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. NCBI Link

Scott KC, White C, Willard HF (2007) An RNA Polymerase III-Dependent Heterochromatin Barrier at Fission Yeast Centromere 1 PLoS ONE. 2: e1099. NCBI Link

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

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

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. NCBI Link

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. NCBI Link

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. NCBI Link

Valley CM, Willard HF (2006) Genomic and epigenomic approaches to the study of X chromosome inactivation. Curr Opin Genet Dev. 16: 240-245. NCBI Link

Scott KC, Merrett SL, Willard HF (2006) A heterochromatin barrier partitions the fission yeast centromere into discrete chromatin domains. Current Biol. 16: 119-129. NCBI Link

Rudd MK, Wray GA, Willard HF (2006) The evolutionary dynamics of {alpha}-satellite. Genome Res. 116: 89-96. NCBI Link

Higgins AW, Gustashaw KM, Willard HF (2005) Engineered human dicentric chromosomes show centromere plasticity. Chromosome Res. 13: 745-762. NCBI Link

Chadwick LH, Willard HF (2005) Genetic and parent-of-origin influences on X chromosome choice in X ce heterozygous mice. Mammalian Genome. 16: 691-699. NCBI Link

Basu J, Willard HF (2005) Artificial and engineered chromosomes: non-integrating vectors for gene therapy. Trends Mol. Med. 11: 251-258. NCBI Link

Ross MT,........ Willard HF et al. (2005) The DNA sequence of the human X chromosome. Nature. 434: 325-337. NCBI Link

Carrel L, Willard HF (2005) X-inactivation profile reveals extensive variability in X-linked gene expression in females. Nature. 434: 400-404. NCBI Link

Basu J, Stromberg G, Compitello 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: 587-596. NCBI Link

Rudd MK, Willard HF (2004) Analysis of the centromeric regions of the human genome assembly. Trends Genet. 20: 529-533. NCBI Link

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: R89. NCBI Link

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: 17450-17455. NCBI Link

Rudd MK, Schueler MG, Willard HF (2004) Characterization and functional annotation of human centromeres. Cold Spring Harb. Symp. Quant. Biol. 68: 141-149.

Chadwick BP, Willard HF (2003) Barring gene expression after XIST: maintaining facultative heterochromatin on the inactive X. Sem Cell Dev Biol 112: 2167-78. NCBI Link

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. NCBI Link

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. 112: 2167-78. NCBI Link

Chadwick BP, Willard HF (2003) SETting the stage. Eed-Enx 1 leaves an epigenetic signature on the inactive X chromosome. Dev Cell 4: 445-447. NCBI Link

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. NCBI Link

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.

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. NCBI Link

Chadwick BP, Willard HF (2002) Cell cycle-dependent localization of macroH2A in chromatin of the inactive X chromosome. J. Cell Biol. 157: 1113-1123. NCBI Link

Grimes BR, Rhoades AA and Willard HF (2002) Alpha-satellite DNA and vector composition influence rates of human artificial chromosome formation. Mol Ther 5: 798-805.

Percec I, Plenge RM, Nadeau JH, Bartolomei MS, Willard HF (2002) Autosomal dominant mutations affecting X inactivation choice in the mouse. Science 296: 1136-1139. NCBI Link

Chadwick BP, Valley CM, Willard HF (2001) Histone variant macroH2A contains two distinct macrochromatin domains capable of directing macroH2A to the inactive X chromosome. Nucl. Acids Res. 29: 2699-2705. NCBI Link

Chadwick BP, Willard HF (2001) Histone H2A variants and the inactive X chromosome: identification of a second macroH2A variant. Hum. Mol. Genet. 10: 1101-1113. NCBI Link

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: 375-384. NCBI Link

Schueler MG, Higgins AW, Rudd MK, Gustashaw K, Willard HF (2001) Genomic and Genetic Definition of a Functional Human Centromere. Science 294: 109-115. NCBI Link

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, Vogelstein B. New York: McGraw Hill, pp. 1191-1221.

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. NCBI Link

Tsuchiya KD, Willard HF (2000) Chromosomal domains and escape from X inactivation: comparative X inactivation analysis in mouse and human. Mammal. Genome 11: 849-54. NCBI Link

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. NCBI Link

Carrel L, Cottle AA, Goglin KC, Willard HF (1999) A first-generation X-inactivation profile of the human X chromosome. Proc. Natl. Acad. Sci. USA 96: 14440-4 NCBI Link

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. NCBI Link

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 Genet. 20: 227-228.

Willard HF (1998) Centromeres: The 'missing link' in the development of human artificial chromosomes. Curr. Opin. Genet. Dev. 8: 219-225. NCBI Link

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. NCBI Link

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. NCBI Link

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. NCBI Link

Carrel L, Hunt PA, Willard HF (1996) Tissue- and lineage-specific variation in inactive X chromosome expression of the murine Smcx gene. Hum. Mol. Genet. 5: 1361-1366. NCBI Link

Willard HF (1996) Chromosome manipulation: a systematic approach toward understanding human chromosome structure and function. Proc. Natl. Acad. Sci. USA 93: 6847-6850.

Willard HF (1996) X chromosome inactivation, XIST, and pursuit of the X inactivation center. Cell 86: 5-7.

Warburton PE, Willard HF (1996) Evolution of centromeric alpha satellite DNA: molecular organization within and between human and primate chromosomes. In: Human Genome Evolution, Jackson M, Strachan T, Dover G, eds. Oxford: BIOS Scientific Publishers, pp. 121-145.

Wolff DJ, Miller AP, VanDyke DL, Schwartz S, Willard HF (1996) Molecular definition of breakpoints associated with human Xq isochromosomes: implications for mechanisms of formation. Am. J. Hum. Genet. 58: 154-160. NCBI Link

Warburton PE, Willard HF (1995) Interhomologue sequence variation of alpha satellite DNA from human chromosome 17: evidence for concerted evolution along haplotypic lineages. J. Mol. Evol. 41: 1006-1015. NCBI Link

Brown CJ, Willard HF (1994) The human X inactivation centre is not required for maintenance of X chromosome inactivation. Nature 368: 646-648. NCBI Link

Wolff D, Brown CJ, Schwartz S, Duncan AMV, Surti S, Willard HF (1994) Small marker X chromosomes lack the X inactivation center: implications for karyotype/phenotype correlations. Am. J. Hum. Genet. 55: 87-95. NCBI Link

Hendrich BD, Brown CJ, Willard HF (1993) Evolutionary conservation of possible functional domains of the human and murine XIST genes. Hum. Mol. Genet. 2: 663-672. NCBI Link

Warburton PE, Waye JS, Willard HF (1993) Non-random localization of recombination events in human alpha satellite repeat unit variants: implications for higher-order structural characteristics within centromeric heterochromatin. Mol. Cell. Biol. 13: 6520-6529. NCBI Link

Mandel J-L, Monaco AP, Nelson DL, Schlessinger D, Willard HF (1992) Genome analysis and the human X chromosome. Science 258:103-109. NCBI Link

Brown CJ, Hendrich BD, Rupert JL, Lafreniere RG, Xing Y, Lawrence J, Willard HF (1992) The human XIST gene: analysis of a 17 kb inactive X-specific RNA that contains conserved repeats and is highly localized within the nucleus. Cell 71: 527-542. NCBI Link

Haaf T, Warburton PE, Willard HF (1992) Integration of human alpha satellite DNA into simian chromosomes: centromere protein binding and disruption of normal chromosome segregation. Cell 70:681-696. NCBI Link

Brown CJ, Ballabio A, Rupert JL, Lafreniere RG, Grompe M, Tonlorenzi R, Willard HF (1991) A gene from the region of the human X inactivation centre is expressed exclusively from the inactive X chromosome. Nature 349:38-44. NCBI Link

Brown CJ, Lafreniere RG, Powers VE, Sebastio G, Ballabio A, Pettigrew AL, Ledbetter DH, Levy E, Craig IW, Willard HF (1991) Localization of the X inactivation centre on the human X chromosome in Xq13. Nature 349:82-84. NCBI Link

Borsani G, Tonlorenzi R, Simmler MC, Dandolo L, Arnaud D, Capra V, Grompe M, Pizzuti A, Muzny D, Lawrence C, Willard HF, Avner P, Ballabio A (1991) Characterization of a murine gene expressed from the inactive X chromosome. Nature 351:325-329. NCBI Link

Warburton PE, Greig GM, Haaf T, Willard HF (1991) PCR amplification of chromosome-specific alpha satellite DNA: definition of centromeric STS markers and polymorphic analysis. Genomics 11:324-333. NCBI Link

Willard HF (1991) Evolution of alpha satellite. Curr. Opin. Genet. Devel.1:509-514. NCBI Link

Wevrick R, Willard HF (1991) Physical map of the centromeric region of human chromosome 7: relationship between two distinct alpha satellite arrays. Nucl. Acids Res. 19:2295-2301. NCBI Link

Willard HF (1990) Centromeres of mammalian chromosomes. Trends in Genetics 6:410-416. NCBI Link

Mahtani MM, Willard HF (1990) Pulsed-field gel analysis of alpha satellite DNA at the human X chromosome centromere: high-frequency polymorphisms and array size estimate. Genomics 7:607-613. NCBI Link

Wevrick R, Earnshaw WC, Howard-Peebles PN, Willard HF (1990) Partial deletion of alpha satellite DNA associated with reduced amounts of the centromere protein CENP-B in a mitotically stable human chromosome rearrangement. Mol. Cell. Biol. 10:6374-6380. NCBI Link

Warburton PE, Willard HF (1990) Genomic analysis of sequence variation in tandemly repeated DNA: evidence for localized homogeneous sequence domains within arrays of alpha satellite DNA. J. Mol. Biol. 216:3-16. NCBI Link

Durfy SJ, Willard HF (1990) Concerted evolution of primate alpha satellite DNA: evidence for an ancestral sequence shared by gorilla and human X chromosome alpha satellite. J. Mol. Biol. 216:555-566. NCBI Link

Greig GM, England SB, Bedford HM, Willard HF (1989) Chromosome-specific alpha satellite DNA from the centromere of human chromosome 16. Am. J. Hum. Genet. 45:862-872. NCBI Link

Wevrick R, Willard HF (1989) Long-range organization of tandem arrays of alpha satellite at the centromeres of human chromosomes: High frequency array length polymorphism and meiotic stability. Proc. Natl. Acad. Sci. USA 86:9394-9398. NCBI Link

Durfy SJ, Willard HF (1989) Patterns of intra- and interarray sequence variation in alpha satellite from the human X chromosome: Evidence for short range homogenization of tandemly repeated DNA sequences. Genomics 5:810-821. NCBI Link

Mahtani MM, Willard HF (1988) A primary genetic map of the pericentromeric region of the human X chromosome. Genomics 2:294-301. NCBI Link

Willard HF, Waye JS (1987) Chromosome-specific subsets of human alpha satellite DNA: Analysis of sequence divergence within and between chromosomal subsets and evidence for an ancestral pentameric repeat. J. Mol. Evol. 25:207-214. NCBI Link

Willard HF, Waye JS (1987) Hierarchical order in chromosome-specific human alpha satellite DNA. Trends in Genetics 3:192-198.

Waye JS, Willard HF (1986) Structure, organization, and sequence of alpha satellite DNA from human chromosome 17: evidence for evolution by unequal crossing-over and an ancestral pentamer repeat shared with the human X chromosome. Mol. Cell. Biol. 6:3156-3165. NCBI Link

Willard HF, Waye JS, Skolnick MH, Schwartz CE, Powers VE, England SB (1986) Restriction fragment length polymorphisms at the centromeres of human chromosomes using chromosome-specific alpha satellite DNA: implications for development of centromere-based genetic linkage maps. Proc. Natl. Acad. Sci. USA 83:5611-5615. NCBI Link

Waye JS, Willard HF (1985) Chromosome-specific alpha satellite DNA: Nucleotide sequence analysis of the 2.0 kilobasepair repeat from the human X chromosome. Nucl. Acids Res. 12:2731-2743. NCBI Link

Willard HF (1985) Chromosome-specific organization of human alpha satellite DNA. Am. J. Hum. Genet. 37:524-532. NCBI Link