Human Induced Pluripotent Stem Cell–Derived Podocytes Mature Into Vascularized Glomeruli Upon Experimental Transplantation

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Human Induced Pluripotent Stem Cell–Derived Podocytes Mature into Vascularized Glomeruli upon Experimental Transplantation

† Sazia Sharmin,* Atsuhiro Taguchi,* Yusuke Kaku,* Yasuhiro Yoshimura,* Tomoko Ohmori,* ‡ † ‡ Tetsushi Sakuma, Masashi Mukoyama, Takashi Yamamoto, Hidetake Kurihara,§ and | Ryuichi Nishinakamura*

*Department of Kidney Development, Institute of Molecular Embryology and Genetics, and †Department of Nephrology, Faculty of Life Sciences, Kumamoto University, Kumamoto, Japan; ‡Department of Mathematical and Life Sciences, Graduate School of Science, Hiroshima University, Hiroshima, Japan; §Division of Anatomy, Juntendo University School of Medicine, Tokyo, Japan; and |Japan Science and Technology Agency, CREST, Kumamoto, Japan

ABSTRACT Glomerular podocytes express proteins, such as nephrin, that constitute the slit diaphragm, thereby contributing to the filtration process in the kidney. Glomerular development has been analyzed mainly in mice, whereas analysis of human kidney development has been minimal because of limited access to embryonic kidneys. We previously reported the induction of three-dimensional primordial glomeruli from human induced pluripotent stem (iPS) cells. Here, using transcription activator–like effector nuclease-mediated homologous recombination, we generated human iPS cell lines that express green fluorescent protein (GFP) in the NPHS1 locus, which encodes nephrin, and we show that GFP expression facilitated accurate visualization of nephrin-positive podocyte formation in vitro. These induced human podocytes exhibited apicobasal polarity, with nephrin proteins accumulated close to the basal domain, and possessed primary processes that were connected with slit diaphragm–like structures. Micro- array analysis of sorted iPS cell–derived podocytes identified well conserved marker gene expression previously showninmouseandhumanpodocytesin vivo. Furthermore, we developed a novel transplantation method using spacers that release the tension of host kidney capsules, thereby allowing the effective formation of glomeruli from human iPS cell–derived nephron progenitors. The human glomeruli were vascularized with the host mouse endothelial cells, and iPS cell–derived podocytes with numerous cell processes accumulated around the fenestrated endothelial cells. Therefore, the podocytes generated from iPS cells retain the podocyte-specific molecular and structural features, which will be useful for dissecting human glomerular development and diseases.

J Am Soc Nephrol 27: 1778–1791, 2016. doi: 10.1681/ASN.2015010096

The glomerulus is the filtering apparatus of the microscopy. The molecular nature of the slit dia- kidney and contains three types of cells: podocytes, phragm was initially revealed by identification of vascular endothelial cells, and mesangial cells. NPHS1 as the gene responsible for Finnish-type Podocytes cover the basal domains of the endothelial cells via the basement membrane and play a Received January 26, 2015. Accepted August 26, 2015. fi 1,2 major role in the ltration process. Podocytes S.S. and A.T. contributed equally to this work. possess multiple cytoplasmic protrusions. The pri- Published online ahead of print. Publication date available at mary processes are complicated by the further www.jasn.org. stemming of smaller protrusions (secondary pro- Correspondence: Dr. Ryuichi Nishinakamura, Department of cesses or foot processes), which interdigitate with Kidney Development, Institute of Molecular Embryology and those from neighboring podocytes. The gaps be- Genetics, Kumamoto University, 2-2-1 Honjo, Chuo-ku, Kumamoto tween these foot processes are connected with the 860-0811, Japan. Email: [email protected] slit diaphragm, which is detectable only by electron Copyright © 2016 by the American Society of Nephrology

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contributes to the formation of the slit diaphragm.2 The molecular mechanisms underlying podocyte development have been extensively studied in mice. However, because of limited access to human embryos, relatively little is known regarding transcription profiles of podocytes and glomerulogenesis in humans.4,11,12 We have recently induced the nephron progenitors from mouse embryonic stem (ES) cells and human induced pluripotent stem (iPS) cells by redefining the in vivo origin of the nephron progenitors.13 The induced progenitor aggregates readily form three-dimensional primordial glomeruli and renal tubules upon Wnt stimulation in vitro. To analyze the detailed structures and transcription profiles of the induced podocytes, we have here inserted the GFP gene into the NPHS1 locus of human iPS cells via homologous recombination using transcription activator–like effector nucleases (TALENs)14 and generated glomeruli with green fluorescent protein (GFP)-tagged podocytes.

Figure 1. Successful generation of NPHS1-GFP iPS cells by homologous recombination. (A) Strategy for targeting the human NPHS1 locus. The GFP cassette was RESULTS inserted upstream of the NPHS1 start codon. The puromycin-resistance cassette (PURO) is flanked by loxP sites. Positions for primers and probes for screening are Fluorescent Visualization of Human indicated. E, EcoRV;N,NheI. (B) Southern blot of control (+/+) and NPHS1-GFP (GFP/+) clones. Genomic DNA was digested and hybridized with the indicated probes. Glomerular Podocytes Generated from NPHS1-GFP iPS Cells To visualize developing human podocytes in congenital nephrotic syndrome.3 The nephrin protein enco- vitro, we inserted a gene encoding GFP into the NPHS1 locus ded by NPHS1 intercalates with those from neighboring cells, of human iPS cells by homologous recombination (Figure thus forming a molecular mesh that hinders serum proteins of 1A). We first constructed a pair of plasmids expressing TAL- high molecular weight from leaking into the urine.4,5 To date, ENs targeted in close proximity to the NPHS1 start codon. many slit diaphragm–associated proteins have been identified, When tested in HEK 293 cells, these plasmids efficiently including NPHS2 (encoding podocin) and NEPH1, mutations deleted the NPHS1 gene (Supplemental Figure 1A). We then that cause proteinuria in humans and/or mice.6,7 introduced these TALEN plasmids, along with a targeting vector Podocytes are derived from nephron progenitors that reside containing the GFP gene and the homology arms, into human in the embryonic kidney and express transcription factor Six2.8 iPS cells. This resulted in efficient homologous recombination Upon Wnt stimulation, the nephron progenitors undergo and isolation of heterozygous GFP knock-in (NPHS1-GFP) mesenchymal-to-epithelial transition and form a tubule.9 clones (Figure 1B, Supplemental Figure 1, B and C). This tubule changes its shape; one end forms the glomerulus We differentiated these NPHS1-GFP iPS clones toward the with podocytes inside, which is surrounded by a Bowman’s nephron progenitors and subsequently combined them with capsule. Meanwhile, vascular endothelial cells and mesangial murine embryonic spinal cord, which is a potent inducer of cells migrate into the developing glomeruli, thus connecting tubulogenesis, as we previously reported.13 Four days after the glomeruli with circulation.2 In these processes, several recombination, spotty GFP signals could be observed, and transcription factors, including Wt1, regulate expression of the number and intensity of GFP signals increased thereafter nephrin in podocytes.10 Apical junctions are initially formed until day 9 (Figure 2A, Supplemental Figure 2A). We observed between the presumptive podocytes, but the apical domain GFP signals in all the examined samples from seven indepen- loses its direct contact with that of the neighboring cells, dent experiments (a total of 50 samples). Some of the signals thus forming the characteristic podocyte shape. Nephrin is started in a crescent shape and gradually changed into round eventually localized to the site close to the basal domain and structures (Figure 2A, lower panels), which suggests that

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A D3 DD54 6D most of the GFP signals overlapped with those of WT1 (Figure 2B). We ordered these glomeruli according to GFP intensity, which is likely to reﬂect the chronologic order of development. Weakly GFP-positive (and WT1-positive) limbs appeared at one end of the tubules, which elongated to surround the renal tubules. GFP intensity increased when the podocyte layers were D7 D8 D9 convoluted. At day 9, strongly GFP-positive round glomeruli were formed. These histologic changes are consistent with the previous observations of human glomeruli in aborted fetuses.15 Thus, we succeeded in visualizing human podocyte development and glomerulogenesis in vitro. Interestingly, some, but not all, of the Bowman’s capsule cells were positive for GFP B D3 D6 D6 D9 and nephrin (Supplemental Figure 2B), suggesting that these cells are not completely speciﬁed yet. Indeed, transient nephrin HE expression in some capsule cells was reported in vivo.16

Induced Podocytes Exhibit Apicobasal Polarity and

GFP Basally Localized Nephrin We analyzed day 9 sections at higher resolution to examine the apicobasal polarityof the induced podocytes. GFP was detected in the nuclei and cytoplasm of most cells in the round glomeruli

WT1/GFP (Figure 3A) because we did not attach any localization signal to GFP when generating NPHS1-GFP iPS cells. Nephrin pro- C Ecad/cad6 Ecad/WT1 Ecad/cad6 Ecad/WT1 teins were distributed in a linear fashion in the iPS cell–derived glomeruli and at one end of the WT1-positive podocyte layer

Figure 2. Fluorescent visualization of human glomerular podocytes generated from NPHS1-GFP iPS cells. (A) Morphologic A Merged Merged Nephrin GFP changes of GFP-positive glomeruli during differentiation in vitro. The nephron progenitors induced from NPHS1-GFP iPS cells were combined with murine embryonic spinal cord and cultured for the indicated time. Lower panels: higher magnification of the areas marked by rectangles in the upper panels. Note the shape B Merged Merged Nephrin WT1 changes of the glomerulus (arrowheads). Scale bars: 500 mm. (B) Histologic sections of glomeruli developing in vitro. Tissues at day 3, 6, and 9 after recombination with the spinal cord were analyzed. Top panels: Hematoxylin-eosin (HE) staining. Middle Merged Merged Nephrin COL panels: GFP (green) staining. Bottom panels: dual staining with C GFP and WT1. Nuclei were stained with 49,6-diamidino-2- phenylindole (DAPI: blue). Scale bars: 20 mm. (C) Presumptive S-shaped bodies observed at day 4 (left two panels) and day 6 (right two panels). Serial sections were stained with E-cadherin D Merged Merged Nephrin PODXL (Ecad: magenta)/cadherin-6 (cad6: green) and E-cadherin (magenta)/WT1 (green). Arrowheads: WT1-positive presumptive glomerular regions. Scale bars: 20 mm. human glomerular formation in vitro may be visualized. Figure 3. Induced podocytes exhibit apicobasal polarity and basally Therefore, we examined glomerulogenesis using sections of localized nephrin. (A) Nephrin (magenta) and GFP (green) staining of the explants. At day 3, only tubular structures were observed the induced glomerulus at day 9. (B) Nephrin (magenta) and WT1 (green) staining. (C) Nephrin (magenta) and type IV collagen (COL: and GFP-positive cells were undetectable (Figure 2B). At day 4, green) staining. (D) Nephrin (magenta) and podocalyxin (PODXL: structures that resembled S-shaped bodies were observed, in green) staining. The left columns are at lower magnification to show fi which proximo-distal speci cation occurred toward the pre- a whole glomerulus. The right two columns are singly stained, while sumptive distal (E-cadherin+) and proximal (cadherin-6+) re- the left two columns represent merged images. Arrows: nephrin nal tubules and glomerular podocytes (WT1+) (Figure 2C). At proteins localized to the basal domain; arrowheads: nephrin-positive day 6, various forms of primordial glomeruli were observed, and dot-like or filamentous structures. Scale bars: 10 mm.

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(Figure 3, A and B). These expression pat- AB terns significantly overlapped with those of type IV collagen, which was accumulated on the basal side of the podocytes (Figure 3C). In contrast, podocalyxin, an apical marker, was expressed in a manner mutu- ally exclusive of nephrin (Figure 3D). Therefore, the induced podocytes exhibited a well established apicobasal polarity and nephrin proteins were properly localized at the basal side, where the pre- CD sumptive slit diaphragm should be formed. We also observed nephrin-positive dots on the lateral side of the podocytes (Figure 3A, arrowheads), as reported in human developing podocytes in vivo.15 We found that these dots actually represent the filamentous structures encompassing the basal to the lateral side of the podocytes (Figure 3, B and C, arrowheads). Although further investigation is required, this may reflect the EF transit state of nephrin proteins shifting from the apical to the basal domain of the induced podocytes. * * Induced Podocytes Possess Primary Processes with the Slit Diaphragm– * Like Structures We further analyzed the morphology of the induced glomeruli by electron microscopy. Both scanning and transmission electron GH microscopy showed well organized glomeruli surrounded by Bowman’scapsules(Fig- ure 4, A and B). Interestingly, numerous microvilli were detected in the apical domain of the induced podocytes (Figure 4, C and D). Similar microvilli were reported in developing in vivo podocytes in humans.17,18 The podocytes were attached to each other at sites close to the basal region (Figure 4D). Inspection of the basal Figure 4. Induced podocytes possess primary processes with the immature slit di- side of the induced podocytes by scanning aphragm–like structures. (A and B) Induced glomerulus covered with a Bowman’s microscopy identified multiple protrusions capsule shown by (A) scanning and (B) transmission electron microscopy. (C) Induced (Figure 4E), which were confirmed by podocytes observed by scanning electron microscopy. Multiple microvilli are ob- transmission microscopy (Figure 4F). served on the apical surface (arrowheads). (D) Aligned podocytes, which attach to Higher magnification clearly showed each other at sites close to the basal region, shown by transmission electron bridging structures between the protru- microscopy. Multiple microvilli are observed on the apical surface (arrowheads). (E) sions, which may represent an immature Primary processes shown by scanning electron microscopy (asterisks). Podocytes from form of the slit diaphragm (Figure 4, G the basal side are shown. (F) Primary processes shown by transmission electron mi- – – and H, Supplemental Figure 3, A C). croscopy (asterisks). (G) Slit diaphragm like structures between the primary processes fi in vitro (arrows), shown by scanning electron microscopy. (H) Primary processes with slit Thus, this is the rst generation of diaphragm–like structures (arrows), shown by transmission electron microscopy. Scale mammalian podocytes with slit dia- – bars: A and B: 10 mm; C–F: 2 mm; G and H: 0.2 mm. phragm like structures from pluripotent stem cells. However, because typical inter- digitation of the protrusions is lacking,

J Am Soc Nephrol 27: 1778–1791, 2016 Podocytes from Human iPS Cells 1781 BASIC RESEARCH www.jasn.org they are likely to represent primary processes but not second- heavy chain 9 (Myh9), causative genes for hereditary kidney ary processes (foot processes). diseases,28–30 were also included. Transcription factors that have important roles in podocyte development, including Induction of Podocytes from Human NPHS1-GFP iPS WT1, MAFB, FOXD1, and TCF21, as well as vascular attractants Cells Enables Their Efficient Isolation such as VEGFA and semaphorin, were also expressed.1,2,31 In- We next tried to purify the GFP-positive podocytes at day 9 by terestingly, when these selected overlapping genes were used for FACS. Of the induced cells, 7.45%60.72% (mean6SEM from the cluster analysis against the microarray data from various five independent induction experiments) were positive for organs described above, kidney and fetal kidney were segregated GFP (Figure 5A, left panel). We also found that the monoclo- as separate clusters, suggesting the kidney-biased features of the nal antibody against the extracellular domain of nephrin overlapping gene set (Supplemental Figure 4B). (48E11),19 in combination with the anti-podocalyxin anti- We also identified genes common to GFP-positive podo- body, was useful for sorting developing podocytes. Of the cytes and adult human glomeruli (Figure 5D, Supplemental GFP-positive cells, 94.0% were positive for both nephrin Table 2), and genes common to GFP-positive podocytes and and podocalyxin (Figure 5A, middle panel), while most of mouse adult podocytes (Figure 5D, Supplemental Table 3). the GFP-negative cells (97.5%) were negative for both markers The former includes BMP7,32 while the latter includes (Figure 5A, right panel). Thus, GFP faithfully mimics nephrin NEPH1 (KIRREL), FOXC2, ROBO2,andEPHRIN-B1.7,33–36 expression and podocytes were enriched in the GFP-positive These results indicated that the typical transcriptional profiles population. Quantitative RT-PCR analysis of sorted cells con- are well conserved among our podocytes generated in vitro as firmed the differential expression of several podocyte markers, well as mouse and human podocytes in vivo. In addition, ex- such as NPHS2 (encoding podocin) and synaptopodin (Figure tracellular matrix components characteristic of glomeruli at 5B). When the sorted GFP-positive cells were cultured for 3 the capillary loop stage, laminina5/b2/g1 isoforms (corre- days, the cells expressed WT1 in nuclei and podocalyxin on the sponding to laminin 521) and type IV collagen a4/a5,37 cell surface (Figure 5C). Nephrin and GFP were detected on were detected, the latter of which is the causative gene for the cell surface membrane and in the cytoplasm, respectively, Alport syndrome. These data indicate that the transition to at day 7 of culture, although expression levels were lower than these mature forms from immature laminin 111 and collagen before the start of the culture. These results indicate that in- a1/a2 has already occurred in vitro. duction from NPHS1-GFP iPS cells enables efficient isolation Taken together, our podocytes induced in vitro possessed of developing human podocytes. the typical features of those in vivo, not only in morphology butalsointranscriptionprofiles, further supporting the GFP-Positive–Induced Podocytes Show Transcriptional authenticity of our human iPS cell induction protocol. In Profiles That Overlap with Those of Mouse and Human addition, genes exclusively expressed in the GFP-positive Podocytes In Vivo podocytes are worthy of further investigation because they To obtain comprehensive transcription profiles of the iPS cell– may include genes specific to developing human derived podocytes, we performed microarray analysis at day 9. podocytes, a possibility that has not been addressed to date We detected 2985 probes that were enriched in GFP-positive (Figure 5D, Supplemental Table 4). podocytes compared with GFP-negative cells. Of these, the top 300 genes were used for unbiased cluster analysis against Transplanted iPS Cell–Derived Nephron Progenitors microarray data from a wide variety of human tissues (Sup- Form Vascularized Glomeruli plemental Figure 4, A and C).20 Genes enriched in the GFP- We next examined whether glomeruli generated from iPS cells positive podocytes had variable tissue specificity. For example, integrated with the vascular endothelial cells. The iPS cell–de- NPHS2 was selectively expressed in the kidney or fetal kidney rived nephron progenitor spheres were induced by spinal cord tissues. However, synaptopodin and FOXC2 were sorted into for 1 day in vitro to initiate tubulogenesis and were then trans- the ubiquitously expressing cluster. Dendrin was assigned to a planted beneath the kidney capsule of immunodeficient mice. cluster enriched in the neuronal tissues. These results suggest a We also cotransplanted mixed aggregates of human umbilical single molecule is not sufficient to confirm the identity of vein endothelial cells (HUVECs) and mesenchymal stem cells podocytes. Therefore, we compared our gene list of GFP- (MSCs) because these cells are useful for the generation of positive human podocytes with the published microarray vascularized organ buds in vitro.38,39 When these aggregates analyses of adult human glomeruli and adult podocytes were transplanted using a conventional method that we used from Mafb-GFP transgenic mice.11,21 Overall, 190 probes for the transplantation of mouse ES cell–derived nephron pro- were overlapping among the three gene sets (Figure 5D, Sup- genitors,13 minimal nephron differentiation was observed at plemental Table 1, Table 1). These included typical slit dia- 10 days after transplantation (n=4) (Figure 6A). Because phragm–related genes, such as NPHS1, NPHS2, CD2AP,22 human iPS cell–derived aggregates were larger (approximately chloride intracellular channel protein 5 (CLIC5),23 and den- 1000 mm in diameter) than those from mouse ES cells (ap- drin,24,25 and basolateral adhesion molecules such as claudin 5 proximately 600 mm) and were instantly flattened upon trans- and integrin a3.26,27Phospholipase «1 and nonmuscle myosin plantation (Supplemental Figure 5A), we hypothesized that

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among the grafted tissue (Figure 6D), suggesting that the iPS cell–derived podocytes possess the potential to attract vasculature. This is also consistent with microarray data showing VEGFA expression in our induced podocytes. At day 20 after transplantation, we observed enlarged transplanted tissues beneath the capsule (Supplemental Figure 5B). Histologic examination revealed excessive growth of stromal cells of human origin, which were presumably derived from nonrenal tissues that were coinduced with nephron progenitors from iPS cells (n=4) (Figure 6E, Supplemental Figure 5C). Nonetheless, glomeruli were formed and the blood vessels were well integrated into the glomeruli (Figure 6, F and G). Moreover, 90% (135 of 150) of the glomeruli contained red blood cells (Figure 6H). Indeed, some of the glomeruli showed an enlarged Bowman’sspaceandcontained eosin-positive precipitation (Figure 6I), which might imply a small amount of urine production. Interestingly, endothelial cells in the induced glomeruli were of mouse origin (Figure 6G, Supplemental Figure 5D). HUVEC-derived endothelial cells were not integrated into the iPS cell– derived glomeruli but were located sepa- rately from the sites of nephron formation (Supplemental Figure 5E). Therefore, HU- VEC may not be competent to interact with human podocytes. The anti-human specificpodocalyxin Figure 5. Induction of podocytes from human NPHS1-GFP iPS cells enables their antibody stained the apical domains of fi ef cient isolation. (A) FACS analysis of induced tissues at day 9. Almost 8% of cells are the iPS cell–derived podocytes, but not positive for GFP in this representative experiment (left panel). Nephrin and podoca- those of the host mouse podocytes (Sup- lyxin (PODXL) expression in the GFP-positive or -negative fraction (middle and right panel, respectively). (B) Quantitative RT-PCR analysis of GFP-positive and -negative plemental Figure 5F). Nephrin protein in fractions. Average and SEM from three independent experiments are shown. b-ACT, induced podocytes was localized at b-actin; SYNPO, synaptopodin. (C) Immunostaining of podocytes cultured for the in- the basal side that faced the vascular endo- dicated times after sorting GFP-positive cells. Scale bars: 5 mm. (D) Venn diagram of thelial cells (Figure 6J), suggesting the the transcription profiles of podocytes. Microarray data of GFP-positive podocytes are emergence of filtering apparatus. Electron compared with those of human adult glomeruli and murine podocytes. microscopic analyses of two additional samples at day 20 showed that iPS cell– derived podocytes accumulated around, mechanical tension of the capsule may have hampered neph- and were closely associated with, endothelial cells (Figure ron differentiation. Therefore, we inserted two agarose rods of 7A). The induced podocytes developed numerous complex 1100 mm diameter in a V-shaped position to release tension cell processes, as well as a linear basement membrane, at in- and secure a space for the transplanted aggregates (Figure 6B). terfaces with endothelial cells (Figure 7B). The distances be- We also soaked the rods with VEGF to enhance vasculogene- tween the cell processes of some podocytes were enlarged, and sis.31 As a result, we observed immature glomerular formation slit diaphragm–like structures were formed between the pro- at day 10 in the transplants, accompanied by blood vessels cesses located above the basement membrane (Figure 7C). integrating into these glomeruli (n=5) (Figure 6, C and D). Each of these diaphragms appeared as an electron-dense line The vessels were preferentially clustered in the glomeruli (approximately 35 nm wide, 10 nm thick) bridging adjacent

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Table 1. Genes common to iPS cell–derived podocytes in vitro, human glomeruli, and mouse podocytes in vivo Gene iPS Cell–Derived Human Mouse Gene Name Symbol Podocytes Glomeruli Podocytes CLIC5 Chloride intracellular channel 5 141.94 6.48 9.10 NPHS1 Nephrosis 1, congenital, Finnish type (nephrin) 141.30 10.41 22.75 CLIC5 Chloride intracellular channel 5 107.11 6.48 9.10 NPHS2 Nephrosis 2, idiopathic, steroid-resistant (podocin) 99.93 9.99 19.95 PODXL Podocalyxin-like 78.48 9.34 26.60 PTPRO Protein tyrosine phosphatase, receptor type, O 62.91 24.68 26.69 CLDN5 Claudin 5 61.39 14.33 5.39 ITGA3 Integrin, a3 (antigen CD49C, a3 subunit of VLA-3 receptor) 51.41 9.33 9.98 CTGF Connective tissue growth factor 38.19 8.31 4.48 NPR1 Natriuretic peptide receptor 1 33.39 4.83 7.31 TGFBR3 Transforming growth factor, b receptor III 32.29 2.06 2.82 FOXD1 Forkhead box D1 27.96 3.87 18.67 MAFB v-maf avian musculoaponeurotic ﬁbrosarcoma oncogene homolog B 26.91 14.07 8.56 BCAM Basal cell adhesion molecule (Lutheran blood group) 22.95 2.04 2.71 RAPGEF3 Rap guanine nucleotide exchange factor (GEF) 3 22.71 5.02 2.04 SEMA3G Sema domain, Ig domain, short basic domain, secreted, (semaphorin) 3G 21.56 31.46 6.33 WT1 Wilms tumor 1 20.93 14.02 19.15 DDN Dendrin 19.22 9.91 4.25 HSPB8 Heat shock 22-kDa protein 8 18.38 6.53 4.66 ST3GAL6 ST3 b-galactoside a-2,3-sialyltransferase 6 17.31 9.95 4.91 TCF21 Transcription factor 21 16.82 31.46 14.60 ELF4 E74-like factor 4 (ets domain transcription factor) 15.79 3.60 4.34 MAFB v-maf avian musculoaponeurotic ﬁbrosarcoma oncogene homologue B 15.26 14.07 8.56 NTNG1 Netrin G1 15.12 2.66 1.89 LAMA5 Laminin, a5 14.59 2.27 2.32 ARHGAP29 Rho GTPase activating protein 29 14.09 2.40 1.98 PLCE1 Phospholipase C, «1 14.02 29.93 33.24 PLCE1 Phospholipase C, «1 14.00 29.93 33.24 VAMP5 Vesicle-associated membrane protein 5 13.44 7.08 2.64 ANXA1 Annexin A1 13.32 11.09 10.61 HTRA1 HtrA serine peptidase 1 13.06 11.95 4.24 MPP5 Membrane protein, palmitoylated 5 (MAGUK p55 subfamily member 5) 12.96 2.90 3.20 VEGFA Vascular endothelial growth factor A 12.93 2.33 6.20 VEGFA Vascular endothelial growth factor A 12.80 2.33 6.20 CORO2B Coronin, actin binding protein, 2B 11.74 7.54 4.86 MXRA8 Matrix-remodelling associated 8 11.08 6.58 2.98 HSPA12A Heat shock 70-kDa protein 12A 10.65 3.66 4.43 SYNPO Synaptopodin 9.66 19.26 7.40 PDLIM5 PDZ and LIM domain 5 9.32 2.87 2.30 NTNG1 Netrin G1 8.98 2.66 1.89 MAGI2 Membrane associated guanylate kinase, WW and PDZ domain containing 2 8.25 25.02 16.28 LEPREL1 Leprecan-like 1 7.46 3.79 3.14 RAPGEF3 Rap guanine nucleotide exchange factor (GEF) 3 7.31 5.02 2.04 DAG1 Dystroglycan 1 (dystrophin-associated glycoprotein 1) 6.89 5.78 4.12 FOXC1 Forkhead box C1 6.45 6.92 2.71 SPARC Secreted protein, acidic, cysteine-rich (osteonectin) 6.28 9.20 8.65 PDPN Podoplanin 6.28 8.99 17.51 TJP1 Tight junction protein 1 6.20 4.15 8.42 TM4SF1 Transmembrane 4 L six family member 1 6.06 3.66 5.99 AMIGO2 Adhesion molecule with Ig-like domain 2 6.02 17.63 3.22 TRAM2 Translocation associated membrane protein 2 5.74 11.62 2.20 LEPROT Leptin receptor overlapping transcript 5.72 2.76 1.96 FAM65A Family with sequence similarity 65, member A 5.47 6.51 13.42 PLOD2 Procollagen-lysine, 2-oxoglutarate 5-dioxygenase 2 5.33 5.50 5.84 MYO1E Myosin IE 5.25 6.36 9.88

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Table 1. Continued

Gene iPS Cell–Derived Human Mouse Gene Name Symbol Podocytes Glomeruli Podocytes PDLIM5 PDZ and LIM domain 5 5.18 2.87 2.30 TM4SF1 Transmembrane 4 L six family member 1 4.86 3.66 5.99 MYH9 Myosin, heavy chain 9, nonmuscle 4.67 4.38 4.07 MGAT5 Mannosyl (a-1,6-)-glycoprotein b-1,6-N-acetyl-glucosaminyltransferase 4.65 14.53 11.89 THSD7A Thrombospondin, type I, domain containing 7A 4.65 2.58 13.87 SYNPO Synaptopodin 4.62 19.26 7.40 LAMB2 Laminin, b2 (laminin S) 4.53 5.00 3.54 LAMB2 Laminin, b2 (laminin S) 4.52 5.00 3.54 PDLIM2 PDZ and LIM domain 2 (mystique) 4.39 7.40 1.86 KANK1 KN motif and ankyrin repeat domains 1 4.17 3.44 4.34 EHD4 EH-domain containing 4 4.13 4.54 4.46 INF2 Inverted formin, FH2 and WH2 domain containing 4.01 1.68 1.82 MYOF Myoferlin 3.93 6.07 6.55 CD151 CD151 molecule (Raph blood group) 3.91 4.13 2.25 FYN FYN oncogene related to SRC, FGR, and YES 3.88 4.72 7.95 KANK1 KN motif and ankyrin repeat domains 1 3.87 3.44 4.34 TAPBP TAP binding protein (tapasin) 3.84 2.95 1.90 MYH9 Myosin, heavy chain 9, nonmuscle 3.75 4.38 4.07 ST6GAL1 ST6 b-galactosamide a-2,6-sialyltranferase 1 3.72 1.52 5.98 ITM2C Integral membrane protein 2C 3.65 1.52 2.11 SPATS2L Spermatogenesis associated, serine-rich 2-like 3.59 1.86 2.55 F2R Coagulation factor II (thrombin) receptor 3.54 9.13 3.53 INF2 Inverted formin, FH2 and WH2 domain containing 3.52 1.68 1.82 CPD Carboxypeptidase D 3.48 2.05 1.86 AHNAK AHNAK nucleoprotein 3.46 5.70 2.26 SEC14L1 SEC14-like 1 (Saccharomyces cerevisiae) 3.36 3.67 1.88 SEC14L1 SEC14-like 1 (S. cerevisiae) 3.33 3.67 1.88 ATP8A1 ATPase, aminophospholipid transporter (APLT), class I, type 8A, member 1 3.31 2.13 2.39 NES Nestin 3.30 14.80 12.04 CD59 CD59 molecule, complement regulatory protein 3.28 2.00 8.21 NLK Nemo-like kinase 3.23 3.01 4.63 ARAF v-raf murine sarcoma 3611 viral oncogene homologue 3.21 1.87 2.10 CD2AP CD2-associated protein 3.20 1.58 3.52 MXRA7 Matrix-remodelling associated 7 3.13 3.78 1.82 NES Nestin 3.13 14.80 12.04 IL13RA1 Interleukin 13 receptor, a1 3.11 3.60 1.64 TAGLN2 Transgelin 2 3.07 2.49 1.70 NEDD9 Neural precursor cell expressed, developmentally downregulated 9 3.07 3.71 2.65 VAMP2 Vesicle-associated membrane protein 2 (synaptobrevin 2) 3.06 1.79 5.21 HEXIM1 Hexamethylene bis-acetamide inducible 1 2.95 1.56 1.78 TAPBP TAP binding protein (tapasin) 2.94 2.95 1.90 LAMC1 Laminin, g1 (formerly LAMB2) 2.93 2.69 2.29 MXRA7 Matrix-remodelling associated 7 2.92 3.78 1.82 CTNNAL1 Catenin (cadherin-associated protein), a-like 1 2.91 2.13 5.97 MYO1D Myosin ID 2.91 12.33 9.98 TRIB2 Tribbles pseudokinase 2 2.90 7.43 3.31 PTPN21 Protein tyrosine phosphatase, nonreceptor type 21 2.89 1.70 2.12 IL13RA1 Interleukin 13 receptor, a1 2.89 3.60 1.64 MXRA7 Matrix-remodeling associated 7 2.87 3.78 1.82 SPTAN1 Spectrin, a, nonerythrocytic 1 2.86 8.98 1.89 SGMS1 Sphingomyelin synthase 1 2.77 2.28 2.04 APLP2 Amyloid b(A4) precursor-like protein 2 2.72 1.69 2.06 SPTAN1 Spectrin, a, nonerythrocytic 1 2.71 8.98 1.89 MXRA7 Matrix-remodeling associated 7 2.71 3.78 1.82

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Table 1. Continued

Gene iPS Cell–Derived Human Mouse Gene Name Symbol Podocytes Glomeruli Podocytes PTGER4 Prostaglandin E receptor 4 (subtype EP4) 2.68 22.84 4.09 KDM4C Lysine (K)-speciﬁc demethylase 4C 2.66 1.81 1.87 LEPROT Leptin receptor overlapping transcript 2.64 2.76 1.96 SIRPA Signal-regulatory protein a 2.64 8.40 3.15 CTNNAL1 Catenin (cadherin-associated protein), a-like 1 2.62 2.13 5.97 LRRC1 Leucine rich repeat containing 1 2.61 2.58 3.07 IGFBP7 Insulin-like growth factor binding protein 7 2.59 1.51 1.68 CYP1B1 Cytochrome P450, family 1, subfamily B, polypeptide 1 2.58 4.53 2.56 SEL1L Sel-1 suppressor of lin-12–like (Caenorhabditis elegans) 2.58 1.98 1.86 GRK5 G protein–coupled receptor kinase 5 2.58 17.01 4.68 SERINC3 Serine incorporator 3 2.57 2.85 1.52 SEL1L Sel-1 suppressor of lin-12–like (C. elegans) 2.55 1.98 1.86 SERINC3 Serine incorporator 3 2.53 2.85 1.52 C1orf21 Chromosome 1 open reading frame 21 2.51 5.03 4.99 Genes are aligned in the order of fold induction in the GFP-positive versus -negative population of podocytes induced from NPHS1-GFP iPS cells (fold induction .2.5). Data on human glomeruli and mouse podocytes are from Ref. 11 and 21, respectively.

cell processes of the iPS cell–derived podocytes (Figure 7D). unnecessary for the purification of podocytes from patient- This feature was also observed in vivo and differed from the derived iPS cells, and possibly from more complex in vivo immature ladder-like structure that was seen between adjacent tissues. podocytes cultured exclusively in vitro without transplanta- It is surprising that well organized glomeruli are formed tion (Figure 4). Endothelial cells also produced basement withoutthe other two componentsof glomeruli: mesangial and membrane, but it was not fused to that of the podocytes in vascular endothelial cells. These two cell types are not derived most cases, thus forming double-layered structures (Figure from nephron progenitors, as shown by cell lineage analysis in 7E). Interestingly, endothelial cells were fenestrated with mice,8,44,45 and indeed we did not detect these lineages in the residual diaphragm, a characteristic feature of embryonic induced glomeruli (Supplemental Figure 3D). Thus, glomer- glomerular endothelial cells (Figure 7F).40 Furthermore, an uli can self-organize their structures solely from the podocytes electron-dense substance was detected in the Bowman’s space derived from nephron progenitors, without any inductive sig- (Figure 7C), as in Figure 6I, implying the possible presence of nals from mesangial cells or the vasculature. However, further filtration. Taken together, glomeruli generated from human maturation will be required to reproduce hereditary glomer- iPS cells were vascularized and had many morphologic fea- ular diseases. We developed a new transplantation technique tures present in glomeruli in vivo. using agarose rods to secure a space against the tension evoked by kidney capsules. This technical improvement led to the successful generation, for the first time, of vascularized glo- DISCUSSION meruli derived from human iPS cells. The induced podocytes exhibited complex cell processes with slit diaphragm–like We have inserted GFP into the NPHS1 locus of human iPS cells structures, and linear basement membrane that ran along and successfully differentiated them toward three-dimen- that of the endothelial cells was formed. Furthermore, endo- sional glomeruli. The GFP-positive–induced podocytes pos- thelial cells were fenestrated, which is a characteristic feature sessed apicobasal polarity and were equipped with primary of glomerular endothelial cells. Most experiments used processes and slit diaphragm–like structures. Furthermore, agarose rods soaked with VEGF to potentially accelerate vas- sorted podocytes exhibited typical transcription profiles that culogenesis; however, the absence of VEGF in the rods also overlap with those in vivo.Thesefindings underscore the au- caused the formation of vascularized glomeruli (Supplemental thenticity of our induction protocol. NPHS1 promoter–driven Figure 5G). Thus, we can at least conclude that the human iPS GFP expression is a good indicator of glomerulus formation. cell–derived podocytes expressed sufficient attractants, in- Several groups have reported the induction of kidney tissues in cluding VEGF, to recruit endothelial cells. vitro,13,41–43 and our iPS cell lines will be useful for assessing It is noteworthy that the integrated endothelial cells were of the induction efficiency of glomeruli by each protocol. In mouse origin from the host animals but were not derived from addition, we successfully sorted human podocytes using a HUVECs, although both vascular sources were initially located combination of anti-nephrin and anti-podocalyxin antibod- in proximity to the iPS cell–derived transplants. Therefore, ies. These reagents will make genetic GFP integration human podocytes recruited mouse endothelial cells despite

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maturation of induced podocytes. How- ever, we observed an excessive growth of stromal, presumably nonrenal, cells in the transplants. Thus, it will be essential to develop methods to purify nephron progenitors for transplantation. At the same time, it is necessary to induce genuine stromal cells because both interstitial cells and mesangial cells are derived from renal stromal progenitors.45 At present, we have no evidence that proper mesangial cells exist in our vascularized glomeruli. Ideally, human endothelial and mesangial cells that correspond to those in the developing kidney should be combined. Although further induction studies, as well as imaging tech- niques to visualize the slit diaphragm with a higher resolution,49 are needed to achieve this goal, our results will accelerate the un- derstanding of human podocyte biology both in developmental and diseased states.

Figure 6. Transplanted iPS cell–derived nephron progenitors form vascularized glomeruli. (A) Hematoxylin-eosin sections of tissues at 10 days after transplantation using CONCISE METHODS a conventional method. Right panel: magnified image of the square in the right panel. kid, kidney of the host mouse. (B) Method for transplantation using solid agarose rods. NPHS1-TALEN Right panel: macroscopic view of transplanted tissue under the kidney capsule. Ag, Generation of agarose rods. (C) Hematoxylin-eosin sections of the transplanted tissue at day 10 in Plasmids and Validation of Their the presence of the rods. Right panel: magnified images of the square. (D) Vascular- Activity NPHS1 ized glomeruli at day 10. Staining of WT1 and CD31. Right panel: magnified image of -TALENs were designed to bind the fol- the square in the left panel. (E) Hematoxylin-eosin section of the transplanted tissue at lowing sequences to cleave close to the starting day 20. Middle and right panel: magnified images of the squares in the panels on their codon of NPHS1:59-TGGCCCTGGGGAC- left, respectively. *Stromal cells. kid, kidney of the host mouse. (F) Vascularized glo- GACGC-39 for the left TALEN and 59-TCAG- meruli formed upon transplantation at day 20. Left panel: magnified images of panel CAGCCCCAGGAGCA-39 for the right TALEN E. Right panel: magnified image of the square in the left panel. Note the enlarged (the underlined sequence corresponds to ’ Bowman s space. (G) The endothelial cells are of mouse origin. Staining of WT1 the latter two thirds of the starting codon). fi (magenta) and MECA-32, a marker for mouse-speci c endothelial cells (green). (H) Platinum Gate TALEN Kit (Addgene; Kit Hematoxylin-eosin staining showing red blood cells in the induced glomeruli. (I) He- #1000000043) was used to construct a TALEN matoxylin-eosin staining showing the eosin-positive precipitates in the Bowman’s expression vector as described previously.50 A space. (J) Staining of nephrin (magenta) and CD31 (green). Right panel shows the CAG – basal localization of nephrin. Scale bars: A, C–F, I: 100 mm; B: 1 mm; G, H, J: 10 mm. promoter driven vector was used as the destination vector. To evaluate mutagenic efficiency, NPHS1-TALENs were transfected into species differences, while HUVECs may not be competent to HEK293 cells using Lipofectamine 2000 (Life Technologies). The interact with human podocytes. Even when we performed target region was amplified using the following primers: 59-AAA- transplantation without HUVECs or MSCs, we observed vas- GAAAAGCAGGTGGCAGA-39 and 59-AAAGGGCAGAGGG- cularized glomeruli, suggesting that paracrine effects of these TTTGTCT-39. When the fragment was denatured and annealed, we cells may also be minimal (Supplemental Figure 5H). The observed a clear band shift that indicated formation of a mismatched presence of double-layered basement membrane might be duplex resulting from deletions or insertions.51 The amplified frag- caused by the incomplete fusion between those derived from ment was then cloned into pCRII-TOPO (Invitrogen) and 10 clones human podocytes and mouse endothelial cells, as observed were sequenced (Supplemental Figure 1A). when mouse embryonic kidney was transplanted onto a quail chorioallantoic membrane.46 Therefore, the identification of Generation of NPHS1-GFP iPS Cells optimal sources for human endothelial cells is necessary. The human iPS cell line (201B7) was maintained on mouse embryonic While it is difficult to estimate the gestational age on the fibroblasts as described elsewhere.52 The cells were pretreated with basis of the morphology of the individual glomeruli,47,48 wait- Y27632 (10 mM) 1 hour before electroporation and were dissociated ing for a longer period after transplantation may help further into single cells with collagenase-containing trypsin solution (CTK:

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AB were correctly targeted as determined by PCR and Southern blotting analyses. The primers used for PCR screening of 59 recombination were 59-AGCACCAGCTACTTGGGAGA-39 and 59-AAGTC- en GTGCTGCTTCATGTG-39 (product size: 1347 bp), and those for 39 recombination were 59-GCCTGAAGAACGAGAATCCAGC-39 and 59–ATCTCCCCACACCTTCTCCT-39 (product size: 1150 bp). bm PCR amplifications were performed using GoTaq DNA polymerase (Promega) by denaturation at 95°C for 5 minutes, followed by 35 C * D * cycles of 95°C for 30 seconds, 58°C for 60 seconds, and 72°C for 30 seconds, and a final extension at 72°C for 7 minutes. The digoxigenin- labeled probes for Southern blot analysis were amplified using * the PCR dig probe synthesis kit (Roche Diagnostics) and the 9 9 bm following primers: 5 -CAAGTCATGGGGCTGTTTTT-3 and bm 59-TCACATGCTTGACTCCTTGC-39 for the 59 probe and 59-GGC- CCTTTTCCTCTAGAACG-39 and 59-AATTGGGTCCCAGA- EF TGTTCA-39 for the 39 probe. GFP faithfully mimicked NPHS1 expression in the podocytes, without deletion of the puromycin- resistance cassette.

bm Kidney Induction from NPHS1-GFP iPS Cells bm The NPHS1-GFP iPS clones were induced to nephron progenitors en en by a method previously established in our laboratory.13 Subsequently the induced nephron progenitor aggregates were cultured at the air- – Figure 7. iPS cell derived glomeruli in the transplants exhibited fluid interface on a polycarbonate filter (0.8 mm; Whatman) supplied many morphologic features of those in vivo. (A) Induced podo- with DMEM containing 10% FCS and with mouse embryonic spinal cytes surrounding the vascular endothelial cells and extending cord taken from E12.5 embryos.13 Upon confirmation of the capacity many cell processes, shown by transmission electron microcopy. of multiple clones to differentiate into glomeruli, clone #3 was main- (B) Complex cell processes of podocytes formed between the 54 cells and above the basement membrane. (C and D) Formation of tained in a feeder-free condition as described earlier and mainly slit diaphragm–like structures (arrows) between the cell processes used for the detailed analyses. Fifty of 50 analyzed aggregates from of induced podocytes. Note the electron-dense substance in the seven independent experiments showed GFP expression. Among Bowman’s capsule (asterisk). (E) Formation of double-layered these, the fluorescent images of four aggregates were taken daily basement membranes, each derived from endothelial cells (white using a macrozoom laser confocal microscope AZ-C1 (Nikon). arrowheads) and induced podocytes. (F) Fenestrated endothelial cells with diaphragms (black arrowheads). bm, basement mem- Immunohistochemical Analysis brane derived from induced podocytes; en, endothelial cells. Samples were fixed in 10% formalin, embedded in paraffin, and cut Scale bars: A: 1 mm; B, E: 0.5 mm; C, D, F: 0.2 mm. into 6-mm thick sections. Antigen retrieval in citrate buffer was performed before staining. Alternatively, the samples were fixed in 4% ReproCELL) followed by Accutase (Millipore); 59 and 39 homology formaldehyde for frozen sections. The following primary antibodies arms (0.87 kb and 0.92 kb, respectively) of NPHS1 were then ampli- were used: guinea pig anti-nephrin (Progen Biotechnik, catalog no. fied from the genomic DNA of the iPS cells using the following GP-N2); chick anti-GFP (Abcam, catalog no. ab13970); rabbit anti- primers: 59-ggatccGGGAGACCACCTTGATCTGA-39 and 59-gctagc- GFP (Life Technologies, catalog no. A-11122); rabbit anti-WT1 (C19) CACAGGTCCCCCTACTGTGA-39 for the 59 homology arm (Santa Cruz Biotechnology, catalog no. sc-192); rabbit anti-type IV and 59-ggcgcgccT GCTGACTGAAGGTGAGTGG-39 and 59- collagen (Rockland Immunochemicals, catalog no. 600–401–106); gcggccgcGGCCCTTAGAAGGGTACTGG-39 for the 39 homology mouse anti–E-cadherin (BD Biosciences, catalog no. 610181); rabbit arm. Upon sequence verification, these arms were cloned in the anti-cadherin-6 (kind gift from Gregory Dressler)55;goatanti- BamHI-NheIandAscI-NotI sites, respectively, of the Oct4-eGFP- podocalyxin (R&D Systems, catalog no. AF1658); rabbit anti-CD31 PGK-PURO vector (obtained from Addgene),14 such that the starting (Abcam, catalog no. Ab28364); rat anti-CD31 (BD Biosciences, cat- codon of NPHS1 was replaced with that of GFP. This targeting vector alog no. 557355); anti-human CD31 antibody (Thermo Fisher Sci- (5 mg), as well as the pair of TALEN plasmids (5 mgeach),waselec- entific, catalog no. MA5-15336); anti-mouse endothelial cell antigen troporated into the dissociated human iPS cells using Super Electro- (MECA-32: Novus Biologicals, catalog no. 100-77668); and anti- porator NEPA21 (Neppagene) under the following conditions: two human nuclear antibody STEM101 (Takara Bio, catalog no. poring pulses (125 V, 2.5 msec) followed by five transfer pulses (20 V, Y40400). Secondary antibodies were conjugated with Alexa 488 or 50 msec). Subsequently the cells were plated onto puromycin- 568 (Life Technologies). Immunofluorescence was visualized with an resistant DR4 feeders and puromycin (0.25 mg/ml) was added 2 LSM780 confocal microscope (Carl Zeiss) or a TCS SP8 confocal days after electroporation.53 Six of 12 puromycin-resistant clones microscope (Leica). Seven samples at day 9 obtained from seven

1788 Journal of the American Society of Nephrology J Am Soc Nephrol 27: 1778–1791, 2016 www.jasn.org BASIC RESEARCH independent induction experiments in vitro were serially sectioned Benjamini-Hochberg procedure, fold induction .1.5, and raw ex- and showed consistent results. pression level .150. Similarly, gene sets of adult human glomeruli were compared with those of tubules and interstitial cells Electron Microscopy (GSE21785).11 Adult podocytes from Mafb-GFP transgenic mice The tissues were fixed in 2.5% glutaraldehyde containing 1% tannic (GSE17142) were compared with adult kidney cortex acid buffered with 0.1 M phosphate buffer (PB, pH 7.4), postfixed (GSE42713).21,56 Furthermore, genes enriched in GFP-positive po- with 1% OsO4 in 0.1 M PB for 1 hour at 4°C, dehydrated in a graded docytes or the gene list common to all three comparisons (Supple- ethanol series, and embedded in epoxy resin. Ultrathin sections were mental Table 1, Table 1) were applied for cluster analysis against the stained with 4% uranyl acetate and lead citrate and then examined array from a variety of human tissues (GSE14938).20 The data were with a JEM 1230 electron microscope (JEOL, Tokyo, Japan). For analyzed using unsupervised hierarchical clustering with Canberra or scanning electron microscopy, samples were fixed in 2.5% glutaral- Chebyshev distance, respectively, and average linkage method using dehyde buffered with 0.1 M PB for 24 hours at 4°C. Samples were GeneSpringGX13. sliced with a razor blade into small pieces (1 mm thickness) and postfixed in 2% OsO4 for 2 hours. Subsequently, samples were de- Transplantation of iPS Cell–Derived Kidney Tissues hydrated, substituted with t-butyl alcohol, and dried at 220°C in a We used the parental iPS cell line (201B7) to avoid the potential vacuum. The samples were made electrically conductive by mount- impairment of differentiation potentials caused by excessive pas- ing on aluminum slabs with a carbon paste, followed by coating in an sages. The iPS cell–derived nephron progenitor aggregates were osmium plasma coater (Filgen Inc., Nagoya, Japan) to a thickness of cultured with E12.5 mouse spinal cords in vitro for 1 day to initiate approximately 10 nm, and then observed with a Hitachi S-4800 tubulogenesis.13 The 1:1 mixed spheres of HUVECs and MSCs (Tokyo, Japan). (both from Ronza) were generated in U-bottom low cell-binding plates (Thermo Fisher Scientific; total, 50,000 cells/sphere). Two Flow Cytometry spacer rods were made in Hematocrit Tubes (DRUMMOND, 1- Induced kidney tissues were dissociated by incubation with 0.25% 000-7500-C/5) with 4% agarose (SIGMA A6013) dissolved in trypsin/EDTA for 5 minutes. The percentage of cells in the GFP- PBS. The rods were soaked with recombinant VEGF (5 ng/ml dis- positive fraction was examined in five independent induction experi- solved; R&D Systems, 293-VE) in DMEM/F12 (Life Technologies, ments. In two of these experiments, further antibody staining was 11320-033) containing 10% bovine serum overnight. The host kid- performed. After blocking in normal mouse serum (Thermo Fisher ney capsule was incised at approximately 2 mm from the caudal end Scientific), staining was carried out in buffer comprising 1% BSA, 13 of the kidneys and the rods were carefully inserted in the lateral side

Hank balanced saline solution, and 0.035% NaHCO3. The extracellular with forceps. Inserted rods were arranged to make a V-shaped free domain of nephrin was stained with a mouse monoclonal antibody space and briefly cauterized with capsule membrane by electric (48E11: a kind gift from K. Tryggvason),19 followed by anti-mouse cautery to prevent them shifting from their initial positions. Fi- IgG1 phycoerythrin (eBioscience, catalog no. 12-4015-82). Podocalyxin nally, two iPS-derived aggregates with mouse embryonic spinal was stained with a goat anti-podocalyxin antibody (R&D Systems, cords and two HUVEC/MSC mixed spheres were inserted from catalog no. AF 1658) followed by donkey anti-goat Alexa-633 (Life the incised window by a 20-gauge plastic indwelling needle con- Technologies, catalog no. A21082). Data were obtained using a FACS nected by a P-200 Gilson pipette. Immunodeficient NOD/SCID/ SORPAria (BD Biosciences) and analyzed with FlowJo software (TreeS- JAK3null mice57 were used as the host animal and were anesthe- tar, Inc.). Sorted podocytes were cultured in RPMI1640 with 10% FCS tized with chloral hydrate (Wako Chemicals; 400 mg/kg body wt). on coverslips coated with laminin-521 (BioLamina). Culture experi- All animal experiments were performed in accordance with insti- ments were performed four times. tutional guidelines and approved by the licensing committee of Kumamoto University (A27–018). Quantitative RT-PCR and Microarray Analysis RNA was isolated using an RNeasy Plus Micro Kit (Qiagen) and then reverse-transcribed with random primers using the Superscript VILO cDNA synthesis kit (Life Technologies). Quantitative PCR was carried ACKNOWLEDGMENTS out using the Dice Real Time System Thermal Cycler (Takara Bio) and Thunderbird SYBR qPCR Mix (Toyobo). The primer sequences are We thank K. Tryggvason for providing the anti-nephrin antibody listed in Supplemental Table 5. All the samples were normalized (48E11); S. Okada for providing the immunodeficient mice; against b-actin expression. Three pairs of GFP-positive and -negative R. Matoba and M. Hiratsuka for microarray analysis; and S. Fujimura, fractions were subjected to microarray analyses, which were per- S. Tanigawa, and the members of the Liaison Laboratory Re- formed using an Agilent Sure Print G3 human gene expression search Promotion Center in Kumamoto University for technical (8360K) microarray (Agilent Technologies). The microarray data assistance. have been deposited in the National Center for Biotechnology Infor- This study was supported by the Japan Science and Technology mation Gene Expression Omnibus (GSE66969). Enriched genes in Agency (CREST); a KAKENHI grant (26253051) from the Ministry of the GFP-positive fraction were selected against the GFP-negative frac- Education, Culture, Sports, Science, and Technology, Japan; and a tion, based on the following criteria: increase call at P,0.05 by the grant from Japan Agency for Medical Research and Development.

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DISCLOSURES 17. Brièare N, Magny P: Scanning electron microscopic observations of None. human fetal kidney maturing in vivo and in serum-free organ culture. Anat Rec 235: 461–474, 1993 18. Hyodo T, Naguro T, Kameie T, Iino A, Miyagawa I: Scanning and transmission electron-microscopic study of the development of the REFERENCES podocyte in the human fetus. Pediatr Nephrol 11: 133–139, 1997 19. Ruotsalainen V, Reponen P, Khoshnoodi J, Kilpeläinen P, Tryggvason 1. Quaggin SE, Kreidberg JA: Development of the renal glomerulus: K: Monoclonal antibodies to human nephrin. Hybrid Hybridomics 23: Good neighbors and good fences. Development 135: 609–620, 55–63, 2004 fi 2008 20. She X, Rohl CA, Castle JC, Kulkarni AV, Johnson JM, Chen R: De nition, 2. Schell C, Wanner N, Huber TB: Glomerular development—shaping the conservation and epigenetics of housekeeping and tissue-enriched multi-cellular filtration unit. Semin Cell Dev Biol 36: 39–49, 2014 genes. BMC Genomics 10: 269, 2009 3. 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J Am Soc Nephrol 27: 1778–1791, 2016 Podocytes from Human iPS Cells 1791 Supplemental data

Supplementary Figure 1

Generation of NPHS1-GFP iPS cells

(A) TALEN activity tested in HEK 293 cells. The targeted region was PCR-amplified and cloned. Deletions in the NPHS1 locus were detected in four clones out of 10 that were sequenced.

(B) PCR screening of human iPS cell homologous recombinants

Supplementary Figure 2

Human glomeruli generated from NPHS1-GFP iPS cells

(A) Morphological changes of GFP-positive glomeruli during differentiation in vitro. A different aggregate from the one shown in Figure 2 is presented. Lower panels: higher magnification of the areas marked by rectangles in the upper panels. Note the shape changes of the glomerulus (arrowheads). Scale bars: 500 µm.

(B) Some, but not all, of the Bowman’s capsule cells were positive for nephrin (48E11 antibody: magenta) and GFP (green). Scale bars: 10 µm.

Supplementary Figure 3

Histology of human podocytes generated in vitro

(A) Transmission electron microscopy of the foot processes. Lower magnification of

Figure 4H. Scale bars: 500 nm.

(B) (C) The slit diaphragm between the foot processes. Higher magnification of the

1 regions marked by rectangles in panel A. Scale bar: 100 nm.

(D) Absence of mesangial or vascular endothelial cells in the induced glomeruli.

Anti-PDGFRβ and CD31 antibodies were used to detect the two lineages, respectively, and no positive signals were observed in the glomeruli. Podocytes are positive for WT1.

Nuclei are counterstained with Nuclear Fast Red. Scale bars: 20 µm.

Supplementary Figure 4

Cluster analysis of gene expression in various human tissues

(A) Unbiased cluster analysis across various human tissues using the top 300 genes enriched in GFP-positive podocytes.

(B) Unbiased cluster analysis across various human tissues using 190 probes overlapping in podocytes in vitro, adult human glomeruli, and adult mouse podocytes.

Supplementary Figure 5

Transplanted iPS-derived nephron progenitors form vascularized glomeruli

(A) Flattening of the iPS cell-derived tissue under the kidney capsule, immediately after transplantation by the conventional method. The graph shows areas of the transplanted tissues before (pre) and immediately after transplantation (post). Note the tissue area increased because of flattening in the absence of agarose rods (Rod-).

(B) iPS cell-derived tissue at day 20 after transplantation using agarose rods.

(C) The stromal cells were of human origin. Frozen sections were stained with an anti-human nuclear antibody (magenta). Right panel: higher magnified image of the rectangle in the left panel. The magenta signal was detected in most cells, except for the

2 kidney tissues of the host mouse (*) and the endothelial cells of mouse origin (green:

MECA-32).

(D) The endothelial cells in the induced glomeruli were of mouse origin. Staining of mouse-specific endothelial antigen (green; MECA-32) and nephrin (magenta).

(E) HUVEC were not integrated into iPS cell-derived glomeruli. Staining with anti-human specific CD31 antibody (magenta) and WT1 (green). Right two panels: higher magnified images of the rectangles in the left panels.

(F) Podocalyxin is expressed in the apical domains of the iPS cell-derived podocytes.

Staining with the anti-human specific podocalyxin antibody (green) and WT1 (magenta).

Middle panel: higher magnified image of the human iPS cell-derived glomerulus (upper rectangle in the left panel). Right panel: higher magnified image of the glomerulus (the lower rectangle in the left panel) in the kidney tissues of the host mouse (*). Note the absence of signal in mouse podocytes, confirming the species specificity of the antibody.

(G) HE staining of vascularized glomeruli upon transplantation without VEGF in the agarose rods (day 20). Right panel: staining of CD31 (green) and nephrin (magenta).

Two independent transplantation experiments showed consistent results.

(H) HE staining of vascularized glomeruli upon transplantation without HUVEC and

MSC (day 20). Right panel: staining of CD31 (green) and nephrin (magenta). Two independent transplantation experiments showed consistent results.

Scale bars: A, B: 1 mm, C–H: 20 µm

Supplementary Table 1

Genes common to iPS-derived podocytes in vitro, human glomeruli, and mouse

3 podocytes in vivo. Continued gene list of Table 1 (fold induction: between 1.5 and 2.5)

Supplementary Table 2

Genes enriched in human podocytes in vitro and adult human glomeruli in vivo

Supplementary Table 3

Genes enriched in human podocytes in vitro and mouse podocytes in vivo

Supplementary Table 4

Genes enriched in podocytes in vitro, but not in adult human glomeruli or mouse podocytes in vivo

Supplementary Table 5

Primer sequences used for qPCR analysis

4 Supplementary Figure 1

B Clone 1 2 3 6 7 8 9 10 11 12 13 14

Left arm

Right arm

C Clone 12 3 9 19 7 Clone 12 3 7 13 1

Wild type Wild type 27.3kb 9.0kb Mutant 6.3kb Mutant 2.7kb

5′ Probe 3′ Probe NheI EcoRV

Supplementary Figure2 A D3 D4 D5 D6

D7 D8 D9

B Merged Nephrin GFP

Merged Nephrin GFP Supplementary Figure 3 A

B C

D PDGFRB CD31 CD31 WT1 Supplementary Figure 4 A

MAFB ITGB3

NPHS2 DDN

SYNPO FOXC2 TCF21 KIRREL

PODXL NPHS1 B C

[Fetal [Fetal [Fetal [Fetal [Kidney](n Gene [Kidney](n Gene [Kidney](n Gene [Kidney](n Gene kidney](no kidney](no kidney](no kidney](no ormalized) symbol ormalized) symbol ormalized) symbol ormalized) symbol rmalized) rmalized) rmalized) rmalized) -0.232 -0.244 CPNE5 0.936 0.204 WT1 0.133 0.000 FRMD3 -0.169 0.075 SQRDL ST6GALN -0.290 -0.425 CD22 0.048 -0.025 DDN 0.019 0.002 GNG13 -0.098 0.197 AC2 RAB11FIP 0.127 0.110 ST3GAL6 -1.056 -0.963 PLEKHB1 0.072 0.024 FUT1 0.210 0.093 1 0.242 0.676 CGNL1 -1.131 -1.018 HEPACAM 0.393 0.290 SYNPO 0.171 0.000 MYO5C 0.369 0.292 VEGFA -0.686 -0.597 PPP2R2B 0.334 0.195 NRIP2 0.644 0.511 PLA2R1 0.390 -0.040 MAFB -0.210 -0.644 OLFM3 2.000 0.487 FOXD2 0.152 0.200 TSPAN8 CCDC109 -0.386 -0.676 -0.203 -0.095 GABRA2 -0.064 0.055 CARD6 0.047 0.331 VAMP8 B -0.151 0.132 IL6R -0.807 -0.559 PCP4 0.032 0.047 CD7 0.211 0.442 AQP3 -0.376 -0.363 C2 -0.602 -0.724 NECAB1 0.759 0.665 SOST 0.146 0.068 KRT19P2 -0.342 -0.434 LILRB3 -0.951 -0.708 DSC1 0.369 0.236 CCBL1 0.187 0.122 KRT19 -0.077 -0.165 AIF1 -1.357 0.000 MYH13 0.361 0.123 BMP7 0.044 -0.032 VAMP5 ARHGAP2 0.316 0.219 -1.027 -1.328 SERPIND1 0.547 0.407 STON2 0.168 -0.048 BST2 9 -0.368 -0.054 CHI3L1 -0.883 -0.860 PAPPA 0.226 0.034 DPPA2 0.025 -0.034 ELF4 -0.533 -0.123 G0S2 -0.405 0.060 ENPP4 0.147 0.114 SYTL1 0.046 -0.237 KLRB1 -0.138 -0.149 F13A1 -0.074 -0.161 NEBL 0.298 0.177 GPR62 0.339 0.491 PLCG2 ZMIZ1- -0.065 -0.253 IL1R1 -0.042 -0.164 SPOCK2 0.212 0.108 -0.375 -0.220 IL10RA AS1 0.186 0.130 ITGB3 0.175 -0.007 CTTNBP2 0.230 0.162 TRADD -0.332 -0.240 FAM46C 0.646 0.475 BCAM 0.053 -0.206 CRTAC1 0.372 0.323 FOXC2 0.003 0.097 LAMB3 0.367 0.060 DSC2 -0.230 -0.206 GPR98 0.248 0.224 DHRS2 -0.003 0.057 KIAA0040 0.057 -0.040 CTGF -0.400 0.365 MRO 0.126 0.146 TPK1 0.027 0.064 MFNG -0.939 -0.759 ANXA1 -0.597 -0.380 CPEB1 0.110 0.438 FOXL1 0.215 0.501 GADD45A LOC28408 -0.451 -0.523 TMEM40 -0.483 -0.268 DNAJC6 0.218 0.123 -0.124 -0.044 KRT4 0 -0.182 -0.215 ANXA3 -0.086 -0.114 CORO2B 0.282 0.201 CLDN5 -0.119 -0.145 TESC -0.122 0.031 ADM 0.066 0.151 HSPA12A 0.324 0.257 CRB2 0.047 0.116 CCDC122 -0.164 -0.210 GPRC5A -0.032 -0.277 PPFIA4 0.231 0.188 ATOH7 0.388 0.301 SBSPON -0.171 -0.232 GPR126 0.807 0.516 PTPRO 0.216 0.251 SECTM1 -0.087 -0.125 ZNF750 0.027 -0.025 FRY 0.094 0.000 FAM81A 0.248 0.245 DRD4 0.321 0.820 SFXN2 TMEM178 -0.265 -0.014 PRUNE2 0.227 0.000 0.240 0.183 TBXA2R 0.178 0.022 ZNF185 A -0.139 -0.033 CHP2 -0.085 0.258 FGF1 0.327 0.000 MMP23B 0.141 0.209 SPINT2 0.143 0.305 ADRA2C -0.171 0.000 MLC1 0.267 0.162 DNAJB13 0.337 0.285 ITGA3 ADAMTSL -0.080 -0.090 S100A7 -0.355 -0.321 DLG2 0.280 0.263 0.080 0.091 SLC48A1 4 -0.122 -0.103 TPPP2 -0.424 -0.390 CAMK2A 0.096 0.109 SLAMF8 0.098 0.097 SEMA3B -0.471 -0.415 HSPB8 0.449 0.562 AIF1L 0.068 0.027 GFRA3 0.125 0.290 S100A2 -0.478 -0.510 HSD11B1 -0.114 -0.149 KLK6 0.187 0.108 PTGS1 0.171 0.106 EPS8L1 -0.266 -0.331 TPPP3 -0.588 -0.248 FA2H 0.084 0.064 POU2AF1 0.614 0.438 PODXL MAGI2- -0.263 -0.026 HTRA1 0.713 0.178 0.189 0.030 LEFTY1 0.078 0.000 MYH3 IT1 -0.095 0.622 GPX3 0.359 0.382 NTNG1 0.239 0.015 GPA33 1.021 0.817 NPHS1 0.665 0.952 ENPEP -0.276 -0.024 ARC 0.156 0.064 KIRREL2 0.061 -0.011 B3GALT4 0.566 0.990 SULT1C2 0.314 0.027 C15orf27 -0.026 -0.178 SLC16A1 0.066 -0.013 TFF3 -0.220 -0.285 PDE4DIP 0.020 -0.156 FOXD1 -0.040 0.079 LINC01127 0.314 0.000 MSLN -0.126 0.105 FAM134B 0.252 0.066 ARHGEF3 -0.025 -0.043 RSPH10B 0.491 0.022 EMID1 -0.352 -0.016 KLK7 0.374 0.069 PDE6B 0.196 0.039 TSPAN2 0.156 0.000 MRGPRF -0.321 0.389 GPD1 -0.045 0.024 SORL1 0.701 0.226 TCF21 0.342 0.249 LAMA5 -0.211 -0.424 DMBT1 -0.043 -0.015 RFPL1 0.401 0.142 PLCE1 0.052 -0.082 RAMP1 -0.135 -0.269 LRRC36 -0.167 0.005 RFPL2 0.517 0.178 NPR1 0.110 0.286 TNFRSF8 -0.721 -0.788 FAM81B -0.094 -0.184 ABLIM2 0.376 0.119 TGFBR3 0.169 0.106 RPH3AL 0.530 0.349 CLIC5 0.406 0.000 MPP5 0.719 0.054 KIRREL 0.183 0.281 CRYGN -0.169 -0.415 TRIM54 0.331 0.310 RAPGEF3 -0.140 -0.217 S100A4 0.200 0.124 SEMA3G 1.179 1.115 NPHS2 0.391 0.043 TYRO3 -0.184 -0.417 GPR133 -0.055 0.049 NOL3 -0.487 -0.558 PLAC1 0.147 0.023 TMCC3 -0.011 -0.110 SDPR 0.283 0.249 GJA3 -0.577 0.000 MYBPC2 -0.466 -0.134 CST2 -0.020 -0.127 OLFML2A 0.341 0.251 TMEM139 -0.438 -0.517 TNNI1 0.023 0.000 PNKD -0.281 0.033 GAS6 0.340 0.247 CARD10 -0.447 -0.495 TNNT2 -0.059 0.050 RIC3 -0.199 -0.061 ITIH5 0.081 0.000 SLC51A -0.583 -0.469 PDLIM5 -0.053 0.290 ENPP5 0.067 -0.205 CPZ -1.108 -0.928 LAMP3 -0.048 0.259 FAAH -0.703 -0.595 FAM129A Supplementary Figure 5

2.5 A 2 B 1.5 1 0.5 0 Rod- pre Rod+ pre Rod- post Rod+ post C hnuclei hnuclei /MECA-32 DNephrin /MECA-32 Nephrin /MECA-32

* E hCD31/WT1 hCD31 /WT1 hCD31 /WT1 E

F WT1/PODXL WT1/PODXL WT1/PODXL

G Nephrin/CD31

H Nephrin/CD31 Supplementary Table1 Genes common to iPS cell-derived podocytes in vitro, human glomeruli, and mouse podocytes in vivo Continued gene list of Table 1 (fold induction: between 1.5 and 2.5)

Gene Gene Name iPS-derived Human Mouse Symbol podocytes glomeruli podocytes BNIP2 BCL2/adenovirus E1B 19kDa interacting protein 2 2.48 2.21 3.41 ITM2C Integral membrane protein 2C 2.41 1.52 2.11 COL4A4 Collagen, type IV, alpha 4 2.40 1.60 4.42 ACTN4 Actinin, alpha 4 2.39 1.96 3.48 ITGAV Integrin, alpha V 2.39 4.59 3.16 NFE2L1 Nuclear factor, erythroid 2-like 1 2.35 1.99 3.22 GSN Gelsolin 2.34 3.13 2.62 ARHGEF18 Rho/Rac guanine nucleotide exchange factor (GEF) 18 2.34 2.07 3.17 VAMP2 Vesicle-associated membrane protein 2 (synaptobrevin 2) 2.32 1.79 5.21 GSN gelsolin 2.28 3.13 2.62 AXL AXL receptor tyrosine kinase 2.27 5.74 2.82 CYP1B1 cytochrome P450, family 1, subfamily B, polypeptide 1 2.21 4.53 2.56 NOMO1 NODAL modulator 1 2.13 1.99 1.55 AHNAK AHNAK nucleoprotein 2.13 5.70 2.26 USP9X Ubiquitin specific peptidase 9, X-linked 2.11 2.19 1.76 OPTN Optineurin 2.11 2.43 7.32 COL4A5 Collagen, type IV, alpha 5 2.06 11.05 2.51 SYNPO Synaptopodin 2.06 19.26 7.40 TENC1 Tensin like C1 domain containing phosphatase (tensin 2) 2.05 5.36 4.18 C1orf21 Chromosome 1 open reading frame 21 2.05 5.03 4.99 RECK Reversion-inducing-cysteine-rich protein with kazal motifs 2.03 12.76 2.17 CAPN2 Calpain 2, (m/II) large subunit 2.00 3.57 2.21 ITGB5 Integrin, beta 5 1.99 3.46 6.75 RSRP1 Arginine/serine-rich protein 1 1.99 1.93 1.60 DOCK4 Dedicator of cytokinesis 4 1.98 5.68 11.99 ARHGEF17 Rho guanine nucleotide exchange factor (GEF) 17 1.97 1.74 2.30 ANXA2 Annexin A2 1.95 6.81 5.28 RAPGEF3 Rap guanine nucleotide exchange factor (GEF) 3 1.94 5.02 2.04 SULF1 Sulfatase 1 1.90 18.37 16.56 AEBP1 AE binding protein 1 1.90 1.58 3.16 SEPT2 Septin 2 1.87 2.14 2.88 GALC Galactosylceramidase 1.86 4.65 2.48 TSC22D1 TSC22 domain family, member 1 1.85 2.28 1.66 PDIA6 Protein disulfide isomerase family A, member 6 1.84 1.58 1.78 FAT1 FAT atypical cadherin 1 1.84 3.64 5.05 YIPF6 Yip1 domain family, member 6 1.82 2.56 1.54 ARNT2 Aryl-hydrocarbon receptor nuclear translocator 2 1.82 2.86 1.68 DCTD dCMP deaminase 1.81 3.18 3.21 PXDN Peroxidasin homolog (Drosophila) 1.77 3.25 2.02 PXDN Peroxidasin homolog (Drosophila) 1.75 3.25 2.02 CBLB Cbl proto-oncogene B, E3 ubiquitin protein ligase 1.74 7.80 9.30 MXD4 MAX dimerization protein 4 1.74 1.52 1.75 RCAN1 Regulator of calcineurin 1 1.73 1.98 1.53 ARAF v-raf murine sarcoma 3611 viral oncogene homolog 1.71 1.87 2.10 HLA-F Major histocompatibility complex, class I, F 1.70 1.98 1.83 BNIP2 BCL2/adenovirus E1B 19kDa interacting protein 2 1.70 2.21 3.41 CREB3L2 cAMP responsive element binding protein 3-like 2 1.69 3.00 3.05 ARL1 ADP-ribosylation factor-like 1 1.69 1.93 1.55 BCL2 B-cell CLL/lymphoma 2 1.68 1.64 1.73 VAMP3 Vesicle-associated membrane protein 3 1.67 1.79 2.26 KRCC1 Lysine-rich coiled-coil 1 1.65 2.09 1.51 ITGB1 Integrin, beta 1 (fibronectin receptor, beta polypeptide, antigen 1.64 2.45 3.28 CD29 includes MDF2, MSK12) TUBB2A Tubulin, beta 2A class IIa 1.64 2.16 1.99 IQGAP1 IQ motif containing GTPase activating protein 1 1.62 2.43 2.07 MXD4 MAX dimerization protein 4 1.62 1.52 1.75 MAN1A2 Mannosidase, alpha, class 1A, member 2 1.61 1.71 4.86 CETN3 Centrin, EF-hand protein, 3 1.60 2.98 1.54 CREBL2 cAMP responsive element binding protein-like 2 1.60 1.60 1.77 CPQ carboxypeptidase Q 1.59 2.09 3.22 BCL2 B-cell CLL/lymphoma 2 1.59 1.64 1.73 MAN1A2 Mannosidase, alpha, class 1A, member 2 1.59 1.71 4.86 BTBD1 BTB (POZ) domain containing 1 1.57 2.87 1.51 HOXA9 Homeobox A9 1.56 2.12 1.53 KTN1 kinectin 1 (kinesin receptor) 1.54 1.65 1.65 IQGAP2 IQ motif containing GTPase activating protein 2 1.53 2.50 3.67 VAMP3 vesicle-associated membrane protein 3 1.52 1.79 2.26 OSBPL9 oxysterol binding protein-like 9 1.52 2.30 1.57

Supplementary Table 2 Genes enriched in human podocytes in vitro and adult human glomeruli in vivo

Gene Gene Name iPS-derived Human Symbol podocytes glomeruli CHI3L1 chitinase 3-like 1 (cartilage glycoprotein-39) 131.10 41.99 SLAMF8 SLAM family member 8 103.02 1.55 FGF1 fibroblast growth factor 1 (acidic) 94.31 13.72 SQRDL sulfide quinone reductase-like (yeast) 70.11 3.28 FAM134B family with sequence similarity 134, member B 62.00 1.58 ITIH5 inter-alpha-trypsin inhibitor heavy chain family, member 5 34.25 8.22 IL1R1 interleukin 1 receptor, type I 32.61 1.91 SPOCK2 sparc/osteonectin, cwcv and kazal-like domains proteoglycan 31.60 11.92 (testican) 2 GPRC5A G protein-coupled receptor, class C, group 5, member A 30.79 8.54 PTGS1 prostaglandin-endoperoxide synthase 1 (prostaglandin G/H synthase 28.85 1.53 and cyclooxygenase) IL1R1 interleukin 1 receptor, type I 27.24 1.91 SLC48A1 solute carrier family 48 (heme transporter), member 1 27.22 4.07 BST2 bone marrow stromal cell antigen 2 26.22 11.36 PPFIA4 protein tyrosine phosphatase, receptor type, f polypeptide (PTPRF), 26.20 4.98 interacting protein (liprin), alpha 4 FRY furry homolog (Drosophila) 25.60 14.14 KLK6 kallikrein-related peptidase 6 24.70 22.43 ITGB3 integrin, beta 3 (platelet glycoprotein IIIa, antigen CD61) 24.05 3.92 PLA2R1 phospholipase A2 receptor 1, 180kDa 23.25 11.32 AIF1 allograft inflammatory factor 1 23.06 6.97 SERINC5 serine incorporator 5 22.21 4.80 BST2 bone marrow stromal cell antigen 2 22.01 11.36 FRY furry homolog (Drosophila) 21.47 14.14 NEBL nebulette 20.26 9.40 KLK6 kallikrein-related peptidase 6 20.23 22.43 ARHGEF3 Rho guanine nucleotide exchange factor (GEF) 3 19.78 9.53 TBXA2R thromboxane A2 receptor 18.69 2.47 SEMA3B sema domain, immunoglobulin domain (Ig), short basic domain, 17.53 6.13 secreted, (semaphorin) 3B FRY furry homolog (Drosophila) 17.33 14.14 GAS6 growth arrest-specific 6 17.11 3.17 NEBL nebulette 16.74 9.40 SLC48A1 solute carrier family 48 (heme transporter), member 1 16.59 4.07 TNNT2 troponin T type 2 (cardiac) 15.54 26.37 TPPP3 tubulin polymerization-promoting protein family member 3 15.48 23.31 RAMP1 receptor (G protein-coupled) activity modifying protein 1 15.07 2.27 CCBL1 cysteine conjugate-beta lyase, cytoplasmic 14.59 2.75 RAMP1 receptor (G protein-coupled) activity modifying protein 1 13.96 2.27 WT1-AS WT1 antisense RNA 11.98 2.63 S100A4 S100 calcium binding protein A4 11.88 4.45 SPINT2 serine peptidase inhibitor, Kunitz type, 2 11.76 1.53 KLK7 kallikrein-related peptidase 7 11.63 10.70 SORL1 sortilin-related receptor, L(DLR class) A repeats containing 11.48 1.93 CARD10 caspase recruitment domain family, member 10 11.02 2.08 BMP7 bone morphogenetic protein 7 10.78 7.13 MYO5C myosin VC 10.25 3.86 PDE4DIP phosphodiesterase 4D interacting protein 10.24 5.55 NOL3 nucleolar protein 3 (apoptosis repressor with CARD domain) 10.14 2.03 OLFML2A olfactomedin-like 2A 10.02 5.78 PPP2R2B protein phosphatase 2, regulatory subunit B, beta 9.39 3.37 B3GALT4 UDP-Gal:betaGlcNAc beta 1,3-galactosyltransferase, polypeptide 4 8.98 2.37 BMP7 bone morphogenetic protein 7 8.87 7.13 TPK1 thiamin pyrophosphokinase 1 8.78 2.56 ENPP4 ectonucleotide pyrophosphatase/phosphodiesterase 4 (putative) 8.75 1.92 KIAA0040 KIAA0040 8.61 3.59 NRIP2 nuclear receptor interacting protein 2 8.55 7.45 PDE4DIP phosphodiesterase 4D interacting protein 8.54 5.55 TNNI1 troponin I type 1 (skeletal, slow) 8.52 11.29 ZNF185 zinc finger protein 185 (LIM domain) 8.39 7.27 TYRO3 TYRO3 protein tyrosine kinase 8.27 5.94 SLC27A3 solute carrier family 27 (fatty acid transporter), member 3 7.49 2.72 PPM1H protein phosphatase, Mg2+/Mn2+ dependent, 1H 7.48 1.50 CELSR2 cadherin, EGF LAG seven-pass G-type receptor 2 7.36 2.23 BMP7 bone morphogenetic protein 7 7.31 7.13 PIEZO1 piezo-type mechanosensitive ion channel component 1 7.21 6.31 DDR1 discoidin domain receptor tyrosine kinase 1 7.02 1.80 PON2 paraoxonase 2 7.02 3.69 PDE4DIP phosphodiesterase 4D interacting protein 6.83 5.55 FRY furry homolog (Drosophila) 6.81 14.14 CFH complement factor H 6.58 5.00 FRMD1 FERM domain containing 1 6.55 1.67 TBXAS1 thromboxane A synthase 1 (platelet) 6.49 2.77 KBTBD11 kelch repeat and BTB (POZ) domain containing 11 6.46 3.37 PLTP phospholipid transfer protein 6.41 15.88 F3 coagulation factor III (thromboplastin, tissue factor) 6.34 8.00 DUSP1 dual specificity phosphatase 1 6.26 2.09 DSG2 desmoglein 2 6.23 1.67 NRGN neurogranin (protein kinase C substrate, RC3) 6.20 1.80 CFH complement factor H 6.20 5.00 LIMK2 LIM domain kinase 2 6.19 3.42 ARAP2 ArfGAP with RhoGAP domain, ankyrin repeat and PH domain 2 6.11 3.35 KAT2B K(lysine) acetyltransferase 2B 6.09 3.43 SPON2 spondin 2, extracellular matrix protein 6.09 1.74 GUCY1B3 guanylate cyclase 1, soluble, beta 3 5.98 3.20 LIMK2 LIM domain kinase 2 5.95 3.42 KCNH2 potassium voltage-gated channel, subfamily H (eag-related), 5.92 2.16 member 2 SERPINB9 serpin peptidase inhibitor, clade B (ovalbumin), member 9 5.88 4.36 PTGDS prostaglandin D2 synthase 21kDa (brain) 5.82 15.98 HOXD1 homeobox D1 5.81 2.02 PDE4DIP phosphodiesterase 4D interacting protein 5.75 5.55 KCNH2 potassium voltage-gated channel, subfamily H (eag-related), 5.71 2.16 member 2 PCOLCE2 procollagen C-endopeptidase enhancer 2 5.68 38.10 DUSP5 dual specificity phosphatase 5 5.59 2.31 GUCY1B3 guanylate cyclase 1, soluble, beta 3 5.56 3.20 OSTM1 osteopetrosis associated transmembrane protein 1 5.55 1.52 CFH complement factor H 5.22 5.00 MAN2A2 mannosidase, alpha, class 2A, member 2 5.22 2.06 BAIAP2 BAI1-associated protein 2 5.09 1.85 TGFBR2 transforming growth factor, beta receptor II (70/80kDa) 5.06 6.26 NOTCH2NL notch 2 N-terminal like 5.06 1.78 PDE4DIP phosphodiesterase 4D interacting protein 4.99 5.55 PPP2R5A protein phosphatase 2, regulatory subunit B', alpha 4.85 1.79 CAND2 cullin-associated and neddylation-dissociated 2 (putative) 4.85 7.17 NOTCH2NL notch 2 N-terminal like 4.78 1.78 SYNGR2 synaptogyrin 2 4.71 1.57 DCAF8 DDB1 and CUL4 associated factor 8 4.63 2.70 CAPRIN2 caprin family member 2 4.62 1.73 SMAD6 SMAD family member 6 4.60 7.97 NOTCH2 notch 2 4.57 4.06 CA10 carbonic anhydrase X 4.53 14.80 CAPRIN2 caprin family member 2 4.52 1.73 ADORA1 adenosine A1 receptor 4.42 3.40 SYNGR2 synaptogyrin 2 4.42 1.57 DYNLT3 dynein, light chain, Tctex-type 3 4.35 1.67 STON1 stonin 1 4.32 5.99 VWA1 von Willebrand factor A domain containing 1 4.28 1.94 RPS6KA2 ribosomal protein S6 kinase, 90kDa, polypeptide 2 4.21 4.81 SHROOM2 shroom family member 2 4.20 3.12 PVRL2 poliovirus receptor-related 2 (herpesvirus entry mediator B) 4.08 4.82 MANSC1 MANSC domain containing 1 4.07 4.85 SULT1A2 sulfotransferase family, cytosolic, 1A, phenol-preferring, member 2 4.02 2.23 CHST2 carbohydrate (N-acetylglucosamine-6-O) sulfotransferase 2 4.01 3.11 SNCA synuclein, alpha (non A4 component of amyloid precursor) 4.00 4.90 ATP2C1 ATPase, Ca++ transporting, type 2C, member 1 3.99 1.95 TPD52 tumor protein D52 3.97 1.90 CD9 CD9 molecule 3.94 2.47 AGRN agrin 3.93 4.43 JMJD7 jumonji domain containing 7 3.91 2.12 SULT1A4 sulfotransferase family, cytosolic, 1A, phenol-preferring, member 4 3.87 1.93 SORT1 sortilin 1 3.87 2.37 GALNT2 polypeptide N-acetylgalactosaminyltransferase 2 3.85 1.93 PAMR1 peptidase domain containing associated with muscle regeneration 1 3.84 7.28 SLC29A1 solute carrier family 29 (equilibrative nucleoside transporter), 3.82 2.01 member 1 SEMA3C sema domain, immunoglobulin domain (Ig), short basic domain, 3.82 2.79 secreted, (semaphorin) 3C RASSF7 Ras association (RalGDS/AF-6) domain family (N-terminal) member 3.69 1.51 7 TRIM37 tripartite motif containing 37 3.69 2.03 PVRL2 poliovirus receptor-related 2 (herpesvirus entry mediator B) 3.68 4.82 TMCO3 transmembrane and coiled-coil domains 3 3.58 2.58 P4HTM prolyl 4-hydroxylase, transmembrane (endoplasmic reticulum) 3.58 2.02 LRP10 low density lipoprotein receptor-related protein 10 3.56 2.21 IRF1 interferon regulatory factor 1 3.54 2.01 TRIM37 tripartite motif containing 37 3.42 2.03 PLEKHG3 pleckstrin homology domain containing, family G (with Rho Gef 3.42 1.64 domain) member 3 CTSD cathepsin D 3.38 1.63 DCAF8 DDB1 and CUL4 associated factor 8 3.37 2.70 LMO2 LIM domain only 2 (rhombotin-like 1) 3.36 3.18 REEP5 receptor accessory protein 5 3.36 1.82 PLEKHG3 pleckstrin homology domain containing, family G (with RhoGef 3.35 1.64 domain) member 3 CALCOCO1 calcium binding and coiled-coil domain 1 3.33 2.12 TSPAN4 tetraspanin 4 3.32 2.11 SERPINB1 serpin peptidase inhibitor, clade B (ovalbumin), member 1 3.31 2.07 C1QL1 complement component 1, q subcomponent-like 1 3.30 1.50 PLXNB2 plexin B2 3.29 1.65 ETHE1 ethylmalonic encephalopathy 1 3.29 3.00 RBPMS RNA binding protein with multiple splicing 3.27 2.73 JUP junction plakoglobin 3.26 1.89 DKK3 dickkopf WNT signaling pathway inhibitor 3 3.25 6.23 AMOTL2 angiomotin like 2 3.24 2.18 CCPG1 cell cycle progression 1 3.24 2.25 CDC42EP2 CDC42 effector protein (Rho GTPase binding) 2 3.24 2.80 MAPK10 mitogen-activated protein kinase 10 3.23 1.55 EFEMP1 EGF containing fibulin-like extracellular matrix protein 1 3.22 5.11 TSPAN4 tetraspanin 4 3.21 2.11 SEPT8 septin 8 3.19 3.16 SLC38A10 solute carrier family 38, member 10 3.16 1.62 GMDS GDP-mannose 4,6-dehydratase 3.14 7.25 PLXND1 plexin D1 3.12 2.52 PLXNB2 plexin B2 3.11 1.65 B9D1 B9 protein domain 1 3.11 1.70 CAST calpastatin 3.08 2.37 SGSH N-sulfoglucosamine sulfohydrolase 3.08 3.08 DUSP14 dual specificity phosphatase 14 3.08 4.99 TMEM80 transmembrane protein 80 3.06 2.14 TRAPPC10 trafficking protein particle complex 10 3.04 2.37 EFNB2 ephrin-B2 3.03 12.12 PSEN2 presenilin 2 3.02 2.82 LBH limb bud and heart development 3.01 3.88 BGN biglycan 3.01 6.14 CLEC16A C-type lectin domain family 16, member A 2.99 1.80 ABCA3 ATP-binding cassette, sub-family A (ABC1), member 3 2.98 2.57 N4BP2L2 NEDD4 binding protein 2-like 2 2.98 2.20 PLA2G15 phospholipase A2, group XV 2.95 1.72 MYL9 myosin, light chain 9, regulatory 2.93 22.66 DYNLT3 dynein, light chain, Tctex-type 3 2.91 1.67 WASF3 WAS protein family, member 3 2.89 2.62 B9D1 B9 protein domain 1 2.88 1.70 PXDC1 PX domain containing 1 2.86 1.75 MID2 midline 2 2.85 2.42 LAMP2 lysosomal-associated membrane protein 2 2.83 2.41 STAT1 signal transducer and activator of transcription 1, 91kDa 2.83 3.37 DDX17 DEAD (Asp-Glu-Ala-Asp) box helicase 17 2.79 1.53 PLEC plectin 2.79 1.94 NACC2 NACC family member 2, BEN and BTB (POZ) domain containing 2.79 4.93 GGCX gamma-glutamyl carboxylase 2.78 1.78 ANGPTL4 angiopoietin-like 4 2.75 2.43 SPATA20 spermatogenesis associated 20 2.74 1.92 LYST lysosomal trafficking regulator 2.73 3.39 STX7 syntaxin 7 2.72 2.02 NDNF neuron-derived neurotrophic factor 2.72 26.53 NOTCH2 notch 2 2.71 4.06 SUCO SUN domain containing ossification factor 2.69 3.01 LGR4 leucine-rich repeat containing G protein-coupled receptor 4 2.68 1.65 CAST calpastatin 2.66 2.37 IL4R interleukin 4 receptor 2.66 2.27 STX7 syntaxin 7 2.66 2.02 ATP6AP1 ATPase, H+ transporting, lysosomal accessory protein 1 2.65 2.15 ADCK2 aarF domain containing kinase 2 2.65 1.94 NPEPL1 aminopeptidase-like 1 2.64 1.68 LOXL1 lysyl oxidase-like 1 2.63 10.88 NCSTN nicastrin 2.61 2.21 ARPC5 actin related protein 2/3 complex, subunit 5, 16kDa 2.60 1.89 ZDHHC6 zinc finger, DHHC-type containing 6 2.59 9.80 BAIAP2 BAI1-associated protein 2 2.59 1.85 STXBP3 syntaxin binding protein 3 2.59 1.60 BAIAP2 BAI1-associated protein 2 2.59 1.85 LGALS8 lectin, galactoside-binding, soluble, 8 2.59 4.52 TSC22D3 TSC22 domain family, member 3 2.58 4.51 CLSTN1 calsyntenin 1 2.57 1.64 MYL4 myosin, light chain 4, alkali; atrial, embryonic 2.57 1.63 CD46 CD46 molecule, complement regulatory protein 2.55 1.56 NPTN neuroplastin 2.54 1.51 B3GNT1 UDP-GlcNAc:betaGal beta-1,3-N-acetylglucosaminyltransferase 1 2.54 3.62 PPAP2B phosphatidic acid phosphatase type 2B 2.53 2.31 GULP1 GULP, engulfment adaptor PTB domain containing 1 2.52 4.75 CDC25B cell division cycle 25B 2.52 2.83 TDRD7 tudor domain containing 7 2.52 2.45 GULP1 GULP, engulfment adaptor PTB domain containing 1 2.51 4.75 NPTN neuroplastin 2.49 1.51 TXNIP thioredoxin interacting protein 2.48 2.05 AACS acetoacetyl-CoA synthetase 2.47 1.65 ACYP2 acylphosphatase 2, muscle type 2.47 1.98 AACS acetoacetyl-CoA synthetase 2.47 1.65 HK2 hexokinase 2 2.45 5.03 CDKN1B cyclin-dependent kinase inhibitor 1B (p27, Kip1) 2.45 4.06 NPEPL1 aminopeptidase-like 1 2.44 1.68 DENND3 DENN/MADD domain containing 3 2.44 1.68 VPS8 vacuolar protein sorting 8 homolog (S. cerevisiae) 2.41 1.87 CITED2 Cbp/p300-interacting transactivator, with Glu/Asp-rich carboxy- 2.40 2.71 terminal domain, 2 YPEL5 yippee-like 5 (Drosophila) 2.40 1.77 ARF3 ADP-ribosylation factor 3 2.40 2.14 KIAA1598 KIAA1598 2.40 2.01 CASP9 caspase 9, apoptosis-related cysteine peptidase 2.39 1.74 CDIPT CDP-diacylglycerol--inositol 3-phosphatidyltransferase 2.37 1.69 TM9SF1 transmembrane 9 superfamily member 1 2.36 2.01 EPHA2 EPH receptor A2 2.35 1.92 CITED2 Cbp/p300-interacting transactivator, with Glu/Asp-rich carboxy- 2.34 2.71 terminal domain, 2 NDRG2 NDRG family member 2 2.34 5.75 NRIP1 nuclear receptor interacting protein 1 2.34 1.89 LAMP2 lysosomal-associated membrane protein 2 2.33 2.41 CDKN1B cyclin-dependent kinase inhibitor 1B (p27, Kip1) 2.33 4.06 C1GALT1C1 C1GALT1-specific chaperone 1 2.33 1.78 PSAP prosaposin 2.32 1.57 ATP13A3 ATPase type 13A3 2.31 1.69 FXYD1 FXYD domain containing ion transport regulator 1 2.31 2.33 ERBB4 v-erb-b2 avian erythroblastic leukemia viral oncogene homolog 4 2.29 3.94 PSAP prosaposin 2.29 1.57 MBD2 methyl-CpG binding domain protein 2 2.29 1.65 VPS13C vacuolar protein sorting 13 homolog C (S. cerevisiae) 2.29 2.12 RNF6 ring finger protein (C3H2C3 type) 6 2.27 1.65 TACC1 transforming, acidic coiled-coil containing protein 1 2.26 1.77 ZSCAN18 zinc finger and SCAN domain containing 18 2.26 2.48 MOAP1 modulator of apoptosis 1 2.26 1.75 CHFR checkpoint with forkhead and ring finger domains, E3 ubiquitin 2.26 1.63 protein ligase ATXN1 ataxin 1 2.26 2.77 PPFIA1 protein tyrosine phosphatase, receptor type, f polypeptide (PTPRF), 2.25 2.38 interacting protein (liprin), alpha 1 TMEM9B TMEM9 domain family, member B 2.25 2.38 IQSEC1 IQ motif and Sec7 domain 1 2.23 2.79 PDLIM1 PDZ and LIM domain 1 2.23 2.54 RNASET2 ribonuclease T2 2.22 1.83 GRN granulin 2.22 1.69 ZHX2 zinc fingers and homeoboxes 2 2.21 1.79 HLA-DMA major histocompatibility complex, class II, DM alpha 2.19 2.64 IFI16 interferon, gamma-inducible protein 16 2.19 4.37 LGALS3BP lectin, galactoside-binding, soluble, 3 binding protein 2.19 3.29 TRAPPC10 trafficking protein particle complex 10 2.18 2.37 WBP2 WW domain binding protein 2 2.17 1.65 SIPA1L1 signal-induced proliferation-associated 1 like 1 2.17 1.61 CBX7 chromobox homolog 7 2.17 3.39 EFR3A EFR3 homolog A (S. cerevisiae) 2.17 1.52 NDRG2 NDRG family member 2 2.16 5.75 MED15 mediator complex subunit 15 2.16 1.55 PRDX6 peroxiredoxin 6 2.16 1.60 C9orf3 chromosome 9 open reading frame 3 2.15 2.53 NBPF10 neuroblastoma breakpoint family, member 10 2.14 3.15 PTPN13 protein tyrosine phosphatase, non-receptor type 13 (APO-1/CD95 2.13 3.72 (Fas)-associated phosphatase) RCHY1 ring finger and CHY zinc finger domain containing 1, E3 ubiquitin 2.13 1.69 protein ligase ANKRD28 ankyrin repeat domain 28 2.11 2.88 WWC3 WWC family member 3 2.11 3.93 CD200 CD200 molecule 2.10 8.82 KDM7A lysine (K)-specific demethylase 7A 2.10 2.10 SH3BP5 SH3-domain binding protein 5 (BTK-associated) 2.10 1.74 SUN1 Sad1 and UNC84 domain containing 1 2.09 2.94 ABR active BCR-related 2.07 2.30 GABARAPL2 GABA(A) receptor-associated protein-like 2 2.07 1.63 RNASET2 ribonuclease T2 2.07 1.83 STAT6 signal transducer and activator of transcription 6, interleukin-4 2.06 1.52 induced CBX6 chromobox homolog 6 2.06 1.82 OBFC1 oligonucleotide/oligosaccharide-binding fold containing 1 2.06 3.18 MPPE1 metallophosphoesterase 1 2.05 2.00 GOLGA1 golgin A1 2.05 1.61 SIDT2 SID1 transmembrane family, member 2 2.04 1.96 BBS1 Bardet-Biedl syndrome 1 2.04 1.59 TBC1D5 TBC1 domain family, member 5 2.04 1.59 PTPMT1 protein tyrosine phosphatase, mitochondrial 1 2.04 1.68 NRIP1 nuclear receptor interacting protein 1 2.03 1.89 HLA-DMA major histocompatibility complex, class II, DM alpha 2.02 2.64 RHEB Ras homolog enriched in brain 2.01 2.04 CD200 CD200 molecule 2.01 8.82 IER5 immediate early response 5 2.01 3.79 CUL3 cullin 3 2.01 2.26 PC pyruvate carboxylase 2.00 1.74 DCAF6 DDB1 and CUL4 associated factor 6 2.00 1.93 RNF38 ring finger protein 38 2.00 2.61 SIK3 SIK family kinase 3 2.00 1.98 ATP13A3 ATPase type 13A3 2.00 1.69 CRELD1 cysteine-rich with EGF-like domains 1 2.00 3.14 ADO 2-aminoethanethiol (cysteamine) dioxygenase 2.00 2.36 HEY1 hes-related family bHLH transcription factor with YRPW motif 1 2.00 1.97 C10orf10 chromosome 10 open reading frame 10 1.99 2.38 APPBP2 amyloid beta precursor protein (cytoplasmic tail) binding protein 2 1.97 2.19 GLTSCR1L GLTSCR1-like 1.97 2.02 TM2D1 TM2 domain containing 1 1.96 3.50 TMEM50A transmembrane protein 50A 1.95 2.08 SLC7A1 solute carrier family 7 (cationic amino acid transporter, y+ system), 1.95 2.37 member 1 ARHGEF10 Rho guanine nucleotide exchange factor (GEF) 10 1.95 4.76 DNAJB2 DnaJ (Hsp40) homolog, subfamily B, member 2 1.95 1.84 PLD3 phospholipase D family, member 3 1.94 3.30 CLTB clathrin, light chain B 1.93 1.72 TACC2 transforming, acidic coiled-coil containing protein 2 1.92 3.80 LIMS2 LIM and senescent cell antigen-like domains 2 1.92 2.39 SENP6 SUMO1/sentrin specific peptidase 6 1.92 1.72 SYNJ2 synaptojanin 2 1.92 11.16 DST dystonin 1.92 3.78 UBXN2B UBX domain protein 2B 1.92 1.64 ID4 inhibitor of DNA binding 4, dominant negative helix-loop-helix protein 1.91 3.84 TM2D1 TM2 domain containing 1 1.91 3.50 EFCAB14 EF-hand calcium binding domain 14 1.91 1.67 BACE1 beta-site APP-cleaving enzyme 1 1.90 4.22 PNMAL1 paraneoplastic Ma antigen family-like 1 1.90 4.44 MPPE1 metallophosphoesterase 1 1.89 2.00 RNF103 ring finger protein 103 1.89 2.11 SAP18 Sin3A-associated protein, 18kDa 1.89 2.55 IKBKB inhibitor of kappa light polypeptide gene enhancer in B-cells, kinase 1.89 2.89 beta HLA-A major histocompatibility complex, class I, A 1.89 1.92 OBSL1 obscurin-like 1 1.88 2.95 OTUD4 OTU deubiquitinase 4 1.88 2.07 EMC1 ER membrane protein complex subunit 1 1.87 1.65 SUN1 Sad1 and UNC84 domain containing 1 1.87 2.94 ENDOD1 endonuclease domain containing 1 1.86 3.18 TXNDC15 thioredoxin domain containing 15 1.86 1.70 CLIP3 CAP-GLY domain containing linker protein 3 1.86 4.11 MBOAT7 membrane bound O-acyltransferase domain containing 7 1.85 1.71 CYLD cylindromatosis (turban tumor syndrome) 1.85 2.06 IFT122 intraflagellar transport 122 homolog (Chlamydomonas) 1.85 1.68 YPEL5 yippee-like 5 (Drosophila) 1.85 1.77 PERP PERP, TP53 apoptosis effector 1.85 1.88 UNC50 unc-50 homolog (C. elegans) 1.84 1.84 RALGAPA1 Ral GTPase activating protein, alpha subunit 1 (catalytic) 1.84 2.67 ACSL3 acyl-CoA synthetase long-chain family member 3 1.84 2.30 SLC35A5 solute carrier family 35, member A5 1.83 1.77 SNX1 sorting nexin 1 1.82 4.24 TFE3 transcription factor binding to IGHM enhancer 3 1.82 2.48 LANCL1 LanC lantibiotic synthetase component C-like 1 (bacterial) 1.82 3.64 RABGAP1L RAB GTPase activating protein 1-like 1.82 3.73 UGGT2 UDP-glucose glycoprotein glucosyltransferase 2 1.81 3.76 PCMT1 protein-L-isoaspartate (D-aspartate) O-methyltransferase 1.81 2.68 KIAA0195 KIAA0195 1.81 1.57 ZNF24 zinc finger protein 24 1.81 1.86 TMCO3 transmembrane and coiled-coil domains 3 1.81 2.58 HLA-A major histocompatibility complex, class I, A 1.81 1.92 MAP1LC3B microtubule-associated protein 1 light chain 3 beta 1.80 1.54 ZC3H14 zinc finger CCCH-type containing 14 1.80 1.63 TMED3 transmembrane emp24 protein transport domain containing 3 1.80 3.97 ANXA11 annexin A11 1.80 1.94 TBC1D22A TBC1 domain family, member 22A 1.80 1.65 VPS54 vacuolar protein sorting 54 homolog (S. cerevisiae) 1.79 1.66 WWOX WW domain containing oxidoreductase 1.78 2.49 UVRAG UV radiation resistance associated 1.78 2.17 APMAP adipocyte plasma membrane associated protein 1.78 3.03 SNX6 sorting nexin 6 1.77 2.73 RASIP1 Ras interacting protein 1 1.77 1.60 LARP4B La ribonucleoprotein domain family, member 4B 1.77 1.69 HLA-G major histocompatibility complex, class I, G 1.77 1.57 HIST1H2BK histone cluster 1, H2bk 1.77 8.91 MAP1LC3B microtubule-associated protein 1 light chain 3 beta 1.76 1.54 TOR1AIP1 torsin A interacting protein 1 1.76 1.78 MAP2K4 mitogen-activated protein kinase kinase 4 1.76 1.58 CRIP2 cysteine-rich protein 2 1.76 4.82 VPS16 vacuolar protein sorting 16 homolog (S. cerevisiae) 1.76 1.54 CTSV cathepsin V 1.75 2.06 ADAM17 ADAM metallopeptidase domain 17 1.75 2.42 PTPRA protein tyrosine phosphatase, receptor type, A 1.75 3.10 ATF1 activating transcription factor 1 1.75 1.61 BACE1 beta-site APP-cleaving enzyme 1 1.75 4.22 IDE insulin-degrading enzyme 1.74 1.57 PALLD palladin, cytoskeletal associated protein 1.74 3.08 INSIG2 insulin induced gene 2 1.74 1.77 ZNF451 zinc finger protein 451 1.74 2.52 PCMT1 protein-L-isoaspartate (D-aspartate) O-methyltransferase 1.74 2.68 ATP9B ATPase, class II, type 9B 1.73 1.55 PVR poliovirus receptor 1.73 3.29 MEF2A myocyte enhancer factor 2A 1.73 2.00 COLGALT2 collagen beta(1-O)galactosyltransferase 2 1.73 4.06 ELF1 E74-like factor 1 (ets domain transcription factor) 1.73 2.57 CTSO cathepsin O 1.73 2.72 SENP6 SUMO1/sentrin specific peptidase 6 1.72 1.72 PPFIA1 protein tyrosine phosphatase, receptor type, f polypeptide (PTPRF), 1.72 2.38 interacting protein (liprin), alpha 1 RAD23B RAD23 homolog B (S. cerevisiae) 1.72 1.72 PIGG phosphatidylinositol glycan anchor biosynthesis, class G 1.72 2.03 PIK3R4 phosphoinositide-3-kinase, regulatory subunit 4 1.71 1.99 HGSNAT heparan-alpha-glucosaminide N-acetyltransferase 1.71 2.09 METTL3 methyltransferase like 3 1.71 2.46 UBE3C ubiquitin protein ligase E3C 1.71 1.77 TMEM87A transmembrane protein 87A 1.71 1.89 H1F0 H1 histone family, member 0 1.71 3.44 COL4A6 collagen, type IV, alpha 6 1.71 2.41 SLC24A1 solute carrier family 24 (sodium/potassium/calcium exchanger), 1.70 2.11 member 1 RAD23B RAD23 homolog B (S. cerevisiae) 1.70 1.72 DMD dystrophin 1.70 5.23 XPOT exportin, tRNA 1.69 2.84 FAXDC2 fatty acid hydroxylase domain containing 2 1.69 1.83 PRDM4 PR domain containing 4 1.69 1.54 PCMT1 protein-L-isoaspartate (D-aspartate) O-methyltransferase 1.69 2.68 BEX4 brain expressed, X-linked 4 1.69 2.91 RPN2 ribophorin II 1.68 1.60 APOL2 apolipoprotein L, 2 1.68 1.72 GSTA4 glutathione S-transferase alpha 4 1.68 2.67 IP6K2 inositol hexakisphosphate kinase 2 1.68 3.91 GAS1 growth arrest-specific 1 1.68 9.85 SNX6 sorting nexin 6 1.68 2.73 SNX19 sorting nexin 19 1.68 2.24 MKRN1 makorin ring finger protein 1 1.67 1.65 FBXL2 F-box and leucine-rich repeat protein 2 1.67 1.99 ERAP1 endoplasmic reticulum aminopeptidase 1 1.67 1.90 FAM208A family with sequence similarity 208, member A 1.67 3.24 TCF25 transcription factor 25 (basic helix-loop-helix) 1.66 1.52 TRAPPC12 trafficking protein particle complex 12 1.66 1.57 TCTN3 tectonic family member 3 1.66 1.63 MKRN2 makorin ring finger protein 2 1.66 2.50 BACE1 beta-site APP-cleaving enzyme 1 1.66 4.22 PHF11 PHD finger protein 11 1.66 3.03 MBTPS1 membrane-bound transcription factor peptidase, site 1 1.65 2.11 TSC2 tuberous sclerosis 2 1.65 2.56 CLN5 ceroid-lipofuscinosis, neuronal 5 1.65 2.07 USO1 USO1 vesicle transport factor 1.64 1.87 RANGAP1 Ran GTPase activating protein 1 1.64 1.99 SALL2 spalt-like transcription factor 2 1.64 2.22 SNED1 sushi, nidogen and EGF-like domains 1 1.64 1.58 NOTCH2NL notch 2 N-terminal like 1.64 1.78 EMC2 ER membrane protein complex subunit 2 1.64 2.11 VPS52 vacuolar protein sorting 52 homolog (S. cerevisiae) 1.64 1.82 BCAR3 breast cancer anti-estrogen resistance 3 1.63 2.80 THADA thyroid adenoma associated 1.63 1.54 MBTPS1 membrane-bound transcription factor peptidase, site 1 1.63 2.11 NPC1 Niemann-Pick disease, type C1 1.63 2.99 BCL2L2 BCL2-like 2 1.63 2.29 TMEM189 transmembrane protein 189 1.63 1.68 CDYL chromodomain protein, Y-like 1.63 2.82 ZNF24 zinc finger protein 24 1.62 1.86 AGGF1 angiogenic factor with G patch and FHA domains 1 1.62 1.83 RGS10 regulator of G-protein signaling 10 1.62 5.42 VPS4B vacuolar protein sorting 4 homolog B (S. cerevisiae) 1.61 2.37 PISD phosphatidylserine decarboxylase 1.61 2.07 OVCA2 ovarian tumor suppressor candidate 2 1.61 1.77 KHNYN KH and NYN domain containing 1.61 1.62 ELF1 E74-like factor 1 (ets domain transcription factor) 1.60 2.57 ASMTL acetylserotonin O-methyltransferase-like 1.60 2.19 OBFC1 oligonucleotide/oligosaccharide-binding fold containing 1 1.60 3.18 LAMP2 lysosomal-associated membrane protein 2 1.60 2.41 MFAP3 microfibrillar-associated protein 3 1.60 1.84 NID2 nidogen 2 (osteonidogen) 1.60 1.62 PPP1R3D protein phosphatase 1, regulatory subunit 3D 1.59 2.72 HS2ST1 heparan sulfate 2-O-sulfotransferase 1 1.59 1.77 ZNF274 zinc finger protein 274 1.59 1.92 ISYNA1 inositol-3-phosphate synthase 1 1.58 2.15 OSBPL2 oxysterol binding protein-like 2 1.58 2.03 C9orf156 chromosome 9 open reading frame 156 1.58 1.65 PAK2 p21 protein (Cdc42/Rac)-activated kinase 2 1.58 2.07 TULP4 tubby like protein 4 1.58 2.09 ISYNA1 inositol-3-phosphate synthase 1 1.58 2.15 MAN1B1 mannosidase, alpha, class 1B, member 1 1.58 1.60 ATRN attractin 1.57 2.57 TPRA1 transmembrane protein, adipocyte asscociated 1 1.57 1.61 RDX radixin 1.57 2.34 LARP4B La ribonucleoprotein domain family, member 4B 1.57 1.69 KATNB1 katanin p80 (WD repeat containing) subunit B 1 1.57 1.65 ZBTB1 zinc finger and BTB domain containing 1 1.57 2.42 CHD7 chromodomain helicase DNA binding protein 7 1.57 2.68 HEG1 heart development protein with EGF-like domains 1 1.56 4.03 FYCO1 FYVE and coiled-coil domain containing 1 1.56 2.00 TMED10 transmembrane emp24-like trafficking protein 10 (yeast) 1.56 2.16 CHCHD7 coiled-coil-helix-coiled-coil-helix domain containing 7 1.56 2.15 CCSER2 coiled-coil serine-rich protein 2 1.56 2.41 GPR107 G protein-coupled receptor 107 1.56 1.84 TEX261 testis expressed 261 1.56 1.56 GTDC1 glycosyltransferase-like domain containing 1 1.56 2.18 SLC10A3 solute carrier family 10, member 3 1.55 2.27 PIGT phosphatidylinositol glycan anchor biosynthesis, class T 1.55 2.19 HLA-DMB major histocompatibility complex, class II, DM beta 1.55 1.83 SRF serum response factor (c-fos serum response element-binding 1.55 1.99 transcription factor) C8orf33 chromosome 8 open reading frame 33 1.55 3.98 FAM13B family with sequence similarity 13, member B 1.55 5.32 IQCE IQ motif containing E 1.55 2.16 PARD3 par-3 family cell polarity regulator 1.54 5.43 PANK3 pantothenate kinase 3 1.54 1.63 RBM23 RNA binding motif protein 23 1.54 2.09 PANK3 pantothenate kinase 3 1.54 1.63 MOB4 MOB family member 4, phocein 1.53 1.97 SACM1L SAC1 suppressor of actin mutations 1-like (yeast) 1.53 1.82 INPP5K inositol polyphosphate-5-phosphatase K 1.53 1.84 ZNF609 zinc finger protein 609 1.53 1.60 SYS1 Sys1 golgi trafficking protein 1.53 2.08 CREM cAMP responsive element modulator 1.52 2.09 TMEM66 transmembrane protein 66 1.52 1.52 EIF2AK1 eukaryotic translation initiation factor 2-alpha kinase 1 1.51 1.80 PALLD palladin, cytoskeletal associated protein 1.51 3.08 PGAM1 phosphoglycerate mutase 1 (brain) 1.51 1.51 CHST12 carbohydrate (chondroitin 4) sulfotransferase 12 1.51 2.57 HNRNPH2 heterogeneous nuclear ribonucleoprotein H2 (H') 1.51 1.59 KIAA0247 KIAA0247 1.51 1.59 ERLIN2 ER lipid raft associated 2 1.51 1.55 FAM127B family with sequence similarity 127, member B 1.50 2.00

Supplementary Table 3 Genes enriched in human podocytes in vitro and mouse podocytes in vivo

Gene Gene Name iPS-derived Mouse Symbol podocytes podocytes FA2H fatty acid 2-hydroxylase 96.90 2.28 ENPEP glutamylaminopeptidase (aminopeptidase A) 70.50 8.32 IL6R interleukin 6 receptor 54.84 2.07 GJA3 gap junction protein, alpha 3, 46kDa 50.83 2.58 NPNT nephronectin 38.77 5.96 TSPAN2 tetraspanin 2 27.98 25.56 CGNL1 cingulin-like 1 20.91 1.96 ZBTB7C zinc finger and BTB domain containing 7C 18.81 2.64 FAM81A family with sequence similarity 81, member A 18.67 3.25 GADD45A growth arrest and DNA-damage-inducible, alpha 14.28 7.26 MMP23B matrix metallopeptidase 23B 14.21 5.03 ADM adrenomedullin 13.69 18.61 TMEM150C transmembrane protein 150C 13.31 8.01 KIRREL kin of IRRE like (Drosophila) 11.84 4.49 AIF1L allograft inflammatory factor 1-like 10.81 4.33 FOXC2 forkhead box C2 (MFH-1, mesenchyme forkhead 1) 10.68 19.02 KIRREL kin of IRRE like (Drosophila) 10.11 4.49 CTTNBP2 cortactin binding protein 2 8.88 3.21 ANXA3 annexin A3 8.79 4.73 AIF1L allograft inflammatory factor 1-like 8.15 4.33 ST6GALNA ST6 (alpha-N-acetyl-neuraminyl-2,3-beta-galactosyl-1,3)-N- 8.04 10.19 C3 acetylgalactosaminide alpha-2,6-sialyltransferase 3 HS3ST6 heparan sulfate (glucosamine) 3-O-sulfotransferase 6 7.93 2.60 ROBO2 roundabout, axon guidance receptor, homolog 2 7.92 15.38 (Drosophila) WIPF3 WAS/WASL interacting protein family, member 3 7.32 5.69 CDC14A cell division cycle 14A 6.88 14.32 RASL11A RAS-like, family 11, member A 6.46 9.22 EPB41L5 erythrocyte membrane protein band 4.1 like 5 6.43 3.16 HS3ST3B1 heparan sulfate (glucosamine) 3-O-sulfotransferase 3B1 6.33 2.23 SRGAP1 SLIT-ROBO Rho GTPase activating protein 1 6.06 24.97 PCED1B PC-esterase domain containing 1B 5.83 3.22 SLC9A3R2 solute carrier family 9, subfamily A (NHE3, cation proton 5.58 2.98 antiporter 3), member 3 regulator 2 PARD6B par-6 family cell polarity regulator beta 5.57 1.88 SH3BGRL2 SH3 domain binding glutamate-rich protein like 2 5.16 3.65 MYLK myosin light chain kinase 5.09 3.03 ARHGAP23 Rho GTPase activating protein 23 5.05 2.60 MYLK myosin light chain kinase 5.03 3.03 EPHX1 epoxide hydrolase 1, microsomal (xenobiotic) 4.82 5.68 NOS1AP nitric oxide synthase 1 (neuronal) adaptor protein 4.66 1.52 ST6GALNA ST6 (alpha-N-acetyl-neuraminyl-2,3-beta-galactosyl-1,3)-N- 4.49 10.19 C3 acetylgalactosaminide alpha-2,6-sialyltransferase 3 ACPP acid phosphatase, prostate 4.46 11.09 EFNB1 ephrin-B1 4.23 4.93 RHPN1 rhophilin, Rho GTPase binding protein 1 4.17 3.31 ARHGEF16 Rho guanine nucleotide exchange factor (GEF) 16 4.15 2.35 LRRFIP1 leucine rich repeat (in FLII) interacting protein 1 4.06 4.90 ST3GAL1 ST3 beta-galactoside alpha-2,3-sialyltransferase 1 4.03 9.85 IFNAR1 interferon (alpha, beta and omega) receptor 1 3.93 1.60 LRRFIP1 leucine rich repeat (in FLII) interacting protein 1 3.85 4.90 ARHGAP28 Rho GTPase activating protein 28 3.85 25.97 ARHGAP28 Rho GTPase activating protein 28 3.84 25.97 NRP1 neuropilin 1 3.70 1.94 C19orf33 chromosome 19 open reading frame 33 3.67 1.74 TSPAN12 tetraspanin 12 3.66 1.84 PLXNB1 plexin B1 3.60 3.07 BTG2 BTG family, member 2 3.60 3.34 SLC6A6 solute carrier family 6 (neurotransmitter transporter), 3.58 1.56 member 6 EZR ezrin 3.54 1.72 SEMA3F sema domain, immunoglobulin domain (Ig), short basic 3.53 1.72 domain, secreted, (semaphorin) 3F DNAJB9 DnaJ (Hsp40) homolog, subfamily B, member 9 3.44 1.88 VASN vasorin 3.41 2.97 FNDC3B fibronectin type III domain containing 3B 3.40 4.62 UNC5C unc-5 homolog C (C. elegans) 3.40 4.82 KREMEN1 kringle containing transmembrane protein 1 3.33 2.44 LATS2 large tumor suppressor kinase 2 3.30 4.00 KREMEN1 kringle containing transmembrane protein 1 3.25 2.44 DUSP3 dual specificity phosphatase 3 3.25 2.46 LRRFIP1 leucine rich repeat (in FLII) interacting protein 1 3.25 4.90 ABCA2 ATP-binding cassette, sub-family A (ABC1), member 2 3.24 2.49 METRNL meteorin, glial cell differentiation regulator-like 3.20 1.96 TPCN1 two pore segment channel 1 3.16 1.56 MSI2 musashi RNA-binding protein 2 3.11 1.79 AKT2 v-akt murine thymoma viral oncogene homolog 2 3.02 1.74 ARHGAP24 Rho GTPase activating protein 24 3.01 3.05 MID1IP1 MID1 interacting protein 1 3.01 1.96 ST6GALNA ST6 (alpha-N-acetyl-neuraminyl-2,3-beta-galactosyl-1,3)-N- 2.97 8.05 C6 acetylgalactosaminide alpha-2,6-sialyltransferase 6 AIF1L allograft inflammatory factor 1-like 2.95 4.33 TMEM245 transmembrane protein 245 2.94 2.01 COL18A1 collagen, type XVIII, alpha 1 2.93 1.96 UCP2 uncoupling protein 2 (mitochondrial, proton carrier) 2.93 1.52 ITM2B integral membrane protein 2B 2.92 1.63 DLC1 deleted in liver cancer 1 2.88 2.67 RBMS1 RNA binding motif, single stranded interacting protein 1 2.86 2.63 LOXL3 lysyl oxidase-like 3 2.83 1.55 ITM2B integral membrane protein 2B 2.82 1.63 GABPB2 GA binding protein transcription factor, beta subunit 2 2.79 2.02 DCDC2 doublecortin domain containing 2 2.77 4.52 GPC3 glypican 3 2.77 3.39 RBMS1 RNA binding motif, single stranded interacting protein 1 2.76 2.63 TANC1 tetratricopeptide repeat, ankyrin repeat and coiled-coil 2.74 1.94 containing 1 UACA uvealautoantigen with coiled-coil domains and ankyrin 2.70 8.09 repeats TPCN1 two pore segment channel 1 2.69 1.56 LOXL3 lysyl oxidase-like 3 2.68 1.55 PLEKHG1 pleckstrin homology domain containing, family G (with Rho 2.66 7.10 Gef domain) member 1 TRIM3 tripartite motif containing 3 2.64 1.81 PDE3A phosphodiesterase 3A, cGMP-inhibited 2.60 2.65 TSPAN5 tetraspanin 5 2.59 4.40 RBMS1 RNA binding motif, single stranded interacting protein 1 2.57 2.63 DACH1 dachshund family transcription factor 1 2.56 10.33 RNF141 ring finger protein 141 2.55 1.50 TMBIM1 transmembrane BAX inhibitor motif containing 1 2.52 3.46 MBLAC2 metallo-beta-lactamase domain containing 2 2.49 2.12 FNDC3B fibronectin type III domain containing 3B 2.47 4.62 GPC3 glypican 3 2.31 3.39 DACH1 dachshund family transcription factor 1 2.30 10.33 SYDE2 synapse defective 1, Rho GTPase, homolog 2 (C. elegans) 2.28 1.81 SBF2 SET binding factor 2 2.26 2.08 AMIGO1 adhesion molecule with Ig-like domain 1 2.23 1.63 MBNL2 muscleblind-like splicing regulator 2 2.22 2.12 ARMCX3 armadillo repeat containing, X-linked 3 2.19 2.29 CRYAB crystallin, alpha B 2.18 1.79 SCARB2 scavenger receptor class B, member 2 2.13 1.64 CAB39 calcium binding protein 39 2.11 1.71 CMTM7 CKLF-like MARVEL transmembrane domain containing 7 2.09 7.01 GNPDA1 glucosamine-6-phosphate deaminase 1 2.06 2.08 ARF6 ADP-ribosylation factor 6 2.05 1.90 NDFIP1 Nedd4 family interacting protein 1 2.05 2.20 ORMDL3 ORMDL sphingolipid biosynthesis regulator 3 2.05 2.26 CRTC1 CREB regulated transcription coactivator 1 2.00 1.83 SDC4 syndecan 4 2.00 4.43 CHRNB1 cholinergic receptor, nicotinic, beta 1 (muscle) 1.99 6.09 YIPF4 Yip1 domain family, member 4 1.99 1.63 PHACTR2 phosphatase and actin regulator 2 1.97 2.55 MSI2 musashi RNA-binding protein 2 1.96 1.79 TOLLIP toll interacting protein 1.95 1.90 ATP6V0A1 ATPase, H+ transporting, lysosomal V0 subunit a1 1.95 2.38 MIB1 mindbomb E3 ubiquitin protein ligase 1 1.95 1.75 SBDS Shwachman-Bodian-Diamond syndrome 1.94 2.35 IKZF2 IKAROS family zinc finger 2 (Helios) 1.94 1.57 SBDS Shwachman-Bodian-Diamond syndrome 1.92 2.35 ORMDL3 ORMDL sphingolipid biosynthesis regulator 3 1.91 2.26 NDFIP1 Nedd4 family interacting protein 1 1.89 2.20 CAB39 calcium binding protein 39 1.88 1.71 PTPRJ protein tyrosine phosphatase, receptor type, J 1.88 1.98 ZFYVE27 zinc finger, FYVE domain containing 27 1.87 1.80 CMTM7 CKLF-like MARVEL transmembrane domain containing 7 1.86 7.01 WASL Wiskott-Aldrich syndrome-like 1.86 2.18 AKT2 v-akt murine thymoma viral oncogene homolog 2 1.85 1.74 EXOC2 exocyst complex component 2 1.84 1.58 C1QTNF7 C1q and tumor necrosis factor related protein 7 1.83 4.38 PHC3 polyhomeotic homolog 3 (Drosophila) 1.83 1.92 DOCK11 dedicator of cytokinesis 11 1.82 3.93 TOM1L2 target of myb1-like 2 (chicken) 1.79 1.72 PRR13 proline rich 13 1.79 1.87 TTC14 tetratricopeptide repeat domain 14 1.78 1.75 NFAT5 nuclear factor of activated T-cells 5, tonicity-responsive 1.78 1.86 PLEKHA5 pleckstrin homology domain containing, family A member 5 1.77 3.10 CFLAR CASP8 and FADD-like apoptosis regulator 1.77 2.03 HIPK3 homeodomain interacting protein kinase 3 1.75 1.92 INTS3 integrator complex subunit 3 1.72 1.96 PDCD6IP programmed cell death 6 interacting protein 1.71 1.66 NRP1 neuropilin 1 1.70 1.94 DCBLD2 discoidin, CUB and LCCL domain containing 2 1.69 4.91 WTIP Wilms tumor 1 interacting protein 1.68 1.63 PHACTR4 phosphatase and actin regulator 4 1.65 3.15 LMTK3 lemur tyrosine kinase 3 1.64 1.50 RRAS2 related RAS viral (r-ras) oncogene homolog 2 1.64 1.77 DCBLD2 discoidin, CUB and LCCL domain containing 2 1.63 4.91 IMPACT impact RWD domain protein 1.63 1.81 P2RX4 purinergic receptor P2X, ligand-gated ion channel, 4 1.62 2.21 ZFP91 ZFP91 zinc finger protein 1.62 1.95 MYOM2 myomesin 2 1.61 18.85 RRAS2 related RAS viral (r-ras) oncogene homolog 2 1.61 1.77 PAPD4 PAP associated domain containing 4 1.60 1.92 CACNA2D2 calcium channel, voltage-dependent, alpha 2/delta subunit 2 1.60 2.66 PHACTR2 phosphatase and actin regulator 2 1.59 2.55 BRD2 bromodomain containing 2 1.59 1.67 DNASE1L1 deoxyribonuclease I-like 1 1.58 2.65 RGS3 regulator of G-protein signaling 3 1.58 1.52 SETD7 SET domain containing (lysine methyltransferase) 7 1.57 6.14 LGALSL lectin, galactoside-binding-like 1.57 5.44 SIGMAR1 sigma non-opioid intracellular receptor 1 1.55 1.68 DCBLD2 discoidin, CUB and LCCL domain containing 2 1.55 4.91 ZFP91 ZFP91 zinc finger protein 1.54 1.95 KDM6A lysine (K)-specific demethylase 6A 1.53 1.51 INO80D INO80 complex subunit D 1.51 1.58 UBE2H ubiquitin-conjugating enzyme E2H 1.50 2.73

Supplementary Table 4 Genes enriched in podocytes in vitro, but not in adult human glomeruli or mouse podocytes in vivo

Gene Gene Name iPS-derived Symbol podocytes LAMP3 lysosomal-associated membrane protein 3 248.08 DSC1 desmocollin 1 144.03 KLRB1 killer cell lectin-like receptor subfamily B, member 1 136.55 SERPIND1 serpin peptidase inhibitor, clade D (heparin cofactor), member 1 133.81 GSDMA gasdermin A 102.44 ST6GALNA ST6 (alpha-N-acetyl-neuraminyl-2,3-beta-galactosyl-1,3)-N-acetylgalactosaminide alpha-2,6- 98.02 C2 sialyltransferase 2 RFPL2 ret finger protein-like 2 94.82 SDPR serum deprivation response 88.06 AADACL4 arylacetamide deacetylase-like 4 73.71 POU2AF1 POU class 2 associating factor 1 58.33 HSD11B1 hydroxysteroid (11-beta) dehydrogenase 1 56.90 LINC01127 long intergenic non-protein coding RNA 1127 54.22 HEPN1 hepatocellular carcinoma, down-regulated 1 48.14 SOST sclerostin 47.27 RFPL1 ret finger protein-like 1 47.12 AQP3 aquaporin 3 (Gill blood group) 45.83 TFF3 trefoil factor 3 (intestinal) 44.84 RSPH10B radial spoke head 10 homolog B (Chlamydomonas) 44.40 DAPL1 death associated protein-like 1 41.83 STON2 stonin 2 40.72 MRGPRF MAS-related GPR, member F 40.47 LINC00488 long intergenic non-protein coding RNA 488 39.38 TFF3 trefoil factor 3 (intestinal) 39.14 TRIM54 tripartite motif containing 54 38.65 PRUNE2 prune homolog 2 (Drosophila) 36.34 MAGI2-IT1 MAGI2 intronic transcript 1 (non-protein coding) 35.46 CAMK2A calcium/calmodulin-dependent protein kinase II alpha 35.22 AQP3 aquaporin 3 (Gill blood group) 34.83 LOC284080 uncharacterized LOC284080 34.46 PLAC1 placenta-specific 1 32.27 HEPACAM hepatic and glial cell adhesion molecule 31.99 TNFRSF8 tumor necrosis factor receptor superfamily, member 8 31.78 FRMD3 FERM domain containing 3 31.59 OLFM3 olfactomedin 3 31.35 TESC tescalcin 31.33 HEPACAM hepatic and glial cell adhesion molecule 31.33 DAPL1 death associated protein-like 1 29.49 FRMD3 FERM domain containing 3 29.02 C21orf37 chromosome 21 open reading frame 37 28.78 CPZ carboxypeptidase Z 28.75 KIRREL2 kin of IRRE like 2 (Drosophila) 28.64 LAMB3 laminin, beta 3 28.34 FAM46C family with sequence similarity 46, member C 28.23 FRMD3 FERM domain containing 3 28.18 DLG2 discs, large homolog 2 (Drosophila) 27.79 C15orf27 chromosome 15 open reading frame 27 27.21 GRIK1-AS1 GRIK1 antisense RNA 1 25.52 PDE6B phosphodiesterase 6B, cGMP-specific, rod, beta 25.10 PAPPA pregnancy-associated plasma protein A, pappalysin 1 24.65 ABLIM2 actin binding LIM protein family, member 2 24.36 LOC100506 uncharacterized LOC100506688 23.99 688 CRYGN crystallin, gamma N 22.99 SBSPON somatomedin B and thrombospondin, type 1 domain containing 22.78 CST2 cystatin SA 22.63 GPA33 glycoprotein A33 (transmembrane) 22.51 ATOH7 atonal homolog 7 (Drosophila) 22.32 ARC activity-regulated cytoskeleton-associated protein 22.14 S100A7 S100 calcium binding protein A7 21.65 G0S2 G0/G1 switch 2 21.64 LINC00961 long intergenic non-protein coding RNA 961 21.32 DMBT1 deleted in malignant brain tumors 1 21.17 FAM81B family with sequence similarity 81, member B 21.08 C2 complement component 2 20.73 DRD4 dopamine receptor D4 20.71 AQP7P1 aquaporin 7 pseudogene 1 20.26 TPPP2 tubulin polymerization-promoting protein family member 2 19.13 DHRS2 dehydrogenase/reductase (SDR family) member 2 19.08 HSBP1L1 heat shock factor binding protein 1-like 1 18.61 IL10RA interleukin 10 receptor, alpha 18.47 TMEM178A transmembrane protein 178A 18.23 SYTL1 synaptotagmin-like 1 18.10 LOC100128 uncharacterized LOC100128340 18.08 340 MRO maestro 17.97 DSC2 desmocollin 2 17.90 NECAB1 N-terminal EF-hand calcium binding protein 1 17.88 NEBL-AS1 NEBL antisense RNA 1 17.29 C2orf91 chromosome 2 open reading frame 91 17.28 MRGPRF MAS-related GPR, member F 17.26 SYTL1 synaptotagmin-like 1 17.13 DNAJB13 DnaJ (Hsp40) homolog, subfamily B, member 13 17.04 LINC01197 long intergenic non-protein coding RNA 1197 16.90 CARD6 caspase recruitment domain family, member 6 16.88 VAMP8 vesicle-associated membrane protein 8 16.60 RASSF10 Ras association (RalGDS/AF-6) domain family (N-terminal) member 10 16.49 VSIG8 V-set and immunoglobulin domain containing 8 16.44 KRT19P2 keratin 19 pseudogene 2 16.42 KRT4 keratin 4 16.42 FREM2 FRAS1 related extracellular matrix protein 2 16.35 FOXL1 forkhead box L1 16.06 CD7 CD7 molecule 16.00 CCDC73 coiled-coil domain containing 73 15.67 MSLN mesothelin 15.64 C15orf59 chromosome 15 open reading frame 59 15.37 KRT19 keratin 19 15.27 ZNF750 zinc finger protein 750 15.11 CHP2 calcineurin-like EF-hand protein 2 15.02 PRUNE2 prune homolog 2 (Drosophila) 15.02 GABRA2 gamma-aminobutyric acid (GABA) A receptor, alpha 2 14.49 FAM129A family with sequence similarity 129, member A 14.41 MYBPC2 myosin binding protein C, fast type 14.35 CATIP ciliogenesis associated TTC17 interacting protein 14.20 DKFZP761C uncharacterized protein DKFZp761C1711 14.14 1711 TSPAN8 tetraspanin 8 14.09 ABLIM2 actin binding LIM protein family, member 2 14.08 RAB11FIP1 RAB11 family interacting protein 1 (class I) 14.00 MYH13 myosin, heavy chain 13, skeletal muscle 13.78 LOC101929 uncharacterized LOC101929494 13.57 494 SFXN2 sideroflexin 2 13.29 SLC51A solute carrier family 51, alpha subunit 13.10 CRTAC1 cartilage acidic protein 1 13.02 GPR133 G protein-coupled receptor 133 12.98 SECTM1 secreted and transmembrane 1 12.92 RASSF10 Ras association (RalGDS/AF-6) domain family (N-terminal) member 10 12.75 LEFTY1 left-right determination factor 1 12.74 GADL1 glutamate decarboxylase-like 1 12.73 MFNG MFNG O-fucosylpeptide 3-beta-N-acetylglucosaminyltransferase 12.59 GFRA3 GDNF family receptor alpha 3 12.34 GPR62 G protein-coupled receptor 62 12.23 EPS8L1 EPS8-like 1 12.21 MIR4697HG MIR4697 host gene (non-protein coding) 12.18 DPPA2 developmental pluripotency associated 2 11.77 FAAH fatty acid amide hydrolase 11.74 RAB11FIP1 RAB11 family interacting protein 1 (class I) 11.71 PNKD paroxysmal nonkinesigenic dyskinesia 11.68 CPNE5 copine V 11.62 CARD6 caspase recruitment domain family, member 6 11.49 ANKRD65 ankyrin repeat domain 65 11.36 LOC100127 uncharacterized LOC100127909 11.24 909 PLCG2 phospholipase C, gamma 2 (phosphatidylinositol-specific) 11.14 GPR98 G protein-coupled receptor 98 11.09 CCDC109B coiled-coil domain containing 109B 10.93 LOC101929 uncharacterized LOC101929687 10.72 687 CRB2 crumbs family member 2 10.72 TMEM139 transmembrane protein 139 10.59 CPEB1 cytoplasmic polyadenylation element binding protein 1 10.58 ZMIZ1-AS1 ZMIZ1 antisense RNA 1 10.55 GNG13 guanine nucleotide binding protein (G protein), gamma 13 10.53 ADRA2C adrenoceptor alpha 2C 10.40 TFF3 trefoil factor 3 (intestinal) 10.29 LILRB3 leukocyte immunoglobulin-like receptor, subfamily B (with TM and ITIM domains), member 3 10.21 PCP4 Purkinje cell protein 4 9.94 KLHL30 kelch-like family member 30 9.75 PLEKHB1 pleckstrin homology domain containing, family B (evectins) member 1 9.63 SMIM5 small integral membrane protein 5 9.60 GPX3 glutathione peroxidase 3 (plasma) 9.60 RAB37 RAB37, member RAS oncogene family 9.56 FOXD2 forkhead box D2 9.54 SULT1C2 sulfotransferase family, cytosolic, 1C, member 2 9.53 MLC1 megalencephalic leukoencephalopathy with subcortical cysts 1 9.53 DNAJC6 DnaJ (Hsp40) homolog, subfamily C, member 6 9.51 TMCC3 transmembrane and coiled-coil domain family 3 9.42 KIAA2022 KIAA2022 9.34 MYH3 myosin, heavy chain 3, skeletal muscle, embryonic 9.31 FUT1 fucosyltransferase 1 (galactoside 2-alpha-L-fucosyltransferase, H blood group) 9.25 GPX3 glutathione peroxidase 3 (plasma) 9.21 CD22 CD22 molecule 9.15 CCDC122 coiled-coil domain containing 122 9.14 SOWAHA sosondowah ankyrin repeat domain family member A 9.11 TRADD TNFRSF1A-associated via death domain 9.11 S100A2 S100 calcium binding protein A2 9.07 GPD1 glycerol-3-phosphate dehydrogenase 1 (soluble) 9.02 SLC16A1 solute carrier family 16 (monocarboxylate transporter), member 1 8.95 SYPL2 synaptophysin-like 2 8.91 GPR126 G protein-coupled receptor 126 8.88 LRRC36 leucine rich repeat containing 36 8.85 C2 complement component 2 8.84 ASPG asparaginase 8.83 ENPP5 ectonucleotide pyrophosphatase/phosphodiesterase 5 (putative) 8.79 F13A1 coagulation factor XIII, A1 polypeptide 8.76 EMID1 EMI domain containing 1 8.70 RIC3 RIC3 acetylcholine receptor chaperone 8.64 ADAMTSL4 ADAMTS-like 4 8.49 UNC5B-AS1 UNC5B antisense RNA 1 8.40 RPH3AL rabphilin 3A-like (without C2 domains) 8.34 TMEM40 transmembrane protein 40 8.27 XKR4 XK, Kell blood group complex subunit-related family, member 4 8.25 SYTL4 synaptotagmin-like 4 8.22 FRAS1 Fraser syndrome 1 8.22 ANXA9 annexin A9 8.16 FAM174B family with sequence similarity 174, member B 8.14 INMT indolethylamine N-methyltransferase 8.13 SYPL2 synaptophysin-like 2 8.11 DUSP23 dual specificity phosphatase 23 8.11 CLEC4M C-type lectin domain family 4, member M 8.07 C15orf52 chromosome 15 open reading frame 52 7.98 LRFN2 leucine rich repeat and fibronectin type III domain containing 2 7.94 DOC2B double C2-like domains, beta 7.90 STAC2 SH3 and cysteine rich domain 2 7.85 NMRK1 nicotinamide riboside kinase 1 7.83 NMRK1 nicotinamide riboside kinase 1 7.79 CILP2 cartilage intermediate layer protein 2 7.78 RASSF5 Ras association (RalGDS/AF-6) domain family member 5 7.68 GGT5 gamma-glutamyltransferase 5 7.67 WSCD2 WSC domain containing 2 7.64 NCCRP1 non-specific cytotoxic cell receptor protein 1 homolog (zebrafish) 7.63 FDXR ferredoxin reductase 7.56 XKR4 XK, Kell blood group complex subunit-related family, member 4 7.55 LOC727808 uncharacterized LOC727808 7.50 NDUFA4L2 NADH dehydrogenase (ubiquinone) 1 alpha subcomplex, 4-like 2 7.46 TCP11L2 t-complex 11, testis-specific-like 2 7.43 S100A6 S100 calcium binding protein A6 7.40 RAB6B RAB6B, member RAS oncogene family 7.26 XK X-linked Kx blood group (McLeod syndrome) 7.26 PAX8 paired box 8 7.19 PRUNE2 prune homolog 2 (Drosophila) 7.18 DEPTOR DEP domain containing MTOR-interacting protein 7.18 SHISA2 shisa family member 2 7.16 HKDC1 hexokinase domain containing 1 7.13 FAAH fatty acid amide hydrolase 7.09 C1QL4 complement component 1, q subcomponent-like 4 7.08 ABCA4 ATP-binding cassette, sub-family A (ABC1), member 4 7.04 EPSTI1 epithelial stromal interaction 1 (breast) 7.03 MT1M metallothionein 1M 7.00 MIR503HG MIR503 host gene (non-protein coding) 6.97 MGST1 microsomal glutathione S-transferase 1 6.94 GMPR guanosine monophosphate reductase 6.90 CILP cartilage intermediate layer protein, nucleotide pyrophosphohydrolase 6.86 CLDN1 claudin 1 6.86 FMO4 flavin containing monooxygenase 4 6.84 CCDC178 coiled-coil domain containing 178 6.84 LIM2 lens intrinsic membrane protein 2, 19kDa 6.84 C11orf52 chromosome 11 open reading frame 52 6.82 PNPLA7 patatin-like phospholipase domain containing 7 6.75 PPARGC1B peroxisome proliferator-activated receptor gamma, coactivator 1 beta 6.73 GNAO1 guanine nucleotide binding protein (G protein), alpha activating activity polypeptide O 6.73 PIGZ phosphatidylinositol glycan anchor biosynthesis, class Z 6.72 LRRN2 leucine rich repeat neuronal 2 6.69 ABCC3 ATP-binding cassette, sub-family C (CFTR/MRP), member 3 6.68 SUSD2 sushi domain containing 2 6.66 PAQR7 progestin and adipoQ receptor family member VII 6.61 SGK1 serum/glucocorticoid regulated kinase 1 6.57 CPXM1 carboxypeptidase X (M14 family), member 1 6.57 SLC9A3R1 solute carrier family 9, subfamily A (NHE3, cation proton antiporter 3), member 3 regulator 1 6.53 LMNTD2 lamin tail domain containing 2 6.52 GFRA2 GDNF family receptor alpha 2 6.50 CXCL1 chemokine (C-X-C motif) ligand 1 (melanoma growth stimulating activity, alpha) 6.48 COLEC11 collectin sub-family member 11 6.45 FAM83G family with sequence similarity 83, member G 6.35 SULT1C2 sulfotransferase family, cytosolic, 1C, member 2 6.29 TRIM8 tripartite motif containing 8 6.29 DOK7 docking protein 7 6.23 UTF1 undifferentiated embryonic cell transcription factor 1 6.21 IP6K3 inositol hexakisphosphate kinase 3 6.21 PAPLN papilin, proteoglycan-like sulfated glycoprotein 6.20 LRRC4C leucine rich repeat containing 4C 6.14 CFHR3 complement factor H-related 3 6.09 MT1X metallothionein 1X 6.03 KREMEN2 kringle containing transmembrane protein 2 6.02 ROM1 retinal outer segment membrane protein 1 6.01 PFKP phosphofructokinase, platelet 5.99 RERG RAS-like, estrogen-regulated, growth inhibitor 5.99 TRIB1 tribbles pseudokinase 1 5.95 FAM183A family with sequence similarity 183, member A 5.94 FRAS1 Fraser syndrome 1 5.94 LINC00472 long intergenic non-protein coding RNA 472 5.91 PFKP phosphofructokinase, platelet 5.88 ITGB8 integrin, beta 8 5.85 DPF3 D4, zinc and double PHD fingers, family 3 5.83 DACH2 dachshund family transcription factor 2 5.81 SLC15A1 solute carrier family 15 (oligopeptide transporter), member 1 5.81 PPARGC1B peroxisome proliferator-activated receptor gamma, coactivator 1 beta 5.81 IRX4 iroquois homeobox 4 5.79 ALPL alkaline phosphatase, liver/bone/kidney 5.76 SH3RF1 SH3 domain containing ring finger 1 5.74 ULK4 unc-51 like kinase 4 5.74 GCNT1 glucosaminyl (N-acetyl) transferase 1, core 2 5.69 KANK3 KN motif and ankyrin repeat domains 3 5.66 THBD thrombomodulin 5.65 CTSL cathepsin L 5.65 UBTD1 ubiquitin domain containing 1 5.63 NKAIN4 Na+/K+ transporting ATPase interacting 4 5.63 STON1- STON1-GTF2A1L readthrough 5.62 GTF2A1L WASH5P WAS protein family homolog 5 pseudogene 5.62 NRG3 neuregulin 3 5.62 KRT80 keratin 80 5.60 SLC6A13 solute carrier family 6 (neurotransmitter transporter), member 13 5.58 TMEM130 transmembrane protein 130 5.57 SLC7A7 solute carrier family 7 (amino acid transporter light chain, y+L system), member 7 5.56 CPEB4 cytoplasmic polyadenylation element binding protein 4 5.54 IL4I1 interleukin 4 induced 1 5.51 LOC100131 uncharacterized LOC100131043 5.51 043 CCNB3 cyclin B3 5.50 NFE2L3 nuclear factor, erythroid 2-like 3 5.49 SLC2A1 solute carrier family 2 (facilitated glucose transporter), member 1 5.45 STXBP2 syntaxin binding protein 2 5.42 FAM183B acyloxyacyl hydrolase (neutrophil) 5.41 CPAMD8 C3 and PZP-like, alpha-2-macroglobulin domain containing 8 5.39 ST8SIA5 ST8 alpha-N-acetyl-neuraminide alpha-2,8-sialyltransferase 5 5.37 GRTP1 growth hormone regulated TBC protein 1 5.37 PRSS16 protease, serine, 16 (thymus) 5.36 PTPLB protein tyrosine phosphatase-like (proline instead of catalytic arginine), member b 5.34 C11orf71 chromosome 11 open reading frame 71 5.33 VEPH1 ventricular zone expressed PH domain-containing 1 5.29 NRG3 neuregulin 3 5.28 CCBE1 collagen and calcium binding EGF domains 1 5.26 ICOSLG inducible T-cell co-stimulator ligand 5.24 SNX25 sorting nexin 25 5.22 MLXIPL MLX interacting protein-like 5.20 ITGA6 integrin, alpha 6 5.18 SPATA13 spermatogenesis associated 13 5.18 KISS1R KISS1 receptor 5.18 LOC102725 uncharacterized LOC102725171 5.15 171 NTN4 netrin 4 5.13 ADAP1 ArfGAP with dual PH domains 1 5.10 SPATA13 spermatogenesis associated 13 5.07 MUSTN1 musculoskeletal, embryonic nuclear protein 1 5.07 CECR5-AS1 CECR5 antisense RNA 1 5.05 SMTNL2 smoothelin-like 2 5.03 CDK20 cyclin-dependent kinase 20 5.00 PLCD3 phospholipase C, delta 3 5.00 IL10RB interleukin 10 receptor, beta 5.00 CDS1 CDP-diacylglycerol synthase (phosphatidate cytidylyltransferase) 1 4.99 C1orf226 chromosome 1 open reading frame 226 4.99 S100A1 S100 calcium binding protein A1 4.98 WWC2-AS2 WWC2 antisense RNA 2 4.94 FZD6 frizzled class receptor 6 4.90 SPATA13 spermatogenesis associated 13 4.90 RASAL1 RAS protein activator like 1 (GAP1 like) 4.87 GRHL1 grainyhead-like 1 (Drosophila) 4.81 AP1S3 adaptor-related protein complex 1, sigma 3 subunit 4.81 SELV selenoprotein V 4.76 FOSB FBJ murine osteosarcoma viral oncogene homolog B 4.74 FAM20C family with sequence similarity 20, member C 4.73 SLC7A8 solute carrier family 7 (amino acid transporter light chain, L system), member 8 4.72 MFSD6 major facilitator superfamily domain containing 6 4.72 TMEM64 transmembrane protein 64 4.70 CDS1 CDP-diacylglycerol synthase (phosphatidate cytidylyltransferase) 1 4.69 HIPK2 homeodomain interacting protein kinase 2 4.69 NUDT18 nudix (nucleoside diphosphate linked moiety X)-type motif 18 4.68 ITGA6 integrin, alpha 6 4.68 AFAP1L2 actin filament associated protein 1-like 2 4.67 LYPD1 LY6/PLAUR domain containing 1 4.65 PARD6G par-6 family cell polarity regulator gamma 4.61 TNXB tenascin XB 4.61 FREM1 FRAS1 related extracellular matrix 1 4.58 ABCC9 ATP-binding cassette, sub-family C (CFTR/MRP), member 9 4.57 CDK20 cyclin-dependent kinase 20 4.56 MROH1 maestro heat-like repeat family member 1 4.56 LRRC56 leucine rich repeat containing 56 4.55 LST1 leukocyte specific transcript 1 4.50 LMNA lamin A/C 4.50 RARRES3 retinoic acid receptor responder (tazarotene induced) 3 4.50 FAM101B family with sequence similarity 101, member B 4.49 LINC00173 long intergenic non-protein coding RNA 173 4.48 SLC22A18 solute carrier family 22, member 18 4.46 SLC16A3 solute carrier family 16 (monocarboxylate transporter), member 3 4.46 APLN apelin 4.42 SEPP1 selenoprotein P, plasma, 1 4.41 ANKH ANKH inorganic pyrophosphate transport regulator 4.39 SEMA4G sema domain, immunoglobulin domain (Ig), transmembrane domain (TM) and short 4.37 cytoplasmic domain, (semaphorin) 4G LOC102725 uncharacterized LOC102725171 4.35 171 TSTD1 thiosulfate sulfurtransferase (rhodanese)-like domain containing 1 4.35 OSTF1 osteoclast stimulating factor 1 4.34 SELV selenoprotein V 4.31 ANKH ANKH inorganic pyrophosphate transport regulator 4.31 CHMP4C charged multivesicular body protein 4C 4.30 S100A16 S100 calcium binding protein A16 4.30 LSR lipolysis stimulated lipoprotein receptor 4.29 LRRC9 leucine rich repeat containing 9 4.26 PTPN6 protein tyrosine phosphatase, non-receptor type 6 4.25 ACOX2 acyl-CoA oxidase 2, branched chain 4.25 TTC18 tetratricopeptide repeat domain 18 4.24 KCTD16 potassium channel tetramerization domain containing 16 4.20 NKAIN2 Na+/K+ transporting ATPase interacting 2 4.19 LYPD1 LY6/PLAUR domain containing 1 4.19 CPXM2 carboxypeptidase X (M14 family), member 2 4.19 HOXA4 homeobox A4 4.19 AP1M2 adaptor-related protein complex 1, mu 2 subunit 4.17 ASS1 argininosuccinate synthase 1 4.14 LMNA lamin A/C 4.13 KCNQ1 potassium voltage-gated channel, KQT-like subfamily, member 1 4.11 RAB37 RAB37, member RAS oncogene family 4.10 TCTA T-cell leukemia translocation altered 4.09 CLDN7 claudin 7 4.08 FAM83H family with sequence similarity 83, member H 4.08 IGFALS insulin-like growth factor binding protein, acid labile subunit 4.06 SESN2 sestrin 2 4.06 PAWR PRKC, apoptosis, WT1, regulator 4.05 PKP2 plakophilin 2 4.04 SIAE sialic acid acetylesterase 4.03 BNC1 basonuclin 1 4.03 IGSF1 immunoglobulin superfamily, member 1 4.02 ZDHHC12 zinc finger, DHHC-type containing 12 4.00 SLC16A10 solute carrier family 16 (aromatic amino acid transporter), member 10 3.98 HIPK2 homeodomain interacting protein kinase 2 3.98 SLC15A3 solute carrier family 15 (oligopeptide transporter), member 3 3.97 FOXD2-AS1 FOXD2 antisense RNA 1 (head to head) 3.97 NAGLU N-acetylglucosaminidase, alpha 3.97 ZDHHC12 zinc finger, DHHC-type containing 12 3.96 ADHFE1 alcohol dehydrogenase, iron containing, 1 3.96 WFIKKN1 WAP, follistatin/kazal, immunoglobulin, kunitz and netrin domain containing 1 3.94 EML3 echinoderm microtubule associated protein like 3 3.92 RASD2 RASD family, member 2 3.92 FRMD4B FERM domain containing 4B 3.92 NOXA1 NADPH oxidase activator 1 3.92 IQSEC2 IQ motif and Sec7 domain 2 3.92 ELL2 elongation factor, RNA polymerase II, 2 3.92 ZBED5-AS1 ZBED5 antisense RNA 1 3.91 EXOC3L2 exocyst complex component 3-like 2 3.90 LYPD1 LY6/PLAUR domain containing 1 3.90 LOC100996 phosphodiesterase 4D interacting protein-like 3.89 724 DICER1- DICER1 antisense RNA 1 3.89 AS1 ZDHHC2 zinc finger, DHHC-type containing 2 3.89 FOXJ1 forkhead box J1 3.88 MAFK v-maf avian musculoaponeurotic fibrosarcoma oncogene homolog K 3.86 HES4 hes family bHLH transcription factor 4 3.86 BDNF-AS BDNF antisense RNA 3.85 CASC2 cancer susceptibility candidate 2 (non-protein coding) 3.85 PPP1R3E protein phosphatase 1, regulatory subunit 3E 3.85 FOXO4 forkhead box O4 3.85 SMPDL3B sphingomyelin phosphodiesterase, acid-like 3B 3.84 FOS FBJ murine osteosarcoma viral oncogene homolog 3.84 LINC00957 long intergenic non-protein coding RNA 957 3.83 KLHDC9 kelch domain containing 9 3.82 SELT selenoprotein T 3.79 CXADR coxsackie virus and adenovirus receptor 3.78 ADI1 acireductone dioxygenase 1 3.77 ASS1 argininosuccinate synthase 1 3.77 HEY2 hes-related family bHLH transcription factor with YRPW motif 2 3.76 RGS9 regulator of G-protein signaling 9 3.76 TMEM64 transmembrane protein 64 3.75 LOC730183 uncharacterized LOC730183 3.74 CYB561 cytochrome b561 3.74 AGPAT9 1-acylglycerol-3-phosphate O-acyltransferase 9 3.73 NUP210 nucleoporin 210kDa 3.73 C17orf97 chromosome 17 open reading frame 97 3.72 CMTM4 CKLF-like MARVEL transmembrane domain containing 4 3.69 MAOA monoamine oxidase A 3.69 PNPLA2 patatin-like phospholipase domain containing 2 3.68 C6orf165 chromosome 6 open reading frame 165 3.68 TEAD4 TEA domain family member 4 3.67 SLC6A12 solute carrier family 6 (neurotransmitter transporter), member 12 3.66 IGFBP4 insulin-like growth factor binding protein 4 3.66 C16orf74 chromosome 16 open reading frame 74 3.66 ABTB1 ankyrin repeat and BTB (POZ) domain containing 1 3.66 LLGL2 lethal giant larvae homolog 2 (Drosophila) 3.66 FOLR1 folate receptor 1 (adult) 3.66 TRNP1 TMF1-regulated nuclear protein 1 3.65 PPP1R1C protein phosphatase 1, regulatory (inhibitor) subunit 1C 3.64 AAED1 AhpC/TSA antioxidant enzyme domain containing 1 3.63 CXADR coxsackie virus and adenovirus receptor 3.62 S1PR1 sphingosine-1-phosphate receptor 1 3.62 MAG myelin associated glycoprotein 3.62 WNK2 WNK lysine deficient protein kinase 2 3.61 CFD complement factor D (adipsin) 3.61 IQSEC2 IQ motif and Sec7 domain 2 3.60 TM7SF2 transmembrane 7 superfamily member 2 3.59 PARP9 poly (ADP-ribose) polymerase family, member 9 3.59 SELT selenoprotein T 3.58 RSPH3 radial spoke 3 homolog (Chlamydomonas) 3.57 WNK2 WNK lysine deficient protein kinase 2 3.57 ATP8A2 ATPase, aminophospholipid transporter, class I, type 8A, member 2 3.55 DIAPH3 diaphanous-related formin 3 3.54 FZD9 frizzled class receptor 9 3.53 SLC4A11 solute carrier family 4, sodium borate transporter, member 11 3.53 TRABD2B TraB domain containing 2B 3.53 FAM46B family with sequence similarity 46, member B 3.52 LMX1B LIM homeobox transcription factor 1, beta 3.52 TBC1D10C TBC1 domain family, member 10C 3.52 VMAC vimentin-type intermediate filament associated coiled-coil protein 3.50 TST thiosulfate sulfurtransferase (rhodanese) 3.50 LHPP phospholysine phosphohistidine inorganic pyrophosphate phosphatase 3.50 GRID1 glutamate receptor, ionotropic, delta 1 3.49 ATP1A1 ATPase, Na+/K+ transporting, alpha 1 polypeptide 3.49 LOC646762 uncharacterized LOC646762 3.46 SNX25 sorting nexin 25 3.45 POGLUT1 protein O-glucosyltransferase 1 3.44 SEPP1 selenoprotein P, plasma, 1 3.44 ARSD arylsulfatase D 3.43 ADAMTS9 ADAM metallopeptidase with thrombospondin type 1 motif, 9 3.43 TP53TG3 TP53 target 3 3.42 GHDC GH3 domain containing 3.42 CARF calcium responsive transcription factor 3.41 MLXIP MLX interacting protein 3.41 ME3 malic enzyme 3, NADP(+)-dependent, mitochondrial 3.40 INSIG1 insulin induced gene 1 3.38 MMP15 matrix metallopeptidase 15 (membrane-inserted) 3.38 S100A13 S100 calcium binding protein A13 3.37 SCN2B sodium channel, voltage-gated, type II, beta subunit 3.37 SHANK2 SH3 and multiple ankyrin repeat domains 2 3.36 IGSF9B immunoglobulin superfamily, member 9B 3.36 ATP1A4 ATPase, Na+/K+ transporting, alpha 4 polypeptide 3.35 PQLC3 PQ loop repeat containing 3 3.34 ACCS 1-aminocyclopropane-1-carboxylate synthase homolog (Arabidopsis)(non-functional) 3.32 FMOD fibromodulin 3.31 LOC100049 uncharacterized LOC100049716 3.31 716 ADI1 acireductone dioxygenase 1 3.31 LOC102723 uncharacterized LOC102723456 3.31 456 ZNF396 zinc finger protein 396 3.30 CCIN calicin 3.28 DNAJB12 DnaJ (Hsp40) homolog, subfamily B, member 12 3.28 ATP1A4 ATPase, Na+/K+ transporting, alpha 4 polypeptide 3.27 LRRC8A leucine rich repeat containing 8 family, member A 3.26 ECHDC2 enoyl CoA hydratase domain containing 2 3.26 SLC25A26 solute carrier family 25 (S-adenosylmethionine carrier), member 26 3.26 ACSL1 acyl-CoA synthetase long-chain family member 1 3.25 ESYT2 extended synaptotagmin-like protein 2 3.25 WDR66 WD repeat domain 66 3.24 C5orf15 chromosome 5 open reading frame 15 3.24 ARRDC1 arrestin domain containing 1 3.24 CDH3 cadherin 3, type 1, P-cadherin (placental) 3.24 SNX8 sorting nexin 8 3.24 PAXIP1- PAXIP1 antisense RNA 1 (head to head) 3.23 AS1 NKAIN4 Na+/K+ transporting ATPase interacting 4 3.22 CANX calnexin 3.22 LYPD6B LY6/PLAUR domain containing 6B 3.22 CASZ1 castor zinc finger 1 3.22 LOC100129 uncharacterized LOC100129846 3.22 846 ADAT2 adenosine deaminase, tRNA-specific 2 3.22 CA11 carbonic anhydrase XI 3.21 OR7E12P olfactory receptor, family 7, subfamily E, member 12 pseudogene 3.21 MARCH3 membrane-associated ring finger (C3HC4) 3, E3 ubiquitin protein ligase 3.20 C1orf213 chromosome 1 open reading frame 213 3.20 ADAMTSL4 ADAMTS-like 4 3.19 LOC100653 uncharacterized LOC100653296 3.19 296 THAP8 THAP domain containing 8 3.18 IMPDH1 IMP (inosine 5'-monophosphate) dehydrogenase 1 3.17 SCAMP1- SCAMP1 antisense RNA 1 3.16 AS1 PRSS23 protease, serine, 23 3.15 GPHN gephyrin 3.15 KMO kynurenine 3-monooxygenase (kynurenine 3-hydroxylase) 3.14 GAS7 growth arrest-specific 7 3.13 COMTD1 catechol-O-methyltransferase domain containing 1 3.12 TMEM30A transmembrane protein 30A 3.11 HAGLR HOXD antisense growth-associated long non-coding RNA 3.10 ZMAT1 zinc finger, matrin-type 1 3.09 SMCO4 single-pass membrane protein with coiled-coil domains 4 3.09 WDR86 WD repeat domain 86 3.09 EMILIN2 elastin microfibril interfacer 2 3.09 JDP2 Jun dimerization protein 2 3.08 EIF3M eukaryotic translation initiation factor 3, subunit M 3.08 FBXO21 F-box protein 21 3.08 PCBD1 pterin-4 alpha-carbinolamine dehydratase/dimerization cofactor of hepatocyte nuclear factor 3.06 1 alpha C5orf15 chromosome 5 open reading frame 15 3.06 TPP1 tripeptidyl peptidase I 3.06 ZNRF3 zinc and ring finger 3 3.05 RNF139 ring finger protein 139 3.04 OCA2 oculocutaneous albinism II 3.04 TMEM191A transmembrane protein 191A (pseudogene) 3.04 LDLRAP1 low density lipoprotein receptor adaptor protein 1 3.04 KLHL29 kelch-like family member 29 3.03 F11R F11 receptor 3.02 ADPRH ADP-ribosylarginine hydrolase 3.02 JKAMP JNK1/MAPK8-associated membrane protein 3.01 ODF3B outer dense fiber of sperm tails 3B 3.01 PIM2 pim-2 oncogene 3.01 MAOB monoamine oxidase B 3.00 LRRC4 leucine rich repeat containing 4 3.00 CCDC9 coiled-coil domain containing 9 3.00 LOC254896 uncharacterized LOC254896 3.00 SCARNA9 small Cajal body-specific RNA 9 2.99 SIRT5 sirtuin 5 2.99 EXD3 exonuclease 3'-5' domain containing 3 2.99 CBR4 carbonyl reductase 4 2.98 TCTN1 tectonic family member 1 2.98 GPSM1 G-protein signaling modulator 1 2.96 SRGAP2C SLIT-ROBO Rho GTPase activating protein 2C 2.95 AJUBA ajuba LIM protein 2.95 KCP kielin/chordin-like protein 2.95 RBM38 RNA binding motif protein 38 2.94 SNORD116- small nucleolar RNA, C/D box 116-19 2.94 19 KIF1A kinesin family member 1A 2.94 ITFG1 integrin alpha FG-GAP repeat containing 1 2.94 TEX41 testis expressed 41 (non-protein coding) 2.93 SKAP1 src kinase associated phosphoprotein 1 2.93 PPP1R9A protein phosphatase 1, regulatory subunit 9A 2.91 MYLK2 myosin light chain kinase 2 2.91 PAN2 PAN2 poly(A) specific ribonuclease subunit 2.91 MVB12A multivesicular body subunit 12A 2.91 LOC100507 uncharacterized LOC100507547 2.91 547 SLC37A4 solute carrier family 37 (glucose-6-phosphate transporter), member 4 2.91 TSPAN3 tetraspanin 3 2.91 RAB3IL1 RAB3A interacting protein (rabin3)-like 1 2.91 LINC01003 long intergenic non-protein coding RNA 1003 2.91 MAP3K5 mitogen-activated protein kinase kinase kinase 5 2.90 C6orf57 chromosome 6 open reading frame 57 2.90 CDKN2C cyclin-dependent kinase inhibitor 2C (p18, inhibits CDK4) 2.89 TOLLIP- TOLLIP antisense RNA 1 (head to head) 2.89 AS1 HRASLS5 HRAS-like suppressor family, member 5 2.87 LHPP phospholysine phosphohistidine inorganic pyrophosphate phosphatase 2.87 KCNE3 potassium voltage-gated channel, Isk-related family, member 3 2.87 GPR162 G protein-coupled receptor 162 2.86 SPR sepiapterin reductase (7,8-dihydrobiopterin:NADP+ oxidoreductase) 2.85 PTGR2 prostaglandin reductase 2 2.84 ZNF524 zinc finger protein 524 2.84 STX1A syntaxin 1A (brain) 2.83 TCEAL6 transcription elongation factor A (SII)-like 6 2.83 LOC100288 uncharacterized LOC100288911 2.83 911 DTWD2 DTW domain containing 2 2.82 PHYKPL 5-phosphohydroxy-L-lysine phospho-lyase 2.82 RNF208 ring finger protein 208 2.82 ATP11A ATPase, class VI, type 11A 2.82 GLB1L2 galactosidase, beta 1-like 2 2.81 RNF208 ring finger protein 208 2.81 SNHG20 small nucleolar RNA host gene 20 (non-protein coding) 2.81 DAK dihydroxyacetone kinase 2 homolog (S. cerevisiae) 2.81 SH3KBP1 SH3-domain kinase binding protein 1 2.81 DNALI1 dynein, axonemal, light intermediate chain 1 2.80 AMFR autocrine motility factor receptor, E3 ubiquitin protein ligase 2.80 ABLIM2 actin binding LIM protein family, member 2 2.80 CLDN3 claudin 3 2.80 SLC9A7 solute carrier family 9, subfamily A (NHE7, cation proton antiporter 7), member 7 2.80 PEBP4 phosphatidylethanolamine-binding protein 4 2.80 C10orf25 chromosome 10 open reading frame 25 2.79 COX7A1 cytochrome c oxidase subunit VIIa polypeptide 1 (muscle) 2.79 KIAA1522 KIAA1522 2.79 SMIM1 small integral membrane protein 1 (Vel blood group) 2.78 PCSK4 proprotein convertase subtilisin/kexin type 4 2.78 DOK1 docking protein 1, 62kDa (downstream of tyrosine kinase 1) 2.77 RILPL2 Rab interacting lysosomal protein-like 2 2.77 ARHGEF15 Rho guanine nucleotide exchange factor (GEF) 15 2.76 FRS2 fibroblast growth factor receptor substrate 2 2.76 ABTB1 ankyrin repeat and BTB (POZ) domain containing 1 2.76 HIST1H2AC histone cluster 1, H2ac 2.76 TNXB tenascin XB 2.75 TCEAL6 transcription elongation factor A (SII)-like 6 2.75 NOL4L nucleolar protein 4-like 2.75 PPM1J protein phosphatase, Mg2+/Mn2+ dependent, 1J 2.74 KIAA1671 KIAA1671 2.74 FSIP2 fibrous sheath interacting protein 2 2.74 SKAP2 src kinase associated phosphoprotein 2 2.73 LPIN3 lipin 3 2.73 ANAPC16 anaphase promoting complex subunit 16 2.72 C8orf58 chromosome 8 open reading frame 58 2.72 MTR 5-methyltetrahydrofolate-homocysteine methyltransferase 2.72 C6orf203 chromosome 6 open reading frame 203 2.72 AK8 adenylate kinase 8 2.71 CST5 cystatin D 2.71 RELL1 RELT-like 1 2.71 SNX30 sorting nexin family member 30 2.71 LMNA lamin A/C 2.71 EXTL1 exostosin-like glycosyltransferase 1 2.71 FAM63A family with sequence similarity 63, member A 2.70 NIPA1 non imprinted in Prader-Willi/Angelman syndrome 1 2.70 LOC728061 hCG2003663 2.70 LTBR lymphotoxin beta receptor (TNFR superfamily, member 3) 2.70 TMEM179B transmembrane protein 179B 2.70 ARID3B AT rich interactive domain 3B (BRIGHT-like) 2.70 ZFP36L2 ZFP36 ring finger protein-like 2 2.70 RIBC1 RIB43A domain with coiled-coils 1 2.69 TMEM30A transmembrane protein 30A 2.69 SH3GLB2 SH3-domain GRB2-like endophilin B2 2.69 GRB7 growth factor receptor-bound protein 7 2.69 BAI1 brain-specific angiogenesis inhibitor 1 2.69 ARAP1 ArfGAP with RhoGAP domain, ankyrin repeat and PH domain 1 2.68 TANGO2 transport and golgi organization 2 homolog (Drosophila) 2.68 WDSUB1 WD repeat, sterile alpha motif and U-box domain containing 1 2.68 KRT18 keratin 18 2.68 PCMTD2 protein-L-isoaspartate (D-aspartate) O-methyltransferase domain containing 2 2.68 ADAMTS1 ADAM metallopeptidase with thrombospondin type 1 motif, 1 2.68 TMEM132A transmembrane protein 132A 2.68 CORO2A coronin, actin binding protein, 2A 2.67 TMCO4 transmembrane and coiled-coil domains 4 2.67 TYROBP TYRO protein tyrosine kinase binding protein 2.67 CAPN7 calpain 7 2.67 TSPAN33 tetraspanin 33 2.67 HSD11B2 hydroxysteroid (11-beta) dehydrogenase 2 2.67 LOC100507 uncharacterized LOC100507547 2.67 547 F11R F11 receptor 2.67 PCYOX1L prenylcysteine oxidase 1 like 2.66 SPSB1 splA/ryanodine receptor domain and SOCS box containing 1 2.66 PPP1R16A protein phosphatase 1, regulatory subunit 16A 2.66 PIGP phosphatidylinositol glycan anchor biosynthesis, class P 2.66 SUMF1 sulfatase modifying factor 1 2.66 FURIN furin (paired basic amino acid cleaving enzyme) 2.66 CCDC96 coiled-coil domain containing 96 2.65 MYO1F myosin IF 2.65 ZNRF3 zinc and ring finger 3 2.65 NAT8L N-acetyltransferase 8-like (GCN5-related, putative) 2.64 SNTB1 syntrophin, beta 1 (dystrophin-associated protein A1, 59kDa, basic component 1) 2.64 VAPA VAMP (vesicle-associated membrane protein)-associated protein A, 33kDa 2.64 BFSP1 beaded filament structural protein 1, filensin 2.64 PNPLA4 patatin-like phospholipase domain containing 4 2.63 SLC26A11 solute carrier family 26 (anion exchanger), member 11 2.62 NEFH neurofilament, heavy polypeptide 2.62 SLC9A7 solute carrier family 9, subfamily A (NHE7, cation proton antiporter 7), member 7 2.62 RNF13 ring finger protein 13 2.62 CNDP2 CNDP dipeptidase 2 (metallopeptidase M20 family) 2.62 TMEM170B transmembrane protein 170B 2.62 FBXL14 F-box and leucine-rich repeat protein 14 2.62 GSDMD gasdermin D 2.62 CHRD chordin 2.61 KRT18P55 keratin 18 pseudogene 55 2.61 LHX1 LIM homeobox 1 2.61 IFNAR2 interferon (alpha, beta and omega) receptor 2 2.61 SLC16A5 solute carrier family 16 (monocarboxylate transporter), member 5 2.61 MTR 5-methyltetrahydrofolate-homocysteine methyltransferase 2.61 MLLT4 myeloid/lymphoid or mixed-lineage leukemia (trithorax homolog, Drosophila); translocated to, 2.60 4 ABCA7 ATP-binding cassette, sub-family A (ABC1), member 7 2.60 USP40 ubiquitin specific peptidase 40 2.60 FOXI2 forkhead box I2 2.60 CEP170B centrosomal protein 170B 2.60 CCNDBP1 cyclin D-type binding-protein 1 2.60 LOC101927 uncharacterized LOC101927285 2.60 285 CCDC103 coiled-coil domain containing 103 2.59 ADAMTS1 ADAM metallopeptidase with thrombospondin type 1 motif, 1 2.59 CD164L2 CD164 sialomucin-like 2 2.58 ABHD14B abhydrolase domain containing 14B 2.58 RBKS ribokinase 2.58 PDK3 pyruvate dehydrogenase kinase, isozyme 3 2.57 DSP desmoplakin 2.57 C17orf58 chromosome 17 open reading frame 58 2.57 ELOVL1 ELOVL fatty acid elongase 1 2.57 KRT8P12 keratin 8 pseudogene 12 2.56 BOK BCL2-related ovarian killer 2.56 TOM1L1 target of myb1 (chicken)-like 1 2.56 PHLDB3 pleckstrin homology-like domain, family B, member 3 2.56 SDSL serine dehydratase-like 2.56 TMEM191B transmembrane protein 191B 2.55 DMXL1 Dmx-like 1 2.55 ITCH itchy E3 ubiquitin protein ligase 2.55 ZDHHC24 zinc finger, DHHC-type containing 24 2.55 SH3KBP1 SH3-domain kinase binding protein 1 2.55 SH3D19 SH3 domain containing 19 2.55 MROH8 maestro heat-like repeat family member 8 2.55 RBM47 RNA binding motif protein 47 2.54 KRT18 keratin 18 2.54 MT1X metallothionein 1X 2.54 MLLT4 myeloid/lymphoid or mixed-lineage leukemia (trithorax homolog, Drosophila); translocated to, 2.53 4 PIGV phosphatidylinositol glycan anchor biosynthesis, class V 2.53 SNTG2 syntrophin, gamma 2 2.53 ITPR1 inositol 1,4,5-trisphosphate receptor, type 1 2.53 GLCE glucuronic acid epimerase 2.52 SH3TC1 SH3 domain and tetratricopeptide repeats 1 2.52 PTH1R parathyroid hormone 1 receptor 2.52 RNF208 ring finger protein 208 2.52 C1GALT1 core 1 synthase, glycoprotein-N-acetylgalactosamine 3-beta-galactosyltransferase, 1 2.52 ARHGEF5 Rho guanine nucleotide exchange factor (GEF) 5 2.52 STT3B STT3B, subunit of the oligosaccharyltransferase complex (catalytic) 2.52 CTBS chitobiase, di-N-acetyl- 2.51 TPCN2 two pore segment channel 2 2.51 IFNAR2 interferon (alpha, beta and omega) receptor 2 2.51 TANGO2 transport and golgi organization 2 homolog (Drosophila) 2.51 FUCA2 fucosidase, alpha-L- 2, plasma 2.51 CDK18 cyclin-dependent kinase 18 2.51 MBP myelin basic protein 2.51 TRMT2B tRNA methyltransferase 2 homolog B (S. cerevisiae) 2.50 PLAU plasminogen activator, urokinase 2.50 CAPN5 calpain 5 2.50 ATP6V0B ATPase, H+ transporting, lysosomal 21kDa, V0 subunit b 2.50 POLM polymerase (DNA directed), mu 2.50 PRMT6 protein arginine methyltransferase 6 2.50 CCDC85C coiled-coil domain containing 85C 2.50 PIGS phosphatidylinositol glycan anchor biosynthesis, class S 2.50 STK36 serine/threonine kinase 36 2.49 ESAM endothelial cell adhesion molecule 2.49 C9orf171 chromosome 9 open reading frame 171 2.49 TAX1BP1 Tax1 (human T-cell leukemia virus type I) binding protein 1 2.49 PIGO phosphatidylinositol glycan anchor biosynthesis, class O 2.48 POPDC2 popeye domain containing 2 2.48 KLHL14 kelch-like family member 14 2.48 PAG1 phosphoprotein membrane anchor with glycosphingolipid microdomains 1 2.48 MT1E metallothionein 1E 2.47 VPS53 vacuolar protein sorting 53 homolog (S. cerevisiae) 2.47 PPP1R26 protein phosphatase 1, regulatory subunit 26 2.47 MTHFS 5,10-methenyltetrahydrofolate synthetase (5-formyltetrahydrofolate cyclo-ligase) 2.47 LOC100270 uncharacterized LOC100270746 2.47 746 PIGS phosphatidylinositol glycan anchor biosynthesis, class S 2.47 NT5DC1 5'-nucleotidase domain containing 1 2.47 KAZN kazrin, periplakin interacting protein 2.47 TNNI3 troponin I type 3 (cardiac) 2.46 CCZ1 CCZ1 vacuolar protein trafficking and biogenesis associated homolog (S. cerevisiae) 2.46 SQSTM1 sequestosome 1 2.46 MARCH2 membrane-associated ring finger (C3HC4) 2, E3 ubiquitin protein ligase 2.46 PPP1R21 protein phosphatase 1, regulatory subunit 21 2.46 KIAA1804 mixed lineage kinase 4 2.45 NLRX1 NLR family member X1 2.45 ATG16L2 autophagy related 16-like 2 (S. cerevisiae) 2.45 CYB561D2 cytochrome b561 family, member D2 2.44 KIAA0319L KIAA0319-like 2.44 CCZ1 CCZ1 vacuolar protein trafficking and biogenesis associated homolog (S. cerevisiae) 2.44 GALNS galactosamine (N-acetyl)-6-sulfate sulfatase 2.44 TBC1D17 TBC1 domain family, member 17 2.43 SLC20A2 solute carrier family 20 (phosphate transporter), member 2 2.43 TMEM106B transmembrane protein 106B 2.43 DPPA4 developmental pluripotency associated 4 2.43 CELF2 CUGBP, Elav-like family member 2 2.43 SERTAD1 SERTA domain containing 1 2.43 KCTD11 potassium channel tetramerization domain containing 11 2.43 TMEM42 transmembrane protein 42 2.42 AIFM2 apoptosis-inducing factor, mitochondrion-associated, 2 2.42 SIRT5 sirtuin 5 2.42 ARHGAP5- ARHGAP5 antisense RNA 1 (head to head) 2.42 AS1 RELB v-rel avian reticuloendotheliosis viral oncogene homolog B 2.42 FAH fumarylacetoacetate hydrolase (fumarylacetoacetase) 2.42 ZBTB10 zinc finger and BTB domain containing 10 2.42 ORAI3 ORAI calcium release-activated calcium modulator 3 2.41 TRPC6 transient receptor potential cation channel, subfamily C, member 6 2.41 CTSLP2 cathepsin L pseudogene 2 2.41 TMEM17 transmembrane protein 17 2.41 WASH1 WAS protein family homolog 1 2.41 GTF3C1 general transcription factor IIIC, polypeptide 1, alpha 220kDa 2.41 TTC30B tetratricopeptide repeat domain 30B 2.40 AGTRAP angiotensin II receptor-associated protein 2.40 TMEM63C transmembrane protein 63C 2.40 JAG1 jagged 1 2.40 SDCCAG8 serologically defined colon cancer antigen 8 2.40 SLC40A1 solute carrier family 40 (iron-regulated transporter), member 1 2.40 EPHX3 epoxide hydrolase 3 2.39 FAM214A family with sequence similarity 214, member A 2.39 CREBRF CREB3 regulatory factor 2.39 PHLPP2 PH domain and leucine rich repeat protein phosphatase 2 2.39 FAM212B family with sequence similarity 212, member B 2.38 DENND1B DENN/MADD domain containing 1B 2.38 FUCA2 fucosidase, alpha-L- 2, plasma 2.38 AMFR autocrine motility factor receptor, E3 ubiquitin protein ligase 2.38 CCDC51 coiled-coil domain containing 51 2.38 RNF13 ring finger protein 13 2.38 TRIOBP TRIO and F-actin binding protein 2.38 CYBA cytochrome b-245, alpha polypeptide 2.38 POLD4 polymerase (DNA-directed), delta 4, accessory subunit 2.38 PFKFB4 6-phosphofructo-2-kinase/fructose-2,6-biphosphatase 4 2.37 ANGPT1 angiopoietin 1 2.37 CACNA1H calcium channel, voltage-dependent, T type, alpha 1H subunit 2.37 CYHR1 cysteine/histidine-rich 1 2.37 SEC14L1P1 SEC14-like 1 pseudogene 1 2.37 CYP1A2 cytochrome P450, family 1, subfamily A, polypeptide 2 2.37 ATP6AP1L ATPase, H+ transporting, lysosomal accessory protein 1-like 2.37 LOC283270 uncharacterized LOC283270 2.37 TRMT10A tRNA methyltransferase 10 homolog A (S. cerevisiae) 2.37 ACBD5 acyl-CoA binding domain containing 5 2.36 HAGH hydroxyacylglutathione hydrolase 2.36 CYB5D2 cytochrome b5 domain containing 2 2.36 CYSRT1 cysteine-rich tail protein 1 2.36 C1GALT1 core 1 synthase, glycoprotein-N-acetylgalactosamine 3-beta-galactosyltransferase, 1 2.36 KIRREL3 kin of IRRE like 3 (Drosophila) 2.36 PPP1R9A protein phosphatase 1, regulatory subunit 9A 2.36 DENND1A DENN/MADD domain containing 1A 2.35 LOC729887 uncharacterized LOC729887 2.35 ENDOG endonuclease G 2.35 ACAD10 acyl-CoA dehydrogenase family, member 10 2.35 MMP14 matrix metallopeptidase 14 (membrane-inserted) 2.35 TOR1AIP2 torsin A interacting protein 2 2.35 TMEM143 transmembrane protein 143 2.35 CAMKK1 calcium/calmodulin-dependent protein kinase kinase 1, alpha 2.35 SLC35F5 solute carrier family 35, member F5 2.34 OR7E37P olfactory receptor, family 7, subfamily E, member 37 pseudogene 2.34 SNX8 sorting nexin 8 2.34 PCK2 phosphoenolpyruvate carboxykinase 2 (mitochondrial) 2.34 PPCS phosphopantothenoylcysteine synthetase 2.34 RABEPK Rab9 effector protein with kelch motifs 2.34 GPR137 G protein-coupled receptor 137 2.34 MYOZ3 myozenin 3 2.33 ZNF385A zinc finger protein 385A 2.33 CCZ1 CCZ1 vacuolar protein trafficking and biogenesis associated homolog (S. cerevisiae) 2.33 PRKAA2 protein kinase, AMP-activated, alpha 2 catalytic subunit 2.33 OR7E47P olfactory receptor, family 7, subfamily E, member 47 pseudogene 2.33 SERF2 small EDRK-rich factor 2 2.33 MGC4859 uncharacterized LOC79150 2.33 VSX1 visual system homeobox 1 2.32 EXOC8 exocyst complex component 8 2.32 OPLAH 5-oxoprolinase (ATP-hydrolysing) 2.32 MZF1 myeloid zinc finger 1 2.32 SIL1 SIL1 nucleotide exchange factor 2.32 SPPL2A signal peptide peptidase like 2A 2.32 VCPIP1 valosin containing protein (p97)/p47 complex interacting protein 1 2.32 ESCO1 establishment of sister chromatid cohesion N-acetyltransferase 1 2.32 MRC2 mannose receptor, C type 2 2.32 LRFN4 leucine rich repeat and fibronectin type III domain containing 4 2.32 MIF4GD MIF4G domain containing 2.32 RPTOR regulatory associated protein of MTOR, complex 1 2.31 DBP D site of albumin promoter (albumin D-box) binding protein 2.31 NSUN7 NOP2/Sun domain family, member 7 2.31 TRAPPC6A trafficking protein particle complex 6A 2.31 GPRC5C G protein-coupled receptor, class C, group 5, member C 2.31 CPVL carboxypeptidase, vitellogenic-like 2.30 FGF11 fibroblast growth factor 11 2.30 ARHGAP5- ARHGAP5 antisense RNA 1 (head to head) 2.30 AS1 CMAS cytidine monophosphate N-acetylneuraminic acid synthetase 2.30 EPS8L2 EPS8-like 2 2.30 ERAP2 endoplasmic reticulum aminopeptidase 2 2.30 NME3 NME/NM23 nucleoside diphosphate kinase 3 2.29 PPP1R21 protein phosphatase 1, regulatory subunit 21 2.29 LRRC10B leucine rich repeat containing 10B 2.29 FLVCR1 feline leukemia virus subgroup C cellular receptor 1 2.29 RNF14 ring finger protein 14 2.29 PTPRS protein tyrosine phosphatase, receptor type, S 2.29 ATP6V1C1 ATPase, H+ transporting, lysosomal 42kDa, V1 subunit C1 2.28 PRICKLE2 prickle homolog 2 (Drosophila) 2.28 MGST2 microsomal glutathione S-transferase 2 2.28 ACAA1 acetyl-CoA acyltransferase 1 2.28 TRAM2-AS1 TRAM2 antisense RNA 1 (head to head) 2.28 LOC100130 uncharacterized LOC100130093 2.28 093 ORC4 origin recognition complex, subunit 4 2.28 KAL1 Kallmann syndrome 1 sequence 2.28 LOC389765 kinesin family member 27 pseudogene 2.28 TOMM34 translocase of outer mitochondrial membrane 34 2.28 NDFIP2 Nedd4 family interacting protein 2 2.27 ORC4 origin recognition complex, subunit 4 2.27 MYOM1 myomesin 1 2.27 CYSRT1 cysteine-rich tail protein 1 2.27 RAB26 RAB26, member RAS oncogene family 2.27 SELT selenoprotein T 2.27 FBXO9 F-box protein 9 2.27 CCDC30 coiled-coil domain containing 30 2.27 CLIP4 CAP-GLY domain containing linker protein family, member 4 2.27 UQCRB ubiquinol-cytochrome c reductase binding protein 2.26 LINC00087 long intergenic non-protein coding RNA 87 2.26 RHBDD2 rhomboid domain containing 2 2.26 PAG1 phosphoprotein membrane anchor with glycosphingolipid microdomains 1 2.26 TMED4 transmembrane emp24 protein transport domain containing 4 2.25 LAD1 ladinin 1 2.25 SELT selenoprotein T 2.25 CLPTM1 cleft lip and palate associated transmembrane protein 1 2.25 TTC12 tetratricopeptide repeat domain 12 2.25 FHIT fragile histidine triad 2.25 LANCL3 LanC lantibiotic synthetase component C-like 3 (bacterial) 2.25 SLC30A3 solute carrier family 30 (zinc transporter), member 3 2.25 LOC100131 uncharacterized LOC100131564 2.24 564 HCST hematopoietic cell signal transducer 2.24 LINC00847 long intergenic non-protein coding RNA 847 2.24 FBXL17 F-box and leucine-rich repeat protein 17 2.24 CCNL2 cyclin L2 2.24 LOC100130 uncharacterized LOC100130027 2.24 027 POMT2 protein-O-mannosyltransferase 2 2.24 MLLT4 myeloid/lymphoid or mixed-lineage leukemia (trithorax homolog, Drosophila); translocated to, 2.24 4 HEXA hexosaminidase A (alpha polypeptide) 2.24 LRFN4 leucine rich repeat and fibronectin type III domain containing 4 2.24 DSP desmoplakin 2.24 HPS1 Hermansky-Pudlak syndrome 1 2.24 PAOX polyamine oxidase (exo-N4-amino) 2.24 UBL3 ubiquitin-like 3 2.24 GAA glucosidase, alpha; acid 2.23 ERBB2IP erbb2 interacting protein 2.23 YAF2 YY1 associated factor 2 2.22 SLC25A4 solute carrier family 25 (mitochondrial carrier; adenine nucleotide translocator), member 4 2.22 RREB1 ras responsive element binding protein 1 2.22 JKAMP JNK1/MAPK8-associated membrane protein 2.22 FIGN fidgetin 2.22 PRR5- PRR5-ARHGAP8 readthrough 2.22 ARHGAP8 RRM2B ribonucleotide reductase M2 B (TP53 inducible) 2.22 TMEM201 transmembrane protein 201 2.22 LINC01125 long intergenic non-protein coding RNA 1125 2.21 DIRC2 disrupted in renal carcinoma 2 2.21 SRCRB4D scavenger receptor cysteine rich domain containing, group B (4 domains) 2.21 LRIG2 leucine-rich repeats and immunoglobulin-like domains 2 2.21 SCUBE1 signal peptide, CUB domain, EGF-like 1 2.21 EMC10 ER membrane protein complex subunit 10 2.21 CBLN3 cerebellin 3 precursor 2.21 ERC2 ELKS/RAB6-interacting/CAST family member 2 2.21 ATP6V1C1 ATPase, H+ transporting, lysosomal 42kDa, V1 subunit C1 2.21 LITAF lipopolysaccharide-induced TNF factor 2.21 MLLT4 myeloid/lymphoid or mixed-lineage leukemia (trithorax homolog, Drosophila); translocated to, 2.21 4 LIFR leukemia inhibitory factor receptor alpha 2.21 PPT2 palmitoyl-protein thioesterase 2 2.21 PYROXD2 pyridine nucleotide-disulphide oxidoreductase domain 2 2.21 CCHCR1 coiled-coil alpha-helical rod protein 1 2.20 HOXA7 homeobox A7 2.20 LCLAT1 lysocardiolipin acyltransferase 1 2.20 LOC642852 uncharacterized LOC642852 2.20 KLF16 Kruppel-like factor 16 2.20 SH3D21 SH3 domain containing 21 2.20 LIPT2 lipoyl(octanoyl) transferase 2 (putative) 2.20 KLK1 kallikrein 1 2.20 NDRG1 N-myc downstream regulated 1 2.19 TMEM107 transmembrane protein 107 2.19 NAV2 neuron navigator 2 2.19 CTBS chitobiase, di-N-acetyl- 2.19 GAREM GRB2 associated, regulator of MAPK1 2.19 LINC01144 long intergenic non-protein coding RNA 1144 2.19 CERS2 ceramide synthase 2 2.19 NIPSNAP3A nipsnap homolog 3A (C. elegans) 2.19 SIRT5 sirtuin 5 2.19 LOC441204 uncharacterized LOC441204 2.19 DGKQ diacylglycerol kinase, theta 110kDa 2.19 ALDH5A1 aldehyde dehydrogenase 5 family, member A1 2.18 PPP1R13B protein phosphatase 1, regulatory subunit 13B 2.18 TMEM169 transmembrane protein 169 2.18 ECHDC3 enoyl CoA hydratase domain containing 3 2.18 NEU1 sialidase 1 (lysosomal sialidase) 2.18 GINM1 glycoprotein integral membrane 1 2.18 MED14OS MED14 opposite strand 2.18 TMEM38A transmembrane protein 38A 2.18 RNPEPL1 arginyl aminopeptidase (aminopeptidase B)-like 1 2.17 KIAA1161 KIAA1161 2.17 PPP3CC protein phosphatase 3, catalytic subunit, gamma isozyme 2.17 ARID3A AT rich interactive domain 3A (BRIGHT-like) 2.17 EFCAB2 EF-hand calcium binding domain 2 2.17 ACOT8 acyl-CoA thioesterase 8 2.17 HEYL hes-related family bHLH transcription factor with YRPW motif-like 2.16 FUOM fucose mutarotase 2.16 LCOR ligand dependent nuclear receptor corepressor 2.16 CYB561D1 cytochrome b561 family, member D1 2.16 C21orf2 chromosome 21 open reading frame 2 2.16 KLHL21 kelch-like family member 21 2.16 WDR78 WD repeat domain 78 2.16 VANGL1 VANGL planar cell polarity protein 1 2.16 ANXA2P1 annexin A2 pseudogene 1 2.15 ZBED6CL ZBED6 C-terminal like 2.15 SCRN2 secernin 2 2.15 PRKAA1 protein kinase, AMP-activated, alpha 1 catalytic subunit 2.15 SMIM11 small integral membrane protein 11 2.15 GPR108 G protein-coupled receptor 108 2.15 COMMD8 COMM domain containing 8 2.14 SGTB small glutamine-rich tetratricopeptide repeat (TPR)-containing, beta 2.14 KIRREL2 kin of IRRE like 2 (Drosophila) 2.14 ZNF148 zinc finger protein 148 2.14 HOOK1 hook microtubule-tethering protein 1 2.13 IQCA1 IQ motif containing with AAA domain 1 2.13 PTCD3 pentatricopeptide repeat domain 3 2.13 EBAG9 estrogen receptor binding site associated, antigen, 9 2.13 DCUN1D2 DCN1, defective in cullin neddylation 1, domain containing 2 2.13 UNC119 unc-119 homolog (C. elegans) 2.12 PSMD3 proteasome (prosome, macropain) 26S subunit, non-ATPase, 3 2.12 TRAK1 trafficking protein, kinesin binding 1 2.12 EVA1A eva-1 homolog A (C. elegans) 2.12 PITRM1 pitrilysin metallopeptidase 1 2.12 MSMO1 methylsterol monooxygenase 1 2.12 DNAJC11 DnaJ (Hsp40) homolog, subfamily C, member 11 2.12 PCNX pecanex homolog (Drosophila) 2.12 IMPAD1 inositol monophosphatase domain containing 1 2.12 ENDOV endonuclease V 2.12 IQCA1 IQ motif containing with AAA domain 1 2.12 GGA2 golgi-associated, gamma adaptin ear containing, ARF binding protein 2 2.11 FBXL17 F-box and leucine-rich repeat protein 17 2.11 CXXC4 CXXC finger protein 4 2.11 SLC39A11 solute carrier family 39, member 11 2.11 UFSP1 UFM1-specific peptidase 1 (non-functional) 2.11 LOC100133 uncharacterized LOC100133985 2.11 985 LRRC73 leucine rich repeat containing 73 2.10 AUH AU RNA binding protein/enoyl-CoA hydratase 2.10 PDE9A phosphodiesterase 9A 2.10 GIN1 gypsy retrotransposon integrase 1 2.10 ZNF14 zinc finger protein 14 2.10 LRPAP1 low density lipoprotein receptor-related protein associated protein 1 2.10 TMEM129 transmembrane protein 129 2.10 IARS2 isoleucyl-tRNA synthetase 2, mitochondrial 2.10 DNAJC3 DnaJ (Hsp40) homolog, subfamily C, member 3 2.10 UBXN6 UBX domain protein 6 2.09 ZNF577 zinc finger protein 577 2.09 LRRTM1 leucine rich repeat transmembrane neuronal 1 2.09 HPS1 Hermansky-Pudlak syndrome 1 2.09 LOC101927 uncharacterized LOC101927974 2.09 974 COMMD8 COMM domain containing 8 2.09 CELF2 CUGBP, Elav-like family member 2 2.09 KLK8 kallikrein-related peptidase 8 2.09 MAFG v-maf avian musculoaponeurotic fibrosarcoma oncogene homolog G 2.09 TMEM125 transmembrane protein 125 2.09 GSTM4 glutathione S-transferase mu 4 2.08 WIPI1 WD repeat domain, phosphoinositide interacting 1 2.08 GLB1L galactosidase, beta 1-like 2.08 TEX264 testis expressed 264 2.08 MYL5 myosin, light chain 5, regulatory 2.08 SMUG1 single-strand-selective monofunctional uracil-DNA glycosylase 1 2.08 TTC12 tetratricopeptide repeat domain 12 2.08 PTDSS2 phosphatidylserine synthase 2 2.08 C1orf56 chromosome 1 open reading frame 56 2.08 COX14 cytochrome c oxidase assembly homolog 14 (S. cerevisiae) 2.08 TMED7- TMED7-TICAM2 readthrough 2.08 TICAM2 BBX bobby sox homolog (Drosophila) 2.08 LRRC41 leucine rich repeat containing 41 2.07 APOPT1 apoptogenic 1, mitochondrial 2.07 PLA2G16 phospholipase A2, group XVI 2.07 C7orf55 chromosome 7 open reading frame 55 2.07 CITED4 Cbp/p300-interacting transactivator, with Glu/Asp-rich carboxy-terminal domain, 4 2.07 SCNN1A sodium channel, non-voltage-gated 1 alpha subunit 2.07 PDDC1 Parkinson disease 7 domain containing 1 2.07 BSG basigin (Ok blood group) 2.06 TTC32 tetratricopeptide repeat domain 32 2.06 FBXL3 F-box and leucine-rich repeat protein 3 2.06 LOC441081 POM121 membrane glycoprotein (rat) pseudogene 2.06 OR7E156P olfactory receptor, family 7, subfamily E, member 156 pseudogene 2.06 LOC100128 uncharacterized LOC100128398 2.06 398 LOC730101 uncharacterized LOC730101 2.06 HS6ST2 heparan sulfate 6-O-sulfotransferase 2 2.06 BBX bobby sox homolog (Drosophila) 2.06 SNX4 sorting nexin 4 2.06 KLHL25 kelch-like family member 25 2.06 HS6ST1 heparan sulfate 6-O-sulfotransferase 1 2.06 KRT8 keratin 8 2.06 MAP3K8 mitogen-activated protein kinase kinase kinase 8 2.05 RNF170 ring finger protein 170 2.05 SAT1 spermidine/spermine N1-acetyltransferase 1 2.05 LOC101928 uncharacterized LOC101928837 2.05 837 MB21D2 Mab-21 domain containing 2 2.05 FUK fucokinase 2.05 DAZAP2 DAZ associated protein 2 2.05 CORO2A coronin, actin binding protein, 2A 2.04 ANKRD13C ankyrin repeat domain 13C 2.04 ATOX1 antioxidant 1 copper chaperone 2.04 KBTBD3 kelch repeat and BTB (POZ) domain containing 3 2.04 TRMT2B tRNA methyltransferase 2 homolog B (S. cerevisiae) 2.04 IGSF1 immunoglobulin superfamily, member 1 2.04 LURAP1L leucine rich adaptor protein 1-like 2.04 SLC17A5 solute carrier family 17 (acidic sugar transporter), member 5 2.04 ANKRD13A ankyrin repeat domain 13A 2.04 LINC00467 long intergenic non-protein coding RNA 467 2.03 PRKCZ protein kinase C, zeta 2.03 RAB14 RAB14, member RAS oncogene family 2.03 FAM174A family with sequence similarity 174, member A 2.03 MCFD2 multiple coagulation factor deficiency 2 2.03 EMC10 ER membrane protein complex subunit 10 2.03 NFKBIE nuclear factor of kappa light polypeptide gene enhancer in B-cells inhibitor, epsilon 2.03 LMBRD1 LMBR1 domain containing 1 2.03 TXNDC11 thioredoxin domain containing 11 2.02 HSD17B14 hydroxysteroid (17-beta) dehydrogenase 14 2.02 UBR3 ubiquitin protein ligase E3 component n-recognin 3 (putative) 2.02 MYC v-myc avian myelocytomatosis viral oncogene homolog 2.02 LCA5 Leber congenital amaurosis 5 2.02 CST3 cystatin C 2.02 TMEM205 transmembrane protein 205 2.02 SLC9A8 solute carrier family 9, subfamily A (NHE8, cation proton antiporter 8), member 8 2.02 DNPH1 2'-deoxynucleoside 5'-phosphate N-hydrolase 1 2.02 FBLN2 fibulin 2 2.02 UFSP1 UFM1-specific peptidase 1 (non-functional) 2.02 GBA glucosidase, beta, acid 2.01 IDH3A isocitrate dehydrogenase 3 (NAD+) alpha 2.01 TMBIM4 transmembrane BAX inhibitor motif containing 4 2.01 WDFY3- WDFY3 antisense RNA 2 2.01 AS2 MAP3K2 mitogen-activated protein kinase kinase kinase 2 2.01 ERLEC1 endoplasmic reticulum lectin 1 2.01 UBXN4 UBX domain protein 4 2.01 TRAK1 trafficking protein, kinesin binding 1 2.01 C9orf9 chromosome 9 open reading frame 9 2.00 VTI1A vesicle transport through interaction with t-SNAREs 1A 2.00 BNIP3 BCL2/adenovirus E1B 19kDa interacting protein 3 2.00 C5orf28 chromosome 5 open reading frame 28 2.00 TMEM259 transmembrane protein 259 2.00 PRELID2 PRELI domain containing 2 2.00 FBXO16 F-box protein 16 2.00 HMGCS1 3-hydroxy-3-methylglutaryl-CoA synthase 1 (soluble) 2.00 NAPEPLD N-acyl phosphatidylethanolamine phospholipase D 2.00 GOLGA4 golgin A4 2.00 C18orf8 chromosome 18 open reading frame 8 2.00 SLC25A23 solute carrier family 25 (mitochondrial carrier; phosphate carrier), member 23 2.00 C1orf186 chromosome 1 open reading frame 186 1.99 SLC39A13 solute carrier family 39 (zinc transporter), member 13 1.99 SOWAHC sosondowah ankyrin repeat domain family member C 1.99 FAM134A family with sequence similarity 134, member A 1.99 GINM1 glycoprotein integral membrane 1 1.99 EFNA1 ephrin-A1 1.99 PIGQ phosphatidylinositol glycan anchor biosynthesis, class Q 1.99 TP53I11 tumor protein p53 inducible protein 11 1.99 SAT1 spermidine/spermine N1-acetyltransferase 1 1.99 EXOC6 exocyst complex component 6 1.98 MBTPS2 membrane-bound transcription factor peptidase, site 2 1.98 WASH1 WAS protein family homolog 1 1.98 ARSA arylsulfatase A 1.98 TRNT1 tRNA nucleotidyl transferase, CCA-adding, 1 1.98 NIPAL3 NIPA-like domain containing 3 1.98 RALGPS1 Ral GEF with PH domain and SH3 binding motif 1 1.98 NICN1 nicolin 1 1.98 ANKRD10 ankyrin repeat domain 10 1.97 THAP6 THAP domain containing 6 1.97 PCYOX1 prenylcysteine oxidase 1 1.97 LOC728705 uncharacterized LOC728705 1.97 DNAJC16 DnaJ (Hsp40) homolog, subfamily C, member 16 1.97 DIRC2 disrupted in renal carcinoma 2 1.97 TMEM150A transmembrane protein 150A 1.96 ZFYVE21 zinc finger, FYVE domain containing 21 1.96 PDK4 pyruvate dehydrogenase kinase, isozyme 4 1.96 PLA2G12A phospholipase A2, group XIIA 1.96 ESRP2 epithelial splicing regulatory protein 2 1.95 PCED1A PC-esterase domain containing 1A 1.95 ZBED9 zinc finger, BED-type containing 9 1.95 MICU2 mitochondrial calcium uptake 2 1.95 TSPAN9 tetraspanin 9 1.95 NAIF1 nuclear apoptosis inducing factor 1 1.95 MYCN v-myc avian myelocytomatosis viral oncogene neuroblastoma derived homolog 1.95 FBXO17 F-box protein 17 1.95 ZFHX2 zinc finger homeobox 2 1.95 KIAA1468 KIAA1468 1.95 ATP6V1H ATPase, H+ transporting, lysosomal 50/57kDa, V1 subunit H 1.95 C18orf25 chromosome 18 open reading frame 25 1.95 SLC3A2 solute carrier family 3 (amino acid transporter heavy chain), member 2 1.95 ZNF226 zinc finger protein 226 1.95 FBXL17 F-box and leucine-rich repeat protein 17 1.95 ATP6V0D1 ATPase, H+ transporting, lysosomal 38kDa, V0 subunit d1 1.94 ANKRA2 ankyrin repeat, family A (RFXANK-like), 2 1.94 FLJ37453 uncharacterized LOC729614 1.94 TMEM18 transmembrane protein 18 1.94 IQCD IQ motif containing D 1.94 PYROXD1 pyridine nucleotide-disulphide oxidoreductase domain 1 1.94 SMOC2 SPARC related modular calcium binding 2 1.94 USP6NL USP6 N-terminal like 1.94 NR1H2 nuclear receptor subfamily 1, group H, member 2 1.94 ASB10 ankyrin repeat and SOCS box containing 10 1.94 NADSYN1 NAD synthetase 1 1.94 SFT2D2 SFT2 domain containing 2 1.94 HECTD2 HECT domain containing E3 ubiquitin protein ligase 2 1.94 STRBP spermatid perinuclear RNA binding protein 1.94 MARVELD2 MARVEL domain containing 2 1.93 ZBTB20 zinc finger and BTB domain containing 20 1.93 TNRC6C trinucleotide repeat containing 6C 1.93 NAPA N-ethylmaleimide-sensitive factor attachment protein, alpha 1.93 NPAS3 neuronal PAS domain protein 3 1.93 ARFGEF2 ADP-ribosylation factor guanine nucleotide-exchange factor 2 (brefeldin A-inhibited) 1.93 STAG3 stromal antigen 3 1.93 OCEL1 occludin/ELL domain containing 1 1.93 CENPBD1 CENPB DNA-binding domains containing 1 1.93 LRPPRC leucine-rich pentatricopeptide repeat containing 1.93 PITRM1 pitrilysin metallopeptidase 1 1.92 RNASEH2C ribonuclease H2, subunit C 1.92 TMX3 thioredoxin-related transmembrane protein 3 1.92 BNIP3L BCL2/adenovirus E1B 19kDa interacting protein 3-like 1.92 SLBP stem-loop binding protein 1.92 TMEM115 transmembrane protein 115 1.92 C14orf37 chromosome 14 open reading frame 37 1.92 ABCD1 ATP-binding cassette, sub-family D (ALD), member 1 1.92 LRRC41 leucine rich repeat containing 41 1.92 PRKCD protein kinase C, delta 1.92 SPTSSA serine palmitoyltransferase, small subunit A 1.92 PCYOX1 prenylcysteine oxidase 1 1.92 HEXDC hexosaminidase (glycosyl hydrolase family 20, catalytic domain) containing 1.92 LOC148413 uncharacterized LOC148413 1.91 SLC39A10 solute carrier family 39 (zinc transporter), member 10 1.91 THBS3 thrombospondin 3 1.91 ECE1 endothelin converting enzyme 1 1.91 PLEKHM1 pleckstrin homology domain containing, family M (with RUN domain) member 1 1.91 CASC4 cancer susceptibility candidate 4 1.91 RTN3 reticulon 3 1.91 TMEM161A transmembrane protein 161A 1.91 BEND3 BEN domain containing 3 1.91 PCED1A PC-esterase domain containing 1A 1.91 LOC100129 uncharacterized LOC100129550 1.91 550 GKAP1 G kinase anchoring protein 1 1.90 SLC35B3 solute carrier family 35 (adenosine 3'-phospho 5'-phosphosulfate transporter), member B3 1.90 C6orf226 chromosome 6 open reading frame 226 1.90 MARCH9 membrane-associated ring finger (C3HC4) 9 1.90 GBAP1 glucosidase, beta, acid pseudogene 1 1.90 EDEM3 ER degradation enhancer, mannosidase alpha-like 3 1.90 TOMM34 translocase of outer mitochondrial membrane 34 1.90 FUCA1 fucosidase, alpha-L- 1, tissue 1.90 ASH1L-AS1 ASH1L antisense RNA 1 1.90 LRRC23 leucine rich repeat containing 23 1.90 OPA3 optic atrophy 3 (autosomal recessive, with chorea and spastic paraplegia) 1.90 PLEKHG4 pleckstrin homology domain containing, family G (with RhoGef domain) member 4 1.90 TTYH2 tweety family member 2 1.90 TEAD3 TEA domain family member 3 1.89 TMEM106B transmembrane protein 106B 1.89 HOXA-AS3 HOXA cluster antisense RNA 3 1.89 THEM4 thioesterase superfamily member 4 1.89 C6orf226 chromosome 6 open reading frame 226 1.89 CRYZ crystallin, zeta (quinone reductase) 1.89 TMF1 TATA element modulatory factor 1 1.89 ABHD12 abhydrolase domain containing 12 1.89 QPRT quinolinate phosphoribosyltransferase 1.89 SLX1A SLX1 structure-specific endonuclease subunit homolog A (S. cerevisiae) 1.89 TMEM175 transmembrane protein 175 1.89 TCAIM T cell activation inhibitor, mitochondrial 1.89 LYPLAL1 lysophospholipase-like 1 1.89 CFL2 cofilin 2 (muscle) 1.89 LOH12CR2 loss of heterozygosity, 12, chromosomal region 2 (non-protein coding) 1.89 TMEM38B transmembrane protein 38B 1.88 SUMF2 sulfatase modifying factor 2 1.88 SCAMP2 secretory carrier membrane protein 2 1.88 CMTM6 CKLF-like MARVEL transmembrane domain containing 6 1.88 MAP3K2 mitogen-activated protein kinase kinase kinase 2 1.88 NPW neuropeptide W 1.88 MRM1 mitochondrial rRNA methyltransferase 1 homolog (S. cerevisiae) 1.88 PPP2CB protein phosphatase 2, catalytic subunit, beta isozyme 1.88 RAB7A RAB7A, member RAS oncogene family 1.88 PURA purine-rich element binding protein A 1.88 C16orf58 chromosome 16 open reading frame 58 1.87 BOLA1 bolA family member 1 1.87 PTK2B protein tyrosine kinase 2 beta 1.87 SLC39A4 solute carrier family 39 (zinc transporter), member 4 1.87 NFATC2IP nuclear factor of activated T-cells, cytoplasmic, calcineurin-dependent 2 interacting protein 1.87 EDEM2 ER degradation enhancer, mannosidase alpha-like 2 1.87 SP6 Sp6 transcription factor 1.87 ATHL1 ATH1, acid trehalase-like 1 (yeast) 1.87 CANX calnexin 1.87 NDOR1 NADPH dependent diflavin oxidoreductase 1 1.87 ENOSF1 enolase superfamily member 1 1.87 ATP6V1D ATPase, H+ transporting, lysosomal 34kDa, V1 subunit D 1.87 HLCS holocarboxylase synthetase (biotin-(proprionyl-CoA-carboxylase (ATP-hydrolysing)) ligase) 1.87 GRINA glutamate receptor, ionotropic, N-methyl D-aspartate-associated protein 1 (glutamate 1.87 binding) MAFG v-maf avian musculoaponeurotic fibrosarcoma oncogene homolog G 1.87 FAM213A family with sequence similarity 213, member A 1.87 C10orf54 chromosome 10 open reading frame 54 1.87 SIRT7 sirtuin 7 1.86 SLC36A1 solute carrier family 36 (proton/amino acid symporter), member 1 1.86 NUPR1L nuclear protein, transcriptional regulator, 1-like 1.86 JMY junction mediating and regulatory protein, p53 cofactor 1.86 CREG1 cellular repressor of E1A-stimulated genes 1 1.86 FAM213A family with sequence similarity 213, member A 1.86 CENPBD1P CENPBD1 pseudogene 1 1.86 1 CNTFR ciliary neurotrophic factor receptor 1.86 MAGIX MAGI family member, X-linked 1.86 ZCCHC2 zinc finger, CCHC domain containing 2 1.86 DVL1 dishevelled segment polarity protein 1 1.86 TMEM19 transmembrane protein 19 1.86 KDSR 3-ketodihydrosphingosine reductase 1.86 RRAGB Ras-related GTP binding B 1.85 SLC25A42 solute carrier family 25, member 42 1.85 ALDH9A1 aldehyde dehydrogenase 9 family, member A1 1.85 CIB1 calcium and integrin binding 1 (calmyrin) 1.85 LOH12CR2 loss of heterozygosity, 12, chromosomal region 2 (non-protein coding) 1.85 GPR143 G protein-coupled receptor 143 1.85 BMP2K BMP2 inducible kinase 1.85 HOOK1 hook microtubule-tethering protein 1 1.85 UGT2B11 UDP glucuronosyltransferase 2 family, polypeptide B11 1.85 ZBTB42 zinc finger and BTB domain containing 42 1.85 B4GALT4 UDP-Gal:betaGlcNAc beta 1,4- galactosyltransferase, polypeptide 4 1.85 RBM7 RNA binding motif protein 7 1.84 C14orf37 chromosome 14 open reading frame 37 1.84 WDR45 WD repeat domain 45 1.84 RNF14 ring finger protein 14 1.84 HSD17B7 hydroxysteroid (17-beta) dehydrogenase 7 1.84 TADA2B transcriptional adaptor 2B 1.84 PHLDB1 pleckstrin homology-like domain, family B, member 1 1.84 GNS glucosamine (N-acetyl)-6-sulfatase 1.84 CMTM3 CKLF-like MARVEL transmembrane domain containing 3 1.84 TMX1 thioredoxin-related transmembrane protein 1 1.84 ATP6AP2 ATPase, H+ transporting, lysosomal accessory protein 2 1.83 ATG10 autophagy related 10 1.83 PSME4 proteasome (prosome, macropain) activator subunit 4 1.83 SLC7A2 solute carrier family 7 (cationic amino acid transporter, y+ system), member 2 1.83 TRAPPC9 trafficking protein particle complex 9 1.83 TCN2 transcobalamin II 1.83 CENPBD1 CENPB DNA-binding domains containing 1 1.83 SUV420H2 suppressor of variegation 4-20 homolog 2 (Drosophila) 1.83 NUDT12 nudix (nucleoside diphosphate linked moiety X)-type motif 12 1.83 PAN3 PAN3 poly(A) specific ribonuclease subunit 1.83 LINC00998 long intergenic non-protein coding RNA 998 1.83 VTI1A vesicle transport through interaction with t-SNAREs 1A 1.83 POMT1 protein-O-mannosyltransferase 1 1.83 PSME4 proteasome (prosome, macropain) activator subunit 4 1.82 BLOC1S3 biogenesis of lysosomal organelles complex-1, subunit 3 1.82 RAB11FIP4 RAB11 family interacting protein 4 (class II) 1.82 CMTM3 CKLF-like MARVEL transmembrane domain containing 3 1.82 LOC100128 uncharacterized LOC100128498 1.82 498 ARSK arylsulfatase family, member K 1.82 RBPMS2 RNA binding protein with multiple splicing 2 1.82 WDR13 WD repeat domain 13 1.82 HIST3H2A histone cluster 3, H2a 1.82 RPL23AP7 ribosomal protein L23a pseudogene 7 1.82 KIF12 kinesin family member 12 1.82 HOXA-AS3 HOXA cluster antisense RNA 3 1.82 SERINC1 serine incorporator 1 1.82 DNPEP aspartyl aminopeptidase 1.82 CXXC5 CXXC finger protein 5 1.81 TTC28-AS1 TTC28 antisense RNA 1 1.81 MEGF8 multiple EGF-like-domains 8 1.81 ASH1L-AS1 ASH1L antisense RNA 1 1.81 CYB5R1 cytochrome b5 reductase 1 1.81 MANEA mannosidase, endo-alpha 1.81 SLMO2 slowmo homolog 2 (Drosophila) 1.81 CCDC103 coiled-coil domain containing 103 1.81 ZNF444 zinc finger protein 444 1.81 HSD17B7 hydroxysteroid (17-beta) dehydrogenase 7 1.81 DIS3L2 DIS3 like 3'-5' exoribonuclease 2 1.81 DSTN destrin (actin depolymerizing factor) 1.81 KLF13 Kruppel-like factor 13 1.81 GMPR2 guanosine monophosphate reductase 2 1.80 KLF10 Kruppel-like factor 10 1.80 HSPA5 heat shock 70kDa protein 5 (glucose-regulated protein, 78kDa) 1.80 SHKBP1 SH3KBP1 binding protein 1 1.80 MAF v-maf avian musculoaponeurotic fibrosarcoma oncogene homolog 1.80 PHLDB1 pleckstrin homology-like domain, family B, member 1 1.80 SPOPL speckle-type POZ protein-like 1.80 MFSD9 major facilitator superfamily domain containing 9 1.80 THEM4 thioesterase superfamily member 4 1.80 FAM3A family with sequence similarity 3, member A 1.80 IDNK idnK, gluconokinase homolog (E. coli) 1.80 SIRT7 sirtuin 7 1.80 HS6ST2 heparan sulfate 6-O-sulfotransferase 2 1.80 CASP10 caspase 10, apoptosis-related cysteine peptidase 1.80 ATG2A autophagy related 2A 1.80 CDH6 cadherin 6, type 2, K-cadherin (fetal kidney) 1.80 LRTOMT leucine rich transmembrane and O-methyltransferase domain containing 1.80 OR7E24 olfactory receptor, family 7, subfamily E, member 24 1.80 ZNF789 zinc finger protein 789 1.79 IGSF1 immunoglobulin superfamily, member 1 1.79 ZBTB46 zinc finger and BTB domain containing 46 1.79 TRIOBP TRIO and F-actin binding protein 1.79 PAFAH2 platelet-activating factor acetylhydrolase 2, 40kDa 1.79 PIGH phosphatidylinositol glycan anchor biosynthesis, class H 1.79 USP37 ubiquitin specific peptidase 37 1.79 MFSD1 major facilitator superfamily domain containing 1 1.79 TMEM141 transmembrane protein 141 1.79 SNX24 sorting nexin 24 1.79 ERLEC1 endoplasmic reticulum lectin 1 1.79 FAM199X family with sequence similarity 199, X-linked 1.79 LYRM2 LYR motif containing 2 1.79 DHRS11 dehydrogenase/reductase (SDR family) member 11 1.79 TSPAN31 tetraspanin 31 1.79 FAM228B family with sequence similarity 228, member B 1.79 UGT2B10 UDP glucuronosyltransferase 2 family, polypeptide B10 1.79 TNFRSF1A tumor necrosis factor receptor superfamily, member 1A 1.79 POLR2E polymerase (RNA) II (DNA directed) polypeptide E, 25kDa 1.79 ENKD1 enkurin domain containing 1 1.79 KCNJ16 potassium inwardly-rectifying channel, subfamily J, member 16 1.79 PICK1 protein interacting with PRKCA 1 1.78 RTN4R reticulon 4 receptor 1.78 DVL1 dishevelled segment polarity protein 1 1.78 SLAIN1 SLAIN motif family, member 1 1.78 CAPNS1 calpain, small subunit 1 1.78 HSD17B12 hydroxysteroid (17-beta) dehydrogenase 12 1.78 SRGAP2B SLIT-ROBO Rho GTPase activating protein 2B 1.78 RNF11 ring finger protein 11 1.78 SENP5 SUMO1/sentrin specific peptidase 5 1.78 FAM71E1 family with sequence similarity 71, member E1 1.78 LMBR1 limb development membrane protein 1 1.78 PRR5 proline rich 5 (renal) 1.77 PDIA3 protein disulfide isomerase family A, member 3 1.77 PIGQ phosphatidylinositol glycan anchor biosynthesis, class Q 1.77 VCPIP1 valosin containing protein (p97)/p47 complex interacting protein 1 1.77 LINC00908 long intergenic non-protein coding RNA 908 1.77 ZSWIM8 zinc finger, SWIM-type containing 8 1.77 PRDX3 peroxiredoxin 3 1.77 FOXP1 forkhead box P1 1.77 LINC00998 long intergenic non-protein coding RNA 998 1.77 MFSD10 major facilitator superfamily domain containing 10 1.77 LINC00467 long intergenic non-protein coding RNA 467 1.77 LAMB2P1 laminin, beta 2 pseudogene 1 1.77 GP1BB glycoprotein Ib (platelet), beta polypeptide 1.77 NHLRC4 NHL repeat containing 4 1.77 PNPO pyridoxamine 5'-phosphate oxidase 1.77 TCP10L t-complex 10-like 1.77 FBXO15 F-box protein 15 1.77 FOXO6 forkhead box O6 1.77 C7orf13 chromosome 7 open reading frame 13 1.77 ZSCAN1 zinc finger and SCAN domain containing 1 1.77 VTI1A vesicle transport through interaction with t-SNAREs 1A 1.77 TTC7A tetratricopeptide repeat domain 7A 1.77 KIAA1919 KIAA1919 1.77 ADD3 adducin 3 (gamma) 1.76 CYB561A3 cytochrome b561 family, member A3 1.76 SC5D sterol-C5-desaturase 1.76 POMGNT1 protein O-linked mannose N-acetylglucosaminyltransferase 1 (beta 1,2-) 1.76 TMED1 transmembrane emp24 protein transport domain containing 1 1.76 ATG2A autophagy related 2A 1.76 PXK PX domain containing serine/threonine kinase 1.76 CTTN cortactin 1.76 SERGEF secretion regulating guanine nucleotide exchange factor 1.76 IPP intracisternal A particle-promoted polypeptide 1.76 GMPR2 guanosine monophosphate reductase 2 1.76 EDEM3 ER degradation enhancer, mannosidase alpha-like 3 1.76 FAM122B family with sequence similarity 122B 1.76 SPSB1 splA/ryanodine receptor domain and SOCS box containing 1 1.76 MAP1LC3B microtubule-associated protein 1 light chain 3 beta pseudogene 1 1.76 P1 WDR34 WD repeat domain 34 1.76 LARP1B La ribonucleoprotein domain family, member 1B 1.76 ALG2 ALG2, alpha-1,3/1,6-mannosyltransferase 1.76 CHMP1B charged multivesicular body protein 1B 1.76 PTS 6-pyruvoyltetrahydropterin synthase 1.75 POMGNT1 protein O-linked mannose N-acetylglucosaminyltransferase 1 (beta 1,2-) 1.75 NUDT14 nudix (nucleoside diphosphate linked moiety X)-type motif 14 1.75 OMA1 OMA1 zinc metallopeptidase 1.75 GSTK1 glutathione S-transferase kappa 1 1.75 LDLR low density lipoprotein receptor 1.75 NT5C 5', 3'-nucleotidase, cytosolic 1.75 DIS3L DIS3 like exosome 3'-5' exoribonuclease 1.75 LOC283861 uncharacterized LOC283861 1.75 VPS36 vacuolar protein sorting 36 homolog (S. cerevisiae) 1.75 TM9SF2 transmembrane 9 superfamily member 2 1.75 NSUN5 NOP2/Sun domain family, member 5 1.75 IER3IP1 immediate early response 3 interacting protein 1 1.75 ARL3 ADP-ribosylation factor-like 3 1.75 TWSG1 twisted gastrulation BMP signaling modulator 1 1.75 SLC25A10 solute carrier family 25 (mitochondrial carrier; dicarboxylate transporter), member 10 1.75 THTPA thiamine triphosphatase 1.75 SHB Src homology 2 domain containing adaptor protein B 1.75 BCAR1 breast cancer anti-estrogen resistance 1 1.75 QSOX2 quiescin Q6 sulfhydryl oxidase 2 1.75 TMBIM4 transmembrane BAX inhibitor motif containing 4 1.75 BNIP3L BCL2/adenovirus E1B 19kDa interacting protein 3-like 1.75 LOC644727 uncharacterized LOC644727 1.74 ATPIF1 ATPase inhibitory factor 1 1.74 MOB3B MOB kinase activator 3B 1.74 B9D2 B9 protein domain 2 1.74 ZNF655 zinc finger protein 655 1.74 GP1BB glycoprotein Ib (platelet), beta polypeptide 1.74 ATAD1 ATPase family, AAA domain containing 1 1.74 ACTR1B ARP1 actin-related protein 1 homolog B, centractin beta (yeast) 1.74 ANXA7 annexin A7 1.74 ZNF469 zinc finger protein 469 1.74 MAP9 microtubule-associated protein 9 1.74 SRRM2- SRRM2 antisense RNA 1 1.74 AS1 SLC22A23 solute carrier family 22, member 23 1.74 PPP2R4 protein phosphatase 2A activator, regulatory subunit 4 1.74 TSTD3 thiosulfate sulfurtransferase (rhodanese)-like domain containing 3 1.74 ZC3H6 zinc finger CCCH-type containing 6 1.74 RINL Ras and Rab interactor-like 1.74 NPHP3 nephronophthisis 3 (adolescent) 1.74 SLC25A20 solute carrier family 25 (carnitine/acylcarnitine translocase), member 20 1.73 TTC8 tetratricopeptide repeat domain 8 1.73 MRS2 MRS2 magnesium transporter 1.73 L3HYPDH L-3-hydroxyproline dehydratase (trans-) 1.73 CCT6B chaperonin containing TCP1, subunit 6B (zeta 2) 1.73 AHCYL1 adenosylhomocysteinase-like 1 1.73 B3GAT3 beta-1,3-glucuronyltransferase 3 (glucuronosyltransferase I) 1.73 PHYHD1 phytanoyl-CoA dioxygenase domain containing 1 1.73 C21orf59 chromosome 21 open reading frame 59 1.73 CYB5B cytochrome b5 type B (outer mitochondrial membrane) 1.73 ATP6V1G1 ATPase, H+ transporting, lysosomal 13kDa, V1 subunit G1 1.73 C21orf2 chromosome 21 open reading frame 2 1.73 ARHGAP5 Rho GTPase activating protein 5 1.73 KLF11 Kruppel-like factor 11 1.73 EMC7 ER membrane protein complex subunit 7 1.73 CRYZ crystallin, zeta (quinone reductase) 1.73 POMP proteasome maturation protein 1.73 STARD3 StAR-related lipid transfer (START) domain containing 3 1.72 RHBDD3 rhomboid domain containing 3 1.72 ASTE1 asteroid homolog 1 (Drosophila) 1.72 AFF4 AF4/FMR2 family, member 4 1.72 ALKBH5 alkB, alkylation repair homolog 5 (E. coli) 1.72 HERPUD1 homocysteine-inducible, endoplasmic reticulum stress-inducible, ubiquitin-like domain 1.72 member 1 COX17 COX17 cytochrome c oxidase copper chaperone 1.72 OMA1 OMA1 zinc metallopeptidase 1.72 VCX2 variable charge, X-linked 2 1.72 IL17RD interleukin 17 receptor D 1.72 LOC339803 uncharacterized LOC339803 1.72 CRADD CASP2 and RIPK1 domain containing adaptor with death domain 1.72 ADAM10 ADAM metallopeptidase domain 10 1.72 BCAS3 breast carcinoma amplified sequence 3 1.72 RASSF8 Ras association (RalGDS/AF-6) domain family (N-terminal) member 8 1.72 SLC12A4 solute carrier family 12 (potassium/chloride transporter), member 4 1.72 GRIN2A glutamate receptor, ionotropic, N-methyl D-aspartate 2A 1.72 YDJC YdjC homolog (bacterial) 1.72 RAB18 RAB18, member RAS oncogene family 1.71 RAB12 RAB12, member RAS oncogene family 1.71 MCEE methylmalonyl CoA epimerase 1.71 NINJ1 ninjurin 1 1.71 AHCYL1 adenosylhomocysteinase-like 1 1.71 RCOR3 REST corepressor 3 1.71 TAOK2 TAO kinase 2 1.71 SPA17 sperm autoantigenic protein 17 1.71 WHAMM WAS protein homolog associated with actin, golgi membranes and microtubules 1.71 SPRTN SprT-like N-terminal domain 1.71 CCDC106 coiled-coil domain containing 106 1.71 CHURC1 churchill domain containing 1 1.71 TMBIM4 transmembrane BAX inhibitor motif containing 4 1.71 NPHP3 nephronophthisis 3 (adolescent) 1.71 ABHD4 abhydrolase domain containing 4 1.71 IFT27 intraflagellar transport 27 homolog (Chlamydomonas) 1.71 MIEN1 migration and invasion enhancer 1 1.71 CPT2 carnitine palmitoyltransferase 2 1.70 CCDC104 coiled-coil domain containing 104 1.70 NIT1 nitrilase 1 1.70 TRAM1 translocation associated membrane protein 1 1.70 BCAP31 B-cell receptor-associated protein 31 1.70 RNF216 ring finger protein 216 1.70 ATP5S ATP synthase, H+ transporting, mitochondrial Fo complex, subunit s (factor B) 1.70 NDUFA7 NADH dehydrogenase (ubiquinone) 1 alpha subcomplex, 7, 14.5kDa 1.70 COQ7 coenzyme Q7 homolog, ubiquinone (yeast) 1.70 WDR35 WD repeat domain 35 1.70 SMPD1 sphingomyelin phosphodiesterase 1, acid lysosomal 1.70 RBL2 retinoblastoma-like 2 1.70 ADIPOR1 adiponectin receptor 1 1.70 SYT7 synaptotagmin VII 1.70 ATP5S ATP synthase, H+ transporting, mitochondrial Fo complex, subunit s (factor B) 1.70 IGSF8 immunoglobulin superfamily, member 8 1.70 ITFG3 integrin alpha FG-GAP repeat containing 3 1.70 TMEM128 transmembrane protein 128 1.70 SEC11C SEC11 homolog C (S. cerevisiae) 1.70 CSRNP1 cysteine-serine-rich nuclear protein 1 1.70 LOC286071 uncharacterized LOC286071 1.70 CTDSP1 CTD (carboxy-terminal domain, RNA polymerase II, polypeptide A) small phosphatase 1 1.69 SMIM19 small integral membrane protein 19 1.69 TRMT1L tRNA methyltransferase 1 homolog (S. cerevisiae)-like 1.69 TRPT1 tRNA phosphotransferase 1 1.69 NAT14 N-acetyltransferase 14 (GCN5-related, putative) 1.69 JMJD8 jumonji domain containing 8 1.69 CDKN2A cyclin-dependent kinase inhibitor 2A 1.69 CLUAP1 clusterin associated protein 1 1.69 KPNA5 karyopherin alpha 5 (importin alpha 6) 1.69 DNAJC30 DnaJ (Hsp40) homolog, subfamily C, member 30 1.69 MFSD1 major facilitator superfamily domain containing 1 1.69 C14orf159 chromosome 14 open reading frame 159 1.69 ATG4D autophagy related 4D, cysteine peptidase 1.69 CTSB cathepsin B 1.69 MAPK3 mitogen-activated protein kinase 3 1.69 PPAPDC2 phosphatidic acid phosphatase type 2 domain containing 2 1.69 TMEM53 transmembrane protein 53 1.69 PTPRU protein tyrosine phosphatase, receptor type, U 1.69 VPS13A vacuolar protein sorting 13 homolog A (S. cerevisiae) 1.69 DAK dihydroxyacetone kinase 2 homolog (S. cerevisiae) 1.69 ALKBH7 alkB, alkylation repair homolog 7 (E. coli) 1.69 CLPX caseinolytic mitochondrial matrix peptidase chaperone subunit 1.69 REEP3 receptor accessory protein 3 1.69 CCDC53 coiled-coil domain containing 53 1.69 ARRDC3 arrestin domain containing 3 1.68 WDR45B WD repeat domain 45B 1.68 HSD17B1 hydroxysteroid (17-beta) dehydrogenase 1 1.68 FBXO33 F-box protein 33 1.68 CCDC6 coiled-coil domain containing 6 1.68 ILF3-AS1 ILF3 antisense RNA 1 (head to head) 1.68 ZDHHC5 zinc finger, DHHC-type containing 5 1.68 PTTG1IP pituitary tumor-transforming 1 interacting protein 1.68 CPNE3 copine III 1.68 MCF2L MCF.2 cell line derived transforming sequence-like 1.68 CDC42EP1 CDC42 effector protein (Rho GTPase binding) 1 1.68 NECAB3 N-terminal EF-hand calcium binding protein 3 1.68 SAMD10 sterile alpha motif domain containing 10 1.68 FUZ fuzzy planar cell polarity protein 1.68 TMEM52 transmembrane protein 52 1.68 PVRL3 poliovirus receptor-related 3 1.68 FBXW7 F-box and WD repeat domain containing 7, E3 ubiquitin protein ligase 1.68 GSTK1 glutathione S-transferase kappa 1 1.67 GLIPR2 GLI pathogenesis-related 2 1.67 PDHB pyruvate dehydrogenase (lipoamide) beta 1.67 CEP19 centrosomal protein 19kDa 1.67 MOSPD1 motile sperm domain containing 1 1.67 NID1 nidogen 1 1.67 COQ4 coenzyme Q4 1.67 LOC286272 uncharacterized LOC286272 1.67 CASC10 cancer susceptibility candidate 10 1.67 C8orf44 chromosome 8 open reading frame 44 1.67 ZNHIT2 zinc finger, HIT-type containing 2 1.67 NSUN5 NOP2/Sun domain family, member 5 1.67 NT5DC1 5'-nucleotidase domain containing 1 1.67 DHRS12 dehydrogenase/reductase (SDR family) member 12 1.67 NHLRC3 NHL repeat containing 3 1.67 ATPIF1 ATPase inhibitory factor 1 1.67 COPS7A COP9 signalosome subunit 7A 1.67 TM2D2 TM2 domain containing 2 1.67 VPS37A vacuolar protein sorting 37 homolog A (S. cerevisiae) 1.67 JTB jumping translocation breakpoint 1.67 THAP7-AS1 THAP7 antisense RNA 1 1.67 SIRT4 sirtuin 4 1.67 ULK1 unc-51 like autophagy activating kinase 1 1.67 PRR5 proline rich 5 (renal) 1.66 ATP6V1A ATPase, H+ transporting, lysosomal 70kDa, V1 subunit A 1.66 AMZ2 archaelysin family metallopeptidase 2 1.66 RRNAD1 ribosomal RNA adenine dimethylase domain containing 1 1.66 PDXK pyridoxal (pyridoxine, vitamin B6) kinase 1.66 RAB12 RAB12, member RAS oncogene family 1.66 LZTS2 leucine zipper, putative tumor suppressor 2 1.66 WDTC1 WD and tetratricopeptide repeats 1 1.66 ZNF277 zinc finger protein 277 1.66 HSD17B7 hydroxysteroid (17-beta) dehydrogenase 7 1.66 ARMC10 armadillo repeat containing 10 1.66 CD164 CD164 molecule, sialomucin 1.66 HDHD2 haloacid dehalogenase-like hydrolase domain containing 2 1.66 ABCA1 ATP-binding cassette, sub-family A (ABC1), member 1 1.66 AIFM1 apoptosis-inducing factor, mitochondrion-associated, 1 1.66 HPS3 Hermansky-Pudlak syndrome 3 1.66 IFT43 intraflagellar transport 43 homolog (Chlamydomonas) 1.66 DCAF12 DDB1 and CUL4 associated factor 12 1.66 ATP13A1 ATPase type 13A1 1.66 PHLPP1 PH domain and leucine rich repeat protein phosphatase 1 1.66 TYW5 tRNA-yW synthesizing protein 5 1.66 CC2D1B coiled-coil and C2 domain containing 1B 1.66 TRUB1 TruB pseudouridine (psi) synthase family member 1 1.65 RFNG RFNG O-fucosylpeptide 3-beta-N-acetylglucosaminyltransferase 1.65 PP7080 uncharacterized LOC25845 1.65 SEC22C SEC22 vesicle trafficking protein homolog C (S. cerevisiae) 1.65 MYL12B myosin, light chain 12B, regulatory 1.65 COG7 component of oligomeric golgi complex 7 1.65 RAB3IP RAB3A interacting protein 1.65 PDXDC1 pyridoxal-dependent decarboxylase domain containing 1 1.65 TMED5 transmembrane emp24 protein transport domain containing 5 1.65 SNX27 sorting nexin family member 27 1.65 GALNT7 polypeptide N-acetylgalactosaminyltransferase 7 1.65 PNPLA8 patatin-like phospholipase domain containing 8 1.65 HPS6 Hermansky-Pudlak syndrome 6 1.65 P4HA2 prolyl 4-hydroxylase, alpha polypeptide II 1.65 PLEKHA3 pleckstrin homology domain containing, family A (phosphoinositide binding specific) member 1.65 3 BIRC6 baculoviral IAP repeat containing 6 1.65 RBBP6 retinoblastoma binding protein 6 1.65 BBS12 Bardet-Biedl syndrome 12 1.65 MAPK6 mitogen-activated protein kinase 6 1.65 ZBTB4 zinc finger and BTB domain containing 4 1.65 HMHA1 histocompatibility (minor) HA-1 1.65 VPS13A vacuolar protein sorting 13 homolog A (S. cerevisiae) 1.65 SIRT3 sirtuin 3 1.65 SLC35C2 solute carrier family 35 (GDP-fucose transporter), member C2 1.65 G6PC3 glucose 6 phosphatase, catalytic, 3 1.65 C12orf5 chromosome 12 open reading frame 5 1.65 VPS37D vacuolar protein sorting 37 homolog D (S. cerevisiae) 1.64 BRMS1L breast cancer metastasis-suppressor 1-like 1.64 RIOK3 RIO kinase 3 1.64 UBXN8 UBX domain protein 8 1.64 USF2 upstream transcription factor 2, c-fos interacting 1.64 ANO2 anoctamin 2, calcium activated chloride channel 1.64 PTK7 protein tyrosine kinase 7 1.64 MFSD3 major facilitator superfamily domain containing 3 1.64 FAM132A family with sequence similarity 132, member A 1.64 ABCD4 ATP-binding cassette, sub-family D (ALD), member 4 1.64 SASH1 SAM and SH3 domain containing 1 1.64 GALNT18 polypeptide N-acetylgalactosaminyltransferase 18 1.64 FAM3A family with sequence similarity 3, member A 1.64 C2orf76 chromosome 2 open reading frame 76 1.64 E2F5 E2F transcription factor 5, p130-binding 1.64 CAMSAP3 calmodulin regulated spectrin-associated protein family, member 3 1.64 SCRN3 secernin 3 1.64 RABGGTA Rab geranylgeranyltransferase, alpha subunit 1.64 WDR81 WD repeat domain 81 1.64 LEPREL2 leprecan-like 2 1.64 DEDD2 death effector domain containing 2 1.64 APRT adenine phosphoribosyltransferase 1.63 PARS2 prolyl-tRNA synthetase 2, mitochondrial (putative) 1.63 CAPZA2 capping protein (actin filament) muscle Z-line, alpha 2 1.63 DENND4C DENN/MADD domain containing 4C 1.63 LOC101929 uncharacterized LOC101929243 1.63 243 KIAA2018 KIAA2018 1.63 DDRGK1 DDRGK domain containing 1 1.63 UHRF1BP1 UHRF1 binding protein 1 1.63 PIGX phosphatidylinositol glycan anchor biosynthesis, class X 1.63 PYGL phosphorylase, glycogen, liver 1.63 RNPC3 RNA-binding region (RNP1, RRM) containing 3 1.63 USP37 ubiquitin specific peptidase 37 1.63 APH1B APH1B gamma secretase subunit 1.63 PLAA phospholipase A2-activating protein 1.63 UGCG UDP-glucose ceramide glucosyltransferase 1.63 SLC50A1 solute carrier family 50 (sugar efflux transporter), member 1 1.63 POC5 POC5 centriolar protein 1.63 SEC22C SEC22 vesicle trafficking protein homolog C (S. cerevisiae) 1.63 C5orf30 chromosome 5 open reading frame 30 1.63 WDR73 WD repeat domain 73 1.63 HM13 histocompatibility (minor) 13 1.63 HOXA-AS2 HOXA cluster antisense RNA 2 1.63 NDUFAF6 NADH dehydrogenase (ubiquinone) complex I, assembly factor 6 1.63 STARD10 StAR-related lipid transfer (START) domain containing 10 1.63 MOSPD2 motile sperm domain containing 2 1.63 PLAA phospholipase A2-activating protein 1.63 UHMK1 U2AF homology motif (UHM) kinase 1 1.63 CBWD5 COBW domain containing 5 1.63 ARMC10 armadillo repeat containing 10 1.63 RNF181 ring finger protein 181 1.63 SLU7 SLU7 splicing factor homolog (S. cerevisiae) 1.63 SLC38A6 solute carrier family 38, member 6 1.62 DUSP28 dual specificity phosphatase 28 1.62 METTL23 methyltransferase like 23 1.62 YAF2 YY1 associated factor 2 1.62 HSP90B3P heat shock protein 90kDa beta (Grp94), member 3, pseudogene 1.62 ARL14EP ADP-ribosylation factor-like 14 effector protein 1.62 SMPD2 sphingomyelin phosphodiesterase 2, neutral membrane (neutral sphingomyelinase) 1.62 ZNF12 zinc finger protein 12 1.62 MRPL57 mitochondrial ribosomal protein L57 1.62 ADD3-AS1 ADD3 antisense RNA 1 1.62 GET4 golgi to ER traffic protein 4 homolog (S. cerevisiae) 1.62 ALKBH5 alkB, alkylation repair homolog 5 (E. coli) 1.62 PYROXD1 pyridine nucleotide-disulphide oxidoreductase domain 1 1.62 RPP25L ribonuclease P/MRP 25kDa subunit-like 1.62 APOBEC3F apolipoprotein B mRNA editing enzyme, catalytic polypeptide-like 3F 1.62 ZBTB44 zinc finger and BTB domain containing 44 1.62 FKRP fukutin related protein 1.62 ADAM10 ADAM metallopeptidase domain 10 1.62 ASXL2 additional sex combs like 2 (Drosophila) 1.62 BCL2L11 BCL2-like 11 (apoptosis facilitator) 1.62 CSRNP1 cysteine-serine-rich nuclear protein 1 1.62 BAG2 BCL2-associated athanogene 2 1.62 HMGCL 3-hydroxymethyl-3-methylglutaryl-CoA lyase 1.61 TP73-AS1 TP73 antisense RNA 1 1.61 WDR78 WD repeat domain 78 1.61 ARMC10 armadillo repeat containing 10 1.61 MAGT1 magnesium transporter 1 1.61 MCF2L MCF.2 cell line derived transforming sequence-like 1.61 SLC26A6 solute carrier family 26 (anion exchanger), member 6 1.61 FBXL15 F-box and leucine-rich repeat protein 15 1.61 NME5 NME/NM23 family member 5 1.61 NDUFC1 NADH dehydrogenase (ubiquinone) 1, subcomplex unknown, 1, 6kDa 1.61 UQCRFS1 ubiquinol-cytochrome c reductase, Rieske iron-sulfur polypeptide 1 1.61 NSMCE1 non-SMC element 1 homolog (S. cerevisiae) 1.61 MARCH6 membrane-associated ring finger (C3HC4) 6, E3 ubiquitin protein ligase 1.61 MLYCD malonyl-CoA decarboxylase 1.61 OXSM 3-oxoacyl-ACP synthase, mitochondrial 1.61 HIBADH 3-hydroxyisobutyrate dehydrogenase 1.61 FOPNL FGFR1OP N-terminal like 1.60 SLC45A4 solute carrier family 45, member 4 1.60 ACADVL acyl-CoA dehydrogenase, very long chain 1.60 SPRTN SprT-like N-terminal domain 1.60 RAB5B RAB5B, member RAS oncogene family 1.60 PCBP2 poly(rC) binding protein 2 1.60 HPCAL1 hippocalcin-like 1 1.60 SCYL1 SCY1-like 1 (S. cerevisiae) 1.60 RUFY2 RUN and FYVE domain containing 2 1.60 NT5C2 5'-nucleotidase, cytosolic II 1.60 EIF2D eukaryotic translation initiation factor 2D 1.60 DNAJA2 DnaJ (Hsp40) homolog, subfamily A, member 2 1.60 HELQ helicase, POLQ-like 1.60 DCXR dicarbonyl/L-xylulose reductase 1.60 PSME1 proteasome (prosome, macropain) activator subunit 1 (PA28 alpha) 1.60 SRGAP3 SLIT-ROBO Rho GTPase activating protein 3 1.60 NUBPL nucleotide binding protein-like 1.60 IFT88 intraflagellar transport 88 homolog (Chlamydomonas) 1.60 ORMDL1 ORMDL sphingolipid biosynthesis regulator 1 1.60 DKFZP564C DKFZP564C152 protein 1.60 152 TEN1 TEN1 CST complex subunit 1.60 TMEM14A transmembrane protein 14A 1.60 UBXN8 UBX domain protein 8 1.60 ESRRA estrogen-related receptor alpha 1.60 IMPA1 inositol(myo)-1(or 4)-monophosphatase 1 1.60 PGK1 phosphoglycerate kinase 1 1.60 PHF10 PHD finger protein 10 1.60 RABGGTB Rab geranylgeranyltransferase, beta subunit 1.60 FBXW4 F-box and WD repeat domain containing 4 1.60 XBP1 X-box binding protein 1 1.60 OSCP1 organic solute carrier partner 1 1.60 SLC9A4 solute carrier family 9, subfamily A (NHE4, cation proton antiporter 4), member 4 1.60 ANKRD39 ankyrin repeat domain 39 1.60 COA5 cytochrome c oxidase assembly factor 5 1.59 RAB18 RAB18, member RAS oncogene family 1.59 A2M-AS1 A2M antisense RNA 1 1.59 SMIM17 small integral membrane protein 17 1.59 DERL1 derlin 1 1.59 FICD FIC domain containing 1.59 LINC00930 long intergenic non-protein coding RNA 930 1.59 NUAK2 NUAK family, SNF1-like kinase, 2 1.59 IGSF8 immunoglobulin superfamily, member 8 1.59 ALG14 ALG14, UDP-N-acetylglucosaminyltransferase subunit 1.59 LOH12CR1 loss of heterozygosity, 12, chromosomal region 1 1.59 MRPL54 mitochondrial ribosomal protein L54 1.59 HM13 histocompatibility (minor) 13 1.59 MTMR10 myotubularin related protein 10 1.59 PSMG3- PSMG3 antisense RNA 1 (head to head) 1.59 AS1 SNHG11 small nucleolar RNA host gene 11 (non-protein coding) 1.59 NPEPPS aminopeptidase puromycin sensitive 1.59 CACFD1 calcium channel flower domain containing 1 1.59 TRIQK triple QxxK/R motif containing 1.59 BEX2 brain expressed X-linked 2 1.59 ALG2 ALG2, alpha-1,3/1,6-mannosyltransferase 1.59 TMEM25 transmembrane protein 25 1.59 ERMP1 endoplasmic reticulum metallopeptidase 1 1.58 FKBP2 FK506 binding protein 2, 13kDa 1.58 CAPZA2 capping protein (actin filament) muscle Z-line, alpha 2 1.58 CUL9 cullin 9 1.58 TFG TRK-fused gene 1.58 RBFA ribosome binding factor A (putative) 1.58 DCUN1D3 DCN1, defective in cullin neddylation 1, domain containing 3 1.58 AIG1 androgen-induced 1 1.58 FAM203A family with sequence similarity 203, member A 1.58 GNPDA2 glucosamine-6-phosphate deaminase 2 1.58 LINC00339 long intergenic non-protein coding RNA 339 1.58 FOXP1 forkhead box P1 1.58 AIMP2 aminoacyl tRNA synthetase complex-interacting multifunctional protein 2 1.58 MAGT1 magnesium transporter 1 1.58 ATP6V1E1 ATPase, H+ transporting, lysosomal 31kDa, V1 subunit E1 1.58 TRMT2A tRNA methyltransferase 2 homolog A (S. cerevisiae) 1.58 FLJ37453 uncharacterized LOC729614 1.58 ABCC10 ATP-binding cassette, sub-family C (CFTR/MRP), member 10 1.58 TCEA3 transcription elongation factor A (SII), 3 1.57 NCKAP1 NCK-associated protein 1 1.57 COASY CoA synthase 1.57 PTTG1IP pituitary tumor-transforming 1 interacting protein 1.57 FAM179B family with sequence similarity 179, member B 1.57 FBXL4 F-box and leucine-rich repeat protein 4 1.57 EFCAB7 EF-hand calcium binding domain 7 1.57 NACC1 nucleus accumbens associated 1, BEN and BTB (POZ) domain containing 1.57 PIGB phosphatidylinositol glycan anchor biosynthesis, class B 1.57 CLPTM1L CLPTM1-like 1.57 RPP38 ribonuclease P/MRP 38kDa subunit 1.57 TMEM260 transmembrane protein 260 1.57 GCLC glutamate-cysteine ligase, catalytic subunit 1.57 OSCP1 organic solute carrier partner 1 1.57 CAAP1 caspase activity and apoptosis inhibitor 1 1.57 SPRYD4 SPRY domain containing 4 1.57 FAM3C family with sequence similarity 3, member C 1.57 XPO5 exportin 5 1.57 ZNF148 zinc finger protein 148 1.57 HSD17B8 hydroxysteroid (17-beta) dehydrogenase 8 1.57 BRE brain and reproductive organ-expressed (TNFRSF1A modulator) 1.56 RBM41 RNA binding motif protein 41 1.56 KLF9 Kruppel-like factor 9 1.56 FDX1 ferredoxin 1 1.56 ESRRA estrogen-related receptor alpha 1.56 PTPRF protein tyrosine phosphatase, receptor type, F 1.56 WDR26 WD repeat domain 26 1.56 WDR73 WD repeat domain 73 1.56 AGA aspartylglucosaminidase 1.56 ZFAND3 zinc finger, AN1-type domain 3 1.56 PLEKHB2 pleckstrin homology domain containing, family B (evectins) member 2 1.56 COMMD7 COMM domain containing 7 1.56 HINT3 histidine triad nucleotide binding protein 3 1.56 MRPS28 mitochondrial ribosomal protein S28 1.56 RSG1 REM2 and RAB-like small GTPase 1 1.56 SCAP SREBF chaperone 1.56 ERMARD ER membrane-associated RNA degradation 1.56 ERC2 ELKS/RAB6-interacting/CAST family member 2 1.56 GPR180 G protein-coupled receptor 180 1.56 ZNF667- ZNF667 antisense RNA 1 (head to head) 1.56 AS1 UFL1 UFM1-specific ligase 1 1.56 FAM160A2 family with sequence similarity 160, member A2 1.56 AMMECR1 Alport syndrome, mental retardation, midface hypoplasia and elliptocytosis chromosomal 1.56 region gene 1 ZNF177 zinc finger protein 177 1.56 ZADH2 zinc binding alcohol dehydrogenase domain containing 2 1.56 ARL2BP ADP-ribosylation factor-like 2 binding protein 1.56 MYEF2 myelin expression factor 2 1.56 ARHGAP5 Rho GTPase activating protein 5 1.56 TMEM8A transmembrane protein 8A 1.56 NUDT22 nudix (nucleoside diphosphate linked moiety X)-type motif 22 1.55 RIPK1 receptor (TNFRSF)-interacting serine-threonine kinase 1 1.55 PDIA3 protein disulfide isomerase family A, member 3 1.55 ABCB10 ATP-binding cassette, sub-family B (MDR/TAP), member 10 1.55 UBE2M ubiquitin-conjugating enzyme E2M 1.55 DEAF1 DEAF1 transcription factor 1.55 RABGGTB Rab geranylgeranyltransferase, beta subunit 1.55 IMPAD1 inositol monophosphatase domain containing 1 1.55 LACTB2 lactamase, beta 2 1.55 FIS1 fission 1 (mitochondrial outer membrane) homolog (S. cerevisiae) 1.55 TRIM41 tripartite motif containing 41 1.55 DTNBP1 dystrobrevin binding protein 1 1.55 USP6 ubiquitin specific peptidase 6 1.55 DEAF1 DEAF1 transcription factor 1.55 ALDH16A1 aldehyde dehydrogenase 16 family, member A1 1.55 TRNT1 tRNA nucleotidyl transferase, CCA-adding, 1 1.55 NPEPPS aminopeptidase puromycin sensitive 1.55 ABHD17C abhydrolase domain containing 17C 1.55 NLRP2 NLR family, pyrin domain containing 2 1.55 INPPL1 inositol polyphosphate phosphatase-like 1 1.54 PLEKHB2 pleckstrin homology domain containing, family B (evectins) member 2 1.54 KRT8 keratin 8 1.54 SELO selenoprotein O 1.54 CYP51A1 cytochrome P450, family 51, subfamily A, polypeptide 1 1.54 SCAF11 SR-related CTD-associated factor 11 1.54 GOLM1 golgi membrane protein 1 1.54 AFG3L2 AFG3-like AAA ATPase 2 1.54 C7orf13 chromosome 7 open reading frame 13 1.54 AP4E1 adaptor-related protein complex 4, epsilon 1 subunit 1.54 IPPK inositol 1,3,4,5,6-pentakisphosphate 2-kinase 1.54 TSNAX translin-associated factor X 1.54 ACAD10 acyl-CoA dehydrogenase family, member 10 1.54 MAGT1 magnesium transporter 1 1.54 POC5 POC5 centriolar protein 1.54 NSMCE1 non-SMC element 1 homolog (S. cerevisiae) 1.54 TMEM134 transmembrane protein 134 1.54 TFPT TCF3 (E2A) fusion partner (in childhood Leukemia) 1.54 MLF2 myeloid leukemia factor 2 1.54 ZNF582- ZNF582 antisense RNA 1 (head to head) 1.54 AS1 ZDHHC4 zinc finger, DHHC-type containing 4 1.54 ZNF264 zinc finger protein 264 1.54 FAM73A family with sequence similarity 73, member A 1.54 ATP7B ATPase, Cu++ transporting, beta polypeptide 1.54 PHTF1 putative homeodomain transcription factor 1 1.54 MKNK2 MAP kinase interacting serine/threonine kinase 2 1.54 GOLM1 golgi membrane protein 1 1.54 SLC39A14 solute carrier family 39 (zinc transporter), member 14 1.54 SIRT3 sirtuin 3 1.54 FOXRED2 FAD-dependent oxidoreductase domain containing 2 1.54 ZCWPW1 zinc finger, CW type with PWWP domain 1 1.54 POR P450 (cytochrome) oxidoreductase 1.54 LINC00938 long intergenic non-protein coding RNA 938 1.53 SNX27 sorting nexin family member 27 1.53 FAM173A family with sequence similarity 173, member A 1.53 HHLA3 HERV-H LTR-associating 3 1.53 IRS2 insulin receptor substrate 2 1.53 NPRL2 nitrogen permease regulator-like 2 (S. cerevisiae) 1.53 MAT2B methionine adenosyltransferase II, beta 1.53 DCAF4 DDB1 and CUL4 associated factor 4 1.53 LINC01347 long intergenic non-protein coding RNA 1347 1.53 LOC441455 makorin ring finger protein 1 pseudogene 1.53 ZNF688 zinc finger protein 688 1.53 UBE2Z ubiquitin-conjugating enzyme E2Z 1.53 TREX1 three prime repair exonuclease 1 1.53 AASDH aminoadipate-semialdehyde dehydrogenase 1.53 BCAP29 B-cell receptor-associated protein 29 1.53 GPAA1 glycosylphosphatidylinositol anchor attachment 1 1.53 PHOSPHO2 phosphatase, orphan 2 1.53 SURF1 surfeit 1 1.53 TCOF1 Treacher Collins-Franceschetti syndrome 1 1.53 CLPB ClpB caseinolytic peptidase B homolog (E. coli) 1.53 TTC7A tetratricopeptide repeat domain 7A 1.53 PLA2G12A phospholipase A2, group XIIA 1.53 TGFB1 transforming growth factor, beta 1 1.52 LARP4 La ribonucleoprotein domain family, member 4 1.52 RNASEK ribonuclease, RNase K 1.52 MIIP migration and invasion inhibitory protein 1.52 POLI polymerase (DNA directed) iota 1.52 DDRGK1 DDRGK domain containing 1 1.52 AK3 adenylate kinase 3 1.52 KIAA1430 KIAA1430 1.52 TMEM104 transmembrane protein 104 1.52 RUNDC1 RUN domain containing 1 1.52 MPI mannose phosphate isomerase 1.52 DNAJC21 DnaJ (Hsp40) homolog, subfamily C, member 21 1.52 ALG12 ALG12, alpha-1,6-mannosyltransferase 1.52 ZADH2 zinc binding alcohol dehydrogenase domain containing 2 1.52 FAM134A family with sequence similarity 134, member A 1.52 GALT galactose-1-phosphate uridylyltransferase 1.52 LEMD2 LEM domain containing 2 1.52 GADD45B growth arrest and DNA-damage-inducible, beta 1.52 DNAJB14 DnaJ (Hsp40) homolog, subfamily B, member 14 1.52 FAM45A family with sequence similarity 45, member A 1.51 FAM45A family with sequence similarity 45, member A 1.51 MED25 mediator complex subunit 25 1.51 CD164 CD164 molecule, sialomucin 1.51 KIAA0141 KIAA0141 1.51 TIMM13 translocase of inner mitochondrial membrane 13 homolog (yeast) 1.51 CLUAP1 clusterin associated protein 1 1.51 DENND4C DENN/MADD domain containing 4C 1.51 IPO13 importin 13 1.51 C11orf30 chromosome 11 open reading frame 30 1.51 KCTD21 potassium channel tetramerization domain containing 21 1.51 EXOC7 exocyst complex component 7 1.51 FBXO25 F-box protein 25 1.51 MSRB2 methionine sulfoxide reductase B2 1.51 C1orf43 chromosome 1 open reading frame 43 1.51 PARP10 poly (ADP-ribose) polymerase family, member 10 1.51 A1BG-AS1 A1BG antisense RNA 1 1.51 SDHD succinate dehydrogenase complex, subunit D, integral membrane protein 1.51 TRIM4 tripartite motif containing 4 1.51 TTC19 tetratricopeptide repeat domain 19 1.51 RNF149 ring finger protein 149 1.51 ADCK5 aarF domain containing kinase 5 1.51 UBE2MP1 ubiquitin-conjugating enzyme E2M pseudogene 1 1.51 NUFIP2 nuclear fragile X mental retardation protein interacting protein 2 1.51 PSENEN presenilin enhancer gamma secretase subunit 1.51 NAMPT nicotinamide phosphoribosyltransferase 1.51 COA5 cytochrome c oxidase assembly factor 5 1.50 MTX2 metaxin 2 1.50 RTCA RNA 3'-terminal phosphate cyclase 1.50 UQCR10 ubiquinol-cytochrome c reductase, complex III subunit X 1.50 ING4 inhibitor of growth family, member 4 1.50 BRAT1 BRCA1-associated ATM activator 1 1.50 REPIN1 replication initiator 1 1.50 TANK TRAF family member-associated NFKB activator 1.50 AKTIP AKT interacting protein 1.50 TMEM181 transmembrane protein 181 1.50 LOC100288 uncharacterized LOC100288069 1.50 069 IRF2BPL interferon regulatory factor 2 binding protein-like 1.50 APOA1BP apolipoprotein A-I binding protein 1.50

Supplementary Table 5

Primer sequences used for qPCR analysis Gene name Forward primer Reverse Primer Actin TGGCACCCAGCACAATGAA CTAAGTCATAGTCCGCCTAGAAGCA WT1 AGGGTACGAGAGCGATAACCACAC CTCAGATGCCGACCGTACAAGA NPHS1 CAACTGGGAGAGACTGGGAGAA AATCTGACAAGACGGAGCA NPHS2 GGAGGCTGAAGCGCAAAGAC GCCATCCTCAGGGACTCAGAAG SYNPO AGCCCAAGGTGACCCCGAAT CCCTGTCACGAGGTGCTGGC PODXL GGCAACGGCATTGAGAACA AGGGTGTTTCCTGTGAGGTTCAG