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Table 1 Methylated target genes identified by methylated-CpG island recovery assay-assisted microarray analysis

From: Methylation of homeobox genes is a frequent and early epigenetic event in breast cancer

Target number

Gene

Description

Gene location

Strand

Position relative to genea

Average ratiob

Count

1*

FOXE1

Forkhead box E1

Chr9: 99655357 to 99658818

+

Downstream

13.76

4

2

SEMA6C

Semaphorin 6C

Chr1: 149370786 to 149385728

-

Downstream

6.40

4

3*

HNF1B c

Hnf1 homeobox B

Chr17: 33120546 to 33179209

-

Upstream

6.06

4

4

OTX1

Orthodenticle homeobox 1

Chr2: 63131468 to 63137816

+

Intragenic

4.83

4

5*

TLX1

T-cell leukemia homeobox 1

Chr10: 102881050 to 102887535

+

Intragenic

4.50

4

6

ESAM

Endothelial cell adhesion molecule

Chr11: 124128228 to 124137433

-

Intragenic

4.16

4

7*

CNTNAP1

Contactin associated protein 1

Chr17: 38088158 to 38105358

+

Intragenic

3.94

4

8*

GFI1

Growth factor independent 1 transcription repressor

Chr1: 92712905 to 92725021

-

Intragenic

3.64

4

9*

RASL10A

Ras-like, family 10, member A

Chr22: 28038921 to 28041748

-

Intragenic

3.42

4

10*

IFNA8

Interferon, alpha 8

Chr9: 21399146 to 21400184

+

Upstream

3.41

4

11

ERGIC2

Ergic and golgi 2

Chr12: 29384845 to 29425410

-

Upstream

3.39

4

12

PCDH7

Protocadherin 7

Chr4: 30331134 to 30757519

+

Intragenic

3.16

4

13

RASGRP2

Ras guanyl releasing protein 2

Chr11: 64250958 to 64269504

-

Intragenic

3.01

4

14

NFATC1

Nuclear factor of activated T-cells 1

Chr18: 75256759 to 75390311

+

Intragenic

2.93

4

15

RLTPR

Rgd motif, leucine-rich repeats, tropomodulin domain and proline-rich containing gene

Chr16: 66236530 to 66248973

+

Intragenic

2.75

4

16

AK123344

Hypothetical gene

Chr10: 102979341 to 102985256

+

Downstream

16.49

3

17

C1orf114

Chromosome 1 open reading frame 114

Chr1: 167630737 to 167663294

-

Upstream

11.13

3

18

TRPS1

Trichorhinophalangeal syndrome I

Chr8: 116489899 to 116750402

-

Intragenic

11.07

3

19

PCDHGA12

Protocadherin gamma subfamily A, 12

Chr5: 140790341 to 140872730

+

Upstream

11.02

3

20

C14orf25

Chromosome 14 open reading frame 25

Chr14: 37150207 to 37580397

+

Upstream

10.34

3

21

CDKN2A

Cyclin-dependent kinase inhibitor 2A (P16)

Chr9: 21957750 to 21984490

-

Downstream

9.86

3

22

SCRT2

Scratch homolog 2, zinc finger protein

Chr20: 590240 to 604823

-

Downstream

9.47

3

23

BCOR

Bcl6 co-repressor

Chrx: 39795443 to 39921526

-

Upstream

9.43

3

24

KHDC1

Kh homology domain containing 1

Chr6: 74007759 to 74076659

-

Upstream

9.00

3

25*

NXPH1

Neurexophilin 1

Chr7: 8440109 to 8759118

+

Intragenic

8.83

3

26

CNR1

Cannabinoid receptor 1 (brain)

Chr6: 88910156 to 88932281

-

Upstream

8.78

3

27

BC039088

Hypothetical gene

Chr5: 43050280 to 43054670

-

Upstream

8.61

3

28

EVX2

Even-skipped homeobox 2

Chr2: 176653080 to 176656936

-

Downstream

8.43

3

29*

MT1E

Metallothionein 1E

Chr16: 55217085 to 55218525

+

Upstream

8.36

3

30*

NR2F2

Nuclear receptor subfamily 2, group F, member 2

Chr15: 94674950 to 94683047

+

Downstream

8.24

3

31

HOXC13

Homeobox C13

Chr12: 52618843 to 52626595

+

Upstream

8.08

3

32

HOXD8

Homeobox D8

Chr2: 176702722 to 176704974

+

Upstream

7.97

3

33

SYCP2L

Synaptonemal complex protein 2-like

Chr6: 10995049 to 11082527

+

Upstream

7.58

3

34

PCDHGB6

Protocadherin gamma subfamily B, 6

Chr5: 140767953 to 140872730

+

Intragenic

7.37

3

35

ACTA1

Actin, alpha 1, skeletal muscle

Chr1: 227633615 to 227636466

-

Upstream

7.36

3

36*

PRDM14

Pr domain containing 14

Chr8: 71126576 to 71146116

-

Intragenic

7.27

3

37*

HOXB13

Homeobox B13

Chr17: 44157124 to 44161110

-

Intragenic

6.91

3

38

OTX2

Orthodenticle homeobox 2

Chr14: 56337177 to 56346937

-

Upstream

6.70

3

39

ZNF711

Zinc finger protein 711

Chrx: 84385652 to 84415025

+

Upstream

6.63

3

40*

NR2E1

Nuclear receptor subfamily 2, group E, member 1

Chr6: 108593954 to 108616706

+

Intragenic

6.44

3

41*

TAC1

Tachykinin, precursor 1

Chr7: 97199206 to 97207720

+

Upstream

6.37

3

42*

CPEB1

Cytoplasmic polyadenylation element binding protein 1

Chr15: 81009005 to 81113783

-

Upstream

5.99

3

43

NKAPL

Nfkb activating protein-like

Chr6: 28335076 to 28336715

+

Upstream

5.86

3

44

NKX6-2

Nk6 homeobox 2

Chr10: 134448309 to 134449527

-

Downstream

5.72

3

45*

TGIF2

Tgfb-induced factor homeobox 2

Chr20: 34635423 to 34655766

+

Upstream

5.60

3

46

CR596471

Hypothetical gene

Chrx: 133511720 to 133522094

+

Upstream

5.56

3

47

EVX2

Even-skipped homeobox 2

Chr2: 176653080 to 176656936

-

Intragenic

5.55

3

48

AX747981

Hypothetical gene

Chr8: 96148213 to 96154324

-

Upstream

5.48

3

49

ST8SIA3

St8 alpha-N-acetyl-neuraminide alpha-2,8-sialyltransferase 3

Chr18: 53170718 to 53187159

+

Intragenic

5.47

3

50

WT1

Wilms tumor 1

Chr11: 32365900 to 32413663

-

Intragenic

5.40

3

51

BARHL2

Barh-like homeobox 2

Chr1: 90950167 to 90955382

-

Upstream

5.39

3

52

IRX1

Iroquois homeobox 1

Chr5: 3649167 to 3654517

+

Intragenic

5.39

3

53

KLF11

Kruppel-like factor 11

Chr2: 10101132 to 10112414

+

Upstream

5.26

3

54

HLXB9

Homeobox Hb9

Chr7: 156479507 to 156496108

-

Upstream

5.25

3

55

NKX2-8

Nk2 homeobox 8

Chr14: 36118966 to 36121537

-

Upstream

5.20

3

56

ZFP91

Zinc finger protein 91 homolog (mouse)

Chr11: 58103162 to 58145091

+

Upstream

5.07

3

57

LHX2

Lim homeobox 2

Chr9: 125813709 to 125835263

+

Intragenic

5.04

3

58

MEIS1

Meis homeobox 1

Chr2: 66516036 to 66653085

+

Downstream

5.00

3

59*

PAX2

Paired box 2

Chr10: 102495457 to 102579688

+

Intragenic

4.95

3

60*

LHX9

Lim homeobox 9

Chr1: 196153139 to 196165896

+

Intragenic

4.81

3

61

GRASP

Grp1-associated scaffold protein

Chr12: 50687014 to 50695938

+

Upstream

4.78

3

62

LHX8

Lim homeobox 8

Chr1: 75366706 to 75399806

+

Intragenic

4.77

3

63

HOXD12

Homeobox D12

Chr2: 176672775 to 176673734

+

Upstream

4.66

3

64

PAX5

Paired box 5

Chr9: 36828530 to 37024476

-

Intragenic

4.56

3

65

LBX1

Ladybird homeobox 1

Chr10: 102976722 to 102978707

-

Downstream

4.53

3

66

EGFR

Epidermal growth factor receptor

Chr7: 55054218 to 55242525

+

Upstream

4.45

3

67

ODZ3

Odz, odd Oz/ten-m homolog 3

Chr4: 183302134 to 183508463

+

Upstream

4.40

3

68

TM7SF4

Transmembrane 7 superfamily member 4

Chr8: 105421230 to 105438092

+

Upstream

4.22

3

69

ZNF311

Zinc finger protein 311

Chr6: 29070573 to 29081014

-

Downstream

4.09

3

70

PCDH19

Protocadherin 19

Chrx: 99433297 to 99551927

-

Intragenic

4.06

3

71

FLJ45983

Hypothetical gene

Chr10: 8132418 to 8135453

-

Upstream

3.84

3

72

AX747375

Hypothetical gene

Chr19: 41955897 to 41958529

+

Downstream

3.83

3

73

PPP2R2C

Protein phosphatase 2 regulatory subunit B, gamma isoform

Chr4: 6373205 to 6525227

-

Intragenic

3.58

3

74

EMX1

Empty spiracles homeobox 1

Chr2: 72998111 to 73015528

+

Intragenic

3.57

3

75

PRKCSH

Protein kinase C substrate 80K-H

Chr19: 11407268 to 11422782

+

Upstream

3.56

3

76

FZD1

Frizzled homolog 1

Chr7: 90731718 to 90736068

+

Intragenic

3.52

3

77

LBX2

Ladybird homeobox 2

Chr2: 74578151 to 74583951

-

Upstream

3.42

3

78

HOXC13

Homeobox C13

Chr12: 52618842 to 52626595

+

Intragenic

3.40

3

79

MT1A

Metallothionein 1A

Chr16: 55230078 to 55231500

+

Upstream

3.36

3

80

DLX5

Distal-less homeobox 5

Chr7: 96487637 to 96492079

-

Intragenic

3.06

3

81

LDOC1L

Leucine zipper, downregulated in cancer 1-like

Chr22: 43267113 to 43272669

-

Intragenic

2.30

3

  1. aFor definitions of upstream, intragenic, and downstream, see Materials and methods (Microarray data analysis). The transcripts that have either upstream, intragenic or downstream peaks in at least three out of six DCIS samples were selected as positive targets. *Gene targets verified by combined bisulfite restriction analysis assays or bisulfite sequencing. bAverage of the log2 ratios assigned to positive peaks across the six DCIS. cHomeobox genes are indicated in bold.