Details for: SART1

Gene ID: 9092

Gene Type:  Protein-coding  - A gene that serves as a template for producing a messenger RNA (mRNA) molecule, which is then translated into a functional protein.

Symbol: SART1

Ensembl ID: ENSG00000175467

Description: spliceosome associated factor 1, recruiter of U4/U6.U5 tri-snRNP

Selected Context(s):  Overall

Cell Significance Landscape

Contexts:

Associated with

Significant Cells

Cell Significance Index (CSI) scores for the chosen context(s)

  • granulocyte CL0000094
    CSI 11.62
    rCSI 17.76%
    PRS 71.16
  • neural progenitor cell CL0011020
    CSI 8.07
    rCSI 35.5%
    PRS 52.23
  • extravillous trophoblast CL0008036
    CSI 7.49
    rCSI 9.26%
    PRS 58.39
  • central memory CD8-positive, alpha-beta T cell CL0000907
    CSI 5.6
    rCSI 3.78%
    PRS 74.93
  • alveolar adventitial fibroblast CL4028006
    CSI 5.06
    rCSI 8%
    PRS 64.16
  • multi-ciliated epithelial cell CL0005012
    CSI 5.04
    rCSI 5.03%
    PRS 55.08
  • CD4-positive, alpha-beta thymocyte CL0000810
    CSI 4.07
    rCSI 3.26%
    PRS 81.86
  • promonocyte CL0000559
    CSI 4.01
    rCSI 6.87%
    PRS 70.65
  • caudal ganglionic eminence derived cortical interneuron CL4023064
    CSI 3.4
    rCSI 6%
    PRS 42.65
  • interstitial cell of Cajal CL0002088
    CSI 3.34
    rCSI 4.25%
    PRS 68.27
  • unswitched memory B cell CL0000970
    CSI 3.29
    rCSI 2.77%
    PRS 78.76
  • hematopoietic stem cell CL0000037
    CSI 3.27
    rCSI 2.17%
    PRS 64.93
  • pro-B cell CL0000826
    CSI 3.09
    rCSI 2.56%
    PRS 64.03
  • epithelial cell of lung CL0000082
    CSI 3.02
    rCSI 2.5%
    PRS 61.04
  • precursor B cell CL0000817
    CSI 2.95
    rCSI 2.59%
    PRS 71.52
  • rod bipolar cell CL0000751
    CSI 2.92
    rCSI 5.25%
    PRS 54.92
  • double negative thymocyte CL0002489
    CSI 2.9
    rCSI 2.01%
    PRS 72.96
  • immature B cell CL0000816
    CSI 2.73
    rCSI 2.03%
    PRS 75.14
  • perivascular cell CL4033054
    CSI 2.65
    rCSI 3.63%
    PRS 67.81
  • CD4-positive, alpha-beta memory T cell CL0000897
    CSI 2.62
    rCSI 1.88%
    PRS 76.04
  • plasmacytoid dendritic cell, human CL0001058
    CSI 2.62
    rCSI 1.83%
    PRS 64.63
  • CD4-positive, CD25-positive, alpha-beta regulatory T cell CL0000792
    CSI 2.48
    rCSI 2.44%
    PRS 77.41
  • ON-bipolar cell CL0000749
    CSI 2.42
    rCSI 3.59%
    PRS 63.37
  • keratinocyte CL0000312
    CSI 2.37
    rCSI 1.99%
    PRS 66.15
  • CD4-positive helper T cell CL0000492
    CSI 2.37
    rCSI 1.79%
    PRS 75.51
  • mesenchymal cell CL0008019
    CSI 2.36
    rCSI 6%
    PRS 56.06
  • mature alpha-beta T cell CL0000791
    CSI 2.31
    rCSI 8.38%
    PRS 81.12
  • CD4-positive, alpha-beta cytotoxic T cell CL0000934
    CSI 2.24
    rCSI 3.08%
    PRS 81.12
  • granulocyte monocyte progenitor cell CL0000557
    CSI 2.24
    rCSI 1.94%
    PRS 66.8
  • promyelocyte CL0000836
    CSI 2.21
    rCSI 3.18%
    PRS 71.33
  • naive T cell CL0000898
    CSI 2.15
    rCSI 1.5%
    PRS 77.02
  • kidney loop of Henle thin descending limb epithelial cell CL1001111
    CSI 2.13
    rCSI 3.02%
    PRS 58.15
  • intermediate monocyte CL0002393
    CSI 2.06
    rCSI 3.12%
    PRS 66.02
  • early lymphoid progenitor CL0000936
    CSI 2.02
    rCSI 1.77%
    PRS 67.36
  • intestine goblet cell CL0019031
    CSI 2.01
    rCSI 1.79%
    PRS 59.79
  • neural crest cell CL0011012
    CSI 2.01
    rCSI 1.59%
    PRS 48.64
  • lung macrophage CL1001603
    CSI 2.01
    rCSI 4.48%
    PRS 69.61
  • ciliated cell CL0000064
    CSI 1.99
    rCSI 3.22%
    PRS 58.43
  • central memory CD4-positive, alpha-beta T cell CL0000904
    CSI 1.98
    rCSI 1.17%
    PRS 79.11
  • mesodermal cell CL0000222
    CSI 1.95
    rCSI 2.34%
    PRS 59.9
  • dendritic cell, human CL0001056
    CSI 1.94
    rCSI 2.99%
    PRS 70.89
  • colon epithelial cell CL0011108
    CSI 1.91
    rCSI 2%
    PRS 58.23
  • activated type II NK T cell CL0000931
    CSI 1.88
    rCSI 2.12%
    PRS 77.88
  • activated CD8-positive, alpha-beta T cell CL0000906
    CSI 1.82
    rCSI 1.79%
    PRS 82.88
  • transit amplifying cell of colon CL0009011
    CSI 1.8
    rCSI 2.12%
    PRS 64.14
  • ciliated epithelial cell CL0000067
    CSI 1.76
    rCSI 1.55%
    PRS 49.67
  • megakaryocyte-erythroid progenitor cell CL0000050
    CSI 1.74
    rCSI 1.57%
    PRS 58.67
  • ionocyte CL0005006
    CSI 1.7
    rCSI 1.82%
    PRS 61.21
  • myeloid leukocyte CL0000766
    CSI 1.66
    rCSI 1.53%
    PRS 63.23
  • lung ciliated cell CL1000271
    CSI 1.63
    rCSI 1.88%
    PRS 51.87
  • ependymal cell CL0000065
    CSI 1.62
    rCSI 3.29%
    PRS 41.24
  • elicited macrophage CL0000861
    CSI 1.62
    rCSI 1.48%
    PRS 70.71
  • goblet cell CL0000160
    CSI 1.6
    rCSI 1.51%
    PRS 61.37
  • neuroblast (sensu Nematoda and Protostomia) CL0000338
    CSI 1.55
    rCSI 1.8%
    PRS 54.99
  • neuroblast (sensu Vertebrata) CL0000031
    CSI 1.53
    rCSI 1.96%
    PRS 58.99
  • pvalb GABAergic cortical interneuron CL4023018
    CSI 1.49
    rCSI 1.86%
    PRS 41.68
  • retinal blood vessel endothelial cell CL0002585
    CSI 1.42
    rCSI 2.27%
    PRS 65.99
  • radial glial cell CL0000681
    CSI 1.41
    rCSI 1.96%
    PRS 60.4
  • common dendritic progenitor CL0001029
    CSI 1.34
    rCSI 1.68%
    PRS 72.22
  • CD14-low, CD16-positive monocyte CL0002396
    CSI 1.32
    rCSI 1.02%
    PRS 62.34
  • mature B cell CL0000785
    CSI 1.29
    rCSI 1.13%
    PRS 72.54
  • kidney connecting tubule epithelial cell CL1000768
    CSI 1.23
    rCSI 3.11%
    PRS 51.3
  • peripheral nervous system neuron CL2000032
    CSI 1.21
    rCSI 1.65%
    PRS 53.66
  • retinal cone cell CL0000573
    CSI 1.2
    rCSI 1.93%
    PRS 51.36
  • placental villous trophoblast CL2000060
    CSI 1.16
    rCSI 1.79%
    PRS 59.97
  • astrocyte of the cerebral cortex CL0002605
    CSI 1.15
    rCSI 2.58%
    PRS 44.33
  • duct epithelial cell CL0000068
    CSI 1.08
    rCSI 1.57%
    PRS 66.45
  • common myeloid progenitor CL0000049
    CSI 1.06
    rCSI 0.86%
    PRS 63.24
  • retina horizontal cell CL0000745
    CSI 1.06
    rCSI 1.61%
    PRS 58.1
  • L2/3-6 intratelencephalic projecting glutamatergic neuron CL4023040
    CSI 0.92
    rCSI 2.24%
    PRS 42.21
  • L5/6 near-projecting glutamatergic neuron CL4030067
    CSI 0.83
    rCSI 2.74%
    PRS 49.05
  • effector memory CD8-positive, alpha-beta T cell, terminally differentiated CL0001062
    CSI 0.71
    rCSI 3.57%
    PRS 74.31
  • pancreatic PP cell CL0002275
    CSI 0.49
    rCSI 1.95%
    PRS 74.35
  • L5 extratelencephalic projecting glutamatergic cortical neuron CL4023041
    CSI 0.42
    rCSI 1.49%
    PRS 41.98

Cell ID: Standard Cell Ontology term used for mapping and comparing cells across experiments. Ensures consistency in analyzing cellular functions across tissues.
Fold Change: Represents the ratio of the current Cell Significance Index to the Cell Significance Index Threshold, indicating how much the gene expression has changed compared to a baseline.
Cell Significance Index: Reflects how strongly a gene is expressed in this specific cell.

Cell ID: Standard Cell Ontology term used for mapping and comparing cells across experiments. Ensures consistency in analyzing cellular functions across tissues.
Fold Change: Represents the ratio of the current Cell Significance Index to the Cell Significance Index Threshold, indicating how much the gene expression has changed compared to a baseline.
Cell Significance Index: Reflects how strongly a gene is expressed in this cell type. Calculated using techniques like effect size estimation and bootstrapping for reliability.

Cell ID: Standard Cell Ontology term used for mapping and comparing cells across experiments. Ensures consistency in analyzing cellular functions across tissues.
Fold Change: Represents the ratio of the current Cell Significance Index to the Cell Significance Index Threshold, indicating how much the gene expression has changed compared to a baseline.
Cell Significance Index: Reflects how strongly a gene is expressed in this cell type. Calculated using techniques like effect size estimation and bootstrapping for reliability.
Network Configuration

Explore relationships of the current gene. Select an Interaction Source: 'ONTOLOGY' for shared pathways (GO/Reactome) or 'STRING' for protein-protein interactions. Further refine by selecting context genes and comparing Cell Significance Index (CSI) scores between baseline and target cell types and their specific contexts.

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Comma-separated if multiple.
Legend:
  • Query Gene
  • Node Color (Target Cell CSI, relative to current network):
    • Very High
    • High
    • Medium
    • Low
    • Very Low
    • CSI N/A
  • Node Size: Proportional to Target Cell CSI magnitude
  • STRING PPI Edge
  • Shared Pathway Edge (ONTOLOGY)

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Other Information

This section provides additional information about the gene, including a description generated by an AI language model and details about associated proteins.

## Summary [SART1](/details-gene/9092) (Spliceosome Associated Factor 1) is a protein-coding gene located on chromosome 11q13.1. It encodes a crucial component of the spliceosome, specifically the U4/U6.U5 tri-snRNP complex, which is essential for pre-mRNA splicing ([Link](https://doi.org/10.1093/emboj/20.10.2553)). Functionally, [SART1](/details-gene/9092) plays a fundamental role in RNA processing and metabolism. Expression data reveals its high significance in metabolically active and proliferating cells, particularly in immune cells such as [granulocytes](/details-cell/CL0000094) and T-lymphocytes, as well as in various progenitor cell types. Notably, beyond its housekeeping role in splicing, [SART1](/details-gene/9092) has been identified as a tumor antigen in squamous cell carcinoma, recognized by cytotoxic T lymphocytes, highlighting a dual role in both core cellular machinery and immuno-oncology ([Link](https://doi.org/10.1084/jem.187.3.277)). ## Cellular Roles and Expression Landscape **Overall**, the expression profile of [SART1](/details-gene/9092) suggests its importance in cells requiring high levels of protein synthesis, differentiation, and turnover, consistent with its fundamental role in mRNA processing. Its highest significance is observed in [granulocytes](/details-cell/CL0000094) (CSI: 11.62), indicating a critical function in the maturation or effector activities of these innate immune cells. The gene is also a prominent marker in various progenitor and developing cell populations, including [neural progenitor cells](/details-cell/CL0011020) (CSI: 8.07) and [hematopoietic stem cells](/details-cell/CL0000037) (CSI: 3.27). This pattern suggests that robust splicing activity, mediated by [SART1](/details-gene/9092), is essential for executing the complex gene expression programs that govern cell fate and differentiation. Furthermore, [SART1](/details-gene/9092) shows significant expression across multiple lymphocyte lineages. This includes high relevance in [central memory CD8-positive, alpha-beta T cells](/details-cell/CL0000907) (CSI: 5.60), developing [CD4-positive, alpha-beta thymocytes](/details-cell/CL0000810) (CSI: 4.07), and various stages of B cell development, such as [pro-B cells](/details-cell/CL0000826) (CSI: 3.09) and [unswitched memory B cells](/details-cell/CL0000970) (CSI: 3.29). This broad expression within the immune system is consistent with its documented role in T cell biology and its recognition as an antigen. ## Pathways and Molecular Function [SART1](/details-gene/9092) is integral to fundamental RNA processing pathways. Its primary molecular functions are [RNA binding](/details-cell/GO:0003723) and [protein binding](/details-cell/GO:0005515), which facilitate its role within the spliceosome. It is a key component of the [U4/u6 x u5 tri-snrnp complex](/details-cell/GO:0046540) and is found in nuclear structures such as [nuclear specks](/details-cell/GO:0016607) and [Cajal bodies](/details-cell/GO:0015030), where splicing factors are concentrated. The gene's involvement is annotated in several core biological processes, most notably [mRNA splicing, via spliceosome](/details-cell/GO:0000398) and [spliceosomal snRNP assembly](/details-cell/GO:0000387). These functions are central to the Reactome pathways [mRNA splicing - major pathway](/details-cell/R-HSA-72163) and the broader [Metabolism of rna](/details-cell/R-HSA-8953854). This essential housekeeping role in gene expression regulation likely underpins its widespread importance across diverse cell types. Intriguingly, beyond its canonical splicing function, [SART1](/details-gene/9092) is also associated with the [positive regulation of cytotoxic t cell differentiation](/details-cell/GO:0045585). This annotation, combined with research identifying it as a tumor rejection antigen ([Link](https://doi.org/10.1084/jem.187.3.277)), suggests that it may either be upregulated during T cell activation or that specific splice variants it produces are critical for T cell effector programming. ## Research Directions The dual identity of [SART1](/details-gene/9092) as both a ubiquitous splicing factor and a specific tumor antigen presents several compelling avenues for future research. The key is to understand how a core cellular component can also be immunogenic and what specific roles it plays in highly specialized cells like granulocytes and T cells. **Testable Hypotheses:** 1. **Role in T Cell Function:** The specific expression of [SART1](/details-gene/9092) in T cells and its link to cytotoxic T cell differentiation suggests it may regulate the alternative splicing of key immune transcripts (e.g., cytokine receptors, transcription factors, or signaling molecules) that are critical for T cell activation, memory formation, and effector function. 2. **Function in Progenitor Cell Fate:** High expression in [neural progenitor cells](/details-cell/CL0011020) and hematopoietic stem cells indicates that [SART1](/details-gene/9092) could be a master regulator of developmental splicing programs. Its activity may be required to generate specific splice isoforms of transcription factors that determine lineage commitment versus self-renewal. 3. **Granulocyte-Specific Splicing:** The top-ranking expression in [granulocytes](/details-cell/CL0000094) is notable. [SART1](/details-gene/9092) may be essential for the rapid, post-transcriptional regulation needed for granulocyte effector functions, such as producing splice variants of inflammatory mediators or receptors involved in phagocytosis and degranulation. **Proposed Experiment:** To test the hypothesis regarding its role in T cell function, a targeted experiment could involve the CRISPR/Cas9-mediated knockdown of [SART1](/details-gene/9092) in primary human CD8+ T cells. Following knockdown, these cells would be activated *in vitro* via TCR stimulation. A combination of deep RNA-sequencing and quantitative proteomics would be used to (a) identify global changes in alternative splicing events, particularly for genes involved in T cell cytotoxicity (e.g., *GZMB*, *PRF1*) and memory formation (e.g., *TCF7*, *IL7R*), and (b) assess the functional consequences on cytokine production (via ELISA) and target cell killing (via cytotoxicity assays). **Therapeutic Potential:** As a direct therapeutic target, inhibiting the core splicing function of [SART1](/details-gene/9092) would likely be highly toxic to healthy tissues. However, its established identity as a tumor antigen makes it an excellent candidate for cancer immunotherapy. The therapeutic strategy would focus on **activation** of the immune system against [SART1](/details-gene/9092)-expressing tumor cells. This could be pursued through peptide-based vaccines designed to elicit a [SART1](/details-gene/9092)-specific cytotoxic T lymphocyte response or through the development of adoptive T-cell therapies, such as CAR-T cells engineered to recognize [SART1](/details-gene/9092)-derived peptides presented on the surface of cancer cells.

Genular Protein ID: 3365242913

Symbol: SNUT1_HUMAN

Name: U4/U6.U5 tri-snRNP-associated protein 1

UniProtKB Accession Codes:

O43290 A6NDN1 Q53GB5

Database IDs:

AGR 1 Allergome 2 AlphaFoldDB 1 Antibodypedia 1 Bgee 1 BioGRID 1 BioGRID-ORCS 1 BioMuta 1 CCDS 1 CORUM 1 CTD 1 ChiTaRS 1 ComplexPortal 1 DIP 1 DNASU 1 DisGeNET 1 EMBL 7 EMDB 25 Ensembl 1 ExpressionAtlas 1 GO 16 GeneCards 1 GeneTree 1 GeneWiki 1 GenomeRNAi 1 GlyGen 1 HGNC 1 HOGENOM 1 HPA 1 InParanoid 1 IntAct 1 InterPro 2 KEGG 1 MANE-Select 1 MIM 1 MINT 1 MassIVE 1 MetOSite 1 NCBI Taxonomy 1 OMA 1 OpenTargets 1 OrthoDB 1 PANTHER 2 PDB 26 PDBsum 26 PIR 1 PRO 1 PathwayCommons 1 PaxDb 1 PeptideAtlas 1 Pfam 2 PharmGKB 1 Pharos 1 PhosphoSitePlus 1 PhylomeDB 1 Proteomes 1 ProteomicsDB 1 Pumba 1 RNAct 1 Reactome 1 RefSeq 1 SIGNOR 1 SMR 1 STRING 1 SignaLink 1 SwissPalm 1 TreeFam 1 UCSC 1 VEuPathDB 1 eggNOG 1 iPTMnet 1 jPOST 1 neXtProt 1

Citations:

PubMed ID: 9449708

Title: A gene encoding antigenic peptides of human squamous cell carcinoma recognized cytotoxic T lymphocytes.

PubMed ID: 9449708

DOI: 10.1084/jem.187.3.277

PubMed ID: 11350945

Title: The 65 and 110 kDa SR-related proteins of the U4/U6*U5 tri-snRNP are essential for the assembly of mature spliceosomes.

PubMed ID: 11350945

DOI: 10.1093/emboj/20.10.2553

PubMed ID: 16554811

Title: Human chromosome 11 DNA sequence and analysis including novel gene identification.

PubMed ID: 16554811

DOI: 10.1038/nature04632

PubMed ID: 15489334

Title: The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC).

PubMed ID: 15489334

DOI: 10.1101/gr.2596504

PubMed ID: 11410364

Title: Intron loss in the SART1 genes of Fugu rubripes and Tetraodon nigroviridis.

PubMed ID: 11410364

DOI: 10.1016/s0378-1119(01)00504-2

PubMed ID: 9856836

Title: Molecular characterization of an autoallergen, Hom s 1, identified by serum IgE from atopic dermatitis patients.

PubMed ID: 9856836

DOI: 10.1046/j.1523-1747.1998.00413.x

PubMed ID: 11991638

Title: Purification and characterization of native spliceosomes suitable for three-dimensional structural analysis.

PubMed ID: 11991638

DOI: 10.1017/s1355838202021088

PubMed ID: 17081983

Title: Global, in vivo, and site-specific phosphorylation dynamics in signaling networks.

PubMed ID: 17081983

DOI: 10.1016/j.cell.2006.09.026

PubMed ID: 16723661

Title: The network of protein-protein interactions within the human U4/U6.U5 tri-snRNP.

PubMed ID: 16723661

DOI: 10.1261/rna.55406

PubMed ID: 17525332

Title: ATM and ATR substrate analysis reveals extensive protein networks responsive to DNA damage.

PubMed ID: 17525332

DOI: 10.1126/science.1140321

PubMed ID: 18220336

Title: Combining protein-based IMAC, peptide-based IMAC, and MudPIT for efficient phosphoproteomic analysis.

PubMed ID: 18220336

DOI: 10.1021/pr0705441

PubMed ID: 18669648

Title: A quantitative atlas of mitotic phosphorylation.

PubMed ID: 18669648

DOI: 10.1073/pnas.0805139105

PubMed ID: 19413330

Title: Lys-N and trypsin cover complementary parts of the phosphoproteome in a refined SCX-based approach.

PubMed ID: 19413330

DOI: 10.1021/ac9004309

PubMed ID: 19690332

Title: Quantitative phosphoproteomic analysis of T cell receptor signaling reveals system-wide modulation of protein-protein interactions.

PubMed ID: 19690332

DOI: 10.1126/scisignal.2000007

PubMed ID: 20068231

Title: Quantitative phosphoproteomics reveals widespread full phosphorylation site occupancy during mitosis.

PubMed ID: 20068231

DOI: 10.1126/scisignal.2000475

PubMed ID: 21269460

Title: Initial characterization of the human central proteome.

PubMed ID: 21269460

DOI: 10.1186/1752-0509-5-17

PubMed ID: 21406692

Title: System-wide temporal characterization of the proteome and phosphoproteome of human embryonic stem cell differentiation.

PubMed ID: 21406692

DOI: 10.1126/scisignal.2001570

PubMed ID: 23186163

Title: Toward a comprehensive characterization of a human cancer cell phosphoproteome.

PubMed ID: 23186163

DOI: 10.1021/pr300630k

PubMed ID: 24275569

Title: An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome.

PubMed ID: 24275569

DOI: 10.1016/j.jprot.2013.11.014

PubMed ID: 25218447

Title: Uncovering global SUMOylation signaling networks in a site-specific manner.

PubMed ID: 25218447

DOI: 10.1038/nsmb.2890

PubMed ID: 25114211

Title: Mapping of SUMO sites and analysis of SUMOylation changes induced by external stimuli.

PubMed ID: 25114211

DOI: 10.1073/pnas.1413825111

PubMed ID: 25092792

Title: UBL5 is essential for pre-mRNA splicing and sister chromatid cohesion in human cells.

PubMed ID: 25092792

DOI: 10.15252/embr.201438679

PubMed ID: 25772364

Title: SUMO-2 orchestrates chromatin modifiers in response to DNA damage.

PubMed ID: 25772364

DOI: 10.1016/j.celrep.2015.02.033

PubMed ID: 25755297

Title: System-wide analysis of SUMOylation dynamics in response to replication stress reveals novel small ubiquitin-like modified target proteins and acceptor lysines relevant for genome stability.

PubMed ID: 25755297

DOI: 10.1074/mcp.o114.044792

PubMed ID: 28112733

Title: Site-specific mapping of the human SUMO proteome reveals co-modification with phosphorylation.

PubMed ID: 28112733

DOI: 10.1038/nsmb.3366

PubMed ID: 30538201

Title: Structural-functional interactions of NS1-BP protein with the splicing and mRNA export machineries for viral and host gene expression.

PubMed ID: 30538201

DOI: 10.1073/pnas.1818012115

Sequence Information:

  • Length: 800
  • Mass: 90255
  • Checksum: B13525E80640DF9E
  • Sequence:
    • MGSSKKHRGE KEAAGTTAAA GTGGATEQPP RHREHKKHKH RSGGSGGSGG ERRKRSRERG 
GERGSGRRGA EAEARSSTHG RERSQAEPSE RRVKREKRDD GYEAAASSKT SSGDASSLSI 
EETNKLRAKL GLKPLEVNAI KKEAGTKEEP VTADVINPMA LRQREELREK LAAAKEKRLL 
NQKLGKIKTL GEDDPWLDDT AAWIERSRQL QKEKDLAEKR AKLLEEMDQE FGVSTLVEEE 
FGQRRQDLYS ARDLQGLTVE HAIDSFREGE TMILTLKDKG VLQEEEDVLV NVNLVDKERA 
EKNVELRKKK PDYLPYAEDE SVDDLAQQKP RSILSKYDEE LEGERPHSFR LEQGGTADGL 
RERELEEIRA KLRLQAQSLS TVGPRLASEY LTPEEMVTFK KTKRRVKKIR KKEKEVVVRA 
DDLLPLGDQT QDGDFGSRLR GRGRRRVSEV EEEKEPVPQP LPSDDTRVEN MDISDEEEGG 
APPPGSPQVL EEDEAELELQ KQLEKGRRLR QLQQLQQLRD SGEKVVEIVK KLESRQRGWE 
EDEDPERKGA IVFNATSEFC RTLGEIPTYG LAGNREEQEE LMDFERDEER SANGGSESDG 
EENIGWSTVN LDEEKQQQDF SASSTTILDE EPIVNRGLAA ALLLCQNKGL LETTVQKVAR 
VKAPNKSLPS AVYCIEDKMA IDDKYSRREE YRGFTQDFKE KDGYKPDVKI EYVDETGRKL 
TPKEAFRQLS HRFHGKGSGK MKTERRMKKL DEEALLKKMS SSDTPLGTVA LLQEKQKAQK 
TPYIVLSGSG KSMNANTITK