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. Author manuscript; available in PMC: 2010 Aug 30.
Published in final edited form as: Genes Chromosomes Cancer. 2009 Jan;48(1):83–97. doi: 10.1002/gcc.20620

Comparative Epigenomics of Human and Mouse Mammary Tumors

Berna Demircan 1, Lisa M Dyer 1, Mallory Gerace 1, Edward K Lobenhofer 2, Keith D Robertson 1, Kevin D Brown 1
PMCID: PMC2929596  NIHMSID: NIHMS76508  PMID: 18836996

Abstract

Gene silencing by aberrant epigenetic chromatin alteration is a well-recognized event contributing to tumorigenesis. While genetically engineered tumor-prone mouse models have proven a powerful tool in understanding many aspects of carcinogenesis, to date few studies have focused on epigenetic alterations in mouse tumors. To uncover epigenetically silenced tumor suppressor genes (TSGs) in mouse mammary tumor cells, we conducted initial genome-wide screening by combining the treatment of cultured cells with the DNA demethylating drug 5-aza-2′-deoxycytidine (5-azadC) and the histone deacetylase inhibitor trichostatin A (TSA) with expression microarray. By conducting this initial screen on EMT6 cells and applying protein function and genomic structure criteria to genes identified as upregulated in response to 5-azadC/TSA, we were able to identify 2 characterized breast cancer TSGs (Timp3 and Rprm) and 4 putative TSGs (Atp1B2, Dusp2, FoxJ1 and Smpd3) silenced in this line. By testing a panel of ten mouse mammary tumor lines, we determined that each of these genes is commonly hypermethylated, albeit with varying frequency. Furthermore, by examining a panel of human breast tumor lines and primary tumors we observed that the human orthologs of ATP1B2, FOXJ1 and SMPD3 are aberrantly hypermethylated in the human disease while DUSP2 was not hypermethylated in primary breast tumors. Finally, we examined hypermethylation of several genes targeted for epigenetic silencing in human breast tumors in our panel of ten mouse mammary tumor lines. We observed that the orthologs of Cdh1, RarB, Gstp1, RassF1 genes were hypermethylated, while neither Dapk1 nor Wif1 were aberrantly methylated in this panel of mouse tumor lines. From this study, we conclude that there is significant, but not absolute, overlap in the epigenome of human and mouse mammary tumors.

INTRODUCTION

Tumorigenesis is a multi-step process stemming from increased oncogene activity in concert with constrained tumor suppressor gene (TSG) activity (Weinberg 1996). Such disease-producing changes in cellular phenotype result from dysregulation of gene expression and/or protein function attributable to genetic and/or epigenetic changes within the genome.

Several well-studied TSGs and other growth regulatory genes undergo de novo hypermethylation and consequential transcriptional silencing in human cancer cells. Aberrant cytosine methylation leading to gene silencing occurs at discreet, clustered 5′-CG-3′ dinucleotides (referred to as CpG islands) within the genome (Bird 1992). This cancer-associated CpG methylation is part of a complex set of epigenetic events that transform chromatin from a transcriptionally-active to an inactive state (Wolffe 2001) and epigenetic gene silencing is now widely recognized as either a causative or correlative event in tumor development (Jones and Baylin 2002; Robertson 2005) and is now widely regarded as one of the ‘hits’ in the Knudsen hypothesis leading to TSG inactivation (Jones and Laird 1999).

In recent years, numerous laboratories have developed tumor-prone mouse models owing to engineered over-expression of oncogenes or lost TSG expression. Clearly, these models have provided a powerful tool in studying tumorigenesis and unique insight into the molecular mechanisms that give rise to cancer in humans. For example, BRCA1, a TSG associated with breast and ovarian tumors in humans (Bishop 1999; Struewing et al., 1997), when conditionally deleted within the mammary epithelium of p53 heterozygous mice greatly increases mammary tumor incidence (Xu et al., 1999). This observation lends strong evidence as to the role BRCA1 plays in breast tumor suppression.

What remains less explored is the role that epigenetics plays during tumorigenesis in mice. While it seems straightforward to presume that gene silencing occurs in a parallel fashion in both human and mouse tumors, mice have not been widely utilized as a platform for either the discovery of epigenetically silenced genes or to study the role that gene silencing plays in the process of tumorigenesis. In one of the few studies to utilize a tumor-prone model system to dissect epigenetic events during disease progression, Yu et al (2005a) used Restriction Landmark Genomic Scanning (RLGS) to investigate aberrant DNA methylation events in a mouse model of T/natural killer acute lymphoblastic leukemia. These investigators observed nonrandom, aberrant DNA methylation occurring during tumorgenesis but was not present in benign preleukemic cells. Further, the inhibitor of DNA binding 4 (Idb4) gene was identified as a putative TSG silenced via promoter hypermethylation in mouse, and this gene was found epigenetically silenced in human leukemia as well. Similarly, Costello and colleagues found that SLC5A8, which encodes a sodium monocarboxylate cotransporter, was epigenetically silenced in both mouse and human brain tumors (Hong et al., 2005). These studies underscore the high potential of mice as a model system to understand the epigenetic basis of cancer and, potentially, other diseases as well.

A list of well-characterized TSG including BRCA1, CDKN2A, SFN, CDH1, and CST6 are targets for epigenetic silencing in breast cancer (Dobrovic and Simpfendorfer 1997; Esteller et al., 2000; Ferguson et al., 2000; Nass et al., 2000; Holst et al., 2003; Ai et al., 2006). However, at present, we are unaware of any genes that have been identified as targets for epigenetic silencing in mouse mammary tumors. To initially address this question, we conducted a screen for epigenetically silenced genes in a mouse mammary tumor line using an approach termed “pharmacological unmasking” (Esteller 2007). This often used approach examines changes in gene expression in response to the global DNA demethylating drug 5-aza-2′-deoxycytidine (5-azadC) and the histone deacetylase (HDAC) inhibitor trichostatin A (TSA) via microarray. Our screen was conducted using a mouse mammary cell line and identified ~1,000 genes that were upregulated ≥ 5-fold in response to 5-azadC/TSA in this line. Based on gene function and genomic architecture in mouse and man, several candidates identified in the initial screen were selected for further study.

MATERIALS and METHODS

Cell Culture and Drug Treatment

Mouse mammary tumor lines 410.4, 410 and 66.1 were obtained from Dr. A. Fulton (University of Maryland, Baltimore, MD), D2F2 and 4T1 from Dr. F. Miller (Karamanos Cancer Foundation, Detroit, MI), LM2 and LM3 from Dr. E. Bal de Kier Joffé (Universidad de Buenos Aires, Argentina) and D1K2 and NeuT from Dr. B. Law (University of Florida, Gainesville, FL). The mouse mammary tumor line EMT6 was purchased from American Type Culture Collection (ATCC, Manassas, VA).

LM2 and LM3 were grown in Eagle’s MEM with 2 mM L-glutamine, 1.5g/L sodium bicarbonate, 0.1 mmol/L nonessential amino acids, and 1.0 mmol/L sodium pyruvate. 4T1 were cultured in RPMI 1640 with 2 mmol/L L-glutamine, 1.5 g/L sodium bicarbonate, 4.5 g/L glucose, 10 mmol/L HEPES, and 1.0 mmol/L sodium pyruvate. 410, 410.4, 66.1, NeuT, D2F2, and D1K2 were maintained in DMEM with 2mM L-glutamine, 0.1 mmol/L nonessential amino acids. EMT6 was cultured in Waymouth’s MB 752/1 medium with 2mM L-glutamine. All cell lines were maintained in medium supplemented with 10% (v/v) heat-inactivated fetal bovine serum (Invitrogen, Carlsbad, CA) at 37°C in 5% CO2 humidified atmosphere. Human breast tumor cell lines were cultured as previously outlined (Ai et al., 2006).

Where indicated, cell lines were treated with 5 μM 5-azadC for 5 days (fresh drug was added every 24 hours) followed by a 24-hour treatment with 100 nM TSA. All drugs were purchased from Sigma.

Human Breast Tumor Specimens

Fifteen fresh-frozen archival breast tumors were obtained from the University of Florida Shands Cancer Center Molecular Tissue Bank. All specimens and pertinent patient information were treated in accordance with policies of the institutional review board (IRB) of the University of Florida Health Sciences Center. Tumors analyzed in this study were examined by a surgical pathologist and identified as invasive breast adenocarcinoma (stage II or III). Where indicated, matched normal breast tissue was obtained from disease-free surgical margin.

Microarray Analysis

Total RNA was extracted from EMT6 cells either untreated or treated with the global DNA demethylating agent 5-aza-2′-deoxycytidine (5-azadC) and the histone deacetylase inhibitor (HDAC) TSA. The quantity and purity of the extracted RNA was evaluated using a Nanodrop ND-100 spectrophometer (Nanodrop Technologies, Wilmington, DE) and its integrity measured using an Agilent Bioanalyzer. Five hundred nanograms (500 ng) of total RNA was amplified and labeled with a fluorescent dye (Cy3) using a Low RNA Input Linear Amplification Labeling kit (Agilent Technologies, Palo Alto, CA). Abundance and quality fluorescently labeled cRNA was also assessed using a Nanodrop ND-1000 spectrophotometer and an Agilent Bioanalyzer. The labeled cRNA (1.65 μg) was hybridized to an Agilent Mouse Whole Genome Oligo Microarray (Agilent Technologies, Inc., Palo Alto, CA) for 17 hrs prior to washing and scanning. Data was extracted from scanned images using Agilent’s Feature Extraction Software.

Gene Expression Analysis by RT-PCR

RT-PCR was carried out according to established protocols (Ai et al., 2006). Briefly, untreated and 5-azadC/TSA treated mouse mammary tumor cells were homogenized in Trizol (Invitrogen), and total RNA purified according to the manufacturer’s instructions. First-strand cDNA synthesis was carried out using SuperScript III reverse transcriptase (Invitrogen). Subsequently, this cDNA was used in PCR reactions using Timp3, Rprm, Smpd3, Atp1B2, Dusp2 and FoxJ1-specific oligonucleotide primers as listed in Supplemental Table I. Mouse glyceraldehyde-3-phosphate dehydrogenase (Gapdh) served as a control to assure equivalent RNA in each reaction. Following PCR, reactions were analyzed by electrophoresis on 10% polyacrylamide gels followed by staining with ethidium bromide.

Extraction and Sodium Bisulfite Modification of Genomic DNA

Genomic DNA (gDNA) was isolated from human breast tumor tissue samples and cultured human and mouse cell lines using a Qiagen Blood and Cell Culture DNA kit (Qiagen, Inc., Valencia, VA) and stored at −20°C before use. DNA was modified with EZ DNA Methylation kit (ZYMO Research Co., Orange, CA) according to the manufacturer’s instructions. Briefly, ~1.0 μg denatured gDNA was treated with sodium bisulfite at 50°C for ~16 hours in the dark. Following this, samples were applied to supplied columns, desulfonation conducted, and DNA was subsequently eluted in a volume of 20 μL with supplied elution buffer. One μL of bisulfite-converted gDNA solution was generally used in subsequent PCR reactions.

Methylation Specific-PCR

Primers used for Methylation Specific-PCR (MSP) analyses are listed in Supplemental Table II. MSP reactions were conducted using Qiagen HotStarTaq DNA polymerase and supplied 1X PCR buffer supplemented with 0.1 mmol/L dNTPs, 2.5 mmol/L MgCl2, and 0.5 μmol/L each of forward and reverse primers and bisulfite-converted gDNA. Following thermocycling, PCR products were loaded 10% polyacrylamide gels and visualized using ethidium bromide.

Human placental gDNA (Biochain Institute, Hayward, CA) and mouse gDNA isolated from tail snips of wildtype C57BL/6 mice were used in primer validation experiments of human and mouse MSP, respectively and where indicated DNA was methylated in vitro using SssI methylase (NEB, Ipswich, MA) as outlined previously (Vo et al., 2004).

Pyrosequencing

PCR reactions were conducted on bisulfite-converted gDNA as described above. Supplemental Table III summarizes pyrosequencing primers and sequence analyzed in each gene. Following PCR amplification using a biotinylated reverse primer, 5–20 μL of PCR product was mixed with 2 μL of streptavidin-coated sepharose beads (GE Heathcare Bio-Sciences AB, Uppsala, Sweden), 40ul of binding buffer (10 mM Tris-HCl, 2 M NaCl, 1mM EDTA, 0.1% Tween 20, pH 7.6), and brought up to 80 μL total volume with dH2O. This mixture was immobilized on streptavidin beads by shaking constantly at 1400 rpm for 10 minutes at RT. The beads were subsequently harvested using a vacuum preparation workstation and appropriate sequencing primer added to a final concentration of 330 nM. The sequencing primers were subsequently annealed to the template by incubating the samples at 80°C for 2 min and cooled at RT for 10 min prior to analysis. Pyrosequencing was conducted using a PyroMark MD system (Biotage AP, Uppsala, Sweden), and sequence analysis conducted using supplied software.

RESULTS

The focus of this investigation was to compare mouse and human mammary cancer for potential overlap in tumor epigenotype across these two species. Our screening strategy entailed utilizing the mouse mammary tumor line EMT6 in a pharmacological unmasking screen (Esteller 2007). Briefly, EMT6 cells were treated with the global DNA demethylating agent 5-aza-2′-deoxycytidine (5-azadC) and the histone deacetylase (HDAC) inhibitor trichostatin A (TSA) to de-repress epigenetically silenced genes. After drug treatment, which was found to have minimal toxicity in EMT6 cells by MTT assay (data not shown), total RNA was harvested from two replicate cultures of drug treated and untreated EMT6 cells and used to probe Agilent Mouse Whole Genome microarrays.

Of the ~44,000 transcripts measured on these microarrays, an average of 54% were detected as significantly above background levels in each of the hybridizations (range: 51 to 57%). The intensity values from the replicate samples for both the treated and untreated cultures had Pearson correlation values greater than 0.97, suggesting that the reproducibility of the results was very high. Reproducibility of the within-group samples, along with the differences between the two groups, were also apparent when the z-score transformed intensity values for the 2,252 transcripts found to be differentially expressed (984 upregulated, 1,268 downregulated) using an absolute fold change criterion of 5-fold and a log ratio p-value of 0.001 were compared (Fig 1).

Figure 1. Hierarchical clustering of the differentially expressed transcripts in all of the biological replicate samples.

Figure 1

Two-way hierarchical clustering using Ward’s Minimum Variance as the heuristic criteria and Euclidean distance as the similarity metric was performed on all 4 samples using the z-score transformed expression values for the 2,339 transcripts that were identified as differentially expressed (fold-change > 5, log ratio p-value < 0.001) when the two groups were compared to each other (984 genes displayed increased expression, 1,355 displayed decreased expression in response to 5-azadC/TSA). The degree of relatedness between each sample is represented by the dendrogram (hierarchical tree) presented in this figure, wherein the height of each branch represents the distance between the 2 objects being connected. The relationship between the color intensity and the expression level is illustrated by the color bar.

Identification of candidate epigenetically silenced genes in the EMT6 mouse mammary tumor cell line

Following the initial screen by expression microarray, we established a series of bioinformatic criteria to narrow our search for epigenetically silenced tumor suppressor genes (TSGs) in the EMT6 cell line. From the entire list of genes measured by the microarray, we first excluded genes that showed a moderate change in expression in response to 5-azadC/TSA. Therefore, we required that a candidate gene demonstrate at least a 5-fold increase with a log ratio p-value of 0.001 to be considered further. This narrowed our focus to 984 genes of interest (Fig 2). (A listing of all 984 genes identified is given in Supplemental Table IV.)

Figure 2.

Figure 2

Flow chart summarizing various experimental steps used following initial microarray screen for epigenetically silenced genes in the EMT6 cell line.

Next, we used protein function and genomic structure criteria to further define potential TSGs that are targets for epigenetic silencing in EMT6 cells. We were principally interested in studying genes that encode proteins with an established tumor suppressor activity(ies), or a function consistent with tumor suppressive activity. Specifically, we included in our study gene products where previously published studies support a gene function that either antagonizes cell growth, activates apoptosis or suppresses anti-apoptotic processes, restrains cellular invasiveness, or inhibits angiogenic processes.

We next excluded from the list of putative/characterized TSGs those genes that do not possess a canonical CpG island within 5′ flanking region in both the human and mouse orthologs using the UC Santa Cruz genome browser (http://genome.ucsc.edu) to investigate genomic architecture of various candidate genes. We used the algorithm within the UCSC genome browser that scores the presence of CpG islands in genomic DNA using criteria originally established by Gardiner-Garden and Frommer (Gardiner-Garden and Frommer 1987). Our rationale for establishing these criteria is that genes which possess both features (i.e., putative TSG activity and the presence of a CpG island that overlaps the translational start site of the gene) are likely to identify high-value candidate targets for epigenetic silencing. Of the 984 genes initially identified as being upregulated ≥5-fold in our expression microarray screen, this bioinformatic investigation identified 37 genes meeting the outlined criteria (see Supplemental Table V). Of this, 26 genes were further investigated in this study.

Our next test was to confirm 5-azadC/TSA-induced upregulation of gene expression by RT-PCR. This revealed 12 genes where the gene upregulation detected by microarray was confirmed by RT-PCR and 14 genes whose upregulation could not be independently confirmed using an independent set of RNA samples isolated from drug treated/untreated EMT6 cells. Methylation-Specific PCR (MSP), a common methodology to examine DNA methylation (Herman et al., 1996), was subsequently used to judge methylation within the CpG islands associated with remaining 12 candidate genes. This analysis revealed that six genes exhibited no methylation as judged by MSP. This may indicate that the 5-azadC/TSA-induced increase in expression of these genes is indirect in nature and/or that silencing of these genes is controlled by chromatin modifications other than CpG methylation (e.g., transcriptionally-repressive histone modifications). The later of these possibilities is quite possible since our drug treatment regimen included the histone deacetylase inhibitor TSA. As a result of these efforts our focus narrowed to six genes: Tissue Inhibitor of Metalloproteinase 3 (Timp3), TP53-dependent G2 Arrest Mediator Candidate also termed Reprimo (Rprm), Sphingomyelin Phosphodiesterase 3 Neutral (Smpd3), ATPase, Na+/K+ Transporting, Beta 2 Polypeptide (Atp1B2), Dual-Specificity Phosphatase 2 (Dusp2), and ForkheadboxJ1 (FoxJ1).

Silencing of Timp3 and Rprm in mouse mammary tumor cell lines

Timp3 and Rprm are well-characterized TSGs silenced in human breast cancer (Bachman et al., 1999; Takahashi et al., 2005). To test how commonly these genes are epigenetically silenced in mouse mammary tumor lines, we acquired 9 distinct cultured mouse mammary tumor lines (designated 410.4, 410, LM3, LM2, D1K2, NeuT, D2F2, 66.1 and 4T1), in addition to the EMT6 cell line. As outlined above, RT-PCR was used to confirm increased expression in response to 5-azadC/TSA; however, due to high-levels of toxicity to this drug cocktail, we found that only four of these cell lines (EMT6, 410.4, 410, and LM3) showed sufficiently low toxicity (>75% viability 48 hrs after 5 μM 5-azadC treatment) to be appropriate subjects for this experiment. These lines were cultured in the presence or absence of 5-azadC and TSA, total RNA harvested and subsequently cDNA synthesized. Results of this RT-PCR experiment clearly indicated that the abundance of Timp3 and Rprm transcripts increased substantially following treatment of each line with 5-azadC/TSA (Fig 3A). Parallel reactions using primers specific for the mouse form of Gapdh confirmed equivalent RNA abundance in each PCR reaction.

Figure 3. Analysis of Timp3 and Rprm silencing in cultured mouse mammary tumor cell lines.

Figure 3

A Indicated mouse mammary tumor lines were either treated or not with a combination of 5-azadC and TSA. Subsequently, total RNA was harvested, cDNA synthesis performed, and PCR reactions conducted using primers specific for the Timp3 (top) or Rprm (middle) genes. Also, PCR reactions using primers specific for the Gapdh gene (bottom) were conducted to assure equivalent RNA abundance in each reaction. B. MSP primers were designed specific to the mouse Timp3 (top) and Rprm (bottom) genes. Primers designed to amplify methylated (M) or unmethylated (U) regions of the CpG island associated with each gene were tested in PCR reactions using mouse genomic DNA either incubated in the presence or absence of SssI methylase prior to bisulfite conversion. C. Genomic DNA harvested from mouse mammary tumor lines EMT6, 410.4, 410, LM3, LM2, NeuT, D2F2, 66.1, 4T1, D1K2 were bisulfite converted and used in MSP reactions with methylated-specific (M) or unmethylated-specific (U) primers. D. Pyrosequencing analysis was conducted on the Timp3 gene using bisulfite-modified genomic DNA from EMT6 and LM3 tumor lines, and mouse genomic DNA as a control. Shown are obtained pyrograms, measured methylation at CpG dinucleotides within the gene region analyzed, and the average CpG methylation within this region of the Timp3 gene. E. Pyrosequencing analysis of the Rprm gene was conducted on EMT6 and LM3 lines and mouse genomic DNA as outlined in D.

We next investigated the methylation status of Timp3 and Rprm in our full panel of mouse mammary tumor line by designing MSP primers using a publically available on-line primer design program (www.mspprimer.org/cgi-mspprimer/design.cgi). All primer sets designed for the analysis for DNA methylation status (both MSP and pyrosequencing analysis) were designed to assay DNA methylation at, or in close proximity to, the transcriptional start site (TSS) of candidate genes since this region is most likely to have undergone dense methylation during the process of epigenetic silencing (Bird 2002; Jones and Baylin 2002).

Primers were designed specific for both the mouse Timp3 and Rprm genes and validated using mouse genomic DNA extracted from tail clippings of C57BL/6 mice. This mouse genomic DNA was either methylated in vitro with SssI methylase or not methylated prior to bisulfite conversion. MSP reactions conducted using this control DNA indicated that our designed primers discriminate between the Timp3 and Rprm genes in either a methylated or unmethylated state (Fig 3B). Subsequently, these MSP primer sets were used to probe the methylation status of the Timp3 and Rprm gene in our panel of 10 mouse mammary tumor cell lines (Fig 3C). With the exception of D2F2, a methylated-specific (M) amplicon was detected when the Timp3 gene was analyzed using bisulfite-converted genomic DNA isolated from each cell line. While amplification of both unmethylated and methylated-specific MSP products is a common finding in MSP analysis (Herman et al., 1996), potentially attributable to heterogeneous promoter methylation within the tumor cell population, this result indicates the presence of methylated CpG within the Timp3 gene in at least a subset of cells of each line except D2F2 where the Timp3 promoter appears to be unmethylated.

As an independent means of quantitatively analyzing DNA methylation, we conducted pyrosequencing on mouse Timp3 and Rprm genes in the EMT6 and LM3 cell lines. Pyrosequencing, like MSP, analyzes bisulfite-converted genomic DNA as a template for PCR; however, pyrosequencing is conducted on PCR products amplified using primers that do not contain internal CpG dinucleotides. Thus, amplification occurs in a manner that is unbiased by DNA methylation status. Subsequently, methylation at distinct CpGs can be determined quantitatively in a pool of genomic DNA (Colella et al., 2003; Tost et al., 2003).

The primer design developed for the mouse Timp3 gene allowed for analysis of 9 CpG dinucleotides within a ~45 bp portion of the CpG island associated with this gene. Pyrosequencing analysis, consistent with MSP results, indicated that Timp3 was methylated in both EMT6 and LM3 cell lines. Moreover, all the CpG dinucleotides investigated using this approach displayed methylation suggesting a high level of methylation density, although CpG methylation was not uniform over the region interrogated. The average CpG methylation measured in this region was 79% in EMT6 and 76% in LM3 (Fig 3D). The pyrosequencing protocol developed for Rprm analyzed 6 CpG dinucleotides over a ~35 bp region of the CpG island within this gene. Similar to Timp3 analysis, all CpG within this region of the Rprm gene exhibit significant methylation with average CpG methylation of 84% and 86% in EMT6 and LM3 cells, respectively (Fig 3E).

Pyrosequencing analysis was also conducted on control (mouse tail) DNA samples (Fig 3D,E). For both Timp3 and Rprm genes, average CpG methylation measured was 7%, indicating very low levels of DNA methylation in this control DNA sample. This result indicates that methylation of the CpG islands associated with the Timp3 and Rprm genes in the mouse genome is a tumor-associated event. Moreover, when our DNA methylation analysis is combined with the results of our RT-PCR experiments using the DNA demethylating agent 5-azadC, we conclude that the Timp3 and Rprm genes are targeted for epigenetic silencing in mouse mammary tumor lines through aberrant gene methylation. Considering most of the 10 distinct cell lines display this feature, silencing of these genes is apparently a common event in cultured mouse mammary tumor cells. Since both Timp3 and Rprm are also targets for epigenetic silencing in human breast tumor line and primary tumors, these results reinforce the validity of our expression microarray screen for epigenetically silenced genes in mouse mammary tumor lines and establish a point of epigenetic overlap between mouse and human mammary gland tumors.

The Atp1B2, Smpd3, Dusp2 and FoxJ1 genes are targets for epigenetic silencing in mouse mammary tumor cell lines

In addition to Timp3 and Rprm, our screen identified four genes (Atp1B2, Smpd3, Dusp2 and FoxJ1) that were strong candidates for epigenetic silencing in the EMT6 line. RT-PCR was conducted on EMT6, 410.4, 410, and LM3 cells cultured in the presence or absence of 5-azadC/TSA using primers designed specifically for the mouse orthologs of these genes. RT-PCR analysis of the Atp1B2 transcript in EMT6 cells showed a clear upregulation of this transcript in response to 5-azadC/TSA (Fig 4A). Conversely, 410.4 and LM3 cells displayed no detectable upregulation of Atp1B2 expression and analysis of 410 cells showed a slight (~2-fold) increase in Atp1B2 expression following 5-azadC/TSA treatment. RT-PCR results showed strong upregulation of Dusp2 expression in 5-azadC/TSA treated EMT6, 410 and 410.4 cells while LM3 cells showed no increased expression of this gene in drug treated LM3 cells. Smpd3 showed marked increases in gene expression in each cell line in response to 5-azadC/TSA. Similarly, FoxJ1 showed clear upregulation in drug treated EMT6 and LM3 cells while expression was more modestly increased in drug treated 410 and 410.4 cells.

Figure 4. Analysis of Atp1B2, Dusp2, Smpd3 and FoxJ1 silencing in mouse mammary tumor lines.

Figure 4

A RT-PCR was conducted on cDNA synthesized from indicated mouse mammary tumor lines treated and untreated with 5-azadC/TSA using primers specific for each indicated gene. B. MSP primers specific for each gene were designed and subsequently validated using bisulfite-modified mouse genomic DNA either methylated or not prior to bisulfite conversion. C. MSP analysis of these four genes using DNA harvested from the indicated mouse mammary cell lines was conducted.

Based on the positive RT-PCR results outlined above, we designed MSP primers for the mouse forms of the Atp1B2, Smpd3, Dusp2 and FoxJ1 genes. As shown (Fig 4B), validation experiments using in vitro methylated/unmethylated mouse genomic DNA confirmed that MSP primer sets were capable of determining the methylation state of each gene. In EMT6 and 410 cells, MSP analysis of the Atp1B2 gene indicated abundant M-specific amplicon indicating that this gene is hypermethylated in these cell lines. This result is consistent with the increased expression of this gene observed in RT-PCR experiments. In 410.4 cells, slight amplification of the M-specific band was noted and no M-specific amplicon was detected when genomic DNA isolated from LM3 cells was assayed, indicating that methylation of this gene is not a prominent feature in this cell line. MSP analysis of the other lines in our panel of mouse mammary tumor cells indicated that only 66.1 cells display prominent amplification of the M-specific product from the Atp1B2 gene. Reduced amplification of the M-specific product was observed in 4T1 and D2F2 cells while no M-specific product was obtained from the other 3 lines (LM2, D1K2 and NeuT) in our panel.

MSP analysis of the Dusp2 gene in EMT6, 410.4 and 410 cells indicated strong amplification of the M-specific product (Fig 4C). When combined with the clear upregulation of the Dusp2 transcript when each of these lines were treated with 5-azadC/TSA, this result indicates that this gene is epigenetically silenced in these lines. Conversely, no aberrant methylation of the Dusp2 gene in LM3 cells was observed. In the remaining six lines in our panel, methylation of the Dusp2 gene was observed in D1K2, NeuT and 66.1 cells and no M-specific amplicon was observed when LM2, D2F2 and 4T1 cells were assayed by MSP.

When the Smpd3 gene was analyzed by MSP, methylation of this gene was observed in all of the cell lines in our panel with the exception of NeuT and D2F2 (Fig 4C). Since we observed upregulation of the Smpd3 in each of the four cell lines treated with 5-azadC/TSA, MSP results are consistent with epigenetic silencing of the Smpd3 gene in cultured mammary tumor lines. Similarly, all ten cultured mammary tumor lines displayed methylation of the FoxJ1 gene as judged by MSP. Again, this is consistent with our observation of increased FoxJ1 expression in 5-azadC/TSA treated cells and indicates that the FoxJ1 is commonly targeted for epigenetic silencing in cultured mouse mammary tumor lines. In sum, these lines of experimentation indicate that the Atp1B2, Dusp2, Smpd3 and FoxJ1 genes are epigenetically silenced in mouse mammary tumor lines; however, the occurrence of these aberrant events is variable within this panel of cultured lines.

Analysis of ATP1B2, DUSP2, SMPD3 and FOXJ1 hypermethylation in human breast cancer lines and primary breast tumors

The ATP1B2, DUSP2, SMPD3 and FOXJ1 genes have not previously been characterized as targets for epigenetic silencing in human breast tumors. Thus, we examined the methylation status of each of these genes in a panel of eight cultured human breast cancer cell lines by designing MSP primer sets for the human ortholog of each gene. Validation experiments using in vitro methylated/unmethylated human placental genomic DNA indicated that each of these primer sets can discriminate between methylated and unmethylated forms of these four genes (Fig 5A). MSP analysis of the ATP1B2 gene in our panel of human breast tumor lines indicated robust amplification of the methylation-specific product in MCF-7, MDA-MB-435 and BT-474 cells while MDA-MB-468 cells show moderate accumulation of the M amplicon and low level amplification of the M product was observed in MDA-MB-231. Other lines in our panel (T-47D, BT-549 and SKBR3) displayed no evidence of ATP1B2 methylation. MSP analysis of the SMPD3 gene determined that MDA-MB-231, MDA-MB-435 and BT-474 cells display methylation of this gene, while the other cell lines show no evidence of SMPD3 methylation. Similarly, DUSP2 was found to be methylated in MCF7, MDA-MB-231 and BT-549 cells but not in T-47D, MDA-MB-468, SKBR3, MDA-MB-435 and BT-474 cells. Finally, MSP analysis of the FOXJ1 gene indicates strong amplification of the M-specific amplicon in MDA-MB-231, MDA-MB-435 and BT-474 cells. Weak amplification of the M-specific FOXJ1 amplicon was observed in MCF-7, MDA-MB-468 and BT-549. No amplification of the FOXJ1 M-specific product was observed in either T-47D or SKBR3 cells.

Figure 5. Analysis of ATP1B2, DUSP2, SMPD3 and FOXJ1 methylation in human breast tumor cell lines.

Figure 5

A MSP primers were designed to the human ortholog of each indicated gene and validated using human placental DNA that was incubated in the presence or absence of SssI methylase prior to bisulfite conversion. B. MSP analysis of indicated genes was conducted on indicated human breast tumor cell lines. C. Pyrosequencing analysis of the ATP1B2 gene was conducted using genomic DNA harvested from MCF-7, BT-474, and human placenta (control). D. Pyrosequencing analysis of the SMPD3 gene in MDA-MB-231, BT-474 and control DNA. E. Pyrosequencing analysis of the DUSP2 gene in MDA-MB-231, BT-549 and control DNA. F. Pyrosequencing analysis of the FOXJ1 gene in MDA-MB-231, BT-474 and control DNA samples.

To confirm the MSP results obtained when the methylation status of the ATP1B2, DUSP2, SMPD3 and FOXJ1 genes was scored by MSP, we conducted pyrosequencing. The pyrosequencing assay developed for the human ATP1B2 gene scores methylation of seven CpG dinucleotides within a ~30 nt region of the CpG island within the 5′ end of the human ATP1B2 gene. When genomic DNA harvested from MCF-7 cells was analyzed by pyrosequencing, we observed low (3%) to moderate (29%) levels of CpG methylation within this cell line in the ATP1B2 gene (Fig 5C). Overall, average CpG methylation measured within the ATP1B2 gene was 17% in this cell line. In BT-474 cells, we observed markedly higher methylation within the ATP1B2 gene with average CpG methylation of 34%. Pyrosequencing of control (human placental) DNA showed a low level of CpG methylation (4.6%) in this gene.

Pyrosequencing analysis of the SMPD3 gene showed high average levels of CpG methylation (56% and 71%) within the SMPD3 gene in MDA-MB-231 and BT-474 cells, respectively (Fig 5D). Conversely, low levels of CpG methylation (0.8%) of the SMPD3 gene was measured in control DNA. Consistent with MSP results, we observed methylation of the CpG island associated with the human DUSP2 gene in MDA-MB-231 (88%) and BT-549 (44%) cells with low (5.5%) CpG methylation measured in control DNA (Fig 5E). We also observed CpG methylation within the human FOXJ1 gene in MDA-MB-231 (40%) and BT-474 (63%) cells and control DNA displayed low (5.5%) CpG methylation within FOXJ1 (Fig 5F). Taken together, both MSP and pyrosequencing analyses indicate that the ATP1B2, DUSP2, SMPD3 and FOXJ1 are aberrantly methylated, albeit at differing frequencies, in cultured human breast tumor lines.

Next, we conducted MSP analysis to examine ATP1B2, DUSP2, SMPD3 and FOXJ1 gene methylation in primary breast tumor/normal mammary tissue samples. Fifteen frozen human breast tumors were obtained and patient-matched disease-free breast tissue were obtained with two of these tumor samples (Tumor72 and Tumor75). Genomic DNA was isolated, bisulfite modified, and used in MSP reactions with primers specific for the ATP1B2, DUSP2, SMPD3 and FOXJ1 genes. In all normal breast tissue samples, we observed exclusive amplification of the unmethylated-specific (U) amplicon, indicating that these genes are not methylated in normal mammary tissue (Fig 6A). When patient-matched breast tumor samples were analyzed by MSP, we observed methylation of the SMPD3 gene in both Tumor72 and Tumor75. Further, methylation of the FOXJ1 gene was observed in Tumor72. No methylation of either DUSP2 or ATP1B2 was observed in either Tumor72 or Tumor75.

Figure 6. Analysis of ATP1B2, DUSP2, SMPD3 and FOXJ1 methylation in primary human breast tumors.

Figure 6

A MSP analysis of the indicated genes was conducted on human primary breast tumors and patient-matched normal breast tissue. B. Representative MSP analysis of six additional human primary breast tumors.

Figure 6B shows a representative subset of additional breast tumors analyzed by MSP. We observed methylation of the SMPD3 gene in Tumor 62, 66 and 70. Overall, we observed SMPD3 methylation in 60% of breast tumors investigated (9 out of 15). We observed methylation of the ATP1B2 gene in Tumor 61, 68 and 70 and, overall, detected a 20% rate (3/15) of ATP1B2 methylation in breast tumors. In contrast, FOXJ1 was found to be commonly methylated in human breast tumors; specifically, we observed that 11 out of 15 tumors surveyed (73%) displayed aberrant FOXJ1 methylation as scored by MSP. Taken together, MSP analysis suggests that promoter hypermethylation the ATP1B2, SMPD3 and FOXJ1 occur in primary human breast tumors with varying occurrence.

Despite the fact that we documented hypermethylation of DUSP2 in cultured human breast cancer lines, this gene was found to be unmethylated in all human breast tumors examined (Fig 6B). DUSP2 encodes a phosphatase (also known as PAC1) that dephosphorylates and inactivates mitogen-activated protein (MAP) kinases (Chu et al., 1996) and is a transcriptional target of p53 in signalling apoptosis (Yin et al., 2003). These properties make this gene an excellent candidate TSG and DUSP2 hypermethylation has been reported in prostate cancer (Yu et al., 2005b). However, we did not observe methylation of this gene in primary tumors leading us to conclude that epigenetic silencing of DUSP2 in human breast cancer is likely a rare event.

Analysis of methylation status of characterized targets for epigenetic silencing in human breast tumors in cultured mouse mammary tumor lines

Finally, we determined the methylation status of well-defined TSGs known to be targets for epigenetic silencing in human breast cancer in our panel of cultured mouse mammary tumor lines. We designed MSP primer sets to assay methylation within the CpG islands associated with the mouse orthologs of the Cdh1, RarB, Gstp1, Dapk1, RassF1 and Wif1 genes. Validation experiments using in vitro methylated/unmethylated mouse genomic DNA indicate that each of these primer sets can delineate gene methylation status (Fig 7A).

Figure 7. Analysis of genes characterized as targets for epigenetic silencing in human breast tumors in a panel of mouse mammary tumor lines.

Figure 7

A MSP primers were designed to the mouse orthologs of the Cdh1, Gstp1, RarB, Wif1, RassF1 and Dapk1 genes. Primer sets were validated using control mouse genomic DNA that was either methylated/unmethylated in vitro prior to bisulfite conversion. B. MSP analysis of indicated genes was conducted on bisulfite-converted DNA isolated from indicated mouse mammary tumor lines. gDNA was harvested from the cell lines and bisulfite-modified. These DNA samples were used in MSP analyses.

Our MSP analyses indicate that RassF1 and RarB were commonly methylated in our panel of mouse mammary tumor lines with 90% and 80% showing significant methylation, respectively (Fig 7B). In contrast, methylation of Cdh1 and Gstp1 were far less commonly encountered in this panel of cell lines with 30% and 10%, respectively, displaying methylation of these genes. In all mouse mammary cell lines, we observed exclusive amplification of the unmethylated-specific products for Dapk1 and Wif1 genes indicating that neither gene is methylated in this panel of mouse cell lines.

DISCUSSION

Comparative epigenomics of mouse and human mammary tumors

By coupling genome-wide screening with bioinformatics analyses and inspection of publically available mouse and human genome sequence, we were able to characterize 6 candidate genes as targets for epigenetic silencing in the EMT6 line. These genes are: Timp3, Rprm, Smpd3, Dusp2, Atp1B2, and FoxJ1. Of these, Timp3 and Rprm have already been characterized as TSGs and targets for epigenetic silencing in human breast and other tumor types (Esteller et al., 2001; Takahashi et al., 2005; Kim et al., 2006;). The fact that our initial screening identified these TSGs as silencing candidates in EMT6 underscores the successful nature of our initial screen. Moreover, since analysis of 10 distinct mouse mammary tumor lines shows that both Timp3 and Rprm are commonly silenced in cell lines of this type, we conclude that both human and mouse mammary tumors share silencing of these tumor suppressor genes in common.

A combination of RT-PCR and MSP confirmed that Smpd3, Dusp2, Atp1B2, and FoxJ1 are silenced in mouse mammary tumor lines. Our findings indicate that Atp1B2 is a target, although likely an uncommon one, for silencing in mouse mammary cell lines. The other 3 genes identified in our screen appear to be more commonly hypermethylated in mouse mammary tumor lines. We observed significant amplification of the M-specific amplicon specific for Dusp2 in 60% of the cell lines tested, Smpd3 in 80%, and FoxJ1 in 100%. When a panel of eight human breast tumor cell lines were assayed for methylation by pyrosequencing and/or MSP, we observed substantial methylation of the ATP1B2 gene in 50% (4/8) cell lines assayed. Also, SMPD3, DUSP2, and FOXJ1 methylation was detected in 38% (3/8) of the cell lines surveyed. Of note, methylation of each of these genes, with the exception of DUSP2, was observed in human primary breast tumors and MSP analysis of normal breast tissue showed methylation of these genes to be a disease-associated event. The fact that DUSP2 methylation was observed in cultured human cell lines but not primary tumors strongly suggests that silencing of this gene is likely a rare event in human breast tumors.

As part of our study we also examined the methylation status of 6 genes known to be targets of epigenetic silencing in human breast tumors in our panel of cultured mouse mammary tumor lines. In this investigation the mouse ortholog of the Cdh1, Gstp1, RarB, Wif1, RassF1, and Dapk1 genes were assayed by MSP. We observed, to various extents, that Cdh1, Gstp1, RarB, and RassF1 were hypermethylated in our panel of mouse tumor lines. However, we obtained no evidence that either the Wif1 or Dapk1 genes are aberrantly methylated in any of these lines.

When the results of our studies on various epiegentically silenced genes in human and mouse cultured mammary tumor cell lines and human primary breast tumors are considered collectively, we conclude that there is significant overlap in the mammary tumor cell epigenome between these two species. Of the twelve genes studied, only Wif1 and Dapk1 were found not be silenced in both species. While Dusp2 was found to be commonly hypermethylated in mouse cell lines, methylation of this gene was less commonly observed in cultured human lines and no evidence for hypermethylation was observed in primary human breast tumors. These results indicate that some genes targeted for epigenetic silencing in human breast tumors are also aberrantly methylated in mouse mammary tumor lines demonstrating considerable but not absolute overlap in the mammary tumor epigenome between these two species.

SMPD3, ATP1B2 and FOXJ1 as novel targets for epigenetic silencing in human breast cancer

Sphingomyelinases (SMases) belong to a multigene family of phosphodiesterases. Each of the three well-characterized SMases (SMPD1, SMPD2, and SMPD3) hydrolyze sphingomyelin to ceramide and phosphocholine. The acid sphingomyelinase SMPD1 is located in the endolysosomal compartment, while the neutral SMases (nSMases) SMPD2 and SMPD3 are located within the endoplasmic reticulum membranes and in the Golgi apparatus, respectively (Stoffel et al., 2007). nSMases are a major intracellular regulator of ceramide, a bioactive lipid.

Several lines of evidence support the view that the ability of nSMases contribute to proliferation of cancer cells and/or to their invasiveness. Of note, the SMPD3 locus is present, as well as six members of the cadherin family, with chromosome 16q22.1, and genetic alterations in this region have been implicated in the progression of various cancers, notably breast cancer (Chalmers et al., 2001; Driouch et al., 1997). Moreover, it has recently been shown that reduction of ceramide levels due to inhibition of nSMase in Hs578T breast adenocarcinoma cells overexpressing caveolin-1 results in a significant enhancement in Akt activation a well-characterized pro-growth, pro-survival mechanism (Wu et al., 2007). Ample evidence also supports a role for ceramide in the activation of apoptosis. For example, Luberto et al (2002) showed inhibition of TNF-α-induced cell death in MCF7 cells using a novel inhibitor of nSMase activity. Data obtained in this study indicates that ceramide induces conformational changes in Bax and activates mitochondrial phosphatase 2A, which, in turn, dephosphorylates Bcl-2. These two events clearly link ceramide and nSMases to the activation of cell death. Similarly, studies link ceramide to the regulation of cell cycle progression by inducing growth arrest by promoting dephosphorylation of the Rb tumor suppressor (Alesse et al., 1998).

In this report, we show that the SMPD3 gene in both mouse and man is a target for epigenetic silencing in mammary tumors. To our knowledge, this is the first report of the SMPD3 gene being targeted for silencing during tumorigenesis. Functionally, this aberrant event will result in decreased ceramide production in response to normal physiological cues promoting dysregulation of both cellular death and growth programs. Thus, this event has clear implications in the process of breast tumorigenesis.

The protein encoded by ATP1B2 is a member of the Na+/K+ ATPase beta chain protein family. Na+/K+ ATPases, which are composed of two subunits (a large catalytic subunit (alpha) and a smaller glycoprotein subunit (beta)), are integral membrane proteins responsible for establishing and maintaining the electrochemical gradients of Na+ and K+ ions across the plasma membrane. This process is essential for osmoregulation, for sodium-coupled transport of a variety of organic and inorganic molecules, and for electrical excitability of nerve and muscle (Dobretsov and Stimers 2005).

The adhesion molecule on glia (AMOG) protein was originally identified as a neural cell adhesion molecule and controls neuron–astrocyte adhesion as well as neural cell migration in vitro (Antonicek et al., 1987), was later found to be encoded by the ATP1B2 gene (Gloor et al., 1990). Others (van den Boom et al., 2006) recently showed that the ATP1B2 gene was targeted for epigenetic silencing via aberrant gene methylation in malignant glioma cells. This finding, supported by the observation that reduced ATP1B2 expression is associated with increased tumor grade (Senner et al., 2003), led these investigators to propose that ATP1B2 silencing could be a significant event in glioma development. Our results indicate that Atp1B2 is epigenetically silenced in a subset of cell lines in our panel of mouse mammary tumor lines and its human ortholog is subject to gene hypermethylation in both human breast tumor cell lines and primary tumors. Given the role that this gene product plays in controlling glial migration, these findings suggest that silencing of ATP1B2 may influence breast tumor cell migration as well. This compelling notion requires further investigation.

FOXJ1 is a member of the fork head gene family of transcription factors originally identified in Drosophila, and a large family of FOX genes are currently characterized in humans (Katoh 2004). In mammalian cells, FOXJ1 has been shown to suppress NFκB transcriptional activity in vitro and suppress inflammation in vivo (Lin et al., 2004). While NFκB is a key regulator of immune response, dysregulation of NFκB often drives deregulated growth, resistance to apoptosis, and increased metastatic potential (Lu and Stark 2004). Since FOXJ1 functions as an antagonist to NFκB activity it is reasonable to suggest that this molecule functions as a TSG, and others have commented that FOXJ1 is an excellent candidate for anti-cancer therapy (Chen 2004). While the role of FOXJ1 in cancer suppression remains to be explored, our finding that this gene is a common target for aberrant hypermethylation in both cultured cells and primary tumors strongly suggests that FOXJ1 silencing is potentially an important event in breast tumorigenesis.

In conclusion, our study has documented considerable, but not absolute, overlap exists between the epigenome of mouse and human mammary tumors. This finding supports the possible utility of using mouse as a model system to understand early epigenetic events during tumorigenesis as well as the general impact of aberrant chromatin remodeling events in cancer. Moreover, our study revealed several putative TSG that are novel targets for epigenetic silencing in human breast cancer.

Table I.

RT-PCR Primers

Primer Designation Human/Mouse Primer Sequence (5′ ➝ 3′) Orientation (Forw/Rev) Annealing Temp (°C) Thermocycles Product Size (bp)
Gapdh-RT M AACGACCCCTTCATTGAC F 57 30 191
TCCACGACATACTCAGCAC R
Timp3-RT M AATTCCTCTGCTCAGGCTCCTGTT F 60 30 429
AGCATGGTCATTTCCTCGGGTTCT R
Rprm-RT M TATTCGCCACGCTGCAGACCTAAT F 60 30 359
AAACCGTGCAGACAGGAGCTAAGA R
Smpd3-RT M ACATCGATTCTCCCACCAACACCT F 60 30 227
AATTCGCACAATGCAGCTGTCCTC R
Atp1b2-RT M TGCTTCTGGAATGTCCTTGACCCT F 60 30 368
TGGGTGGATGAGACCATTTCCTGT R
Dusp2-RT M TGGTCCTGTGGAAATCTTGCCCTA F 60 31 307
AATCAGGTATGCCAGGCAGATGGT R
FoxJ1-RT M AAGGTAACTTTGACTGGGAGGCCA F 60 30 225
AGGATGTGCCCAAGAAGGTCTCAT R

Table II.

Methylation-Specific PCR (MSP) Primers

Primer Designation Human/Mouse Primer Sequence (5′ ➝ 3′) Orientation (Forw/Rev) Annealing Temp (°C) Thermo- cycles Product Size (bp)
Timp3-MSP-M M CGGAGGGGGCGGATTTC F 60 40 84
CACGAATCCGCGAAACCAAC R
Timp3-MSP-U M TGGAGGGGGTGGATTTT F 55 40 184
ACACAAATCCACAAAACCAAC R
Rprm-MSP-M M GTCGGGTATTTGTTCGTTTCGC F 58 40 96
ACTAAATTCATCACGAAAACAAACGAAAACCG R
Rprm-MSP-U M GTTAGTGTTTTGTTGGGTATTTGTTTGTTTTGTG F 58 40 96
ACACTAAATTCATCACAAAAACAAACAAAAACCA R
Smpd3-MSP-M M GGGTTTTTTAGAGACGGTTTTCGC F 59 40 90
GCTCCAAATACTAAATCCGAAAAAAACTCCG R
Smpd3-MSP-U M TGATTGGGGTTTTTTAGAGATGGTTTTTGTG F 58 40 90
CCACTCCAAATACTAAATCCAAAAAAAACTCCA R
Atp1b2-MSP-M M CGTGGAGGGTTTTAGCGC F 58 40 74
AAACACGAAACGAATCCCGACG R
Aatp1b2-MSP-U M GTGTGTGGAGGGTTTTAGTGTG F 57.5 40 74
AACCATCTTAAAAACACAAAACAAATCCCAACA R
Dusp2-MSP-M M GTTCGTTTTTGGTTTTTTGTCGGAGTTAC F 58.5 40 102
AAAACAACGACGAAAATACCGCG R
Dusp2-MSP-U M TGTTGTTTGTTTTTGGTTTTTTGTTGGAGTTATG F 58 40 102
ACAAACAAACAAAAACAACAACAAAAATACCACA R
FoxJ1-MSP-M M GGTTTTTTTAGCGGTTAAATTCGACGC F 59 40 86
GCTAAATCTCACGACGAATACTCG R
FoxJ1-MSP-U M GGTTGGGTTTTTTTAGTGGTTAAATTTGATGTG F 58 40 86
CCCACACTAAATCTCACAACAAATACTCA R
SMPD3-MSP-M H TTAGATTTGCGGTCGGGAGTTC F 58.5 40 100
AACACTAACCTCTAACGACGAACG R
SMPD3-MSP-U H TTGTTGTTAGATTTGTGGTTGGGAGTTTG F 58.5 40 100
AAATTCAACAACACTAACCTCTAACAACAAACA R
ATP1B2-MSP-M H GGCGATCGCGGGTTTC F 59 40 116
CCGATAATAACTCAACCGCGAAAATAAAACG R
ATP1B2-MSP-U H GTATAAAGATTGTGTTTGGTGATTGTGGGTTTTG F 58.5 40 116
ACACCAATAATAACTCAACCACAAAAATAAAACA R
DUSP2-MSP-M H TTATCGTTACGAGAGTTCGGGAC F 58 40 119
CGCAACAACGTACCCAACG R
DUSP2-MSP-U H GGTAGTGTTATTGTTATGAGAGTTTGGGATG F 58.5 40 119
AAATCCCACAACAACATACCCAACA R
FOXJ1-MSP-M H TTTTTAGAACGAACGGATTTATAGC F 59 40 193
ACACTTACCACTCTCCCTAACGAC R
FOXJ1-MSP-U H TTTAGAATGAATGGATTTATAGTGG F 58 40 193
AACACTTACCACTCTCCCTAACAAC R
Ecad-MSP-M M GGTTTAATTCGGTTTTGTTCGATCGTATTC F 58 40 75
GAACTCCCATAACGAACCCG R
Ecad-MSP-U M GTTTGGTTTAATTTGGTTTTGTTTGATTGTATTTG F 58 40 75
ACCAAACTCCCATAACAAACCCA R
RassF1-MSP-M M GGGCGGTCGTTGGAATTTC F 58.5 40 99
CCCTCCGAATAACTCGCTATAAAAAAACG R
RassF1-MSP-U M AGGGGTGGTTGTTGGAATTTTG F 57.5 40 99
CAAAACCCTCCAAATAACTCACTATAAAAAAACA R
Gstp1-MSP-M M GAATTCGTATTTAGTAGGTAGTGAGTTCGTTTC F 58 40 106
CGATACAAAAAACGCCAATCGTAAACG R
Gstp1-MSP-U M TGAATTTGTATTTAGTAGGTAGTGAGTTTGTTTTG F 58 40 106
AACTACCCAATACAAAAAACACCAATCATAAACA R
Rarβ-MSP-M M GATTTTAAGTTTTTTTTTTTAGGCGACGTTC F 56.5 40 89
GAACAAAAAACGTCTAAACACCG R
Rarβ-MSP-U M ATTGATTTTAAGTTTTTTTTTTTAGGTGATGTTTG F 56.5 40 89
CCCAAACAAAAAACATCTAAACACCA R
Dapk1-MSP-M M TTCGGGTCGCGGAGTC F 59 40 110
CAATTCCTCGCCGATATCGTAATAATCG R
Dapk1-MSP-U M GATGGTTTGGGTTGTGGAGTTG F 57.5 40 110
CTACCCAATTCCTCACCAATATCATAATAATCA R
Wif1-MSP-M M GGGTATCGCGGTTTCGTTTTC F 58.5 40 96
CTACTAACAAAAAACGAACACTAAAAACGACCG R
Wif1-MSP-U M GGGTTTATTGGGTATTGTGGTTTTGTTTTTG F 58 40 96
ACTACTAACAAAAAACAAACACTAAAAACAACCA R

Table III.

Pyrosequencing Primers (Amplification and Sequencing)

Primer Designation Human/Mouse Primer Sequence (5′ ➝ 3′) Orientation (Forw/Rev) Annealing Temp (°C) Thermo- cycles Product Size (bp)
Rprm-PYRO M TTGTAGTTAAGGAAGAAGATGTTAAAAATTAT F 50.5 50 344
AACCTAAAACTTTAAAAAAAAAACTAAAAAAC R
GTTTGGGTGTAGTAG SEQ*
YGTATYGYGYGTTTTAGYGTTTYGTTAT SEQ to analyze**
Timp3-PYRO M GATGAATTTTTGTTTAGGTTT F 50 50 195
AAAATCCAAAACTACAAAATAAT R
TTTTTATAAGGATTTGAA SEQ*
YGGTTYGGTYGGTTTYGTTTTGYGTTTYGYGTTTTTYGGTTTYGTT SEQ to analyze**
ATP1B2-PYRO H GGTGGGAAGGGGGTGGGTATTT F 56 50 123
CAACCAAAAATAAAAAACCCCCCAAA R
GGGGTGGGTATTTTT SEQ*
YGYGTTYGGYGATYGYGGGTTTTYGTATT SEQ to analyze**
DUSP2-PYRO H GAAGTTTTTAGGTTAGTTTAGATTG F 53 50 242
CCTCCAACCCCATAACCACC R
GTTTTTGTATGGTGTTT SEQ*
YGGAGGTYGGTTTAYGGGTATTTAATTYGGGTYGTYGYGGA SEQ to analyze**
FOXJ1-PYRO H TTTTAGTTTTAGTTTTTAATTTTAGGTTT F 52 50 290
ACACTTACCACTCTCCCTAAC R
GTTTTAGGAGTAGTTGA SEQ*
YGTTTTAAAGYGTAYGGGATAGGAAGGGGGTGGGGGYGA SEQ to analyze**
SMPD3-PYRO H GGATGTGGTTTTGAAGGAGAGATTT F 53.5 50 305
GAATTCAACAACACTAACCTCTA R
GGGTTGTGAGGAG SEQ*
TYGGATTYGAGAGTYGAGAGTYGTYGTYGTTGYGGTYGTYGTT SEQ to analyze**
*

SEQ: Sequencing primer

**

SEQ to analyze: Gene-specific sequence analyzed by pyrosequencing (Y: Potential methylation site)

Table IV. Microarray Data.

Shown are the 984 genes found to be ≥5-fold upregulated in 5-azadC treated EMT6 cells, genes subject to further study are indicated in magenta.

Primary Sequence Name Sequence Description Accession # Fold Change P-value
6720406K03 hypothetical protein 6720406K03 AK032705 63.47921 2.69E-07
Gcg Glucagon, mRNA (cDNA clone MGC:14058 IMAGE:4218718) AK050538 63.45748 2.73E-07
2900027M19Rik Transmembrane protein 28 (Tmem28), mRNA AK045452 63.31421 2.98E-07
D830044D21Rik RIKEN cDNA D830044D21 gene (D830044D21Rik), mRNA NM_177210 61.82942 2.46E-07
Pkd1l2 Polycystic kidney disease 1 like 2 (Pkd1l2), mRNA NM_029686 60.27456 2.10E-44
Tspan33 Tetraspanin 33 (Tspan33), mRNA NM_146173 58.68657 8.71E-33
4930442J19Rik Adult male testis cDNA, RIKEN full-length enriched library, clone:4930442J19 product: unclassifiable, full insert sequence AK015363 56.56114 9.32E-08
2610024A01Rik RIKEN cDNA 2610024A01 gene (2610024A01Rik), mRNA BC059048 49.58956 4.61E-08
Fgf11 Fibroblast growth factor 11 (Fgf11), mRNA BC066859 48.44208 8.99E-26
AF366264 CDNA sequence AF366264 (AF366264), mRNA NM_153093 47.83025 3.85E-08
Ung2 PREDICTED: similar to uracil-DNA glycosylase 2 [Mus musculus], mRNA sequence AK053890 47.79745 3.05E-17
6430548M08Rik RIKEN cDNA 6430548M08 gene, mRNA (cDNA clone MGC:38801 IMAGE:5359842) AK122299 44.75741 1.10E-15
Serpini1 Serine (or cysteine) peptidase inhibitor, clade I, member 1, mRNA (cDNA clone MGC:6413 IMAGE:3588856) NM_009250 44.68072 1.38E-15
Dpep1 Dipeptidase 1 (renal), mRNA (cDNA clone MGC:6318 IMAGE:2812088) NM_007876 44.65215 2.73E-29
Cabc1 Chaperone, ABC1 activity of bc1 complex like (S. pombe), mRNA (cDNA clone MGC:31744 IMAGE:4922892) NM_023341 41.85896 2.83E-24
1190002H23Rik RIKEN cDNA 1190002H23 gene (1190002H23Rik), mRNA NM_025427 39.94254 9.82E-29
1600029D21Rik RIKEN cDNA 1600029D21 gene (1600029D21Rik), mRNA NM_029639 39.51588 1.83E-22
4930522H14Rik RIKEN cDNA 4930522H14 gene (4930522H14Rik), mRNA NM_026291 39.45048 5.43E-08
TC982356 Unknown 39.22191 3.98E-43
Xist Xist (X inactive specific transcript) mRNA for open reading frame X59289 37.43628 1.34E-23
Dusp2 Dual specificity phosphatase 2 (Dusp2), mRNA NM_010090 37.08016 1.15E-17
BB133117 Transcribed locus BB133117 37.05725 2.84E-13
Nlk Nemo like kinase (Nlk), mRNA AK031461 35.36866 1.11E-08
6430548M08Rik RIKEN cDNA 6430548M08 gene, mRNA (cDNA clone MGC:38801 IMAGE:5359842) NM_172286 33.78417 6.01E-14
BC011467 CDNA sequence BC011467, mRNA (cDNA clone MGC:37865 IMAGE:5100525) BC025823 32.62732 6.73E-17
LOC545748 similar to hypothetical protein LOC280487 AK031095 32.16875 3.93E-17
9630025C22 hypothetical protein 9630025C22 NM_177597 30.76949 9.72E-12
Hspa1a Heat shock protein (hsp68) mRNA, clone MHS213 NM_010479 30.68889 6.14E-11
Timp3 Tissue inhibitor of metalloproteinase 3, mRNA (cDNA clone MGC:25359 IMAGE:4239015) NM_011595 30.407 3.63E-12
H2-DMb1 Histocompatibility 2, class II, locus Mb1, mRNA (cDNA clone MGC:5741 IMAGE:3486844) NM_010387 30.20498 1.12E-44
Arrdc4 Arrestin domain containing 4 (Arrdc4), mRNA NM_025549 30.05997 1.22E-14
9330117B14 Mus musculus adult male diencephalon cDNA, RIKEN full-length enriched library, clone:9330117B14 product: unclassifiable, full insert sequence. AK033923 29.13712 8.04E-09
Sema4f Sema domain, immunoglobulin domain (Ig), TM domain, and short cytoplasmic domain (Sema4f), mRNA NM_011350 29.0931 1.55E-14
Fzd9 Mus musculus 16 days embryo head cDNA, RIKEN full-length enriched library, clone:C130077B15 product: frizzled homolog 9 (Drosophila), full insert sequence. AK021164 28.87872 2.71E-19
Zfpn1a4 Zinc finger protein, subfamily 1A, 4 (Eos) (Zfpn1a4), mRNA AK041640 27.41483 4.91E-20
Osbp2 Oxysterol binding protein 2 (Osbp2), mRNA AK049816 27.39835 3.84E-24
A930035J23Rik PREDICTED: Mus musculus RIKEN cDNA A930035J23 gene (A930035J23Rik), mRNA AK041893 27.09363 3.60E-28
Tmem25 Transmembrane protein 25 (Tmem25), mRNA NM_027865 26.96004 2.00E-25
Selenbp1 Selenium binding protein 1, mRNA (cDNA clone MGC:18519 IMAGE:4209393) NM_009150 26.94885 3.24E-18
Tfpi Tissue factor pathway inhibitor, mRNA (cDNA clone MGC:37332 IMAGE:4975683) NM_011576 26.6853 9.47E-30
Klk16 Kallikrein 16, mRNA (cDNA clone MGC:18522 IMAGE:4165443) NM_008454 26.63172 8.21E-09
Cd248 CD248 antigen, endosialin (Cd248), mRNA NM_054042 26.56997 7.43E-19
Pde2a Phosphodiesterase 2A, cGMP-stimulated, mRNA (cDNA clone IMAGE:4949471) BC006845 26.40924 2.97E-19
Agxt2l1 Alanine-glyoxylate aminotransferase 2-like 1 (Agxt2l1), mRNA AK030395 25.35919 8.75E-09
Slc30a1 Solute carrier family 30 (zinc transporter), member 1 (Slc30a1), mRNA AK048209 25.18293 7.34E-19
8030431J09Rik Ankyrin repeat and BTB (POZ) domain containing 2, mRNA (cDNA clone MGC:62845 IMAGE:6493840) AK033113 24.99684 1.12E-15
1700030G11Rik PREDICTED: RIKEN cDNA 1700030G11 [Mus musculus], mRNA sequence AK006546 24.99246 0.00001
Mlf1 Myeloid leukemia factor 1, mRNA (cDNA clone MGC:41163 IMAGE:1495721) NM_010801 24.45573 7.37E-15
A_51_P306031 Unknown 23.98088 4.47E-25
Chi3l3 Chitinase 3-like 3 (Chi3l3), mRNA NM_009892 23.96213 1.01E-38
Dbccr1 BRINP mRNA for BMP/retinoic acid-inducible neural-specific protein (BRINP) NM_019967 23.93755 3.66E-17
B230362M20Rik RIKEN cDNA B230362M20 gene (B230362M20Rik), mRNA NM_177273 23.77677 1.66E-08
9030221C07Rik Aromatic-preferring amino acid transporter (Arpat) NM_177802 23.37548 0.00001
Sipa1l2 Signal-induced proliferation-associated 1 like 2, mRNA (cDNA clone IMAGE:30360538) AK122501 23.25885 4.47E-36
Gm364 PREDICTED: similar to hypothetical protein [Mus musculus], mRNA sequence AV280269 23.15239 6.48E-38
Selenbp2 Selenium binding protein 2, mRNA (cDNA clone MGC:35927 IMAGE:5096831) NM_019414 23.06403 1.19E-14
Lin7b Lin 7 homolog b (C. elegans), mRNA (cDNA clone MGC:18338 IMAGE:4164883) NM_011698 22.83365 6.72E-27
Tacstd1 Tumor-associated calcium signal transducer 1, mRNA (cDNA clone MGC:11680 IMAGE:3711069) NM_008532 22.5851 4.03E-33
Eml5 Echinoderm microtubule associated protein like 5, mRNA (cDNA clone IMAGE:4953665) BC027154 22.21267 2.27E-28
Fank1 Fibronectin type 3 and ankyrin repeat domains 1 (Fank1), mRNA NM_025850 22.05824 3.68E-22
1700001J03Rik RIKEN cDNA 1700001J03 gene (1700001J03Rik), mRNA BC048528 21.14908 3.13E-26
NAP059666-1 Unknown 21.09742 3.79E-06
Klf2 Kruppel-like factor 2 (lung) (Klf2), mRNA NM_008452 20.99121 2.75E-09
AK054415 Mus musculus 2 days pregnant adult female ovary cDNA, RIKEN full-length enriched library, clone:E330023G08 product: phosphatidylinositol 3-kinase, C2 domain containing, gamma polypeptide, full insert sequence. AK054415 20.9248 0.00003
D330017J20Rik MKIAA1170 protein AK122459 20.91998 8.39E-35
NAP108451-1 Unknown 20.55861 0.00005
Gp2 Glycoprotein 2 (zymogen granule membrane) (Gp2), mRNA NM_025989 20.50423 3.80E-08
Ush2a PREDICTED: Mus musculus similar to usherin isoform B (LOC545397), mRNA AK044721 20.08695 2.99E-08
Abca3 Mus musculus B6-derived CD11 +ve dendritic cells cDNA, RIKEN full-length enriched library, clone:F730004H09 product: ATP-binding cassette, sub-family A (ABC1), member 3, full insert sequence. AK089316 19.91308 2.21E-08
Sned1 Secreted nidogen domain protein (Snep gene), splice form SNEP NM_172463 19.73947 1.44E-21
4930488E11Rik RIKEN cDNA 4930488E11 gene, mRNA (cDNA clone MGC:106799 IMAGE:3373135) NM_207267 19.66398 1.42E-30
Etv5 Ets variant gene 5, mRNA (cDNA clone MGC:28414 IMAGE:4036564) NM_023794 19.61583 5.13E-27
Disp2 Dispatched homolog 2 (Drosophila) (Disp2), mRNA NM_170593 19.59815 5.48E-08
ENSMUST00000064479 Unknown 19.4577 8.91E-11
TC952297 AF302077 neprilysin-like peptidase gamma {Mus musculus}, partial (4%) [TC952297] 19.41319 8.90E-30
Igbp1 Mus musculus 12 days embryo male wolffian duct includes surrounding region cDNA, RIKEN full-length enriched library, clone:6720407F13 product: immunoglobulin (CD79A) binding protein 1, full insert sequence. AK032709 19.30429 2.63E-08
TC1019179 Unknown 19.29024 1.03E-22
Cd248 CD248 antigen, endosialin (Cd248), mRNA NM_054042 19.23851 4.63E-24
Arhgap29 Rho GTPase activating protein 29 (Arhgap29), mRNA NM_172525 19.18384 3.71E-10
E430004F17Rik RIKEN cDNA E430004F17 gene (E430004F17Rik), mRNA NM_183199 19.05738 5.93E-16
Stard10 START domain containing 10 (Stard10), mRNA NM_019990 18.91585 1.20E-12
Fn3k Fructosamine 3 kinase (Fn3k), mRNA NM_022014 18.91504 1.03E-24
Pfn2 Profilin 2, mRNA (cDNA clone MGC:35930 IMAGE:5059568) NM_019410 18.78823 1.79E-18
Ptgfr Prostaglandin F receptor (Ptgfr), mRNA AK086563 18.75204 3.09E-08
Man1a Mannosidase 1, alpha, mRNA (cDNA clone MGC:18448 IMAGE:4223319) NM_008548 18.7403 3.14E-08
Vwf Von Willebrand factor (Vwf) mRNA, exon 1 and 5′UTR AK044921 18.63142 5.82E-08
Kif27 Kinesin family member 27 (Kif27), mRNA NM_175214 18.39237 2.38E-42
Sh2d4b SH2 domain containing 4B (Sh2d4b), mRNA NM_177816 18.37578 5.13E-08
Sh2d1b SH2 domain protein 1B (Sh2d1b), mRNA NM_012009 18.35538 3.50E-08
Arhgap29 Rho GTPase activating protein 29 (Arhgap29), mRNA NM_172525 17.9961 1.04E-10
AI450469 Transcribed locus AI450469 17.7114 4.97E-28
4930504O13Rik RIKEN cDNA 4930504O13 gene (4930504O13Rik), mRNA NM_207527 17.60137 7.87E-08
Anxa6 Mus musculus 10, 11 days embryo whole body cDNA, RIKEN full-length enriched library, clone:2810407D22 product: annexin A6, full insert sequence. AK013026 17.41486 1.20E-08
Mical2 MKIAA0750 protein NM_177282 17.22894 1.87E-11
Lrrc15 Leucine rich repeat containing 15 (Lrrc15), mRNA BC050245 17.22617 1.28E-27
NM_133237 Mus musculus cDNA sequence AB023957 (AB023957), mRNA [NM_133237] NM_133237 17.09971 2.44E-31
Myl2 Myosin, light polypeptide 2, regulatory, cardiac, slow (Myl2), mRNA NM_010861 17.05658 5.16E-06
Trib2 Tribbles homolog 2 (Drosophila), mRNA (cDNA clone MGC:32449 IMAGE:5043179) NM_144551 16.9309 1.31E-20
A930037G23Rik RIKEN cDNA A930037G23 gene, mRNA (cDNA clone MGC:67253 IMAGE:6403474) NM_178787 16.70077 1.01E-25
Derl3 Der1-like domain family, member 3 (Derl3), mRNA NM_024440 16.70039 2.27E-08
Rasl11b RAS-like, family 11, member B, mRNA (cDNA clone MGC:7064 IMAGE:3156762) AK004534 16.64886 4.78E-26
A130052D22 Mus musculus hypothetical protein A130052D22 (A130052D22), mRNA [NM_183306] NM_183306 16.27097 3.94E-29
Soat1 Sterol O-acyltransferase 1, mRNA (cDNA clone IMAGE:4007277) BC025091 16.22614 2.66E-44
Adam30 A disintegrin and metallopeptidase domain 30 (Adam30), mRNA AK077058 16.15904 0.00015
Abtb2 Ankyrin repeat and BTB (POZ) domain containing 2, mRNA (cDNA clone MGC:62845 IMAGE:6493840) NM_178890 16.12632 2.87E-27
Dock4 MKIAA0716 protein AK085106 15.90782 4.44E-37
6720468P15Rik 0 day neonate thymus cDNA, RIKEN full-length enriched library, clone:A430057M21 product: unclassifiable, full insert sequence AK078497 15.87182 4.79E-06
Cxcl4 Chemokine (C-X-C motif) ligand 4 (Cxcl4), mRNA NM_019932 15.81666 7.84E-36
Otub2 OTU domain, ubiquitin aldehyde binding 2 (Otub2), mRNA NM_026580 15.80523 1.53E-15
Thyn1 Thymocyte protein mThy28 AK037947 15.5622 1.23E-07
E130308A19Rik RIKEN cDNA E130308A19 gene (E130308A19Rik), transcript variant 1, mRNA AK038359 15.44643 1.28E-07
0610042C05Rik RIKEN cDNA 0610042C05 gene, mRNA (cDNA clone MGC:37398 IMAGE:4977394) AK076350 15.44004 4.96E-07
Chn2 Beta chimaerin (bch gene) NM_023543 15.41819 2.69E-07
Rab3b RAB3B, member RAS oncogene family (Rab3b), mRNA NM_023537 15.2858 2.35E-15
Anxa6 Annexin A6, mRNA (cDNA clone MGC:6574 IMAGE:3482035) NM_013472 15.14969 1.10E-13
Nedd9 Neural precursor cell expressed, developmentally down-regulated gene 9, mRNA (cDNA clone IMAGE:3499250) NM_017464 15.10892 1.79E-15
Dazl Deleted in azoospermia-like (Dazl), mRNA NM_010021 15.03384 3.11E-43
Glcci1 Glucocorticoid induced transcript 1 (Glcci1), transcript variant 1, mRNA AK042221 14.97325 2.00E-07
Tfdp2 Transcription factor Dp 2, mRNA (cDNA clone IMAGE:30604420) NM_178667 14.96471 2.55E-17
Cacna1g Calcium channel, voltage-dependent, T type, alpha 1G subunit (Cacna1g), mRNA NM_009783 14.95897 1.24E-16
Ifna4 Interferon alpha family, gene 4 (Ifna4), mRNA NM_010504 14.95108 1.76E-07
Notch3 Notch gene homolog 3 (Drosophila) (Notch3), mRNA NM_008716 14.90218 1.13E-28
Pacsin1 Protein kinase C and casein kinase substrate in neurons 1, mRNA (cDNA clone MGC:25285 IMAGE:4527708) NM_011861 14.88882 1.74E-11
Atp1b2 ATPase, Na+/K+ transporting, beta 2 polypeptide (Atp1b2), mRNA NM_013415 14.7351 1.07E-22
Habp2 HYALURONIC ACID BINDING PROTEIN 2; HYADURONIC ACID-BINDING PROTEIN 2. [Source:RefSeq;Acc:NM_146101] [ENSMUST00000026060] 14.65301 2.18E-06
Fabp7 Fatty acid binding protein 7, brain (Fabp7), mRNA NM_021272 14.34666 4.16E-06
D14Ertd171e DNA segment, Chr 14, ERATO Doi 171, expressed, mRNA (cDNA clone MGC:59433 IMAGE:6331997) AK039674 14.3364 2.78E-07
Oas1d 2′-5′olygoadenylate synthetase 1D (Oas1d) NM_133893 14.18881 1.11E-14
Nid1 Nidogen 1 (Nid1), mRNA NM_010917 14.08903 5.04E-23
Mcf2l Mcf.2 transforming sequence-like (Mcf2l), mRNA NM_178076 14.07378 6.06E-26
Hbb-b1 Hemoglobin, beta adult major chain, mRNA (cDNA clone MGC:40691 IMAGE:3988455) NM_008220 14.02514 4.51E-07
Prg4 PREDICTED: proteoglycan 4 [Mus musculus], mRNA sequence AB034730 14.01659 1.28E-07
4930524B15Rik RIKEN cDNA 4930524B15 gene (4930524B15Rik), mRNA NM_026262 13.99617 8.54E-07
4833416E15Rik 0 day neonate head cDNA, RIKEN full-length enriched library, clone:4833416E15 product: unclassifiable, full insert sequence AK014707 13.9541 1.99E-43
Zfp422-rs1 zinc finger protein 422, related sequence 1 AK083028 13.94195 3.70E-34
AI607043 Adult male small intestine cDNA, RIKEN full-length enriched library, clone:2010310C07 product: unclassifiable, full insert sequence AI607043 13.90419 1.08E-12
Dnajc5b DnaJ (Hsp40) homolog, subfamily C, member 5 beta (Dnajc5b), mRNA NM_025489 13.89499 0.00001
Slc2a6 Solute carrier family 2 (facilitated glucose transporter), member 6 (Slc2a6), mRNA NM_172659 13.84051 3.99E-43
Drctnnb1a down-regulated by Ctnnb1, a NM_053090 13.83104 1.14E-34
Sdc4 Syndecan 4, mRNA (cDNA clone MGC:11456 IMAGE:3154160) NM_011521 13.79767 7.42E-23
Ell3 Elongation factor RNA polymerase II-like 3 (Ell3), mRNA NM_145973 13.77959 5.40E-23
Stard10 START domain containing 10 (Stard10), mRNA NM_019990 13.74577 6.06E-19
Mapk8ip1 JNK interacting protein-1b (JIP-1b) AK053819 13.62189 1.54E-12
1110051M20Rik RIKEN cDNA 1110051M20 gene, transcript variant 2, mRNA (cDNA clone MGC:103036 IMAGE:6440075) AK053282 13.62105 1.43E-18
Ppfia4 MKIAA0897 protein XM_129443 13.46761 1.10E-27
Itga6 Integrin alpha 6 (Itga6), mRNA BC024571 13.44888 1.15E-11
Nfatc2 Mus musculus 16 days embryo head cDNA, RIKEN full-length enriched library, clone:C130082H17 product: nuclear factor of activated T-cells, cytoplasmic 2, full insert AK081853 13.39808 9.81E-45
4933401N24Rik RIKEN cDNA 4933401N24 gene (4933401N24Rik), mRNA NM_175367 13.39483 1.76E-41
Mpp7 Membrane protein, palmitoylated 7 (MAGUK p55 subfamily member 7), mRNA (cDNA clone IMAGE:6532780) AK036956 13.36708 1.64E-28
Acpp Acid phosphatase, prostate (Acpp), transcript variant 2, mRNA NM_207668 13.35389 1.35E-17
AK038010 16 days neonate thymus cDNA, RIKEN full-length enriched library, clone:A130071G05 product: unclassifiable, full insert sequence AK038010 13.30008 5.17E-07
XM_156408 Mus musculus similar to RIKEN cDNA 2610524H06 gene (LOC239727), mRNA [XM_156408] XM_156408 13.28002 4.58E-14
Itga6 Integrin alpha 6 (Itga6), mRNA NM_008397 13.26801 5.17E-12
1300013J15Rik RIKEN cDNA 1300013J15 gene, mRNA (cDNA clone MGC:28678 IMAGE:4238749) NM_026183 13.1909 1.26E-20
Hsd17b13 Hydroxysteroid (17-beta) dehydrogenase 13, mRNA (cDNA clone MGC:30360 IMAGE:5132342) AK050378 13.159 8.50E-07
Ankrd13d Ankyrin repeat domain 13 family, member D (Ankrd13d), mRNA NM_026720 13.07501 1.65E-18
Sh3tc1 SH3 domain and tetratricopeptide repeats 1, mRNA (cDNA clone IMAGE:3490511) NM_194344 13.05323 1.40E-45
Gas6 Growth arrest specific 6, mRNA (cDNA clone MGC:6124 IMAGE:3592398) NM_019521 13.04704 3.30E-34
Fank1 Fibronectin type 3 and ankyrin repeat domains 1 (Fank1), mRNA NM_025850 13.03388 2.66E-34
6430590A10Rik Mus musculus RIKEN cDNA 6430590A10 gene (6430590A10Rik), mRNA [NM_177017] NM_177017 12.97531 0.00001
Nck2 Non-catalytic region of tyrosine kinase adaptor protein 2, mRNA (cDNA clone MGC:7093 IMAGE:3157491) AK042129 12.94784 4.72E-07
1700123K08Rik RIKEN cDNA 1700123K08 gene (1700123K08Rik), mRNA NM_029693 12.92483 2.79E-11
Jag1 Jagged1 (Jag1) NM_013822 12.89675 1.67E-18
Cacna1g Calcium channel, voltage-dependent, T type, alpha 1G subunit (Cacna1g), mRNA NM_009783 12.88716 9.48E-20
2410003J06Rik RIKEN cDNA 2410003J06 gene, mRNA (cDNA clone MGC:60956 IMAGE:30007513) AK010362 12.79956 2.82E-16
Pgbd5 PiggyBac transposable element derived 5, mRNA (cDNA clone MGC:106643 IMAGE:30606749) AK013562 12.79654 1.93E-30
Nfe2l3 Nuclear factor, erythroid derived 2, like 3, mRNA (cDNA clone MGC:5702 IMAGE:3485182) NM_010903 12.78367 8.28E-39
D330012F22Rik Mus musculus RIKEN cDNA D330012F22 gene (D330012F22Rik), mRNA. NM_178752 12.76423 3.55E-12
Odz3 ODZ3 (Odz3) NM_011857 12.75018 6.34E-06
Plat Plasminogen activator, tissue, mRNA (cDNA clone MGC:18508 IMAGE:4038404) NM_008872 12.73118 5.16E-23
Hist1h1e Histone 1, H1e (Hist1h1e), mRNA AK047852 12.71911 1.20E-35
Atp6v0e2 ATPase, H+ transporting, lysosomal, V0 subunit E isoform 2 (Atp6v0e2), mRNA NM_133764 12.6221 7.20E-18
Capn5 Calpain 5, mRNA (cDNA clone MGC:25253 IMAGE:4458488) NM_007602 12.51361 5.38E-23
Fzd4 Frizzled homolog 4 (Drosophila), mRNA (cDNA clone MGC:18403 IMAGE:4238940) NM_008055 12.51026 5.20E-28
Nppb Natriuretic peptide precursor type B (Nppb), mRNA NM_008726 12.3018 2.94E-13
4631402N15Rik RIKEN cDNA 4631402N15 gene AK028622 12.293 1.87E-06
Hey1 Hairy-related transcription factor 1 (Hrt1) NM_010423 12.27919 9.21E-35
Cldn19 Claudin 19 (Cldn19), mRNA NM_153105 12.26932 2.86E-06
Olfr488 Olfactory receptor 488 (Olfr488), mRNA NM_146732 12.23274 1.87E-06
4930402F06Rik PREDICTED: hypothetical protein LOC74854 [Mus musculus], mRNA sequence XM_130219 12.21738 2.16E-06
Aldh6a1 Aldehyde dehydrogenase family 6, subfamily A1, mRNA (cDNA clone MGC:28587 IMAGE:4212084) NM_134042 12.14788 1.68E-44
Olfr512 Olfactory receptor 512 (Olfr512), mRNA NM_146724 12.01476 1.44E-06
Baiap2l1 BAI1-associated protein 2-like 1 (Baiap2l1), mRNA NM_025833 11.99713 2.71E-21
Gcnt1 Glucosaminyl (N-acetyl) transferase 1, core 2, mRNA (cDNA clone MGC:11452 IMAGE:3154671) NM_010265 11.98347 1.35E-11
C330006P03Rik Homer homolog 1 (Drosophila) (Homer1), transcript variant L, mRNA AK049142 11.98031 1.54E-10
Wfdc12 WAP four-disulfide core domain 12, mRNA (cDNA clone MGC:32134 IMAGE:4923469) NM_138684 11.9789 4.85E-23
Als2cr15 Amyotrophic lateral sclerosis 2 (juvenile) chromosome region, candidate 15, mRNA (cDNA clone MGC:90771 IMAGE:30355976) NM_027407 11.97403 2.12E-20
Phtf2 Putative homeodomain transcription factor 2, mRNA (cDNA clone MGC:60851 IMAGE:30050876) AK035536 11.97298 4.35E-26
Itga4 Mus musculus 16 days neonate thymus cDNA, RIKEN full-length enriched library, clone:A130050E18 product: integrin alpha 4, full insert sequence. AK037794 11.95943 1.52E-06
Col5a3 Asp6 mRNA for adipocyte-specific protein 6 NM_016919 11.95215 2.56E-07
Egr1 Early growth response 1 (Egr1), mRNA NM_007913 11.93462 4.12E-43
Muc2 Mucin 2, mRNA (cDNA clone MGC:37529 IMAGE:4986546) AK008250 11.89613 5.08E-08
Drctnnb1a down-regulated by Ctnnb1, a NM_053090 11.8877 1.99E-36
Ctss Cathepsin S, mRNA (cDNA clone MGC:6643 IMAGE:3495719) NM_021281 11.87519 2.68E-24
D7Ertd413e Mus musculus DNA segment, Chr 7, ERATO Doi 413, expressed (D7Ertd413e), mRNA [NM_177468] NM_177468 11.87163 2.22E-07
Fcmd Fukuyama type congenital muscular dystrophy homolog (human) (Fcmd), mRNA NM_139309 11.84098 2.11E-06
Rab26 RAB26, member RAS oncogene family, mRNA (cDNA clone MGC:107516 IMAGE:6436242) AK043912 11.75247 1.64E-12
Gm520 Mus musculus similar to hypothetical protein FLJ35880 (LOC245027), mRNA [XM_147082] XM_147082 11.75046 4.73E-11
Rasgef1a PREDICTED: RasGEF domain family, member 1A [Mus musculus], mRNA sequence AK018120 11.73011 8.35E-29
Chrna10 cholinergic receptor, nicotinic, alpha polypeptide 10 AK033068 11.71933 6.66E-15
Ptpn13 Protein tyrosine phosphatase, non-receptor type 13 (Ptpn13), mRNA AK033818 11.71558 4.43E-33
BC039632 CDNA sequence BC039632, mRNA (cDNA clone IMAGE:5124409) BC039632 11.63456 4.31E-15
4632419I22Rik Mus musculus 0 day neonate skin cDNA, RIKEN full-length enriched library, clone:4632419I22 product: unclassifiable, full insert sequence. AK014588 11.6079 3.26E-06
Rab3il1 RAB3A interacting protein (rabin3)-like 1 (Rab3il1), mRNA NM_144538 11.55758 7.46E-24
Rapgef2 Rap guanine nucleotide exchange factor (GEF) 2, mRNA (cDNA clone IMAGE:30545886) BM247313 11.55429 3.38E-06
Cpeb4 Cytoplasmic polyadenylation element binding protein 4 (Cpeb4), mRNA AK079421 11.54925 5.71E-19
Ctgf Connective tissue growth factor, mRNA (cDNA clone MGC:8122 IMAGE:3589136) NM_010217 11.54686 1.93E-08
Wnt6 Wingless-related MMTV integration site 6 (Wnt6), mRNA NM_009526 11.37961 1.23E-17
Atp7a ATPase, Cu++ transporting, alpha polypeptide (Atp7a), mRNA AK033254 11.34701 1.47E-28
Efemp1 Epidermal growth factor-containing fibulin-like extracellular matrix protein 1 (Efemp1), mRNA NM_146015 11.34476 3.71E-35
Cyp2s1 Cytochrome P450, family 2, subfamily s, polypeptide 1, mRNA (cDNA clone IMAGE:4986756) NM_028775 11.3342 8.41E-25
Plat Plasminogen activator, tissue, mRNA (cDNA clone MGC:18508 IMAGE:4038404) NM_008872 11.28647 2.45E-19
Hmg20a High mobility group 20A, mRNA (cDNA clone MGC:12002 IMAGE:3602239) AK032790 11.27672 1.40E-45
NAP002592-003 Unknown 11.26244 2.03E-08
Ppl Periplakin (Ppl), mRNA NM_008909 11.23955 1.47E-09
TC976091 AY129963 GPI-gamma 4; GPIgamma4 {Mus musculus}, partial (72%) [TC976091] 11.21971 2.49E-24
BC067047 PREDICTED: PREX1 protein [Mus musculus], mRNA sequence AK090301 11.18218 2.25E-21
Tekt1 Tektin 1 (Tekt1), mRNA NM_011569 11.17168 2.41E-26
Atrnl1 Attractin like 1, mRNA (cDNA clone IMAGE:4221668) AK050882 11.16275 1.82E-14
4930470F04Rik RAB11 family interacting protein 2 (class I) (Rab11fip2), mRNA AK015532 11.14061 0.00748
Svep1 Sushi, von Willebrand factor type A, EGF and pentraxin domain containing 1, mRNA (cDNA clone IMAGE:3582135) AK035333 11.11646 3.62E-06
AK052909 16 days neonate heart cDNA, RIKEN full-length enriched library, clone:D830036I24 product: weakly similar to CYTOCHROME C OXIDASE POLYPEPTIDE VA, MITOCHONDRIAL PRECURSOR (EC 1.9.3.1) [Rattus norvegicus], full insert sequence AK052909 11.00683 1.38E-15
Lrp4 MKIAA0816 protein AK032360 11.00157 7.64E-06
Khk Ketohexokinase, mRNA (cDNA clone MGC:18722 IMAGE:4235560) NM_008439 10.99577 1.01E-23
Mpp7 Membrane protein, palmitoylated 7 (MAGUK p55 subfamily member 7), mRNA (cDNA clone IMAGE:6532780) BC049662 10.97808 2.54E-27
Muc3 Mucin 3, intestinal, mRNA (cDNA clone IMAGE:4989899) BC058768 10.94339 2.55E-18
3-Mar Membrane-associated ring finger (C3HC4) 3 (March3), mRNA AK085528 10.91169 6.53E-19
Baiap2l1 BAI1-associated protein 2-like 1 (Baiap2l1), mRNA NM_025833 10.90945 1.29E-08
Ptger2 Prostaglandin E receptor 2 (subtype EP2), mRNA (cDNA clone MGC:6041 IMAGE:3487087) NM_008964 10.83013 1.65E-07
Tnfaip6 Tumor necrosis factor alpha induced protein 6, mRNA (cDNA clone MGC:13914 IMAGE:4208738) AK078295 10.7698 2.30E-17
AK037893 Adult male hypothalamus cDNA, RIKEN full-length enriched library, clone:A230098A08 product: hypothetical protein, full insert sequence AK037893 10.7616 3.07E-15
Ppic Peptidylprolyl isomerase C, mRNA (cDNA clone MGC:31015 IMAGE:5026674) NM_008908 10.73102 1.49E-26
Kcnj1 Potassium inwardly-rectifying channel, subfamily J, member 1, mRNA (cDNA clone MGC:18350 IMAGE:4241262) NM_019659 10.65826 0.00005
Zfp423 Early B-cell factor-associated zinc finger protein (Ebfaz) NM_033327 10.59541 6.09E-06
Rora RAR-related orphan receptor alpha, mRNA (cDNA clone MGC:5892 IMAGE:3592667) AK087905 10.59469 9.75E-10
AK050947 9 days embryo whole body cDNA, RIKEN full-length enriched library, clone:D030042L12 product: unclassifiable, full insert sequence AK050947 10.59118 5.88E-06
AK076715 Mus musculus adult male testis cDNA, RIKEN full-length enriched library, clone:4930420A11 product: unclassifiable, full insert sequence. AK076715 10.58991 2.55E-06
Ttn Titin, mRNA (cDNA clone IMAGE:5149168) AK009965 10.56327 0.00001
AK038867 Adult male hypothalamus cDNA, RIKEN full-length enriched library, clone:A230069K02 product: hypothetical protein, full insert sequence AK038867 10.55157 0.00001
Ifrd1 Interferon-related developmental regulator 1 (Ifrd1), mRNA AK078888 10.54876 2.35E-12
Bmp8a Bone morphogenetic protein 8a (Bmp8a), mRNA NM_007558 10.50666 1.72E-12
NP_TR6JSE50FPA olfactory receptor NP_TR6JSE50FPA NM_203509 10.50266 6.41E-23
TC1084975 AE003651 CG13260-PA {Drosophila melanogaster}, partial (10%) [TC1084975] 10.49094 2.76E-07
Tnfaip6 Tumor necrosis factor alpha induced protein 6, mRNA (cDNA clone MGC:13914 IMAGE:4208738) NM_009398 10.44943 2.17E-09
LOC432449 hypothetical gene supported by AK038984 AK038984 10.44738 2.83E-07
2900041A09Rik RIKEN cDNA 2900041A09 gene (2900041A09Rik), mRNA NM_182839 10.44139 8.16E-28
1600015H20Rik RIKEN cDNA 1600015H20 gene, mRNA (cDNA clone IMAGE:3592288) NM_024237 10.43001 4.20E-45
4930412F09Rik Mus musculus adult male testis cDNA, RIKEN full-length enriched library, clone:4930412F09 product: unclassifiable, full insert sequence. AK015119 10.4231 6.86E-26
1700108L22Rik RIKEN cDNA 1700108L22 gene, mRNA (cDNA clone MGC:38253 IMAGE:5324328) AK037099 10.41494 4.96E-24
Tjp3 Tight junction protein 3, mRNA (cDNA clone MGC:11960 IMAGE:3600848) NM_013769 10.41438 5.26E-21
NAP053018-1 Unknown 10.38831 1.26E-44
Ibrdc3 IBR domain containing 3, mRNA (cDNA clone IMAGE:5254400) AK015966 10.37732 9.05E-35
Slco4a1 Solute carrier organic anion transporter family, member 4a1 (Slco4a1), mRNA NM_148933 10.35873 7.30E-26
E430024C06Rik Mus musculus 2 days neonate thymus thymic cells cDNA, RIKEN full-length enriched library, clone:E430003N15 product: unclassifiable, full insert sequence. AK088091 10.34123 3.10E-23
1500026H17Rik Adult male corpora quadrigemina cDNA, RIKEN full-length enriched library, clone:B230317N08 product: unclassifiable, full insert sequence AK005299 10.33636 4.98E-16
3110057O12Rik RIKEN cDNA 3110057O12 gene, mRNA (cDNA clone IMAGE:4007705) NM_026622 10.33092 1.71E-11
Ptprm Protein tyrosine phosphatase, receptor type, M (Ptprm), mRNA AK047905 10.30778 8.35E-06
Hemt1 17 days embryo kidney cDNA, RIKEN full-length enriched library, clone:I920059J21 product:11 beta-hydroxylase homolog [Mus sp], full insert sequence AK078619 10.30203 7.99E-06
Stau1 Staufen (RNA binding protein) homolog 1 (Drosophila), mRNA (cDNA clone MGC:13708 IMAGE:4188394) AK030890 10.2707 2.31E-16
Hist1h1d Histone 1, H1d (Hist1h1d), mRNA NM_145713 10.25591 3.12E-13
D16H22S680E DNA segment, Chr 16, human D22S680E, expressed (D16H22S680E), mRNA AK047383 10.23081 1.29E-32
Rgl1 Ral guanine nucleotide dissociation stimulator,-like 1, mRNA (cDNA clone MGC:18430 IMAGE:4241244) AK032340 10.22274 1.40E-45
8430422M09Rik Mus musculus RIKEN cDNA 8430422M09 gene (8430422M09Rik), mRNA [XM_148115] XM_148115 10.21893 0.00001
Kcnab3 Potassium voltage-gated channel, shaker-related subfamily, beta member 3, mRNA (cDNA clone MGC:27635 IMAGE:4507029) NM_010599 10.2158 1.74E-06
Slc25a21 Solute carrier family 25 (mitochondrial oxodicarboxylate carrier), member 21, mRNA (cDNA clone MGC:67702 IMAGE:5042653) NM_172577 10.21333 5.59E-14
Col20a1 Collagen, type XX, alpha 1, mRNA (cDNA clone IMAGE:5366843) AK006759 10.16397 3.65E-12
Hrbl HIV-1 Rev binding protein-like (Hrbl), transcript variant 1, mRNA AK037434 10.14872 3.47E-17
NM_029423 Mus musculus RIKEN cDNA 4833414G05 gene (4833414G05Rik), mRNA [NM_029423] NM_029423 10.1245 3.34E-13
Kcnma1 Large conductance calcium activated potassium BK channel STREX-1 variant (Slo) AK052354 10.12088 7.76E-06
Hivep3 Clone 5′RACE1 ZAS3 (Krc) mRNA, 5′ UTR AK087330 10.11146 8.57E-37
Isl2 Insulin related protein 2 (islet 2) (Isl2), mRNA NM_027397 10.06446 6.70E-06
A930034L06Rik RIKEN cDNA A930034L06 gene (A930034L06Rik), mRNA NM_175692 10.0482 3.73E-14
Rhbg Rhesus blood group-associated B glycoprotein, mRNA (cDNA clone MGC:35874 IMAGE:4218889) NM_021375 10.00581 2.67E-18
Grtp1 GH regulated TBC protein 1, mRNA (cDNA clone MGC:27905 IMAGE:3500563) NM_025768 9.93642 5.59E-28
Pfkfb2 6-phosphofructo-2-kinase/fructose-2,6-biphosphatase 2, mRNA (cDNA clone MGC:25723 IMAGE:3979400) AK016729 9.92957 3.07E-35
BC049806 CDNA sequence BC049806, mRNA (cDNA clone MGC:76460 IMAGE:30431671) NM_172513 9.90803 1.62E-19
Cdc14a CDC14 cell division cycle 14 homolog A (S. cerevisiae), mRNA (cDNA clone MGC:100044 IMAGE:30356962) AK048250 9.87444 0.0038
Hist1h4i Mus musculus histone 2, H4, mRNA (cDNA clone IMAGE:4205460), partial cds. BC019757 9.86807 1.47E-19
Mcam Melanoma cell adhesion molecule, mRNA (cDNA clone MGC:35823 IMAGE:5372497) NM_023061 9.84222 1.58E-07
Mfi2 Antigen p97 (melanoma associated) identified by monoclonal antibodies 133.2 and 96.5 (Mfi2), mRNA NM_013900 9.83413 4.36E-21
Dock4 MKIAA0716 protein BC051125 9.81909 2.29E-11
Xlr3b X-linked lymphocyte-regulated 3B (Xlr3b), mRNA NM_011727 9.78323 6.44E-14
NAP069339-1 Unknown 9.77056 2.40E-32
2610018I05Rik Mus musculus RIKEN cDNA 2610018I05 gene (2610018I05Rik), mRNA [NM_183279] NM_183279 9.77006 1.01E-17
Nrarp Notch-regulated ankyrin repeat protein (Nrarp), mRNA NM_025980 9.74784 4.38E-07
Eml5 Mus musculus echinoderm microtubule associated protein like 5, mRNA (cDNA clone MGC:40838 IMAGE:5368750), complete cds. BC030361 9.71342 1.76E-27
Slc26a4 Pendrin (Pds) NM_011867 9.7024 2.38E-37
Sdfr1 Stromal cell derived factor receptor 1 (Sdfr1), mRNA AK039616 9.69602 1.98E-06
A430105I05Rik B6-derived CD11 +ve dendritic cells cDNA, RIKEN full-length enriched library, clone:F730031B04 product: Serine-threonine kinase (Fragment) homolog [Rattus norvegicus], full insert sequence AK040531 9.69534 3.56E-18
Dgat2 Diacylglycerol O-acyltransferase 2 (Dgat2), mRNA NM_026384 9.69276 2.73E-35
W29301 2 days pregnant adult female oviduct cDNA, RIKEN full-length enriched library, clone:E230029K24 product: hypothetical protein, full insert sequence W29301 9.68389 1.54E-44
Gng8 Guanine nucleotide binding protein (G protein), gamma 8 subunit (Gng8), mRNA NM_010320 9.68159 4.39E-19
Ppfibp2 Protein tyrosine phosphatase, receptor-type, F interacting protein, binding protein 2, mRNA (cDNA clone IMAGE:3488375) NM_008905 9.67203 0.00004
Aplp2 Amyloid beta (A4) precursor-like protein 2, mRNA (cDNA clone IMAGE:4504805) NM_009691 9.54457 8.29E-26
4921524J17Rik RIKEN cDNA 4921524J17 gene (4921524J17Rik), mRNA AK006764 9.52196 0.00001
Nppb Natriuretic peptide precursor type B (Nppb), mRNA NM_008726 9.5214 2.01E-10
AK082626 0 day neonate cerebellum cDNA, RIKEN full-length enriched library, clone:C230071E14 product: unclassifiable, full insert sequence AK082626 9.49805 0.00002
Ccpg1 Cell cycle progression 1, mRNA (cDNA clone MGC:12151 IMAGE:3711012) NM_028181 9.49574 1.25E-22
4933401N24Rik RIKEN cDNA 4933401N24 gene (4933401N24Rik), mRNA NM_175367 9.48097 1.88E-15
Etv5 Ets variant gene 5, mRNA (cDNA clone MGC:28414 IMAGE:4036564) NM_023794 9.46181 1.65E-37
Upb1 Ureidopropionase, beta, mRNA (cDNA clone IMAGE:4974180) NM_133995 9.45902 7.68E-06
AK041503 PREDICTED: Mus musculus hypothetical protein LOC (LOC545502), mRNA AK041503 9.45271 0.00022
Arc Activity regulated cytoskeletal-associated protein, mRNA (cDNA clone MGC:27890 IMAGE:3498057) NM_018790 9.4495 6.20E-31
4931406B18Rik RIKEN cDNA 4931406B18 gene (4931406B18Rik), mRNA NM_028737 9.44613 0.00004
Atg12 Apg12 NM_026217 9.43954 2.67E-16
Zswim5 Zinc finger, SWIM domain containing 5 (Zswim5), mRNA AK046815 9.43427 0.00003
Pacsin3 Protein kinase C and casein kinase II substrate 3 (Pacsin3) AK014830 9.42759 4.14E-13
AW125688 Expressed sequence AW125688 (AW125688), mRNA NM_001001885 9.37585 5.66E-26
Odz2 ODZ2 (Odz2) AK083775 9.36205 0.00002
Smpd3 Sphingomyelin phosphodiesterase 3, neutral (Smpd3), mRNA NM_021491 9.34228 2.30E-12
Ceacam2 CEA-related cell adhesion molecule 2, mRNA (cDNA clone MGC:18435 IMAGE:4241449) NM_007543 9.33621 5.14E-12
Mmp10 Matrix metallopeptidase 10 (Mmp10), mRNA NM_019471 9.28633 1.13E-07
BC013712 CDNA sequence BC013712, mRNA (cDNA clone IMAGE:3155889) BC013712 9.27602 1.41E-15
Cfh Complement component factor h (Cfh), mRNA NM_009888 9.21081 1.00E-18
H2-Oa Histocompatibility 2, O region alpha locus (H2-Oa), mRNA NM_008206 9.20769 2.53E-16
Cacng5 Calcium channel, voltage-dependent, gamma subunit 5 (Cacng5), mRNA NM_080644 9.20551 4.94E-09
Slc28a2 Purine-selective Na+ nucleoside cotransporter (Cnt2) NM_172980 9.20316 2.97E-09
AK089607 Mus musculus activated spleen cDNA, RIKEN full-length enriched library, clone:F830004H11 product: unclassifiable, full insert sequence. AK089607 9.1922 0.00004
Gcnt1 Glucosaminyl (N-acetyl) transferase 1, core 2, mRNA (cDNA clone MGC:11452 IMAGE:3154671) NM_010265 9.17498 8.19E-10
Serpinb1a EIA (Serpinb1) NM_025429 9.17307 6.72E-18
Ccbl2 Cysteine conjugate-beta lyase 2 (Ccbl2), mRNA NM_173763 9.15601 3.33E-11
Nsccn1 Mus musculus non-selective cation channel 1 (Nsccn1), mRNA [NM_010940] NM_010940 9.15204 0.0001
Hrc Histidine rich calcium binding protein (Hrc), mRNA NM_010473 9.13879 6.63E-24
H2-DMb2 Histocompatibility 2, class II, locus Mb1, mRNA (cDNA clone MGC:5741 IMAGE:3486844) NM_010388 9.10407 1.15E-27
Rgl1 Ral guanine nucleotide dissociation stimulator,-like 1, mRNA (cDNA clone MGC:18430 IMAGE:4241244) NM_016846 9.0778 1.59E-41
Dapk2 Death-associated kinase 2, mRNA (cDNA clone MGC:13742 IMAGE:4016911) NM_010019 9.06365 4.58E-16
Lmtk2 Lemur tyrosine kinase 2, mRNA (cDNA clone IMAGE:6407188) AK039738 9.05942 1.27E-38
Cfh Complement component factor h, mRNA (cDNA clone IMAGE:5132971) M29010 8.96985 2.80E-06
Spry1 Sprouty homolog 1 (Drosophila), mRNA (cDNA clone MGC:18307 IMAGE:3672353) NM_011896 8.96343 8.83E-21
Gm693 PREDICTED: similar to novel protein [Mus musculus], mRNA sequence XM_204104 8.95937 1.91E-21
Ryr1 RyR1 mRNA for skeletal muscle ryanodine receptor AK042482 8.94761 2.14E-13
8430437N05Rik DACH protein (Dach) AK018473 8.93946 0.00004
Arl5c ADP-ribosylation factor-like 12 (Arl12), mRNA NM_207231 8.93395 2.10E-19
Rasip1 Ras interacting protein 1, mRNA (cDNA clone IMAGE:1245387) AK033276 8.93241 7.99E-10
Adam25 Testase 2 (Adam-25) NM_011781 8.91583 0.00004
4930579G22Rik RIKEN cDNA 4930579G22 gene (4930579G22Rik), mRNA NM_026916 8.91564 1.40E-14
NAP062857-1 Unknown 8.89087 0.0001
Pbx3 Pre B-cell leukemia transcription factor 3 (Pbx3), mRNA AK052017 8.85804 0.00004
Kcnip1 A-type potassium channel modulatory protein 1.2 (Kchip1.2) NM_027398 8.84607 0.00004
Pard6b Par-6 (partitioning defective 6) homolog beta (C. elegans), mRNA (cDNA clone MGC:35871 IMAGE:2655658) NM_021409 8.83071 3.85E-15
Flrt3 Strain C57BL/6xCBA fibronectin leucine rich transmembrane protein 3 (Flrt3) NM_178382 8.81882 2.05E-14
AK032440 Adult male olfactory brain cDNA, RIKEN full-length enriched library, clone:6430547C23 product: unclassifiable, full insert sequence AK032440 8.80642 0.00004
Cpn1 Carboxypeptidase N, polypeptide 1, mRNA (cDNA clone MGC:25883 IMAGE:4211169) NM_030703 8.79262 7.85E-18
2810405K02Rik RIKEN cDNA 2810405K02 gene, mRNA (cDNA clone MGC:40622 IMAGE:5401948) NM_025582 8.78415 0.00006
9230110K08Rik Adult male epididymis cDNA, RIKEN full-length enriched library, clone:9230110K08 product: hypothetical HMG-I and HMG-Y DNA-binding domain (A+T-hook) containing protein, full insert sequence AK020331 8.77293 4.72E-29
Igsf1 Immunoglobulin superfamily, member 1, transcript variant 2, mRNA (cDNA clone IMAGE:30249729) NM_183336 8.75732 0.00008
Ndrg4 N-myc downstream regulated gene 4 (Ndrg4), mRNA NM_145602 8.75653 6.57E-36
Mst1r Macrophage stimulating 1 receptor (c-met-related tyrosine kinase) (Mst1r), mRNA NM_009074 8.71853 6.63E-18
Cdon Cell adhesion molecule-related/down-regulated by oncogenes (Cdon), mRNA NM_021339 8.71613 1.34E-16
Serpine1 Serine (or cysteine) peptidase inhibitor, clade E, member 1 (Serpine1), mRNA NM_008871 8.66415 7.69E-24
Atg12 Apg12 NM_026217 8.66365 5.05E-14
AK051813 Mus musculus 12 days embryo eyeball cDNA, RIKEN full-length enriched library, clone:D230004F15 product: unclassifiable, full insert sequence. AK051813 8.6557 6.88E-09
F830010H11Rik Mus musculus activated spleen cDNA, RIKEN full-length enriched library, clone:F830010H11 product: unclassifiable, full insert sequence. AK089716 8.65219 0.00002
Clu Clusterin (Clu), mRNA NM_013492 8.64934 3.24E-12
Akap12 A kinase (PRKA) anchor protein (gravin) 12 (Akap12), mRNA NM_031185 8.6406 1.71E-27
Pacsin1 Protein kinase C and casein kinase substrate in neurons 1, mRNA (cDNA clone MGC:25285 IMAGE:4527708) NM_178365 8.62899 5.37E-08
4930449I04Rik Adult male testis cDNA, RIKEN full-length enriched library, clone:4930449I04 product: hypothetical protein, full insert sequence AK015432 8.6266 3.24E-33
Ap3m2 Adaptor-related protein complex 3, mu 2 subunit, mRNA (cDNA clone MGC:40858 IMAGE:5369288) NM_029505 8.62306 1.02E-10
Olfr1383 Olfactory receptor 1383 (Olfr1383), mRNA NM_207574 8.60579 1.28E-24
Lip1 Lysosomal acid lipase 1 (Lip1), mRNA AK083099 8.59815 2.87E-09
Ero1lb ERO1-like beta (S. cerevisiae) (Ero1lb), mRNA NM_026184 8.59729 5.41E-29
Zfp509 Mus musculus 2 days pregnant adult female ovary cDNA, RIKEN full-length enriched library, clone:E330031I18 product: weakly similar to ZINC FINGER PROTEIN ZFP-93 (ZINC FINGER PROTEIN HZF6) (FRAGMENT) [Homo sapiens], full insert sequence. AK054488 8.5938 1.99E-28
Fgd4 FYVE, RhoGEF and PH domain containing 4 (Fgd4), transcript variant beta, mRNA AK043656 8.53643 5.81E-08
Il1rap Interleukin 1 receptor accessory protein (Il1rap), transcript variant 2, mRNA AK045686 8.52846 5.61E-45
1700109H08Rik RIKEN cDNA 1700109H08 gene, mRNA (cDNA clone MGC:118394 IMAGE:30920581) AK007150 8.52551 1.98E-13
Ptger4 Mus musculus 12 days embryo spinal ganglion cDNA, RIKEN full-length enriched library, clone:D130004L12 product: prostaglandin E receptor 4 (subtype EP4), full insert sequence. AK051137 8.52297 6.25E-26
4933408J17Rik Small nuclear ribonucleoprotein polypeptide F, mRNA (cDNA clone MGC:118343 IMAGE:30917013) AK016739 8.52005 0.00012
Jrkl 0 day neonate thymus cDNA, RIKEN full-length enriched library, clone:A430098H07 product: hypothetical protein, full insert sequence AK079877 8.51387 4.41E-18
Cxcl9 Chemokine (C-X-C motif) ligand 9, mRNA (cDNA clone MGC:6179 IMAGE:3257716) NM_008599 8.50345 1.39E-17
Cfh Complement component factor h (Cfh), mRNA NM_009888 8.49969 1.56E-11
AK042806 Mus musculus 7 days neonate cerebellum cDNA, RIKEN full-length enriched library, clone:A730026C13 product: unclassifiable, full insert sequence. AK042806 8.4994 1.23E-08
Gdpd5 Glycerophosphodiester phosphodiesterase domain containing 5, mRNA (cDNA clone MGC:37148 IMAGE:4953174) NM_201352 8.48545 5.38E-09
Tgm4 Experimental autoimmune prostatitis antigen 1 (Eapa1) NM_177911 8.48033 0.00012
Btbd5 BTB (POZ) domain containing 5, mRNA (cDNA clone MGC:47075 IMAGE:5347183) NM_025707 8.46778 1.71E-15
Rtn4r Reticulon 4 receptor (Rtn4r), mRNA NM_022982 8.45249 7.05E-28
Ibrdc3 IBR domain containing 3, mRNA (cDNA clone IMAGE:5254400) AK040902 8.4511 6.53E-22
Aldh1l2 Aldehyde dehydrogenase 1 family, member L2 (Aldh1l2), mRNA AK085074 8.41092 2.26E-14
LOC433886 hypothetical gene supported by AK049058; BC025881 BC025881 8.39834 0.00022
Kiss1 Golgi transport 1 homolog A (S. cerevisiae) (Golt1a), mRNA NM_178260 8.39812 0.00014
Egr3 Early growth response 3 (Egr3), mRNA AK078299 8.39771 1.35E-12
NAP112027-1 Unknown 8.38015 1.69E-11
Myh3 PREDICTED: myosin, heavy polypeptide 3, skeletal muscle, embryonic [Mus musculus], mRNA sequence M11154 8.3727 4.98E-25
Vil1 Villin 1, mRNA (cDNA clone MGC:18506 IMAGE:4236751) NM_009509 8.36524 5.30E-09
Aplp2 Amyloid beta (A4) precursor-like protein 2, mRNA (cDNA clone IMAGE:4504805) NM_009691 8.36361 8.30E-43
Gats Opposite strand transcription unit to Stag3 (Gats), mRNA BC026208 8.35244 8.31E-10
Olfr1052 Olfactory receptor 1052 (Olfr1052), mRNA NM_147010 8.34985 0.00013
Akap13 PREDICTED: hypothetical protein LOC75547 [Mus musculus], mRNA sequence AK006382 8.31218 1.41E-14
Nanos1 Nanos homolog 1 (Drosophila), mRNA (cDNA clone MGC:102288 IMAGE:6849859) BC056473 8.30066 1.94E-13
Eif2c3 Eukaryotic translation initiation factor 2C, 3 (Eif2c3), mRNA AK087577 8.26887 1.33E-22
AK086157 Mus musculus 15 days embryo head cDNA, RIKEN full-length enriched library, clone:D930008M17 product: unclassifiable, full insert sequence. AK086157 8.26126 0.00012
Art3 ART3 mono(ADP-ribosyl)transferase (art3 gene), splice variant 1 NM_181728 8.2485 9.93E-16
TC1085358 AF179867 STE20-like kinase {Homo sapiens}, partial (56%) [TC1085358] 8.24529 1.54E-22
Eif2c3 Eukaryotic translation initiation factor 2C, 3 (Eif2c3), mRNA BC060127 8.21917 1.00E-29
Cd52 CD52 antigen, mRNA (cDNA clone MGC:40993 IMAGE:1396480) NM_013706 8.21899 0.00035
Slc25a29 Solute carrier family 25 (mitochondrial carrier, palmitoylcarnitine transporter), member 29, mRNA (cDNA clone MGC:7958 IMAGE:3584570) NM_181328 8.20668 2.73E-17
Kif13b MKIAA0639 protein AK031370 8.20576 2.43E-10
Itpka Inositol 1,4,5-trisphosphate 3-kinase A (Itpka), mRNA NM_146125 8.20068 2.20E-07
Efhd1 EF hand domain containing 1, mRNA (cDNA clone MGC:28727 IMAGE:4459208) NM_028889 8.18839 4.79E-24
Ccpg1 Cell cycle progression 1, mRNA (cDNA clone MGC:12151 IMAGE:3711012) BC006717 8.15286 2.83E-33
1810020D17Rik RIKEN cDNA 1810020D17 gene, mRNA (cDNA clone MGC:36240 IMAGE:5027461) AK031619 8.13287 0.00011
Ripk3 Receptor-interacting serine-threonine kinase 3, mRNA (cDNA clone MGC:35896 IMAGE:3590770) NM_019955 8.12366 1.65E-15
Zdhhc15 Zinc finger, DHHC domain containing 15 (Zdhhc15), mRNA BU610239 8.10685 0.00016
Scgb1c1 PREDICTED: similar to RYD5 [Mus musculus], mRNA sequence XM_355976 8.10518 1.20E-23
XM_356367 Mus musculus similar to cyclin-dependent kinase-like 5; serine/threonine kinase 9 (LOC382253), mRNA [XM_356367] XM_356367 8.10486 1.44E-12
Rab3il1 Mus musculus RAB3A interacting protein (rabin3)-like 1 (Rab3il1), mRNA. NM_178881 8.09444 1.48E-10
Egfl5 MKIAA0818 protein NM_172694 8.09398 1.04E-08
Csnd Mus musculus 10 days lactation, adult female mammary gland cDNA, RIKEN full-length enriched library, clone:D730003B11 product: casein delta, full insert sequence. AK052789 8.09332 0.00016
Fkbp6 FK506 binding protein 6 (Fkbp6), mRNA NM_033571 8.09194 5.51E-33
Dll4 Dll-4 mRNA for Delta-4 NM_019454 8.08534 8.83E-21
Sema4g Semaphorin subclass 4 member G (sema4g) NM_011976 8.0833 5.15E-11
Rccd1 Protein regulator of cytokinesis 1 (Prc1), mRNA AK047578 8.0818 4.76E-07
Cul4a Cullin 4A (Cul4a), mRNA AK084008 8.07649 0.00002
Tec Cytoplasmic tyrosine kinase, Dscr28C related (Drosophila), mRNA (cDNA clone MGC:46873 IMAGE:5029430) NM_013689 8.06 3.75E-18
Entpd2 Ectonucleoside triphosphate diphosphohydrolase 2, mRNA (cDNA clone MGC:5806 IMAGE:3582550) NM_009849 8.05603 1.54E-20
Krtap16-4 Keratin associated protein 16-4 (Krtap16-4), mRNA NM_130873 8.05589 0.00004
Aqp11 Aquaporin 11, mRNA (cDNA clone IMAGE:5126527) NM_175105 8.05418 1.48E-42
Cd36 CD36 antigen, mRNA (cDNA clone MGC:6068 IMAGE:3481681) L23108 8.05303 7.36E-26
1700001G17Rik RIKEN cDNA 1700001G17 gene (1700001G17Rik), mRNA NM_026204 8.04819 1.22E-14
Pgr Progesterone receptor (Pgr), mRNA AK036862 8.04787 0.00003
Maf c-maf=c-Maf protein {proto-oncogene} [mice, BALB/c, cerebellum, mRNA, 2736 nt]. S74567 8.03437 0.00007
Sdc4 Syndecan 4, mRNA (cDNA clone MGC:11456 IMAGE:3154160) NM_011521 8.0206 1.74E-10
Aytl1 Hypothetical protein A330042H22 (A330042H22), mRNA NM_173014 8.01654 7.00E-11
2310047O13Rik RIKEN cDNA 2310047O13 gene, mRNA (cDNA clone MGC:28605 IMAGE:4217391) AK049694 8.01005 1.88E-15
Trp73 P73 alpha protein (P73 gene) AK014503 8.00474 0.00017
Tagln3 Transgelin 3 (Tagln3), mRNA NM_019754 8.00047 0.00012
NAP052665-1 Unknown 7.99834 3.51E-13
Cd40 CD40 antigen (Cd40), transcript variant 5, mRNA NM_170701 7.98914 2.13E-26
4732465E10Rik MKIAA0716 protein AK028870 7.98626 1.12E-30
1700109F18Rik Homeo box D4 (Hoxd4), mRNA AK006878 7.96332 0.00015
Slc25a29 Solute carrier family 25 (mitochondrial carrier, palmitoylcarnitine transporter), member 29, mRNA (cDNA clone MGC:7958 IMAGE:3584570) NM_181328 7.95316 5.52E-28
Ebi3 Epstein-Barr virus induced gene 3, mRNA (cDNA clone MGC:11457 IMAGE:3154957) NM_015766 7.94831 1.93E-15
D330017J20Rik Mus musculus RIKEN cDNA D330017J20 gene (D330017J20Rik), mRNA [NM_177204] NM_177204 7.9325 1.86E-28
A930037G23Rik RIKEN cDNA A930037G23 gene, mRNA (cDNA clone MGC:67253 IMAGE:6403474) AK044382 7.92998 0.00002
Wdr47 WD repeat domain 47 (Wdr47), mRNA NM_181400 7.92668 7.41E-15
Ppl Periplakin (Ppl), mRNA BC057152 7.88627 4.33E-13
3110021A11Rik Rab6 interacting protein 2 (Rab6ip2), transcript variant 1, mRNA AK052063 7.84546 0.00001
Dmn MKIAA0353 protein NM_201639 7.82763 3.07E-17
Olfr541 Olfactory receptor 541 (Olfr541), mRNA NM_146962 7.82426 0.0002
Lgr4 PREDICTED: G protein-coupled receptor 48 [Mus musculus], mRNA sequence AK044357 7.82294 1.53E-17
Star Steroidogenic acute regulatory protein (Star), mRNA NM_011485 7.81952 6.22E-16
Plekha7 Pleckstrin homology domain containing, family A member 7, mRNA (cDNA clone MGC:90927 IMAGE:3602481) NM_172743 7.81937 5.45E-12
D330028D13Rik RIKEN cDNA D330028D13 gene, mRNA (cDNA clone MGC:67688 IMAGE:4487036) NM_172727 7.81506 3.42E-10
Rprm Reprimo, TP53 dependent G2 arrest mediator candidate, mRNA (cDNA clone MGC:41066 IMAGE:1434823) NM_023396 7.7925 1.35E-15
Lcn12 Lipocalin 12 (Lcn12), mRNA AK020307 7.78453 0.00011
Spire1 Spire homolog 1 (Drosophila), mRNA (cDNA clone MGC:106458 IMAGE:30535423) AK053505 7.77423 9.18E-16
Glt25d2 Glycosyltransferase 25 domain containing 2, mRNA (cDNA clone MGC:92944 IMAGE:5706069) NM_177756 7.7719 0.00026
Ddit3 DNA-damage inducible transcript 3, mRNA (cDNA clone MGC:18641 IMAGE:4012433) NM_007837 7.77126 3.83E-06
Gab3 Growth factor receptor bound protein 2-associated protein 3 (Gab3), mRNA NM_181584 7.75833 1.55E-13
1110031B11Rik RIKEN cDNA 1110031B11 gene (1110031B11Rik), mRNA NM_026811 7.73455 0.00024
Tcf7 Transcription factor 7, T-cell specific (Tcf7), mRNA NM_009331 7.7187 0.00008
Lcn5 Lipocalin 5 (Lcn5), mRNA NM_007947 7.70424 0.00016
BB064041 Activated spleen cDNA, RIKEN full-length enriched library, clone:F830206M21 product: unclassifiable, full insert sequence BB064041 7.6678 1.96E-26
Tagln Transgelin, mRNA (cDNA clone MGC:6045 IMAGE:3600413) NM_011526 7.64204 1.12E-11
Tktl1 Transketolase-like 1 (Tktl1), mRNA NM_031379 7.63328 0.00021
Mlph Melanophilin (Mlph), mRNA NM_053015 7.6104 0.00021
Armc9 RIKEN cDNA 4930438O05 gene (4930438O05Rik), mRNA NM_030184 7.59315 8.43E-21
9330102E08Rik PREDICTED: hypothetical protein XP_489154 [Mus musculus], mRNA sequence AK033854 7.58577 7.54E-14
Cbara1 Calcium binding atopy-related autoantigen 1 (Cbara1), mRNA AK029884 7.57763 9.77E-14
Hisppd2a 4 days neonate male adipose cDNA, RIKEN full-length enriched library, clone:B430315C20 product: hypothetical Histidine acid phosphatase containing protein, full insert sequence AK046696 7.56076 1.42E-12
TC1052172 Unknown 7.55807 1.37E-10
Dcun1d4 Expressed sequence AI836376, mRNA (cDNA clone IMAGE:3995042) NM_178896 7.55542 4.88E-10
Ralgps1 Ral GEF with PH domain and SH3 binding motif 1, mRNA (cDNA clone MGC:99991 IMAGE:30536315) AK129123 7.52544 1.74E-06
Flrt3 Strain C57BL/6xCBA fibronectin leucine rich transmembrane protein 3 (Flrt3) NM_178382 7.51845 1.29E-33
Efnb2 Ephrin B2 (Efnb2), mRNA NM_010111 7.50936 8.34E-13
Mum1l1 Melanoma associated antigen (mutated) 1-like 1, mRNA (cDNA clone IMAGE:30619674) NM_175541 7.50527 4.98E-06
Zfp185 Zinc finger protein 185, mRNA (cDNA clone MGC:49126 IMAGE:4191053) NM_009549 7.50102 5.10E-06
Epb4.1l5 Erythrocyte protein band 4.1-like 5 (Epb4.1l5), mRNA AK079301 7.49371 1.30E-11
Tnfaip8l2 Tumor necrosis factor, alpha-induced protein 8-like 2 (Tnfaip8l2), mRNA NM_027206 7.48745 3.98E-30
Rad9b RAD9 homolog B (S. cerevisiae) (Rad9b), mRNA NM_144912 7.47671 3.20E-11
Irf8 Interferon consensus sequence binding protein 1, mRNA (cDNA clone MGC:6194 IMAGE:3487214) NM_008320 7.47332 6.42E-32
9430085L16Rik Early B-cell factor 2 (Ebf2), mRNA AK020505 7.46734 0.00033
9630050P21Rik 16 days neonate cerebellum cDNA, RIKEN full-length enriched library, clone:9630050P21 product: unclassifiable, full insert sequence AK036274 7.46671 1.10E-07
Gpr155 G protein-coupled receptor 155, mRNA (cDNA clone IMAGE:4167684) BC026382 7.45823 3.04E-31
Habp4 Hyaluronic acid binding protein 4 (Habp4), mRNA NM_019986 7.45096 7.37E-17
9830169C18Rik RIKEN cDNA 9830169C18 gene, mRNA (cDNA clone MGC:69959 IMAGE:6509090) NM_172581 7.4493 6.68E-15
Lrp1b Low density lipoprotein-related protein 1B (deleted in tumors) (Lrp1b), mRNA AK034373 7.43347 0.00006
Btg2 B-cell translocation gene 2, anti-proliferative (Btg2), mRNA NM_007570 7.43345 2.06E-10
2410066E13Rik RIKEN cDNA 2410066E13 gene (2410066E13Rik), mRNA NM_026629 7.43071 6.08E-15
Got1l1 RIKEN cDNA 1700083M11 gene (1700083M11Rik), mRNA AK006984 7.42792 8.13E-06
Mell1 Neprilysin-like metallopeptidase 1 (Nl1) NM_013783 7.41912 4.75E-11
Pde6c Phosphodiesterase 6C, cGMP specific, cone, alpha prime, mRNA (cDNA clone MGC:38745 IMAGE:5358611) NM_033614 7.40919 0.00024
3-Mar Membrane-associated ring finger (C3HC4) 3 (March3), mRNA AK036622 7.38467 0.00002
TC1013578 AF110520 NG28 {Mus musculus}, partial (30%) [TC1013578] 7.37995 2.09E-06
2310046K01Rik RIKEN cDNA 2310046K01 gene (2310046K01Rik), mRNA NM_027172 7.37995 6.69E-29
Ldhc Lactate dehydrogenase 3, C chain, sperm specific (Ldh3), mRNA NM_013580 7.36571 3.03E-20
Mgst3 Microsomal glutathione S-transferase 3, mRNA (cDNA clone MGC:35738 IMAGE:5372427) NM_025569 7.36371 7.31E-12
AK084877 0 day neonate eyeball cDNA, RIKEN full-length enriched library, clone:E130014I08 product: unclassifiable, full insert sequence AK084877 7.35577 7.22E-23
4732467B22 hypothetical protein 4732467B22 NM_177822 7.33593 6.02E-12
Spib Spi-B transcription factor (Spi-1/PU.1 related) (Spib), mRNA U87620 7.32567 1.70E-08
Aldh1a1 Ahd-2=acetaldehyde dehydrogenase [mice, BALB/c, livers, mRNA Partial, 1632 nt] NM_013467 7.32059 1.68E-29
BB128963 Expressed sequence BB128963, mRNA (cDNA clone IMAGE:3964696) AK088715 7.29587 3.02E-21
Samd8 Sterile alpha motif domain containing 8 (Samd8), mRNA NM_026283 7.28944 2.04E-13
Dnmt3b DNA cytosine methyltransferase 3b6 (Dnmt3b6) mRNA, complete cds; alternatively spliced NM_010068 7.28075 1.55E-06
Txnip Thioredoxin interacting protein, mRNA (cDNA clone MGC:25534 IMAGE:3591421) AK004653 7.27867 6.51E-09
D930040F23Rik Mus musculus adult male olfactory brain cDNA, RIKEN full-length enriched library, clone:6430407I07 product: unclassifiable, full insert sequence. AK032178 7.26988 3.49E-09
Wdr47 WD repeat domain 47 (Wdr47), mRNA NM_181400 7.26652 1.73E-06
Pla2g5 Phospholipase A2, group V, mRNA (cDNA clone MGC:31537 IMAGE:4502326) NM_011110 7.25852 7.82E-12
Tex101 Testis expressed gene 101 (Tex101), mRNA NM_019981 7.22574 4.77E-20
Gprk5 G protein-coupled receptor kinase 5 (GRK5) AK079806 7.217 0.00001
Cry2 Cryptochrome 2 (photolyase-like), mRNA (cDNA clone IMAGE:3593948) NM_009963 7.21232 4.87E-34
Epb4.1l3 Protein 4.1B (Epb4.1l3) AK086340 7.20588 0.00037
Cpeb4 Cytoplasmic polyadenylation element binding protein 4 (Cpeb4), mRNA AY313775 7.18189 1.32E-08
Rasgef1b RasGEF domain family, member 1B (Rasgef1b), transcript variant 1, mRNA NM_181318 7.17597 4.25E-16
Atrnl1 Attractin like 1, mRNA (cDNA clone IMAGE:4221668) AK048968 7.15363 4.73E-28
Ablim1 Actin-binding LIM protein 1, mRNA (cDNA clone IMAGE:5323557) AK031453 7.15289 6.91E-24
D930005D10Rik MKIAA1849 protein NM_178702 7.13213 5.89E-24
Rgl1 Ral guanine nucleotide dissociation stimulator,-like 1, mRNA (cDNA clone MGC:18430 IMAGE:4241244) NM_016846 7.12829 2.93E-40
Olfr295 Olfactory receptor 295 (Olfr295), mRNA NM_146851 7.12443 2.88E-06
Ccpg1 Cell cycle progression 1, mRNA (cDNA clone MGC:12151 IMAGE:3711012) BC006717 7.12363 1.51E-24
Zbtb10 PREDICTED: zinc finger and BTB domain containing 10 [Mus musculus], mRNA sequence AK083443 7.11228 0.00061
Icosl Icos ligand, mRNA (cDNA clone MGC:35971 IMAGE:4217333) NM_015790 7.1024 0.0004
Mycn Neuroblastoma myc-related oncogene 1, mRNA (cDNA clone MGC:6240 IMAGE:3495446) NM_008709 7.10045 0.0004
D030074K08Rik YY1 associated factor 2 (Yaf2), mRNA AK083749 7.09956 0.00016
Slc6a18 Solute carrier family 6 (neurotransmitter transporter), member 18, mRNA (cDNA clone MGC:18346 IMAGE:4235659) AK085473 7.09686 0.00038
Tex12 Testis protein TEX12 (Tex12) NM_025687 7.09649 1.01E-10
Usp49 Ubiquitin specific peptidase 49 (Usp49), mRNA NM_198421 7.09535 1.27E-14
Magi3 Membrane associated guanylate kinase, WW and PDZ domain containing 3 (Magi3), mRNA NM_133853 7.08507 8.81E-28
Mgl1 Macrophage galactose N-acetyl-galactosamine specific lectin 1 (Mgl1), mRNA NM_010796 7.07263 8.28E-25
Pdlim5 PDZ and LIM domain 5, mRNA (cDNA clone MGC:46824 IMAGE:4457868) NM_019809 7.05824 9.20E-15
A630005I04Rik RIKEN cDNA A630005I04 gene (A630005I04Rik), mRNA AK088332 7.03002 2.01E-26
AK089990 RCB-0559 K-1. F1 cDNA, RIKEN full-length enriched library, clone:G430046J05 product: unclassifiable, full insert sequence AK089990 7.0234 9.15E-19
Dmn MKIAA0353 protein NM_201639 7.0219 2.13E-11
Fabp4 Fatty acid binding protein 4, adipocyte (Fabp4), mRNA NM_024406 7.02153 1.00E-18
4930463G05Rik PREDICTED: Mus musculus RIKEN cDNA 4930463G05 gene (4930463G05Rik), mRNA AK015217 7.02125 1.22E-42
Rshl3 PREDICTED: similar to radial spoke head-like 1 [Mus musculus], mRNA sequence AK038980 7.01713 0.00039
Smpd3 Sphingomyelin phosphodiesterase 3, neutral (Smpd3), mRNA NM_021491 7.00037 1.98E-09
AI931714 PREDICTED: hypothetical protein LOC102182 [Mus musculus], mRNA sequence AK045296 7.00021 4.36E-14
Diras1 DIRAS family, GTP-binding RAS-like 1 (Diras1), mRNA NM_145217 6.99843 8.46E-28
Olfr460 Olfactory receptor 460 (Olfr460), mRNA NM_146383 6.98029 0.00004
Lphn3 Latrophilin 3, mRNA (cDNA clone IMAGE:30549106) AK053210 6.97894 0.00044
1700026L06Rik RIKEN cDNA 1700026L06 gene (1700026L06Rik), mRNA BC055108 6.97247 2.83E-27
Kremen1 Kringle containing transmembrane protein 1 (Kremen1), mRNA AK081707 6.95532 2.39E-24
B3gnt4 Beta1,3 N-acetylglucosaminyltransferase-4 (B3gnt4) NM_198611 6.94475 2.02E-12
Hcn3 Hyperpolarization-activated, cyclic nucleotide-gated K+ 3 (Hcn3), mRNA NM_008227 6.93488 8.78E-08
Plxdc1 Plexin domain containing 1 (Plxdc1), mRNA NM_028199 6.93463 8.74E-18
Meig1 Meiosis expressed gene 1 (Meig1), mRNA NM_008579 6.92131 9.92E-14
Cdx1 Caudal type homeo box 1, mRNA (cDNA clone MGC:28644 IMAGE:4224352) NM_009880 6.91978 0.00044
Zp3 Zona pellucida glycoprotein 3 (Zp3), mRNA NM_011776 6.91854 8.16E-36
6530409C15Rik PREDICTED: hypothetical protein LOC76224 [Mus musculus], mRNA sequence AK018330 6.91722 0.00041
4930518F03Rik Mus musculus RIKEN cDNA 4930518F03 gene (4930518F03Rik), mRNA [NM_177191] NM_177191 6.91301 0.00073
Mmp2 Matrix metallopeptidase 2 (Mmp2), mRNA BF147716 6.91006 1.84E-11
Herc5 Iduronato sulfatase AK037442 6.90618 7.13E-07
Neud4 Clone m62 zinc finger protein neuro-d4 (neuro-d4) NM_013874 6.90322 1.87E-11
Slc16a10 Solute carrier family 16 (monocarboxylic acid transporters), member 10, mRNA (cDNA clone MGC:60866 IMAGE:30046388) AK011813 6.89975 2.58E-10
TC947784 SNAG_HUMAN Gamma-soluble NSF attachment protein (SNAP-gamma) (N-ethylmaleimide- sensitive factor attachment protein, gamma). [Human] {Homo sapiens}, partial (13%) [TC947784] 6.88434 0.00048
Tmem64 Transmembrane protein 64, mRNA (cDNA clone MGC:40684 IMAGE:3661112) AK035894 6.87747 0.00049
AK043237 Mus musculus 7 days neonate cerebellum cDNA, RIKEN full-length enriched library, clone:A730074B07 product: unclassifiable, full insert sequence. AK043237 6.86471 0.00065
Gm172 PREDICTED: similar to hypothetical protein [Mus musculus], mRNA sequence AV254918 6.86109 7.31E-13
Apobec2 Apolipoprotein B editing complex 2, mRNA (cDNA clone MGC:41184 IMAGE:1433092) NM_009694 6.85954 0.0007
Bcl2l11 BCL2-like 11 (apoptosis facilitator), transcript variant 1, mRNA (cDNA clone MGC:28730 IMAGE:4459720) BC058175 6.85608 1.80E-13
Gcnt1 Glucosaminyl (N-acetyl) transferase 1, core 2, mRNA (cDNA clone MGC:11452 IMAGE:3154671) NM_173442 6.84626 6.17E-10
Fcgr3 Fc gamma receptor III (Fcgr3) mRNA, Fcgr3-b allele NM_010188 6.844 1.64E-15
Zdhhc6 Zinc finger, DHHC domain containing 6, mRNA (cDNA clone MGC:30394 IMAGE:4459242) NM_025883 6.84083 0.00001
Pbx4 Pre-B-cell leukemia transcription factor 4, mRNA (cDNA clone MGC:40990 IMAGE:1263363) NM_030555 6.83768 6.20E-23
Eif2c3 Eukaryotic translation initiation factor 2C, 3 (Eif2c3), mRNA BC060127 6.80015 7.50E-12
1810057P16Rik RIKEN cDNA 1810057P16 gene (1810057P16Rik), mRNA AK075801 6.77942 5.18E-12
Osbpl8 Oxysterol binding protein-like 8 (Osbpl8), transcript variant 2, mRNA AK036477 6.77762 1.16E-17
Lysmd4 RIKEN cDNA 4930506D23 gene, mRNA (cDNA clone MGC:65434 IMAGE:4504057) AK039294 6.75923 2.83E-18
Fsd1 Fibronectin type 3 and SPRY domain-containing protein (Fsd1), mRNA NM_183178 6.74462 0.0004
Lcmt1 Leucine carboxyl methyltransferase 1, mRNA (cDNA clone IMAGE:3595687) NM_025304 6.72952 3.34E-35
Bach2 BTB and CNC homology 2 (Bach2), mRNA NM_007521 6.72785 5.54E-12
Kif17 Kinesin family member 17 (Kif17), mRNA NM_010623 6.70819 8.92E-14
LOC195281 similar to Proton-associated sugar transporter A (PAST-A) (Deleted in neuroblastoma 5 protein) (DNb-5) XM_111577 6.70613 4.14E-29
D330012F22Rik RIKEN cDNA D330012F22 gene, mRNA (cDNA clone MGC:47349 IMAGE:4458984) AK084531 6.70519 3.43E-07
Arhgef7 Rho guanine nucleotide exchange factor (GEF7) (Arhgef7), mRNA AK129064 6.70342 1.94E-21
D930046H04Rik Mus musculus RIKEN cDNA D930046H04 gene (D930046H04Rik), mRNA [NM_176934] NM_176934 6.70196 0.00005
Tmem24 Transmembrane protein 24, mRNA (cDNA clone IMAGE:5360163) AK004920 6.68313 1.32E-19
Cyp46a1 Cytochrome P450, family 46, subfamily a, polypeptide 1, mRNA (cDNA clone MGC:18311 IMAGE:4195579) NM_010010 6.67773 6.83E-22
Usp22 MKIAA1063 protein AK086502 6.67222 1.15E-09
9530059J11Rik Mus musculus adult male urinary bladder cDNA, RIKEN full-length enriched library, clone:9530059J11 product: weakly similar to 12 DAYS EMBRYO HEAD CDNA, RIKEN FULL-LENGTH ENRICHED LIBRARY, CLONE:3010005E14, FULL INSERT SEQUENCE [Mus musculus], full insert sequence. AK020615 6.66825 1.21E-08
Mgst3 Microsomal glutathione S-transferase 3, mRNA (cDNA clone MGC:35738 IMAGE:5372427) AK008211 6.65726 9.29E-09
9330107J05Rik PREDICTED: hypothetical protein LOC73884 [Mus musculus], mRNA sequence AK047625 6.65622 0.00061
Grb7 Growth factor receptor bound protein 7, mRNA (cDNA clone MGC:5653 IMAGE:3583896) NM_010346 6.65154 6.22E-12
Pros1 Protein S (alpha) (Pros1), mRNA NM_011173 6.6501 9.72E-31
Usf1 M.musculus USF1 (non-coding exon 1). X95315 6.64978 1.88E-09
Grin3b Glutamate receptor, ionotropic, NMDA3B (Grin3b), mRNA NM_130455 6.64959 3.38E-22
A730016F12Rik Mus musculus RIKEN cDNA A730016F12 gene (A730016F12Rik), transcript variant 2, mRNA [NM_175079] NM_175079 6.64561 1.08E-35
Casz1 MKIAA3026 protein AK010559 6.63118 9.68E-36
Nckap1l NCK associated protein 1 like (Nckap1l), mRNA NM_153505 6.62986 7.63E-07
AI324046 Immunoglobulin heavy chain 6 (heavy chain of IgM), mRNA (cDNA clone MGC:18788 IMAGE:4189350) NM_198640 6.62565 0.00096
9530001N24Rik Peroxiredoxin 1 (Prdx1), mRNA AK020533 6.60763 2.92E-11
4732496G21Rik RIKEN cDNA 4732496G21 gene (4732496G21Rik), mRNA AK035094 6.59781 0.00384
Evpl Envoplakin, mRNA (cDNA clone IMAGE:3985477) AK007353 6.59541 1.14E-40
Postn Periostin, osteoblast specific factor, mRNA (cDNA clone MGC:25368 IMAGE:4457222) NM_015784 6.59457 2.74E-12
Lyzs Lysozyme, mRNA (cDNA clone IMAGE:2655292) NM_017372 6.58874 0.00173
Apol2 PREDICTED: hypothetical protein XP_139463 [Mus musculus], mRNA sequence AK036408 6.56783 8.71E-25
Paqr3 Progestin and adipoQ receptor family member III (Paqr3), mRNA NM_198422 6.56779 5.81E-14
4833421E05Rik RIKEN cDNA 4833421E05 gene (4833421E05Rik), mRNA AK014743 6.56384 7.26E-13
4930519L02Rik PREDICTED: hypothetical protein LOC75085 [Mus musculus], mRNA sequence AK015845 6.55752 0.00113
Hrbl HIV-1 Rev binding protein-like (Hrbl), transcript variant 1, mRNA NM_178162 6.55743 8.29E-31
Bai2 Brain-specific angiogenesis inhibitor 2, mRNA (cDNA clone MGC:66520 IMAGE:5705854) NM_173071 6.5429 2.54E-10
Map2k6 MAP Kinase Kinase NM_011943 6.53525 2.77E-11
Ttll1 Mus musculus tubulin tyrosine ligase-like 1 (Ttll1), mRNA. NM_178869 6.52155 6.57E-30
Hand2 BHLH transcription factor dHand NM_010402 6.51319 0.00064
Cpeb4 Cytoplasmic polyadenylation element binding protein 4 (Cpeb4), mRNA AK173229 6.50992 7.59E-11
C920005C14Rik RIKEN cDNA C920005C14 gene (C920005C14Rik), mRNA NM_177391 6.49949 0.0012
Fbp2 Fructose bisphosphatase 2, mRNA (cDNA clone MGC:14060 IMAGE:4205515) NM_007994 6.49091 1.69E-19
AK042798 Mus musculus 7 days neonate cerebellum cDNA, RIKEN full-length enriched library, clone:A730025J02 product: unclassifiable, full insert sequence. AK042798 6.47491 0.00001
D930005D10Rik MKIAA1849 protein NM_178702 6.4664 3.36E-17
Irf6 Interferon regulatory factor 6, mRNA (cDNA clone MGC:5918 IMAGE:3592582) NM_016851 6.46367 8.31E-13
A530064D06Rik RIKEN cDNA A530064D06 gene (A530064D06Rik), mRNA NM_178796 6.46325 0.00093
Fbxl16 PREDICTED: similar to spinal cord injury and regeneration related protein 1 [Mus musculus], mRNA sequence XM_128530 6.46071 5.26E-11
1700013F07Rik PREDICTED: hypothetical protein LOC75504 [Mus musculus], mRNA sequence AK005946 6.44623 1.24E-23
Cdh15 Cadherin 15 (Cdh15), mRNA NM_007662 6.44407 1.52E-22
AK033995 Mus musculus adult male diencephalon cDNA, RIKEN full-length enriched library, clone:9330137C05 product: unclassifiable, full insert sequence. AK033995 6.44198 2.74E-11
Slc36a2 Solute carrier family 36 (proton/amino acid symporter), member 2 (Slc36a2), mRNA NM_153170 6.43572 4.47E-18
Agt Angiotensinogen (serpin peptidase inhibitor, clade A, member 8), mRNA (cDNA clone MGC:19077 IMAGE:4194190) NM_007428 6.43479 0.00014
4921513E08Rik RIKEN cDNA 4921513E08 gene (4921513E08Rik), mRNA NM_025725 6.42815 1.00E-30
Hey2 Basic helix-loop-helix factor 1 (CHF1) AK048343 6.425 0.0015
Krtap6-1 Keratin associated protein 6-1 (Krtap6-1), mRNA D86421 6.42422 0.0014
AK046930 10 days neonate cerebellum cDNA, RIKEN full-length enriched library, clone:B930004K07 product: histocompatibility 2, T region locus 18, full insert sequence AK046930 6.42302 4.29E-12
Rtf1 Gene trap locus 7, mRNA (cDNA clone IMAGE:6740526) XM_283757 6.42082 3.63E-18
Rnf122 Ring finger protein 122, mRNA (cDNA clone MGC:36976 IMAGE:4947596) NM_175136 6.40147 0.00007
5730508B09Rik RIKEN cDNA 5730508B09 gene (5730508B09Rik), mRNA AK017758 6.39187 2.05E-39
D230014K01Rik RIKEN cDNA D230014K01 gene (D230014K01Rik), mRNA NM_172573 6.38975 1.69E-06
Spdef SAM pointed domain containing ets transcription factor, mRNA (cDNA clone MGC:13763 IMAGE:4218741) NM_013891 6.37865 8.43E-08
Adc Arginine decarboxylase (Adc), mRNA NM_172875 6.37788 6.03E-36
Adora2b Adenosine A2b receptor (Adora2b), mRNA NM_007413 6.37729 8.30E-10
TC949982 AF465729 cyclin fold protein 1 variant b {Homo sapiens}, partial (83%) [TC949982] 6.36781 5.12E-29
Ccdc37 RIKEN cDNA C230069K22 gene, mRNA (cDNA clone MGC:90884 IMAGE:5693354) NM_173775 6.36741 4.66E-17
Epas1 Endothelial PAS domain protein 1 (Epas1), mRNA AK087208 6.35043 1.62E-18
9630020I17Rik Adult male aorta and vein cDNA, RIKEN full-length enriched library, clone:A530095L03 product: unclassifiable, full insert sequence AK079318 6.34776 3.78E-06
ENSMUST00000055324 OLFACTORY RECEPTOR GA_X6K02T2PSCP-1637650–1636712 (FRAGMENT). [Source:SPTREMBL;Acc:Q7TRL0] [ENSMUST00000055324] 6.3429 0.00019
AI646023 Expressed sequence AI646023 (AI646023), mRNA NM_198860 6.3422 1.37E-12
Ptpn12 Protein tyrosine phosphatase, non-receptor type 12 (Ptpn12), mRNA AK087609 6.32927 0.00114
Wdr40b WD repeat domain 40B, mRNA (cDNA clone MGC:76797 IMAGE:30431243) NM_178739 6.32111 0.00162
Rap2ip Rap2 interacting protein (Rap2ip), mRNA AK007019 6.3115 4.38E-17
6230427J02Rik RIKEN cDNA 6230427J02 gene (6230427J02Rik), mRNA AK020090 6.30412 4.72E-38
5730420B22Rik RIKEN cDNA 5730420B22 gene, mRNA (cDNA clone IMAGE:5356411) NM_172597 6.29909 2.99E-13
Zhx1 Zinc fingers and homeoboxes protein 1 (Zhx1), mRNA AK051410 6.29396 3.18E-08
Lmtk2 Lemur tyrosine kinase 2, mRNA (cDNA clone IMAGE:6407188) BC058653 6.2919 1.25E-23
2410003J06Rik RIKEN cDNA 2410003J06 gene, mRNA (cDNA clone MGC:60956 IMAGE:30007513) AK010362 6.2909 1.27E-08
Dgat2 Diacylglycerol O-acyltransferase 2 (Dgat2), mRNA NM_026384 6.28587 3.72E-19
Rab3d RAB3D, member RAS oncogene family, mRNA (cDNA clone MGC:6707 IMAGE:3584902) NM_031874 6.28515 3.96E-08
2610031L17Rik RIKEN cDNA 2610031L17 gene, mRNA (cDNA clone MGC:38351 IMAGE:5344332) AK043897 6.2836 0.00003
Jmjd2d Hypothetical protein 4932416A15 (4932416A15), mRNA NM_173433 6.27488 2.19E-07
1700023A16Rik PREDICTED: hypothetical protein LOC69371 [Mus musculus], mRNA sequence AK006257 6.27378 2.95E-13
Tmcc3 Transmembrane and coiled coil domains 3, mRNA (cDNA clone MGC:30586 IMAGE:3660220) NM_172051 6.27294 9.85E-06
Clec2e C-type lectin domain family 2, member e (Clec2e), mRNA NM_153506 6.27057 0.0012
H2-DMa Histocompatibility 2, class II, locus DMa, mRNA (cDNA clone MGC:5743 IMAGE:3591389) NM_010386 6.26974 1.31E-33
Habp4 Hyaluronic acid binding protein 4 (Habp4), mRNA NM_019986 6.26891 6.44E-11
LOC434018 hypothetical gene supported by AK029757 AK029757 6.2591 0.0016
2700038N03Rik RIKEN cDNA 2700038N03 gene, mRNA (cDNA clone MGC:27927 IMAGE:3584484) NM_027356 6.2369 1.38E-30
3526402J09Rik RIKEN cDNA 3526402J09 gene, mRNA (cDNA clone IMAGE:30544810) AK028312 6.23264 3.49E-06
Dlgap1 Discs, large (Drosophila) homolog-associated protein 1 (Dlgap1), transcript variant 2, mRNA AK034215 6.21769 1.54E-06
9230116B18Rik PREDICTED: hypothetical protein LOC78245 [Mus musculus], mRNA sequence AK020348 6.21758 3.02E-09
H2-Ke6 H2-K region expressed gene 6 (H2-Ke6), mRNA NM_013543 6.21346 6.39E-38
Prdx1 Peroxiredoxin 1 (Prdx1), mRNA AK083642 6.21182 3.67E-17
4930550L24Rik Mus musculus RIKEN cDNA 4930550L24 gene (4930550L24Rik), mRNA [NM_023774] NM_023774 6.21139 6.89E-09
Trim62 Tripartite motif-containing 62 (Trim62), mRNA NM_178110 6.20606 8.48E-07
NAP044566-1 Unknown 6.20486 9.69E-13
Brsk2 BR serine/threonine kinase 2 (Brsk2), transcript variant 2, mRNA BC056498 6.18857 0.00023
9630019E01Rik Mus musculus 12 days embryo spinal ganglion cDNA, RIKEN full-length enriched library, clone:D130045C05 product: unclassifiable, full insert sequence. AK051384 6.17974 2.17E-15
Itga6 Mus musculus adult male corpora quadrigemina cDNA, RIKEN full-length enriched library, clone:B230112G01 product: integrin alpha 6, full insert sequence. AK045391 6.17009 2.44E-07
Acoxl Acyl-Coenzyme A oxidase-like (Acoxl), mRNA AV282483 6.16545 1.07E-07
C030014L02 hypothetical protein C030014L02 AK047704 6.15908 0.00132
4921515A04Rik Mus musculus RIKEN cDNA 4921515A04 gene (4921515A04Rik), mRNA [NM_172939] NM_172939 6.15587 2.02E-16
Frat1 Frequently rearranged in advanced T-cell lymphomas (Frat1), mRNA NM_008043 6.14419 9.07E-08
D5Wsu178e DNA segment, Chr 5, Wayne State University 178, expressed (D5Wsu178e), mRNA AK046608 6.14236 5.29E-09
AK012034 Mus musculus 10 days embryo whole body cDNA, RIKEN full-length enriched library, clone:2610315N17 product: unclassifiable, full insert sequence. AK012034 6.1156 2.54E-08
Tax1bp1 Tax1 (human T-cell leukemia virus type I) binding protein 1, mRNA (cDNA clone MGC:11692 IMAGE:3962810) AK051030 6.11319 0.00526
Syce1 PREDICTED: Mus musculus RIKEN cDNA 4933406J07 gene (4933406J07Rik), mRNA AK016694 6.1065 1.56E-15
A430091O22Rik RIKEN cDNA A430091O22 gene (A430091O22Rik), mRNA AK039072 6.10372 0.00337
Scrib Scribbled homolog (Drosophila), mRNA (cDNA clone IMAGE:4459388) AK084936 6.1006 8.84E-14
Rhov Ras homolog gene family, member V (Rhov), mRNA NM_145530 6.0999 1.54E-23
AK040776 Adult male aorta and vein cDNA, RIKEN full-length enriched library, clone:A530024I17 product: unclassifiable, full insert sequence AK040776 6.09753 1.78E-38
D330014H01Rik Mus musculus RIKEN cDNA D330014H01 gene (D330014H01Rik), mRNA [NM_177617] NM_177617 6.08754 1.59E-14
D0H4S114 DNA segment, human D4S114 (D0H4S114), mRNA NM_053078 6.08447 0.00133
Herc1 Hect (homologous to the E6-AP (UBE3A) carboxyl terminus) domain and RCC1 (CHC1)-like domain (RLD) 1, mRNA (cDNA clone MGC:7618 IMAGE:3494753) AK083444 6.08322 4.90E-24
1700065D16Rik Protein kinase ATR (Atr) AK030118 6.07886 0.00004
Ppp1r13b Protein phosphatase 1, regulatory (inhibitor) subunit 13B (Ppp1r13b), mRNA NM_011625 6.0768 9.44E-25
LOC213402 hypothetical LOC213402 XM_136978 6.07669 6.10E-19
Capns2 Hypothetical protein A330042H22 (A330042H22), mRNA AK009171 6.0641 6.40E-07
NM_027559 Mus musculus RIKEN cDNA 4930463G05 gene (4930463G05Rik), mRNA [NM_027559] NM_027559 6.05992 8.61E-10
2810427I04Rik RIKEN cDNA 2810427I04 gene (2810427I04Rik), mRNA NM_028146 6.05072 3.51E-08
Mycbp C-myc binding protein, mRNA (cDNA clone MGC:13820 IMAGE:3985166) NM_019660 6.0454 3.53E-09
D130076A03Rik Mus musculus 12 days embryo spinal ganglion cDNA, RIKEN full-length enriched library, clone:D130076A03 product: unclassifiable, full insert sequence. AK084007 6.04495 1.01E-16
AA919669 Adult male hippocampus cDNA, RIKEN full-length enriched library, clone:2900016J10 product: unclassifiable, full insert sequence AA919669 6.04404 1.08E-19
Ifrd1 Interferon-related developmental regulator 1 (Ifrd1), mRNA NM_013562 6.04347 2.10E-09
Ddit4l DNA-damage-inducible transcript 4-like, mRNA (cDNA clone MGC:47974 IMAGE:5254530) NM_030143 6.04198 0.00184
Gsdm1 Gasdermin 1 (Gsdm1), mRNA NM_021347 6.04098 2.79E-13
3110057O12Rik RIKEN cDNA 3110057O12 gene, mRNA (cDNA clone IMAGE:4007705) NM_026622 6.04095 1.90E-12
Pdpk1 Phosphoinositide-dependent protein kinase-1 beta (Pdk1beta) NM_011062 6.03733 3.77E-31
Zp3 Zona pellucida glycoprotein 3 (Zp3), mRNA NM_011776 6.02738 1.28E-12
TC984987 NDK6_HUMAN Nucleoside diphosphate kinase 6(NDK 6) (NDP kinase 6) (nm23-H6) (Inhibitor of p53-induced apoptosis-alpha) (IPIA-alpha). [Human] {Homo sapiens}, partial (14%) [TC984987] 6.02678 0.00023
1700011E04Rik PREDICTED: Mus musculus RIKEN cDNA 1700011E04 gene (1700011E04Rik), mRNA AK005860 6.02661 6.11E-06
Sstr2 Somatostatin receptor 2 (Sstr2), mRNA NM_009217 6.0182 1.48E-11
Pgm2l1 Phosphoglucomutase 2-like 1, mRNA (cDNA clone MGC:29299 IMAGE:5004308) NM_027629 6.01695 1.89E-15
BE646931 Transcribed locus BE646931 6.00062 1.38E-15
Ccdc37 ZXD family zinc finger C, mRNA (cDNA clone MGC:7160 IMAGE:3256958) AK021139 5.99479 5.23E-06
Timd4 T-cell immunoglobulin and mucin domain containing 4, mRNA (cDNA clone MGC:41117 IMAGE:1195144) NM_178759 5.98633 0.00165
Ap3m2 Adaptor-related protein complex 3, mu 2 subunit, mRNA (cDNA clone MGC:40858 IMAGE:5369288) NM_029505 5.97689 1.24E-08
Htr2b 5-hydroxytryptamine (serotonin) receptor 2B, mRNA (cDNA clone MGC:36132 IMAGE:5344328) NM_008311 5.97448 0.00018
Tfdp2 DP-3=protein regulating cell cycle transcription factor DRTF1/E2F [mice, pl-2, F9 EC, mRNA, 1380 nt]. S79780 5.97116 4.82E-07
Rybp RING1 and YY1 binding protein (Rybp), mRNA NM_019743 5.97057 9.26E-38
Sema3b Sema domain, immunoglobulin domain (Ig), short basic domain, secreted, (semaphorin) 3B (Sema3b), mRNA NM_009153 5.9703 2.43E-22
Dcn Decorin (Dcn), mRNA NM_007833 5.96766 2.08E-06
Lhx2 LIM-homeodomain protein MLHX2 (Lhx2) NM_010710 5.96324 1.93E-11
Fbxo33 F-box only protein 33, mRNA (cDNA clone IMAGE:3661715) BC020022 5.96152 1.82E-17
Cfh Complement component factor h (Cfh), mRNA M12660 5.95652 0.00006
Psd2 Pleckstrin and Sec7 domain containing 2 (Psd2), mRNA NM_028707 5.95334 6.23E-14
Ncoa1 Nuclear receptor coactivator 1 (Ncoa1), mRNA AA738860 5.94761 1.72E-25
Robo2 MKIAA1568 protein BC055333 5.94552 0.00019
AW125753 Expressed sequence AW125753 (AW125753), mRNA NM_029007 5.94357 1.90E-15
Cfh Complement component factor h (Cfh), mRNA NM_009888 5.94161 6.84E-07
1700014D04Rik PREDICTED: hypothetical protein LOC74224 [Mus musculus], mRNA sequence AK077007 5.93902 9.40E-15
6-Sep Mus musculus 2 days neonate thymus thymic cells cDNA, RIKEN full-length enriched library, clone:C920001C06 product: hypothetical protein, full insert sequence. AK083302 5.93827 0.00283
Pdpk1 Phosphoinositide-dependent protein kinase-1 beta (Pdk1beta) NM_011062 5.93442 2.75E-36
BC048651 CDNA sequence BC048651 (BC048651), mRNA NM_207258 5.93022 1.19E-13
Metrnl Meteorin, glial cell differentiation regulator-like (Metrnl), mRNA AK053718 5.9294 3.74E-21
LOC229608 Histone 2, H2bb (Hist2h2bb), mRNA BC019122 5.92838 4.58E-08
Arl5b ADP-ribosylation factor-like 8 (Arl8), mRNA NM_029466 5.92634 1.03E-09
Olfr987 Olfactory receptor 987 (Olfr987), mRNA CB174485 5.92041 0.00266
Malat1 Adult male stomach cDNA, RIKEN full-length enriched library, clone:2210407K09 product: receptor (calcitonin) activity modifying protein 2, full insert sequence AK020483 5.91958 6.65E-15
Mall CDNA sequence BC012256 (BC012256), mRNA NM_145532 5.91692 8.22E-08
Phtf2 Putative homeodomain transcription factor 2, mRNA (cDNA clone MGC:60851 IMAGE:30050876) NM_172992 5.91568 1.02E-30
D330028D13Rik RIKEN cDNA D330028D13 gene, mRNA (cDNA clone MGC:67688 IMAGE:4487036) NM_172727 5.91051 1.54E-22
Rhpn2 Rhophilin, Rho GTPase binding protein 2 (Rhpn2), mRNA NM_027897 5.90122 2.02E-22
3526402J09Rik 8 days embryo whole body cDNA, RIKEN full-length enriched library, clone:5730590C14 product: unclassifiable, full insert sequence AK019982 5.89752 1.80E-24
Tigd5 Tigger transposable element derived 5 (Tigd5), mRNA NM_178646 5.89512 9.40E-16
C230078M08Rik RIKEN cDNA C230078M08 gene (C230078M08Rik), mRNA NM_176995 5.88686 6.61E-06
D930001I22Rik RIKEN cDNA D930001I22 gene, mRNA (cDNA clone MGC:107602 IMAGE:6750721) AK052945 5.86474 5.89E-43
Tcf15 Transcription factor 15, mRNA (cDNA clone MGC:41210 IMAGE:5143248) NM_009328 5.86455 2.79E-14
Wif1 Wnt inhibitory factor 1, mRNA (cDNA clone MGC:13706 IMAGE:3984128) NM_011915 5.85749 1.02E-06
Egfl5 MKIAA0818 protein AK122378 5.85619 1.22E-10
Fcer1g Fc receptor, IgE, high affinity I, gamma polypeptide, mRNA (cDNA clone MGC:36077 IMAGE:5065647) AI326608 5.85071 2.00E-10
Mtap2 Mus musculus microtubule-associated protein 2 (Mtap2), mRNA [NM_008632] NM_008632 5.85052 5.98E-13
Uros Uroporphyrinogen III synthase (Uros), mRNA NM_009479 5.84947 1.39E-29
3110098I04Rik Transmembrane protein 49, mRNA (cDNA clone MGC:7590 IMAGE:3493738) AK019443 5.84922 1.23E-18
Snai3 Snail-related zinc finger protein SMUC (Smuc) NM_013914 5.84401 6.78E-12
Arzc Mus musculus hypothetical protein (Arzc) mRNA, complete cds. AY344585 5.83844 1.81E-12
BE987854 Bone marrow macrophage cDNA, RIKEN full-length enriched library, clone:I830003K21 product: unclassifiable, full insert sequence BE987854 5.8367 3.15E-09
Npc1l1 Mus musculus NPC1-like 1 (Npc1l1), mRNA. XM_137497 5.83041 2.18E-14
Etv1 Ets variant gene 1, mRNA (cDNA clone MGC:6207 IMAGE:3257346) NM_007960 5.82656 0.00003
AK041974 3 days neonate thymus cDNA, RIKEN full-length enriched library, clone:A630050C01 product: unclassifiable, full insert sequence AK041974 5.82616 0.00025
Atp11b ATPase, Class VI, type 11B (Atp11b), mRNA AK041684 5.82607 3.61E-08
Pgm2l1 Phosphoglucomutase 2-like 1, mRNA (cDNA clone MGC:29299 IMAGE:5004308) NM_027629 5.81964 0.0001
Ppp1r13b Protein phosphatase 1, regulatory (inhibitor) subunit 13B (Ppp1r13b), mRNA NM_011625 5.81761 4.07E-12
2310068J10Rik Adult male tongue cDNA, RIKEN full-length enriched library, clone:2310068J10 product: unclassifiable, full insert sequence AK075917 5.81515 5.55E-09
Nos3 Nitric oxide synthase 3, endothelial cell (Nos3), mRNA NM_008713 5.80909 1.30E-07
Spsb1 SplA/ryanodine receptor domain and SOCS box containing 1 (Spsb1), mRNA NM_029035 5.80869 5.16E-13
8430416H19Rik RIKEN cDNA 8430416H19 gene (8430416H19Rik), mRNA NM_198636 5.80743 1.09E-11
Gpr111 PREDICTED: similar to G-protein coupled receptor 111 [Mus musculus], mRNA sequence AY255579 5.79653 0.00225
Pla2g4f RIKEN cDNA 4732472I07 gene (4732472I07Rik), mRNA BC039947 5.7812 5.59E-15
Rfx2 Regulatory factor X, 2 (influences HLA class II expression), mRNA (cDNA clone MGC:6105 IMAGE:3497910) NM_009056 5.77832 1.13E-11
Dpf3 D4, zinc and double PHD fingers, family 3 (Dpf3), mRNA NM_058212 5.77719 9.83E-15
Zfp654 Zinc finger protein 654, mRNA (cDNA clone IMAGE:5346461) NM_028059 5.77371 1.78E-35
Rnut1 RNA, U transporter 1, mRNA (cDNA clone MGC:74414 IMAGE:30251661) NM_178374 5.76909 7.92E-16
Mpp7 Membrane protein, palmitoylated 7 (MAGUK p55 subfamily member 7), mRNA (cDNA clone IMAGE:6532780) AK017344 5.76872 3.39E-11
Cx3cl1 Mus musculus mRNA, complete cds, clone:1–53. AB030188 5.75894 0.00037
Map2k6 Mus musculus 15 days embryo head cDNA, RIKEN full-length enriched library, clone:D930047J04 product: mitogen activated protein kinase kinase 6, full insert sequence. AK086722 5.74548 2.97E-08
Gsto2 Glutathione S-transferase omega 2 (Gsto2), mRNA NM_026619 5.73514 3.96E-07
A030009H04Rik RIKEN cDNA A030009H04 gene, mRNA (cDNA clone MGC:35887 IMAGE:5352453) NM_020591 5.73374 2.63E-12
Atox1 ATX1 (antioxidant protein 1) homolog 1 (yeast), mRNA (cDNA clone MGC:41183 IMAGE:1328907) AK089832 5.73109 2.04E-11
Map2k6 MAP Kinase Kinase NM_011943 5.72903 0.00035
Bmp2k BMP2 inducible kinase (Bmp2k), mRNA NM_080708 5.72563 2.28E-11
St3gal5 ST3 beta-galactoside alpha-2,3-sialyltransferase 5 (St3gal5), mRNA AI464789 5.72266 0.0001
Rasd1 RAS, dexamethasone-induced 1, mRNA (cDNA clone MGC:36188 IMAGE:4989312) NM_009026 5.71641 9.16E-11
C330001K17Rik RIKEN cDNA C330001K17 gene (C330001K17Rik), mRNA AK031175 5.70009 1.16E-08
Olig2 Olig2 bHLH protein BC051967 5.69272 0.00135
Zfp422-rs1 zinc finger protein 422, related sequence 1 NM_029952 5.68877 7.93E-14
Ccni Cyclin I, mRNA (cDNA clone MGC:5636 IMAGE:3583418) AK083496 5.68864 0.00337
AK028719 10 days neonate skin cDNA, RIKEN full-length enriched library, clone:4732444B01 product: unclassifiable, full insert sequence AK028719 5.68121 0.00073
5730469D23Rik RIKEN cDNA 5730469D23 gene, mRNA (cDNA clone MGC:66555 IMAGE:6825308) AK017687 5.67016 6.96E-16
Tspan1 Tetraspan 1 (Tspan1), mRNA NM_133681 5.66843 7.35E-08
Ebf1 Early B-cell factor 1 (Ebf1), mRNA AK036716 5.66713 0.00329
Susd2 Sushi domain containing 2 (Susd2), mRNA NM_027890 5.66242 2.96E-08
XM_131928 Mus musculus myeloid/lymphoid or mixed-lineage leukemia 3 (Mll3), mRNA [XM_131928] XM_131928 5.65966 0.00033
Trfr2 Transferrin receptor 2, mRNA (cDNA clone MGC:18814 IMAGE:4196597) NM_015799 5.65881 2.39E-07
Tspyl4 Mus musculus TSPY-like 4 (Tspyl4), mRNA. NM_133745 5.65493 1.21E-11
Hgf Hepatocyte growth factor AK082461 5.65072 0.00111
Txnip Thioredoxin interacting protein, mRNA (cDNA clone MGC:25534 IMAGE:3591421) AK004653 5.6496 1.05E-06
Angptl2 Angiopoietin-like 2 (Angptl2), mRNA AK035967 5.64678 9.32E-12
Dos Downstream of Stk11 (Dos), mRNA AK035824 5.6364 2.23E-11
Mars2 Methionine-tRNA synthetase 2 (mitochondrial), mRNA (cDNA clone MGC:106216 IMAGE:3153138) NM_175439 5.62867 1.35E-27
Hamp1 Prohepcidin (hepc1) NM_032541 5.62597 1.61E-09
BC049806 CDNA sequence BC049806, mRNA (cDNA clone MGC:76460 IMAGE:30431671) NM_172513 5.62361 6.88E-34
Sbsn suprabasin NM_172205 5.62177 3.83E-18
FHOS2 formin-family protein FHOS2 NM_175276 5.62016 0.00001
Als2 Amyotrophic lateral sclerosis 2 (juvenile) homolog (human), mRNA (cDNA clone MGC:27807 IMAGE:3257574) NM_028717 5.61726 6.41E-30
5330420D20Rik RIKEN cDNA 5330420D20 gene (5330420D20Rik), mRNA NM_027442 5.61363 0.00005
Gdap5 Dishevelled associated activator of morphogenesis 1 (Daam1), mRNA Y17854 5.61177 1.02E-26
E030003N13Rik Fibroblast growth factor receptor substrate 2, mRNA (cDNA clone MGC:64739 IMAGE:6830555) AK051267 5.61074 0.00191
Zfp59 Zinc finger protein 59 (Zfp59), mRNA AK042484 5.60473 0.00217
Rnd1 Rho family GTPase 1, mRNA (cDNA clone MGC:58446 IMAGE:6535763) BC048531 5.60436 2.78E-39
Ypel4 Ypel4 mRNA for yippee-like 4 AK043577 5.60118 0.00005
Hbld2 HESB like domain containing 2 (Hbld2), mRNA NM_026921 5.59811 1.03E-15
Plekhk1 Membrane-bound factor MBF1 (Mbf) AY036116 5.59772 2.32E-19
Asb11 Mus musculus adult male colon cDNA, RIKEN full-length enriched library, clone:9030022F06 product: ankyrin repeat and SOCS box-containing protein 11, full insert sequence. AK078851 5.59544 0.00382
TC996139 Unknown 5.59089 1.46E-07
Bai2 Brain-specific angiogenesis inhibitor 2, mRNA (cDNA clone MGC:66520 IMAGE:5705854) NM_173071 5.57838 0.00002
Rnf17 Ring finger protein 17 (Rnf17), transcript variant 1, mRNA NM_013894 5.57648 0.00408
Scoc Short coiled-coil protein, mRNA (cDNA clone MGC:28845 IMAGE:4506964) NM_019708 5.57413 1.02E-21
Lama3 Laminin-5, alpha3B chain X84014 5.56979 1.47E-30
Bmp6 Bone morphogenetic protein 6 (Bmp6), mRNA NM_007556 5.5686 6.55E-09
Dffa DNA fragmentation factor, alpha subunit (Dffa), transcript variant 2, mRNA AK051011 5.55806 2.27E-19
Nfatc4 Nuclear factor of activated T-cells, cytoplasmic, calcineurin-dependent 4, mRNA (cDNA clone MGC:25538 IMAGE:3670623) AK014164 5.55789 1.20E-09
Klc3 Kinesin light chain 3 (Klc3), mRNA NM_146182 5.55558 2.85E-09
Kif17 Kinesin family member 17 (Kif17), mRNA NM_010623 5.5494 0.00022
Btbd10 BTB (POZ) domain containing 10, mRNA (cDNA clone IMAGE:3484902) AK052335 5.54822 4.06E-10
Slc25a35 Solute carrier family 25, member 35 (Slc25a35), mRNA NM_028048 5.54317 9.37E-07
AK076567 Adult male testis cDNA, RIKEN full-length enriched library, clone:4921513I01 product: unclassifiable, full insert sequence AK076567 5.54169 4.56E-17
Kif17 Kinesin family member 17 (Kif17), mRNA NM_010623 5.53068 7.65E-13
Trpt1 TRNA phosphotransferase 1 (Trpt1), mRNA NM_153597 5.52301 1.26E-44
C030032O16Rik Putative homeodomain transcription factor 1 (Phtf1), mRNA AK021115 5.52287 0.00004
NAP050738-1 Unknown 5.51793 0.00143
Slc40a1 Solute carrier family 40 (iron-regulated transporter), member 1, mRNA (cDNA clone MGC:6489 IMAGE:2647365) NM_016917 5.51526 5.26E-10
Mll3 Mixed-lineage leukemia 3 protein (Mll3) XM_355579 5.51391 1.75E-19
Ablim3 Actin binding LIM protein family, member 3, mRNA (cDNA clone MGC:106450 IMAGE:30534236) NM_198649 5.51387 4.79E-12
Dmxl1 Dmx-like 1, mRNA (cDNA clone MGC:28386 IMAGE:4022096) AK082851 5.51375 9.31E-09
Hivep3 Clone 5′RACE1 ZAS3 (Krc) mRNA, 5′ UTR AK013707 5.51275 3.18E-26
Fhl2 Four and a half LIM domains 2, mRNA (cDNA clone MGC:29060 IMAGE:5066565) AK081770 5.51184 0.00028
Arl5b ADP-ribosylation factor-like 8 (Arl8), mRNA NM_029466 5.50628 6.74E-06
Usp53 Ubiquitin specific peptidase 53, mRNA (cDNA clone IMAGE:4236151) AK030317 5.5048 6.73E-16
Eaf2 Testosterone regulated apoptosis inducer and tumor suppressor (Traits) NM_134111 5.49779 6.28E-07
Lmbr1 Limb region 1, mRNA (cDNA clone MGC:28737 IMAGE:4481088) AK030631 5.49689 1.41E-08
F730023N20 hypothetical protein F730023N20 NM_183158 5.48167 0.00469
Vamp1 Vesicle-associated membrane protein 1 (Vamp1), mRNA NM_009496 5.47901 7.04E-22
Abcc10 ATP-binding cassette, sub-family C (CFTR/MRP), member 10 (Abcc10), transcript variant mrp7B, mRNA NM_145140 5.47864 1.39E-10
BG080473 Transcribed locus BG080473 5.47538 0.00524
Ppp1r9a Protein phosphatase 1, regulatory (inhibitor) subunit 9A, mRNA (cDNA clone MGC:59493 IMAGE:6331919) NM_181595 5.47417 4.82E-07
TC1012618 RGS2_MOUSE Regulator of G-protein signaling 2 (RGS2). [Mouse] {Mus musculus}, partial (43%) [TC1012618] 5.46503 6.76E-10
Smarcad1 MRNA of enhancer-trap-locus 1 AK088163 5.4628 3.56E-06
Malat1 Adult male stomach cDNA, RIKEN full-length enriched library, clone:2210407K09 product: receptor (calcitonin) activity modifying protein 2, full insert sequence BC004722 5.46155 1.11E-09
Btg2 B-cell translocation gene 2, anti-proliferative (Btg2), mRNA NM_007570 5.46006 1.64E-08
Malat1 Adult male stomach cDNA, RIKEN full-length enriched library, clone:2210407K09 product: receptor (calcitonin) activity modifying protein 2, full insert sequence AK090111 5.45534 2.16E-21
Kcnk7 Dpkch3 mRNA for double-pore K channel 3 AF110521 5.45419 0.00021
XM_358570 Mus musculus LOC381415 (LOC381415), mRNA [XM_358570] XM_358570 5.4508 0.00001
Tnrc15 Trinucleotide repeat containing 15 (Tnrc15), mRNA AK034426 5.44852 9.38E-14
D16H22S680E DNA segment, Chr 16, human D22S680E, expressed (D16H22S680E), mRNA NM_138583 5.4478 6.43E-27
Cxxc5 CXXC finger 5 (Cxxc5), mRNA NM_133687 5.44654 0.00002
Pbx4 Pre-B-cell leukemia transcription factor 4, mRNA (cDNA clone MGC:40990 IMAGE:1263363) NM_030555 5.4461 4.76E-13
Gm1418 PREDICTED: similar to Ig kappa chain precursor V region - mouse [Mus musculus], mRNA sequence J04610 5.44288 0.0003
Slc25a2 Mutant ornithine transporter 2 (Ornt2) AK077159 5.43766 3.03E-14
Hist3h2bb Histone 3, H2bb (Hist3h2bb), mRNA NM_206882 5.43583 1.97E-33
Ralgps2 Ral GEF with PH domain and SH3 binding motif 2 (Ralgps2), mRNA NM_023884 5.43465 0.0001
Lao1 L-amino acid oxidase 1 (Lao1), mRNA NM_133892 5.43168 0.00322
Hist1h2bm Histone 1, H2bm (Hist1h2bm), mRNA NM_178200 5.42961 1.25E-36
Zfp422-rs1 zinc finger protein 422, related sequence 1 AK020746 5.42824 3.08E-44
Spire1 Spire homolog 1 (Drosophila), mRNA (cDNA clone MGC:106458 IMAGE:30535423) NM_176832 5.42537 9.41E-07
Matn3 Matrilin 3 (Matn3), mRNA NM_010770 5.42402 0.00001
Zfp422-rs1 zinc finger protein 422, related sequence 1 NM_029952 5.42321 2.97E-17
Pcdh10 OL-protocadherin isoform (Pcadh10) mRNA, complete cds; alternatively spliced AK122503 5.42181 5.94E-28
NAP042178-1 Unknown 5.41306 8.92E-21
Ranbp6 RAN binding protein 6 (Ranbp6), mRNA AK083089 5.41102 0.00015
Myh3 PREDICTED: myosin, heavy polypeptide 3, skeletal muscle, embryonic [Mus musculus], mRNA sequence XM_354614 5.39493 3.95E-11
Vill Villin-like (Vill), mRNA U72681 5.39006 1.44E-17
Ephb3 Eph receptor B3, mRNA (cDNA clone MGC:18409 IMAGE:3673003) NM_010143 5.38235 1.15E-08
Xpr1 Xenotropic and polytropic murine retrovirus receptor (Xpr1) NM_011273 5.38141 1.81E-13
Bank1 B-cell scaffold protein with ankyrin repeats 1 (Bank1), mRNA AK041242 5.37887 0.00416
TC1066872 S3A1_HUMAN Splicing factor 3 subunit 1 (Spliceosome associated protein 114) (SAP 114) (SF3a120). [Human] {Homo sapiens}, partial (3%) [TC1066872] 5.3751 0.00537
Wdr37 MKIAA0982 protein AK044740 5.36656 0.00008
Mtap2 Microtubule-associated protein 2, mRNA (cDNA clone MGC:63400 IMAGE:6412797) BC052446 5.36496 7.87E-18
AB112350 NAPE-PLD mRNA for N-acyl-phosphatidylethanolamine-hydrolyzing phospholipase D NM_178728 5.36332 3.14E-16
Malat1 Adult male stomach cDNA, RIKEN full-length enriched library, clone:2210407K09 product: receptor (calcitonin) activity modifying protein 2, full insert sequence AK020134 5.36328 6.17E-11
AI464131 MKIAA1161 protein BC036141 5.36291 4.74E-15
H1f0 H1 histone family, member 0, mRNA (cDNA clone MGC:19309 IMAGE:4166167) NM_008197 5.35386 2.93E-07
4933404M02Rik RIKEN cDNA 4933404M02 gene (4933404M02Rik), mRNA AK017113 5.35281 0.00005
Adarb1 Adenosine deaminase, RNA-specific, B1 (Adarb1), transcript variant 1, mRNA AF403109 5.3483 1.65E-15
Tcf7 Transcription factor 7, T-cell specific (Tcf7), mRNA NM_009331 5.34121 1.40E-08
Strbp Spermatid perinuclear RNA binding protein, mRNA (cDNA clone MGC:25337 IMAGE:4502840) AK081780 5.33831 0.00004
Foxj1 Forkhead Box j1 (Foxj1) NM_008240 5.33683 3.52E-08
Vill Villin-like (Vill), mRNA NM_011700 5.33523 2.17E-10
A_51_P170529 Unknown 5.32731 0.00598
2310047O13Rik RIKEN cDNA 2310047O13 gene, mRNA (cDNA clone MGC:28605 IMAGE:4217391) AK083925 5.32579 5.06E-32
Ccdc28a Coiled-coil domain containing 28A (Ccdc28a), mRNA NM_144820 5.32149 2.42E-10
NAP057224-1 Unknown 5.32046 8.05E-06
Arl5b ADP-ribosylation factor-like 8 (Arl8), mRNA NM_029466 5.31107 0.00004
Zfp40 Zinc finger protein 40 (Zfp40), mRNA NM_009555 5.30822 4.36E-10
Thnsl1 Threonine synthase-like 1 (bacterial), mRNA (cDNA clone IMAGE:5037172) NM_177588 5.30806 7.21E-20
4932411G14Rik RIKEN cDNA 4932411G14 gene (4932411G14Rik), mRNA NM_177711 5.30751 3.12E-11
Map3k14 Mitogen-activated protein kinase kinase kinase 14 (Map3k14), mRNA NM_016896 5.30704 5.21E-18
1700110N18Rik PREDICTED: hypothetical protein LOC73569 [Mus musculus], mRNA sequence XM_283372 5.3057 1.79E-09
ENSMUST00000050440 Unknown 5.30328 1.74E-27
Rapgef6 Rap guanine nucleotide exchange factor (GEF) 6, mRNA (cDNA clone IMAGE:5321951) AK040089 5.30322 2.62E-11
Arhgap26 MKIAA0621 protein AK129176 5.30199 0.00026
Wnt9a Wingless-type MMTV integration site 9A (Wnt9a), mRNA BC066165 5.3003 0.00003
Hist1h2ba Histone 1, H2ba (Hist1h2ba), mRNA NM_175663 5.29806 2.18E-27
Lincr lung-inducible neuralized-related C3HC4 RING domain protein NM_153408 5.29767 1.15E-07
B230105J10 hypothetical protein B230105J10 AK045359 5.29449 1.35E-06
TC984274 Unknown 5.29254 2.02E-17
5730553K21 5′-AMP-ACTIVATED PROTEIN KINASE, BETA-2 SUBUNIT (AMPK BETA-2 CHAIN) homolog [Rattus norvegicus] [5730553K21] 5.29004 4.53E-13
Nek1 PREDICTED: Mus musculus NIMA (never in mitosis gene a)-related expressed kinase 1 (Nek1), mRNA AK049625 5.28832 1.68E-17
AK086484 Mus musculus 15 days embryo head cDNA, RIKEN full-length enriched library, clone:D930031J16 product: microtubule-associated protein 2, full insert sequence. AK086484 5.28708 1.64E-12
Rb1 Retinoblastoma 1 (Rb1), mRNA NM_009029 5.27516 1.41E-07
Olfml3 Olfactomedin-like 3 (Olfml3), mRNA NM_133859 5.26799 3.38E-15
Etv3 ETS-domain transcriptional repressor PE1 (PE1) NM_012051 5.26705 2.17E-29
Ptpn13 Protein tyrosine phosphatase, non-receptor type 13 (Ptpn13), mRNA NM_011204 5.26698 1.95E-21
Tfdp2 Transcription factor Dp 2, mRNA (cDNA clone IMAGE:30604420) NM_178667 5.26621 2.69E-06
TC970961 AF169692 protocadherin-9 {Homo sapiens}, partial (9%) [TC970961] 5.2653 5.85E-08
Plekhk1 Membrane-bound factor MBF1 (Mbf) AY036116 5.25979 2.22E-16
Tmcc3 Transmembrane and coiled coil domains 3, mRNA (cDNA clone MGC:30586 IMAGE:3660220) AK083129 5.25708 2.98E-07
Prph1 Peripherin 1 (Prph1), mRNA NM_013639 5.25389 4.55E-08
Tbc1d1 TBC1 domain family, member 1, mRNA (cDNA clone IMAGE:3500261) NM_019636 5.2511 2.50E-15
Map3k1 MAP kinase kinase kinase 1 (Mekk1) NM_011945 5.25036 0.00004
1810053B23Rik 10 day old male pancreas cDNA, RIKEN full-length enriched library, clone:1810053B23 product: unclassifiable, full insert sequence AK007854 5.24933 0.00479
Ppp1r2 Protein phosphatase 1, regulatory (inhibitor) subunit 2 (Ppp1r2), mRNA AK037804 5.2492 8.39E-28
Rnf24 Ring finger protein 24, mRNA (cDNA clone MGC:106607 IMAGE:6406985) NM_178607 5.24766 3.12E-31
Bhlhb8 Basic helix-loop-helix domain containing, class B, 8, mRNA (cDNA clone MGC:19046 IMAGE:4189225) NM_010800 5.24479 8.19E-09
A230105L22Rik Cytochrome P450, family 4, subfamily f, polypeptide 18, mRNA (cDNA clone MGC:19144 IMAGE:4218398) AK020714 5.24224 0.00468
Herc1 Hect (homologous to the E6-AP (UBE3A) carboxyl terminus) domain and RCC1 (CHC1)-like domain (RLD) 1, mRNA (cDNA clone MGC:7618 IMAGE:3494753) BC004027 5.23497 5.02E-14
Upk3b Uroplakin 3B, mRNA (cDNA clone IMAGE:30297822) NM_175309 5.22777 0.00453
Pja2 Praja 2, RING-H2 motif containing (Pja2), transcript variant 2, mRNA AK122282 5.22623 2.64E-30
Usp2 Ubiquitin specific peptidase 2, mRNA (cDNA clone MGC:27630 IMAGE:4506362) NM_016808 5.22394 9.53E-08
Pak3 P21-activated kinase 3 (pak3 gene) NM_008778 5.22033 7.00E-06
0610008C08Rik RIKEN cDNA 0610008C08 gene (0610008C08Rik), mRNA AK029224 5.21977 0.00125
6330416L11Rik RIKEN cDNA 6330416L11 gene (6330416L11Rik), mRNA NM_027518 5.21346 3.01E-07
Homer1 Homer homolog 1 (Drosophila) (Homer1), transcript variant L, mRNA AK035139 5.20561 0.00057
Armc8 Armadillo repeat containing 8, mRNA (cDNA clone MGC:47965 IMAGE:1230321) AK078280 5.20529 6.30E-35
Lipg Lipase, endothelial, mRNA (cDNA clone MGC:13719 IMAGE:3981856) NM_010720 5.20391 0.00001
Dnahc2 Axonemal dynein heavy chain (partial, ID mdhc5) Z83813 5.20226 0.00081
Cotl1 Coactosin-like 1 (Dictyostelium) (Cotl1), mRNA NM_028071 5.20139 3.70E-07
2700050L05Rik RIKEN cDNA 2700050L05 gene (2700050L05Rik), transcript variant 1, mRNA NM_145995 5.19216 5.70E-15
Rab11fip4 MKIAA1821 protein AK122559 5.19174 0.00128
Eml5 Echinoderm microtubule associated protein like 5, mRNA (cDNA clone IMAGE:4953665) AK047762 5.19141 0.00003
Ptpn13 Protein tyrosine phosphatase, non-receptor type 13 (Ptpn13), mRNA NM_011204 5.19002 1.54E-26
3-Mar Membrane-associated ring finger (C3HC4) 3 (March3), mRNA NM_177115 5.18876 3.33E-22
Synpo2 Adult male urinary bladder cDNA, RIKEN full-length enriched library, clone:9530006G20 product: unclassifiable, full insert sequence AK035258 5.18793 9.93E-06
Zfp36 Zinc finger protein 36, mRNA (cDNA clone MGC:29129 IMAGE:5053664) NM_011756 5.18672 4.78E-06
Epb4.1l5 Erythrocyte protein band 4.1-like 5 (Epb4.1l5), mRNA NM_145506 5.18616 1.68E-43
Pip5k3 Phosphatidylinositol-3-phosphate/phosphatidylinositol 5-kinase, type III (Pip5k3), mRNA NM_011086 5.18443 1.80E-10
Tec Cytoplasmic tyrosine kinase, Dscr28C related (Drosophila), mRNA (cDNA clone MGC:46873 IMAGE:5029430) NM_013689 5.1779 7.22E-11
C130065N10Rik PREDICTED: hypothetical protein XP_488897 [Mus musculus], mRNA sequence AK048435 5.17772 1.71E-43
Pctk2 PCTAIRE-motif protein kinase 2 (Pctk2), mRNA NM_146239 5.17294 1.41E-12
A130019P10Rik Eukaryotic translation initiation factor 4E binding protein 2, mRNA (cDNA clone MGC:13950 IMAGE:4218820) AK037456 5.17158 7.10E-27
C920006C10Rik RIKEN cDNA C920006C10 gene (C920006C10Rik), mRNA AK052176 5.17083 1.37E-21
Camkk1 Calcium/calmodulin-dependent protein kinase kinase 1, alpha, mRNA (cDNA clone MGC:27706 IMAGE:4924656) NM_018883 5.17034 0.0001
Tusc2 Tumor suppressor candidate 2 (Tusc2), mRNA NM_019742 5.16901 1.18E-25
AK048541 Mus musculus 16 days embryo head cDNA, RIKEN full-length enriched library, clone:C130071D07 product: ATPase, Cu++ transporting, alpha polypeptide, full insert sequence. AK048541 5.16848 1.68E-13
Golph3 Mus musculus 10 days neonate skin cDNA, RIKEN full-length enriched library, clone:4733401N08 product: golgi phosphoprotein 3, full insert sequence. AK014644 5.16802 3.09E-33
Fa2h Fatty acid 2-hydroxylase, mRNA (cDNA clone IMAGE:4983638) NM_178086 5.16055 2.45E-15
Sesn3 Sestrin 3 (Sesn3), mRNA NM_030261 5.15972 4.03E-09
D030022P07Rik RIKEN cDNA 4930451A13 gene, mRNA (cDNA clone MGC:25849 IMAGE:4194266) AK045169 5.15849 2.37E-16
Spock2 Sparc/osteonectin, cwcv and kazal-like domains proteoglycan 2 (Spock2), mRNA NM_052994 5.15198 5.72E-12
Elavl3 ELAV (embryonic lethal, abnormal vision, Drosophila)-like 3 (Hu antigen C) (Elavl3), mRNA NM_010487 5.14832 3.61E-06
LOC620630 similar to Contactin 5 precursor (Neural recognition molecule NB-2) XM_146640 5.14694 0.0002
Plxna3 Plexin A3 (Plxna3), mRNA NM_008883 5.14348 1.78E-10
2200001I15Rik RIKEN cDNA 2200001I15 gene (2200001I15Rik), mRNA NM_183278 5.12983 9.01E-11
Stk38l Serine/threonine kinase 38 like, mRNA (cDNA clone MGC:70096 IMAGE:30135051) NM_172734 5.12867 1.65E-13
NAP092202-001 Unknown 5.12151 9.54E-32
Dock4 MKIAA0716 protein AK049953 5.10798 0.0001
Nek1 Mus musculus 13 days embryo male testis cDNA, RIKEN full-length enriched library, clone:6030407P11 product: NIMA (never in mitosis gene a)-related expressed kinase 1, full insert sequence. AK031330 5.1072 1.46E-15
Rgs2 Regulator of G-protein signaling 2, mRNA (cDNA clone MGC:36187 IMAGE:5365271) NM_009061 5.10596 4.25E-10
Mapk8ip1 JNK interacting protein-1b (JIP-1b) NM_011162 5.10563 1.85E-29
Pard6b Par-6 (partitioning defective 6) homolog beta (C. elegans), mRNA (cDNA clone MGC:35871 IMAGE:2655658) BC025147 5.1056 2.16E-21
H1f0 H1 histone family, member 0, mRNA (cDNA clone MGC:19309 IMAGE:4166167) NM_008197 5.10476 2.71E-06
Dmp1 Dentin matrix protein 1 (Dmp1), mRNA NM_016779 5.10461 0.0003
Usf1 M.musculus USF1 (non-coding exon 1). X95315 5.1015 0.00002
2810453I06Rik RIKEN cDNA 2810453I06 gene (2810453I06Rik), mRNA NM_026050 5.08664 2.86E-23
Ndrg3 N-myc downstream regulated gene 3, mRNA (cDNA clone MGC:21649 IMAGE:4500970) NM_013865 5.08321 3.05E-26
2810013E07Rik RIKEN cDNA 2810013E07 gene, mRNA (cDNA clone MGC:67555 IMAGE:6402150) AK036255 5.08237 0.0059
6330406L22Rik RIKEN cDNA 6330406L22 gene, mRNA (cDNA clone MGC:59510 IMAGE:6333971) AK041380 5.07511 0.00009
Tubgcp5 Mus musculus 12 days embryo embryonic body between diaphragm region and neck cDNA, RIKEN full-length enriched library, clone:9430028E06 product: unclassifiable, full insert sequence. AK034715 5.07129 0.00549
AK037544 Mus musculus 16 days neonate thymus cDNA, RIKEN full-length enriched library, clone:A130024K02 product: hypothetical Proteinase inhibititor I4, serpin containing protein, full insert sequence. AK037544 5.07051 5.67E-11
Creb3l3 CAMP responsive element binding protein 3-like 3 (Creb3l3), mRNA NM_145365 5.06499 2.30E-13
Mfap3l Microfibrillar-associated protein 3-like, mRNA (cDNA clone MGC:106286 IMAGE:5025281) AK122332 5.0638 0.00006
Ddah2 Dimethylarginine dimethylaminohydrolase 2, mRNA (cDNA clone MGC:5866 IMAGE:3158175) NM_016765 5.06334 6.63E-30
Il1rap Interleukin 1 receptor accessory protein (Il1rap), transcript variant 2, mRNA NM_134103 5.06296 4.72E-21
Cox7a1 Cytochrome c oxidase, subunit VIIa 1 (Cox7a1), mRNA NM_009944 5.06261 3.37E-27
Unc13a Glycosyltransferase 25 domain containing 1 (Glt25d1), mRNA AK083196 5.06174 6.78E-08
Mdm1 Transformed mouse 3T3 cell double minute 1 (Mdm1), transcript variant 2, mRNA NM_148922 5.0613 3.82E-13
Flywch1 FLYWCH-type zinc finger 1 (Flywch1), mRNA BC025645 5.06097 8.55E-16
2410116G06Rik RIKEN cDNA 2410116G06 gene (2410116G06Rik), mRNA NM_026630 5.05623 7.14E-06
AK087708 Mus musculus 2 days pregnant adult female ovary cDNA, RIKEN full-length enriched library, clone:E330009M13 product: unclassifiable, full insert sequence. AK087708 5.05545 0.00228
Hist1h2bk Histone 1, H2bk (Hist1h2bk), mRNA NM_175665 5.0523 1.47E-33
Dffa DNA fragmentation factor, alpha subunit (Dffa), transcript variant 2, mRNA NM_010044 5.05211 4.21E-08
Gpr155 G protein-coupled receptor 155, mRNA (cDNA clone IMAGE:4167684) AY255560 5.05096 5.00E-13
Cntn3 Contactin 3 (Cntn3), mRNA NM_008779 5.05051 8.40E-08
Trpc6 Transient receptor potential cation channel, subfamily C, member 6 (Trpc6), mRNA NM_013838 5.04168 0.00005
Xpr1 Xenotropic and polytropic murine retrovirus receptor (Xpr1) AK047436 5.0407 5.27E-29
BC052046 CDNA sequence BC052046 (BC052046), mRNA NM_183177 5.03591 1.47E-06
Scnn1g Sodium channel, nonvoltage-gated 1 gamma, mRNA (cDNA clone MGC:29058 IMAGE:5042876) NM_011326 5.0338 0.00021
Rab11fip1 MFLJ00294 protein AK008412 5.03263 6.56E-16
Zfpm1 Zinc finger protein, multitype 1 (Zfpm1), mRNA NM_009569 5.02713 4.89E-09
Serpinb1c Serine (or cysteine) peptidase inhibitor, clade B, member 1c (Serpinb1c), mRNA NM_173051 5.02704 1.26E-06
Ablim3 Actin binding LIM protein family, member 3, mRNA (cDNA clone MGC:106450 IMAGE:30534236) NM_198649 5.01635 0.00514
Hic2 Hypermethylated in cancer 2 protein (Hic2) NM_178922 5.01624 5.56E-27
Vkorc1 Vitamin K epoxide reductase complex, subunit 1, mRNA (cDNA clone MGC:25747 IMAGE:3991412) NM_178600 5.01352 1.91E-06
3-Mar Membrane-associated ring finger (C3HC4) 3 (March3), mRNA AK018160 5.01164 0.00268
Kcnh1 Potassium voltage-gated channel, subfamily H (eag-related), member 1 (Kcnh1), mRNA NM_010600 5.00508 2.36E-08
Rab37 RAB37, member of RAS oncogene family (Rab37), mRNA NM_021411 5.00277 1.08E-09
Hist3h2a Histone 3, H2a, mRNA (cDNA clone MGC:70265 IMAGE:6493838) NM_178218 5.00243 0.00002
Atp6v1b2 ATPase, H+ transporting, V1 subunit B, isoform 2, mRNA (cDNA clone MGC:21587 IMAGE:4500843) BB633273 5.00183 2.44E-07

Table V.

Confirmed/Putative TSGs identified in our screen that contain CpG islands in both human and mouse forms of the gene.

Gene CpG Island
M/H
RT-PCR* MSP**
M/H
Tspan33 +/+ + −/−
Cabc1 +/+ + −/−
Dusp2 +/+ + +/+
Nlk +/+ + −/−
Timp3 +/+ + +/+
Dbccr1 +/+ −/−
Klf2 +/+ −/−
Dock4 +/+ −/−
Cacna1g +/+ + −/−
Atp1b2 +/+ + +/+
Egr1 +/+ −/−
Smpd3 +/+ + +/+
Akap12 +/+ −/−
Rprm +/+ + +/+
Zfp185 +/+ −/−
Cry2 +/+ −/−
Scrib +/+ −/−
Btg2 +/+ −/−
Foxj1 +/+ + +/+
Sesn3 +/+ −/−
Kremen1 +/+ + −/−
Bach2 +/+ + −/−
Rb1 +/+ −/−
Ddit3 +/+ −/−
Irf6 +/+ −/−
Bcl2l11 +/+ −/−
Tusc2 +/+ ND ND
Tfdp2 +/+ ND ND
Ptpn13 +/+ ND ND
Ephb3 +/+ ND ND
Pcdh10 +/+ ND ND
Eaf2 +/+ ND ND
Sema3b +/+ ND ND
Lrp1b +/+ ND ND
Rad9b +/+ ND ND
Cdh15 +/+ ND ND
Dapk2 +/+ ND ND

M: mouse, H: human

ND: not done

*

For RT-PCR assays, + = increased gene expression in 5-azadC/TSA treated EMT6 cells

**

For MSP assays, + = gene methylation in any mouse mammary or human breast tumor line studied

Acknowledgments

The authors would like to thank Drs. Lingbao Ai and Wan-Ju Kim for technical assistance and advise during the completion of this study and Dr. Martha Campbell-Thompson for supplying breast tumors.

Supported, in part, by a grant from the NIH (R21 CA102220) to KDB.

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