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Gary C. Schoenwolf

Gary C. Schoenwolf, PhD

Languages spoken: English

Academic Information

Departments Emeritus - Neurobiology

Academic Office Information

Research Interests

  • Developmental Biology
  • Morphogenesis
  • Cell Communication
  • Cell Signaling
  • Inner Ear
  • Growth Factors
  • Birth Defects
  • Congenital Anomalies

The Schoenwolf laboratory addresses the general question of how intercellular and intracellular signaling results in the generation of pattern during vertebrate embryogenesis. Although we work on a number of diverse organ rudiments and developmental events using multiple experimental organisms, our current work is focused on normal and abnormal development of the inner ear. Our research combines molecular, cellular, embryological, transgenic, imaging, and in vitro and in vivo approaches. This work is relevant for understanding birth defects, cancer, and aging.

Dr. Schoenwolf is also actively involved in mentoring and teaching grant writing.

Education History

Other Training University of Utah
University of New Mexico
Postdoctoral Fellow
Doctoral Training University of Illinois
University of Washington, Friday Harbor Laboratories
Graduate Training University of Illinois
Elmhurst College

Selected Publications

Journal Article

  1. Ohta S, Wang B, Mansour SL, Schoenwolf GC (2016). SHH ventralizes the otocyst by maintaining basal PKA activity and regulating GLI3 signaling. Dev Biol, 420(1), 100-109.
  2. Ohta S, Wang B, Mansour SL, Schoenwolf GC (2016). BMP regulates regional gene expression in the dorsal otocyst through canonical and non-canonical intracellular pathways. Development, 143(12), 2228-37.
  3. Olaya-Sanchez D, Sanchez-Guardado LO, Ohta S, Chapman SC, Schoenwolf GC, Puelles L, Hidalgo-Sanchez M (2016). Fgf3 and Fgf16 expression patterns define spatial and temporal domains in the developing chick inner ear. Brain Struct Funct.
  4. Hung IH, Schoenwolf GC, Lewandoski M, Ornitz DM (2016). A combined series of Fgf9 and Fgf18 mutant alleles identifies unique and redundant roles in skeletal development. Dev Biol, 411(1), 72-84.
  5. Schoenwolf GC (2013). Getting published well requires fulfilling editors' and reviewers' needs and desires. Dev Growth Differ, 55(9), 735-43.
  6. Nishita J, Ohta S, Bleyl SB, Schoenwolf GC (2011). Detection of isoform-specific fibroblast growth factor receptors by whole-mount in situ hybridization in early chick embryos. Dev Dyn, 240(6), 1537-47.
  7. Ohta S, Mansour SL, Schoenwolf GC (2010). BMP/SMAD signaling regulates the cell behaviors that drive the initial dorsal-specific regional morphogenesis of the otocyst. Dev Biol, 347(2), 369-81.
  8. Ohta S, Schoenwolf GC, Yamada G (2010). The cessation of gastrulation: BMP signaling and EMT during and at the end of gastrulation. Cell Adh Migr, 4(3), 440-6.
  9. Urness LD, Paxton CN, Wang X, Schoenwolf GC, Mansour SL (2010). FGF signaling regulates otic placode induction and refinement by controlling both ectodermal target genes and hindbrain Wnt8a. Dev Biol, 340(2), 595-604.
  10. Bleyl SB, Saijoh Y, Bax NA, Gittenberger-de Groot AC, Wisse LJ, Chapman SC, Hunter J, Shiratori H, Hamada H, Yamada S, Shiota K, Klewer SE, Leppert MF, Schoenwolf GC (2010). Dysregulation of the PDGFRA gene causes inflow tract anomalies including TAPVR: integrating evidence from human genetics and model organisms. Hum Mol Genet, 19(7), 1286-301.
  11. Paxton CN, Bleyl SB, Chapman SC, Schoenwolf GC (2010). Identification of differentially expressed genes in early inner ear development. Gene Expr Patterns, 10(1), 31-43.
  12. Bleyl SB, Moshrefi A, Shaw GM, Saijoh Y, Schoenwolf GC, Pennacchio LA, Slavotinek AM (2007). Candidate genes for congenital diaphragmatic hernia from animal models: sequencing of FOG2 and PDGFRalpha reveals rare variants in diaphragmatic hernia patients. Eur J Hum Genet, 15(9), 950-8.
  13. Chapman SC, Matsumoto K, Cai Q, Schoenwolf GC (2007). Specification of germ layer identity in the chick gastrula. BMC Dev Biol, 7, 91.
  14. Chapman SC, Cai Q, Bleyl SB, Schoenwolf GC (2006). Restricted expression of Fgf16 within the developing chick inner ear. Dev Dyn, 235(8), 2276-81.
  15. Decastro M, Saijoh Y, Schoenwolf GC (2006). Optimized cationic lipid-based gene delivery reagents for use in developing vertebrate embryos. Dev Dyn, 235(8), 2210-9.
  16. Chapman SC, Sawitzke AL, Campbell DS, Schoenwolf GC (2005). A three-dimensional atlas of pituitary gland development in the zebrafish. J Comp Neurol, 487(4), 428-40.
  17. Anderson-Berry A, OBrien EA, Bleyl SB, Lawson A, Gundersen N, Ryssman D, Sweeley J, Dahl MJ, Drake CJ, Schoenwolf GC, Albertine KH (2005). Vasculogenesis drives pulmonary vascular growth in the developing chick embryo. Dev Dyn, 233(1), 145-53.
  18. Ladher RK, Wright TJ, Moon AM, Mansour SL, Schoenwolf GC (2005). FGF8 initiates inner ear induction in chick and mouse. Genes Dev, 19(5), 603-13.
  19. Chapman SC, Lawson A, Macarthur WC, Wiese RJ, Loechel RH, Burgos-Trinidad M, Wakefield JK, Ramabhadran R, Mauch TJ, Schoenwolf GC (2005). Ubiquitous GFP expression in transgenic chickens using a lentiviral vector. Development, 132(5), 935-40.
  20. Sawitzke AL, Chapman SC, Bleyl SB, Schoenwolf GC (2005). Improvements in histological quality and signal retention following in situ hybridization in early chick embryos using plastic resin and recolorization. Biotech Histochem, 80(1), 35-41.
  21. Lopez-Sanchez C, Garcia-Martinez V, Lawson A, Chapman SC, Schoenwolf GC (2004). Rapid triple-labeling method combining in situ hybridization and double immunocytochemistry. Dev Dyn, 230(2), 309-15.
  22. Wright TJ, Ladher R, McWhirter J, Murre C, Schoenwolf GC, Mansour SL (2004). Mouse FGF15 is the ortholog of human and chick FGF19, but is not uniquely required for otic induction. Dev Biol, 269(1), 264-75.
  23. Chapman SC, Brown R, Lees L, Schoenwolf GC, Lumsden A (2004). Expression analysis of chick Wnt and frizzled genes and selected inhibitors in early chick patterning. Dev Dyn, 229(3), 668-76.
  24. Antin PB, Fallon JF, Schoenwolf GC (2004). The chick embryo rules (still)! Dev Dyn, 229(3), 413.
  25. Schoenwolf GC (2004). In honor of James A. Weston and the neural crest. Dev Dyn, 229(1), 1.
  26. Schoenwolf GC (2003). Trink: His life, his philosophy. Dev Dyn, 228(4), 591-593.
  27. Schoenwolf GC (2003). In memory of John Philip Trinkaus. Dev Dyn, 228(4), 585.
  28. Chapman SC, Schubert FR, Schoenwolf GC, Lumsden A (2003). Anterior identity is established in chick epiblast by hypoblast and anterior definitive endoderm. Development, 130(21), 5091-101.
  29. Colas JF, Schoenwolf GC (2003). Assessing the contributions of gene products to the form-shaping events of neurulation: a transgenic approach in chick. Genesis, 37(2), 64-75.
  30. Wright TJ, Hatch EP, Karabagli H, Karabagli P, Schoenwolf GC, Mansour SL (2003). Expression of mouse fibroblast growth factor and fibroblast growth factor receptor genes during early inner ear development. Dev Dyn, 228(2), 267-72.
  31. Schoenwolf GC (2003). What's cool in Dev Bio. Dev Dyn, 228(2), 151.
  32. Colas JF, Schoenwolf GC (2003). Differential expression of two cell adhesion molecules, Ephrin-A5 and Integrin alpha6, during cranial neurulation in the chick embryo. Dev Neurosci, 25(5), 357-65.
  33. Schoenwolf GC (2003). Book look. Dev Dyn, 227(4), 469.
  34. Lawson A, Schoenwolf GC (2003). Epiblast and primitive-streak origins of the endoderm in the gastrulating chick embryo. Development, 130(15), 3491-501.
  35. Colas JF, Schoenwolf GC (2003). Localization of cartilage linking protein 1 during primary neurulation in the chick embryo. Brain Res Dev Brain Res, 141(1-2), 141-8.
  36. Kirby ML, Lawson A, Stadt HA, Kumiski DH, Wallis KT, McCraney E, Waldo KL, Li YX, Schoenwolf GC (2003). Hensen's node gives rise to the ventral midline of the foregut: implications for organizing head and heart development. Dev Biol, 253(2), 175-88.
  37. Wright, TJ, E Hatch, H Karbagli, GC Schoenwolf, SL Mansour (2003). Expression of mouse fibroglast growth factor and fibroblast growth factor receptor genes during early inner ear development. Dev Dyn, 228, 267-272.
  38. Karabagli H, Karabagli P, Ladher RK, Schoenwolf GC (2002). Comparison of the expression patterns of several fibroblast growth factors during chick gastrulation and neurulation. Anat Embryol (Berl), 205(5-6), 365-70.
  39. Karabagli H, Karabagli P, Ladher RK, Schoenwolf GC (2002). Survey of fibroblast growth factor expression during chick organogenesis. Anat Rec, 268(1), 1-6.
  40. Lopez-Sanchez C, Climent V, Schoenwolf GC, Alvarez IS, Garcia-Martinez V (2002). Induction of cardiogenesis by Hensen's node and fibroblast growth factors. Cell Tissue Res, 309(2), 237-49.
  41. Chapman SC, Schubert FR, Schoenwolf GC, Lumsden A (2002). Analysis of spatial and temporal gene expression patterns in blastula and gastrula stage chick embryos. Dev Biol, 245(1), 187-99.
  42. Schoenwolf GC (2002). Principles of developmental biology. J Musculoskelet Neuronal Interact, 2(3), 268-9.
  43. Gallego-Diaz V, Schoenwolf GC, Alvarez IS (2002). The effects of BMPs on early chick embryos suggest a conserved signaling mechanism for epithelial and neural induction among vertebrates. Brain Res Bull, 57(3-4), 289-91.
  44. Healy KH, Schoenwolf GC, Darnell DK (2001). Cell interactions underlying notochord induction and formation in the chick embryo. Dev Dyn, 222(2), 165-77.
  45. Lawson A, Schoenwolf GC (2001). Cell populations and morphogenetic movements underlying formation of the avian primitive streak and organizer. Genesis, 29(4), 188-95.
  46. Chapman SC, Collignon J, Schoenwolf GC, Lumsden A (2001). Improved method for chick whole-embryo culture using a filter paper carrier. Dev Dyn, 220(3), 284-9.
  47. Lawson A, Schoenwolf GC (2001). New insights into critical events of avian gastrulation. Anat Rec, 262(3), 238-52.
  48. Lawson A, Colas JF, Schoenwolf GC (2001). Classification scheme for genes expressed during formation and progression of the avian primitive streak. Anat Rec, 262(2), 221-6.
  49. Lawson A, Anderson H, Schoenwolf GC (2001). Cellular mechanisms of neural fold formation and morphogenesis in the chick embryo. Anat Rec, 262(2), 153-68.
  50. Lopez-Sanchez C, Garcia-Martinez V, Schoenwolf GC (2001). Localization of cells of the prospective neural plate, heart and somites within the primitive streak and epiblast of avian embryos at intermediate primitive-streak stages. Cells Tissues Organs, 169(4), 334-46.
  51. Ladher RK, Anakwe KU, Gurney AL, Schoenwolf GC, Francis-West PH (2000). Identification of synergistic signals initiating inner ear development. Science, 290(5498), 1965-7.
  52. Stark MR, Rao MS, Schoenwolf GC, Yang G, Smith D, Mauch TJ (2000). Frizzled-4 expression during chick kidney development. Mech Dev, 98(1-2), 121-5.
  53. Yuan S, Schoenwolf GC (2000). Islet-1 marks the early heart rudiments and is asymmetrically expressed during early rotation of the foregut in the chick embryo. Anat Rec, 260(2), 204-7.
  54. Colas JF, Schoenwolf GC (2000). Subtractive hybridization identifies chick-cripto, a novel EGF-CFC ortholog expressed during gastrulation, neurulation and early cardiogenesis. Gene, 255(2), 205-17.
  55. Lawson A, Colas JF, Schoenwolf GC (2000). Ectodermal markers delineate the neural fold interface during avian neurulation. Anat Rec, 260(1), 106-9.
  56. Colas JF, Lawson A, Schoenwolf GC (2000). Evidence that translation of smooth muscle alpha-actin mRNA is delayed in the chick promyocardium until fusion of the bilateral heart-forming regions. Dev Dyn, 218(2), 316-30.
  57. Stark MR, Biggs JJ, Schoenwolf GC, Rao MS (2000). Characterization of avian frizzled genes in cranial placode development. Mech Dev, 93(1-2), 195-200.
  58. Mauch TJ, Yang G, Wright M, Smith D, Schoenwolf GC (2000). Signals from trunk paraxial mesoderm induce pronephros formation in chick intermediate mesoderm. Dev Biol, 220(1), 62-75.
  59. Schoenwolf GC (2000). Molecular genetic control of axis patterning during early embryogenesis of vertebrates. Ann N Y Acad Sci, 919, 246-60.
  60. Darnell DK, Schoenwolf GC (2000). Transplantation chimeras. Use in analyzing mechanisms of avian development. Methods Mol Biol, 135, 367-71.
  61. Darnell DK, Garcia-Martinez V, Lopez-Sanchez C, Yuan S, Schoenwolf GC (2000). Dynamic labeling techniques for fate mapping, testing cell commitment, and following living cells in avian embryos. Methods Mol Biol, 135, 305-21.
  62. Darnell DK, Schoenwolf GC (2000). Culture of avian embryos. Methods Mol Biol, 135, 31-8.
  63. Darnell DK, Schoenwolf GC (2000). The chick embryo as a model system for analyzing mechanisms of development. Methods Mol Biol, 135, 25-9.
  64. Yuan S, Schoenwolf GC (1999). The spatial and temporal pattern of C-Lmx1 expression in the neuroectoderm during chick neurulation. Mech Dev, 88(2), 243-7.
  65. Lawson A, Schoenwolf GC, England MA, Addai FK, Ahima RS (1999). Programmed cell death and the morphogenesis of the hindbrain roof plate in the chick embryo. Anat Embryol (Berl), 200(5), 509-19.
  66. Davidson BP, Kinder SJ, Steiner K, Schoenwolf GC, Tam PP (1999). Impact of node ablation on the morphogenesis of the body axis and the lateral asymmetry of the mouse embryo during early organogenesis. Dev Biol, 211(1), 11-26.
  67. Yuan S, Schoenwolf GC (1999). Reconstitution of the organizer is both sufficient and required to re-establish a fully patterned body plan in avian embryos. Development, 126(11), 2461-73.
  68. Darnell DK, Stark MR, Schoenwolf GC (1999). Timing and cell interactions underlying neural induction in the chick embryo. Development, 126(11), 2505-14.
  69. Thayer JM, Schoenwolf GC (1998). Early expression of Osteopontin in the chick is restricted to rhombomeres 5 and 6 and to a subpopulation of neural crest cells that arise from these segments. Anat Rec, 250(2), 199-209.
  70. Yuan S, Schoenwolf GC (1998). De novo induction of the organizer and formation of the primitive streak in an experimental model of notochord reconstitution in avian embryos. Development, 125(2), 201-13.
  71. Hackett DA, Smith JL, Schoenwolf GC (1997). Epidermal ectoderm is required for full elevation and for convergence during bending of the avian neural plate. Dev Dyn, 210(4), 397-406.
  72. Rodriguez-Gallardo L, Climent V, Garcia-Martinez V, Schoenwolf GC, Alvarez IS (1997). Targeted over-expression of FGF in chick embryos induces formation of ectopic neural cells. Int J Dev Biol, 41(5), 715-23.
  73. Sausedo RA, Smith JL, Schoenwolf GC (1997). Role of nonrandomly oriented cell division in shaping and bending of the neural plate. J Comp Neurol, 381(4), 473-88.
  74. Darnell DK, Schoenwolf GC (1997). Vertical induction of engrailed-2 and other region-specific markers in the early chick embryo. Dev Dyn, 209(1), 45-58.
  75. Inagaki T, Schoenwolf GC, Walker ML (1997). Experimental model: change in the posterior fossa with surgically induced spina bifida aperta in mouse. Pediatr Neurosurg, 26(4), 185-9.
  76. Garcia-Martinez V, Darnell DK, Lopez-Sanchez C, Sosic D, Olson EN, Schoenwolf GC (1997). State of commitment of prospective neural plate and prospective mesoderm in late gastrula/early neurula stages of avian embryos. Dev Biol, 181(1), 102-15.
  77. Garton HJ, Schoenwolf GC (1996). Improving the efficacy of fluorescent labeling for histological tracking of cells in early mammalian and avian embryos. Anat Rec, 244(1), 112-17.
  78. Yuan S, Darnell DK, Schoenwolf GC (1995). Identification of inducing, responding, and suppressing regions in an experimental model of notochord formation in avian embryos. Dev Biol, 172(2), 567-84.
  79. Moury JD, Schoenwolf GC (1995). Cooperative model of epithelial shaping and bending during avian neurulation: autonomous movements of the neural plate, autonomous movements of the epidermis, and interactions in the neural plate/epidermis transition zone. Dev Dyn, 204(3), 323-37.
  80. Schoenwolf GC, Yuan S (1995). Experimental analyses of the rearrangement of ectodermal cells during gastrulation and neurulation in avian embryos. Cell Tissue Res, 280(2), 243-51.
  81. Darnell DK, Schoenwolf GC (1995). Dorsoventral patterning of the avian mesencephalon/metencephalon: role of the notochord and floor plate in suppressing Engrailed-2. J Neurobiol, 26(1), 62-74.
  82. Yuan S, Darnell DK, Schoenwolf GC (1995). Mesodermal patterning during avian gastrulation and neurulation: experimental induction of notochord from non-notochordal precursor cells. Dev Genet, 17(1), 38-54.
  83. Smith JL, Gesteland KM, Schoenwolf GC (1994). Prospective fate map of the mouse primitive streak at 7.5 days of gestation. Dev Dyn, 201(3), 279-89.
  84. Sulik K, Dehart DB, Iangaki T, Carson JL, Vrablic T, Gesteland K, Schoenwolf GC (1994). Morphogenesis of the murine node and notochordal plate. Dev Dyn, 201(3), 260-78.
  85. Sausedo RA, Schoenwolf GC (1994). Quantitative analyses of cell behaviors underlying notochord formation and extension in mouse embryos. Anat Rec, 239(1), 103-12.
  86. Smith JL, Schoenwolf GC, Quan J (1994). Quantitative analyses of neuroepithelial cell shapes during bending of the mouse neural plate. J Comp Neurol, 342(1), 144-51.
  87. Garcia-Martinez V, Alvarez IS, Schoenwolf GC (1993). Locations of the ectodermal and nonectodermal subdivisions of the epiblast at stages 3 and 4 of avian gastrulation and neurulation. J Exp Zool, 267(4), 431-46.
  88. Garcia-Martinez V, Schoenwolf GC (1993). Primitive-streak origin of the cardiovascular system in avian embryos. Dev Biol, 159(2), 706-19.
  89. Sausedo RA, Schoenwolf GC (1993). Cell behaviors underlying notochord formation and extension in avian embryos: quantitative and immunocytochemical studies. Anat Rec, 237(1), 58-70.
  90. Inagaki T, Schoenwolf GC (1993). Axis development in avian embryos: the ability of Hensen's node to self-differentiate, as analyzed with heterochronic grafting experiments. Anat Embryol (Berl), 188(1), 1-11.
  91. Inagaki T, Garcia-Martinez V, Schoenwolf GC (1993). Regulative ability of the prospective cardiogenic and vasculogenic areas of the primitive streak during avian gastrulation. Dev Dyn, 197(1), 57-68.
  92. Ellis LC, Smith AM, Alvarez IS, Schoenwolf GC (1993). Monoclonal antibodies identifying subsets of ectodermal, mesodermal, and endodermal cells in gastrulating and neurulating avian embryos. Anat Rec, 235(4), 591-603.
  93. Schoenwolf GC (1992). Morphological and mapping studies of the paranodal and postnodal levels of the neural plate during chick neurulation. Anat Rec, 233(2), 281-90.
  94. Darnell DK, Schoenwolf GC, Ordahl CP (1992). Changes in dorsoventral but not rostrocaudal regionalization of the chick neural tube in the absence of cranial notochord, as revealed by expression of engrailed-2. Dev Dyn, 193(4), 389-96.
  95. Alvarez IS, Schoenwolf GC (1992). Expansion of surface epithelium provides the major extrinsic force for bending of the neural plate. J Exp Zool, 261(3), 340-8.
  96. Garcia-Martinez V, Schoenwolf GC (1992). Positional control of mesoderm movement and fate during avian gastrulation and neurulation. Dev Dyn, 193(3), 249-56.
  97. Schoenwolf GC, Garcia-Martinez V, Dias MS (1992). Mesoderm movement and fate during avian gastrulation and neurulation. Dev Dyn, 193(3), 235-48.
  98. Schoenwolf GC, Alvarez IS (1991). Specification of neurepithelium and surface epithelium in avian transplantation chimeras. Development, 112(3), 713-22.
  99. Smith JL, Schoenwolf GC (1991). Further evidence of extrinsic forces in bending of the neural plate. J Comp Neurol, 307(2), 225-36.
  100. Alvarez IS, Schoenwolf GC (1991). Patterns of neurepithelial cell rearrangement during avian neurulation are determined prior to notochordal inductive interactions. Dev Biol, 143(1), 78-92.
  101. Dias MS, Schoenwolf GC (1990). Formation of ectopic neurepithelium in chick blastoderms: age-related capacities for induction and self-differentiation following transplantation of quail Hensen's nodes. Anat Rec, 228(4), 437-48.
  102. Schoenwolf GC, Sheard P (1990). Fate mapping the avian epiblast with focal injections of a fluorescent-histochemical marker: ectodermal derivatives. J Exp Zool, 255(3), 323-39.
  103. Schoenwolf GC, Alvarez IS (1989). Roles of neuroepithelial cell rearrangement and division in shaping of the avian neural plate. Development, 106(3), 427-39.
  104. Smith JL, Schoenwolf GC (1989). Notochordal induction of cell wedging in the chick neural plate and its role in neural tube formation. J Exp Zool, 250(1), 49-62.
  105. Schoenwolf GC, Bortier H, Vakaet L (1989). Fate mapping the avian neural plate with quail/chick chimeras: origin of prospective median wedge cells. J Exp Zool, 249(3), 271-8.
  106. Schoenwolf GC, Sheard P (1989). Shaping and bending of the avian neural plate as analysed with a fluorescent-histochemical marker. Development, 105(1), 17-25.
  107. Schoenwolf GC, Everaert S, Bortier H, Vakaet L (1989). Neural plate- and neural tube-forming potential of isolated epiblast areas in avian embryos. Anat Embryol (Berl), 179(6), 541-9.
  108. Schoenwolf GC (1988). Microsurgical analyses of avian neurulation: separation of medial and lateral tissues. J Comp Neurol, 276(4), 498-507.
  109. Smith JL, Schoenwolf GC (1988). Role of cell-cycle in regulating neuroepithelial cell shape during bending of the chick neural plate. Cell Tissue Res, 252(3), 491-500.
  110. Schoenwolf GC, Folsom D, Moe A (1988). A reexamination of the role of microfilaments in neurulation in the chick embryo. Anat Rec, 220(1), 87-102.
  111. Fineman RM, Schoenwolf GC (1987). Animal model: dysmorphogenesis and death in a chicken embryo model. Am J Med Genet, 27(3), 543-52.
  112. Schoenwolf GC, Powers ML (1987). Shaping of the chick neuroepithelium during primary and secondary neurulation: role of cell elongation. Anat Rec, 218(2), 182-95.
  113. Smith JL, Schoenwolf GC (1987). Cell cycle and neuroepithelial cell shape during bending of the chick neural plate. Anat Rec, 218(2), 196-206.
  114. Fineman RM, Schoenwolf GC, Huff M, Davis PL (1986). Causes of windowing-induced dysmorphogenesis (neural tube defects and early amnion deficit spectrum) in chicken embryos. Am J Med Genet, 25(3), 489-505.
  115. Desmond ME, Schoenwolf GC (1986). Evaluation of the roles of intrinsic and extrinsic factors in occlusion of the spinal neurocoel during rapid brain enlargement in the chick embryo. J Embryol Exp Morphol, 97, 25-46.
  116. Schoenwolf GC, Desmond ME (1986). Timing and positioning of reopening of the occluded spinal neurocele in the chick embryo. J Comp Neurol, 246(4), 459-66.
  117. Pacheco MA, Marks RW, Schoenwolf GC, Desmond ME (1986). Quantification of the initial phases of rapid brain enlargement in the chick embryo. Am J Anat, 175(4), 403-11.
  118. Schoenwolf GC, Chandler NB, Smith JL (1985). Analysis of the origins and early fates of neural crest cells in caudal regions of avian embryos. Dev Biol, 110(2), 467-79.
  119. Desmond ME, Schoenwolf GC (1985). Timing and positioning of occlusion of the spinal neurocele in the chick embryo. J Comp Neurol, 235(4), 479-87.
  120. Schoenwolf GC (1985). Shaping and bending of the avian neuroepithelium: morphometric analyses. Dev Biol, 109(1), 127-39.
  121. Schoenwolf GC, Franks MV (1984). Quantitative analyses of changes in cell shapes during bending of the avian neural plate. Dev Biol, 105(2), 257-72.
  122. Schoenwolf GC, Desmond ME (1984). Neural tube occlusion precedes rapid brain enlargement. J Exp Zool, 230(3), 405-7.
  123. Schoenwolf GC, Desmond ME (1984). Descriptive studies of occlusion and reopening of the spinal canal of the early chick embryo. Anat Rec, 209(2), 251-63.
  124. Schoenwolf GC (1984). Histological and ultrastructural studies of secondary neurulation in mouse embryos. Am J Anat, 169(4), 361-76.
  125. Schoenwolf GC, Nichols DH (1984). Histological and ultrastructural studies on the origin of caudal neural crest cells in mouse embryos. J Comp Neurol, 222(4), 496-505.
  126. Chandler NB, Schoenwolf GC (1983). Wrinkle-free plastic sections for light microscopy. Stain Technol, 58(4), 238-40.
  127. Fisher M, Schoenwolf GC (1983). The use of early chick embryos in experimental embryology and teratology: improvements in standard procedures. Teratology, 27(1), 65-72.
  128. Schoenwolf GC, Fisher M (1983). Analysis of the effects of Streptomyces hyaluronidase on formation of the neural tube. J Embryol Exp Morphol, 73, 1-15.
  129. Schoenwolf, GC, NB Chandler (1983). A novel sectioning technique for use in descriptive embryology: Bridging the gap between paraffin and plastic serial sections. Anat Rec, 206, 221-225.
  130. Schoenwolf GC (1982). A simple method for fabricating inexpensive storage boxes for 6.35 mm-thick glass knives. Stain Technol, 57(3), 185-90.
  131. Costanzo R, Watterson RL, Schoenwolf GC (1982). Evidence that secondary neurulation occurs autonomously in the chick embryo. J Exp Zool, 219(2), 233-40.
  132. Schoenwolf GC, Singh U (1981). Changes in the surface morphologies of the cells in the bursa cloacalis (bursa of Fabricius) and thymus during ontogeny of the chick embryo. Anat Rec, 201(2), 303-16.
  133. Schoenwolf GC (1981). Morphogenetic processes involved in the remodeling of the tail region of the chick embryo. Anat Embryol (Berl), 162(2), 183-97.
  134. Schoenwolf GC, Bell LA, Watterson RL (1981). Vasculogenesis of the bursa cloacalis (bursa of Fabricius) of the chick embryo. J Morphol, 167(1), 35-42.
  135. Waterman RE, Schoenwolf GC (1980). The ultrastructure of oral (buccopharyngeal) membrane formation and rupture in the chick embryo. Anat Rec, 197(4), 441-70.
  136. Schoenwolf GC, Delongo J (1980). Ultrastructure of secondary neurulation in the chick embryo. Am J Anat, 158(1), 43-63.
  137. Schoenwolf GC, Kelley RO (1980). Characterization of intercellular junctions in the caudal portion of the developing neural tube of the chick embryo. Am J Anat, 158(1), 29-41.
  138. Schoenwolf GC (1979). Observations on closure of the neuropores in the chick embryo. Am J Anat, 155(4), 445-65.
  139. Schoenwolf GC (1979). Histological and ultrastructural observations of tail bud formation in the chick embryo. Anat Rec, 193(1), 131-47.
  140. Schoenwolf GC (1978). Effects of complete tail bud extirpation on early development of the posterior region of the chick embryo. Anat Rec, 192(2), 289-95.
  141. Schoenwolf, GC (1978). An SEM study of posterior spinal cord development region of the chick embryo. Scan Electron Microsc, II, 739-746.
  142. Keller, RE, GC Schoenwolf (1977). An SEM study of cellular morphology, contact, and arrangement, as related to gastrulation in Xenopus laevis. Wilehm Roux Arch Dev Biol, 182, 165-186.
  143. Schoenwolf, GC (1977). Tail (end) bud contributions to the posterior region of the chick embryo. J Exp Zool, 201, 227-246.


  1. Francis-West, P, RK Ladher, GC Schoewolf (2002). Development of Sensory Organs. [Review]. Sci Prog, 84, 151-173.
  2. Schoenwolf GC (2001). Cutting, pasting and painting: experimental embryology and neural development. [Review]. Nat Rev Neurosci, 2(11), 763-71.
  3. Colas JF, Schoenwolf GC (2001). Towards a cellular and molecular understanding of neurulation. [Review]. Dev Dyn, 221(2), 117-45.
  4. Schoenwolf, GC (2001). Principles fo development biology. [Review]. J Musculoskelet Neuronal Interact, 2, 268-269.
  5. Schoenwolf, GC (2000). Molecular genetic control of axis patterning during early embryogenesis of vertebrates. [Review]. Ann N Y Acad Sci, 919, 246-260.
  6. Schoenwolf GC, Smith JL (2000). Gastrulation and early mesodermal patterning in vertebrates. [Review]. Methods Mol Biol, 135, 113-25.
  7. Schoenwolf GC, Smith JL (2000). Mechanisms of neurulation. [Review]. Methods Mol Biol, 136, 125-34.
  8. Inagaki T, Smith JL, Walker ML, Schoenwolf GC (2000). Neural tube defects. [Review]. Methods Mol Biol, 136, 161-6.
  9. Smith JL, Schoenwolf GC (1998). Getting organized: new insights into the organizer of higher vertebrates. [Review]. Curr Top Dev Biol, 40, 79-110.
  10. Smith JL, Schoenwolf GC (1997). Neurulation: coming to closure. [Review]. Trends Neurosci, 20(11), 510-7.
  11. Schoenwolf GC, Garcia-Martinez V (1995). Primitive-streak origin and state of commitment of cells of the cardiovascular system in avian and mammalian embryos. [Review]. Cell Mol Biol Res, 41(4), 233-40.
  12. Schoenwolf GC (1994). Formation and patterning of the avian neuraxis: one dozen hypotheses. [Review]. Ciba Found Symp, 181, 25-38; discussion 38-50.
  13. Schoenwolf GC, Alvarez IS (1992). Role of cell rearrangement in axial morphogenesis. [Review]. Curr Top Dev Biol, 27, 129-73.
  14. Schoenwolf GC (1991). Cell movements driving neurulation in avian embryos. [Review]. Development, Suppl 2, 157-68.
  15. Schoenwolf GC, Smith JL (1990). Mechanisms of neurulation: traditional viewpoint and recent advances. [Review]. Development, 109(2), 243-70.
  16. Sulik, KK, GC Schoenwolf (1985). Highlights of craniofacial morphogenesis, as revealed by scanning electron microscopy. [Review]. Scan Electron Microsc, IV, 1735-1752.
  17. Schoenwolf, GC (1983). The chick ephiblast: A model for examining epithelial morphogenesis. [Review]. Scan Electron Microsc, III, 1371-1385.
  18. Schoenwolf, GC (1982). On the morphogenesis of the early rudiments of the developing central nervous system. [Review]. Scan Electron Microsc, I, 289-308.


  1. Schoenwolf, GC and Mathews, WW (2003). Atlas of Descriptive Embryology, 6th Edition. Upper Saddle River, New Jersey: Prentice Hall.
  2. Schoenwolf, GC (2001). Laboratory Studies of Vertebrate and Invertebrate Embryos. Guide and Atlas of Descriptive and Experimental Developmental, 8th Edition. Upper Saddle River, New Jersey: Prentice Hall.
  3. Mathews, WW and GC Schoenwolf (1998). Atlas of Descriptive Embryology, 5th Edition. Upper Saddle River, New Jersey: Prentice Hall.
  4. Schoenwolf, GC (1995). Laboratory Studies of Vertebrate and Invertebrate Embryos. Guide and Atlas of Descriptive and Experimental Development, 7th Edition. Englewood, New Jersey: Prentice Hall.
  5. Schoenwolf, GC, RL Watterson (1989). Laboratory Studies of Chick, Pig, and Frog Embryos. Guide and Atlas of Vertebrate Embryology, 6th Edition. New York: Macmillan Publishing Co.
  6. Schoenwolf, GC (1986). Scanning Electron Microscopy Studies of Embryogenesis. Chicago, Illinois: Scanning Electron Microscopy, Inc.
  7. Watterson, RL, GC Schoenwolf (1984). Laboratory Studies of Chick, Pig, and Frog Embryos, Guide and Atlas of Vertebrate Embryology, 5 Edition. New York: Macmillan Publishing Co.
  8. Watterson, RL, GC Schoenwolf, RM Sweeney (1979). Laboratory Studies of Chick, Pig, and Frog Embryos, 4th Edition. Minneapolis, Minnesota: Burgess Publishing co.

Book Chapter

  1. Ladher, RK and GC Schoenwolf (2003). Neural Induction. In M. Jacobson and M. Rao (Eds.), Development Neurobiology, 4th Edition. New York, In press: Plenum Presss.
  2. Lawson, A, S Lewin, GC Schoenwolf (2003). Neural Tube. In J. Opitz, G. Neri, S. Lewin (Eds.), Human Development and Malformations. Evolutionary, Genetic, and Morphogenetic Perspectives. New York, In press: John Wiley and Sons.
  3. Dias, M, GC Schoenwolf (2001). The molecular biology of early neural development. In D.L. McLone, A.E. Marlin, D.H. Reigel, R.M. Scott, M.L. Walker, P. Steinbok, W.R. Cheek (Eds.), Pediatric Neurosurgery. Surgery of the Developing Nervous System 4th Edition (pp. 73-86). W.B. Saunders Co., Publisher.
  4. Tam PPL, Goldman D, Camus A, Schoenwolf GC (2000). Early events of somitogenesis in higher vertebrates: Allocation of precursor cells at gastrulation and the organization of a meristic pattern in the paraxial mesoderm. In Ordahl CP (Ed.), Current Topics in Development Biology (47, pp. 1-32). New York: Academic Press.
  5. Inagaki, T, GC Schoenwolf, Y Yamanouchi, K Kawamoto (1999). Experimental analysis: The cause of neural tube defects and the Chiari II type malformation and the important role for normal neurulation in their prevention. In S. Matsumoto and H. Sato (Eds.), Spina Bifida (pp. 245-249). Tokyo: Springer-Verlag.
  6. Schoenwolf, GC (1999). The avian embryo: A model for descriptive and experimental embryology. In S.A. Moody (Ed.), Cell Lineage and Fate Determination (pp. 429-437). New York: Academic Press.
  7. Tam, PPL and GC Schoenwolf (1999). Cardiac fate maps: Lineage allocation, morphogenetic movement and cell commitment. In R.P. Harvey and N. Rosenthal (Eds.), Heart Development (pp. 3-18). New York: Academic Press.
  8. Schoenwolf, GC (1997). Reptiles and Birds. In S.F. Gilbert, A.M. Raunio (Eds.), Embryology: Constructing the Organism (pp. 437-458). Sunderland, Massachusetts: Sinauer Associates, Inc.
  9. Darnell, DK and GC Schoenwolf (1996). Modern techniques for cell labeling in avian and murine embryos. In G.P. Daston (Ed.), Molecular and Cellular Methods in Developmental Toxicology (pp. 231-273). Boca Raton, Florida.
  10. Schoenwolf, GC (1991). Neurepithelial cell behavior during avian neurulation. In J. Gerhart (Ed.), Cell- Cell Interactions in Early Development, 49th Symposium of the Society for Developmental Biology (pp. 63-78). New York: Wiley-Liss, Inc.
  11. Schoenwolf, GC (1991). Cell movements in the epiblast during gastrulation and neurulation in avian embryos. In Keller, R.E., W. Clark, Jr., F. Griffin (Eds.), Gastrulation: Movements, Patterns, and Molecules, Bodega Marine Laboratory Marine Series (3, pp. 1-28). New York: Plenum Press.
  12. Schoenwolf, GC (1991). Cell movements in the epiblast during gastrulation and neurulation in avian embryos. In Keller, R.E., W. Clark, Jr., F. Griffin (Eds.), Gastrulation: Movements, Patterns, and Molecules, Bodega Marine Laboratory Marine Series (3, pp. 1-28). New York: Plenum Press.
  13. Schoenwolf, GC, JL Smith (1990). Epithlial cell wedging: A fundamental cell behavior contributing to hinge point formation during epithelial morphogenesis. In R.E. Keller, D. Fristrom, (Eds.), Control of Morphogenesis by Specific Cell Behaviors Seminars in Development Biology (1, pp. 325-334). London: W.B. Saunders Co.

Case Report

  1. Bleyl SB, Byrne JL, South ST, Dries DC, Stevenson DA, Rope AF, Vianna-Morgante AM, Schoenwolf GC, Kivlin JD, Brothman A, Carey JC (2007). Brachymesomelic dysplasia with Peters anomaly of the eye results from disruptions of the X chromosome near the SHOX and SOX3 genes. Am J Med Genet A, 143A(23), 2785-95.


  1. Schoenwolf GC (2005). Lost in (color) space? Dev Dyn, 234(2), 243.
  2. Schoenwolf GC (2004). DD achieves Zen. Dev Dyn, 231(3), 461.
  3. Schoenwolf GC (2004). A special focus on the scientific contributions of Ursula K. Abbott and avian developmental genetics. Dev Dyn, 231(1), 1-3.
  4. Antin PB, Schoenwolf GC, Olsen NA (2004). A bigger bang for your buck: enhanced access to your chick data. Dev Dyn, 230(3), 391.


  1. G C Schoenwolf (1996). 1996 Senate Testimony on FY '96 Biomedical Funding [Abstract]. Congressional Record, 142.