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. Author manuscript; available in PMC: 2016 Jul 9.
Published in final edited form as: Nature. 2015 Jun 24;523(7559):177–182. doi: 10.1038/nature14581

Glypican1 identifies cancer exosomes and facilitates early detection of cancer

Sonia A Melo 1,#,*, Linda B Luecke 1,*, Christoph Kahlert 1,*, Agustin F Fernandez 2, Seth T Gammon 3, Judith Kaye 1, Valerie S LeBleu 1, Elizabeth A Mittendorf 4, Juergen Weitz 5, Nuh Rahbari 5, Christoph Reissfelder 5, Christian Pilarsky 5, Mario F Fraga 2,6, David Piwnica-Worms 3, Raghu Kalluri 1,§
PMCID: PMC4825698  NIHMSID: NIHMS764674  PMID: 26106858

Summary

Exosomes are lipid bilayer-enclosed extracellular vesicles (EVs) that contain proteins and nucleic acids. They are secreted by all cells and circulate in the blood. Specific detection and isolation of cancer cell-derived exosomes in circulation is currently lacking. Using mass spectrometry analyses, we identified a cell surface proteoglycan, glypican-1 (GPC1), specifically enriched on cancer cell-derived exosomes. GPC1+ circulating exosomes (crExos) were monitored and isolated using flow cytometry from the serum of cancer patients and mice with cancer. GPC1+ crExos were detected in the serum of patients with pancreas cancer with absolute specificity and sensitivity, distinguishing healthy subjects and patients with a benign pancreas disease from patients with early and late stage pancreas cancer. Levels of GPC1+ crExos correlate with tumor burden and survival in patients pre- and post-surgical tumor resection. GPC1+ crExos from patients and from mice with spontaneous pancreas tumors driven by oncogenic KRAS contained RNA with specific KRAS mutation, and it emerges as a reliable biomarker for the detection of PanIN lesions despite negative signal by MRI in mice. GPC1+ crExos may serve as a potential non-invasive diagnostic and screening tool to detect early stages of pancreas cancer to facilitate possible curative surgical therapy.

Introduction

Exosomes are secreted membrane enclosed vesicles (extracellular vesicles or EVs) of a size range of 50 to 150 nm diameter1. Formed during the inward budding of late endosomes, they develop into intracellular multivesicular endosomes (MVEs)2. During this process, nucleic acids and proteins are encapsulated into exosomes36. Exosomes are released into the extracellular space and enter the circulation7,8. Various cell types such as immune cells8, platelets9 or endothelial cells10, release exosomes into the blood stream. Several exosomes-enriched proteins have been described and include members of the tetraspanin family (CD9, CD63 and CD81), members of the endosomal sorting complexes required for transport (ESCRT; TSG101, Alix) and heat shock proteins (Hsp60, Hsp70, Hsp90)11. Epithelial tumor cells secrete exosomes carrying the epithelial cell adhesion molecule (EpCAM)1214. Melanoma-derived exosomes contain the tumor-associated antigen Mart-1 and tyrosinase-related protein-2 (TYRP2)1517. Exosomes from gastric, breast and pancreas cancer carry members of the human epidermal growth factor receptor (HER) family1820. None of these markers are however specific to cancer-derived exosomes. Identification and isolation of cancer specific exosomes in body fluids would aid in the detection and monitoring of cancer and enable specific identification of DNA, RNA and protein content without contamination from non-cancer exosomes. Such possibility could enable the early monitoring of cancer and aid therapy decision.

GPC1 is a cancer exosomes specific protein

EVs from cancer cells (MDA-MB-231, triple negative human metastatic breast carcinoma), fibroblasts (HDF, human dermal fibroblasts; NIH/3T3, mouse embryonic fibroblasts) and non-tumorigenic epithelial cells (MCF10A, human mammary epithelial cells; E10, mouse lung epithelial cells) were isolated using established ultracentrifugation methods21,22, and called exosomes based on the following observations. NanoSight® nanoparticle tracking analysis and transmission electron microscopy (TEM) revealed a range of 105±5 nm and 112±4 nm in diameter, respectively (Extended Data Fig. 1a,b)23. Presence of CD9 on exosomes was shown by immunogold and TEM (Extended Data Fig. 1c) and of flotillin1 and CD81 by immunoblot (Extended Data Fig. 1d, Extended Data Fig. 8)23. The exosomes proteome was evaluated using ultra performance liquid chromatography – mass spectrometry (UPLC-MS) (Extended Data Table 1)24. Proteins were identified in exosomes derived from different cell types (HDF, NIH/3T3, E10, MCF10A and MDA-MB-231), including the exosomes markers such as TSG101, CD9 and CD63 (total number of proteins identified in exosomes from different cell types were: HDF: 261, NIH/3T3: 171, E10: 232, MCF10A: 214 and MDA-MB-231: 242; Extended Data Table 2). Bioinformatic analyses revealed 48 proteins (25 cytoplasmic, 7 nuclear, 5 transmembrane, 1 membrane-anchored and 7 secreted) as exclusively present in the cancer cell-derived exosomes (MDA-MB-231; Fig. 1a, Extended Data Table 1). Glypican-1 (GPC1) is a membrane anchored protein reported previously as overexpressed in a variety of cancers, including breast and pancreas cancer25,26,27. GPC1 transcripts and protein levels were elevated in several breast and pancreas cancer cell lines compared to non-tumorigenic cells (Extended Data Fig. 1e,f & Extended Data Fig. 8). Therefore, in this study we focused on probing the utility of GPC1 as marker of cancer exosomes. In contrast to exosomes derived from non-tumorigenic cell lines, GPC1 protein was only detected in cancer cell-derived exosomes by immunoblot (Extended Data Fig. 1g & Extended Data Fig. 8). Additionally, GPC1+ exosomes were detected by immunogold TEM in cancer exosomes (T3M4 pancreas cancer line) but not in non-cancer exosomes (HMLE; Fig. 1b).

Figure 1. GPC1 is present specifically on cancer exosomes.

Figure 1

a, Venn diagram of exosomal proteins from NIH/3T3 (blue), MCF 10A (red), HDF (green), E10 (yellow) and MDA-MB-231 (purple) cells. 48 proteins were exclusively detected in exosomes from MDA-MB-231 cells (n=3 protein samples, technical replicates). b, Transmission electron micrographs (TEM; left image) and immunogold TEM (right image) of GPC1. Upper right: digitally zoomed inset (n=2 experiments). c, Diagram of flow cytometry experiment to detect GPC1 on the surface of exosomes bound to beads. d, TEM of bead-bound exosomes and immunogold labeling of GPC1 (n=2 biological replicates). e, Percent beads with GPC1+ exosomes from cancer cells (red) and non-tumorigenic cells (black). f, Flow cytometry analyses of percent beads with GPC1+ exosomes from indicated cell lines (n=2 biological replicates). Negative control: secondary antibody only.

We performed FACS analysis of exosomes coupled to aldehyde/sulphate beads to detect GPC1 protein on their surface (Fig. 1c). Immunogold and TEM identified cancer exosomes at the surface of beads with GPC1 expression while non-tumorigenic exosomes did not exhibit GPC1 expression (Fig. 1d). Exosomes purified using sucrose gradients or ultracentrifugation from cell lines showed GPC1 expression specifically when derived from cancer cell lines (Fig. 1c, e–f; Extended Data Fig. 1h & Extended Data Fig. 8).

To determine whether GPC1+ exosomes could be isolated from the blood of tumor-bearing mice, we implanted MDA-MB-231 human breast cancer cells in the mammary fat pads of nude mice. The mice were bled prior to cancer cell inoculation, and repeatedly again when tumors reached an average volume of 300, 550, 1000 and 1350 mm3, and circulating exosomes (crExos) were isolated and assessed for the presence of GPC1 (Fig. 2a). The relative percentage of GPC1+ crExos increased proportionally with tumor growth and correlated with tumor burden (Fig. 2b–c; r=0.98, P=0.004). To further confirm the cancer cell origin of GPC1+ crExos, we engineered MDA-MB-231 cells to stably express CD63-GFP fusion protein. CD63 is an established exosomal marker22 and exosomes from these cells (MDA-MB-231-CD63GFP) in culture were uniformly positive for GFP (Fig. 2d). crExos were also collected from mice with orthotopic MDA-MB-231-CD63GFP tumors (~1500 mm3 in size), and a subpopulation of the crExos were GFP+ (Fig. 2e). GPC1 expression was exclusively detected in the GFP+ crExos fraction but not in the GFP crExos (Fig. 2f).

Figure 2. GPC1+ crExosomes are derived from cancer cells in tumor-bearing mice.

Figure 2

a, Longitudinal blood collection: nude mice with orthotopic MDA-MB-231 tumors (n=4 mice). b, Percentage of beads with GPC1+ exosomes harvested from systemic circulation (crExos) plotted against average tumor volume (n=4 mice, each sample analyzed in technical triplicates for GPC1). ANOVA, post-hoc Tamhane T2, ** P<0.01, *** P<0.001. Data is shown as mean ± standard deviation. c, Correlation between tumor volume and percent beads with GPC1+ crExos (Pearson correlation test). d, NanoSight® analyses of exosomes from cultured MDA-MB-231 CD63-GFP cells. Black: all exosomes; green: CD63-GFP+ exosomes (n=3 technical replicates). e, NanoSight® analyses of crExos from mice with MDA-MB-231 CD63-GFP orthotopic tumor. Black: all exosomes; green: CD63-GFP+ exosomes (n=3 technical replicates). f, Flow cytometry analyses of beads with exosomes from cultured MDA-MB-231 (upper left) and MDA-MB-231 CD63-GFP (upper middle) cells, and from crExos of mice with MDA-MB-231 CD63-GFP orthotopic tumors (lower left). Staining of CD63-GFP+ (cancer cell derived) and CD63-GFP (host derived) crExos for GPC1 (APC+, lower right; n=3 biological replicates and 3 technical replicates). Percent positive beads are listed. Negative control: secondary antibody alone (right upper panel).

GPC1+ circulating exosomes as a biomarker

Exosomes derived from cancer cell lines and circulating cancer cell-derived exosomes from tumor-bearing mice were almost exclusively positive for GPC1 (Fig. 1e and Fig. 2f). Next, we isolated crExos from patients with breast cancer (n=32), pancreatic ductal adenocarcinoma (PDAC, n=190) and healthy donors (n=100) (Patient data is shown in Extended Data Table 3). TEM of crExos isolated from the serum revealed a lipid bilayer and CD9 positivity (Extended Data Fig. 2a–b). crExos isolated using sucrose gradient also showed expression of exosomes marker, flotillin1 (Extended Data Fig. 2c & Extended Data Fig. 8)23,22. The relative concentration of crExos was significantly higher in the sera of cancer patients compared to healthy individuals (Extended Data Fig. 2d), and the average size of PDAC crExos was significantly smaller compared to all other crExos (Extended Data Fig. 2e). Analyses of sera from healthy individuals revealed baseline positivity for GPC1 in crExos, ranging from 0.3 to 4.7% (average of 2.3%; Fig. 3a). We observed that 24 out of 32 (75% of patients) breast cancer patients demonstrated crExos GPC1+ level higher that healthy individuals (P<0.0001; Fig. 3a). Any specific correlation between the level of GPC1+ crExos and breast cancer subtypes was not appreciated in this patient cohort (Luminal A, Luminal B or Triple Negative subtypes; Extended Data Fig. 2f), albeit group size may be too small to conclusively probe such correlation. In contrast, all 190 PDAC crExos revealed levels of GPC1+ crExos higher than levels noted in serum of healthy individuals (P<0.0001; Fig. 3a, Extended Data Fig. 7a–b). These results indicated a strong correlation between GPC1+ crExos and cancer, particularly for PDAC in this analysis. These results encouraged us to perform further analyses to potentially inform on the utility of GPC1+ crExos as a detection and monitoring tool for PDAC.

Figure 3. GPC1+ crExos is a non-invasive biomarker for pancreas cancer.

Figure 3

a, Percent beads with GPC1+ crExos in healthy donors (n=100), breast cancer patients (n=32) and patients with PDAC (n=190; ANOVA, post-hoc Tamhane T2, **** P<0.0001). b, Frequency of KRAS mutation in 47 tumor specimens and representative DNA sequencing chromatograms. c, TEM of GPC1 immunogold of crExos from three PDAC patients following flow cytometry isolation of GPC1+ (left column) and GPC1 (right column) crExos (n=3, 3 technical replicates). d, Ct value for KRAS G12D/G12V, KRAS WT mRNA and 18S rRNA expression in GPC1+ (red and blue) and GPC1 (black) crExos (after flow cytometry isolation; n=2, 2 biological replicates and 3 technical replicates each) e, Percent beads with GPC1+ crExos, discovery cohort (healthy donors, n=100; patients with a benign pancreatic disease (BPD), n=26; pancreatic cancer precursor lesion (PCPL), n=5; and PDAC patients, n=190; ANOVA, post-hoc Tamhane T2, ** P<0.01, **** P<0.0001; 3 technical replicates. f, ROC curve of discovery cohort. g, Percent beads with GPC1+ crExos, validation cohort (healthy donors, n=20; BPD, n=6; PDAC, n=56; ANOVA, post-hoc Tukey-Kramer test, **** P<0.0001; 3 technical replicates). h, ROC curve of validation cohort. Data is shown as mean ± standard deviation.

GPC1+ crExos contain oncogenic KrasG12D

Exosomes contain DNA and RNA28. KRAS is a frequently mutated gene in pancreas cancer and mutant transcripts have been found in circulation3,29,30. Primary tumor samples of 47 patients with PDAC were sequenced to assess oncogenic KRAS status. Sixteen PDAC tumors contained wild-type KRAS allele, 14 revealed KRASG12Dmutation, 11 with KRASG12Vallele, 5 with KRASG12R allele and 1 with KRASG12V/C mutation (Fig. 3b). Sufficient amount of corresponding serum was available from 10 patients with KrasG12D mutations and 5 with KrasG12V mutation. GPC1+ crExos and GCP1 crExos from the same patient were subjected to immunogold TEM to confirm specific GPC1 expression (Fig. 3c). All 15 GPC1+ crExos with tumor validated oncogenic Kras mutation revealed identical mutation by qPCR analysis of exosomal mRNA using specific primers (Fig. 3d). Wild-type Kras mRNA was found both in GPC1+ and GPC1 crExos, while mutant Kras transcript was only detected in the GPC1+ crExos (Fig. 3d). These results provide further support for the cancer cell origin of the GPC1+ crExos and highlight their potential utility identifying cancer specific genetic defects.

GPC1+ crExos detect early stage pancreas cancer

Further analysis of sera from the discovery cohort of patients revealed that the levels of GPC1+ crExos could distinguish patients with histologically validated pancreatic cancer precursor lesions (PCPL, n=5; Extended Data Table 3) from healthy individuals and patients with benign pancreatic disease (BPD, n=26; Extended Data Table 3, Extended Data Table 4, Fig. 3e). Specifically, the levels of GPC1+ crExos in the PCPL group (intraductal papillary mucinous neoplasm, IPMN n=5) was consistently higher than the levels of GPC1+ crExos in the healthy donor group, as well as in the BPD group (which includes 18 patients with pancreatitis and 8 with cystic adenomas; Fig. 3e). All patients in PCPL group presented with specific clinical symptoms and exhibited a macroscopic mass imaging using MRI or CT. The BPD group exhibited similar GPC1+ crExos levels (average 2.1% GPC1+ crExos) as healthy individuals (Fig. 3e).

We compared the specificity and sensitivity of GPC1+ crExos detection levels to CA 19-9 levels, a circulating protein currently used as a tumor biomarker for patients with pancreatic adenocarcinoma31. CA 19-9 levels were elevated in the serum of patients with PDAC when compared to healthy donors, but CA 19-9 levels were also significantly elevated in the serum of patients with BPD (P<0.0001; Extended Data Fig. 2g). Importantly, CA 19-9 serum levels failed to distinguish patients with PCPL from healthy donors (Extended Data Fig. 2g). When comparing patients with pancreatic cancer from stage I to stage IV, to healthy individuals and patients with BPD, the receiver operating characteristic (ROC) curves show that GPC1+ crExos revealed a near perfect classifier with an AUC of 1.0 (95% CI: 0.988 – 1.0) exhibiting a sensitivity of 100% (95% CI: 98.1–100%) and specificity of 100% (95% CI: 97.1–100%); with a positive predictive value of 100% (95% CI: 98.1–100%) and a negative predictive value of 100% (95%: 86.8–100%; Fig. 3f and Extended Data Fig. 3a–f). In contrast, CA 19–9 was inferior in distinguishing between patients with pancreas cancer and healthy controls (AUC of 0.739, 95% CI: 70.2–82.6%, P<0.001; Fig. 3f and Extended Data Fig. 3a–f). Of note, neither the concentration of exosomes nor the size of exosomes was a valid parameter to stratify patients with pancreas cancer versus control (Fig. 3f and Extended Data Fig. 3a–f). GPC1+ crExos showed a sensitivity and specificity of 100% in each stage of pancreas cancer (carcinoma-in-situ, stage I as well as stages II–IV), supporting its utility as a biomarker for all stages of pancreas cancer progression and its potential use for early detection of pancreas cancer.

Next, the discovery findings were validated using an independent patient cohort, composed of 6 patients with histologically validated BPD (chronic pancreatitis), 56 patients with PDAC and 20 healthy individuals (Extended Data Table 3). GPC1+ crExos distinguished patients with PDAC from healthy individuals and patients with BPD (Fig. 3g). The BPD group exhibited similar GPC1+ crExos levels as healthy individuals (Fig. 3g). In complete agreement with the discovery cohort, receiver operating characteristic (ROC) curves indicated that GPC1+ crExos (from PDAC, BPD patients and healthy individuals) revealed a near perfect classifier with an AUC of 1.0 (95% CI: 0.956 – 1.0) a sensitivity of 100% (95% CI: 93.6–100%), a specificity of 100% (95% CI: 86.8–100%), a positive predictive value of 100% (95% CI: 93.6–100%), and a negative predictive value of 100% (95%: 86.3–100%; Fig.3h, Extended Data Fig. 3g).

GPC1+ crExos inform pancreas cancer burden

GPC1+ crExos levels correlated with tumor burden in mice (Fig. 2b–c). We next sought to evaluate whether GPC1+ crExos levels could inform on metastatic disease burden of patients with PDAC (Extended Data Table 3). GPC1+ crExos of PDAC patients with distant metastatic disease showed significantly higher levels of bead bound-GPC1+ crExos (average 58.5%) when compared to patients with metastatic disease restricted to lymph nodes (average 50.5%) or no metastases (average 39.9%; Extended Data Fig. 4a). Further, we evaluated GPC1+ crExos in serum of PDAC patients at pre- and post surgery stages (post operative day 7; PDAC n=29, PCPL n=4 and BPD n=4; Fig. 4a). Twenty-eight out of 29 PDAC patients and all PCPL patients with longitudinal blood collections showed a significant decrease in GPC1+ crExos levels following surgical resection (PDAC: P<0.0001; PCPL: P<0.001; Fig. 4b). In contrast, CA 19-9 levels decreased in only 19 out of 29 PDAC patients and in none of the PCPL patients (PDAC: P=0.003; PCPL: P=0.81; Extended Data Fig. 4b). In 4 BPD patients, the levels of neither GPC1+ crExos nor CA 19-9 showed a difference (Fig. 4b and Extended Data Fig. 4b).

Figure 4. Levels of circulating GPC1+ exosomes inform pancreas cancer resection outcome.

Figure 4

a, Longitudinal blood collection pre- (pre-op) and post-operatively (day 7). b, Percent beads with GPC1+ crExos from patients with BPD (n=4), PCPL (n=4) or PDAC (n=29) (paired two-tailed Student’s t-test, ** P<0.01, **** P<0.0001). Data is shown as mean ± standard deviation. c–d, Kaplan–Meier curves (log-rank test) displaying overall (c) and disease-specific survival (d) of patients with GPC1+ crExos drop ≥ median (blue) and GPC1+ crExos drop < median (green) drop after resection. e–f, Kaplan–Meier curves (log-rank test) displaying overall (e) and disease-specific survival (f) of patients with a CA 19-9 drop ≥ median (blue) and a CA 19-9 drop < median (green) drop after resection.

To determine the prognostic relevance of GPC1+ crExos in this longitudinal study cohort (Fig. 4a), patients were dichotomized into 2 groups. Group 1 was defined by a decrease of GPC1+ crExos greater or equal (≥) to the median decrease in GPC1+ crExos, and group 2 was defined by a decrease of GPC1+ crExos less (<) than the median decrease of GPC1+ crExos. Group 1 presented with improved overall (26.2 months) and disease specific (27.7 months) survival when compared to group 2 (15.5 months for both overall and disease specific), indicating that a greater decrease in GPC1+ crExos levels after surgery is associated with increased survival (Fig. 4c–d). While a decrease in CA 19-9 levels is noted when comparing pre- and post-resection blood draws, this decrease did not significantly associate with overall and disease-specific survival (Fig. 4e–f and Extended Data Fig. 4b). Using a Cox regression model for a multivariate test to include the decrease in GPC1+ crExos, median age, AJCC stage, tumor grade and CA 19-9 levels, only GPC1+ crExos revealed to be an independent prognostic and predictive marker for disease-specific survival (hazard ratio: 5.353, CI: 1.651–17.358, P=0.005; Extended Data Fig. 4c–d).

Next, we evaluated whether an ELISA for circulating GPC1 could function with the same specificity and sensitivity as GPC1+ crExos. Serum samples of the validation cohort (20 healthy individuals, 6 BPD patients and 56 PDAC patients) were analyzed for circulating GPC1. While GPC1 levels were significantly higher in the PDAC patients when compared to BPD patients and healthy individuals, the sensitivity and specificity of the assay was lower when compared to GPC1+ crExos. The GPC1 ELISA was similar to circulating CA 19-9 assay. Receiver operating characteristic curve (ROC) indicated that GPC1 circulating protein shows an AUC of 0.781 (95% CI: 0.675–0.865) a sensitivity of 82.14% (95% CI: 69.6–91.9%), a specificity of 75% (95% CI: 54.9–90.6%), a positive predictive value of 0.04 (95% CI: 0 – 17.4%), and a negative predictive value of 100% (95%: 94.2 – 100%; Extended Data Fig. 4e–f).

GPC1+ crExos to detect early PanIN lesions

In light of the highly specific and sensitive detection of GPC1+ crExos in pancreas cancer, we next evaluated time course of GPC1+ crExos appearance in the serum relative to pancreas tumor burden. To this end, we employed a genetically engineered mouse model (GEMM) for PDAC. The Ptf1acre/+; LSL-KrasG12D/+; Tgfbr2L/L mice (PKT mice)32 develop PDAC with full penetrance that reliably recapitulates the clinical and histopathological features of the human disease32,33. The PKT mice consistently progress from pancreatic intraepithelial neoplasia (PanIN) at 4.5 weeks of age and die at 8 weeks of age due to PDAC32,33 (Extended Data Fig. 5a). In a longitudinal study, we bled PKT and littermate control mice repeatedly at 4, 5, 6, 7 and 8 weeks of age (n=7 PKT mice and n=6 control mice; Fig. 5a). Three out of 7 PKT mice were euthanized by week 7 along with 4 out of 6 controls, while the remaining 3 PKT mice and 2 controls were euthanized at week 8. At 4 weeks of age PKT mice showed on average an 8.4% GPC1+ crExos, and this increased proportionally with time (and tumor burden) and severity of disease (histopathology), whereas control mice showed an average of 1.2% GPC1+ crExos and this level remained constant with time (Fig. 5b). crExos sizes and concentration did not consistently correlate with disease over time (Extended Data Fig. 5b,c). Magnetic resonance imaging (MRI), an established imaging modality used for the evaluation of PDAC34, was performed at the same time points when mice were bled to measure GPC1+ crExos (Fig. 5c). When evaluated as a group, GPC1+ crExos levels appeared prior to MRI detectable pancreatic masses (Fig. 5c–d and Extended Data Fig. 5d). GPC1+ crExos size and concentration minimally correlated with pancreas cancer (Extended Data Fig. 5b–c), whereas GPC1+ crExos levels correlated with tumor volume determined by MRI, and appeared to lead the growth of the tumor (Pearson correlation test, r=0.67, P=0.0005, 95% CI: 0.3504–0.8462; Fig. 5c,d and Extended Data Fig. 5d). Importantly, no elevation of GPC1+ crExos was noted in mice cerulein-induced acute pancreatitis, supporting GPC1+ crExos is elevation is pancreas cancer specific (Extended Data Fig. 5e). ROC curve analysis for GPC1+ crExos showed an AUC of 1.0 (95% CI: 0.75–1.0) in PKT mice compared to healthy littermate control mice at all ages evaluated (Fig. 5e and Extended Data Fig. 5f).

Figure 5. GPC1+ circulating exosomes predict pancreas cancer in GEMM.

Figure 5

a, Longitudinal blood collection from control and PKT mice: 4 (n=6 and n=7, respectively), 5 (n=6 and n=7), 6 (n=6 and n=6), 7 (n=6 and n=6) and 8 (n=2 and n=3) weeks of age. b, Percent beads with GPC1+ crExos from PKT (red) and control (blue) mice; ANOVA, post-hoc Tukey-Kramer test, **** P<0.0001; 3 technical replicates. c, MRI with tumor encircled in red. d, Tumor volume and percent GPC1+ crExos in PKT mice at indicated age (ANOVA, post-hoc Tamhane T2), *P<0.05, ** P<0.01, *** P<0.001**** P<0.0001; 3 technical replicates) e, ROC curve analysis of 4 weeks-old control mice (n=6) and mice with acute pancreatitis (n=4) vs. 4 weeks-old PKT mice (n=7). f, Cross sectional study: blood collected from 16 or 20 days old control (n=6) and PKT (n=7) mice. g, Percent beads with GPC1+ crExos from control and PKT (16 – 20 days-old) mice (paired two-tailed Student’s t-test, P<0.0001; 3 technical replicates). h–i, Correlation between histological score (h) or age (i) and percent beads with GPC1+ crExos (Pearson correlation test). j, Relative percent PanIN lesions and representative staining for phosphorylated ERK. Data is shown as mean ± standard deviation.

A cross-sectional study was also initiated to assay tumor burden and GPC1+ crExos in PKT mice, as early as 16 and 20 days of age (Fig. 5f). Mice were imaged by MRI, bled and euthanized at these early time points, when mice present with pre-PanIN to early PanIN lesions (Fig. 5f, Extended Data Fig. 6a and Extended Data Table 5). GPC1+ crExos were detected in all PKT mice (PKT: 8.3% average, control: 1.8% average; Fig. 5g and Extended Data Table 5). Histological analysis of PKT mice confirmed pre-PanIN lesions in 4 out of 7 PKT mice, and despite no observed histological lesions in 3 out of 7 PKT mice, GPC1+ crExos predicted future pancreas cancer emergence (Extended Data Table 5). Moreover, we did not observe pancreas-associated masses by MRI in 16 and 20 days old PKT mice. Both histopathological score and age of PKT mice correlated with GPC1+ crExos levels (Fig. 5h–i). Of note, in 4 out of 7 PKT mice with no observed histological lesions, downstream signals for Kras activation, such as phosphorylated ERK (pERK), was detected in the pancreas tissue (Fig. 5j and Extended Data Fig. 6a). We also observed exclusive detection of mutant KrasG12D mRNA in GPC1+ crExos compared to GPC1 crExos (Extended Data Fig. 6b).

Discussion

Tumor-derived exosomes are enriched in GPC1, and GPC1+ crExos exclusively carry mutant Kras mRNA. We show that GPC1+ crExos is a reliable biomarker for detection of early pancreas cancer. GPC1+ crExos are a prognostic marker superior to CA 19-9, the currently employed serum biomarker in pancreas cancer. GPC1+ crExos lead MR imaging as they can be detected in circulation prior to MRI-detectable lesions in GEMM of pancreas cancer.

Routine screening of the general population for a rare cancer such as PDAC using MRI or CT would be prohibitively expensive and associated with a high false positive rate35. GPC1+ crExos detect possibility of pancreas cancer in 16-day-old mice with unremarkable pancreas histology and negative MRI. These results suggest the utility of GPC1+ crExos as a detection and monitoring tool for pancreas cancer, with an emphasis on its application in early detection, a clinical challenge that remains to be addressed.

While oncogenic KRAS mutations are likely driver mutations for pancreas cancer and are detected in early PanIN-1 lesions, it is still estimated that 15 to 20 years may lapse before early PanIN lesions become metastatic PDAC3638. While this could offer a window for early detection of this disease, PDAC currently presents late with rather non-specific clinical symptoms. Therefore, as many as 80% of patients present with regional and distant metastasis at diagnosis39. Although our understanding of the molecular and genetic details associated with the progression of non-malignant precursor lesions to PDAC is steadily increasing, such knowledge has yet to translate into the development of biomarkers that do not require invasive biopsies. Patients with pancreas cancer exhibit elevated serum levels of CA 19-9, CEA, CA-50, SPan-1, peanut agglutinin, DU-PAN-2, a-fetoprotein, tissue polypeptide antigen and pancreatic oncofetal antigen40. While each of these markers exhibits some utility in tracking biopsy-diagnosed disease they are not tumor specific because they are also elevated in patients with benign diseases of the pancreas. The lack of specific serum-based biomarkers with high degree of sensitivity and specificity and retroperitoneal position of the pancreas further challenges the early detection of pancreas cancer41,42. Pancreatico-duodenectomy (Whipple procedure) can be curative for PDAC patients if tumors are detected early with clear surgical margins43. Due to the late diagnosis of pancreas cancer, only around 15% of patients present with surgically resectable tumors44. Studies comparing stage of disease with outcome following surgery suggests that death rates for PDAC would be reduced if the disease were diagnosed at an earlier stage45.

Specific isolation of exosomes from the serum of cancer patients remains a challenge due to the lack of specific markers that can be used to identify and distinguish cancer exosomes from exosomes produced by other cells. Genetic profiling studies on circulating DNA from cancer patients are cofounded by the fact that the isolated DNA represents all cells of the body, thus making mutation and genetic defects challenging46,47. We previously demonstrated that the DNA in circulation is mainly contained in exosomes6. Therefore a marker for cancer-derived exosomes will significantly increase the sensitivity of detection for low frequency mutations in the circulation. Such mutations are frequently associated with tumor complexity and therapy resistance and relapse, providing a highly specific and sensitive non-invasive tool for genetic monitoring of a tumor. As a proof of concept GPC1+ crExos identified KRAs mutations with 100% correlation with KRAS mutations in the tumor.

Our results provide evidence for GPC1 as a pan-specific marker of cancer exosomes. GPC1 is a cell surface proteoglycan that interacts with many proteins and has diverse functions48. Many cancer cells overexpress glypican-1 with most abundant increase observed in pancreas cancer cells lines and tissue2527. Many studies have suggested a role of GPC1 as a positive regulator of cancer progression using orthotopic and GEMMs of PDAC49,50. GPC1+ crExos is an attractive candidate for detection and isolation of exosomes in circulation of cancer patients for further genetic analysis of cancer specific alterations in DNA, microRNA, RNA and proteins. Such opportunity offers an unprecedented opportunity for informative early detection of pancreas cancer and help design potential curative surgical options.

Methods

Patient samples and tissue collection

The study was conducted according to the Reporting Recommendations for Tumor Marker Prognostic Studies (REMARK) criteria. The studies using human samples were designed as an explorative study. As there was no interventional approach in this study, a priori power calculation was not applicable. Instead, the number of patients included was assessed based on previous studies investigating the diagnostic relevance of circulating biologicals in pancreatic cancer51.

Serum samples and tissue samples from patients with pancreas cancer, serum samples only from patients with a benign pancreas disease and from healthy donors, who had no evidence of acute or chronic or malignant disease and had no surgery within the past 12 months, were received from the department of General, Visceral and Transplantation Surgery from the University of Heidelberg and from the University Hospital of Dresden after approval by the local institutional review board (Heidelberg: 323/2004, Dresden: EK357112012). The cases were obtained under an IRB-exempt protocol of the MD Anderson Cancer Center (IRB no. PA14-0154). Serum samples from patients with breast cancer were collected at the MD Anderson Cancer Center after approval of the institutional review board (IRB no. LAB10-0690). A written consent for the serum sampling and tumor sampling was obtained pre-operatively from all patients and prior to serum collection from each healthy donor with disclosure of planned analyses regarding potential prognostic markers. The patients included in this study were all consecutive patients who underwent a surgical procedure at the University Hospital of Heidelberg, Germany, at the University Hospital of Dresden, Germany (pancreas disease) or at the MD Anderson Cancer Center (breast cancer). All samples were randomly selected from larger cohorts and were analyzed in a blinded fashion. Unblinding of clinical parameters and corresponding experimental data was performed only after finishing all experiments. Inclusion criteria of patients were a minimum of 18 years of age, histologically verified pancreas cancer (pancreatic ductal adenocarcinoma), histologically verified benign pancreas disease or breast cancer in a resection specimen, and a negative medical history for any other malignant disease. All blood samples were taken before treatment. Inclusion criteria for healthy control donors were a negative medical history for any malignant disease.

On the day of surgery, 10 ml serum separator tubes were used to collect blood samples before surgical incision. The blood samples were then centrifuged at 2,500g for 10 min to extract the serum, and the serum was stored at −80°C until analyzed. Likewise, blood samples were collected on day 7 after surgery for 29 patients with an adenocarcinoma of the pancreas (PDAC), 4 patients with chronic pancreatitis and 4 patients with an intraductal papillary mucinous neoplasm (IPMN).

Patient characteristics and clinical specimens

The pancreatic discovery cohort from the University Hospital of Heidelberg included 190 patients with an adenocarcinoma of the pancreas (PDAC), 18 patients with pancreatitis, 8 patients with a benign serous cystadenoma and 5 patients with an intraductal papillary mucinous neoplasm (IPMN). Patients were subjected to surgery between 2006 and 2012 at the Department of General, Visceral, and Transplantation Surgery, University of Heidelberg. Clinical information included age, gender, AJCC tumor stage, tumor size (pT), presence and number of lymph node metastases (pN), tumor grade (G), and treatment with (neo-)/adjuvant chemotherapy. The pancreatic cohort from the University Hospital of Dresden included 56 patients with an adenocarcinoma of the pancreas (PDAC), 6 patients with chronic pancreatitis, and 20 healthy donors. Patients were subjected to surgery between 2007 and 2013 at the Department of Gastrointestinal, Thoracic and Vascular Surgery, University of Dresden. Clinical information included age, gender, AJCC tumor stage, tumor size (pT), presence and number of lymph node metastases (pN), tumor grade (G), and treatment with (neo-)/adjuvant chemotherapy. The breast cancer cohort consisted of 32 women with breast cancer. All breast cancer patients were treated at the MD Anderson Cancer Center, Houston, Texas. Clinical information included age, gender, AJCC tumor stage, tumor size (pT), presence and number of lymph node metastases (pN), tumor grade, and treatment with (neo-)/adjuvant chemotherapy.

Animal Studies

Nude mice (nu/nu) (purchased from Jackson Laboratory) underwent breast pad injections with 0.5 million MDA-MB-231 cells or MDA-MB-231-CD63GFP cells in 20 μl of PBS injected per breast pad. Blood was collected retro-orbitally and exosomes were isolated prior to injection and at tumor volumes of 300, 550, 1000 and 1350mm3. Mice were euthanized when the tumor size reached 1500 mm3 or when severe disease symptoms were present.

The disease progression and genotyping for the Ptf1acre/+; LSL-KrasG12D/+; Tgfbr2L/L (PKT) and the Pdx1cre/+; LSL-KrasG12D/+; p53R172H/+ (KPC) mice was previously described32,33,52. In the PKT longitudinal cohort, retro-orbital blood collections were performed at 4, 5, 6, 7 and 8 weeks of age. Mice were euthanized at 8 weeks of age or sooner if severe disease symptoms were noted. Histopathological analysis of mouse pancreas specimen was performed following previously defined criteria4. Four C57BL/6 adult mice were subjected to repeated cerulean injection to induce acute pancreatitis (five hourly repeated i.p. injections of 50 μg cerulein per kg of body weight) and euthanized 24 hrs after injection the last injection. Histological analyses of pancreas of mice was performed according to Hingorani SR et al.52, and an histological score was attributed according to the type of lesions detected: Score 1: PanIN1a, Score 2: PanIN1 a/b, Score 3: PanIN2, Score 4: PanIN3, Score 5: Ductal adenocarcinoma. All mice were housed under standard housing conditions at the MD Anderson Cancer Center (MDACC) animal facilities, and all animal procedures were reviewed and approved by the MDACC institutional animal care and use committee.

Cell lines

The following human cells lines were used: HMLE (American Type Culture Collection – ATCC, Manassas, VA), BJ (ATCC), HDF (ATCC), HMEL (ATCC), MCF-7 (ATCC), MDA-MB-231 (ATTC), Panc-1 (ATTC), SW480 (ATCC), HCT-116 (ATCC), MIA Paca2 (ATCC) and T3M4 cells (Cell Bank, RIKEN BioResource Centre, Japan). The following murine cells lines were used: NIH/3T3 (ATCC), E10 (ATCC), NMuMG (ATTC), 4T1 (ATTC) and B16F10 cells (ATTC). HDF cells were cultured in DMEM supplemented with 20% (v/v) fetal bovine serum (FBS), 100 U.ml−1 penicillin and 100 μg.ml−1 streptomycin. HMLE cells and MCF10A cells were grown in DMEM/F12 supplemented with 5% (v/v) horse serum, 100 U.ml−1 penicillin, 100 μg.ml−1 streptomycin, 20 ng.ml−1 EGF, 0.5 mg.ml−1 hydrocortisone, 100 ng.ml−1 cholera toxin and 10 μg.ml−1 insulin. HMEL, MCF-7, MDA-MB-231, HCT-116, SW480, 4T1, NIH/3T3, E10, U87 and B16F10 cells were maintained in DMEM supplemented with 10% (v/v) FBS, 100 U.ml−1 penicillin and 100 μg.ml−1 streptomycin. Panc-1, MIA Paca2 and T3M4 cells were cultured in RPMI-1640 supplemented with 10% (v/v) FBS, 100 U.ml−1 penicillin and 100 μg.ml−1 streptomycin. NMuMG cells were grown in DMEM supplemented with 10% (v/v) FBS, 100 U.ml−1 penicillin, 100 μg.ml−1 streptomycin and 10 μg.ml−1 insulin. All cell lines were kept in a humidifying atmosphere at 5% CO2 at 37°C. MDA-MB-231-CD63GFP cells were engineered by transfection with a plasmid encoding a CD63-GFP fusion protein expressed under the control of a CMV promoter (p-CMV6-CD63-GFP from Origene, RG217238). Transfections were performed using Lipofectamine 2000 reagent (Invitrogen).

Exosomes isolation from cells

Exosomes were obtained from supernatant of cells as previously described with some modifications6. Briefly, cells were grown in T225 cm2 flasks in FBS-depleted of exosomes RPMI media until they reached a confluency of 80–90%. Next, the media was collected and centrifuged at 800g for 5 min, followed by a centrifugation step of 2,000g for 10 min to discard cellular debris. Then, the media was filtered using a 0.2 μm pore filter (Syringe filter, Cat. No. 6786-1302, GE Healthcare, UK). The collected media was then ultracentrifuged at 100,000g for 2 hrs at 4°C. The exosomes pellet was washed with 35 ml 1X PBS, followed by a second step of ultracentrifugation at 100,000g for 2 hrs at 4°C. Afterwards, the supernatant was discarded. Exosomes used for RNA extraction were resuspended in 500 μl of Trizol; exosomes used for protein extraction were resuspended in 250 μl of lysis buffer (8M Urea/2.5%SDS, 5 μg.ml−1 leupeptin, 1 μg.ml−1 pepstatin and 1mM phenylmethylsulphonyl fluoride). Exosomes used for flow cytometry analysis (FACS), transmission electron microscopy (TEM; see sections below) and immunogold staining were resuspended in 100 μl 1X PBS. Ten microliters of these exosomes sample were used for NanoSight LM10 (NanoSight Ltd., Minton Park, Amesbury, UK) analysis after dilution 1:100 in 1X PBS.

Exosomes isolation from human serum samples

As previously described, 250 μl of cell-free serum samples were thawed on ice6. Serum was diluted in 11 ml 1X PBS and filtered through a 0.2 μm pore filter. Afterwards, the samples were ultracentrifuged at 150,000g overnight at 4°C. Next, the exosomes pellet was washed in 11 ml 1X PBS followed by a second step of ultracentrifugation at 150,000g at 4°C for 2 hrs. The supernatant was discarded and pelleted exosomes were resuspended in 500 μl of Trizol for RNA analyses; or in 250 μl of lysis buffer (8M Urea/2.5%SDS, 5 μg.ml−1 leupeptin, 1 μg.ml−1 pepstatin and 1mM phenylmethylsulphonyl fluoride) for protein analyses. Exosomes used for flow cytometry analysis (FACS), transmission electron microscopy (TEM; see sections below) and immunogold staining were resuspended in 100 μl 1X PBS. Ten microliters of this exosomes sample were used for NanoSight LM10 (NanoSight Ltd., Minton Park, Amesbury, UK) analysis after Nano dilution 1:100 in 1X.

Immunogold Labeling and Electron Microscopy

Fixed specimens at an optimal concentration were placed onto a 400 mesh carbon/formvar coated grids and allowed to absorb to the formvar for a minimum of 1 minute. For immunogold staining the grids were placed into a blocking buffer for a block/permeabilization step for 1 hr. Without rinsing, the grids were immediately placed into the primary antibody at the appropriate dilution overnight at 4°C (1:300 anti-CD9 ab92726, Abcam and anti-GPC1 PIPA528055, Thermo Scientific). As controls, some of the grids were not exposed to the primary antibody. The following day, all the grids were rinsed with PBS then floated on drops of the appropriate secondary antibody attached with 10 nm gold particles (AURION, Hatfield, PA) for 2 hrs at room temperature. Grids were rinsed with PBS and were placed in 2.5% Glutaraldehyde in 0.1M Phosphate buffer for 15 min. After rinsing in PBS and distilled water the grids were allowed to dry and stained for contrast using uranyl acetate. The samples were viewed with a Tecnai Bio Twin transmission electron microscope (FEI, Hillsboro, OR) and images were taken with an AMT CCD Camera (Advanced Microscopy Techniques, Danvers, MA).

Sucrose gradient

Sucrose density gradients were performed to purify exosomes. Exosomes were resuspended in 2 ml of HEPES/sucrose stock solution (2.5M sucrose, 20mM HEPES/NaOH solution, pH 7.4). The exosomes suspension was overlaid with a linear sucrose gradient (2.0-0.25M sucrose, 20mM HEPES/NaOH, pH 7.4) in a SW41 tube (Beckman). The gradients were ultracentrifuged for 16 hrs at 210,000g at 4°C. Gradient fractions of 1 ml were collected from top to bottom and densities of each fractions were evaluated using a refractometer. Next, the exosomes pellets were washed in 1X PBS followed by a second step of ultracentrifugation at 150,000g at 4°C for 2 hrs. Exosomes pellets were resuspended in Laemmli buffer and/or PBS for further immunoblot and flow cytometry analysis.

Flow cytometry analysis of exosomes-bound beads

Exosomes were attached to 4 μm aldehyde/sulfate latex beads (Invitrogen, Carlsbad, CA, USA) by mixing 30 μg exosomes in a 10 μl volume of beads for 15 min at room temperature with continuous rotation. This suspension was diluted to 1 ml with 1X PBS and left for 30 min rotating at room temperature. The reaction was stopped with 100mM glycine and 2% BSA in 1X PBS and left 30 min rotating at room temperature. Exosomes-bound beads were washed 1 time in 1X PBS/2% BSA and centrifuge for 1 min at 10,000 rpm, blocked with 10% BSA with rotation at room temperature for 30 min, washed a second time in 1X PBS/2% BSA and centrifuged for 1 min at 10,000 rpm, and incubated with anti-GPC1 (PIPA528055, Thermo-Scientific, 3 μl of antibody in 20 μl of 2%BSA/1X PBS) during 30 min rotating at 4°C. Beads were centrifuged for 1 min at 10,000 rpm, the supernatant was discarded and beads were washed in 1X PBS/2% BSA and centrifuged for 1 min at 10,000 rpm. Alexa-488 or Alexa-594-tagged secondary antibodies (Life Technologies, NY 14072, USA, 3 μl of antibody in 20 μl of 2%BSA/1X PBS) were used during 30 min with rotation at 4°C. Secondary antibody incubation alone was used as control and to gate the beads with GPC1+ bound exos. The percent positive bead was calculated relative to the total number of beads analyzed per sample (100,000 events). This percentage was therein referred to as the percent beads with GPC1+ exosomes.

Ultra performance liquid chromatography – mass spectrometry (UPLC-MS)

Exosomes were mixed with 200 μl of methanol spiked with the Internal Standard tryptophan-d5. After brief vortex mixing, the samples were incubated for 1 hr at −20°C. After centrifugation at 16,000g for 15 min at 4°C, 190 μl of the supernatants was collected and the solvent removed. The dried extracts were then reconstituted in 15 μl of methanol, of which 10 μl were transferred to microtubes and derivatized. Chromatographic separation and mass spectrometric detection conditions employed are summarized below:

System SQD
Column type UPLC BEH C18, 1.0 x 100 mm, 1.7 μm
Flow rate 0.14 ml/min
Solvent A H2O + 10mM Ammonium Bicarbonate (+NH4OH until pH: 8.8)
Solvent B ACN
(%B), time 2%, 0 min
(%B), time 8%, 6.5 min
(%B), time 20%, 10 min
(%B), time 30%, 11 min
(%B), time 99.9%, 12 min
(%B), time 2%, 14 min
Column temperature 40°C
Injection volume 1 μl
Ionisation ES+
Source temperature 120°C
Nebulisation N2 flow 600 l / hour
Nebulisation N2 temperature 350°C
Cone N2 flow 10 l / hour
Capillary voltage 3.2 kV
Cone voltage 30 V

The mass range, 50 – 1000 m/z, was calibrated with cluster ions of sodium formate. An appropriate test mixture of standard compounds was analyzed before and after the entire set of randomized duplicated sample injections, in order to examine the retention time stability and sensitivity of the LC/MS system throughout the course of the run. Data were processed using the TargetLynx application manager for MassLynx 4.1 software (Waters Corp., Milford, USA). A set of predefined retention time, mass-to-charge ratio pairs, Rt-m/z, corresponding to metabolites included in the analysis are fed into the program. Associated extracted ion chromatograms (mass tolerance window = 0.05 Da) are then peak-detected and noise-reduced in both the LC and MS domains such that only true metabolite related features are processed by the software. A list of chromatographic peak areas is then generated for each sample injection, using the Rt-m/z data pairs (retention time tolerance = 6 s) as identifiers. Normalization factors were calculated for each metabolite by dividing their intensities in each sample by the recorded intensity of the internal standard in that same sample. Visualization of disjoint and overlapping protein data sets was carried out by drawing a VennDiagram of the 5 protein data sets using an R package53.

Cancer Antigen CA 19-9 Human and GPC1 ELISAs

Serum CA 19-9 and GPC1 protein levels in patients with pancreatic cancer, pancreatic cancer precursor lesion, or benign pancreatic disease, and in healthy donors were assessed using the Cancer Antigen CA 19-9 Human ELISA Kit (Abcam, ab108642) and the GPC1 Human ELISA kit (ABIN840422), according to the manufacturer’s directions.

Western blot analyses

Cells were lysed in RIPA buffer containing 5 μg.ml−1 leupeptin, 1 μg.ml−1 pepstatin and 1mM phenylmethylsulphonyl fluoride. Exosomes were lysed in 8M Urea/2.5% SDS containing 5 μg.ml−1 leupeptin, 1 μg.ml−1 pepstatin and 1mM phenylmethylsulphonyl fluoride. Sample loading was normalized according to Bradford relative protein quantification and proteins separated following an electrophoretic gradient across polyacrylamide gels. Wet electrophoretic transfer was used to transfer the proteins in the gel onto PVDF membranes (ImmobilonP). The protein blot was blocked for 1hr at room temperature with 5% non-fat dry milk in PBS/0,05% Tween and incubated overnight at 4°C with the following primary antibodies: 1:300 anti-GPC1, PIPA528055 (Thermo-Scientific); 1:300 anti-β-Actin A3854 (Sigma-Aldrich); 1:300 anti-CD81 sc-166029 (Santa-Cruz); 1:300 anti-Flottilin1 sc-25506 (Santa-Cruz). Afterwards, HRP conjugated secondary antibodies were incubated for 1 hr at room temperature. Washes after antibody incubations were done on an orbital shaker, four times at 10 min intervals, with 1X PBS 0.05% Tween20. Blots were developed with chemiluminescent reagents from Pierce.

RNA extraction of cells and exosomes

RNA of cells and exosomes was isolated using Trizol Plus RNA purification kit (Life Technologies, 12183555) according to manufactures protocol. RNA was quantified using a Nanodrop® ND-1000 (Thermo Fischer Scientific).

Quantitative real-time-PCR (qRT-PCR)

qRT-PCR was performed on DNase treated RNA using the SuperScript® III Platinum® One-Step Quantitative RT-PCR System (Cat. no. 11732-088, Invitrogen, Life Technologies, Grand Island, NY 14072 USA) according to the manufacturer’s directions on a 7300 Sequence Detector System (Applied Biosystems). 150 ng of RNA extracted from 2.5 × 108 exosomes was used as qPCR input. Primers for KRAS G12D mRNA and KRAS G12V mRNA (both Sigma-Aldrich Corp. St. Louis, MO, USA) were designed as reported previously54. Briefly, the altered base of KRAS G12D and KRAS G12V mutation was kept at the 3′ end of the forward primer. An additional base mutation was included two positions before the Kras mutation in order to increase the specificity of the amplification of the mutant Kras allele. Forward primer sequences for KRAS G12D mRNA: F-5′-ACTTGTGGTAGTTGGAGCAGA-3′. Forward primer sequences for KRAS G12V mRNA: F-5′-ACTTGTGGTAGTTGGAGCAGT-3′. Forward primer sequences for KRAS wild-type mRNA: F-5′-ACTTGTGGTAGTTGGAGCTGG-3′. Reverse primer for all KRAS: R-5′-TTGGATCATATTCGTCCACAA-3′. GPC1 mRNA primer pairs (Cat. No. PPH06045A) and 18S mRNA primer pairs (Cat. No. QF00530467) were purchased as ready specific primer pairs from Qiagen (Qiagen, Hilden, Germany). Threshold cycle55 (Ct) the fractional cycle number at which the amount of amplified target reached a fixed threshold, was determined and expression was measured using the 2−ΔCt formula, as previously reported56.

DNA extraction from human primary pancreatic cancer tumors and crExos

Immediately after resection, pancreatic tumor samples were snap-frozen in liquid nitrogen and stored at −80°C until further processing. A 10 μm reference section of each sample was cut and stained with hematoxylin and eosin by standard methods to evaluate the proportion of tumor tissue and adjacent tumor stroma. Samples with a tumor stroma proportion > 30 % were excluded into this study. DNA isolation was performed using a commercial DNA Extraction Kit (DNeasy Blood & Tissue Kit, Cat. No. 69506, Qiagen, Germany) according to the manufacturer’s protocol. The amount of DNA from tumor samples was quantified using a Nanodrop® 1000 Spectrophotometer (Thermo Fisher Scientific, Wilmington, DE 19810, USA).

Polymerase chain reaction (PCR) and Sanger Sequencing

PCR was performed in a 25 μl reaction tube consisting of 10 μl template DNA, 1 μM of each primer, 2.5mM dNTP, 2.5 10 x PCR Buffer, 25mM Mg solution, 0.5 μl H2O and 2.5 μl Taq Polymerase. Amplification was carried out in a T100 ThermoCycler (Bio-Rad) under following conditions: 94°C for 1 min, 2 cycles of 94 °C for 10 s, 67°C for 30 s, 70 °C for 30 s; 2 cycles of 94 °C for 10 s, 64°C for 30 s, 70 °C for 30 s; 2 cycles of 94 °C for 10 s, 61°C for 30 s, 70 °C for 30 s; 35 cycles of 94 °C for 10 s, 59°C for 30 s, 70 °C for 30 s; endless 4 °C. KRAS amplicon were generated using the following primers: forward 5′-AAGGCCTGCTGAAAATGACTG-3′, 5′-AGAATGGTCCTGCACCAGTAA-3′. PCR products were purified using the QIAquick PCR purification kit (Qiagen, Hilden, Germany). Subsequently, sequencing reaction was performed using BigDye terminator kit (v3.1, Life Technologies, USA) according to the manufacturer’s instructions. Sequencing products were separated on an ABI 3730 automated sequencer (Life Technologies, USA). KRAS mutation status was evaluated using Finch TV (Geospiza, Inc., Seattle, WA 98119, USA).

MRI imaging

MRI studies were conducted using a 7T small animal MR system, the Biospec USR70/30 (Bruker Biospin MRI, Billerica, MA) is based on an actively-shielded 7T magnet with a 30-cm bore and cryo-refrigeration. The system is equipped with 6 cm inner-diameter gradients that deliver a maximum gradient field of 950 mT/m. A 3.5 cm inner-diameter linear birdcage coil transmits and receives the MR signal. For image acquisition, T2 weighted, respiratory gated, multi-slice imaging will be performed with respiration held to under 25 breaths per minute to minimize motion artifacts in the abdomen. For mice where fat signal might mask the T2 weighted image the fat-suppression pulse module will be utilized. Acquisition parameters were minimally modified from Schmid A et. Al.57. The RARE-T2 weighted pulse sequence was modified to include an effective Te of 56 ms with a total TR of 2265 ms. Between 18 and 20 coronal slices were acquired per mouse with a slice thickness of 0.75 mm and slice spacing of 1 mm. In plane, pixel sizes of 0.156 mm × 0.156 mm with a matrix size of 256×192 (40 mm × 30 mm FOV) was chosen to minimize in plane partial volume effects, maintain a FOV sufficient to cover the abdomen, while also providing sufficient throughput for the experiment.

To measure tumor burden, the region of suspected lesions are drawn blinded on each slice after images intensities were normalized. The volume is calculated by addition of delineated region of interest in mm2 x 1 mm slice distance.

Statistical analysis

The GraphPad Prism version 6.0 (GraphPad Software, La Jolla, CA 92037 USA) and MedCalc statistical software version 13.0 (MedCalc Software bvba, Acacialaan 22, 8400 Ostend, Belgium) were used for all calculations. Unpaired Student’s t-test was applied to calculate expression differences of the qPCR results (dCt-values). Analysis of variance (ANOVA) tests were performed to calculate differences of multiple serum factors in murine and human serum samples. As a post-hoc test, a Tukey-Kramer test was applied for pairwise comparison of subgroups when the ANOVA test was positive in case of equal variance. Tamhane T2 test was applied for pairwise comparison of subgroups when the ANOVA test was positive in case of unequal variances. A paired two-tailed Student’s t-test was applied to assay differences in percent bead with GPC1+ crExos and CA 19-9 in the longitudinal cohort between pre-operative and postoperative blood samples. Receiver operating characteristic curves (ROC) were used to determine the sensitivity, specificity, positive and negative predictive value and to compare area under the curves (AUC) of serum factors using the Delong method58. The cut-off value was determined using the Youden-Index. Univariate analysis using the log-rank test was conducted to visualize (Kaplan Meier curves) and assess disease-specific survival (time from diagnosis to cancer-related death or last follow-up) in the longitudinal cohort of patients with pancreatic cancer. A multivariate analysis using the Cox proportional hazards regression model was performed to evaluate the effect of a decrease of percent bead with GPC1+ crExos in addition to age (continuous variable), AJCC tumor stage, and tumor grade (G) and CA 19-9 levels (U.ml−1). Correlation analysis between murine tumor burden and percent bead with GPC1+ crExos was performed using the Spearman correlation test. Figures were prepared by using GraphPad Prism (GraphPad Software, La Jolla, CA 92037 USA) and MedCalc statistical software version 13.0 (MedCalc Software bvba, Acacialaan 22, 8400 Ostend, Belgium). All presented P values are two-sided and P < 0.05 was considered to be statistically significant.

Extended Data

Extended Data Figure 1. Exosomes isolation.

Extended Data Figure 1

a, Exosomes concentration and size distribution by NanoSight® analysis of culture supernatant from NIH/3T3, MCF10A, HDF, MDA-MB-231 and E10 cells. Size mode: 105 nanometers (nm; 3 technical replicates). b, Transmission electron microscopy (TEM) micrograph of MDA-MB-231-derived exosomes. Upper right image shows a digitally zoomed inset. c, TEM micrograph of MDA-MB-231-derived exosomes following immunogold labeling for CD9. Gold particles are depicted as black dots. Upper right image shows a digitally zoomed inset. d, Immunoblot of flotillin1 and CD81 in exosomal proteins extracted from culture supernatant of E10, NIH/3T3, MDA-MB-231, MCF10A and HDF cells. e, RT–qPCR measurement of GPC1 mRNA levels in HMEL, HDF, HMLE, MCF7, MDA-MB-231, T3M4, Panc-1, MIA Paca2. Results are shown as mean ± standard deviation; n=3, 3 biological replicates, with 3 technical replicates each. f, Immunoblot of GPC1 in HMEL, HDF, HMLE, MCF7, MDA-MB-231, T3M4, Panc-1 and MIA Paca2 cell lysates (upper panel). β-actin was used as a loading control (lower panel). g, Immunoblot of GPC1 in exosomal protein lysates derived from the culture supernatant of 3 non-tumorigenic cell lines (HDF, HMEL, HMLE) and 5 tumorigenic cell lines (MCF7, MDA-MB-231, T3M4, Panc-1, MIA Paca2) (upper panel). Immunoblot of flotillin1 was used as loading control (lower panel). h, Immunoblot of flotillin1 in exosomal protein lysates from the culture supernatant of MDA-MB-231 and T3M4 following sucrose gradient purification. The protein content is assayed in each of the density layers listed.

Extended Data Figure 2. NanoSight® analysis in human serum samples.

Extended Data Figure 2

a, Transmission electron microscopy (TEM) micrograph of cancer patient serum-derived exosomes. Upper right image shows a digitally zoomed inset. b, TEM micrograph of cancer patient serum-derived exosomes following immunogold labeling for CD9. Gold particles are depicted as black dots. Upper right image shows a digitally zoomed inset. c, Immunoblot of flotillin1 of exosomal protein lysates from serum of cancer patient following exosomes purification by sucrose gradient. The protein content is assayed in each of the density layers listed. d, Exosomes concentration by NanoSight® analysis showing the number of exosomes per 1 ml of serum derived from healthy donors (n=100), from breast cancer patients (n=32) and from patients with a pancreatic ductal adenocarcinoma (PDAC, n=190). ANOVA, post-hoc Tamhane T2, * P<0.05, **** P<0.0001; 3 technical replicates. e, Exosomes size distribution by NanoSight® analysis showing the mode size of exosomes in 1 ml of serum derived from healthy donors (n=100), from breast cancer patients (n=32) and from patients with a pancreatic ductal adenocarcinoma (PDAC, n=190). ANOVA, post-hoc Tukey-Kramer test, *** P<0.001; 3 technical replicates. f, Scatter dot plots depicting the percentage of beads with GPC1+ bound exosomes purified from the serum of breast cancer patients. The patients are subdivided into three subtypes: Luminal A, Luminal B and Triple Negative breast cancer. g, Scatter plots depicting the serum CA 19-9 concentration (U/mL), evaluated by ELISA, in healthy donors (n=100), patients with benign pancreas disease (BPD, n=26), pancreas cancer precursor lesion (PCPL, n=5) and pancreatic ductal adenocarcinoma (PDAC, n=190). Discovery cohort, ANOVA, post-hoc Tamhane T2 * P<0.05; **** P<0.0001; 3 technical replicates. Data is presented as the mean ± standard deviation.

Extended Data Figure 3. Tumor stage-specific analysis.

Extended Data Figure 3

a, Table associated with receiver Operating Characteristic (ROC) curve analysis depicted in Figure 1f. b–f, ROC curve analysis for percent GPC1+ crExos (red line), CA 19-9 serum levels (blue scattered line), exosomes concentration (black line) and exosomes size (scattered black line) in patients with carcinoma in situ (CIS) or stage I pancreatic cancer (n=5) (a), stage IIa pancreatic cancer (n=18) (b), stage IIb pancreatic cancer (n=117) (c), stage III pancreatic cancer (n=11) (d), and stage IV pancreas cancer (n=41) (e), compared to control (healthy donors (n=100) and patients with a benign pancreatic disease (n=26), total n=126). g, Table associated with ROC curve analysis depicted in Figure 1h. AUC: Area under the curve, CI: confidence interval.

Extended Data Figure 4. Longitudinal human study.

Extended Data Figure 4

a, Scatter plots of percent beads with GPC1+ crExos by flow cytometry in patients with pancreas cancer. Patients are divided based on metastatic disease (non-metastatic lesions, lymph node metastases and distant metastases). ANOVA, post-hoc Tukey-Kramer test, * P<0.05; 3 technical replicates. b, Scatter plots depicting serum CA 19-9 levels (U/ml) in patients with benign pancreas disease (BPD) (n=4), cancer precursor lesion (PCPL) (n=4), and pancreatic ductal adenocarcinoma (PDAC) (n=29) on the preoperative day and postoperative day 7 in patients. Paired two-tailed Student’s t-test, ** P<0.01; 3 technical replicates. c–d, Multivariate analysis (Cox proportional hazards regression model) of prognostic parameters for overall (c) and disease-specific (d) survival of patients with pancreas cancer in the longitudinal cohort (n=29). e, Scatter plots depicting serum GPC1 (ng/ml) levels by ELISA in patients with benign pancreas disease (BPD, n=6), pancreatic ductal adenocarcinoma (PDAC, n=56) and healthy controls (n=20). ANOVA, post-hoc Tukey-Kramer test, **** P<0.0001; 3 technical replicates. f, ROC curve for circulating GPC1 protein (red line) in patients with pancreas cancer (n=56) vs. control (healthy donors (n=20) and patients with a benign pancreatic disease (n=26), total n=6). AUC: Area under the curve, CI: confidence interval.

Extended Data Figure 5. PDAC GEMM longitudinal study.

Extended Data Figure 5

a, Schematic diagram depicting the spontaneous development and progression of pancreatic cancer in Ptf1acre/+;LSL-KrasG12D/+;Tgfbr2L/L (PKT) mice and H&E of the pancreas at the indicated time points showing healthy pancreas, PanIN lesions, and PDAC lesions. Scale bars: 100 μm. b–c, Exosomes size (b) and concentration (c) assayed by NanoSight® analysis from the serum of PKT mice (E: experimental, red) and control mice (C: control, blue) at 4, 5, 6,7 and 8 weeks of age. ANOVA, post-hoc Tukey-Kramer test, * P<0.05; 3 technical replicates. d, Graph depicting the time wise progression of tumor volume measured by MRI and the % beads with GPC1+ bound crExos in individual PKT mice (blue: tumor volume, red: % GPC1+ crExos). e, Percent beads with GPC1+ crExos on beads from control mice (n=3) and mice with cerulein-induced acute pancreatitis (n=4). Two-tailed Student’s t-test, ns: not significant; 3 technical replicates. f, Results from ROC curves for percent beads with GPC1+ bound crExos, exosomes concentration and exosomes size in 4, 5, 6 and 7 weeks old PKT mice (n=7) vs. control (including age-matched littermate healthy control (n=6) and mice with induced acute pancreatitis (n=4), n=10)). Data is presented as the mean ± standard deviation.

Extended Data Figure 6. PDAC GEMM cross sectional studies.

Extended Data Figure 6

a, Representative micrographs of H&E stained pancreas from 16 days old control mice (left panel) and PKT mice presenting with (right panel, encircled) and without (middle panel) PanIN lesions. Scale bars: 100 μm. b, Ct values following qPCR analyses for oncogenic KRASG12D, wild-type KRAS and 18S internal control RNA from exosomes of 44–48 days old PKT mice serum segregated using FACS for GPC1+ bead bound exos (red) and GPC1 bead bound exos (blue). Data is presented as the mean ± standard deviation.

Extended Data Figure 7. Raw scatter dot plot depicting flow cytometry analyses of beads with GPC1+ bound exosomes.

Extended Data Figure 7

a, Scatter plots and histogram of flow cytometry analyses of serum exosomes on beads of a representative healthy control (left panels are secondary antibody only; right panels are GPC1 antibody and secondary antibody). b, Scatter plots and histogram of flow cytometry analysis of serum exosomes on beads of a representative pancreas cancer sample (left panels are secondary antibody only; right panels are with GPC1 antibody and secondary antibody).

Extended Data Figure 8.

Extended Data Figure 8

Uncropped Western blots 

Extended Data Table 1. List of the 48 proteins exclusive to MDA-MB-231 exosomes.

Listing of the 48 proteins exclusively detected in exosomes from MDA-MB-231 cells determined by ultra performance liquid chromatography – mass spectrometry (UPLC-MS) and comparative analyses of exosomes derived from NIH/3T3, MCF 10A, HDF, E10 and MDA-MB-231 cells. The proteins are grouped based on cellular location.

Protein Name Gene ID Cellular Location
ATP-binding cassette sub-family A member 6 ABCA6 Transmembrane
Tetraspanin-4 TSPAN4 Transmembrane
SLIT and NTRK-like protein 4 SLITRK4 Transmembrane
Putative protocadherin beta-18 PCDHB18 Transmembrane
Myeloid cell surface antigen CD33 CD33 Transmembrane
Glypican-1 GPC1 Membrane anchored
Protein Name Gene ID Cellular Location
Histone H2A type 2-A HIST1H2AA Nucleus
Histone H2A type 1-A HIST1H1AA Nucleus
Histone H3.3 H3F3A Nucleus
Histone H3.1 HIST1H3A Nucleus
Zinc finger protein 37 homolog ZFP37 Nucleus
Hypermethylated in cancer 2 protein HIC2 Nucleus
Zinc finger protein 12 ZSCAN12 Nucleus
Protein Name Gene ID Cellular Location
Laminin subunit beta-1 LAMB1 Secreted
Tubulointerstitial nephritis antigen-like TINAGL1 Secreted
Peroxiredoxin-4 PRDX4 Secreted
Collagen alpha-2(IV) chain COL4A2 Secreted
Putative protein C3P1 C3P1 Secreted
Collagen alpha-1(II) chain COL2A1 Secreted
Hemicentin-1 HMCN1 Secreted
Protein Name Gene ID Cellular Location
Putative rhophilin-2-like protein RHPN2P1 Not specified
Ankyrin repeat domain-containing protein 62 ANKRD62 Not specified
Tripartite motif-containing protein 42 TRIM42 Not specified !
Protein Name Gene ID Cellular Location
Junction plakoglobin JUP Cytoplasm
Tubulin beta-2B chain TUBB2B Cytoplasm
Endoribonuclease Dicer DICER1 Cytoplasm
E3 ubiquitin-protein ligase TRIM71 TRIM71 Cytoplasm
Katanin p60 ATPase-containing subunit A-like 2 KATNAL2 Cytoplasm
Protein S100-A6 S100A6 Cytoplasm
5′-nucleotidase domain-containing protein 3 NT5DC3 Cytoplasm
Valine--tRNA ligase VARS Cytoplasm
Kazrin KAZN Cytoplasm
ELAV-like protein 4 ELAVL4 Cytoplasm
RING finger protein 166 RNF166 Cytoplasm
FERM and PDZ domain-containing protein 1 FRMPD1 Cytoplasm
78 kDa glucose-regulated protein HSPA5 Cytoplasm
Trafficking protein particle complex subunit 6A TRAPPC6A Cytoplasm
Squalene monooxygenase SQLE Cytoplasm
Tumor susceptibility gene 101 protein TSG101 Cytoplasm
Vacuolar protein sorting 28 homolog VPS28 Cytoplasm
Prostaglandin F2 receptor negative regulator PTGFRN Cytoplasm
Isobutyryl-CoA dehydrogenase, mitochondrial ACAD8 Cytoplasm
26S protease regulatory subunit 6B PSMC4 Cytoplasm
Elongation factor 1-gamma EEF1G Cytoplasm
Titin TTN Cytoplasm
Tyrosine-protein phosphatase type 13 PTPN13 Cytoplasm
Triosephosphate isomerase TPI1 Cytoplasm
Carboxypeptidase E CPE Cytoplasm

Extended Data Table 2.

Exosomes proteins assayed by ultra performance liquid chromatography – mass spectrometry (UPLC-MS) in indicated cell lines.

E10
prot_hit prot_acc prot_desc prot_score prot_mass prot_matches prot_sequences_sig prot_cover prot_pi
1 P04264 Keratin, type II cytoskeletal 1 OS=Homo sapiens GN=KRT1 PE=1 SV=6 2721 65999 68 30 57.8 8.15
2 P35527 Keratin, type I cytoskeletal 9 OS=Homo sapiens GN=KRT9 PE=1 SV=3 2381 62027 59 24 68.2 5.14
3 P13645 Keratin, type I cytoskeletal 10 OS=Homo sapiens GN=KRT10 PE=1 SV=6 2146 58792 66 29 57.9 5.13
4 P02768 Serum albumin OS=Homo sapiens GN=ALB PE=1 SV=2 2032 69321 104 9 13.5 5.92
5 P07996 Thrombospondin-1 OS=Homo sapiens GN=THBS1 PE=1 SV=2 1823 129300 57 29 37.9 4.71
6 P02765 Alpha-2-HS-glycoprotein OS=Homo sapiens GN=AHSG PE=1 SV=1 1784 39300 50 3 8.7 5.43
7 P01023 Alpha-2-macroglobulin OS=Homo sapiens GN=A2M PE=1 SV=3 1719 163188 62 8 7.5 6.03
8 P20742 Pregnancy zone protein OS=Homo sapiens GN=PZP PE=1 SV=4 1562 163760 48 4 2.5 5.97
9 P02751 Fibronectin OS=Homo sapiens GN=FN1 PE=1 SV=4 1186 262460 37 26 17.6 5.46
10 P12111 Collagen alpha-3(VI) chain OS=Homo sapiens GN=COL6A3 PE=1 SV=5 1113 343457 51 39 20.2 6.26
11 P35908 Keratin, type II cytoskeletal 2 epidermal OS=Homo sapiens GN=KRT2 PE=1 SV=2 1057 65393 35 21 48.4 8.07
12 P01024 Complement C3 OS=Homo sapiens GN=C3 PE=1 SV=2 978 187030 43 18 14.7 6.02
13 P00734 Prothrombin OS=Homo sapiens GN=F2 PE=1 SV=2 846 69992 21 6 8 5.64
14 P60709 Actin, cytoplasmic 1 OS=Homo sapiens GN=ACTB PE=1 SV=1 820 41710 23 12 47.5 5.29
15 P08779 Keratin, type I cytoskeletal 16 OS=Homo sapiens GN=KRT16 PE=1 SV=4 762 51236 31 18 48 4.99
16 P19823 Inter-alpha-trypsin inhibitor heavy chain H2 OS=Homo sapiens GN=ITIH2 PE=1 SV=2 708 106397 27 8 9.3 6.4
17 P02533 Keratin, type I cytoskeletal 14 OS=Homo sapiens GN=KRT14 PE=1 SV=4 579 51529 25 15 37.1 5.09
18 P0C0L4 Complement C4-A OS=Homo sapiens GN=C4A PE=1 SV=1 492 192650 20 9 5.6 6.65
19 P04259 Keratin, type II cytoskeletal 6B OS=Homo sapiens GN=KRT6B PE=1 SV=5 446 60030 18 10 25.5 8.09
20 P12109 Collagen alpha-1(VI) chain OS=Homo sapiens GN=COL6A1 PE=1 SV=3 420 108462 11 7 11.3 5.26
21 P04406 Glyceraldehyde-3-phosphate dehydrogenase OS=Homo sapiens GN=GAPDH PE=1 SV=3 419 36030 12 6 28.4 8.57
22 Q07954 Prolow-density lipoprotein receptor-related protein 1 OS=Homo sapiens GN=LRP1 PE=1 SV=2 419 504276 22 10 5.8 5.16
23 P02538 Keratin, type II cytoskeletal 6A OS=Homo sapiens GN=KRT6A PE=1 SV=3 419 60008 18 10 27.1 8.09
24 P02788 Lactotransferrin OS=Homo sapiens GN=LTF PE=1 SV=6 416 78132 16 3 4.8 8.5
25 P68032 Actin, alpha cardiac muscle 1 OS=Homo sapiens GN=ACTC1 PE=1 SV=1 408 41992 13 6 26.8 5.23
26 P02771 Alpha-fetoprotein OS=Homo sapiens GN=AFP PE=1 SV=1 395 68633 9 3 5.6 5.48
27 Q08380 Galectin-3-binding protein OS=Homo sapiens GN=LGALS3BP PE=1 SV=1 392 65289 15 10 26.7 5.13
28 P68363 Tubulin alpha-1B chain OS=Homo sapiens GN=TUBA1B PE=1 SV=1 364 50120 18 9 38.4 4.94
29 P13647 Keratin, type II cytoskeletal 5 OS=Homo sapiens GN=KRT5 PE=1 SV=3 360 62340 17 11 23.4 7.59
30 Q04695 Keratin, type I cytoskeletal 17 OS=Homo sapiens GN=KRT17 PE=1 SV=2 357 48076 20 10 32.6 4.97
31 Q9BYX7 Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 347 41989 8 3 13.6 5.91
32 P06396 Gelsolin OS=Homo sapiens GN=GSN PE=1 SV=1 340 85644 11 7 12.5 5.9
33 P04114 Apolipoprotein B-100 OS=Homo sapiens GN=APOB PE=1 SV=2 327 515283 16 12 2.4 6.58
34 P03956 Interstitial collagenase OS=Homo sapiens GN=MMP1 PE=1 SV=3 318 53973 15 11 33.5 6.47
35 Q06033 Inter-alpha-trypsin inhibitor heavy chain H3 OS=Homo sapiens GN=ITIH3 PE=1 SV=2 298 99787 10 5 5.1 5.49
36 P69905 Hemoglobin subunit alpha OS=Homo sapiens GN=HBA1 PE=1 SV=2 297 15248 13 3 33.1 8.72
37 P24821 Tenascin OS=Homo sapiens GN=TNC PE=1 SV=3 292 240700 12 10 9.4 4.79
38 P07437 Tubulin beta chain OS=Homo sapiens GN=TUBB PE=1 SV=2 286 49639 12 7 34.2 4.78
39 P68371 Tubulin beta-2C chain OS=Homo sapiens GN=TUBB2C PE=1 SV=1 278 49799 10 7 27.4 4.79
40 P02774 Vitamin D-binding protein OS=Homo sapiens GN=GC PE=1 SV=1 273 52929 12 6 12 5.4
41 Q562R1 Beta-actin-like protein 2 OS=Homo sapiens GN=ACTBL2 PE=1 SV=2 271 41976 8 3 16 5.39
42 P23142 Fibulin-1 OS=Homo sapiens GN=FBLN1 PE=1 SV=4 269 77162 10 5 7.4 5.07
43 P13646 Keratin, type I cytoskeletal 13 OS=Homo sapiens GN=KRT13 PE=1 SV=4 256 49557 13 6 10.9 4.91
44 P02647 Apolipoprotein A–I OS=Homo sapiens GN=APOA1 PE=1 SV=1 255 30759 5 1 6 5.56
45 P14618 Pyruvate kinase isozymes M1/M2 OS=Homo sapiens GN=PKM2 PE=1 SV=4 228 57900 9 6 20.3 7.96
46 P04004 Vitronectin OS=Homo sapiens GN=VTN PE=1 SV=1 215 54271 6 1 4.2 5.55
47 P19012 Keratin, type I cytoskeletal 15 OS=Homo sapiens GN=KRT15 PE=1 SV=3 210 49181 14 6 10.3 4.71
48 Q15582 Transforming growth factor-beta-induced protein ig-h3 OS=Homo sapiens GN=TGFBI PE=1 SV=1 196 74634 8 7 16.1 7.62
49 P36955 Pigment epithelium-derived factor OS=Homo sapiens GN=SERPINF1 PE=1 SV=4 188 46283 7 3 12 5.97
50 Q99715 Collagen alpha-1(XII) chain OS=Homo sapiens GN=COL12A1 PE=1 SV=2 176 332941 8 5 4.9 5.38
51 P07900 Heat shock protein HSP 90-alpha OS=Homo sapiens GN=HSP90AA1 PE=1 SV=5 175 84607 8 4 12.8 4.94
52 P02675 Fibrinogen beta chain OS=Homo sapiens GN=FGB PE=1 SV=2 174 55892 7 4 6.7 8.54
53 P02042 Hemoglobin subunit delta OS=Homo sapiens GN=HBD PE=1 SV=2 171 16045 9 2 12.9 7.85
54 P11717 Cation-independent mannose-6-phosphate receptor OS=Homo sapiens GN=IGF2R PE=1 SV=3 163 274199 5 2 0.8 5.64
55 P51884 Lumican OS=Homo sapiens GN=LUM PE=1 SV=2 161 38405 9 3 12.4 6.16
56 P08697 Alpha-2-antiplasmin OS=Homo sapiens GN=SERPINF2 PE=1 SV=3 158 54531 2 1 2.2 5.87
57 P12259 Coagulation factor V OS=Homo sapiens GN=F5 PE=1 SV=4 153 251546 9 6 3.5 5.68
58 P08107 Heat shock 70 kDa protein 1A/1B OS=Homo sapiens GN=HSPA1A PE=1 SV=5 147 70009 3 3 6.2 5.48
59 P17066 Heat shock 70 kDa protein 6 OS=Homo sapiens GN=HSPA6 PE=1 SV=2 147 70984 3 3 6.2 5.81
60 P19013 Keratin, type II cytoskeletal 4 OS=Homo sapiens GN=KRT4 PE=1 SV=4 146 57250 6 4 7.3 6.25
61 Q8TAA3 Proteasome subunit alpha type-7-like OS=Homo sapiens GN=PSMA8 PE=1 SV=3 144 28512 4 3 19.5 9.07
62 Q5XKE5 Keratin, type II cytoskeletal 79 OS=Homo sapiens GN=KRT79 PE=1 SV=2 138 57800 7 3 6.9 6.75
63 P0C0S8 Histone H2A type 1 OS=Homo sapiens GN=HIST1H2AG PE=1 SV=2 136 14083 3 2 21.5 10.9
64 Q99816 Tumor susceptibility gene 101 protein OS=Homo sapiens GN=TSG101 PE=1 SV=2 135 43916 4 2 21.8 8.2
65 P23526 Adenosylhomocysteinase OS=Homo sapiens GN=AHCY PE=1 SV=4 134 47685 3 2 9.5 5.92
66 P00747 Plasminogen OS=Homo sapiens GN=PLG PE=1 SV=2 132 90510 7 4 5.3 7.04
67 O95678 Keratin, type II cytoskeletal 75 OS=Homo sapiens GN=KRT75 PE=1 SV=2 132 59524 7 3 7.8 7.6
68 P13639 Elongation factor 2 OS=Homo sapiens GN=EEF2 PE=1 SV=4 132 95277 6 4 10.3 6.41
69 Q9UBG0 C-type mannose receptor 2 OS=Homo sapiens GN=MRC2 PE=1 SV=2 128 166568 2 2 1.6 5.54
70 Q15063 Periostin OS=Homo sapiens GN=POSTN PE=1 SV=2 127 93255 6 3 10.3 7.27
71 P62805 Histone H4 OS=Homo sapiens GN=HIST1H4A PE=1 SV=2 127 11360 5 4 38.8 11.36
72 P01008 Antithrombin-III OS=Homo sapiens GN=SERPINC1 PE=1 SV=1 126 52569 13 3 14.9 6.32
73 P19827 Inter-alpha-trypsin inhibitor heavy chain H1 OS=Homo sapiens GN=ITIH1 PE=1 SV=3 124 101326 4 3 2.9 6.31
74 P11142 Heat shock cognate 71 kDa protein OS=Homo sapiens GN=HSPA8 PE=1 SV=1 123 70854 4 4 7.6 5.37
75 P08729 Keratin, type II cytoskeletal 7 OS=Homo sapiens GN=KRT7 PE=1 SV=5 123 51354 4 2 5.1 5.4
76 P02794 Ferritin heavy chain OS=Homo sapiens GN=FTH1 PE=1 SV=2 122 21212 4 3 16.4 5.3
77 P05787 Keratin, type II cytoskeletal 8 OS=Homo sapiens GN=KRT8 PE=1 SV=7 122 53671 4 2 5.6 5.52
78 P01031 Complement C5 OS=Homo sapiens GN=C5 PE=1 SV=4 122 188186 5 3 2.6 6.11
79 Q5VTE0 Putative elongation factor 1-alpha-like 3 OS=Homo sapiens GN=EEF1A1P5 PE=5 SV=1 121 50153 4 3 14.1 9.15
80 P05452 Tetranectin OS=Homo sapiens GN=CLEC3B PE=1 SV=3 121 22522 5 3 28.7 5.52
81 P49327 Fatty acid synthase OS=Homo sapiens GN=FASN PE=1 SV=3 119 273254 5 3 3.3 6.01
82 P07195 L-lactate dehydrogenase B chain OS=Homo sapiens GN=LDHB PE=1 SV=2 118 36615 2 2 6.9 5.71
83 Q9H4B7 Tubulin beta-1 chain OS=Homo sapiens GN=TUBB1 PE=1 SV=1 112 50295 5 3 11.8 5.05
84 P15531 Nucleoside diphosphate kinase A OS=Homo sapiens GN=NME1 PE=1 SV=1 111 17138 1 1 11.2 5.83
85 Q7Z3Y8 Keratin, type I cytoskeletal 27 OS=Homo sapiens GN=KRT27 PE=1 SV=2 109 49792 7 2 7.6 4.98
86 P08238 Heat shock protein HSP 90-beta OS=Homo sapiens GN=HSP90AB1 PE=1 SV=4 109 83212 4 2 6.2 4.97
87 P08123 Collagen alpha-2(I) chain OS=Homo sapiens GN=COL1A2 PE=1 SV=7 105 129235 2 2 2.3 9.08
88 Q86YZ3 Hornerin OS=Homo sapiens GN=HRNR PE=1 SV=2 101 282228 1 1 0.6 10.05
89 P02452 Collagen alpha-1(I) chain OS=Homo sapiens GN=COL1A1 PE=1 SV=5 99 138857 4 2 3 5.6
90 P04075 Fructose-bisphosphate aldolase A OS=Homo sapiens GN=ALDOA PE=1 SV=2 98 39395 2 2 6.9 8.3
91 P02787 Serotransferrin OS=Homo sapiens GN=TF PE=1 SV=3 97 77014 4 2 1.7 6.81
92 P02748 Complement component C9 OS=Homo sapiens GN=C9 PE=1 SV=2 95 63133 3 2 3.8 5.43
93 Q8TEV9_REVERSED Smith-Magenis syndrome chromosomal region candidate gene 8 protein OS=Homo sapiens GN=SMCR8 PE=1 SV=2 - REVERSED 91 104956 12 1 1.2 5.36
94 P02461 Collagen alpha-1(III) chain OS=Homo sapiens GN=COL3A1 PE=1 SV=4 90 138479 2 2 1.6 6.21
95 P00450 Ceruloplasmin OS=Homo sapiens GN=CP PE=1 SV=1 88 122128 3 3 2.9 5.44
96 P62979 Ubiquitin-40S ribosomal protein S27a OS=Homo sapiens GN=RPS27A PE=1 SV=2 88 17953 4 3 19.9 9.68
97 Q14697 Neutral alpha-glucosidase AB OS=Homo sapiens GN=GANAB PE=1 SV=3 84 106807 2 2 3.9 5.74
98 Q08431 Lactadherin OS=Homo sapiens GN=MFGE8 PE=1 SV=2 83 43095 2 1 3.9 8.47
99 A6NMY6 Putative annexin A2-like protein OS=Homo sapiens GN=ANXA2P2 PE=5 SV=2 82 38635 2 2 6.2 6.49
100 P07355 Annexin A2 OS=Homo sapiens GN=ANXA2 PE=1 SV=2 82 38580 3 2 10.3 7.57
101 Q9BYX7_REVERSED Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 - REVERSED 80 41989 4 1 3.5 5.92
102 Q8NI35 InaD-like protein OS=Homo sapiens GN=INADL PE=1 SV=3 79 196247 8 1 0.3 4.84
103 P15169 Carboxypeptidase N catalytic chain OS=Homo sapiens GN=CPN1 PE=1 SV=1 79 52253 1 1 3.3 6.86
104 P28074 Proteasome subunit beta type-5 OS=Homo sapiens GN=PSMB5 PE=1 SV=3 79 28462 2 2 8.7 6.43
105 P49720 Proteasome subunit beta type-3 OS=Homo sapiens GN=PSMB3 PE=1 SV=2 78 22933 3 2 16.6 6.14
106 Q04756 Hepatocyte growth factor activator OS=Homo sapiens GN=HGFAC PE=1 SV=1 77 70636 1 1 2.4 6.99
107 Q12931 Heat shock protein 75 kDa, mitochondrial OS=Homo sapiens GN=TRAP1 PE=1 SV=3 76 80060 1 1 2 8.3
108 P05121 Plasminogen activator inhibitor 1 OS=Homo sapiens GN=SERPINE1 PE=1 SV=1 76 45031 4 2 7 6.68
109 P20618 Proteasome subunit beta type-1 OS=Homo sapiens GN=PSMB1 PE=1 SV=2 74 26472 2 2 13.3 8.27
110 Q14624 Inter-alpha-trypsin inhibitor heavy chain H4 OS=Homo sapiens GN=ITIH4 PE=1 SV=4 73 103293 4 1 1.9 6.51
111 Q5D862 Filaggrin-2 OS=Homo sapiens GN=FLG2 PE=1 SV=1 72 247928 1 1 0.5 8.45
112 P06733 Alpha-enolase OS=Homo sapiens GN=ENO1 PE=1 SV=2 71 47139 3 2 10.1 7.01
113 P02790 Hemopexin OS=Homo sapiens GN=HPX PE=1 SV=2 70 51643 7 1 5.4 6.55
114 P35442 Thrombospondin-2 OS=Homo sapiens GN=THBS2 PE=1 SV=2 70 129908 4 2 3.2 4.62
115 P13929 Beta-enolase OS=Homo sapiens GN=ENO3 PE=1 SV=4 69 46902 2 2 7.6 7.59
116 Q8WW52_REVERSED Protein FAM151A OS=Homo sapiens GN=FAM151A PE=2 SV=2 - REVERSED 69 63987 2 1 1.4 6.2
117 P12110 Collagen alpha-2(VI) chain OS=Homo sapiens GN=COL6A2 PE=1 SV=4 69 108512 4 2 3.4 5.85
118 P04083 Annexin A1 OS=Homo sapiens GN=ANXA1 PE=1 SV=2 68 38690 1 1 3.8 6.57
119 P01009 Alpha-1-antitrypsin OS=Homo sapiens GN=SERPINA1 PE=1 SV=3 68 46707 4 1 3.3 5.37
120 P22303 Acetylcholinesterase OS=Homo sapiens GN=ACHE PE=1 SV=1 67 67753 1 1 2.1 5.87
121 P55058 Phospholipid transfer protein OS=Homo sapiens GN=PLTP PE=1 SV=1 66 54705 1 1 2.2 6.53
122 P00488 Coagulation factor XIII A chain OS=Homo sapiens GN=F13A1 PE=1 SV=4 66 83215 1 1 2.2 5.75
123 Q03692 Collagen alpha-1(X) chain OS=Homo sapiens GN=COL10A1 PE=1 SV=2 66 66117 1 1 1.9 9.68
124 P05546 Heparin cofactor 2 OS=Homo sapiens GN=SERPIND1 PE=1 SV=3 65 57034 2 1 2.4 6.41
125 P01266 Thyroglobulin OS=Homo sapiens GN=TG PE=1 SV=5 65 304594 3 3 0.8 5.4
126 P00338 L-lactate dehydrogenase A chain OS=Homo sapiens GN=LDHA PE=1 SV=2 63 36665 2 1 8.1 8.44
127 O15111_REVERSED Inhibitor of nuclear factor kappa-B kinase subunit alpha OS=Homo sapiens GN=CHUK PE=1 SV=2 -REVERSED 63 84585 7 1 1.6 6.28
128 Q8NAV2_REVERSED Uncharacterized protein C8orf58 OS=Homo sapiens GN=C8orf58 PE=2 SV=2 - REVERSED 63 39636 3 1 4.1 8.54
129 P28070 Proteasome subunit beta type-4 OS=Homo sapiens GN=PSMB4 PE=1 SV=4 62 29185 2 2 7.2 5.72
130 P07686 Beta-hexosaminidase subunit beta OS=Homo sapiens GN=HEXB PE=1 SV=3 61 63071 1 1 2.9 6.29
131 P05543 Thyroxine-binding globulin OS=Homo sapiens GN=SERPINA7 PE=1 SV=2 61 46295 2 1 2.4 5.87
132 Q9HDC9 Adipocyte plasma membrane-associated protein OS=Homo sapiens GN=APMAP PE=1 SV=2 60 46451 3 1 1.7 5.82
133 P25787 Proteasome subunit alpha type-2 OS=Homo sapiens GN=PSMA2 PE=1 SV=2 60 25882 1 1 6 6.92
134 P35125 Ubiquitin carboxyl-terminal hydrolase 6 OS=Homo sapiens GN=USP6 PE=1 SV=2 60 158557 12 1 0.4 7.87
135 A6NIZ1 Ras-related protein Rap-1b-like protein OS=Homo sapiens PE=2 SV=1 59 20912 1 1 6.5 5.37
136 B9A064 Immunoglobulin lambda-like polypeptide 5 OS=Homo sapiens GN=IGLL5 PE=2 SV=2 59 23049 2 1 3.7 9.08
137 Q99456 Keratin, type I cytoskeletal 12 OS=Homo sapiens GN=KRT12 PE=1 SV=1 58 53478 6 2 3.6 4.7
138 Q86Y46 Keratin, type II cytoskeletal 73 OS=Homo sapiens GN=KRT73 PE=1 SV=1 58 58887 5 4 6.7 6.93
139 P20774 Mimecan OS=Homo sapiens GN=OGN PE=1 SV=1 58 33901 2 2 9.7 5.46
140 Q9Y240 C-type lectin domain family 11 member A OS=Homo sapiens GN=CLEC11A PE=1 SV=1 57 35672 1 1 5.3 5.06
141 P10909 Clusterin OS=Homo sapiens GN=CLU PE=1 SV=1 57 52461 2 1 7.8 5.89
142 Q9NPY3_REVERSED Complement component C1q receptor OS=Homo sapiens GN=CD93 PE=1 SV=3 - REVERSED 57 68515 2 1 1.1 5.27
143 P81605 Dermcidin OS=Homo sapiens GN=DCD PE=1 SV=2 56 11277 2 2 22.7 6.08
144 P03973 Antileukoproteinase OS=Homo sapiens GN=SLPI PE=1 SV=2 55 14316 1 1 9.1 9.11
145 Q6YHK3 CD109 antigen OS=Homo sapiens GN=CD109 PE=1 SV=2 55 161587 2 2 1.8 5.59
146 P04180 Phosphatidylcholine-sterol acyltransferase OS=Homo sapiens GN=LCAT PE=1 SV=1 55 49546 1 1 2.5 5.71
147 Q9BXJ4 Complement C1q tumor necrosis factor-related protein 3 OS=Homo sapiens GN=C1QTNF3 PE=2 SV=1 55 26977 1 1 5.3 6.04
148 P16070 CD44 antigen OS=Homo sapiens GN=CD44 PE=1 SV=3 53 81487 1 1 1.6 5.13
149 P02649 Apolipoprotein E OS=Homo sapiens GN=APOE PE=1 SV=1 53 36132 1 1 2.8 5.65
150 P29353_REVERSED SHC-transforming protein 1 OS=Homo sapiens GN=SHC1 PE=1 SV=4 - REVERSED 51 62782 6 1 1.4 6.01
151 P35580 Myosin-10 OS=Homo sapiens GN=MYH10 PE=1 SV=3 51 228858 2 2 1.2 5.44
152 P28066 Proteasome subunit alpha type-5 OS=Homo sapiens GN=PSMA5 PE=1 SV=3 50 26394 1 1 4.1 4.74
153 O95445 Apolipoprotein M OS=Homo sapiens GN=APOM PE=1 SV=2 49 21239 4 1 6.9 5.66
154 P04745 Alpha-amylase 1 OS=Homo sapiens GN=AMY1A PE=1 SV=2 49 57731 1 1 2.3 6.47
155 Q13454 Tumor suppressor candidate 3 OS=Homo sapiens GN=TUSC3 PE=2 SV=1 49 39650 1 1 2 9.93
156 P49721 Proteasome subunit beta type-2 OS=Homo sapiens GN=PSMB2 PE=1 SV=1 49 22822 3 1 19.9 6.51
157 P32119 Peroxiredoxin-2 OS=Homo sapiens GN=PRDX2 PE=1 SV=5 49 21878 1 1 9.1 5.66
158 Q6ZU45 Putative C-type lectin domain-containing protein NCRNA00083 OS=Homo sapiens GN=NCRNA00083 PE=5 SV=1 49 25796 1 1 3.4 6.3
159 O43264_REVERSED Centromere/kinetochore protein zw10 homolog OS=Homo sapiens GN=ZW10 PE=1 SV=3 - REVERSED 49 88773 1 1 0.8 5.89
160 Q9Y490 Talin-1 OS=Homo sapiens GN=TLN1 PE=1 SV=3 48 269599 2 2 1 5.77
161 Q96CD0_REVERSED F-box/LRR-repeat protein 8 OS=Homo sapiens GN=FBXL8 PE=1 SV=1 - REVERSED 48 40490 11 1 1.9 7.03
162 P06702 Protein S100-A9 OS=Homo sapiens GN=S100A9 PE=1 SV=1 47 13234 1 1 13.2 5.71
163 Q6P1X5_REVERSED Transcription initiation factor TFIID subunit 2 OS=Homo sapiens GN=TAF2 PE=1 SV=3 - REVERSED 47 136883 1 1 0.5 8.46
164 Q16853 Membrane primary amine oxidase OS=Homo sapiens GN=AOC3 PE=1 SV=3 47 84568 1 1 1.2 6.05
165 P00558 Phosphoglycerate kinase 1 OS=Homo sapiens GN=PGK1 PE=1 SV=3 47 44586 3 1 8.4 8.3
166 P02749 Beta-2-glycoprotein 1 OS=Homo sapiens GN=APOH PE=1 SV=3 46 38273 1 1 2.6 8.34
167 A8MPT4 Glutathione S-transferase theta-4 OS=Homo sapiens GN=GSTT4 PE=3 SV=2 45 27941 2 1 2.5 6.25
168 Q8N6V9_REVERSED Testis-expressed sequence 9 protein OS=Homo sapiens GN=TEX9 PE=2 SV=1 - REVERSED 45 44798 2 1 1.5 6.2
169 Q9Y2U8_REVERSED Inner nuclear membrane protein Man1 OS=Homo sapiens GN=LEMD3 PE=1 SV=2 - REVERSED 45 99935 2 1 0.7 7.31
170 P05154 Plasma serine protease inhibitor OS=Homo sapiens GN=SERPINA5 PE=1 SV=3 45 45646 2 1 1.5 9.3
171 Q8NDM7_REVERSED WD repeat-containing protein 96 OS=Homo sapiens GN=WDR96 PE=2 SV=3 - REVERSED 45 191861 1 1 0.5 5.71
172 Q6NXT2 Histone H3.3C OS=Homo sapiens GN=H3F3C PE=1 SV=3 44 15204 1 1 5.2 11.1
173 O95248 Myotubularin-related protein 5 OS=Homo sapiens GN=SBF1 PE=1 SV=3 44 208184 1 1 0.4 6.46
174 Q9UK61_REVERSED Uncharacterized protein C3orf63 OS=Homo sapiens GN=C3orf63 PE=1 SV=3 - REVERSED 44 188914 1 1 0.4 5.55
175 Q9Y6X9_REVERSED MORC family CW-type zinc finger protein 2 OS=Homo sapiens GN=MORC2 PE=1 SV=2 - REVERSED 43 117750 2 1 1.6 8.6
176 P06746 DNA polymerase beta OS=Homo sapiens GN=POLB PE=1 SV=3 43 38154 1 1 1.8 9.01
177 Q92820 Gamma-glutamyl hydrolase OS=Homo sapiens GN=GGH PE=1 SV=2 43 35941 1 1 4.1 6.67
178 Q9TNN7 HLA class I histocompatibility antigen, Cw-5 alpha chain OS=Homo sapiens GN=HLA-C PE=2 SV=1 43 40886 1 1 3.6 7.11
179 Q12968_REVERSED Nuclear factor of activated T-cells, cytoplasmic 3 OS=Homo sapiens GN=NFATC3 PE=1 SV=1 - REVERSED 41 115522 3 1 0.7 5.91
180 Q96M89 Coiled-coil domain-containing protein 138 OS=Homo sapiens GN=CCDC138 PE=1 SV=1 41 76171 2 1 2.3 8.75
181 Q9Y5E4_REVERSED Protocadherin beta-5 OS=Homo sapiens GN=PCDHB5 PE=1 SV=1 - REVERSED 41 86369 2 1 1 4.87
182 Q9NU02_REVERSED Ankyrin repeat domain-containing protein 5 OS=Homo sapiens GN=ANKRD5 PE=2 SV=2 - REVERSED 41 86610 2 1 1.8 8.51
183 Q7Z5M8 Abhydrolase domain-containing protein 12B OS=Homo sapiens GN=ABHD12B PE=2 SV=1 41 40750 4 1 1.7 8.57
184 Q15020_REVERSED Squamous cell carcinoma antigen recognized by T-cells 3 OS=Homo sapiens GN=SART3 PE=1 SV=1 - REVERSED 41 109865 4 1 0.6 5.45
185 P10643 Complement component C7 OS=Homo sapiens GN=C7 PE=1 SV=2 41 93457 2 1 2 6.09
186 P61769 Beta-2-microglobulin OS=Homo sapiens GN=B2M PE=1 SV=1 40 13706 1 1 8.4 6.06
187 Q9BY43 Charged multivesicular body protein 4a OS=Homo sapiens GN=CHMP4A PE=1 SV=3 40 25083 2 1 6.3 4.65
188 O60312 Probable phospholipid-transporting ATPase VA OS=Homo sapiens GN=ATP10A PE=2 SV=2 40 167582 2 1 0.6 8.7
189 Q8NBM4 Ubiquitin-associated domain-containing protein 2 OS=Homo sapiens GN=UBAC2 PE=2 SV=1 40 38938 2 1 2.3 9.21
190 Q99798 Aconitate hydratase, mitochondrial OS=Homo sapiens GN=ACO2 PE=1 SV=2 40 85372 1 1 1 7.36
191 Q14849 StAR-related lipid transfer protein 3 OS=Homo sapiens GN=STARD3 PE=1 SV=2 40 50471 1 1 1.6 8.52
192 Q8NFY9 Kelch repeat and BTB domain-containing protein 8 OS=Homo sapiens GN=KBTBD8 PE=2 SV=2 39 68778 1 1 1.8 5.88
193 Q96K21 Zinc finger FYVE domain-containing protein 19 OS=Homo sapiens GN=ZFYVE19 PE=1 SV=3 39 51514 1 1 1.7 5.57
194 P02760 Protein AMBP OS=Homo sapiens GN=AMBP PE=1 SV=1 39 38974 1 1 2 5.95
195 P53675_REVERSED Clathrin heavy chain 2 OS=Homo sapiens GN=CLTCL1 PE=1 SV=2 - REVERSED 39 186910 1 1 0.5 5.57
196 P0C869 Cytosolic phospholipase A2 beta OS=Homo sapiens GN=PLA2G4B PE=1 SV=2 39 87922 2 1 1 5.64
197 Q8WUM4 Programmed cell death 6-interacting protein OS=Homo sapiens GN=PDCD6IP PE=1 SV=1 39 95963 1 1 1.6 6.13
198 P05156 Complement factor I OS=Homo sapiens GN=CFI PE=1 SV=2 39 65707 1 1 1.2 7.72
199 P00736 Complement C1r subcomponent OS=Homo sapiens GN=C1R PE=1 SV=2 39 80067 2 1 3.4 5.82
200 Q00535_REVERSED Cyclin-dependent kinase 5 OS=Homo sapiens GN=CDK5 PE=1 SV=3 - REVERSED 38 33283 1 1 3.1 7.9
201 Q9ULT0 Tetratricopeptide repeat protein 7A OS=Homo sapiens GN=TTC7A PE=1 SV=3 38 96123 2 1 1.2 6.03
202 P25786 Proteasome subunit alpha type-1 OS=Homo sapiens GN=PSMA1 PE=1 SV=1 38 29537 1 1 4.6 6.15
203 Q9Y2C9_REVERSED Toll-like receptor 6 OS=Homo sapiens GN=TLR6 PE=2 SV=2 - REVERSED 38 91820 1 1 1 6.83
204 Q14C86_REVERSED GTPase-activating protein and VPS9 domain-containing protein 1 OS=Homo sapiens GN=GAPVD1 PE=1 SV=2 - REVERSED 38 164876 2 1 0.5 5.09
205 Q5T1B0 Axonemal dynein light chain domain-containing protein 1 OS=Homo sapiens GN=AXDND1 PE=1 SV=1 38 117953 1 1 0.6 5.49
206 Q9P2G4_REVERSED Uncharacterized protein KIAA1383 OS=Homo sapiens GN=KIAA1383 PE=1 SV=2 - REVERSED 38 100283 1 1 0.8 6.77
207 Q6NUP7 Serine/threonine-protein phosphatase 4 regulatory subunit 4 OS=Homo sapiens GN=PPP4R4 PE=1 SV=1 38 99389 2 1 0.7 7.96
208 Q16348 Solute carrier family 15 member 2 OS=Homo sapiens GN=SLC15A2 PE=2 SV=2 38 81730 3 1 1.5 8.4
209 Q9P2E9 Ribosome-binding protein 1 OS=Homo sapiens GN=RRBP1 PE=1 SV=4 38 152381 1 1 0.4 8.69
210 P04920 Anion exchange protein 2 OS=Homo sapiens GN=SLC4A2 PE=1 SV=4 38 136923 1 1 0.5 5.9
211 Q99819 Rho GDP-dissociation inhibitor 3 OS=Homo sapiens GN=ARHGDIG PE=2 SV=2 38 25082 1 1 2.7 5.45
212 O95071_REVERSED E3 ubiquitin-protein ligase UBR5 OS=Homo sapiens GN=UBR5 PE=1 SV=2 - REVERSED 38 309158 2 1 0.3 5.59
213 P22352 Glutathione peroxidase 3 OS=Homo sapiens GN=GPX3 PE=1 SV=2 37 25537 1 1 5.3 8.26
214 P29401 Transketolase OS=Homo sapiens GN=TKT PE=1 SV=3 37 67835 1 1 2.9 7.58
215 P35443 Thrombospondin-4 OS=Homo sapiens GN=THBS4 PE=1 SV=2 37 105802 1 1 1 4.44
216 Q14028_REVERSED Cyclic nucleotide-gated cation channel beta-1 OS=Homo sapiens GN=CNGB1 PE=1 SV=2 - REVERSED 37 139590 2 1 0.6 4.76
217 Q14573_REVERSED Inositol 1,4,5-trisphosphate receptor type 3 OS=Homo sapiens GN=ITPR3 PE=1 SV=2 - REVERSED 37 303912 2 1 0.4 6.05
218 Q9C0B2 Uncharacterized protein KIAA1751 OS=Homo sapiens GN=KIAA1751 PE=2 SV=2 37 86902 2 1 1.6 5.39
219 Q6ZUT9_REVERSED DENN domain-containing protein 5B OS=Homo sapiens GN=DENND5B PE=1 SV=2 - REVERSED 37 144927 2 1 1 6.29
220 O94991 SLIT and NTRK-like protein 5 OS=Homo sapiens GN=SLITRK5 PE=1 SV=2 37 107418 2 1 1.4 6.48
221 O94868_REVERSED FCH and double SH3 domains protein 2 OS=Homo sapiens GN=FCHSD2 PE=1 SV=3 - REVERSED 37 84224 2 1 1.9 5.55
222 Q4G0X9_REVERSED Coiled-coil domain-containing protein 40 OS=Homo sapiens GN=CCDC40 PE=2 SV=2 - REVERSED 37 130033 1 1 0.5 5.21
223 Q9UC07_REVERSED Zinc finger protein 69 OS=Homo sapiens GN=ZNF69 PE=2 SV=2 - REVERSED 37 65718 1 1 1.1 9.13
224 Q9Y496_REVERSED Kinesin-like protein KIF3A OS=Homo sapiens GN=KIF3A PE=1 SV=4 - REVERSED 37 79991 1 1 0.9 6.16
225 Q14997 Proteasome activator complex subunit 4 OS=Homo sapiens GN=PSME4 PE=1 SV=2 37 211199 1 1 0.3 6.45
226 Q92841 Probable ATP-dependent RNA helicase DDX17 OS=Homo sapiens GN=DDX17 PE=1 SV=1 37 72326 1 1 0.9 8.82
227 Q8N6K7 Sterile alpha motif domain-containing protein 3 OS=Homo sapiens GN=SAMD3 PE=2 SV=2 37 61197 7 1 2.1 6.67
228 Q9NU22 Midasin OS=Homo sapiens GN=MDN1 PE=1 SV=2 36 632420 1 1 0.2 5.46
229 A8MW92_REVERSED PHD finger protein 20-like protein 1 OS=Homo sapiens GN=PHF20L1 PE=1 SV=2 - REVERSED 36 114938 1 1 1.3 6.4
230 O95613 Pericentrin OS=Homo sapiens GN=PCNT PE=1 SV=4 36 377806 2 1 0.6 5.4
231 P10242_REVERSED Transcriptional activator Myb OS=Homo sapiens GN=MYB PE=1 SV=2 - REVERSED 36 72296 3 1 1.3 6.35
232 Q7L8A9 Vasohibin-1 OS=Homo sapiens GN=VASH1 PE=1 SV=1 36 40931 1 1 1.9 9.5
HDF
prot_hit prot_acc prot_desc prot_score prot_mass prot_matches prot_sequences_sig prot_cover prot_pi
1 P02751 Fibronectin OS=Homo sapiens GN=FN1 PE=1 SV=4 16183 262460 423 86 66 5.46
2 P07996 Thrombospondin-1 OS=Homo sapiens GN=THBS1 PE=1 SV=2 3990 129300 115 42 49.4 4.71
3 P12111 Collagen alpha-3(VI) chain OS=Homo sapiens GN=COL6A3 PE=1 SV=5 3014 343457 118 59 33.3 6.26
4 P02768 Serum albumin OS=Homo sapiens GN=ALB PE=1 SV=2 2557 69321 114 8 11.8 5.92
5 P02765 Alpha-2-HS-glycoprotein OS=Homo sapiens GN=AHSG PE=1 SV=1 2385 39300 60 4 8.7 5.43
6 P04264 Keratin, type II cytoskeletal 1 OS=Homo sapiens GN=KRT1 PE=1 SV=6 2277 65999 62 25 51.1 8.15
7 P13645 Keratin, type I cytoskeletal 10 OS=Homo sapiens GN=KRT10 PE=1 SV=6 2109 58792 64 27 54.8 5.13
8 P03956 Interstitial collagenase OS=Homo sapiens GN=MMP1 PE=1 SV=3 1904 53973 57 20 60.8 6.47
9 P07093 Glia-derived nexin OS=Homo sapiens GN=SERPINE2 PE=1 SV=1 1820 43974 53 17 59.8 9.35
10 P35908 Keratin, type II cytoskeletal 2 epidermal OS=Homo sapiens GN=KRT2 PE=1 SV=2 1740 65393 52 23 56.2 8.07
11 P01023 Alpha-2-macroglobulin OS=Homo sapiens GN=A2M PE=1 SV=3 1729 163188 56 11 7.5 6.03
12 P24821 Tenascin OS=Homo sapiens GN=TNC PE=1 SV=3 1664 240700 47 31 25.3 4.79
13 P35527 Keratin, type I cytoskeletal 9 OS=Homo sapiens GN=KRT9 PE=1 SV=3 1625 62027 36 20 62.6 5.14
14 P08253 72 kDa type IV collagenase OS=Homo sapiens GN=MMP2 PE=1 SV=2 1609 73835 50 24 60.5 5.26
15 P20742 Pregnancy zone protein OS=Homo sapiens GN=PZP PE=1 SV=4 1393 163760 42 5 4 5.97
16 P43235 Cathepsin K OS=Homo sapiens GN=CTSK PE=1 SV=1 1336 36942 26 9 53.5 8.72
17 P01024 Complement C3 OS=Homo sapiens GN=C3 PE=1 SV=2 1296 187030 53 13 15.4 6.02
18 P60709 Actin, cytoplasmic 1 OS=Homo sapiens GN=ACTB PE=1 SV=1 951 41710 31 14 47.5 5.29
19 P12109 Collagen alpha-1(VI) chain OS=Homo sapiens GN=COL6A1 PE=1 SV=3 940 108462 21 13 19.6 5.26
20 P98160 Basement membrane-specific heparan sulfate proteoglycan core protein OS=Homo sapiens GN=HSPG2 PE=1 SV=4 933 468532 48 30 13.6 6.06
21 P00734 Prothrombin OS=Homo sapiens GN=F2 PE=1 SV=2 818 69992 18 8 10.9 5.64
22 P19823 Inter-alpha-trypsin inhibitor heavy chain H2 OS=Homo sapiens GN=ITIH2 PE=1 SV=2 799 106397 24 8 9.3 6.4
23 P06396 Gelsolin OS=Homo sapiens GN=GSN PE=1 SV=1 746 85644 26 11 14.3 5.9
24 P26022 Pentraxin-related protein PTX3 OS=Homo sapiens GN=PTX3 PE=1 SV=3 727 41949 16 8 30.2 4.94
25 P36955 Pigment epithelium-derived factor OS=Homo sapiens GN=SERPINF1 PE=1 SV=4 680 46283 23 5 12.2 5.97
26 Q08380 Galectin-3-binding protein OS=Homo sapiens GN=LGALS3BP PE=1 SV=1 667 65289 21 12 30.3 5.13
27 Q15582 Transforming growth factor-beta-induced protein ig-h3 OS=Homo sapiens GN=TGFBI PE=1 SV=1 604 74634 17 14 28.8 7.62
28 Q99715 Collagen alpha-1(XII) chain OS=Homo sapiens GN=COL12A1 PE=1 SV=2 550 332941 23 19 11.7 5.38
29 Q14393 Growth arrest-specific protein 6 OS=Homo sapiens GN=GAS6 PE=1 SV=2 527 79625 16 10 19.4 5.84
30 P68032 Actin, alpha cardiac muscle 1 OS=Homo sapiens GN=ACTC1 PE=1 SV=1 521 41992 19 9 26.8 5.23
31 Q07954 Prolow-density lipoprotein receptor-related protein 1 OS=Homo sapiens GN=LRP1 PE=1 SV=2 455 504276 16 13 4.5 5.16
32 P02771 Alpha-fetoprotein OS=Homo sapiens GN=AFP PE=1 SV=1 433 68633 13 4 6.7 5.48
33 P12110 Collagen alpha-2(VI) chain OS=Homo sapiens GN=COL6A2 PE=1 SV=4 430 108512 19 12 18.1 5.85
34 P02788 Lactotransferrin OS=Homo sapiens GN=LTF PE=1 SV=6 429 78132 18 3 4.8 8.5
35 P0C0L4 Complement C4-A OS=Homo sapiens GN=C4A PE=1 SV=1 405 192650 15 8 5.6 6.65
36 P08254 Stromelysin-1 OS=Homo sapiens GN=MMP3 PE=1 SV=2 403 53943 15 9 24.9 5.77
37 P69905 Hemoglobin subunit alpha OS=Homo sapiens GN=HBA1 PE=1 SV=2 386 15248 18 5 34.5 8.72
38 P02533 Keratin, type I cytoskeletal 14 OS=Homo sapiens GN=KRT14 PE=1 SV=4 376 51529 13 9 23.5 5.09
39 P35442 Thrombospondin-2 OS=Homo sapiens GN=THBS2 PE=1 SV=2 331 129908 13 8 9.2 4.62
40 P04406 Glyceraldehyde-3-phosphate dehydrogenase OS=Homo sapiens GN=GAPDH PE=1 SV=3 323 36030 9 5 26.3 8.57
41 P05121 Plasminogen activator inhibitor 1 OS=Homo sapiens GN=SERPINE1 PE=1 SV=1 323 45031 12 9 41.5 6.68
42 Q562R1 Beta-actin-like protein 2 OS=Homo sapiens GN=ACTBL2 PE=1 SV=2 320 41976 11 4 13 5.39
43 Q8WUJ3 Protein KIAA1199 OS=Homo sapiens GN=KIAA1199 PE=1 SV=2 308 152900 13 9 12 7.98
44 O00391 Sulfhydryl oxidase 1 OS=Homo sapiens GN=QSOX1 PE=1 SV=3 301 82526 9 6 14.5 9.13
45 P07355 Annexin A2 OS=Homo sapiens GN=ANXA2 PE=1 SV=2 298 38580 8 6 31 7.57
46 Q06033 Inter-alpha-trypsin inhibitor heavy chain H3 OS=Homo sapiens GN=ITIH3 PE=1 SV=2 294 99787 11 4 8.8 5.49
47 P07339 Cathepsin D OS=Homo sapiens GN=CTSD PE=1 SV=1 282 44524 13 7 25.2 6.1
48 Q08431 Lactadherin OS=Homo sapiens GN=MFGE8 PE=1 SV=2 281 43095 12 8 26.6 8.47
49 P08123 Collagen alpha-2(I) chain OS=Homo sapiens GN=COL1A2 PE=1 SV=7 262 129235 7 7 8.4 9.08
50 P15144 Aminopeptidase N OS=Homo sapiens GN=ANPEP PE=1 SV=4 254 109471 12 6 13 5.31
51 P08758 Annexin A5 OS=Homo sapiens GN=ANXA5 PE=1 SV=2 249 35914 9 5 19.1 4.94
52 P13647 Keratin, type II cytoskeletal 5 OS=Homo sapiens GN=KRT5 PE=1 SV=3 246 62340 14 7 14.6 7.59
53 P02774 Vitamin D-binding protein OS=Homo sapiens GN=GC PE=1 SV=1 244 52929 9 5 12 5.4
54 P51884 Lumican OS=Homo sapiens GN=LUM PE=1 SV=2 239 38405 14 6 21.9 6.16
55 Q13219 Pappalysin-1 OS=Homo sapiens GN=PAPPA PE=1 SV=3 238 180856 6 4 6.1 5.76
56 P14618 Pyruvate kinase isozymes M1/M2 OS=Homo sapiens GN=PKM2 PE=1 SV=4 238 57900 7 7 21.3 7.96
57 P11142 Heat shock cognate 71 kDa protein OS=Homo sapiens GN=HSPA8 PE=1 SV=1 234 70854 5 5 10.2 5.37
58 P04114 Apolipoprotein B-100 OS=Homo sapiens GN=APOB PE=1 SV=2 231 515283 15 6 2.2 6.58
59 P98095 Fibulin-2 OS=Homo sapiens GN=FBLN2 PE=1 SV=2 221 126489 7 4 8.4 4.73
60 P04259 Keratin, type II cytoskeletal 6B OS=Homo sapiens GN=KRT6B PE=1 SV=5 216 60030 12 5 10.1 8.09
61 Q15063 Periostin OS=Homo sapiens GN=POSTN PE=1 SV=2 215 93255 11 7 15.1 7.27
62 P00747 Plasminogen OS=Homo sapiens GN=PLG PE=1 SV=2 212 90510 9 5 8.5 7.04
63 P12259 Coagulation factor V OS=Homo sapiens GN=F5 PE=1 SV=4 210 251546 10 6 4.2 5.68
64 Q16363 Laminin subunit alpha-4 OS=Homo sapiens GN=LAMA4 PE=1 SV=4 210 202397 4 3 4.1 5.89
65 P04004 Vitronectin OS=Homo sapiens GN=VTN PE=1 SV=1 209 54271 4 1 3.1 5.55
66 P55268 Laminin subunit beta-2 OS=Homo sapiens GN=LAMB2 PE=1 SV=2 207 195854 11 6 6.4 6.07
67 P02647 Apolipoprotein A-I OS=Homo sapiens GN=APOA1 PE=1 SV=1 204 30759 4 1 6 5.56
68 P23142 Fibulin-1 OS=Homo sapiens GN=FBLN1 PE=1 SV=4 202 77162 11 5 10.2 5.07
69 Q9Y6C2 EMILIN-1 OS=Homo sapiens GN=EMILIN1 PE=1 SV=2 198 106601 5 4 8.8 5.07
70 Q99816 Tumor susceptibility gene 101 protein OS=Homo sapiens GN=TSG101 PE=1 SV=2 60 43916 15 5 19.2 5.2
71 P05543 Thyroxine-binding globulin OS=Homo sapiens GN=SERPINA7 PE=1 SV=2 197 46295 6 3 8.2 5.87
72 Q13510 Acid ceramidase OS=Homo sapiens GN=ASAH1 PE=1 SV=5 197 44631 5 5 18.5 7.52
73 P02452 Collagen alpha-1(I) chain OS=Homo sapiens GN=COL1A1 PE=1 SV=5 188 138857 8 4 5.2 5.6
74 P11047 Laminin subunit gamma-1 OS=Homo sapiens GN=LAMC1 PE=1 SV=3 179 177489 6 4 3.8 5.01
75 P02042 Hemoglobin subunit delta OS=Homo sapiens GN=HBD PE=1 SV=2 176 16045 11 2 12.9 7.85
76 P35443 Thrombospondin-4 OS=Homo sapiens GN=THBS4 PE=1 SV=2 171 105802 6 5 7.3 4.44
77 P49747 Cartilage oligomeric matrix protein OS=Homo sapiens GN=COMP PE=1 SV=2 167 82808 6 5 7.1 4.36
78 P19827 Inter-alpha-trypsin inhibitor heavy chain H1 OS=Homo sapiens GN=ITIH1 PE=1 SV=3 166 101326 3 3 2.9 6.31
79 P06733 Alpha-enolase OS=Homo sapiens GN=ENO1 PE=1 SV=2 165 47139 4 3 12 7.01
80 P13929 Beta-enolase OS=Homo sapiens GN=ENO3 PE=1 SV=4 163 46902 2 2 7.6 7.59
81 P01031 Complement C5 OS=Homo sapiens GN=C5 PE=1 SV=4 162 188186 7 4 3 6.11
82 P02538 Keratin, type II cytoskeletal 6A OS=Homo sapiens GN=KRT6A PE=1 SV=3 159 60008 9 4 9.9 8.09
83 P02100 Hemoglobin subunit epsilon OS=Homo sapiens GN=HBE1 PE=1 SV=2 159 16192 7 2 12.9 8.67
84 P07858 Cathepsin B OS=Homo sapiens GN=CTSB PE=1 SV=3 159 37797 5 4 15 5.88
85 Q6YHK3 CD109 antigen OS=Homo sapiens GN=CD109 PE=1 SV=2 158 161587 6 5 5.3 5.59
86 P42785 Lysosomal Pro-X carboxypeptidase OS=Homo sapiens GN=PRCP PE=1 SV=1 155 55764 2 1 4.2 6.75
87 P02649 Apolipoprotein E OS=Homo sapiens GN=APOE PE=1 SV=1 154 36132 6 3 9.1 5.65
88 P02794 Ferritin heavy chain OS=Homo sapiens GN=FTH1 PE=1 SV=2 154 21212 5 4 38.3 5.3
89 P02675 Fibrinogen beta chain OS=Homo sapiens GN=FGB PE=1 SV=2 154 55892 3 2 3.9 8.54
90 Q92626 Peroxidasin homolog OS=Homo sapiens GN=PXDN PE=1 SV=2 153 165170 5 4 4.9 6.79
91 P08697 Alpha-2-antiplasmin OS=Homo sapiens GN=SERPINF2 PE=1 SV=3 150 54531 2 1 2.2 5.87
92 P00736 Complement C1r subcomponent OS=Homo sapiens GN=C1R PE=1 SV=2 150 80067 6 5 13 5.82
93 Q9Y4K0 Lysyl oxidase homolog 2 OS=Homo sapiens GN=LOXL2 PE=1 SV=1 148 86668 3 3 5.7 5.95
94 Q13103 Secreted phosphoprotein 24 OS=Homo sapiens GN=SPP2 PE=1 SV=1 147 24322 2 1 5.7 8.59
95 P34931 Heat shock 70 kDa protein 1-like OS=Homo sapiens GN=HSPA1L PE=1 SV=2 145 70331 3 3 7 5.76
96 P04083 Annexin A1 OS=Homo sapiens GN=ANXA1 PE=1 SV=2 141 38690 3 3 16.2 6.57
97 Q9Y490 Talin-1 OS=Homo sapiens GN=TLN1 PE=1 SV=3 140 269599 7 4 4.2 5.77
98 P68363 Tubulin alpha-1B chain OS=Homo sapiens GN=TUBA1B PE=1 SV=1 139 50120 8 4 19.7 4.94
99 P15169 Carboxypeptidase N catalytic chain OS=Homo sapiens GN=CPN1 PE=1 SV=1 139 52253 2 1 3.3 6.86
100 P07686 Beta-hexosaminidase subunit beta OS=Homo sapiens GN=HEXB PE=1 SV=3 137 63071 8 3 15.8 6.29
101 P02748 Complement component C9 OS=Homo sapiens GN=C9 PE=1 SV=2 135 63133 3 2 3.8 5.43
102 P19013 Keratin, type II cytoskeletal 4 OS=Homo sapiens GN=KRT4 PE=1 SV=4 130 57250 8 4 7.3 6.25
103 P07437 Tubulin beta chain OS=Homo sapiens GN=TUBB PE=1 SV=2 129 49639 6 4 18.5 4.78
104 Q9Y240 C-type lectin domain family 11 member A OS=Homo sapiens GN=CLEC11A PE=1 SV=1 128 35672 3 1 8 5.06
105 P51888 Prolargin OS=Homo sapiens GN=PRELP PE=1 SV=1 124 43782 6 4 15.2 9.47
106 Q9UBG0 C-type mannose receptor 2 OS=Homo sapiens GN=MRC2 PE=1 SV=2 123 166568 1 1 1.1 5.54
107 P00742 Coagulation factor X OS=Homo sapiens GN=F10 PE=1 SV=2 122 54697 5 3 7.6 5.68
108 Q86YZ3 Hornerin OS=Homo sapiens GN=HRNR PE=1 SV=2 115 282228 1 1 0.7 10.05
109 P13646 Keratin, type I cytoskeletal 13 OS=Homo sapiens GN=KRT13 PE=1 SV=4 112 49557 5 4 6.1 4.91
110 P09871 Complement C1s subcomponent OS=Homo sapiens GN=C1S PE=1 SV=1 111 76635 5 4 6.8 4.86
111 P10619 Lysosomal protective protein OS=Homo sapiens GN=CTSA PE=1 SV=2 106 54431 4 3 12.5 6.16
112 P62979 Ubiquitin-40S ribosomal protein S27a OS=Homo sapiens GN=RPS27A PE=1 SV=2 100 17953 7 3 28.2 9.68
113 Q02809 Procollagen-lysine,2-oxoglutarate 5-dioxygenase 1 OS=Homo sapiens GN=PLOD1 PE=1 SV=2 98 83497 4 2 8.4 6.47
114 P07602 Proactivator polypeptide OS=Homo sapiens GN=PSAP PE=1 SV=2 96 58074 4 3 7.6 5.06
115 P07900 Heat shock protein HSP 90-alpha OS=Homo sapiens GN=HSP90AA1 PE=1 SV=5 96 84607 5 2 5.9 4.94
116 P49746 Thrombospondin-3 OS=Homo sapiens GN=THBS3 PE=1 SV=1 94 104135 7 2 8.1 4.43
117 P08294 Extracellular superoxide dismutase [Cu-Zn] OS=Homo sapiens GN=SOD3 PE=1 SV=2 94 25835 1 1 5.4 6.14
118 P00450 Ceruloplasmin OS=Homo sapiens GN=CP PE=1 SV=1 94 122128 3 1 2 5.44
119 P08473 Neprilysin OS=Homo sapiens GN=MME PE=1 SV=2 93 85460 2 2 3.2 5.54
120 P04745 Alpha-amylase 1 OS=Homo sapiens GN=AMY1A PE=1 SV=2 91 57731 2 2 5.5 6.47
121 P08238 Heat shock protein HSP 90-beta OS=Homo sapiens GN=HSP90AB1 PE=1 SV=4 91 83212 3 2 4 4.97
122 P21333 Filamin-A OS=Homo sapiens GN=FLNA PE=1 SV=4 89 280564 5 3 2.6 5.7
123 Q8NI35 InaD-like protein OS=Homo sapiens GN=INADL PE=1 SV=3 89 196247 8 1 0.3 4.84
124 P00558 Phosphoglycerate kinase 1 OS=Homo sapiens GN=PGK1 PE=1 SV=3 88 44586 2 1 7.9 8.3
125 P48723 Heat shock 70 kDa protein 13 OS=Homo sapiens GN=HSPA13 PE=1 SV=1 88 51895 3 3 8.1 5.52
126 Q8TEV9_REVERSED Smith-Magenis syndrome chromosomal region candidate gene 8 protein OS=Homo sapiens GN=SMCR8 PE=1 SV=2 - REVERSED 86 104956 10 1 1.2 5.36
127 P09486 SPARC OS=Homo sapiens GN=SPARC PE=1 SV=1 85 34610 3 2 16.5 4.73
128 P68371 Tubulin beta-2C chain OS=Homo sapiens GN=TUBB2C PE=1 SV=1 85 49799 5 3 15.7 4.79
129 P27487 Dipeptidyl peptidase 4 OS=Homo sapiens GN=DPP4 PE=1 SV=2 85 88222 2 1 3 5.67
130 P09238 Stromelysin-2 OS=Homo sapiens GN=MMP10 PE=1 SV=1 84 54117 4 2 6.1 5.49
131 Q8N436 Inactive carboxypeptidase-like protein X2 OS=Homo sapiens GN=CPXM2 PE=2 SV=3 84 85816 2 2 4 6.4
132 O95497 Pantetheinase OS=Homo sapiens GN=VNN1 PE=1 SV=2 81 56975 1 1 3.7 5.32
133 Q06828 Fibromodulin OS=Homo sapiens GN=FMOD PE=1 SV=2 80 43152 1 1 3.5 5.66
134 O00560 Syntenin-1 OS=Homo sapiens GN=SDCBP PE=1 SV=1 79 32424 4 2 21.1 7.05
135 P23526 Adenosylhomocysteinase OS=Homo sapiens GN=AHCY PE=1 SV=4 79 47685 4 3 12 5.92
136 P33908 Mannosyl-oligosaccharide 1,2-alpha-mannosidase IA OS=Homo sapiens GN=MAN1A1 PE=1 SV=3 78 72922 3 2 6.4 6.04
137 O14773 Tripeptidyl-peptidase 1 OS=Homo sapiens GN=TPP1 PE=1 SV=2 77 61210 1 1 4.8 6.01
138 P01008 Antithrombin-III OS=Homo sapiens GN=SERPINC1 PE=1 SV=1 77 52569 5 2 7.3 6.32
139 P16035 Metalloproteinase inhibitor 2 OS=Homo sapiens GN=TIMP2 PE=1 SV=2 77 24383 4 2 12.7 7.45
140 P11717 Cation-independent mannose-6-phosphate receptor OS=Homo sapiens GN=IGF2R PE=1 SV=3 77 274199 4 3 2.2 5.64
141 Q12860 Contactin-1 OS=Homo sapiens GN=CNTN1 PE=1 SV=1 75 113249 3 2 3.4 5.62
142 Q92743 Serine protease HTRA1 OS=Homo sapiens GN=HTRA1 PE=1 SV=1 75 51255 1 1 2.9 8.09
143 P05067 Amyloid beta A4 protein OS=Homo sapiens GN=APP PE=1 SV=3 74 86888 2 1 2.6 4.73
144 Q8TAA3 Proteasome subunit alpha type-7-like OS=Homo sapiens GN=PSMA8 PE=1 SV=3 74 28512 1 1 5.5 9.07
145 Q5D862 Filaggrin-2 OS=Homo sapiens GN=FLG2 PE=1 SV=1 72 247928 2 2 1 8.45
146 O00300 Tumor necrosis factor receptor superfamily member 11B OS=Homo sapiens GN=TNFRSF11B PE=1 SV=3 71 45996 1 1 4.2 8.66
147 P02790 Hemopexin OS=Homo sapiens GN=HPX PE=1 SV=2 71 51643 4 1 1.9 6.55
148 O15111_REVERSED Inhibitor of nuclear factor kappa-B kinase subunit alpha OS=Homo sapiens GN=CHUK PE=1 SV=2 - REVERSED 71 84585 6 1 1.6 6.28
149 P28799 Granulins OS=Homo sapiens GN=GRN PE=1 SV=2 69 63500 4 2 6.2 6.43
150 O75508 Claudin-11 OS=Homo sapiens GN=CLDN11 PE=1 SV=2 69 21978 1 1 6.8 8.22
151 P81605 Dermcidin OS=Homo sapiens GN=DCD PE=1 SV=2 68 11277 2 2 20 6.08
152 P55058 Phospholipid transfer protein OS=Homo sapiens GN=PLTP PE=1 SV=1 68 54705 1 1 2.2 6.53
153 P02749 Beta-2-glycoprotein 1 OS=Homo sapiens GN=APOH PE=1 SV=3 68 38273 3 1 4.3 8.34
154 P60033 CD81 antigen OS=Homo sapiens GN=CD81 PE=1 SV=1 67 25792 2 1 18.2 5.09
155 Q8WXE9 Stonin-2 OS=Homo sapiens GN=STON2 PE=1 SV=1 67 101102 2 1 1 5.2
156 O95445 Apolipoprotein M OS=Homo sapiens GN=APOM PE=1 SV=2 66 21239 5 1 6.9 5.66
157 Q7Z304 MAM domain-containing protein 2 OS=Homo sapiens GN=MAMDC2 PE=2 SV=3 66 77506 3 2 3.9 5.05
158 Q12931 Heat shock protein 75 kDa, mitochondrial OS=Homo sapiens GN=TRAP1 PE=1 SV=3 65 80060 3 1 2.8 8.3
159 P19883 Follistatin OS=Homo sapiens GN=FST PE=1 SV=2 65 37981 2 1 7 5.53
160 P61224 Ras-related protein Rap-1b OS=Homo sapiens GN=RAP1B PE=1 SV=1 64 20812 2 2 12.5 5.65
161 P20908 Collagen alpha-1(V) chain OS=Homo sapiens GN=COL5A1 PE=1 SV=3 64 183447 1 1 1.2 4.94
162 P10599 Thioredoxin OS=Homo sapiens GN=TXN PE=1 SV=3 64 11730 1 1 12.4 4.82
163 P08133 Annexin A6 OS=Homo sapiens GN=ANXA6 PE=1 SV=3 62 75826 2 2 4.2 5.42
164 Q9NZP8 Complement C1r subcomponent-like protein OS=Homo sapiens GN=C1RL PE=1 SV=2 62 53464 1 1 2.5 6.75
165 Q03692 Collagen alpha-1(X) chain OS=Homo sapiens GN=COL10A1 PE=1 SV=2 61 66117 1 1 1.9 9.68
166 P26447 Protein S100-A4 OS=Homo sapiens GN=S100A4 PE=1 SV=1 61 11721 2 2 16.8 5.85
167 P14550 Alcohol dehydrogenase [NADP+] OS=Homo sapiens GN=AKR1A1 PE=1 SV=3 61 36550 1 1 3.1 6.32
168 P15121 Aldose reductase OS=Homo sapiens GN=AKR1B1 PE=1 SV=3 61 35830 1 1 3.2 6.51
169 P22105 Tenascin-X OS=Homo sapiens GN=TNXB PE=1 SV=3 60 464034 1 1 0.3 5.19
170 Q9BYX7_REVERSED Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 - REVERSED 57 41989 9 1 3.5 5.92
171 P05546 Heparin cofactor 2 OS=Homo sapiens GN=SERPIND1 PE=1 SV=3 57 57034 2 1 3.8 6.41
172 P02461 Collagen alpha-1(III) chain OS=Homo sapiens GN=COL3A1 PE=1 SV=4 57 138479 3 1 2.9 6.21
173 P07585 Decorin OS=Homo sapiens GN=DCN PE=1 SV=1 57 39722 2 1 9.7 8.75
174 Q03135 Caveolin-1 OS=Homo sapiens GN=CAV1 PE=1 SV=4 56 20458 1 1 7.9 5.65
175 O43264_REVERSED Centromere/kinetochore protein zw10 homolog OS=Homo sapiens GN=ZW10 PE=1 SV=3 - REVERSED 55 88773 2 1 0.8 5.89
176 P62937 Peptidyl-prolyl cis-trans isomerase A OS=Homo sapiens GN=PPIA PE=1 SV=2 55 18001 2 1 13.9 7.68
177 P07737 Profilin-1 OS=Homo sapiens GN=PFN1 PE=1 SV=2 55 15045 2 2 12.1 8.44
178 P02760 Protein AMBP OS=Homo sapiens GN=AMBP PE=1 SV=1 55 38974 1 1 2 5.95
179 B9A064 Immunoglobulin lambda-like polypeptide 5 OS=Homo sapiens GN=IGLL5 PE=2 SV=2 54 23049 1 1 3.7 9.08
180 P18206 Vinculin OS=Homo sapiens GN=VCL PE=1 SV=4 54 123722 2 1 2 5.5
181 P10643 Complement component C7 OS=Homo sapiens GN=C7 PE=1 SV=2 54 93457 2 2 2 6.09
182 Q9UK55 Protein Z-dependent protease inhibitor OS=Homo sapiens GN=SERPINA10 PE=1 SV=1 53 50674 4 1 4.1 8.28
183 Q8NBM4 Ubiquitin-associated domain-containing protein 2 OS=Homo sapiens GN=UBAC2 PE=2 SV=1 52 38938 5 1 2.3 9.21
184 Q14624 Inter-alpha-trypsin inhibitor heavy chain H4 OS=Homo sapiens GN=ITIH4 PE=1 SV=4 52 103293 6 1 2.5 6.51
185 Q86VH2 Kinesin-like protein KIF27 OS=Homo sapiens GN=KIF27 PE=2 SV=1 51 160184 1 1 0.6 6.9
186 P28300 Protein-lysine 6-oxidase OS=Homo sapiens GN=LOX PE=1 SV=2 51 46915 1 1 3.1 8.36
187 P05452 Tetranectin OS=Homo sapiens GN=CLEC3B PE=1 SV=3 51 22522 1 1 10.4 5.52
188 Q8NAV2_REVERSED Uncharacterized protein C8orf58 OS=Homo sapiens GN=C8orf58 PE=2 SV=2 - REVERSED 51 39636 3 1 4.1 8.54
189 P08962 CD63 antigen OS=Homo sapiens GN=CD63 PE=1 SV=2 51 25619 1 1 4.2 8.14
190 P61769 Beta-2-microglobulin OS=Homo sapiens GN=B2M PE=1 SV=1 51 13706 1 1 8.4 6.06
191 P15311 Ezrin OS=Homo sapiens GN=EZR PE=1 SV=4 50 69370 1 1 1.5 5.94
192 P04439 HLA class I histocompatibility antigen, A-3 alpha chain OS=Homo sapiens GN=HLA-A PE=1 SV=2 50 40815 3 2 9.3 5.66
193 Q86Y46 Keratin, type II cytoskeletal 73 OS=Homo sapiens GN=KRT73 PE=1 SV=1 50 58887 8 3 8.5 6.93
194 P26038 Moesin OS=Homo sapiens GN=MSN PE=1 SV=3 50 67778 2 1 3.3 6.08
195 O60312 Probable phospholipid-transporting ATPase VA OS=Homo sapiens GN=ATP10A PE=2 SV=2 50 167582 3 1 0.6 8.7
196 Q9HDC9 Adipocyte plasma membrane-associated protein OS=Homo sapiens GN=APMAP PE=1 SV=2 50 46451 1 1 1.7 5.82
197 Q16394 Exostosin-1 OS=Homo sapiens GN=EXT1 PE=1 SV=2 49 86200 3 1 8.6 9.16
198 Q15751_REVERSED Probable E3 ubiquitin-protein ligase HERC1 OS=Homo sapiens GN=HERC1 PE=1 SV=2 - REVERSED 48 531891 1 1 0.2 5.69
199 P02787 Serotransferrin OS=Homo sapiens GN=TF PE=1 SV=3 47 77014 2 1 1.7 6.81
200 Q13454 Tumor suppressor candidate 3 OS=Homo sapiens GN=TUSC3 PE=2 SV=1 47 39650 1 1 2 9.93
201 P00338 L-lactate dehydrogenase A chain OS=Homo sapiens GN=LDHA PE=1 SV=2 47 36665 1 1 3 8.44
202 P60903 Protein S100-A10 OS=Homo sapiens GN=S100A10 PE=1 SV=2 47 11196 1 1 17.5 6.82
203 P29353_REVERSED SHC-transforming protein 1 OS=Homo sapiens GN=SHC1 PE=1 SV=4 - REVERSED 46 62782 5 1 1.4 6.01
204 P01042 Kininogen-1 OS=Homo sapiens GN=KNG1 PE=1 SV=2 46 71912 1 1 1.4 6.34
205 Q76LX8 A disintegrin and metalloproteinase with thrombospondin motifs 13 OS=Homo sapiens GN=ADAMTS13 PE=1 SV=1 45 153505 1 1 0.7 6.96
206 Q9UHL4 Dipeptidyl peptidase 2 OS=Homo sapiens GN=DPP7 PE=1 SV=3 45 54307 1 1 2.4 5.91
207 P07225 Vitamin K-dependent protein S OS=Homo sapiens GN=PROS1 PE=1 SV=1 45 75074 2 1 2.4 5.48
208 Q99456 Keratin, type I cytoskeletal 12 OS=Homo sapiens GN=KRT12 PE=1 SV=1 45 53478 3 1 3.2 4.7
209 Q96CD0_REVERSED F-box/LRR-repeat protein 8 OS=Homo sapiens GN=FBXL8 PE=1 SV=1 - REVERSED 44 40490 6 1 1.9 7.03
210 Q6ZU45 Putative C-type lectin domain-containing protein NCRNA00083 OS=Homo sapiens GN=NCRNA00083 PE=5 SV=1 44 25796 1 1 3.4 6.3
211 Q3KQU3 MAP7 domain-containing protein 1 OS=Homo sapiens GN=MAP7D1 PE=1 SV=1 44 92764 1 1 0.7 10.12
212 Q5U5X0 LYR motif-containing protein 7 OS=Homo sapiens GN=LYRM7 PE=1 SV=1 44 11947 1 1 5.8 9.67
213 Q8N7B9_REVERSED EF-hand calcium-binding domain-containing protein 3 OS=Homo sapiens GN=EFCAB3 PE=1 SV=1 - REVERSED 44 50114 1 1 1.4 9.31
214 Q8NB25_REVERSED Protein FAM184A OS=Homo sapiens GN=FAM184A PE=2 SV=3 - REVERSED 44 132883 1 1 0.5 5.59
215 P06732 Creatine kinase M-type OS=Homo sapiens GN=CKM PE=1 SV=2 44 43074 1 1 1.6 6.77
216 Q93096_REVERSED Protein tyrosine phosphatase type IVA 1 OS=Homo sapiens GN=PTP4A1 PE=1 SV=2 - REVERSED 44 19802 1 1 3.5 9.17
217 Q8IWC1 MAP7 domain-containing protein 3 OS=Homo sapiens GN=MAP7D3 PE=1 SV=2 44 98368 1 1 0.7 9.34
218 P11137 Microtubule-associated protein 2 OS=Homo sapiens GN=MAP2 PE=1 SV=4 42 199404 2 1 1.1 4.82
219 Q16348 Solute carrier family 15 member 2 OS=Homo sapiens GN=SLC15A2 PE=2 SV=2 42 81730 3 1 1.5 8.4
220 P15531 Nucleoside diphosphate kinase A OS=Homo sapiens GN=NME1 PE=1 SV=1 42 17138 1 1 11.2 5.83
221 Q14566_REVERSED DNA replication licensing factor MCM6 OS=Homo sapiens GN=MCM6 PE=1 SV=1 - REVERSED 41 92831 1 1 0.9 5.28
222 O75830 Serpin I2 OS=Homo sapiens GN=SERPINI2 PE=2 SV=1 41 46116 1 1 3.5 5.08
223 O60568 Procollagen-lysine,2-oxoglutarate 5-dioxygenase 3 OS=Homo sapiens GN=PLOD3 PE=1 SV=1 41 84731 1 1 1.9 5.69
224 Q8NHP8 Putative phospholipase B-like 2 OS=Homo sapiens GN=PLBD2 PE=1 SV=2 41 65430 2 1 3.2 6.34
225 Q99466_REVERSED Neurogenic locus notch homolog protein 4 OS=Homo sapiens GN=NOTCH4 PE=1 SV=2 - REVERSED 41 209480 2 1 0.4 5.43
226 P17936 Insulin-like growth factor-binding protein 3 OS=Homo sapiens GN=IGFBP3 PE=1 SV=2 41 31654 1 1 3.8 9.03
227 Q14789_REVERSED Golgin subfamily B member 1 OS=Homo sapiens GN=GOLGB1 PE=1 SV=2 - REVERSED 41 375790 1 1 0.3 4.96
228 Q9BXJ4 Complement C1q tumor necrosis factor-related protein 3 OS=Homo sapiens GN=C1QTNF3 PE=2 SV=1 40 26977 1 1 2.8 6.04
229 P25789 Proteasome subunit alpha type-4 OS=Homo sapiens GN=PSMA4 PE=1 SV=1 40 29465 2 2 6.5 7.57
230 Q8WZ42 Titin OS=Homo sapiens GN=TTN PE=1 SV=2 40 3813810 5 1 0.1 6.01
231 P02750 Leucine-rich alpha-2-glycoprotein OS=Homo sapiens GN=LRG1 PE=1 SV=2 40 38154 1 1 1.7 6.45
232 Q9GZR7 ATP-dependent RNA helicase DDX24 OS=Homo sapiens GN=DDX24 PE=1 SV=1 40 96271 1 1 0.7 9.14
233 Q92616_REVERSED Translational activator GCN1 OS=Homo sapiens GN=GCN1L1 PE=1 SV=6 - REVERSED 40 292572 6 1 0.5 7.28
234 Q03923_REVERSED Zinc finger protein 85 OS=Homo sapiens GN=ZNF85 PE=2 SV=3 - REVERSED 40 68691 1 1 1.5 9.45
235 Q8IVF5_REVERSED T-lymphoma invasion and metastasis-inducing protein 2 OS=Homo sapiens GN=TIAM2 PE=2 SV=4 - REVERSED 40 189985 2 1 0.4 6.8
236 A2RRP1_REVERSED Neuroblastoma-amplified sequence OS=Homo sapiens GN=NBAS PE=1 SV=2 - REVERSED 40 268401 1 1 0.3 5.65
237 P55083 Microfibril-associated glycoprotein 4 OS=Homo sapiens GN=MFAP4 PE=1 SV=2 39 28630 1 1 7.1 5.38
238 O14782 Kinesin-like protein KIF3C OS=Homo sapiens GN=KIF3C PE=1 SV=3 39 89371 3 1 1.1 8.28
239 O14556 Glyceraldehyde-3-phosphate dehydrogenase, testis-specific OS=Homo sapiens GN=GAPDHS PE=1 SV=2 39 44473 1 1 3.4 8.38
240 P05156 Complement factor I OS=Homo sapiens GN=CFI PE=1 SV=2 39 65707 1 1 1.2 7.72
241 Q6IWH7 Anoctamin-7 OS=Homo sapiens GN=ANO7 PE=1 SV=1 39 105463 1 1 1 8.33
242 Q14C86_REVERSED GTPase-activating protein and VPS9 domain-containing protein 1 OS=Homo sapiens GN=GAPVD1 PE=1 SV=2 - REVERSED 39 164876 2 1 1.1 5.09
243 P02792 Ferritin light chain OS=Homo sapiens GN=FTL PE=1 SV=2 38 20007 2 1 19.4 5.51
244 Q15022 Polycomb protein SUZ12 OS=Homo sapiens GN=SUZ12 PE=1 SV=3 38 83003 1 1 0.8 8.98
245 P33981 Dual specificity protein kinase TTK OS=Homo sapiens GN=TTK PE=1 SV=2 38 97011 1 1 0.7 8.41
246 P27105 Erythrocyte band 7 integral membrane protein OS=Homo sapiens GN=STOM PE=1 SV=3 38 31711 1 1 4.2 7.71
247 P28066 Proteasome subunit alpha type-5 OS=Homo sapiens GN=PSMA5 PE=1 SV=3 38 26394 1 1 4.1 4.74
248 Q12968_REVERSED Nuclear factor of activated T-cells, cytoplasmic 3 OS=Homo sapiens GN=NFATC3 PE=1 SV=1 - REVERSED 37 115522 3 1 0.7 5.91
249 O00754 Lysosomal alpha-mannosidase OS=Homo sapiens GN=MAN2B1 PE=1 SV=3 37 113672 1 1 1.1 6.84
250 P24347 Stromelysin-3 OS=Homo sapiens GN=MMP11 PE=1 SV=3 36 54556 1 1 1.4 6.38
251 A6H8Y1 Transcription factor TFIIIB component B~~ homolog OS=Homo sapiens GN=BDP1 PE=1 SV=3 36 293705 2 1 0.3 5.06
252 O95071_REVERSED E3 ubiquitin-protein ligase UBR5 OS=Homo sapiens GN=UBR5 PE=1 SV=2 - REVERSED 36 309158 1 1 0.3 5.59
253 Q9UDT6 CAP-Gly domain-containing linker protein 2 OS=Homo sapiens GN=CLIP2 PE=1 SV=1 36 115767 4 1 1.4 6.29
254 Q9C093_REVERSED Sperm flagellar protein 2 OS=Homo sapiens GN=SPEF2 PE=1 SV=2 - REVERSED 36 209680 2 1 0.4 5.4
255 Q9NW08 DNA-directed RNA polymerase III subunit RPC2 OS=Homo sapiens GN=POLR3B PE=1 SV=2 36 127702 2 1 0.6 8.77
256 Q5VWK0 Neuroblastoma breakpoint family member 6 OS=Homo sapiens GN=NBPF6 PE=2 SV=2 36 72194 1 1 1.9 4.84
257 O15264 Mitogen-activated protein kinase 13 OS=Homo sapiens GN=MAPK13 PE=1 SV=1 36 42063 4 1 1.9 8.48
258 P21580_REVERSED Tumor necrosis factor alpha-induced protein 3 OS=Homo sapiens GN=TNFAIP3 PE=1 SV=1 - REVERSED 36 89556 1 1 0.9 8.61
259 P10242_REVERSED Transcriptional activator Myb OS=Homo sapiens GN=MYB PE=1 SV=2 - REVERSED 36 72296 1 1 1.3 6.35
260 Q96IF1 Protein ajuba OS=Homo sapiens GN=JUB PE=1 SV=1 36 56897 2 1 1.5 6.86
261 Q96RD9_REVERSED Fc receptor-like protein 5 OS=Homo sapiens GN=FCRL5 PE=1 SV=3 - REVERSED 36 106371 1 1 0.9 6.72
MCF 10A
prot_hit prot_acc prot_desc prot_score prot_mass prot_matches prot_sequences_sig prot_cover prot_pi
1 P02751 Fibronectin OS=Homo sapiens GN=FN1 PE=1 SV=4 3184 262460 89 44 34.2 5.46
2 P04264 Keratin, type II cytoskeletal 1 OS=Homo sapiens GN=KRT1 PE=1 SV=6 3099 65999 85 33 58.1 8.15
3 P13645 Keratin, type I cytoskeletal 10 OS=Homo sapiens GN=KRT10 PE=1 SV=6 2577 58792 82 31 61.6 5.13
4 P12111 Collagen alpha-3(VI) chain OS=Homo sapiens GN=COL6A3 PE=1 SV=5 2447 343457 101 65 32.7 6.26
5 P35527 Keratin, type I cytoskeletal 9 OS=Homo sapiens GN=KRT9 PE=1 SV=3 2240 62027 64 25 59.9 5.14
6 P02768 Serum albumin OS=Homo sapiens GN=ALB PE=1 SV=2 2090 69321 94 11 11.8 5.92
7 P35908 Keratin, type II cytoskeletal 2 epidermal OS=Homo sapiens GN=KRT2 PE=1 SV=2 1977 65393 59 28 58.5 8.07
8 Q99715 Collagen alpha-1(XII) chain OS=Homo sapiens GN=COL12A1 PE=1 SV=2 1875 332941 84 55 28.3 5.38
9 P01023 Alpha-2-macroglobulin OS=Homo sapiens GN=A2M PE=1 SV=3 1827 163188 60 9 7.7 6.03
10 P07996 Thrombospondin-1 OS=Homo sapiens GN=THBS1 PE=1 SV=2 1802 129300 68 33 38.1 4.71
11 P02765 Alpha-2-HS-glycoprotein OS=Homo sapiens GN=AHSG PE=1 SV=1 1724 39300 54 5 8.7 5.43
12 P20742 Pregnancy zone protein OS=Homo sapiens GN=PZP PE=1 SV=4 1283 163760 43 6 4.3 5.97
13 P01024 Complement C3 OS=Homo sapiens GN=C3 PE=1 SV=2 756 187030 35 17 15.6 6.02
14 P12109 Collagen alpha-1(VI) chain OS=Homo sapiens GN=COL6A1 PE=1 SV=3 726 108462 23 10 18.9 5.26
15 P02533 Keratin, type I cytoskeletal 14 OS=Homo sapiens GN=KRT14 PE=1 SV=4 718 51529 32 14 32.8 5.09
16 P60709 Actin, cytoplasmic 1 OS=Homo sapiens GN=ACTB PE=1 SV=1 676 41710 28 10 47.5 5.29
17 P00734 Prothrombin OS=Homo sapiens GN=F2 PE=1 SV=2 607 69992 17 6 10.5 5.64
18 P08779 Keratin, type I cytoskeletal 16 OS=Homo sapiens GN=KRT16 PE=1 SV=4 585 51236 29 13 32.1 4.99
19 P19823 Inter-alpha-trypsin inhibitor heavy chain H2 OS=Homo sapiens GN=ITIH2 PE=1 SV=2 534 106397 15 7 9.3 6.4
20 P02788 Lactotransferrin OS=Homo sapiens GN=LTF PE=1 SV=6 529 78132 20 3 4.8 8.5
21 Q08380 Galectin-3-binding protein OS=Homo sapiens GN=LGALS3BP PE=1 SV=1 520 65289 20 13 30.4 5.13
22 P24821 Tenascin OS=Homo sapiens GN=TNC PE=1 SV=3 429 240700 21 11 13.8 4.79
23 P04259 Keratin, type II cytoskeletal 6B OS=Homo sapiens GN=KRT6B PE=1 SV=5 426 60030 21 12 21.3 8.09
24 P02538 Keratin, type II cytoskeletal 6A OS=Homo sapiens GN=KRT6A PE=1 SV=3 401 60008 21 12 21.1 8.09
25 P13647 Keratin, type II cytoskeletal 5 OS=Homo sapiens GN=KRT5 PE=1 SV=3 387 62340 23 13 23.7 7.59
26 P0C0L4 Complement C4-A OS=Homo sapiens GN=C4A PE=1 SV=1 356 192650 15 8 5.2 6.65
27 Q15582 Transforming growth factor-beta-induced protein ig-h3 OS=Homo sapiens GN=TGFBI PE=1 SV=1 344 74634 18 10 25.2 7.62
28 P68032 Actin, alpha cardiac muscle 1 OS=Homo sapiens GN=ACTC1 PE=1 SV=1 332 41992 16 6 26.8 5.23
29 P13646 Keratin, type I cytoskeletal 13 OS=Homo sapiens GN=KRT13 PE=1 SV=4 330 49557 16 6 12.9 4.91
30 P06396 Gelsolin OS=Homo sapiens GN=GSN PE=1 SV=1 324 85644 13 5 11.4 5.9
31 P04406 Glyceraldehyde-3-phosphate dehydrogenase OS=Homo sapiens GN=GAPDH PE=1 SV=3 324 36030 10 5 28.4 8.57
32 P68363 Tubulin alpha-1B chain OS=Homo sapiens GN=TUBA1B PE=1 SV=1 318 50120 14 10 28.8 4.94
33 Q04695 Keratin, type I cytoskeletal 17 OS=Homo sapiens GN=KRT17 PE=1 SV=2 309 48076 19 8 22 4.97
34 P02771 Alpha-fetoprotein OS=Homo sapiens GN=AFP PE=1 SV=1 303 68633 10 4 6.7 5.48
35 P69905 Hemoglobin subunit alpha OS=Homo sapiens GN=HBA1 PE=1 SV=2 291 15248 14 4 33.1 8.72
36 P02774 Vitamin D-binding protein OS=Homo sapiens GN=GC PE=1 SV=1 282 52929 11 5 11.6 5.4
37 Q00610 Clathrin heavy chain 1 OS=Homo sapiens GN=CLTC PE=1 SV=5 280 191493 7 6 6.2 5.48
38 P19012 Keratin, type I cytoskeletal 15 OS=Homo sapiens GN=KRT15 PE=1 SV=3 256 49181 17 6 12.1 4.71
39 P36955 Pigment epithelium-derived factor OS=Homo sapiens GN=SERPINF1 PE=1 SV=4 254 46283 6 3 9.1 5.97
40 P02452 Collagen alpha-1(I) chain OS=Homo sapiens GN=COL1A1 PE=1 SV=5 243 138857 10 6 9.6 5.6
41 P04114 Apolipoprotein B-100 OS=Homo sapiens GN=APOB PE=1 SV=2 237 515283 15 9 2.2 6.58
42 P12110 Collagen alpha-2(VI) chain OS=Homo sapiens GN=COL6A2 PE=1 SV=4 235 108512 15 7 15.3 5.85
43 P07355 Annexin A2 OS=Homo sapiens GN=ANXA2 PE=1 SV=2 220 38580 7 6 23.9 7.57
44 Q562R1 Beta-actin-like protein 2 OS=Homo sapiens GN=ACTBL2 PE=1 SV=2 213 41976 10 3 16 5.39
45 P68366 Tubulin alpha-4A chain OS=Homo sapiens GN=TUBA4A PE=1 SV=1 208 49892 10 7 19 4.95
46 P04004 Vitronectin OS=Homo sapiens GN=VTN PE=1 SV=1 196 54271 6 1 4.2 5.55
47 P23142 Fibulin-1 OS=Homo sapiens GN=FBLN1 PE=1 SV=4 180 77162 7 4 5.7 5.07
48 P07437 Tubulin beta chain OS=Homo sapiens GN=TUBB PE=1 SV=2 180 49639 8 5 19.6 4.78
49 Q5XKE5 Keratin, type II cytoskeletal 79 OS=Homo sapiens GN=KRT79 PE=1 SV=2 179 57800 8 4 6.9 6.75
50 P62805 Histone H4 OS=Homo sapiens GN=HIST1H4A PE=1 SV=2 173 11360 7 5 50.5 11.36
51 P02042 Hemoglobin subunit delta OS=Homo sapiens GN=HBD PE=1 SV=2 171 16045 10 2 12.9 7.85
52 P05787 Keratin, type II cytoskeletal 8 OS=Homo sapiens GN=KRT8 PE=1 SV=7 170 53671 7 4 5.8 5.52
53 Q07954 Prolow-density lipoprotein receptor-related protein 1 OS=Homo sapiens GN=LRP1 PE=1 SV=2 167 504276 9 4 2.6 5.16
54 P49747 Cartilage oligomeric matrix protein OS=Homo sapiens GN=COMP PE=1 SV=2 165 82808 6 4 5.8 4.36
55 P02647 Apolipoprotein A-I OS=Homo sapiens GN=APOA1 PE=1 SV=1 164 30759 4 1 8.2 5.56
56 P19013 Keratin, type II cytoskeletal 4 OS=Homo sapiens GN=KRT4 PE=1 SV=4 163 57250 8 4 7.3 6.25
57 P08697 Alpha-2-antiplasmin OS=Homo sapiens GN=SERPINF2 PE=1 SV=3 145 54531 2 1 2.2 5.87
58 P01008 Antithrombin-III OS=Homo sapiens GN=SERPINC1 PE=1 SV=1 144 52569 11 5 10.3 6.32
59 P68371 Tubulin beta-2C chain OS=Homo sapiens GN=TUBB2C PE=1 SV=1 140 49799 7 4 16.2 4.79
60 Q06033 Inter-alpha-trypsin inhibitor heavy chain H3 OS=Homo sapiens GN=ITIH3 PE=1 SV=2 139 99787 6 3 4.2 5.49
61 P12259 Coagulation factor V OS=Homo sapiens GN=F5 PE=1 SV=4 135 251546 8 4 3.2 5.68
62 P23526 Adenosylhomocysteinase OS=Homo sapiens GN=AHCY PE=1 SV=4 133 47685 3 2 9 5.92
63 P51884 Lumican OS=Homo sapiens GN=LUM PE=1 SV=2 130 38405 6 2 9.8 6.16
64 Q8NI35 InaD-like protein OS=Homo sapiens GN=INADL PE=1 SV=3 129 196247 11 1 0.3 4.84
65 P19827 Inter-alpha-trypsin inhibitor heavy chain H1 OS=Homo sapiens GN=ITIH1 PE=1 SV=3 120 101326 3 2 3.8 6.31
66 Q05707 Collagen alpha-1(XIV) chain OS=Homo sapiens GN=COL14A1 PE=1 SV=3 119 193394 11 4 7.6 5.16
67 P12035 Keratin, type II cytoskeletal 3 OS=Homo sapiens GN=KRT3 PE=1 SV=3 118 64378 6 4 5.6 6.12
68 P15169 Carboxypeptidase N catalytic chain OS=Homo sapiens GN=CPN1 PE=1 SV=1 116 52253 2 1 3.3 6.86
69 P02675 Fibrinogen beta chain OS=Homo sapiens GN=FGB PE=1 SV=2 114 55892 5 3 5.3 8.54
70 P02748 Complement component C9 OS=Homo sapiens GN=C9 PE=1 SV=2 113 63133 2 2 3.8 5.43
71 P11717 Cation-independent mannose-6-phosphate receptor OS=Homo sapiens GN=IGF2R PE=1 SV=3 111 274199 4 2 0.8 5.64
72 P00747 Plasminogen OS=Homo sapiens GN=PLG PE=1 SV=2 110 90510 6 2 4.3 7.04
73 P81605 Dermcidin OS=Homo sapiens GN=DCD PE=1 SV=2 110 11277 4 3 22.7 6.08
74 P05543 Thyroxine-binding globulin OS=Homo sapiens GN=SERPINA7 PE=1 SV=2 105 46295 4 2 4.8 5.87
75 P0C0S8 Histone H2A type 1 OS=Homo sapiens GN=HIST1H2AG PE=1 SV=2 105 14083 3 2 21.5 10.9
76 Q86YZ3 Hornerin OS=Homo sapiens GN=HRNR PE=1 SV=2 102 282228 2 2 1.3 10.05
77 P02749 Beta-2-glycoprotein 1 OS=Homo sapiens GN=APOH PE=1 SV=3 101 38273 4 1 5.2 8.34
78 Q08431 Lactadherin OS=Homo sapiens GN=MFGE8 PE=1 SV=2 101 43095 2 1 3.9 8.47
79 P01031 Complement C5 OS=Homo sapiens GN=C5 PE=1 SV=4 101 188186 6 2 2.6 6.11
80 Q5D862 Filaggrin-2 OS=Homo sapiens GN=FLG2 PE=1 SV=1 100 247928 2 2 1 8.45
81 P07195 L-lactate dehydrogenase B chain OS=Homo sapiens GN=LDHB PE=1 SV=2 99 36615 2 2 6 5.71
82 P01009 Alpha-1-antitrypsin OS=Homo sapiens GN=SERPINA1 PE=1 SV=3 98 46707 4 1 3.3 5.37
83 P55072 Transitional endoplasmic reticulum ATPase OS=Homo sapiens GN=VCP PE=1 SV=4 96 89266 5 2 8.8 5.14
84 O43854 EGF-like repeat and discoidin I-like domain-containing protein 3 OS=Homo sapiens GN=EDIL3 PE=1 SV=1 93 53730 4 4 7.5 7.08
85 P08238 Heat shock protein HSP 90-beta OS=Homo sapiens GN=HSP90AB1 PE=1 SV=4 92 83212 3 2 4.1 4.97
86 O14818 Proteasome subunit alpha type-7 OS=Homo sapiens GN=PSMA7 PE=1 SV=1 91 27870 3 3 13.7 8.6
87 O15111_REVERSED Inhibitor of nuclear factor kappa-B kinase subunit alpha OS=Homo sapiens GN=CHUK PE=1 SV=2 - REVERSED 87 84585 7 1 1.6 6.28
88 P08107 Heat shock 70 kDa protein 1A/1B OS=Homo sapiens GN=HSPA1A PE=1 SV=5 82 70009 4 3 8.3 5.48
89 Q86Y46 Keratin, type II cytoskeletal 73 OS=Homo sapiens GN=KRT73 PE=1 SV=1 81 58887 6 4 6.7 6.93
90 P14618 Pyruvate kinase isozymes M1/M2 OS=Homo sapiens GN=PKM2 PE=1 SV=4 81 57900 6 3 14.1 7.96
91 P02461 Collagen alpha-1(III) chain OS=Homo sapiens GN=COL3A1 PE=1 SV=4 80 138479 3 2 2.3 6.21
92 P04075 Fructose-bisphosphate aldolase A OS=Homo sapiens GN=ALDOA PE=1 SV=2 80 39395 4 2 10.7 8.3
93 P61204 ADP-ribosylation factor 3 OS=Homo sapiens GN=ARF3 PE=1 SV=2 79 20588 1 1 8.3 6.84
94 Q14764 Major vault protein OS=Homo sapiens GN=MVP PE=1 SV=4 77 99266 4 3 5.6 5.34
95 P02787 Serotransferrin OS=Homo sapiens GN=TF PE=1 SV=3 77 77014 3 2 1.7 6.81
96 Q5QNW6 Histone H2B type 2-F OS=Homo sapiens GN=HIST2H2BF PE=1 SV=3 76 13912 3 2 15.9 10.31
97 P07900 Heat shock protein HSP 90-alpha OS=Homo sapiens GN=HSP90AA1 PE=1 SV=5 76 84607 3 2 4.9 4.94
98 Q8N7M0_REVERSED Tctex1 domain-containing protein 1 OS=Homo sapiens GN=TCTEX1D1 PE=2 SV=2 - REVERSED 75 20717 13 1 3.9 9.04
99 P04083 Annexin A1 OS=Homo sapiens GN=ANXA1 PE=1 SV=2 74 38690 2 1 8.4 6.57
100 P11142 Heat shock cognate 71 kDa protein OS=Homo sapiens GN=HSPA8 PE=1 SV=1 73 70854 4 2 8.7 5.37
101 P04180 Phosphatidylcholine-sterol acyltransferase OS=Homo sapiens GN=LCAT PE=1 SV=1 70 49546 1 1 2.5 5.71
102 P06702 Protein S100-A9 OS=Homo sapiens GN=S100A9 PE=1 SV=1 70 13234 3 1 24.6 5.71
103 Q9BXJ4 Complement C1q tumor necrosis factor-related protein 3 OS=Homo sapiens GN=C1QTNF3 PE=2 SV=1 68 26977 2 2 8.1 6.04
104 P02753 Retinol-binding protein 4 OS=Homo sapiens GN=RBP4 PE=1 SV=3 66 22995 2 1 16.4 5.76
105 P29353_REVERSED SHC-transforming protein 1 OS=Homo sapiens GN=SHC1 PE=1 SV=4 - REVERSED 66 62782 11 1 1.4 6.01
106 Q08554 Desmocollin-1 OS=Homo sapiens GN=DSC1 PE=1 SV=2 66 99924 1 1 1.7 5.25
107 Q16853 Membrane primary amine oxidase OS=Homo sapiens GN=AOC3 PE=1 SV=3 66 84568 1 1 1.2 6.05
108 P00338 L-lactate dehydrogenase A chain OS=Homo sapiens GN=LDHA PE=1 SV=2 66 36665 2 1 6 8.44
109 P00488 Coagulation factor XIII A chain OS=Homo sapiens GN=F13A1 PE=1 SV=4 65 83215 1 1 1.2 5.75
110 Q5VTE0 Putative elongation factor 1-alpha-like 3 OS=Homo sapiens GN=EEF1A1P5 PE=5 SV=1 64 50153 3 2 5.2 9.15
111 Q9H4B7 Tubulin beta-1 chain OS=Homo sapiens GN=TUBB1 PE=1 SV=1 64 50295 3 1 9.5 5.05
112 P62979 Ubiquitin-40S ribosomal protein S27a OS=Homo sapiens GN=RPS27A PE=1 SV=2 63 17953 1 1 10.3 9.68
113 P35443 Thrombospondin-4 OS=Homo sapiens GN=THBS4 PE=1 SV=2 62 105802 5 2 3.9 4.44
114 P02760 Protein AMBP OS=Homo sapiens GN=AMBP PE=1 SV=1 61 38974 2 1 2 5.95
115 P21333 Filamin-A OS=Homo sapiens GN=FLNA PE=1 SV=4 60 280564 3 2 1.3 5.7
116 P35579 Myosin-9 OS=Homo sapiens GN=MYH9 PE=1 SV=4 59 226392 2 1 1.5 5.5
117 Q15063 Periostin OS=Homo sapiens GN=POSTN PE=1 SV=2 59 93255 3 1 5.9 7.27
118 P02649 Apolipoprotein E OS=Homo sapiens GN=APOE PE=1 SV=1 59 36132 4 2 8.2 5.65
119 Q14624 Inter-alpha-trypsin inhibitor heavy chain H4 OS=Homo sapiens GN=ITIH4 PE=1 SV=4 59 103293 3 1 2.5 6.51
120 P08758 Annexin A5 OS=Homo sapiens GN=ANXA5 PE=1 SV=2 58 35914 4 2 10.6 4.94
121 P02786 Transferrin receptor protein 1 OS=Homo sapiens GN=TFRC PE=1 SV=2 58 84818 1 1 1.6 6.18
122 A6NIZ1 Ras-related protein Rap-1b-like protein OS=Homo sapiens PE=2 SV=1 57 20912 1 1 6.5 5.37
123 Q93070 Ecto-ADP-ribosyltransferase 4 OS=Homo sapiens GN=ART4 PE=2 SV=2 57 35854 8 1 2.2 9.31
124 Q9Y240 C-type lectin domain family 11 member A OS=Homo sapiens GN=CLEC11A PE=1 SV=1 57 35672 1 1 5.3 5.06
125 P32119 Peroxiredoxin-2 OS=Homo sapiens GN=PRDX2 PE=1 SV=5 55 21878 1 1 9.1 5.66
126 Q99456 Keratin, type I cytoskeletal 12 OS=Homo sapiens GN=KRT12 PE=1 SV=1 55 53478 4 1 3.2 4.7
127 P01591 Immunoglobulin J chain OS=Homo sapiens GN=IGJ PE=1 SV=4 54 18087 1 1 6.3 5.12
128 Q6ZU45 Putative C-type lectin domain-containing protein NCRNA00083 OS=Homo sapiens GN=NCRNA00083 PE=5 SV=1 53 25796 1 1 3.4 6.3
129 Q86TI0_REVERSED TBC1 domain family member 1 OS=Homo sapiens GN=TBC1D1 PE=1 SV=2 - REVERSED 53 133000 1 1 0.7 6.52
130 Q99436 Proteasome subunit beta type-7 OS=Homo sapiens GN=PSMB7 PE=1 SV=1 53 29946 1 1 3.6 7.57
131 P15144 Aminopeptidase N OS=Homo sapiens GN=ANPEP PE=1 SV=4 53 109471 3 1 3.8 5.31
132 P28066 Proteasome subunit alpha type-5 OS=Homo sapiens GN=PSMA5 PE=1 SV=3 52 26394 2 1 12 4.74
133 Q07092 Collagen alpha-1(XVI) chain OS=Homo sapiens GN=COL16A1 PE=1 SV=2 52 157653 1 1 0.7 8.14
134 Q9BYX7_REVERSED Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 - REVERSED 51 41989 3 1 3.5 5.92
135 P05452 Tetranectin OS=Homo sapiens GN=CLEC3B PE=1 SV=3 51 22522 1 1 5.9 5.52
136 P35125 Ubiquitin carboxyl-terminal hydrolase 6 OS=Homo sapiens GN=USP6 PE=1 SV=2 50 158557 11 1 0.4 7.87
137 O95445 Apolipoprotein M OS=Homo sapiens GN=APOM PE=1 SV=2 50 21239 2 1 6.9 5.66
138 P02790 Hemopexin OS=Homo sapiens GN=HPX PE=1 SV=2 50 51643 4 1 3.2 6.55
139 Q13454 Tumor suppressor candidate 3 OS=Homo sapiens GN=TUSC3 PE=2 SV=1 50 39650 2 1 2.3 9.93
140 Q8NA29_REVERSED Major facilitator superfamily domain-containing protein 2A OS=Homo sapiens GN=MFSD2A PE=2 SV=1 - REVERSED 50 60131 1 1 1.3 6.55
141 Q9TNN7 HLA class I histocompatibility antigen, Cw-5 alpha chain OS=Homo sapiens GN=HLA-C PE=2 SV=1 49 40886 1 1 3.6 7.11
142 P12036 Neurofilament heavy polypeptide OS=Homo sapiens GN=NEFH PE=1 SV=4 49 112411 2 1 1.6 5.99
143 Q8N1N4 Keratin, type II cytoskeletal 78 OS=Homo sapiens GN=KRT78 PE=1 SV=2 49 56830 2 1 2.7 5.79
144 Q9Y490 Talin-1 OS=Homo sapiens GN=TLN1 PE=1 SV=3 49 269599 1 1 0.6 5.77
145 Q9UBR2 Cathepsin Z OS=Homo sapiens GN=CTSZ PE=1 SV=1 49 33846 1 1 3.3 6.7
146 P49720 Proteasome subunit beta type-3 OS=Homo sapiens GN=PSMB3 PE=1 SV=2 48 22933 2 1 12.2 6.14
147 P05546 Heparin cofactor 2 OS=Homo sapiens GN=SERPIND1 PE=1 SV=3 48 57034 1 1 2.4 6.41
148 P01042 Kininogen-1 OS=Homo sapiens GN=KNG1 PE=1 SV=2 48 71912 1 1 1.4 6.34
149 Q96Q89_REVERSED Kinesin-like protein KIF20B OS=Homo sapiens GN=KIF20B PE=1 SV=3 - REVERSED 48 210500 3 1 0.5 5.54
150 P03956 Interstitial collagenase OS=Homo sapiens GN=MMP1 PE=1 SV=3 48 53973 2 1 3 6.47
151 Q8NDM7_REVERSED WD repeat-containing protein 96 OS=Homo sapiens GN=WDR96 PE=2 SV=3 - REVERSED 47 191861 1 1 0.5 5.71
152 Q71DI3 Histone H3.2 OS=Homo sapiens GN=HIST2H3A PE=1 SV=3 46 15379 4 1 14.7 11.27
153 P13639 Elongation factor 2 OS=Homo sapiens GN=EEF2 PE=1 SV=4 46 95277 1 1 1.2 6.41
154 Q9Y6X9_REVERSED MORC family CW-type zinc finger protein 2 OS=Homo sapiens GN=MORC2 PE=1 SV=2 - REVERSED 46 117750 1 1 0.7 8.6
155 P28074 Proteasome subunit beta type-5 OS=Homo sapiens GN=PSMB5 PE=1 SV=3 46 28462 1 1 4.6 6.43
156 Q9ULC6 Protein-arginine deiminase type-1 OS=Homo sapiens GN=PADI1 PE=1 SV=2 46 74618 1 1 1.4 6.07
157 Q8NBM4 Ubiquitin-associated domain-containing protein 2 OS=Homo sapiens GN=UBAC2 PE=2 SV=1 46 38938 3 1 2.3 9.21
158 Q2VWP7 Protogenin OS=Homo sapiens GN=PRTG PE=2 SV=1 46 126996 1 1 0.6 7.37
159 Q9NPY3_REVERSED Complement component C1q receptor OS=Homo sapiens GN=CD93 PE=1 SV=3 - REVERSED 45 68515 1 1 1.1 5.27
160 Q9NRY4 Glucocorticoid receptor DNA-binding factor 1 OS=Homo sapiens GN=GRLF1 PE=1 SV=2 45 172121 1 1 0.5 6.33
161 Q9NU02_REVERSED Ankyrin repeat domain-containing protein 5 OS=Homo sapiens GN=ANKRD5 PE=2 SV=2 - REVERSED 45 86610 5 1 1.8 8.51
162 B9A064 Immunoglobulin lambda-like polypeptide 5 OS=Homo sapiens GN=IGLL5 PE=2 SV=2 45 23049 1 1 3.7 9.08
163 Q5T764_REVERSED Interferon-induced protein with tetratricopeptide repeats 1B OS=Homo sapiens GN=IFIT1B PE=2 SV=1 - REVERSED 45 54958 2 1 3.4 7.69
164 P07339 Cathepsin D OS=Homo sapiens GN=CTSD PE=1 SV=1 45 44524 1 1 1.9 6.1
165 P02794 Ferritin heavy chain OS=Homo sapiens GN=FTH1 PE=1 SV=2 45 21212 2 1 9.3 5.3
166 Q9UKT8_REVERSED F-box/WD repeat-containing protein 2 OS=Homo sapiens GN=FBXW2 PE=1 SV=2 - REVERSED 45 51479 2 1 3.1 6.11
167 P29803_REVERSED Pyruvate dehydrogenase E1 component subunit alpha, testis-specific form, mitochondrial OS=Homo sapiens GN=PDHA2 PE=1 SV=1 - REVERSED 44 42906 3 1 4.1 8.76
168 Q9BY43 Charged multivesicular body protein 4a OS=Homo sapiens GN=CHMP4A PE=1 SV=3 44 25083 3 1 6.3 4.65
169 Q00013 55 kDa erythrocyte membrane protein OS=Homo sapiens GN=MPP1 PE=1 SV=2 44 52264 2 1 1.5 6.91
170 Q13477_REVERSED Mucosal addressin cell adhesion molecule 1 OS=Homo sapiens GN=MADCAM1 PE=1 SV=2 - REVERSED 43 40130 2 1 2.6 5
171 Q8NFM7_REVERSED Interleukin-17 receptor D OS=Homo sapiens GN=IL17RD PE=1 SV=3 - REVERSED 43 82358 1 1 0.9 6.81
172 Q92609 TBC1 domain family member 5 OS=Homo sapiens GN=TBC1D5 PE=1 SV=1 43 88949 1 1 1 6.1
173 Q96CD0_REVERSED F-box/LRR-repeat protein 8 OS=Homo sapiens GN=FBXL8 PE=1 SV=1 - REVERSED 43 40490 9 1 1.9 7.03
174 P04745 Alpha-amylase 1 OS=Homo sapiens GN=AMY1A PE=1 SV=2 43 57731 1 1 2.3 6.47
175 Q9NZM4 Glioma tumor suppressor candidate region gene 1 protein OS=Homo sapiens GN=GLTSCR1 PE=1 SV=2 42 158393 2 1 0.5 6.17
176 P10643 Complement component C7 OS=Homo sapiens GN=C7 PE=1 SV=2 42 93457 2 1 1.1 6.09
177 Q6ZUT9_REVERSED DENN domain-containing protein 5B OS=Homo sapiens GN=DENND5B PE=1 SV=2 - REVERSED 41 144927 2 1 0.5 6.29
178 Q86UQ0_REVERSED Zinc finger protein 589 OS=Homo sapiens GN=ZNF589 PE=1 SV=1 - REVERSED 41 41163 2 1 1.6 8.62
179 P43652 Afamin OS=Homo sapiens GN=AFM PE=1 SV=1 41 69024 1 1 1.2 5.64
180 Q96RD9_REVERSED Fc receptor-like protein 5 OS=Homo sapiens GN=FCRL5 PE=1 SV=3 - REVERSED 41 106371 1 1 0.9 6.72
181 Q16563 Synaptophysin-like protein 1 OS=Homo sapiens GN=SYPL1 PE=1 SV=1 41 28547 1 1 4.2 8.69
182 Q58FF6 Putative heat shock protein HSP 90-beta 4 OS=Homo sapiens GN=HSP90AB4P PE=5 SV=1 40 58228 2 1 5 4.65
183 Q9ULT0 Tetratricopeptide repeat protein 7A OS=Homo sapiens GN=TTC7A PE=1 SV=3 40 96123 3 1 1.2 6.03
184 P98160 Basement membrane-specific heparan sulfate proteoglycan core protein OS=Homo sapiens GN=HSPG2 PE=1 SV=4 39 468532 1 1 0.2 6.06
185 O14782 Kinesin-like protein KIF3C OS=Homo sapiens GN=KIF3C PE=1 SV=3 39 89371 3 1 1.1 8.28
186 Q99816 Tumor susceptibility gene 101 protein OS=Homo sapiens GN=TSG101 PE=1 SV=2 60 43916 3 1 2.4 9.4
187 P11166 Solute carrier family 2, facilitated glucose transporter member 1 OS=Homo sapiens GN=SLC2A1 PE=1 SV=2 39 54049 1 1 1.6 8.93
188 Q00535_REVERSED Cyclin-dependent kinase 5 OS=Homo sapiens GN=CDK5 PE=1 SV=3 - REVERSED 39 33283 2 1 3.1 7.9
189 Q8N3E9 1-phosphatidylinositol-4,5-bisphosphate phosphodiesterase delta-3 OS=Homo sapiens GN=PLCD3 PE=1 SV=3 39 89202 5 1 1.9 6.52
190 P43235 Cathepsin K OS=Homo sapiens GN=CTSK PE=1 SV=1 38 36942 1 1 6.1 8.72
191 P22303 Acetylcholinesterase OS=Homo sapiens GN=ACHE PE=1 SV=1 38 67753 1 1 2.1 5.87
192 Q9P2G4_REVERSED Uncharacterized protein KIAA1383 OS=Homo sapiens GN=KIAA1383 PE=1 SV=2 - REVERSED 38 100283 1 1 0.8 6.77
193 Q9BZP6 Acidic mammalian chitinase OS=Homo sapiens GN=CHIA PE=1 SV=1 38 52237 1 1 1.7 5.54
194 Q9ULI3_REVERSED Protein HEG homolog 1 OS=Homo sapiens GN=HEG1 PE=1 SV=3 - REVERSED 38 147369 5 1 0.9 5.81
195 O43264_REVERSED Centromere/kinetochore protein zw10 homolog OS=Homo sapiens GN=ZW10 PE=1 SV=3 - REVERSED 38 88773 1 1 0.8 5.89
196 P48436_REVERSED Transcription factor SOX-9 OS=Homo sapiens GN=SOX9 PE=1 SV=1 - REVERSED 38 56102 1 1 1.6 6.33
197 Q5TAH2 Sodium/hydrogen exchanger 11 OS=Homo sapiens GN=SLC9A11 PE=2 SV=1 38 128969 1 1 0.6 6.5
198 Q03518 Antigen peptide transporter 1 OS=Homo sapiens GN=TAP1 PE=1 SV=2 37 87163 10 1 1.9 8.24
199 P06746 DNA polymerase beta OS=Homo sapiens GN=POLB PE=1 SV=3 37 38154 1 1 1.8 9.01
200 Q7Z5M8 Abhydrolase domain-containing protein 12B OS=Homo sapiens GN=ABHD12B PE=2 SV=1 37 40750 3 1 1.7 8.57
201 Q15020_REVERSED Squamous cell carcinoma antigen recognized by T-cells 3 OS=Homo sapiens GN=SART3 PE=1 SV=1 - REVERSED 37 109865 3 1 0.6 5.45
202 Q9NX14 NADH dehydrogenase [ubiquinone] 1 beta subcomplex subunit 11, mitochondrial OS=Homo sapiens GN=NDUFB11 PE=1 SV=1 37 17306 1 1 5.2 5.17
203 Q9H115_REVERSED Beta-soluble NSF attachment protein OS=Homo sapiens GN=NAPB PE=1 SV=2 - REVERSED 37 33535 1 1 4 5.32
204 Q14573_REVERSED Inositol 1,4,5-trisphosphate receptor type 3 OS=Homo sapiens GN=ITPR3 PE=1 SV=2 - REVERSED 37 303912 3 1 0.4 6.05
205 P00742 Coagulation factor X OS=Homo sapiens GN=F10 PE=1 SV=2 37 54697 1 1 1.8 5.68
206 P26006 Integrin alpha-3 OS=Homo sapiens GN=ITGA3 PE=1 SV=4 37 118680 1 1 0.9 6.51
207 Q6DHV7_REVERSED Adenosine deaminase-like protein OS=Homo sapiens GN=ADAL PE=2 SV=2 - REVERSED 37 40239 1 1 3.1 5.89
208 P21462_REVERSED fMet-Leu-Phe receptor OS=Homo sapiens GN=FPR1 PE=1 SV=3 - REVERSED 37 38420 2 1 2 9.23
209 Q9UGM3 Deleted in malignant brain tumors 1 protein OS=Homo sapiens GN=DMBT1 PE=1 SV=2 36 260569 1 1 0.2 5.18
210 Q14CM0_REVERSED FERM and PDZ domain-containing protein 4 OS=Homo sapiens GN=FRMPD4 PE=1 SV=1 - REVERSED 36 144289 3 1 0.6 5.12
211 O00754 Lysosomal alpha-mannosidase OS=Homo sapiens GN=MAN2B1 PE=1 SV=3 36 113672 1 1 1.1 6.84
212 P05109 Protein S100-A8 OS=Homo sapiens GN=S100A8 PE=1 SV=1 36 10828 1 1 7.5 6.51
213 O75691_REVERSED Small subunit processome component 20 homolog OS=Homo sapiens GN=UTP20 PE=1 SV=3 - REVERSED 36 318182 1 1 0.3 7.09
214 Q96ED9_REVERSED Protein Hook homolog 2 OS=Homo sapiens GN=HOOK2 PE=1 SV=3 - REVERSED 36 83155 3 1 1.5 5.36
MDA-MB-231
prot_hit prot_acc prot_desc prot_score prot_mass prot_matches prot_sequences_sig prot_cover prot_pi
1 P04264 Keratin, type II cytoskeletal 1 OS=Homo sapiens GN=KRT1 PE=1 SV=6 3363 65999 84 31 57.3 8.15
2 P13645 Keratin, type I cytoskeletal 10 OS=Homo sapiens GN=KRT10 PE=1 SV=6 2875 58792 85 31 58.7 5.13
3 P02751 Fibronectin OS=Homo sapiens GN=FN1 PE=1 SV=4 2436 262460 88 40 25.6 5.46
4 P02765 Alpha-2-HS-glycoprotein OS=Homo sapiens GN=AHSG PE=1 SV=1 2048 39300 52 4 7.1 5.43
5 P35527 Keratin, type I cytoskeletal 9 OS=Homo sapiens GN=KRT9 PE=1 SV=3 1994 62027 55 23 65.7 5.14
6 P35908 Keratin, type II cytoskeletal 2 epidermal OS=Homo sapiens GN=KRT2 PE=1 SV=2 1865 65393 55 25 57.9 8.07
7 P02768 Serum albumin OS=Homo sapiens GN=ALB PE=1 SV=2 1781 69321 112 8 11.8 5.92
8 P01023 Alpha-2-macroglobulin OS=Homo sapiens GN=A2M PE=1 SV=3 1613 163188 57 6 5.2 6.03
9 P20742 Pregnancy zone protein OS=Homo sapiens GN=PZP PE=1 SV=4 1346 163760 41 4 3.8 5.97
10 P55072 Transitional endoplasmic reticulum ATPase OS=Homo sapiens GN=VCP PE=1 SV=4 906 89266 35 21 37.1 5.14
11 Q5QNW6 Histone H2B type 2-F OS=Homo sapiens GN=HIST2H2BF PE=1 SV=3 881 13912 22 5 35.7 10.31
12 P60709 Actin, cytoplasmic 1 OS=Homo sapiens GN=ACTB PE=1 SV=1 647 41710 25 13 47.5 5.29
13 P00734 Prothrombin OS=Homo sapiens GN=F2 PE=1 SV=2 613 69992 15 6 9.5 5.64
14 P07996 Thrombospondin-1 OS=Homo sapiens GN=THBS1 PE=1 SV=2 580 129300 27 13 13.2 4.71
15 P02533 Keratin, type I cytoskeletal 14 OS=Homo sapiens GN=KRT14 PE=1 SV=4 575 51529 25 15 32.4 5.09
16 P01024 Complement C3 OS=Homo sapiens GN=C3 PE=1 SV=2 509 187030 23 11 8.8 6.02
17 P19823 Inter-alpha-trypsin inhibitor heavy chain H2 OS=Homo sapiens GN=ITIH2 PE=1 SV=2 468 106397 15 8 9.3 6.4
18 Q6FI13 Histone H2A type 2-A OS=Homo sapiens GN=HIST2H2AA3 PE=1 SV=3 456 14087 14 4 49.2 10.9
19 P13647 Keratin, type II cytoskeletal 5 OS=Homo sapiens GN=KRT5 PE=1 SV=3 450 62340 22 13 25.3 7.59
20 P07437 Tubulin beta chain OS=Homo sapiens GN=TUBB PE=1 SV=2 441 49639 20 13 45 4.78
21 Q07954 Prolow-density lipoprotein receptor-related protein 1 OS=Homo sapiens GN=LRP1 PE=1 SV=2 439 504276 15 8 3.6 5.16
22 P98160 Basement membrane-specific heparan sulfate proteoglycan core protein OS=Homo sapiens GN=HSPG2 PE=1 SV=4 439 468532 20 13 4.7 6.06
23 P08779 Keratin, type I cytoskeletal 16 OS=Homo sapiens GN=KRT16 PE=1 SV=4 394 51236 19 10 18 4.99
24 P02788 Lactotransferrin OS=Homo sapiens GN=LTF PE=1 SV=6 380 78132 16 3 4.8 8.5
25 P0C0L4 Complement C4-A OS=Homo sapiens GN=C4A PE=1 SV=1 367 192650 15 7 5.6 6.65
26 P62805 Histone H4 OS=Homo sapiens GN=HIST1H4A PE=1 SV=2 363 11360 15 6 51.5 11.36
27 P04259 Keratin, type II cytoskeletal 6B OS=Homo sapiens GN=KRT6B PE=1 SV=5 349 60030 15 8 14 8.09
28 P11142 Heat shock cognate 71 kDa protein OS=Homo sapiens GN=HSPA8 PE=1 SV=1 346 70854 12 7 16.7 5.37
29 Q86YZ3 Hornerin OS=Homo sapiens GN=HRNR PE=1 SV=2 344 282228 3 3 2.1 10.05
30 P02538 Keratin, type II cytoskeletal 6A OS=Homo sapiens GN=KRT6A PE=1 SV=3 339 60008 15 8 15.6 8.09
31 P68032 Actin, alpha cardiac muscle 1 OS=Homo sapiens GN=ACTC1 PE=1 SV=1 333 41992 17 9 26.8 5.23
32 P02771 Alpha-fetoprotein OS=Homo sapiens GN=AFP PE=1 SV=1 310 68633 9 3 5.6 5.48
33 Q9BQE3 Tubulin alpha-1C chain OS=Homo sapiens GN=TUBA1C PE=1 SV=1 285 49863 18 6 31.8 4.96
34 P02774 Vitamin D-binding protein OS=Homo sapiens GN=GC PE=1 SV=1 277 52929 10 6 11.8 5.4
35 P04004 Vitronectin OS=Homo sapiens GN=VTN PE=1 SV=1 277 54271 6 1 4.2 5.55
36 Q9BVA1 Tubulin beta-2B chain OS=Homo sapiens GN=TUBB2B PE=1 SV=1 270 49921 17 10 36.9 4.78
37 Q04695 Keratin, type I cytoskeletal 17 OS=Homo sapiens GN=KRT17 PE=1 SV=2 257 48076 16 9 15 4.97
38 P68366 Tubulin alpha-4A chain OS=Homo sapiens GN=TUBA4A PE=1 SV=1 256 49892 13 5 22.5 4.95
39 P13646 Keratin, type I cytoskeletal 13 OS=Homo sapiens GN=KRT13 PE=1 SV=4 254 49557 12 6 9.2 4.91
40 P69905 Hemoglobin subunit alpha OS=Homo sapiens GN=HBA1 PE=1 SV=2 231 15248 8 3 33.1 8.72
41 P04114 Apolipoprotein B-100 OS=Homo sapiens GN=APOB PE=1 SV=2 227 515283 14 7 1.9 6.58
42 P04406 Glyceraldehyde-3-phosphate dehydrogenase OS=Homo sapiens GN=GAPDH PE=1 SV=3 222 36030 5 2 17.6 8.57
43 Q5XKE5 Keratin, type II cytoskeletal 79 OS=Homo sapiens GN=KRT79 PE=1 SV=2 221 57800 10 4 6.9 6.75
44 P23142 Fibulin-1 OS=Homo sapiens GN=FBLN1 PE=1 SV=4 214 77162 11 4 7.3 5.07
45 P08238 Heat shock protein HSP 90-beta OS=Homo sapiens GN=HSP90AB1 PE=1 SV=4 193 83212 8 4 10.4 4.97
46 Q562R1 Beta-actin-like protein 2 OS=Homo sapiens GN=ACTBL2 PE=1 SV=2 189 41976 12 4 16 5.39
47 P06396 Gelsolin OS=Homo sapiens GN=GSN PE=1 SV=1 185 85644 6 5 9.5 5.9
48 Q96QV6 Histone H2A type 1-A OS=Homo sapiens GN=HIST1H2AA PE=1 SV=3 183 14225 6 3 29.8 10.86
49 P36955 Pigment epithelium-derived factor OS=Homo sapiens GN=SERPINF1 PE=1 SV=4 181 46283 6 3 8.6 5.97
50 Q5VTE0 Putative elongation factor 1-alpha-like 3 OS=Homo sapiens GN=EEF1A1P5 PE=5 SV=1 175 50153 8 6 21 9.15
51 Q06033 Inter-alpha-trypsin inhibitor heavy chain H3 OS=Homo sapiens GN=ITIH3 PE=1 SV=2 168 99787 7 3 4 5.49
52 P07900 Heat shock protein HSP 90-alpha OS=Homo sapiens GN=HSP90AA1 PE=1 SV=5 165 84607 9 4 10.7 4.94
53 P08107 Heat shock 70 kDa protein 1A/1B OS=Homo sapiens GN=HSPA1A PE=1 SV=5 162 70009 4 3 6.2 5.48
54 P17066 Heat shock 70 kDa protein 6 OS=Homo sapiens GN=HSPA6 PE=1 SV=2 162 70984 4 3 6.2 5.81
55 P14618 Pyruvate kinase isozymes M1/M2 OS=Homo sapiens GN=PKM2 PE=1 SV=4 157 57900 7 6 16 7.96
56 P19013 Keratin, type II cytoskeletal 4 OS=Homo sapiens GN=KRT4 PE=1 SV=4 155 57250 8 4 7.3 6.25
57 P05556 Integrin beta-1 OS=Homo sapiens GN=ITGB1 PE=1 SV=2 155 88357 7 4 9.6 5.27
58 Q9UPY3 Endoribonuclease, Dicer OS=Homo sapiens GN=DICER PE=1 SV=2 154 218682 9 4 6.6 8.38
59 P62979 Ubiquitin-40S ribosomal protein S27a OS=Homo sapiens GN=RPS27A PE=1 SV=2 154 17953 8 5 34 9.68
60 P02042 Hemoglobin subunit delta OS=Homo sapiens GN=HBD PE=1 SV=2 150 16045 9 2 12.9 7.85
61 P08697 Alpha-2-antiplasmin OS=Homo sapiens GN=SERPINF2 PE=1 SV=3 148 54531 2 1 2.2 5.87
62 P02647 Apolipoprotein A-I OS=Homo sapiens GN=APOA1 PE=1 SV=1 146 30759 3 1 6 5.56
63 Q7Z3Y8 Keratin, type I cytoskeletal 27 OS=Homo sapiens GN=KRT27 PE=1 SV=2 142 49792 6 3 7.4 4.98
64 Q9P2B2 Prostaglandin F2 receptor negative regulator OS=Homo sapiens GN=PTGFRN PE=1 SV=2 140 98495 5 5 5.5 6.16
65 P13497 Bone morphogenetic protein 1 OS=Homo sapiens GN=BMP1 PE=1 SV=2 133 111178 3 2 3.1 6.48
66 P02675 Fibrinogen beta chain OS=Homo sapiens GN=FGB PE=1 SV=2 132 55892 4 3 4.7 8.54
67 Q9Y6L7 Tolloid-like protein 2 OS=Homo sapiens GN=TLL2 PE=1 SV=1 132 113484 2 2 2.2 5.6
68 P00747 Plasminogen OS=Homo sapiens GN=PLG PE=1 SV=2 131 90510 6 3 4.4 7.04
69 Q8WUM4 Programmed cell death 6-interacting protein OS=Homo sapiens GN=PDCD6IP PE=1 SV=1 126 95963 6 3 8.3 6.13
70 Q71DI3 Histone H3.2 OS=Homo sapiens GN=HIST2H3A PE=1 SV=3 121 15379 6 4 44.9 11.27
71 P84243 Histone H3.3 OS=Homo sapiens GN=H3F3A PE=1 SV=2 111 15318 6 4 44.9 11.27
72 Q86Y46 Keratin, type II cytoskeletal 73 OS=Homo sapiens GN=KRT73 PE=1 SV=1 110 58887 9 4 8 6.93
73 P12259 Coagulation factor V OS=Homo sapiens GN=F5 PE=1 SV=4 109 251546 6 4 2.3 5.68
74 P01008 Antithrombin-III OS=Homo sapiens GN=SERPINC1 PE=1 SV=1 108 52569 7 3 10.1 6.32
75 P06733 Alpha-enolase OS=Homo sapiens GN=ENO1 PE=1 SV=2 108 47139 3 3 10.8 7.01
76 P19827 Inter-alpha-trypsin inhibitor heavy chain H1 OS=Homo sapiens GN=ITIH1 PE=1 SV=3 106 101326 4 2 2.9 6.31
77 Q8NI35 InaD-like protein OS=Homo sapiens GN=INADL PE=1 SV=3 106 196247 10 1 0.3 4.84
78 Q8TAA3 Proteasome subunit alpha type-7-like OS=Homo sapiens GN=PSMA8 PE=1 SV=3 106 28512 3 2 19.5 9.07
79 P02452 Collagen alpha-1(I) chain OS=Homo sapiens GN=COL1A1 PE=1 SV=5 104 138857 2 2 1.6 5.6
80 P68431 Histone H3.1 OS=Homo sapiens GN=HIST1H3A PE=1 SV=2 103 15394 6 4 44.9 11.13
81 P07942 Laminin subunit beta-1 OS=Homo sapiens GN=LAMB1 PE=1 SV=2 102 197909 3 2 1.9 4.83
82 P05787 Keratin, type II cytoskeletal 8 OS=Homo sapiens GN=KRT8 PE=1 SV=7 102 53671 5 3 7.2 5.52
83 P51884 Lumican OS=Homo sapiens GN=LUM PE=1 SV=2 101 38405 3 2 8.3 6.16
84 Q00610 Clathrin heavy chain 1 OS=Homo sapiens GN=CLTC PE=1 SV=5 97 191493 2 2 2.9 5.48
85 P00558 Phosphoglycerate kinase 1 OS=Homo sapiens GN=PGK1 PE=1 SV=3 93 44586 3 2 8.4 8.3
86 P15531 Nucleoside diphosphate kinase A OS=Homo sapiens GN=NME1 PE=1 SV=1 92 17138 1 1 11.2 5.83
87 P04075 Fructose-bisphosphate aldolase A OS=Homo sapiens GN=ALDOA PE=1 SV=2 92 39395 5 2 12.4 8.3
88 P05452 Tetranectin OS=Homo sapiens GN=CLEC3B PE=1 SV=3 91 22522 3 3 21.8 5.52
89 P02787 Serotransferrin OS=Homo sapiens GN=TF PE=1 SV=3 91 77014 3 2 1.7 6.81
90 A6NIZ1 Ras-related protein Rap-1b-like protein OS=Homo sapiens PE=2 SV=1 89 20912 2 1 6.5 5.37
91 Q9GZM7 Tubulointerstitial nephritis antigen-like OS=Homo sapiens GN=TINAGL1 PE=1 SV=1 88 52353 1 1 2.8 6.54
92 P11021 78 kDa glucose-regulated protein OS=Homo sapiens GN=HSPA5 PE=1 SV=2 87 72288 3 2 3.1 5.07
93 P16870 Carboxypeptidase E OS=Homo sapiens GN=CPE PE=1 SV=1 87 53117 2 2 4.8 5.03
94 P81605 Dermcidin OS=Homo sapiens GN=DCD PE=1 SV=2 86 11277 2 2 20 6.08
95 P15169 Carboxypeptidase N catalytic chain OS=Homo sapiens GN=CPN1 PE=1 SV=1 84 52253 1 1 3.3 6.86
96 P11717 Cation-independent mannose-6-phosphate receptor OS=Homo sapiens GN=IGF2R PE=1 SV=3 83 274199 4 2 1.2 5.64
97 P07195 L-lactate dehydrogenase B chain OS=Homo sapiens GN=LDHB PE=1 SV=2 81 36615 2 1 7.5 5.71
98 P06702 Protein S100-A9 OS=Homo sapiens GN=S100A9 PE=1 SV=1 80 13234 2 1 24.6 5.71
99 Q5D862 Filaggrin-2 OS=Homo sapiens GN=FLG2 PE=1 SV=1 79 247928 2 1 0.9 8.45
100 P28066 Proteasome subunit alpha type-5 OS=Homo sapiens GN=PSMA5 PE=1 SV=3 78 26394 1 1 7.9 4.74
101 Q92743 Serine protease HTRA1 OS=Homo sapiens GN=HTRA1 PE=1 SV=1 78 51255 3 2 6.5 8.09
102 P23526 Adenosylhomocysteinase OS=Homo sapiens GN=AHCY PE=1 SV=4 78 47685 2 2 6.3 5.92
103 P02760 Protein AMBP OS=Homo sapiens GN=AMBP PE=1 SV=1 77 38974 2 1 2 5.95
104 P05543 Thyroxine-binding globulin OS=Homo sapiens GN=SERPINA7 PE=1 SV=2 75 46295 3 1 2.4 5.87
105 P43686 26S protease regulatory subunit 6B OS=Homo sapiens GN=PSMC4 PE=1 SV=2 74 47337 2 1 2.9 5.09
106 Q7RTS7 Keratin, type II cytoskeletal 74 OS=Homo sapiens GN=KRT74 PE=1 SV=2 74 57830 4 2 4.2 7.59
108 Q99456 Keratin, type I cytoskeletal 12 OS=Homo sapiens GN=KRT12 PE=1 SV=1 73 53478 7 2 3.6 4.7
109 P60174 Triosephosphate isomerase OS=Homo sapiens GN=TPI1 PE=1 SV=2 72 26653 1 1 5.2 6.45
110 P05156 Complement factor I OS=Homo sapiens GN=CFI PE=1 SV=2 71 65707 2 2 3.4 7.72
111 Q12931 Heat shock protein 75 kDa, mitochondrial OS=Homo sapiens GN=TRAP1 PE=1 SV=3 71 80060 1 1 2 8.3
112 P08962 CD63 OS=HOMO SAPIENS GN=CD63 PE=4 SV=2 71 25637 3 3 16.4 5.3
113 P02749 Beta-2-glycoprotein 1 OS=Homo sapiens GN=APOH PE=1 SV=3 67 38273 3 1 5.2 8.34
114 B9A064 Immunoglobulin lambda-like polypeptide 5 OS=Homo sapiens GN=IGLL5 PE=2 SV=2 66 23049 2 1 3.7 9.08
115 P12109 Collagen alpha-1(VI) chain OS=Homo sapiens GN=COL6A1 PE=1 SV=3 66 108462 1 1 0.9 5.26
116 P04745 Alpha-amylase 1 OS=Homo sapiens GN=AMY1A PE=1 SV=2 66 57731 2 1 2.3 6.47
117 P35052 Glypican-1 OS=Homo sapiens GN=GPC1 PE=1 SV=2 66 61641 4 1 10.9 7.07
118 P08133 Annexin A6 OS=Homo sapiens GN=ANXA6 PE=1 SV=3 65 75826 4 2 5.9 5.42
119 P01031 Complement C5 OS=Homo sapiens GN=C5 PE=1 SV=4 65 188186 4 2 1.8 6.11
120 P29353_REVERSED SHC-transforming protein 1 OS=Homo sapiens GN=SHC1 PE=1 SV=4 - REVERSED 65 62782 10 1 1.4 6.01
121 O15111_REVERSED Inhibitor of nuclear factor kappa-B kinase subunit alpha OS=Homo sapiens GN=CHUK PE=1 SV=2 - REVERSED 65 84585 6 1 1.6 6.28
123 P01009 Alpha-1-antitrypsin OS=Homo sapiens GN=SERPINA1 PE=1 SV=3 64 46707 3 2 3.3 5.37
124 P13639 Elongation factor 2 OS=Homo sapiens GN=EEF2 PE=1 SV=4 64 95277 3 2 6.9 6.41
125 A6NMY6 Putative annexin A2-like protein OS=Homo sapiens GN=ANXA2P2 PE=5 SV=2 63 38635 2 1 6.2 6.49
126 Q99816 Tumor susceptibility gene 101 protein OS=Homo sapiens GN=TSG101 PE=1 SV=2 60 43916 1 1 2.6 6.06
127 P02461 Collagen alpha-1(III) chain OS=Homo sapiens GN=COL3A1 PE=1 SV=4 59 138479 2 2 1.5 6.21
128 Q14624 Inter-alpha-trypsin inhibitor heavy chain H4 OS=Homo sapiens GN=ITIH4 PE=1 SV=4 58 103293 5 2 3.3 6.51
129 P02649 Apolipoprotein E OS=Homo sapiens GN=APOE PE=1 SV=1 57 36132 1 1 2.8 5.65
130 P26641 Elongation factor 1-gamma OS=Homo sapiens GN=EEF1G PE=1 SV=3 56 50087 2 1 5 6.25
131 P35125 Ubiquitin carboxyl-terminal hydrolase 6 OS=Homo sapiens GN=USP6 PE=1 SV=2 55 158557 13 1 0.4 7.87
132 P02790 Hemopexin OS=Homo sapiens GN=HPX PE=1 SV=2 54 51643 6 1 5.4 6.55
133 Q9BYX7_REVERSED Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 - REVERSED 54 41989 6 1 3.5 5.92
135 P14923 Junction plakoglobin OS=Homo sapiens GN=JUP PE=1 SV=3 53 81693 1 1 2.7 5.75
136 Q14764 Major vault protein OS=Homo sapiens GN=MVP PE=1 SV=4 52 99266 1 1 1.9 5.34
138 P01042 Kininogen-1 OS=Homo sapiens GN=KNG1 PE=1 SV=2 52 71912 1 1 1.4 6.34
139 Q8N139 ATP-binding cassette sub-family A member 6 OS=Homo sapiens GN=ABCA6 PE=1 SV=2 52 184167 2 1 0.4 7.05
141 Q96Q89_REVERSED Kinesin-like protein KIF20B OS=Homo sapiens GN=KIF20B PE=1 SV=3 - REVERSED 51 210500 2 1 0.5 5.54
142 Q2Q1W2_REVERSED Tripartite motif-containing protein 71 OS=Homo sapiens GN=TRIM71 PE=2 SV=1 - REVERSED 51 93326 1 1 1.2 7.62
143 Q9BY43 Charged multivesicular body protein 4a OS=Homo sapiens GN=CHMP4A PE=1 SV=3 51 25083 2 1 3.2 4.65
144 Q8NAV2_REVERSED Uncharacterized protein C8orf58 OS=Homo sapiens GN=C8orf58 PE=2 SV=2 - REVERSED 51 39636 1 1 1.9 8.54
145 P02748 Complement component C9 OS=Homo sapiens GN=C9 PE=1 SV=2 51 63133 1 1 1.6 5.43
146 O00754 Lysosomal alpha-mannosidase OS=Homo sapiens GN=MAN2B1 PE=1 SV=3 51 113672 1 1 1.1 6.84
147 O14817 Tetraspanin-4 OS=Homo sapiens GN=TSPAN4 PE=1 SV=1 50 26100 2 1 10.5 6.07
148 P11047 Laminin subunit gamma-1 OS=Homo sapiens GN=LAMC1 PE=1 SV=3 50 177489 3 3 1.9 5.01
149 P21333 Filamin-A OS=Homo sapiens GN=FLNA PE=1 SV=4 50 280564 2 1 1.6 5.7
150 A8MT19_REVERSED Putative rhophilin-2-like protein OS=Homo sapiens PE=5 SV=2 - REVERSED 49 65903 1 1 1.2 6.53
151 P12111 Collagen alpha-3(VI) chain OS=Homo sapiens GN=COL6A3 PE=1 SV=5 48 343457 3 1 1.2 6.26
152 Q00013 55 kDa erythrocyte membrane protein OS=Homo sapiens GN=MPP1 PE=1 SV=2 47 52264 2 1 1.5 6.91
153 Q8IW52 SLIT and NTRK-like protein 4 OS=Homo sapiens GN=SLITRK4 PE=2 SV=1 47 94271 1 1 0.8 7.95
154 O95445 Apolipoprotein M OS=Homo sapiens GN=APOM PE=1 SV=2 47 21239 1 1 3.7 5.66
155 Q13162 Peroxiredoxin-4 OS=Homo sapiens GN=PRDX4 PE=1 SV=1 46 30521 1 1 4.1 5.86
156 P00488 Coagulation factor XIII A chain OS=Homo sapiens GN=F13A1 PE=1 SV=4 46 83215 2 2 4 5.75
157 P08572 Collagen alpha-2(IV) chain OS=Homo sapiens GN=COL4A2 PE=1 SV=4 46 167449 1 1 0.6 8.89
158 O14782 Kinesin-like protein KIF3C OS=Homo sapiens GN=KIF3C PE=1 SV=3 46 89371 4 1 1.1 8.28
159 P35443 Thrombospondin-4 OS=Homo sapiens GN=THBS4 PE=1 SV=2 45 105802 2 1 2.8 4.44
161 P49747 Cartilage oligomeric matrix protein OS=Homo sapiens GN=COMP PE=1 SV=2 44 82808 2 1 2.8 4.36
162 Q14573_REVERSED Inositol 1,4,5-trisphosphate receptor type 3 OS=Homo sapiens GN=ITPR3 PE=1 SV=2 - REVERSED 44 303912 1 1 0.4 6.05
163 Q8IWC1 MAP7 domain-containing protein 3 OS=Homo sapiens GN=MAP7D3 PE=1 SV=2 44 98368 2 1 2.1 9.34
164 Q3KQU3 MAP7 domain-containing protein 1 OS=Homo sapiens GN=MAP7D1 PE=1 SV=1 44 92764 1 1 0.7 10.12
165 Q5U5X0 LYR motif-containing protein 7 OS=Homo sapiens GN=LYRM7 PE=1 SV=1 44 11947 1 1 5.8 9.67
166 Q8N7B9_REVERSED EF-hand calcium-binding domain-containing protein 3 OS=Homo sapiens GN=EFCAB3 PE=1 SV=1 - REVERSED 44 50114 1 1 1.4 9.31
167 Q8NB25_REVERSED Protein FAM184A OS=Homo sapiens GN=FAM184A PE=2 SV=3 - REVERSED 44 132883 1 1 0.5 5.59
168 P06732 Creatine kinase M-type OS=Homo sapiens GN=CKM PE=1 SV=2 44 43074 1 1 1.6 6.77
169 Q93096_REVERSED Protein tyrosine phosphatase type IVA 1 OS=Homo sapiens GN=PTP4A1 PE=1 SV=2 - REVERSED 44 19802 1 1 3.5 9.17
170 Q6YHK3 CD109 antigen OS=Homo sapiens GN=CD109 PE=1 SV=2 44 161587 1 1 0.6 5.59
171 Q8IYT4 Katanin p60 ATPase-containing subunit A-like 2 OS=Homo sapiens GN=KATNAL2 PE=2 SV=3 43 61214 2 1 2.2 7.22
172 A8MPT4 Glutathione S-transferase theta-4 OS=Homo sapiens GN=GSTT4 PE=3 SV=2 43 27941 1 1 2.5 6.25
173 Q8N6V9_REVERSED Testis-expressed sequence 9 protein OS=Homo sapiens GN=TEX9 PE=2 SV=1 - REVERSED 43 44798 1 1 1.5 6.2
174 Q9Y2U8_REVERSED Inner nuclear membrane protein Man1 OS=Homo sapiens GN=LEMD3 PE=1 SV=2 - REVERSED 43 99935 1 1 0.7 7.31
175 P05154 Plasma serine protease inhibitor OS=Homo sapiens GN=SERPINA5 PE=1 SV=3 43 45646 1 1 1.5 9.3
176 Q92609 TBC1 domain family member 5 OS=Homo sapiens GN=TBC1D5 PE=1 SV=1 42 88949 1 1 1 6.1
177 Q9H2G4 Testis-specific Y-encoded-like protein 2 OS=Homo sapiens GN=TSPYL2 PE=1 SV=1 42 79387 1 1 1 4.45
178 Q00535_REVERSED Cyclin-dependent kinase 5 OS=Homo sapiens GN=CDK5 PE=1 SV=3 - REVERSED 42 33283 1 1 3.1 7.9
179 P06703 Protein S100-A6 OS=Homo sapiens GN=S100A6 PE=1 SV=1 41 10173 2 1 8.9 5.33
180 Q6ZMU1 Putative protein C3P1 OS=Homo sapiens GN=C3P1 PE=5 SV=3 41 40171 2 1 5.5 6.32
183 Q86UY8_REVERSED 5~-nucleotidase domain-containing protein 3 OS=Homo sapiens GN=NT5DC3 PE=2 SV=1 - REVERSED 41 63379 2 1 1.5 8.47
184 O75460_REVERSED Serine/threonine-protein kinase/endoribonuclease IRE1 OS=Homo sapiens GN=ERN1 PE=1 SV=2 - REVERSED 41 109666 1 1 0.6 5.98
186 Q9Y6X9_REVERSED MORC family CW-type zinc finger protein 2 OS=Homo sapiens GN=MORC2 PE=1 SV=2 - REVERSED 40 117750 1 1 0.7 8.6
187 Q96TA0_REVERSED Putative protocadherin beta-18 OS=Homo sapiens GN=PCDHB18 PE=5 SV=1 - REVERSED 40 80423 1 1 0.8 4.87
188 P02458 Collagen alpha-1(II) chain OS=Homo sapiens GN=COL2A1 PE=1 SV=3 40 141698 1 1 0.7 6.58
191 Q8NBM4 Ubiquitin-associated domain-containing protein 2 OS=Homo sapiens GN=UBAC2 PE=2 SV=1 40 38938 1 1 2.3 9.21
193 Q7Z5M8 Abhydrolase domain-containing protein 12B OS=Homo sapiens GN=ABHD12B PE=2 SV=1 40 40750 5 1 1.7 8.57
194 Q15020_REVERSED Squamous cell carcinoma antigen recognized by T-cells 3 OS=Homo sapiens GN=SART3 PE=1 SV=1 - REVERSED 40 109865 5 1 0.6 5.45
196 O60312 Probable phospholipid-transporting ATPase VA OS=Homo sapiens GN=ATP10A PE=2 SV=2 39 167582 2 1 0.6 8.7
197 Q9Y6Q3_REVERSED Zinc finger protein 37 homolog OS=Homo sapiens GN=ZFP37 PE=2 SV=3 - REVERSED 39 71164 1 1 1.7 9.26
198 Q96RW7 Hemicentin-1 OS=Homo sapiens GN=HMCN1 PE=1 SV=2 39 613001 2 1 0.1 6.07
199 Q13183_REVERSED Solute carrier family 13 member 2 OS=Homo sapiens GN=SLC13A2 PE=2 SV=1 - REVERSED 38 64368 2 1 1.4 6.61
200 Q9UK41 Vacuolar protein sorting-associated protein 28 homolog OS=Homo sapiens GN=VPS28 PE=1 SV=1 38 25409 2 1 6.8 5.37
201 Q14C86_REVERSED GTPase-activating protein and VPS9 domain-containing protein 1 OS=Homo sapiens GN=GAPVD1 PE=1 SV=2 - REVERSED 38 164876 1 1 0.5 5.09
206 P26640 Valyl-tRNA synthetase OS=Homo sapiens GN=VARS PE=1 SV=4 38 140387 1 1 1.2 7.53
207 Q96JB3 Hypermethylated in cancer 2 protein OS=Homo sapiens GN=HIC2 PE=1 SV=2 38 66114 1 1 1.6 5.95
208 O43309 Zinc finger and SCAN domain-containing protein 12 OS=Homo sapiens GN=ZSCAN12 PE=2 SV=1 38 70178 1 1 2 6.28
209 Q96K21 Zinc finger FYVE domain-containing protein 19 OS=Homo sapiens GN=ZFYVE19 PE=1 SV=3 38 51514 2 1 1.7 5.57
210 Q8WZ42_REVERSED Titin OS=Homo sapiens GN=TTN PE=1 SV=2 - REVERSED 38 3813810 2 1 0.1 6.01
211 O75865_REVERSED Trafficking protein particle complex subunit 6A OS=Homo sapiens GN=TRAPPC6A PE=1 SV=2 - REVERSED 38 17593 1 1 3.8 5.14
213 P04083 Annexin A1 OS=Homo sapiens GN=ANXA1 PE=1 SV=2 37 38690 1 1 2.3 6.57
214 P28070 Proteasome subunit beta type-4 OS=Homo sapiens GN=PSMB4 PE=1 SV=4 37 29185 1 1 3.8 5.72
216 P02753 Retinol-binding protein 4 OS=Homo sapiens GN=RBP4 PE=1 SV=3 37 22995 2 1 10 5.76
217 Q12923 Tyrosine-protein phosphatase non-receptor type 13 OS=Homo sapiens GN=PTPN13 PE=1 SV=2 37 276733 1 1 0.5 5.99
218 P05109 Protein S100-A8 OS=Homo sapiens GN=S100A8 PE=1 SV=1 37 10828 1 1 11.8 6.51
220 Q5SYB0 FERM and PDZ domain-containing protein 1 OS=Homo sapiens GN=FRMPD1 PE=1 SV=1 37 173328 1 1 0.4 5.14
221 A6NC57_REVERSED Ankyrin repeat domain-containing protein 62 OS=Homo sapiens GN=ANKRD62 PE=2 SV=4 - REVERSED 37 106379 1 1 0.7 6.26
222 Q8IWZ5_REVERSED Tripartite motif-containing protein 42 OS=Homo sapiens GN=TRIM42 PE=2 SV=2 - REVERSED 37 82690 1 1 0.8 8.31
223 Q9UKU7_REVERSED Isobutyryl-CoA dehydrogenase, mitochondrial OS=Homo sapiens GN=ACAD8 PE=1 SV=1 - REVERSED 37 45040 1 1 1.4 8.16
224 Q674X7_REVERSED Kazrin OS=Homo sapiens GN=KAZN PE=1 SV=2 - REVERSED 37 86298 1 1 0.8 6.59
225 Q14534 Squalene monooxygenase OS=Homo sapiens GN=SQLE PE=2 SV=3 37 63882 1 1 1 8.8
226 P20138 Myeloid cell surface antigen CD33 OS=Homo sapiens GN=CD33 PE=1 SV=2 37 39800 1 1 1.9 8.69
227 A2RRP1_REVERSED Neuroblastoma-amplified sequence OS=Homo sapiens GN=NBAS PE=1 SV=2 - REVERSED 37 268401 1 1 0.3 5.65
231 O95714_REVERSED E3 ubiquitin-protein ligase HERC2 OS=Homo sapiens GN=HERC2 PE=1 SV=2 - REVERSED 37 526895 6 1 0.4 5.88
232 Q9UGM3 Deleted in malignant brain tumors 1 protein OS=Homo sapiens GN=DMBT1 PE=1 SV=2 36 260569 1 1 0.2 5.18
233 P26378 ELAV-like protein 4 OS=Homo sapiens GN=ELAVL4 PE=1 SV=2 36 41743 2 1 3.7 9.45
234 Q96A37 RING finger protein 166 OS=Homo sapiens GN=RNF166 PE=2 SV=1 36 26105 1 1 4.2 8.42
236 P22352 Glutathione peroxidase 3 OS=Homo sapiens GN=GPX3 PE=1 SV=2 36 25537 1 1 5.3 8.26
237 Q9Y240 C-type lectin domain family 11 member A OS=Homo sapiens GN=CLEC11A PE=1 SV=1 36 35672 1 1 5.3 5.06
240 P25789 Proteasome subunit alpha type-4 OS=Homo sapiens GN=PSMA4 PE=1 SV=1 36 29465 1 1 3.4 7.57
241 P10242_REVERSED Transcriptional activator Myb OS=Homo sapiens GN=MYB PE=1 SV=2 - REVERSED 36 72296 2 1 1.3 6.35
242 P02794 Ferritin heavy chain OS=Homo sapiens GN=FTH1 PE=1 SV=2 35 21226 1 1 2.5 8.2
NIH 3T3
prot_hit prot_acc prot_desc prot_score prot_mass prot_matches prot_sequences_sig prot_cover prot_pi
1 P04264 Keratin, type II cytoskeletal 1 OS=Homo sapiens GN=KRT1 PE=1 SV=6 2826 65999 70 31 55.3 8.15
2 P13645 Keratin, type I cytoskeletal 10 OS=Homo sapiens GN=KRT10 PE=1 SV=6 2816 58792 83 29 57.9 5.13
3 P02768 Serum albumin OS=Homo sapiens GN=ALB PE=1 SV=2 2011 69321 114 8 13.5 5.92
4 P01023 Alpha-2-macroglobulin OS=Homo sapiens GN=A2M PE=1 SV=3 1949 163188 70 8 5.7 6.03
5 P02765 Alpha-2-HS-glycoprotein OS=Homo sapiens GN=AHSG PE=1 SV=1 1830 39300 50 4 8.7 5.43
6 P35527 Keratin, type I cytoskeletal 9 OS=Homo sapiens GN=KRT9 PE=1 SV=3 1822 62027 44 22 68.7 5.14
7 P35908 Keratin, type II cytoskeletal 2 epidermal OS=Homo sapiens GN=KRT2 PE=1 SV=2 1641 65393 46 24 58.1 8.07
8 P02751 Fibronectin OS=Homo sapiens GN=FN1 PE=1 SV=4 1564 262460 50 29 18.1 5.46
9 P20742 Pregnancy zone protein OS=Homo sapiens GN=PZP PE=1 SV=4 1547 163760 48 5 4.3 5.97
10 P01024 Complement C3 OS=Homo sapiens GN=C3 PE=1 SV=2 704 187030 25 11 8.9 6.02
11 P07996 Thrombospondin-1 OS=Homo sapiens GN=THBS1 PE=1 SV=2 622 129300 26 15 19.1 4.71
12 P19823 Inter-alpha-trypsin inhibitor heavy chain H2 OS=Homo sapiens GN=ITIH2 PE=1 SV=2 593 106397 18 7 9.2 6.4
13 P00734 Prothrombin OS=Homo sapiens GN=F2 PE=1 SV=2 527 69992 11 6 8 5.64
14 P13647 Keratin, type II cytoskeletal 5 OS=Homo sapiens GN=KRT5 PE=1 SV=3 514 62340 23 16 28.8 7.59
15 P0C0L4 Complement C4-A OS=Homo sapiens GN=C4A PE=1 SV=1 467 192650 18 7 5.2 6.65
16 Q07954 Prolow-density lipoprotein receptor-related protein 1 OS=Homo sapiens GN=LRP1 PE=1 SV=2 415 504276 14 11 3.5 5.16
17 P02788 Lactotransferrin OS=Homo sapiens GN=LTF PE=1 SV=6 406 78132 18 3 4.8 8.5
18 P04259 Keratin, type II cytoskeletal 6B OS=Homo sapiens GN=KRT6B PE=1 SV=5 394 60030 17 11 23 8.09
19 P02538 Keratin, type II cytoskeletal 6A OS=Homo sapiens GN=KRT6A PE=1 SV=3 388 60008 17 12 24.6 8.09
20 P06396 Gelsolin OS=Homo sapiens GN=GSN PE=1 SV=1 360 85644 11 7 12.8 5.9
21 P02533 Keratin, type I cytoskeletal 14 OS=Homo sapiens GN=KRT14 PE=1 SV=4 353 51529 17 9 20.3 5.09
22 P60709 Actin, cytoplasmic 1 OS=Homo sapiens GN=ACTB PE=1 SV=1 315 41710 12 8 33.6 5.29
23 P02771 Alpha-fetoprotein OS=Homo sapiens GN=AFP PE=1 SV=1 288 68633 8 3 6.7 5.48
24 P02774 Vitamin D-binding protein OS=Homo sapiens GN=GC PE=1 SV=1 271 52929 12 5 12 5.4
25 P69905 Hemoglobin subunit alpha OS=Homo sapiens GN=HBA1 PE=1 SV=2 252 15248 13 4 33.1 8.72
26 Q7Z794 Keratin, type II cytoskeletal 1b OS=Homo sapiens GN=KRT77 PE=1 SV=3 248 61864 7 3 4.8 5.73
27 P36955 Pigment epithelium-derived factor OS=Homo sapiens GN=SERPINF1 PE=1 SV=4 233 46283 10 3 12 5.97
28 P04004 Vitronectin OS=Homo sapiens GN=VTN PE=1 SV=1 226 54271 6 1 4.2 5.55
29 P02647 Apolipoprotein A-I OS=Homo sapiens GN=APOA1 PE=1 SV=1 226 30759 4 1 6 5.56
30 P62736 Actin, aortic smooth muscle OS=Homo sapiens GN=ACTA2 PE=1 SV=1 220 41982 7 5 18.8 5.23
31 Q04695 Keratin, type I cytoskeletal 17 OS=Homo sapiens GN=KRT17 PE=1 SV=2 205 48076 12 6 14.6 4.97
32 Q5XKE5 Keratin, type II cytoskeletal 79 OS=Homo sapiens GN=KRT79 PE=1 SV=2 200 57800 7 4 6.9 6.75
33 P13646 Keratin, type I cytoskeletal 13 OS=Homo sapiens GN=KRT13 PE=1 SV=4 198 49557 8 4 6.1 4.91
34 Q06033 Inter-alpha-trypsin inhibitor heavy chain H3 OS=Homo sapiens GN=ITIH3 PE=1 SV=2 196 99787 9 3 6.3 5.49
35 Q86YZ3 Hornerin OS=Homo sapiens GN=HRNR PE=1 SV=2 193 282228 3 2 2.3 10.05
36 P02042 Hemoglobin subunit delta OS=Homo sapiens GN=HBD PE=1 SV=2 189 16045 10 2 12.9 7.85
37 Q15063 Periostin OS=Homo sapiens GN=POSTN PE=1 SV=2 174 93255 7 2 7.4 7.27
38 P23142 Fibulin-1 OS=Homo sapiens GN=FBLN1 PE=1 SV=4 172 77162 9 4 7.3 5.07
39 P02100 Hemoglobin subunit epsilon OS=Homo sapiens GN=HBE1 PE=1 SV=2 160 16192 7 1 12.9 8.67
40 P08697 Alpha-2-antiplasmin OS=Homo sapiens GN=SERPINF2 PE=1 SV=3 143 54531 2 1 2.2 5.87
41 P00747 Plasminogen OS=Homo sapiens GN=PLG PE=1 SV=2 139 90510 6 3 4.3 7.04
42 P02675 Fibrinogen beta chain OS=Homo sapiens GN=FGB PE=1 SV=2 136 55892 5 3 6.7 8.54
43 P04406 Glyceraldehyde-3-phosphate dehydrogenase OS=Homo sapiens GN=GAPDH PE=1 SV=3 134 36030 3 3 17.6 8.57
44 P05556 Integrin beta-1 OS=Homo sapiens GN=ITGB1 PE=1 SV=2 131 88357 4 3 6 5.27
45 Q9BQE3 Tubulin alpha-1C chain OS=Homo sapiens GN=TUBA1C PE=1 SV=1 130 49863 7 4 14.9 4.96
46 P01008 Antithrombin-III OS=Homo sapiens GN=SERPINC1 PE=1 SV=1 130 52569 10 4 14.7 6.32
47 P02452 Collagen alpha-1(I) chain OS=Homo sapiens GN=COL1A1 PE=1 SV=5 124 138857 3 3 2.4 5.6
48 P01031 Complement C5 OS=Homo sapiens GN=C5 PE=1 SV=4 122 188186 6 2 2.9 6.11
49 P11142 Heat shock cognate 71 kDa protein OS=Homo sapiens GN=HSPA8 PE=1 SV=1 122 70854 3 3 5.3 5.37
50 P07437 Tubulin beta chain OS=Homo sapiens GN=TUBB PE=1 SV=2 121 49639 5 4 24.8 4.78
51 P19013 Keratin, type II cytoskeletal 4 OS=Homo sapiens GN=KRT4 PE=1 SV=4 120 57250 7 3 7.3 6.25
52 P08107 Heat shock 70 kDa protein 1A/1B OS=Homo sapiens GN=HSPA1A PE=1 SV=5 119 70009 2 2 4.2 5.48
53 P17066 Heat shock 70 kDa protein 6 OS=Homo sapiens GN=HSPA6 PE=1 SV=2 119 70984 2 2 4.2 5.81
54 P12109 Collagen alpha-1(VI) chain OS=Homo sapiens GN=COL6A1 PE=1 SV=3 117 108462 2 2 2.4 5.26
55 P02748 Complement component C9 OS=Homo sapiens GN=C9 PE=1 SV=2 114 63133 3 2 3.8 5.43
56 P19827 Inter-alpha-trypsin inhibitor heavy chain H1 OS=Homo sapiens GN=ITIH1 PE=1 SV=3 113 101326 4 2 2.9 6.31
57 Q8TAA3 Proteasome subunit alpha type-7-like OS=Homo sapiens GN=PSMA8 PE=1 SV=3 113 28512 2 2 10.9 9.07
58 Q7Z3Y8 Keratin, type I cytoskeletal 27 OS=Homo sapiens GN=KRT27 PE=1 SV=2 111 49792 6 3 7.4 4.98
59 P06702 Protein S100-A9 OS=Homo sapiens GN=S100A9 PE=1 SV=1 109 13234 4 2 43 5.71
60 P05787 Keratin, type II cytoskeletal 8 OS=Homo sapiens GN=KRT8 PE=1 SV=7 109 53671 4 3 5.6 5.52
61 P05109 Protein S100-A8 OS=Homo sapiens GN=S100A8 PE=1 SV=1 109 10828 3 3 31.2 6.51
62 P02787 Serotransferrin OS=Homo sapiens GN=TF PE=1 SV=3 109 77014 5 2 1.7 6.81
63 Q562R1 Beta-actin-like protein 2 OS=Homo sapiens GN=ACTBL2 PE=1 SV=2 108 41976 5 3 13.8 5.39
64 P62979 Ubiquitin-40S ribosomal protein S27a OS=Homo sapiens GN=RPS27A PE=1 SV=2 102 17953 7 3 30.1 9.68
65 P12035 Keratin, type II cytoskeletal 3 OS=Homo sapiens GN=KRT3 PE=1 SV=3 99 64378 5 2 4 6.12
66 Q8NI35 InaD-like protein OS=Homo sapiens GN=INADL PE=1 SV=3 98 196247 10 1 0.3 4.84
67 Q8TEV9_REVERSED Smith-Magenis syndrome chromosomal region candidate gene 8 protein OS=Homo sapiens GN=SMCR8 PE=1 SV=2 - REVERSED 97 104956 10 1 1.2 5.36
68 Q5D862 Filaggrin-2 OS=Homo sapiens GN=FLG2 PE=1 SV=1 96 247928 2 2 1.3 8.45
69 P12111 Collagen alpha-3(VI) chain OS=Homo sapiens GN=COL6A3 PE=1 SV=5 96 343457 3 3 1.1 6.26
70 P49747 Cartilage oligomeric matrix protein OS=Homo sapiens GN=COMP PE=1 SV=2 94 82808 3 2 4.5 4.36
71 A6NIZ1 Ras-related protein Rap-1b-like protein OS=Homo sapiens PE=2 SV=1 91 20912 2 1 6.5 5.37
72 P12259 Coagulation factor V OS=Homo sapiens GN=F5 PE=1 SV=4 89 251546 5 2 2 5.68
73 Q14624 Inter-alpha-trypsin inhibitor heavy chain H4 OS=Homo sapiens GN=ITIH4 PE=1 SV=4 87 103293 5 2 3.3 6.51
74 Q6KB66 Keratin, type II cytoskeletal 80 OS=Homo sapiens GN=KRT80 PE=1 SV=2 86 50494 3 1 5.1 5.58
75 P13497 Bone morphogenetic protein 1 OS=Homo sapiens GN=BMP1 PE=1 SV=2 83 111178 4 3 3.1 6.48
76 Q86Y46 Keratin, type II cytoskeletal 73 OS=Homo sapiens GN=KRT73 PE=1 SV=1 78 58887 7 3 8 6.93
77 P04114 Apolipoprotein B-100 OS=Homo sapiens GN=APOB PE=1 SV=2 77 515283 3 3 0.5 6.58
78 P02790 Hemopexin OS=Homo sapiens GN=HPX PE=1 SV=2 76 51643 6 1 3.2 6.55
79 P35443 Thrombospondin-4 OS=Homo sapiens GN=THBS4 PE=1 SV=2 75 105802 2 2 2.5 4.44
80 P08123 Collagen alpha-2(I) chain OS=Homo sapiens GN=COL1A2 PE=1 SV=7 75 129235 2 2 2.2 9.08
81 P15169 Carboxypeptidase N catalytic chain OS=Homo sapiens GN=CPN1 PE=1 SV=1 73 52253 1 1 3.3 6.86
82 Q92743 Serine protease HTRA1 OS=Homo sapiens GN=HTRA1 PE=1 SV=1 73 51255 2 2 4.4 8.09
83 Q9Y6L7 Tolloid-like protein 2 OS=Homo sapiens GN=TLL2 PE=1 SV=1 72 113484 3 2 2.2 5.6
84 P05452 Tetranectin OS=Homo sapiens GN=CLEC3B PE=1 SV=3 69 22522 2 2 12.9 5.52
85 Q96Q89_REVERSED Kinesin-like protein KIF20B OS=Homo sapiens GN=KIF20B PE=1 SV=3 - REVERSED 68 210500 2 1 0.5 5.54
86 P51884 Lumican OS=Homo sapiens GN=LUM PE=1 SV=2 68 38405 4 1 6.8 6.16
87 P02749 Beta-2-glycoprotein 1 OS=Homo sapiens GN=APOH PE=1 SV=3 67 38273 3 1 5.2 8.34
88 P63104 14-3-3 protein zeta/delta OS=Homo sapiens GN=YWHAZ PE=1 SV=1 67 27728 1 1 4.9 4.73
89 O15111_REVERSED Inhibitor of nuclear factor kappa-B kinase subunit alpha OS=Homo sapiens GN=CHUK PE=1 SV=2 - REVERSED 66 84585 7 1 1.6 6.28
90 Q03692 Collagen alpha-1(X) chain OS=Homo sapiens GN=COL10A1 PE=1 SV=2 64 66117 1 1 1.9 9.68
91 B9A064 Immunoglobulin lambda-like polypeptide 5 OS=Homo sapiens GN=IGLL5 PE=2 SV=2 63 23049 2 1 3.7 9.08
92 Q7RTS7 Keratin, type II cytoskeletal 74 OS=Homo sapiens GN=KRT74 PE=1 SV=2 62 57830 3 2 4.2 7.59
93 Q99456 Keratin, type I cytoskeletal 12 OS=Homo sapiens GN=KRT12 PE=1 SV=1 62 53478 5 2 3.6 4.7
94 Q9BYX7_REVERSED Putative beta-actin-like protein 3 OS=Homo sapiens GN=POTEKP PE=5 SV=1 - REVERSED 61 41989 6 1 3.5 5.92
95 P00450 Ceruloplasmin OS=Homo sapiens GN=CP PE=1 SV=1 59 122128 3 2 2.9 5.44
96 P23526 Adenosylhomocysteinase OS=Homo sapiens GN=AHCY PE=1 SV=4 59 47685 2 1 6.3 5.92
97 P20618 Proteasome subunit beta type-1 OS=Homo sapiens GN=PSMB1 PE=1 SV=2 58 26472 1 1 5.8 8.27
98 P98160 Basement membrane-specific heparan sulfate proteoglycan core protein OS=Homo sapiens GN=HSPG2 PE=1 SV=4 57 468532 2 2 0.6 6.06
99 Q8WUM4 Programmed cell death 6-interacting protein OS=Homo sapiens GN=PDCD6IP PE=1 SV=1 57 95963 2 1 2.2 6.13
100 P35125 Ubiquitin carboxyl-terminal hydrolase 6 OS=Homo sapiens GN=USP6 PE=1 SV=2 56 158557 12 1 0.4 7.87
101 P02679 Fibrinogen gamma chain OS=Homo sapiens GN=FGG PE=1 SV=3 56 51479 1 1 2 5.37
102 P29353_REVERSED SHC-transforming protein 1 OS=Homo sapiens GN=SHC1 PE=1 SV=4 - REVERSED 56 62782 9 1 1.4 6.01
103 P05546 Heparin cofactor 2 OS=Homo sapiens GN=SERPIND1 PE=1 SV=3 55 57034 1 1 2.4 6.41
104 P01344 Insulin-like growth factor II OS=Homo sapiens GN=IGF2 PE=1 SV=1 54 20127 1 1 8.9 9.5
105 P02794 Ferritin heavy chain OS=Homo sapiens GN=FTH1 PE=1 SV=2 54 21212 1 1 5.5 5.3
106 P01009 Alpha-1-antitrypsin OS=Homo sapiens GN=SERPINA1 PE=1 SV=3 52 46707 3 1 3.3 5.37
107 Q16853 Membrane primary amine oxidase OS=Homo sapiens GN=AOC3 PE=1 SV=3 52 84568 1 1 1.2 6.05
108 P20930 Filaggrin OS=Homo sapiens GN=FLG PE=1 SV=3 51 434922 1 1 0.3 9.24
109 Q13454 Tumor suppressor candidate 3 OS=Homo sapiens GN=TUSC3 PE=2 SV=1 49 39650 1 1 2 9.93
110 Q9BXJ4 Complement C1q tumor necrosis factor-related protein 3 OS=Homo sapiens GN=C1QTNF3 PE=2 SV=1 48 26977 2 2 6.5 6.04
111 Q9HAV0 Guanine nucleotide-binding protein subunit beta-4 OS=Homo sapiens GN=GNB4 PE=1 SV=3 48 37543 1 1 2.9 5.6
112 Q9NWT1_REVERSED p21-activated protein kinase-interacting protein 1 OS=Homo sapiens GN=PAK1IP1 PE=1 SV=2 - REVERSED 48 43936 2 1 1.8 9.07
113 Q6ZU45 Putative C-type lectin domain-containing protein NCRNA00083 OS=Homo sapiens GN=NCRNA00083 PE=5 SV=1 48 25796 1 1 3.4 6.3
114 Q92609 TBC1 domain family member 5 OS=Homo sapiens GN=TBC1D5 PE=1 SV=1 47 88949 2 1 1 6.1
115 O95497 Pantetheinase OS=Homo sapiens GN=VNN1 PE=1 SV=2 47 56975 1 1 3.7 5.32
116 O95714_REVERSED E3 ubiquitin-protein ligase HERC2 OS=Homo sapiens GN=HERC2 PE=1 SV=2 - REVERSED 47 526895 4 1 0.2 5.88
117 P11717 Cation-independent mannose-6-phosphate receptor OS=Homo sapiens GN=IGF2R PE=1 SV=3 47 274199 2 1 0.8 5.64
118 Q9NPY3_REVERSED Complement component C1q receptor OS=Homo sapiens GN=CD93 PE=1 SV=3 - REVERSED 46 68515 1 1 1.1 5.27
119 P31151 Protein S100-A7 OS=Homo sapiens GN=S100A7 PE=1 SV=4 46 11464 2 1 31.7 6.28
120 P19961 Alpha-amylase 2B OS=Homo sapiens GN=AMY2B PE=1 SV=1 46 57673 2 1 5.1 6.64
121 P02760 Protein AMBP OS=Homo sapiens GN=AMBP PE=1 SV=1 46 38974 2 1 2 5.95
122 Q9H4M9 EH domain-containing protein 1 OS=Homo sapiens GN=EHD1 PE=1 SV=2 46 60589 2 1 4.9 6.35
123 Q96CD0_REVERSED F-box/LRR-repeat protein 8 OS=Homo sapiens GN=FBXL8 PE=1 SV=1 - REVERSED 45 40490 11 1 1.9 7.03
124 Q08554 Desmocollin-1 OS=Homo sapiens GN=DSC1 PE=1 SV=2 45 99924 1 1 1.7 5.25
125 P49720 Proteasome subunit beta type-3 OS=Homo sapiens GN=PSMB3 PE=1 SV=2 45 22933 2 2 16.6 6.14
126 P05543 Thyroxine-binding globulin OS=Homo sapiens GN=SERPINA7 PE=1 SV=2 44 46295 1 1 2.4 5.87
127 Q9H2G4 Testis-specific Y-encoded-like protein 2 OS=Homo sapiens GN=TSPYL2 PE=1 SV=1 44 79387 1 1 1 4.45
128 P25787 Proteasome subunit alpha type-2 OS=Homo sapiens GN=PSMA2 PE=1 SV=2 44 25882 2 1 9 6.92
129 P02461 Collagen alpha-1(III) chain OS=Homo sapiens GN=COL3A1 PE=1 SV=4 43 138479 3 2 1.4 6.21
130 P81605 Dermcidin OS=Homo sapiens GN=DCD PE=1 SV=2 43 11277 2 1 10 6.08
131 Q9Y6X9_REVERSED MORC family CW-type zinc finger protein 2 OS=Homo sapiens GN=MORC2 PE=1 SV=2 - REVERSED 43 117750 2 1 1.6 8.6
132 Q8NBM4 Ubiquitin-associated domain-containing protein 2 OS=Homo sapiens GN=UBAC2 PE=2 SV=1 43 38938 4 1 2.3 9.21
133 P10643 Complement component C7 OS=Homo sapiens GN=C7 PE=1 SV=2 43 93457 2 1 2 6.09
134 Q12968_REVERSED Nuclear factor of activated T-cells, cytoplasmic 3 OS=Homo sapiens GN=NFATC3 PE=1 SV=1 - REVERSED 43 115522 3 1 0.7 5.91
135 P32119 Peroxiredoxin-2 OS=Homo sapiens GN=PRDX2 PE=1 SV=5 42 21878 1 1 9.1 5.66
136 Q9UGM3 Deleted in malignant brain tumors 1 protein OS=Homo sapiens GN=DMBT1 PE=1 SV=2 42 260569 1 1 0.2 5.18
137 O75460_REVERSED Serine/threonine-protein kinase/endoribonuclease IRE1 OS=Homo sapiens GN=ERN1 PE=1 SV=2 - REVERSED 41 109666 1 1 0.6 5.98
138 Q9UKT8_REVERSED F-box/WD repeat-containing protein 2 OS=Homo sapiens GN=FBXW2 PE=1 SV=2 - REVERSED 41 51479 2 1 3.1 6.11
139 O95445 Apolipoprotein M OS=Homo sapiens GN=APOM PE=1 SV=2 41 21239 2 1 6.9 5.66
140 P00558 Phosphoglycerate kinase 1 OS=Homo sapiens GN=PGK1 PE=1 SV=3 40 44586 2 1 8.4 8.3
141 O14578_REVERSED Citron Rho-interacting kinase OS=Homo sapiens GN=CIT PE=1 SV=2 - REVERSED 40 231286 2 1 0.5 6.16
142 Q7Z5M8 Abhydrolase domain-containing protein 12B OS=Homo sapiens GN=ABHD12B PE=2 SV=1 40 40750 3 1 1.7 8.57
143 Q15020_REVERSED Squamous cell carcinoma antigen recognized by T-cells 3 OS=Homo sapiens GN=SART3 PE=1 SV=1 - REVERSED 40 109865 3 1 0.6 5.45
144 P29803_REVERSED Pyruvate dehydrogenase E1 component subunit alpha, testis-specific form, mitochondrial OS=Homo sapiens GN=PDHA2 PE=1 SV=1 - REVERSED 40 42906 2 1 1.8 8.76
145 Q14914 Prostaglandin reductase 1 OS=Homo sapiens GN=PTGR1 PE=1 SV=2 40 35847 1 1 2.1 8.45
146 Q9NQX4 Myosin-Vc OS=Homo sapiens GN=MYO5C PE=1 SV=2 40 202682 1 1 0.4 7.69
147 Q13183_REVERSED Solute carrier family 13 member 2 OS=Homo sapiens GN=SLC13A2 PE=2 SV=1 - REVERSED 40 64368 2 1 1.4 6.61
148 A8MPT4 Glutathione S-transferase theta-4 OS=Homo sapiens GN=GSTT4 PE=3 SV=2 39 27941 2 1 2.5 6.25
149 Q8N6V9_REVERSED Testis-expressed sequence 9 protein OS=Homo sapiens GN=TEX9 PE=2 SV=1 - REVERSED 39 44798 2 1 1.5 6.2
150 Q9Y2U8_REVERSED Inner nuclear membrane protein Man1 OS=Homo sapiens GN=LEMD3 PE=1 SV=2 - REVERSED 39 99935 2 1 0.7 7.31
151 P05154 Plasma serine protease inhibitor OS=Homo sapiens GN=SERPINA5 PE=1 SV=3 39 45646 2 1 1.5 9.3
152 P11137 Microtubule-associated protein 2 OS=Homo sapiens GN=MAP2 PE=1 SV=4 39 199404 4 1 1.4 4.82
153 Q9BY43 Charged multivesicular body protein 4a OS=Homo sapiens GN=CHMP4A PE=1 SV=3 39 25083 1 1 3.2 4.65
154 P22303 Acetylcholinesterase OS=Homo sapiens GN=ACHE PE=1 SV=1 39 67753 1 1 2.1 5.87
155 P07864 L-lactate dehydrogenase C chain OS=Homo sapiens GN=LDHC PE=2 SV=4 38 36288 3 1 6.6 7.08
156 P28066 Proteasome subunit alpha type-5 OS=Homo sapiens GN=PSMA5 PE=1 SV=3 38 26394 2 1 12 4.74
157 Q14C86_REVERSED GTPase-activating protein and VPS9 domain-containing protein 1 OS=Homo sapiens GN=GAPVD1 PE=1 SV=2 - REVERSED 38 164876 2 1 0.5 5.09
158 O75808 Calpain-15 OS=Homo sapiens GN=SOLH PE=1 SV=1 38 117239 1 1 0.6 6.27
159 Q15751_REVERSED Probable E3 ubiquitin-protein ligase HERC1 OS=Homo sapiens GN=HERC1 PE=1 SV=2 - REVERSED 38 531891 1 1 0.2 5.69
160 Q6R2W3_REVERSED SCAN domain-containing protein 3 OS=Homo sapiens GN=SCAND3 PE=2 SV=1 - REVERSED 38 151570 3 1 0.7 6.3
161 Q6ZNE9_REVERSED RUN and FYVE domain-containing protein 4 OS=Homo sapiens GN=RUFY4 PE=2 SV=2 - REVERSED 37 64309 1 1 1.4 6.43
162 Q8N7B9 EF-hand calcium-binding domain-containing protein 3 OS=Homo sapiens GN=EFCAB3 PE=1 SV=1 37 50114 3 1 2.1 9.31
163 O60312 Probable phospholipid-transporting ATPase VA OS=Homo sapiens GN=ATP10A PE=2 SV=2 37 167582 1 1 0.6 8.7
164 O15264 Mitogen-activated protein kinase 13 OS=Homo sapiens GN=MAPK13 PE=1 SV=1 37 42063 3 1 1.9 8.48
165 Q9H2K8 Serine/threonine-protein kinase TAO3 OS=Homo sapiens GN=TAOK3 PE=1 SV=2 37 105340 5 1 1.9 6.83
166 P48634 Protein PRRC2A OS=Homo sapiens GN=PRRC2A PE=1 SV=3 37 228724 4 1 0.3 9.48
167 Q8IVF6 Ankyrin repeat domain-containing protein 18A OS=Homo sapiens GN=ANKRD18A PE=1 SV=3 37 115525 4 1 0.7 7.92
168 P35580 Myosin-10 OS=Homo sapiens GN=MYH10 PE=1 SV=3 36 228858 1 1 0.6 5.44
169 P10242_REVERSED Transcriptional activator Myb OS=Homo sapiens GN=MYB PE=1 SV=2 - REVERSED 36 72296 1 1 1.3 6.35
170 Q9BV38 WD repeat-containing protein 18 OS=Homo sapiens GN=WDR18 PE=1 SV=2 36 47375 1 1 2.1 6.21
171 Q99816 Tumor susceptibility gene 101 protein OS=Homo sapiens GN=TSG101 PE=1 SV=2 60 43916 1 1 6.4 8.21

Extended Data Table 3.

Demographics of Patients and Healthy Participants 

DISCOVERY COHORT VALIDATION COHORT
No. of participants (n=321) % of participants No. of participants (n=82) % of participants No. of participants (n=32) % of participants
Pancreatic Cancer Pancreatic Cancer Breast cancer
Total 190 59.19% 56 68.29% 32 100%

Sex

 Men 104 54.74% 28 50.00% 0 0%

 Women 86 45.26% 28 50.00% 32 100%

 Median Age (range) 66 (37 – 86) 70 (40 – 85) 57 (30 – 85)

AJCC stage

 0 n.a. - n.a. - 2 6%

 I 2 1.05% 2 3.57% 12 38%

 II n.a - n.a - 17 53%

 IIa 19 10.00% 15 26.79% n.a. -

 IIb 117 61.58% 36 64.29% n.a. -

 III 11 5.79% 0 0.00% 1 3%

 IV 41 21.58% 3 5.36% n.a. -

Tumor grade

 1 1 0.53% 1 1.79% 8 25%

 2 91 47.89% 35 62.5% 13 41%

 3 49 25.79% 19 33.93% 10 31%

 4 1 0.53% 0 0.00% n.a. -

 Unknown 48 25.26% 1 1.79% 1 3%

Tumor resected

 Yes 152 80.00% 54 96.43% 32 100%

 No 38 20.00% 2 3.57% 0 0%

Neoadjuvant Radio- /Chemotherapy

 Received 10 5.26% 0 0.00% 0 0%

 Not received 180 94.74% 56 100.00% 32 100%

Benign Pancreatic disease (BPD)

Total 26 8.15%

Sex

 Men 18 69.23% 3 50.00%

 Women 8 30.77% 3 50.00%

 Median Age (range) 58.5 (31 – 77) 49 (43 – 56)

Diagnosis

 Chronic pancreatits 15 57.69% 6 100%

 Autoimmune pancreatitis 3 11.54% 0 0.00%

 Serous cystadenoma 8 30.77% 0 0.00%

Pancreatic cancer precursor lesion (PCPL)

Total 5 1.55% 0 0.00%

Sex

 Men 2 40.00% 0 0.00%

 Women 3 60.00% 0 0.00%

 Median Age (range) 65 (59 – 74) - - -

Neoplasms

* IPMN 5 100.0% 0 0.00%

Healthy donors

Total 100 31.15% 20 24.39%

n.a. - non applicable 

Abbreviations: American Joint Committee on Cancer (AJCC), Intraductal papillary mucinous neoplasm (IPMN), Pancreatic Intraepithelial Neoplasia (PanIN).

*

The group of IPMNs consist of 2 IPMN associated with a carcinoma in situ, 1 IPMN associated with an early adenocarcinoma of the pancreas (pT1), an IPMN with intermediate dysplasia and an IPMN with low-grade dysplasia.

Extended Data Table 4. Histological report of patients with chronic pancreatitis.

The histopathological report is listed for the 15 patients with chronic pancreatitis in the discovery cohort.

Patient No. PanIN described Histopathological report
1 No Pancreatic tissue with chronic pancreatitis and extensive fibrosis and focal necrosis lipolytic and triptolytic areas.
0 No Diffuse periductal lymphoplasmacytic infiltrates; severe periductal fibrosis and duct obstruction/disappearance; severe interlobular and acinar involvement; severe inflammatory storiform fibrosis and diffuse sclerosis; frequent venulitis and occasional arteritis; scattered and occasionally prominent lymphoid follicles
3 No Chronic pancreatitis with periductal, inter- and intralobular fibrosis
4 No Chronic pancreatitis and extensive fibrosis
5 No Chronic recurrent and acute pancreatitis with plurifocal tryptolytic and lipolytic necrosis
6 No Low-grade chronic pancreatitis with periductal fibrosis, in the present material no evidence of neoplastic events, no evidence of malignancy.
7 No Chronic recurrent pancreatitis with some more pronounced fibrosis and intraductal calcifications. Chronic pancreatitis extends to the pancreas resection margin.
8 No Chronic pancreatitis, cholangitis and papillitis with focally histomorphological aspect of an autoimmune, chronic sclerosing pancreatitis.
9 PanIN 1a Pancreatic parenchyma (head of the pancreas) and peripancreatic fat and connective tissue with chronic recurrent pancreatitis with some areas fibrosis and abscesses. Pancreatic intraepithelial neoplasia (PanIN) Grade 1A.
10 No Pancreatic parenchyma with perilobular fibrosis as well as dilated pancreatic ducts. In addition, peripancreatic fat and connective tissue with fibrosis. The finding represents a chronic pancreatitis.
11 PanIN 1a Chronic-recurrent pancreatitis with pronounced fibrosis and dilated pancreatic ducts with focal inflammatory reactive epithelial cells. Older areas of organized necrosis and focal pancreatic intraepithelial neoplasia (PanIN) Grade 1A. Chronic pancreatitis also affects the pancreas resection margin.
12 No Pancreatitis with focally accentuated, periductal, perilobular and intralobular fibrosis as well as smaller areas of organized fatty necrosis and presence of singel giant cells of foreign body type.
13 No Chronic recurrent and acute pancreatitis with plurifocal tryptolytic and lipolytic necrosis with extensive destruction of the pancreatic parenchyma. Smaller secretion - and obliteration of pancreatic ducts with periductal fibrosis and localized squamous metaplasia. Peri- and interlobular fibrosis of the pancreatic parenchyma. Pancreatitis reaches the resection margin. At present, no neoplastic tissue, no evidence of malignancy.
14 No Pancreatic tissue with some scarring chronic inflammation and chronic pancreatits. In the present material, no evidence of malignancy.
15 No Tumor -free pancreatic tissue (surgical margins) with low periductal and interlobular fibrosis.

Extended Data Table 5. Histopathological findings and scoring in PKT mice in the cross sectional study.

The mouse ID, age, genotype and percent bead with GPC1+ crExos of mice in the cross sectional study are listed. A description of the histopathological findings and associated histological score is listed for PKT mice. Score 1: PanIN1a, Score 2: PanIN1 a/b, Score 3: PanIN2, Score 4: PanIN3, Score 5: Ductal adenocarcinoma (DCA). P: present (lesions were detected). 0: no lesion detected.

Mouse ID Age (days) Sex Genotype GPC1 (%) Normal (% area) Involved (% area) PanIN1a PanIN1b PanIN2 PanIN3 DCA Other (defined) Pathological score
T187 16 M Healthy+control 0.6 100 0 0 0 0 0 0
T188 16 M Healthy+control 1.1 100 0 0 0 0 0 0
T157 20 M Healthy+control 0.2 100 0 0 0 0 0 0
T158 20 M Healthy+control 0.2 100 0 0 0 0 0 0
T163 20 M Healthy+control 0.2 100 0 0 0 0 0 0
T164 20 M Healthy+control 0.4 100 0 0 0 0 0 0
T183 16 M PKT 8.4 98.6 1.4 P 0 0 0 0 1
T190 16 F PKT 5.2 100.0 0.0 0 0 0 0 0 reactive+ducts 0
T191 16 F PKT 6.4 100.0 0.0 0 0 0 0 0 reactive+ducts 0
T156 20 M PKT 8.3 100.0 0.0 0 0 0 0 0 0
T159 20 F PKT 10.8 95.1 4.9 P P P 0 0 3
T160 20 F PKT 9.8 99.3 0.7 P P 0 0 0 2
T165 20 F PKT 9.2 96.1 3.9 P P 0 0 0 2

P : present 

0 : not detected 

Acknowledgments

This work was primarily supported by the Cancer Prevention and Research Institute of Texas and UT MD Anderson Cancer Center. RK is also supported by NIH Grants CA-155370, CA-151925, DK 081576 and Metastasis Research Center at the MD Anderson Cancer Center (P30CA016672). VSL is supported by the NIH/NCI under the award number P30CA016672 and the UT MDACC Khalifa Bin Zayed Al Nahya Foundation. DPW and STG are supported by the NIH P50-CA094056. Institutional Core Grant CA16672 for High Resolution Electron Microscopy Facility. The Flow Cytometry MDACC core facility is partially funded by NIH P30CA16672. The MDACC Small Animal Imaging Facility is partially funded by NIH grants P30-CA016672 and 5U24-CA126577. SAM is a Human Frontiers Science Program Fellow. CK is funded by a Research Fellowship of the Deutsche Forschungsgemeinschaf (DFG). We thank Paul A. Kurywchak for the help with immunoblots and sucrose gradients, and Sushrut Kamerkar for the help with the sucrose gradients. We thank Kenneth Dunner Jr. for the help with the transmission electron microscopy and immunogold techniques.

Footnotes

Supplementary Information is linked to the online version of the paper at www.nature.com/nature.

The authors declare no competing financial interests.

Author contribution: S.A.M. conceptually designed and carried out most of the experiments, generated the data and the figures, and wrote the manuscript; L.B.L. carried out most of the experiments; C.K. performed sequencing of primary tumors, performed most of the statistical analysis of the manuscript and support in manuscript writing; A.F.F. optimized, performed and analyzed the UPLC-MS data; S.T.G. performed mouse MRI and analyzed the data; J.K. GEMMs breeding, bleeding, euthanasia and material collection; V.S.L. GEMMs breeding, bleeding, euthanasia, material collection and support in manuscript editing; E.A.M. provided breast cancer patient samples and patient history; J.W., N.R., C.R. and C.P. collected and provided serum samples and patient history for analysis; M.F.F. optimized, performed and analyzed the UPLC-MS data; D.P.W. mouse MRI interpretation and provided support in data interpretation and analysis; R.K. conceived the idea, conceptually designed the study, supervised the project and wrote the manuscript.

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