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Dr Jonathan Mill  BA PhD
Senior lecturer
Psychiatric Epigenetics

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tel020 7848 0859
fax020 7848 0866

Institute of Psychiatry
Box  P80 / 82
De Crespigny Park
United Kingdom, SE5 8AF

location C308
MRC, SGDP Centre
departmentsSocial, Genetic and Developmental Psychiatry (IRG)
Social, Genetic and Developmental Psychiatry Centre (MRC)
alsoPsychiatric Epigenetics


Jonathan heads the Psychiatric Epigenetics laboratory at the Institute of Psychiatry. His group aims to explore the role of epigenetic changes in mediating risk for mental illness, with a particular focus on:

1) genomewide investigations of DNA methylation in post-mortem brain tissue for disorders such as schizophrenia, bipolar disorder, and Alzheimer’s disorder

2) investigating the role of epigenetic variation in mediating phenotypic variation between genetically-identical individuals (i.e. monozygotic twins, inbred animals)

3) elucidating how external environmental factors may bring about long-term changes in gene expression via epigenetic alterations

4) identifying novel imprinted regions of the genome, and their role in mediating parent-of-origin effects in psychiatric disorders

5) exploring interactions between genotype and epigenotype

Please visit the Psychiatric Epigenetics Group Webpage for more information.

activities and interests

Complex disease epigenomics, genomic imprinting, gene-environment interaction, monozygotic twin discordance

teaching activities

Neuroscience » Masters (MSc)
Social, Genetic & Developmental Psychiatry » Masters (MSc)


Articles in Press
Heijmans BT, Mill J. The seven plagues of epigenetic epidemiology. Int J. Epidemiol. In Press.

Bell JT*, Tsai P-C*, Yang T-P, Pidsley R, Nisbet J, Glass D, Zhai G, Mangino M, Zhai G, Zhang F, Valdes A, Shin S-Y, Dempster EL, Murray RM, Grundberg E, Hedman AK, Nica A, Small KS, The MuTHER Consortium, Dermitzakis ET, McCarthy MI, Mill J, Spector TD*, Deloukas P*. 2012. Epigenome-wide scans identify differentially methylated regions for age and age- related phenotypes in a healthy ageing population. PLoS Genetics, in press.

Pidsley R, Dempster E, Troakes C, Al-Sarraj S, Mill J. Epigenetic and genetic variation at the IGF2/H19 imprinting control region on 11p15.5 is associated with cerebellum weight. Epigenetics, in press.

Boks MP, de Jong NM, Kas MJ, Vinkers CH, Fernandes C, Kahn RS, Mill J, Ophoff RA. Current status and future prospects for epigenetic psychopharmacology. Epigenetics. 2012 Jan 1;7(1).

Dempster EL, Pidsley R, Schalkwyk LC, Owens S, Georgiades A, Kane F, Kalidindi S, Picchioni M, Kravariti E, Toulopoulou T, Murray RM, Mill J. Disease-associated epigenetic changes in monozygotic twins discordant for schizophrenia and bipolar disorder. Hum Mol Genet. 2011 Sep 9.

Pidsley R, Mill J. Research Highlights: epigenetic changes to serotonin receptor gene expression in schizophrenia and bipolar disorder. Epigenomics. 2011 Oct;3(5):537-8.

Voineagu I, Wang X, Johnston P, Lowe JK, Tian Y, Horvath S, Mill J, Cantor RM, Blencowe BJ, Geschwind DH. Transcriptomic analysis of autistic brain reveals convergent molecular pathology. Nature. 474(7351):380-4.

Engmann O, Hortobágyi T, Pidsley R, Troakes C, Bernstein HG, Kreutz MR, Mill J, Nikolic M, Giese KP. Schizophrenia is associated with dysregulation of the Cdk5 activator p35 resulting in deficits of synaptic protein expression and cognition. Brain. 134(Pt 8):2408-21.

Kaminsky Z, Tochigi M, Jia P, Mill J, Kwan A, Ioshikhes I, Vincent J, Kennedy J, Strauss J, Pai S, Wang SC, Petronis A. A multi-tissue analysis identifies HLA Complex Group 9 gene methylation differences in bipolar disorder. Molecular Psychiatry. In Press.

Wong CC, Caspi A, Williams B, Houts R, Craig I, Mill J. A Longitudinal Twin Study of Skewed X Chromosome Inactivation. PloS one. 2011

Wong CC, Mill J, Fernandes C. Drugs and addiction: an introduction to epigenetics. Addiction. 2011 Mar;106(3):480-9

Campbell IC, Mill J, Uher R, Schmidt U. Eating disorders, gene-environment interactions and epigenetics. Neuroscience and Biobehavioral reviews. 2011 Jan;35(3):784-93.

Wong CC, Caspi A, Williams B, Craig IW, Houts R, Ambler A, Moffitt TE, Mill J. A Longitudinal Study of Epigenetic Variation in Twins. Epigenetics. 2010 Aug 4;5(6).

Schalkwyk, L.C., Meaburn, E.L., Smith, R.G., Dempster, E., Jeffries, A.R., Davies, M.N., Plomin, R., and Mill, J. Allelic skewing of DNA methylation is widespread across the genome. Am J Hum Genet. 2010 Feb 12;86(2):196-212.

Wong CC, Caspi A, Williams B, Craig IW, Houts R, Ambler A, Moffitt TE, Mill J. A longitudinal study of epigenetic variation in twins. Epigenetics. 2010 Aug 4;5(6).

Pidsley R, Mill J. Epigenetic studies of psychosis: current findings, methodological approaches, and implications for postmortem research. Biological psychiatry. 2010 Jan 15;69(2):146-56.

Pidsley R, Dempster EL, Mill J. Brain weight in males is correlated with DNA methylation at IGF2. Molecular Psychiatry. 2010 Sep;15(9):880-1.

Meaburn EL, Schalkwyk LC, Mill J. Allele-specific methylation in the human genome Implications for genetic studies of complex disease. Epigenetics. 2010 Oct 9;5(7).

Lundstrom S, Haworth CM, Carlstrom E, Gillberg C, Mill J, Rastam M, Hultman CM, Ronald A, Anckarsater H, Plomin R, Lichtenstein P, Reichenberg A. Trajectories leading to autism spectrum disorders are affected by paternal age: findings from two nationally representative twin studies. Journal of child psychology and psychiatry, and allied disciplines. 2010 Jul;51(7):850-6.

Docherty SJ, Davis OS, Haworth CM, Plomin R, Mill J. DNA methylation profiling using bisulfite-based epityping of pooled genomic DNA. Methods (San Diego, Calif. 2010 Nov;52(3):255-8.

Bell CG, Finer S, Lindgren CM, Wilson GA, Rakyan VK, Teschendorff AE, Akan P, Stupka E, Down TA, Prokopenko I, Morison IM, Mill J, Pidsley R, Deloukas P, Frayling TM, Hattersley AT, McCarthy MI, Beck S, Hitman GA. Integrated genetic and epigenetic analysis identifies haplotype-specific methylation in the FTO type 2 diabetes and obesity susceptibility locus. PloS one. 2010;5(11):e14040.

Smith, R.G., Kember, R.L., Mill, J., Fernandes, C., Schalkwyk, L.C., Buxbaum, J.D., and Reichenberg, A. (2009). Advancing paternal age is associated with deficits in social and exploratory behaviors in the offspring: a mouse model. PLoS One 4, e8456.

Docherty SJ, Davis OS, Haworth CM, Plomin R, Mill J. Bisulfite-based epityping on pooled genomic DNA provides an accurate estimate of average group DNA methylation. Epigenetics Chromatin. 2009 Mar 10;2(1):3.

Rutten, B. and Mill, J. (2009). Epigenetic Mediation of Environmental Influences in Major Psychotic Disorders. Schizophr Bull. 2009 Nov;35(6):1045-56.

Xu, X., Mill, J., Sun, B., Chen, C.K., Huang, Y.S., Wu, Y.Y., and Asherson, P. (2009). Association study of promoter polymorphisms at the dopamine transporter gene in Attention Deficit Hyperactivity Disorder. BMC Psychiatry 9, 3.

Reichenberg, A., Mill, J., and MacCabe, J.H. (2009). Epigenetics, genomic mutations and cognitive function. Cogn Neuropsychiatry 14, 377-90.

Mill, J., Wigg, K., Burcescu, I., Vetro, A., Kiss, E., Kapornai, K., Tamas, Z., Baji, I., Gadoros, J., Kennedy, J.L., Kovacs, M., and Barr, C.L. (2009). Mutation screen and association analysis of the glucocorticoid receptor gene (NR3C1) in childhood-onset mood disorders (COMD). Am J Med Genet B Neuropsychiatr Genet 150B, 866-73.

Mill J, Petronis A. Profiling DNA methylation from small amounts of genomic DNA starting material: efficient sodium bisulfite conversion and subsequent whole-genome amplification. Methods Mol Biol. 2009;507:371-81.

Mill J, Tang T, Kaminsky Z, Khare T, Yazdanpanah S, Bouchard L, Jia P, Assadzadeh A, Flanagan J, Schumacher A, Wang SC, Petronis A. Epigenomic Profiling Reveals DNA Methylation Changes Associated with Major Psychosis. American Journal of Human Genetics. 2008 82(3): 696-711

Mill J, Petronis A. Pre- and peri-natal environmental risks for attention-deficit hyperactivity disorder: the role of epigenetic processes in mediating susceptibility. The Journal of Child Psychology and Psychiatry. 2008 May 19. [Epub ahead of print]

Docherty S, Mill J. Epigenetic mechanisms as mediators of environmental risks for psychiatric disorders. Psychiatry. 2008. 7(12):500-506.

Mill J, Kiss E, Baji I, Kapornai K, Daróczy G, Vetró A, Kennedy J, Kovacs M, Barr C; International Consortium for Childhood-Onset Mood Disorders. Association study of the estrogen receptor alpha gene (ESR1) and childhood-onset mood disorders. Am J Med Genet B Neuropsychiatr Genet. 2008 Oct 5;147B(7):1323-6.

Brookes KJ, Neale B, Xu X, Thapar A, Gill M, Langley K, Hawi Z, Mill J, Taylor E, Franke B, Chen W, Ebstein R, Buitelaar J, Banaschewski T, Sonuga-Barke E, Eisenberg J, Manor I, Miranda A, Oades RD, Roeyers H, Rothenberger A, Sergeant J, Steinhausen HC, Faraone SV, Asherson P. Differential dopamine receptor D4 allele association with ADHD dependent of proband season of birth. Am J Med Genet B Neuropsychiatr Genet. 2008 Jan 5;147(1):94-9.

Mill J. Rodent models: utility for candidate gene studies in human attention-deficit hyperactivity disorder (ADHD). Journal of Neuroscience Methods 2007 Jan 16; [Epub ahead of print]

Mill J, Petronis A. Molecular Studies of Major Depressive Disorder: The Epigenetic Perspective. Molecular Psychiatry. 2007; 12(9):799-814.

Brookes KJ, Mill J, Guindalini C, Curran S, Xu X, Knight J, Chen CK, Huang YS, Sethna V, Taylor E, Chen W, Breen G, Asherson P. A common haplotype of the dopamine transporter gene associated with attention-deficit/hyperactivity disorder and interacting with maternal use of alcohol during pregnancy. Arch Gen Psychiatry. 2006, 63(1):74-81.

Dempster EL, Mill J, Craig IW, Collier DA. The quantification of COMT mRNA in post mortem cerebellum tissue: diagnosis, genotype, methylation and expression. BMC Med Genet. 2006 Feb 16;7(1):10

Mill J, Dempster E, Caspi A, Williams B, Moffitt T, Craig I. Evidence for monozygotic twin (MZ) discordance in methylation level at two CpG sites in the promoter region of the Catechol-O-Methyltransferase (COMT) gene. Am J Med Genet B Neuropsychiatr Genet. 2006 Jun 5;141(4):421-5.

Mill J, Caspi A, Williams B, Craig I, Taylor A, Polo-Tomas M, Berridge C, Poulton R, Moffitt T. Genetic polymorphisms in the dopamine system account for cognitive heterogeneity among children with attention-deficit hyperactivity disorder: Replicated evidence from two birth cohorts. Arch Gen Psychiatry. 2006 Apr;63(4):462-9

Xu X, Mill J, Zhou K, Brookes K, Chen CK, Asherson P. Family-based association study between brain-derived neurotrophic factor gene polymorphisms and attention deficit hyperactivity disorder in UK and Taiwanese samples. Am J Med Genet B Neuropsychiatr Genet. 2006 Oct 16; [Epub ahead of print]

Mill J, Yazdanpanah S, Gückel E, Ziegler S, Kaminsky Z, Petronis A. Whole genome amplification of sodium bisulfite-treated DNA allows the accurate estimate of methylated cytosine density in limited DNA resources. BioTechniques. 2006 41(5): 603-607

Mill, J.; Asherson, P.; Craig, I.; D’Souza, U. M. Transient Expression Analysis of Allelic Variants of a VNTR in the Dopamine Transporter Gene (DAT1). BMC. Genet 2005, 6, 3.

Mill, J.; Xu, X.; Ronald, A.; Curran, S.; Price, T.; Knight, J.; Craig, I.; Sham, P.; Plomin, R.; Asherson, P. Quantitative Trait Locus Analysis of Candidate Gene Alleles Associated With Attention Deficit Hyperactivity Disorder (ADHD) in Five Genes: DRD4, DAT1, DRD5, SNAP-25, and 5HT1B. Am. J Med Genet B Neuropsychiatr. Genet 2005, 133, 68-73.

Xu, X.; Knight, J.; Brookes, K.; Mill, J.; Sham, P.; Craig, I.; Taylor, E.; Asherson, P. DNA Pooling Analysis of 21 Norepinephrine Transporter Gene SNPs With Attention Deficit Hyperactivity Disorder: No Evidence for Association. Am. J Med Genet B Neuropsychiatr. Genet 2005, 134, 115-118.

Xu X, Mill J, Chen CK, Brookes K, Taylor E, Asherson P. Family-based association study of serotonin transporter gene polymorphisms in attention deficit hyperactivity disorder: no evidence for association in UK and Taiwanese samples. Am J Med Genet B Neuropsychiatr Genet. 2005, 139(1):11-3.

Mill J, Sagvolden T, Asherson P. Sequence analysis of Drd2, Drd4, and Dat1 in SHR and WKY rat strains. Behav Brain Funct. 2005 15;1(1):24.

Craig, I. W.; Mill, J.; Craig, G. M.; Loat, C.; Schalkwyk, L. C. Application of Microarrays to the Analysis of the Inactivation Status of Human X-Linked Genes Expressed in Lymphocytes. Eur J Hum Genet 2004, 12, 639-646.

D’Souza, U. M.; Russ, C.; Tahir, E.; Mill, J.; McGuffin, P.; Asherson, P. J.; Craig, I. W. Functional Effects of a Tandem Duplication Polymorphism in the 5’Flanking Region of the DRD4 Gene. Biol Psychiatry 2004, 56, 691-697.

Lowe, N.; Kirley, A.; Hawi, Z.; Sham, P.; Wickham, H.; Kratochvil, C. J.; Smith, S. D.; Lee, S. Y.; Levy, F.; Kent, L.; Middle, F.; Rohde, L. A.; Roman, T.; Tahir, E.; Yazgan, Y.; Asherson, P.; Mill, J.; Thapar, A.; Payton, A.; Todd, R. D.; Stephens, T.; Ebstein, R. P.; Manor, I.; Barr, C. L.; Wigg, K. G.; Sinke, R. J.; Buitelaar, J. K.; Smalley, S. L.; Nelson, S. F.; Biederman, J.; Faraone, S. V.; Gill, M. Joint Analysis of the DRD5 Marker Concludes Association With Attention-Deficit/Hyperactivity Disorder Confined to the Predominantly Inattentive and Combined Subtypes. Am. J Hum Genet 2004, 74, 348-356.

Mill, J.; Richards, S.; Knight, J.; Curran, S.; Taylor, E.; Asherson, P. Haplotype Analysis of SNAP-25 Suggests a Role in the Aetiology of ADHD. Mol. Psychiatry 2004, 9, 801-810.

Mill, J.; Curran, S.; Richards, S.; Taylor, E.; Asherson, P. Polymorphisms in the Dopamine D5 Receptor (DRD5) Gene and ADHD. Am. J Med Genet B Neuropsychiatr. Genet 2004, 125, 38-42.

Caspi A, Sugden K, Moffitt TE, Taylor A, Craig IW, Harrington H, McClay J, Mill J, Martin J, Braithwaite A, Poulton R. Influence of life stress on depression: moderation by a polymorphism in the 5-HTT gene. Science 2003 Jul 18;301(5631):386-9.

Mill, J.; Fisher, N.; Curran, S.; Richards, S.; Taylor, E.; Asherson, P. Polymorphisms in the Dopamine D4 Receptor Gene and Attention-Deficit Hyperactivity Disorder. Neuroreport 2003, 14, 1463-1466.

Chen, C. K.; Chen, S. L.; Mill, J.; Huang, Y. S.; Lin, S. K.; Curran, S.; Purcell, S.; Sham, P.; Asherson, P. The Dopamine Transporter Gene Is Associated With Attention Deficit Hyperactivity Disorder in a Taiwanese Sample. Mol. Psychiatry 2003, 8, 393-396.

Freeman B, Smith N, Curtis C, Huckett L, Mill J, Craig IW. DNA from buccal swabs recruited by mail: evaluation of storage effects on long-term stability and suitability for multiplex polymerase chain reaction genotyping. Behav Genet. 2003, Jan;33(1):67-72.

Caspi A, McClay J, Moffitt TE, Mill J, Martin J, Craig IW, Taylor A, Poulton R. Role of genotype in the cycle of violence in maltreated children. Science. 2002 Aug 2;297(5582):851-4.

Holmes, J.; Payton, A.; Barrett, J.; Harrington, R.; McGuffin, P.; Owen, M.; Ollier, W.; Worthington, J.; Gill, M.; Kirley, A.; Hawi, Z.; Fitzgerald, M.; Asherson, P.; Curran, S.; Mill, J.; Gould, A.; Taylor, E.; Kent, L.; Craddock, N.; Thapar, A. Association of DRD4 in Children With ADHD and Comorbid Conduct Problems. Am. J Med Genet 2002, 114, 150-153.

Mill, J.; Galsworthy, M. J.; Paya-Cano, J. L.; Sluyter, F.; Schalkwyk, L. C.; Plomin, R.; Asherson, P. Home-Cage Activity in Heterogeneous Stock (HS) Mice As a Model of Baseline Activity. Genes Brain Behav. 2002, 1, 166-173.

Mill, J.; Asherson, P.; Browes, C.; D’Souza, U.; Craig, I. Expression of the Dopamine Transporter Gene Is Regulated by the 3’ UTR VNTR: Evidence From Brain and Lymphocytes Using Quantitative RT-PCR. Am. J Med Genet 2002, 114, 975-979.

Mill, J.; Curran, S.; Kent, L.; Gould, A.; Huckett, L.; Richards, S.; Taylor, E.; Asherson, P. Association Study of a SNAP-25 Microsatellite and Attention Deficit Hyperactivity Disorder. Am. J Med Genet 2002, 114, 269-271.

Mill, J. S.; Caspi, A.; McClay, J.; Sugden, K.; Purcell, S.; Asherson, P.; Craig, I.; McGuffin, P.; Braithwaite, A.; Poulton, R.; Moffitt, T. E. The Dopamine D4 Receptor and the Hyperactivity Phenotype: a Developmental-Epidemiological Study. Mol. Psychiatry 2002, 7, 383-391.

Curran, S.; Mill, J.; Tahir, E.; Kent, L.; Richards, S.; Gould, A.; Huckett, L.; Sharp, J.; Batten, C.; Fernando, S.; Ozbay, F.; Yazgan, Y.; Simonoff, E.; Thompson, M.; Taylor, E.; Asherson, P. Association Study of a Dopamine Transporter Polymorphism and Attention Deficit Hyperactivity Disorder in UK and Turkish Samples. Mol. Psychiatry 2001, 6, 425-428.

Mill, J.; Curran, S.; Kent, L.; Richards, S.; Gould, A.; Virdee, V.; Huckett, L.; Sharp, J.; Batten, C.; Fernando, S.; Simanoff, E.; Thompson, M.; Zhao, J.; Sham, P.; Taylor, E.; Asherson, P. Attention Deficit Hyperactivity Disorder (ADHD) and the Dopamine D4 Receptor Gene: Evidence of Association but No Linkage in a UK Sample. Mol. Psychiatry 2001, 6, 440-444.

Curran, S.; Mill, J.; Sham, P.; Rijsdijk, F.; Marusic, K.; Taylor, E.; Asherson, P. QTL Association Analysis of the DRD4 Exon 3 VNTR Polymorphism in a Population Sample of Children Screened With a Parent Rating Scale for ADHD Symptoms. Am. J Med Genet 2001, 105, 387-393.

last updated: Monday, February 27, 2012