite morphogenesis and nerve regeneration. These studies are expected to reveal genetic networks with similar roles in human brain development and thus can lead to a deeper understanding of mechanisms that protect the brain from disease and repair damage arising from injury.
Research Keywords
development of the nervous system
Matthewman C, Miller-Fleming TW, Miller DM, Bianchi L. Ca2+ permeability and Na+ conductance in cellular toxicity caused by hyperactive DEG/ENaC channels. Am. J. Physiol., Cell Physiol [print-electronic]. 2016 Dec 12/1/2016; 311(6): C920-C930. PMID: 27760755, PII: ajpcell.00247.2016, DOI: 10.1152/ajpcell.00247.2016, ISSN: 1522-1563.

Lim MA, Chitturi J, Laskova V, Meng J, Findeis D, Wiekenberg A, Mulcahy B, Luo L, Li Y, Lu Y, Hung W, Qu Y, Ho CY, Holmyard D, Ji N, McWhirter R, Samuel AD, Miller DM, Schnabel R, Calarco JA, Zhen M. Neuroendocrine modulation sustains the C. elegans forward motor state. Elife. 2016 Nov 11/18/2016; 5: PMID: 27855782, PMCID: PMC5120884, DOI: 10.7554/eLife.19887, ISSN: 2050-084X.

Byrne AB, McWhirter RD, Sekine Y, Strittmatter SM, Miller DM, Hammarlund M. Inhibiting poly(ADP-ribosylation) improves axon regeneration. Elife. 2016 Oct 10/4/2016; 5: PMID: 27697151, PMCID: PMC5050021, DOI: 10.7554/eLife.12734, ISSN: 2050-084X.

O'Brien B, Miller DM. Neurodevelopment: Three's a Crowd, Four Is a Receptor Complex. Curr. Biol. 2016 Sep 9/12/2016; 26(17): R799-801. PMID: 27623260, PII: S0960-9822(16)30848-X, DOI: 10.1016/j.cub.2016.07.055, ISSN: 1879-0445.

Miller-Fleming TW, Petersen SC, Manning L, Matthewman C, Gornet M, Beers A, Hori S, Mitani S, Bianchi L, Richmond J, Miller DM. The DEG/ENaC cation channel protein UNC-8 drives activity-dependent synapse removal in remodeling GABAergic neurons. Elife. 2016 Jul 7/12/2016; 5: PMID: 27403890, PMCID: PMC4980115, DOI: 10.7554/eLife.14599, ISSN: 2050-084X.

He S, Philbrook A, McWhirter R, Gabel CV, Taub DG, Carter MH, Hanna IM, Francis MM, Miller DM. Transcriptional Control of Synaptic Remodeling through Regulated Expression of an Immunoglobulin Superfamily Protein. Curr. Biol [print-electronic]. 2015 Oct 10/5/2015; 25(19): 2541-8. PMID: 26387713, PMCID: PMC4596794, PII: S0960-9822(15)00993-8, DOI: 10.1016/j.cub.2015.08.022, ISSN: 1879-0445.

Hardaway JA, Sturgeon SM, Snarrenberg CL, Li Z, Xu XZ, Bermingham DP, Odiase P, Spencer WC, Miller DM, Carvelli L, Hardie SL, Blakely RD. Glial Expression of the Caenorhabditis elegans Gene swip-10 Supports Glutamate Dependent Control of Extrasynaptic Dopamine Signaling. J. Neurosci. 2015 Jun 6/24/2015; 35(25): 9409-23. PMID: 26109664, PMCID: PMC4478255, PII: 35/25/9409, DOI: 10.1523/JNEUROSCI.0800-15.2015, ISSN: 1529-2401.

Gerstein MB, Rozowsky J, Yan KK, Wang D, Cheng C, Brown JB, Davis CA, Hillier L, Sisu C, Li JJ, Pei B, Harmanci AO, Duff MO, Djebali S, Alexander RP, Alver BH, Auerbach R, Bell K, Bickel PJ, Boeck ME, Boley NP, Booth BW, Cherbas L, Cherbas P, Di C, Dobin A, Drenkow J, Ewing B, Fang G, Fastuca M, Feingold EA, Frankish A, Gao G, Good PJ, Guigó R, Hammonds A, Harrow J, Hoskins RA, Howald C, Hu L, Huang H, Hubbard TJ, Huynh C, Jha S, Kasper D, Kato M, Kaufman TC, Kitchen RR, Ladewig E, Lagarde J, Lai E, Leng J, Lu Z, MacCoss M, May G, McWhirter R, Merrihew G, Miller DM, Mortazavi A, Murad R, Oliver B, Olson S, Park PJ, Pazin MJ, Perrimon N, Pervouchine D, Reinke V, Reymond A, Robinson G, Samsonova A, Saunders GI, Schlesinger F, Sethi A, Slack FJ, Spencer WC, Stoiber MH, Strasbourger P, Tanzer A, Thompson OA, Wan KH, Wang G, Wang H, Watkins KL, Wen J, Wen K, Xue C, Yang L, Yip K, Zaleski C, Zhang Y, Zheng H, Brenner SE, Graveley BR, Celniker SE, Gingeras TR, Waterston R. Comparative analysis of the transcriptome across distant species. Nature. 2014 Aug 8/28/2014; 512(7515): 445-8. PMID: 25164755, PMCID: PMC4155737, PII: nature13424, DOI: 10.1038/nature13424, ISSN: 1476-4687.

Spencer WC, McWhirter R, Miller T, Strasbourger P, Thompson O, Hillier LW, Waterston RH, Miller DM. Isolation of specific neurons from C. elegans larvae for gene expression profiling. PLoS ONE. 2014; 9(11): e112102. PMID: 25372608, PMCID: PMC4221280, PII: PONE-D-14-35449, DOI: 10.1371/journal.pone.0112102, ISSN: 1932-6203.

Wang Y, Matthewman C, Han L, Miller T, Miller DM, Bianchi L. Neurotoxic unc-8 mutants encode constitutively active DEG/ENaC channels that are blocked by divalent cations. J. Gen. Physiol. 2013 Aug; 142(2): 157-69. PMID: 23898007, PMCID: PMC3727304, PII: jgp.201310974, DOI: 10.1085/jgp.201310974, ISSN: 1540-7748.

Smith CJ, O'Brien T, Chatzigeorgiou M, Spencer WC, Feingold-Link E, Husson SJ, Hori S, Mitani S, Gottschalk A, Schafer WR, Miller DM. Sensory neuron fates are distinguished by a transcriptional switch that regulates dendrite branch stabilization. Neuron. 2013 Jul 7/24/2013; 79(2): 266-80. PMID: 23889932, PMCID: PMC3795438, PII: S0896-6273(13)00402-9, DOI: 10.1016/j.neuron.2013.05.009, ISSN: 1097-4199.

Schneider J, Skelton RL, Von Stetina SE, Middelkoop TC, van Oudenaarden A, Korswagen HC, Miller DM. UNC-4 antagonizes Wnt signaling to regulate synaptic choice in the C. elegans motor circuit. Development. 2012 Jun; 139(12): 2234-45. PMID: 22619391, PMCID: PMC3357913, PII: 139/12/2234, DOI: 10.1242/dev.075184, ISSN: 1477-9129.

Smith CJ, Watson JD, VanHoven MK, Colón-Ramos DA, Miller DM. Netrin (UNC-6) mediates dendritic self-avoidance. Nat. Neurosci. 2012 May; 15(5): 731-7. PMID: 22426253, PMCID: PMC3337961, PII: nn.3065, DOI: 10.1038/nn.3065, ISSN: 1546-1726.

Petersen SC, Watson JD, Richmond JE, Sarov M, Walthall WW, Miller DM. A transcriptional program promotes remodeling of GABAergic synapses in Caenorhabditis elegans. J. Neurosci. 2011 Oct 10/26/2011; 31(43): 15362-75. PMID: 22031882, PMCID: PMC3229156, PII: 31/43/15362, DOI: 10.1523/JNEUROSCI.3181-11.2011, ISSN: 1529-2401.

Spencer WC, Zeller G, Watson JD, Henz SR, Watkins KL, McWhirter RD, Petersen S, Sreedharan VT, Widmer C, Jo J, Reinke V, Petrella L, Strome S, Von Stetina SE, Katz M, Shaham S, Rätsch G, Miller DM. A spatial and temporal map of C. elegans gene expression. Genome Res [print-electronic]. 2011 Feb; 21(2): 325-41. PMID: 21177967, PMCID: PMC3032935, PII: gr.114595.110, DOI: 10.1101/gr.114595.110, ISSN: 1549-5469.

Lu ZJ, Yip KY, Wang G, Shou C, Hillier LW, Khurana E, Agarwal A, Auerbach R, Rozowsky J, Cheng C, Kato M, Miller DM, Slack F, Snyder M, Waterston RH, Reinke V, Gerstein MB. Prediction and characterization of noncoding RNAs in C. elegans by integrating conservation, secondary structure, and high-throughput sequencing and array data. Genome Res [print-electronic]. 2011 Feb; 21(2): 276-85. PMID: 21177971, PMCID: PMC3032931, PII: gr.110189.110, DOI: 10.1101/gr.110189.110, ISSN: 1549-5469.

Hallem EA, Spencer WC, McWhirter RD, Zeller G, Henz SR, Rätsch G, Miller DM, Horvitz HR, Sternberg PW, Ringstad N. Receptor-type guanylate cyclase is required for carbon dioxide sensation by Caenorhabditis elegans. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2011 Jan 1/4/2011; 108(1): 254-9. PMID: 21173231, PMCID: PMC3017194, PII: 1017354108, DOI: 10.1073/pnas.1017354108, ISSN: 1091-6490.

Albeg A, Smith CJ, Chatzigeorgiou M, Feitelson DG, Hall DH, Schafer WR, Miller DM, Treinin M. C. elegans multi-dendritic sensory neurons: morphology and function. Mol. Cell. Neurosci [print-electronic]. 2011 Jan; 46(1): 308-17. PMID: 20971193, PMCID: PMC3018541, PII: S1044-7431(10)00246-0, DOI: 10.1016/j.mcn.2010.10.001, ISSN: 1095-9327.

Thorne CA, Hanson AJ, Schneider J, Tahinci E, Orton D, Cselenyi CS, Jernigan KK, Meyers KC, Hang BI, Waterson AG, Kim K, Melancon B, Ghidu VP, Sulikowski GA, LaFleur B, Salic A, Lee LA, Miller DM, Lee E. Small-molecule inhibition of Wnt signaling through activation of casein kinase 1a. Nat. Chem. Biol [print-electronic]. 2010 Nov; 6(11): 829-36. PMID: 20890287, PMCID: PMC3681608, PII: nchembio.453, DOI: 10.1038/nchembio.453, ISSN: 1552-4469.

Smith CJ, Watson JD, Spencer WC, O'Brien T, Cha B, Albeg A, Treinin M, Miller DM. Time-lapse imaging and cell-specific expression profiling reveal dynamic branching and molecular determinants of a multi-dendritic nociceptor in C. elegans. Dev. Biol [print-electronic]. 2010 Sep 9/1/2010; 345(1): 18-33. PMID: 20537990, PMCID: PMC2919608, PII: S0012-1606(10)00805-5, DOI: 10.1016/j.ydbio.2010.05.502, ISSN: 1095-564X.

Earls LR, Hacker ML, Watson JD, Miller DM. Coenzyme Q protects Caenorhabditis elegans GABA neurons from calcium-dependent degeneration. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2010 Aug 8/10/2010; 107(32): 14460-5. PMID: 20663955, PMCID: PMC2922603, PII: 0910630107, DOI: 10.1073/pnas.0910630107, ISSN: 1091-6490.

Chatzigeorgiou M, Yoo S, Watson JD, Lee WH, Spencer WC, Kindt KS, Hwang SW, Miller DM, Treinin M, Driscoll M, Schafer WR. Specific roles for DEG/ENaC and TRP channels in touch and thermosensation in C. elegans nociceptors. Nat. Neurosci [print-electronic]. 2010 Jul; 13(7): 861-8. PMID: 20512132, PMCID: PMC2975101, PII: nn.2581, DOI: 10.1038/nn.2581, ISSN: 1546-1726.

Hammock EA, Eagleson KL, Barlow S, Earls LR, Miller DM, Levitt P. Homologs of genes expressed in Caenorhabditis elegans GABAergic neurons are also found in the developing mouse forebrain. Neural Dev. 2010; 5: 32. PMID: 21122108, PMCID: PMC3006369, PII: 1749-8104-5-32, DOI: 10.1186/1749-8104-5-32, ISSN: 1749-8104.

Helmcke KJ, Syversen T, Miller DM, Aschner M. Characterization of the effects of methylmercury on Caenorhabditis elegans. Toxicol. Appl. Pharmacol [print-electronic]. 2009 Oct 10/15/2009; 240(2): 265-72. PMID: 19341752, PMCID: PMC2753706, PII: S0041-008X(09)00123-9, DOI: 10.1016/j.taap.2009.03.013, ISSN: 1096-0333.

Meissner B, Warner A, Wong K, Dube N, Lorch A, McKay SJ, Khattra J, Rogalski T, Somasiri A, Chaudhry I, Fox RM, Miller DM, Baillie DL, Holt RA, Jones SJ, Marra MA, Moerman DG. An integrated strategy to study muscle development and myofilament structure in Caenorhabditis elegans. PLoS Genet [print-electronic]. 2009 Jun; 5(6): e1000537. PMID: 19557190, PMCID: PMC2694363, DOI: 10.1371/journal.pgen.1000537, ISSN: 1553-7404.

Watson JD, Wang S, Von Stetina SE, Spencer WC, Levy S, Dexheimer PJ, Kurn N, Heath JD, Miller DM. Complementary RNA amplification methods enhance microarray identification of transcripts expressed in the C. elegans nervous system. BMC Genomics. 2008; 9: 84. PMID: 18284693, PMCID: PMC2263045, PII: 1471-2164-9-84, DOI: 10.1186/1471-2164-9-84, ISSN: 1471-2164.

Miller DM. Neuroscience. Synapses here and not everywhere. Science. 2007 Aug 8/17/2007; 317(5840): 907-8. PMID: 17702933, PII: 317/5840/907, DOI: 10.1126/science.1147570, ISSN: 1095-9203.

Von Stetina SE, Fox RM, Watkins KL, Starich TA, Shaw JE, Miller DM. UNC-4 represses CEH-12/HB9 to specify synaptic inputs to VA motor neurons in C. elegans. Genes Dev. 2007 Feb 2/1/2007; 21(3): 332-46. PMID: 17289921, PMCID: PMC1785118, PII: 21/3/332, DOI: 10.1101/gad.1502107, ISSN: 0890-9369.

Fox RM, Watson JD, Von Stetina SE, McDermott J, Brodigan TM, Fukushige T, Krause M, Miller DM. The embryonic muscle transcriptome of Caenorhabditis elegans. Genome Biol. 2007; 8(9): R188. PMID: 17848203, PMCID: PMC2375026, PII: gb-2007-8-9-r188, DOI: 10.1186/gb-2007-8-9-r188, ISSN: 1474-760X.

Von Stetina SE, Watson JD, Fox RM, Olszewski KL, Spencer WC, Roy PJ, Miller DM. Cell-specific microarray profiling experiments reveal a comprehensive picture of gene expression in the C. elegans nervous system. Genome Biol. 2007; 8(7): R135. PMID: 17612406, PMCID: PMC2323220, PII: gb-2007-8-7-r135, DOI: 10.1186/gb-2007-8-7-r135, ISSN: 1474-760X.

Varadan V, Miller DM, Anastassiou D. Computational inference of the molecular logic for synaptic connectivity in C. elegans. Bioinformatics. 2006 Jul 7/15/2006; 22(14): e497-506. PMID: 16873513, PII: 22/14/e497, DOI: 10.1093/bioinformatics/btl224, ISSN: 1367-4811.

Von Stetina SE, Treinin M, Miller DM. The motor circuit. Int. Rev. Neurobiol. 2006; 69: 125-67. PMID: 16492464, PII: S0074-7742(05)69005-8, DOI: 10.1016/S0074-7742(05)69005-8, ISSN: 0074-7742.

Touroutine D, Fox RM, Von Stetina SE, Burdina A, Miller DM, Richmond JE. Acr-16 encodes an essential subunit of the levamisole-resistant nicotinic receptor at the Caenorhabditis elegans neuromuscular junction. J. Biol. Chem [print-electronic]. 2005 Jul 7/22/2005; 280(29): 27013-21. PMID: 15917232, PII: M502818200, DOI: 10.1074/jbc.M502818200, ISSN: 0021-9258.

Fox RM, Von Stetina SE, Barlow SJ, Shaffer C, Olszewski KL, Moore JH, Dupuy D, Vidal M, Miller DM. A gene expression fingerprint of C. elegans embryonic motor neurons. BMC Genomics. 2005; 6: 42. PMID: 15780142, PMCID: PMC1079822, PII: 1471-2164-6-42, DOI: 10.1186/1471-2164-6-42, ISSN: 1471-2164.

Sedensky MM, Siefker JM, Koh JY, Miller DM, Morgan PG. A stomatin and a degenerin interact in lipid rafts of the nervous system of Caenorhabditis elegans. Am. J. Physiol., Cell Physiol [print-electronic]. 2004 Aug; 287(2): C468-74. PMID: 15102610, PII: 00182.2003, DOI: 10.1152/ajpcell.00182.2003, ISSN: 0363-6143.

Nass R, Hall DH, Miller DM, Blakely RD. Neurotoxin-induced degeneration of dopamine neurons in Caenorhabditis elegans. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2002 Mar 3/5/2002; 99(5): 3264-9. PMID: 11867711, PMCID: PMC122507, PII: 042497999, DOI: 10.1073/pnas.042497999, ISSN: 0027-8424.

Christensen M, Estevez A, Yin X, Fox R, Morrison R, McDonnell M, Gleason C, Miller DM, Strange K. A primary culture system for functional analysis of C. elegans neurons and muscle cells. Neuron. 2002 Feb 2/14/2002; 33(4): 503-14. PMID: 11856526, PII: S0896627302005913, ISSN: 0896-6273.

Esmaeili B, Ross JM, Neades C, Miller DM, Ahringer J. The C. elegans even-skipped homologue, vab-7, specifies DB motoneurone identity and axon trajectory. Development. 2002 Feb; 129(4): 853-62. PMID: 11861469, ISSN: 0950-1991.

Nass R, Miller DM, Blakely RD. C. elegans: a novel pharmacogenetic model to study Parkinson's disease. Parkinsonism Relat. Disord. 2001 Jul; 7(3): 185-91. PMID: 11331185, PII: S1353-8020(00)00056-0, ISSN: 1873-5126.

Lickteig KM, Duerr JS, Frisby DL, Hall DH, Rand JB, Miller DM. Regulation of neurotransmitter vesicles by the homeodomain protein UNC-4 and its transcriptional corepressor UNC-37/groucho in Caenorhabditis elegans cholinergic motor neurons. J. Neurosci. 2001 Mar 3/15/2001; 21(6): 2001-14. PMID: 11245684, PII: 21/6/2001, ISSN: 1529-2401.

Bianchi L, Miller DM, George AL. Expression of a CIC chloride channel in Caenorhabditis elegans gamma-aminobutyric acid-ergic neurons. Neurosci. Lett. 2001 Feb 2/23/2001; 299(3): 177-80. PMID: 11165764, PII: S0304394001015257, ISSN: 0304-3940.

Winnier AR, Meir JY, Ross JM, Tavernarakis N, Driscoll M, Ishihara T, Katsura I, Miller DM. UNC-4/UNC-37-dependent repression of motor neuron-specific genes controls synaptic choice in Caenorhabditis elegans. Genes Dev. 1999 Nov 11/1/1999; 13(21): 2774-86. PMID: 10557206, PMCID: PMC317130, ISSN: 0890-9369.

Miller DM, Desai NS, Hardin DC, Piston DW, Patterson GH, Fleenor J, Xu S, Fire A. Two-color GFP expression system for C. elegans. BioTechniques. 1999 May; 26(5): 914-8, 920. PMID: 10337485, ISSN: 0736-6205.

Jayanthi LD, Apparsundaram S, Malone MD, Ward E, Miller DM, Eppler M, Blakely RD. The Caenorhabditis elegans gene T23G5.5 encodes an antidepressant- and cocaine-sensitive dopamine transporter. Mol. Pharmacol. 1998 Oct; 54(4): 601-9. PMID: 9765501, ISSN: 0026-895X.

Pflugrad A, Meir JY, Barnes TM, Miller DM. The Groucho-like transcription factor UNC-37 functions with the neural specificity gene unc-4 to govern motor neuron identity in C. elegans. Development. 1997 May; 124(9): 1699-709. PMID: 9165118, ISSN: 0950-1991.

Miller DM, Niemeyer CJ. Expression of the unc-4 homeoprotein in Caenorhabditis elegans motor neurons specifies presynaptic input. Development. 1995 Sep; 121(9): 2877-86. PMID: 7555714, ISSN: 0950-1991.

Miller DM, Shakes DC. Immunofluorescence microscopy. Methods Cell Biol. 1995; 48: 365-94. PMID: 8531735, ISSN: 0091-679X.

Miller DM, Niemeyer CJ, Chitkara P. Dominant unc-37 mutations suppress the movement defect of a homeodomain mutation in unc-4, a neural specificity gene in Caenorhabditis elegans. Genetics. 1993 Nov; 135(3): 741-53. PMID: 7904971, PMCID: PMC1205717, ISSN: 0016-6731.

Miller DM, Shen MM, Shamu CE, Bürglin TR, Ruvkun G, Dubois ML, Ghee M, Wilson L. C. elegans unc-4 gene encodes a homeodomain protein that determines the pattern of synaptic input to specific motor neurons. Nature. 1992 Feb 2/27/1992; 355(6363): 841-5. PMID: 1347150, DOI: 10.1038/355841a0, ISSN: 0028-0836.

Maruyama IN, Miller DM, Brenner S. Myosin heavy chain gene amplification as a suppressor mutation in Caenorhabditis elegans. Mol. Gen. Genet. 1989 Oct; 219(1-2): 113-8. PMID: 2575705, ISSN: 0026-8925.

Miller DM, Stockdale FE, Karn J. Immunological identification of the genes encoding the four myosin heavy chain isoforms of Caenorhabditis elegans. Proc. Natl. Acad. Sci. U.S.A. 1986 Apr; 83(8): 2305-9. PMID: 2422655, PMCID: PMC323285, ISSN: 0027-8424.

Watts FZ, Miller DM, Orr E. Identification of myosin heavy chain in Saccharomyces cerevisiae. Nature. 1985 Jul 7/4/1985; 316(6023): 83-5. PMID: 3892306, ISSN: 0028-0836.

Epstein HF, Miller DM, Ortiz I, Berliner GC. Myosin and paramyosin are organized about a newly identified core structure. J. Cell Biol. 1985 Mar; 100(3): 904-15. PMID: 3972901, PMCID: PMC2113503, ISSN: 0021-9525.

Miller DM, Ortiz I, Berliner GC, Epstein HF. Differential localization of two myosins within nematode thick filaments. Cell. 1983 Sep; 34(2): 477-90. PMID: 6352051, PII: 0092-8674(83)90381-1, ISSN: 0092-8674.