Faculty and Research

Huntington F. Willard, PhD
Nanaline H. Duke Professor
bio  •  lab members  •  lab website

Representative Publications

  • Horvath JE, Weisrock DW, Embry SL, Fiorentino I, Balhoff JP, Kappeler P, Wray GA, Willard HF, Yoder AD (2008). Development and application of a phylogenomic toolkit: Resolving the evolutionary history of Madagascar's lemurs. Genome Research. 18:489-499.

  • Horvath JE, Willard HF (2007). Primate comparative genomics: Lemur biology and evolution. Trends in Genetics. 23:173-182.

  • Scott KC, White C, Willard HF (2007). An RNA polymerase III-dependent heterochromatin barrier at fission yeast centromere 1. PLoS ONE. 2:e1099.

  • Basu J, Willard HF (2006). Human artificial chromosomes: Potential applications and clinical considerations. Pediatr Clin North Am. 53:843-853.

  • Wang Z, Willard HF, Mukherjee S, Furey TS (2006). Evidence of influence of genomic DNA sequence on human X chromosome inactivation. PLoS Comput Biol. 2:979-988.

  • Amos-Landgraf JM, Cottle A, Plenge RM, Friez M, Schwartz CE, Longshore J, Willard HF (2006). X chromosome-inactivation patterns of 1,005 phenotypically unaffected females. Amer J Hum Genet. 79:493-499.

  • Valley CM, Pertz LM, Balakumaran BS, Willard HF (2006). Chromosome-wide, allele-specific analysis of the histone code on the human X chromosome. Hum Mol Genet. 15:2335-2347.
  • Valley CM, Willard HF (2006). Genomic and epigenomic approaches to the study of X chromosome inactivation. Curr Opin Genet Dev. 16:240-245.

  • Scott KC, Merrett SL, Willard HF (2006). A heterochromatin barrier partitions the fission yeast centromere into discrete chromatin domains. Current Biol. 16:119-129.
  • Rudd MK, Wray GA, Willard HF (2006). The evolutionary dynamics of {alpha}-satellite. Genome Res. 116:89-96.
  • Higgins Aw, Gustashaw KM, Willard HF (2005). Engineered human dicentric chromosomes show centromere plasticity. Chromosome Res. 13:745-762.
  • Chadwick LH, Willard HF (2005). Genetic and parent-of-origin influences on X chromosome choise in X ce heterozygous mice. Mammalian Genome. 16:691-699.
  • Schueler MJ, Dunn JM, Bird CP, Ross MT, Viggiano L, Rocchi M, Willard HF, and Green ED (2005). NISC Comparative Sequencing Program. Progressive proximal expansion of the primate X chromosome centromere. Proc. Natl. Acad. Sci. USA 102:10563-10568.
  • Basu J and Willard HF (2005). Artificial and engineered chromosomes: non-integrating vectors for gene therapy. Trends Mol. Med. 11:251-258.
  • Ross MT, . . . Willard HF et al. (2005). The DNA sequence of the human X chromosome. Nature 434: 325-337.
  • Carrel L, Willard HF (2005). X-inactivation profile reveals extensive variability in X-linked gene expression in females. Nature 434: 400-404.
  • Basu J, Stromberg G, Willard HF, Van Bokkelen G (2005) Rapid creation of BAC-based human artificial chromosome vectors by transposition with synthetic alpha-satellite arrays. Nucleic Acids Res. 33(2):587-96.
  • Rudd MK, Willard HF (2004) Analysis of the centromeric regions of the human genome assembly. Trends Genet. 20(11):529-33.
  • Grimes BR, Babcock J, Rudd MK, Chadwick B, Willard HF (2004) Assembly and characterization of heterochromatin and euchromatin on human artificial chromosomes. Genome Biol. 5(11):R89.
  • Chadwick BP, Willard HF. (2004).Multiple spatially distinct types of facultative heterochromatin on the human inactive X chromosome. Proc Natl Acad Sci U S A. 101(50):17450-5.
  • Rudd MK, Schueler MG, Willard HF (2004). Characterization and functional annotation of human centromeres. Cold Spring Harb. Symp. Quant. Biol. 68: 141-149.
  • Rudd MK, Mays RW, Schwartz S, Willard HF (2003). Human artificial chromosomes with alpha satellite-based de novo centromeres show increased frequency of nondisjunction and anaphase lag. Mol. Cell. Biol. 23: 7689-7697.
  • Chadwick BP, Willard, HF (2003). Chromatin of the Barr Body: Histone and non-histone proteins associated with or excluded from the inactive X chromosome. Hum. Mol. Genet. 12: 2167-2178.
  • Percec I, Thorvaldsen JL, Plenge RM, Krapp CJ, Nadeau JH, Willard HF, Bartolomei MS (2003). An ENU mutagenesis screen for epigenetic mutations in the mouse. Genetics 164: 1481-1494.
  • Hall LL, Byron M, Sakai K, Carrel L, Willard HF, Lawrence JB (2002). An ectopic human XIST gene can induce chromosome inactivation in postdifferentiation human HT1080 cells. Proc. Natl. Acad. Sci. USA 99: 8677-8692.
  • Chadwick BP, Willard, HF (2002). Cell cycle-dependent localization of macroH2A in chromatin of the inactive X chromosome. J. Cell Biol. 157: 1113-1123.
  • Plenge, RM, Stevenson, RA, Lubs, HA, Schwartz, CE, Willard, HF (2002). Skewed X chromosome inactivation is a common feature of x-linked mental retardation disorders. Am. J. Hum. Genet. 71: 168-173.
  • Grimes BR, Rhoades AA, Willard HF (2002). Alpha satellite and vector composition influence rates of human artificial chromosome formation. Mol. Therapy 5: 798-805.
  • Percec, I, Pleange, RM, Nadeau, JH, Bartolomei, MS, Willard, HF (2002). Autosomal dominant mutations affecting X inactivation choice in the mouse. Science 296: 1136-1139.
  • Schueler, MG, Higgins, AW, Rudd, MK, Gustashaw, K, Willard HF (2001). Genomic and genetic definition of a functional human centromere. Science 294: 109-115.
  • Chadwick BP, Willard HF (2001). A Novel Chromatin Protein, Distantly Related to Histone H2A, Is Largely Excluded From the Inactive X Chromosome. J. Cell Biol.152(2): 375-384
  • Tsuchiya KD, Willard HF (2000). Chromosomal domains and escape from X inactivation: comparative X inactivation analysis in mouse and human. Mammal. Genome 11: 849-854.
  • Plenge RM, Percec I, Nadeau JH, Willard HF (2000). Expression-based assay of an X-linked gene to examine effects of the X-controlling element (Xce) locus. Mammal. Genome 11: 405-408.
  • Schueler MG, Higgins AW, Nagaraja R, Tentler D, Dahl N, Gustashaw K, Willard HF (2000). Large-insert clone / STS contigs in Xq11-q12, spanning deletions in patients with androgen insensitivity and mental retardation. Genomics 66: 104-109.
  • Willard HF (2000) The sex chromosomes and X chromosome inactivation. In: The Metabolic and Molecular Bases of Inherited Disease, 8th ed. Edited by Scriver CR, Beaudet AL, Sly WS, Valle D, Childs B, Vogelstien B. New York: McGraw-Hill (1191-1221).
  • Willard HF, Hendrich DB (1999). Breaking the silence in Rett syndrome [News & Views]. Nature Genet. 23:127-128
  • Higgins AW, Schueler MG, Willard HF (1999). Chromosome engineering: generation of mono- and dicentric isochromosomes in a somatic cell hybrid system. Chromosoma 108: 256-265.
  • Carrel L, Cottle A, Goglin KC, Willard HF (1999). A first-generation X inactivation profile of the human X chromosome. Proc. Natl. Acad. Sci. USA 96: 14440-14444.
  • Plenge RM, Tranebjaerg L, Jensen PKA, Schwartz C, Willard HF (1999). Evidence that mutations in the X-linked DDP gene cause incompletely penetrant and variable skewed X inactivation. Am. J. Hum. Genet. 64: 759-767.
  • Carrel L, Willard HF (1999). Heterogeneous gene expression from the inactive X chromosome: an X-linked gene that escapes X inactivation in some human cell lines but is inactivated in others. Proc. Natl. Acad. Sci. USA 96: 7364-7369.
  • Puck JM, Willard HF (1998). X inactivation in females with X-linked disease. New Engl. J. Med. 338: 291-295.
  • Mahtani MM, Willard HF (1998). Physical and genetic mapping of the human X chromosome centromere: repression of recombination. Genome Res. 8: 100-110.
  • Willard HF (1998). Centromeres: the ‘missing link’ in the development of human artificial chromosomes. Curr. Opin. Genet. Dev. 8: 219-225.
  • Carrel L, Willard HF (1998). Counting on Xist. Nature Genetics 19: 211-212.
  • Miller AP, Willard HF (1998). Chromosomal basis for X chromosome inactivation: identification of a multigene domain in Xp11.21-11.22 that escapes inactivation. Proc. Natl. Acad. Sci. USA 95: 8709-8714.
  • Sullivan BA, Willard HF (1998). Stable dicentric X chromosomes with two active centromeres. Nature Genetics 20: 227-228.
  • Harrington JJ, Van Bokkelen G, Mays RW, Gustashaw K, Willard HF (1997). Formation of de novo centromeres and construction of first-generation human artificial microchromosomes. Nature Genetics 15: 345-355.
  • Plenge RM, Hendrich BD, Schwartz C, Arena JF, Naumova A, Sapienza C, Winter RM, Willard HF (1997). A promoter mutation in the XIST gene in two unrelated families with skewed X chromosome inactivation. Nature Genetics 17: 353-356.
  • Brown CJ, Carrel L, Willard HF (1997). Expression of genes from the human active and inactive X chromosomes. Am. J. Hum. Genet. 60: 1333-1343.