Stk16 (serine/threonine kinase 16) - Rat Genome Database

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Gene: Stk16 (serine/threonine kinase 16) Rattus norvegicus
Analyze
Symbol: Stk16
Name: serine/threonine kinase 16
RGD ID: 629474
Description: Enables RNA polymerase II cis-regulatory region sequence-specific DNA binding activity and protein kinase activity. Involved in cellular response to transforming growth factor beta stimulus; positive regulation of transcription by RNA polymerase II; and protein autophosphorylation. Predicted to be located in several cellular components, including Golgi-associated vesicle; cytosol; and nucleoplasm. Predicted to be active in Golgi apparatus. Orthologous to human STK16 (serine/threonine kinase 16); INTERACTS WITH (+)-schisandrin B; 17beta-estradiol 3-benzoate; 2,3,7,8-tetrachlorodibenzodioxine.
Type: protein-coding
RefSeq Status: PROVISIONAL
Previously known as: F52; MPSK; myristoylated and palmitoylated serine/threonine-protein kinase; PKL12; protein kinase PKL12; serine/threonine-protein kinase 16; TGF-beta-stimulated factor 1; TSF-1; tyrosine-protein kinase STK16
RGD Orthologs
Human
Mouse
Chinchilla
Bonobo
Dog
Squirrel
Pig
Green Monkey
Naked Mole-Rat
Alliance Genes
More Info more info ...
Latest Assembly: mRatBN7.2 - mRatBN7.2 Assembly
Position:
Rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
mRatBN7.2976,705,591 - 76,708,859 (+)NCBImRatBN7.2mRatBN7.2
mRatBN7.2 Ensembl976,705,602 - 76,708,855 (+)EnsemblmRatBN7.2 Ensembl
UTH_Rnor_SHR_Utx985,150,423 - 85,153,654 (+)NCBIRnor_SHR
UTH_Rnor_SHRSP_BbbUtx_1.0990,279,314 - 90,282,545 (+)NCBIRnor_SHRSP
UTH_Rnor_WKY_Bbb_1.0988,665,529 - 88,668,760 (+)NCBIRnor_WKY
Rnor_6.0982,411,010 - 82,414,249 (+)NCBIRnor6.0Rnor_6.0rn6Rnor6.0
Rnor_6.0 Ensembl982,411,013 - 82,414,240 (+)EnsemblRnor6.0rn6Rnor6.0
Rnor_5.0982,180,271 - 82,183,510 (+)NCBIRnor5.0Rnor_5.0rn5Rnor5.0
RGSC_v3.4974,491,917 - 74,495,148 (+)NCBIRGSC3.4RGSC_v3.4rn4RGSC3.4
RGSC_v3.1974,638,898 - 74,642,126 (+)NCBI
Celera974,276,032 - 74,279,263 (+)NCBICelera
Cytogenetic Map9q33NCBI
JBrowse: View Region in Genome Browser (JBrowse)
Model


Gene Ontology Annotations     Click to see Annotation Detail View

Cellular Component

References

References - curated
# Reference Title Reference Citation
1. Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Gaudet P, etal., Brief Bioinform. 2011 Sep;12(5):449-62. doi: 10.1093/bib/bbr042. Epub 2011 Aug 27.
2. Rat ISS GO annotations from MGI mouse gene data--August 2006 MGD data from the GO Consortium
3. A novel transcriptional factor with Ser/Thr kinase activity involved in the transforming growth factor (TGF)-beta signalling pathway. Ohta S, etal., Biochem J 2000 Sep 1;350 Pt 2:395-404.
4. GOA pipeline RGD automated data pipeline
5. Data Import for Chemical-Gene Interactions RGD automated import pipeline for gene-chemical interactions
Additional References at PubMed
PMID:9712705   PMID:16310770   PMID:18184589  


Genomics

Comparative Map Data
Stk16
(Rattus norvegicus - Norway rat)
Rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
mRatBN7.2976,705,591 - 76,708,859 (+)NCBImRatBN7.2mRatBN7.2
mRatBN7.2 Ensembl976,705,602 - 76,708,855 (+)EnsemblmRatBN7.2 Ensembl
UTH_Rnor_SHR_Utx985,150,423 - 85,153,654 (+)NCBIRnor_SHR
UTH_Rnor_SHRSP_BbbUtx_1.0990,279,314 - 90,282,545 (+)NCBIRnor_SHRSP
UTH_Rnor_WKY_Bbb_1.0988,665,529 - 88,668,760 (+)NCBIRnor_WKY
Rnor_6.0982,411,010 - 82,414,249 (+)NCBIRnor6.0Rnor_6.0rn6Rnor6.0
Rnor_6.0 Ensembl982,411,013 - 82,414,240 (+)EnsemblRnor6.0rn6Rnor6.0
Rnor_5.0982,180,271 - 82,183,510 (+)NCBIRnor5.0Rnor_5.0rn5Rnor5.0
RGSC_v3.4974,491,917 - 74,495,148 (+)NCBIRGSC3.4RGSC_v3.4rn4RGSC3.4
RGSC_v3.1974,638,898 - 74,642,126 (+)NCBI
Celera974,276,032 - 74,279,263 (+)NCBICelera
Cytogenetic Map9q33NCBI
STK16
(Homo sapiens - human)
Human AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCh382219,245,479 - 219,250,337 (+)NCBIGRCh38GRCh38hg38GRCh38
GRCh38.p13 Ensembl2219,245,455 - 219,250,337 (+)EnsemblGRCh38hg38GRCh38
GRCh372220,110,201 - 220,115,059 (+)NCBIGRCh37GRCh37hg19GRCh37
Build 362219,818,436 - 219,821,873 (+)NCBINCBI36Build 36hg18NCBI36
Build 342219,935,728 - 219,939,133NCBI
Celera2213,879,899 - 213,884,766 (+)NCBICelera
Cytogenetic Map2q35NCBI
HuRef2211,963,107 - 211,967,974 (+)NCBIHuRef
CHM1_12220,116,035 - 220,120,902 (+)NCBICHM1_1
T2T-CHM13v2.02219,730,246 - 219,735,104 (+)NCBIT2T-CHM13v2.0
Stk16
(Mus musculus - house mouse)
Mouse AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCm39175,187,463 - 75,192,250 (+)NCBIGRCm39GRCm39mm39
GRCm39 Ensembl175,187,482 - 75,192,250 (+)EnsemblGRCm39 Ensembl
GRCm38175,210,819 - 75,215,606 (+)NCBIGRCm38GRCm38mm10GRCm38
GRCm38.p6 Ensembl175,210,838 - 75,215,606 (+)EnsemblGRCm38mm10GRCm38
MGSCv37175,207,404 - 75,212,181 (+)NCBIGRCm37MGSCv37mm9NCBIm37
MGSCv36175,094,003 - 75,098,754 (+)NCBIMGSCv36mm8
Celera175,701,740 - 75,706,517 (+)NCBICelera
Cytogenetic Map1C4NCBI
Stk16
(Chinchilla lanigera - long-tailed chinchilla)
Chinchilla AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
ChiLan1.0 EnsemblNW_00495545314,182,066 - 14,185,584 (-)EnsemblChiLan1.0
ChiLan1.0NW_00495545314,182,167 - 14,185,569 (-)NCBIChiLan1.0ChiLan1.0
STK16
(Pan paniscus - bonobo/pygmy chimpanzee)
Bonobo AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
PanPan1.12B225,093,621 - 225,098,504 (+)NCBIpanpan1.1PanPan1.1panPan2
PanPan1.1 Ensembl2B225,093,645 - 225,098,504 (+)Ensemblpanpan1.1panPan2
Mhudiblu_PPA_v02B106,502,953 - 106,506,470 (+)NCBIMhudiblu_PPA_v0Mhudiblu_PPA_v0panPan3
STK16
(Canis lupus familiaris - dog)
Dog AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
CanFam3.13725,793,993 - 25,797,632 (+)NCBICanFam3.1CanFam3.1canFam3CanFam3.1
CanFam3.1 Ensembl3725,794,382 - 25,797,587 (+)EnsemblCanFam3.1canFam3CanFam3.1
Dog10K_Boxer_Tasha3726,619,873 - 26,623,533 (+)NCBIDog10K_Boxer_Tasha
ROS_Cfam_1.03725,809,201 - 25,812,862 (+)NCBIROS_Cfam_1.0
ROS_Cfam_1.0 Ensembl3725,809,595 - 25,812,860 (+)EnsemblROS_Cfam_1.0 Ensembl
UMICH_Zoey_3.13725,713,591 - 25,717,250 (+)NCBIUMICH_Zoey_3.1
UNSW_CanFamBas_1.03725,648,706 - 25,652,368 (+)NCBIUNSW_CanFamBas_1.0
UU_Cfam_GSD_1.03725,669,607 - 25,673,268 (+)NCBIUU_Cfam_GSD_1.0
Stk16
(Ictidomys tridecemlineatus - thirteen-lined ground squirrel)
Squirrel AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
HiC_Itri_2NW_024405303175,321,165 - 175,324,867 (+)NCBIHiC_Itri_2
SpeTri2.0 EnsemblNW_0049365691,663,634 - 1,668,521 (+)EnsemblSpeTri2.0
SpeTri2.0NW_0049365691,664,031 - 1,667,648 (+)NCBISpeTri2.0SpeTri2.0SpeTri2.0
STK16
(Sus scrofa - pig)
Pig AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
Sscrofa11.1 Ensembl15121,284,920 - 121,288,473 (+)EnsemblSscrofa11.1susScr11Sscrofa11.1
Sscrofa11.115121,285,065 - 121,288,475 (+)NCBISscrofa11.1Sscrofa11.1susScr11Sscrofa11.1
Sscrofa10.215134,319,991 - 134,323,453 (+)NCBISscrofa10.2Sscrofa10.2susScr3
STK16
(Chlorocebus sabaeus - green monkey)
Green Monkey AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
ChlSab1.110105,149,953 - 105,153,553 (+)NCBIChlSab1.1ChlSab1.1chlSab2
ChlSab1.1 Ensembl10105,150,421 - 105,153,693 (+)EnsemblChlSab1.1ChlSab1.1 EnsemblchlSab2
Vero_WHO_p1.0NW_02366604094,242,891 - 94,246,400 (-)NCBIVero_WHO_p1.0Vero_WHO_p1.0
Stk16
(Heterocephalus glaber - naked mole-rat)
Naked Mole-rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
HetGla_female_1.0 EnsemblNW_0046248235,763,171 - 5,770,587 (-)EnsemblHetGla_female_1.0HetGla_female_1.0 EnsemblhetGla2
HetGla 1.0NW_0046248235,765,284 - 5,770,778 (-)NCBIHetGla_female_1.0HetGla 1.0hetGla2

Variants

.
Variants in Stk16
8 total Variants
miRNA Target Status

Predicted Target Of
Summary Value
Count of predictions:160
Count of miRNA genes:121
Interacting mature miRNAs:139
Transcripts:ENSRNOT00000026194
Prediction methods:Microtar, Miranda, Rnahybrid, Targetscan
Result types:miRGate_prediction

The detailed report is available here: Full Report CSV TAB Printer

miRNA Target Status data imported from miRGate (http://mirgate.bioinfo.cnio.es/).
For more information about miRGate, see PMID:25858286 or access the full paper here.


QTLs in Region (mRatBN7.2)
The following QTLs overlap with this region.    Full Report CSV TAB Printer Gviewer
RGD IDSymbolNameLODP ValueTraitSub TraitChrStartStopSpecies
10054125Srcrt7Stress Responsive Cort QTL 73.330.0011blood corticosterone amount (VT:0005345)plasma corticosterone level (CMO:0001173)9187073594Rat
70186Niddm26Non-insulin dependent diabetes mellitus QTL 263.87blood glucose amount (VT:0000188)blood glucose level area under curve (AUC) (CMO:0000350)92207116986369743Rat
7207814Bmd91Bone mineral density QTL 913.5femur size trait (VT:1000369)femoral neck cross-sectional area (CMO:0001697)92375414483851531Rat
70218Cm28Cardiac mass QTL 288.30.0001heart mass (VT:0007028)heart wet weight (CMO:0000069)92526804479271759Rat
724544Uae9Urinary albumin excretion QTL 94.5urine albumin amount (VT:0002871)urine albumin level (CMO:0000130)925268044114175309Rat
731164Uae25Urinary albumin excretion QTL 253.50.0001urine albumin amount (VT:0002871)urine albumin excretion rate (CMO:0000757)925661188100929786Rat
7411656Foco26Food consumption QTL 269.80.001eating behavior trait (VT:0001431)feed conversion ratio (CMO:0001312)93253550577535505Rat
7411571Bw138Body weight QTL 13814.30.001body mass (VT:0001259)body weight gain (CMO:0000420)93253550577535505Rat
1598834Memor11Memory QTL 112.5exploratory behavior trait (VT:0010471)average horizontal distance between subject and target during voluntary locomotion in an experimental apparatus (CMO:0002674)93696235977814038Rat
8662828Vetf6Vascular elastic tissue fragility QTL 63.9artery integrity trait (VT:0010639)patent ductus arteriosus score (CMO:0002566)93696235992058970Rat
2290450Scl57Serum cholesterol level QTL 574.15blood cholesterol amount (VT:0000180)plasma total cholesterol level (CMO:0000585)93696235995410867Rat
6903941Pur31Proteinuria QTL 310.036urine total protein amount (VT:0000032)urine protein excretion rate (CMO:0000759)94019418885194188Rat
11353949Bp393Blood pressure QTL 393arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)94019418885194188Rat
61352Bp34Blood pressure QTL 345arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)94249534379271511Rat
10058949Gmadr5Adrenal mass QTL 520.014adrenal gland mass (VT:0010420)both adrenal glands wet weight to body weight ratio (CMO:0002411)94279151387976209Rat
11353951Bp394Blood pressure QTL 394arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)94464992189649921Rat
12879506Pur33Proteinuria QTL 33urine total protein amount (VT:0000032)urine total protein excretion rate (CMO:0000756)94464992189649921Rat
11353957Bmd92Bone mineral density QTL 920.01tibia mineral mass (VT:1000283)volumetric bone mineral density (CMO:0001553)94611419991114199Rat
631656Bp108Blood pressure QTL 1085.970.0001arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)94859825193598251Rat
2303170Bp332Blood pressure QTL 3323.730.027arterial blood pressure trait (VT:2000000)mean arterial blood pressure (CMO:0000009)95584784177026453Rat
2303180Bp333Blood pressure QTL 3330.001arterial blood pressure trait (VT:2000000)mean arterial blood pressure (CMO:0000009)95662771378595166Rat
2303180Bp333Blood pressure QTL 3330.001arterial blood pressure trait (VT:2000000)diastolic blood pressure (CMO:0000005)95662771378595166Rat
2303180Bp333Blood pressure QTL 3330.001arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)95662771378595166Rat
1578760Cm53Cardiac mass QTL 533.30.0001heart mass (VT:0007028)heart wet weight (CMO:0000069)956771635101771635Rat
724515Uae16Urinary albumin excretion QTL 168urine albumin amount (VT:0002871)urine albumin excretion rate (CMO:0000757)958163035100929646Rat
1300134Bp185Blood pressure QTL 1853.73arterial blood pressure trait (VT:2000000)mean arterial blood pressure (CMO:0000009)961381434104821652Rat
1581580Uae34Urinary albumin excretion QTL 34urine albumin amount (VT:0002871)urine albumin excretion rate (CMO:0000757)96207227596470995Rat
1578757Pur6Proteinuria QTL 63.30.005urine total protein amount (VT:0000032)urine total protein excretion rate (CMO:0000756)96207227596470995Rat
61385Edpm9Estrogen-dependent pituitary mass QTL 93.430.05pituitary gland mass (VT:0010496)pituitary gland wet weight (CMO:0000853)963869687108869687Rat
731171Glom6Glomerulus QTL 62.80.0003kidney glomerulus morphology trait (VT:0005325)count of superficial glomeruli not directly contacting the kidney surface (CMO:0001002)964573531109573531Rat
1354626Bvd1Brain ventricular dilatation QTL 13.730.001brain ventricle morphology trait (VT:0000822)hydrocephalus severity score (CMO:0001881)975712843111552878Rat
1581517Bp284Blood pressure QTL 284arterial blood pressure trait (VT:2000000)systolic blood pressure (CMO:0000004)97665062976771824Rat

Markers in Region
RH130189  
Rat AssemblyChrPosition (strand)SourceJBrowse
mRatBN7.2976,708,639 - 76,708,830 (+)MAPPERmRatBN7.2
Rnor_6.0982,414,034 - 82,414,224NCBIRnor6.0
Rnor_5.0982,183,295 - 82,183,485UniSTSRnor5.0
RGSC_v3.4974,494,938 - 74,495,128UniSTSRGSC3.4
Celera974,279,053 - 74,279,243UniSTS
RH 3.4 Map9757.89UniSTS
Cytogenetic Map9q33UniSTS


Expression


RNA-SEQ Expression
High: > 1000 TPM value   Medium: Between 11 and 1000 TPM
Low: Between 0.5 and 10 TPM   Below Cutoff: < 0.5 TPM

alimentary part of gastrointestinal system circulatory system endocrine system exocrine system hemolymphoid system hepatobiliary system integumental system musculoskeletal system nervous system renal system reproductive system respiratory system appendage
High
Medium 3 36 57 41 19 41 1 4 74 35 41 11 1
Low 7 7 7 7
Below cutoff

Sequence


Reference Sequences
RefSeq Acc Id: ENSRNOT00000026194   ⟹   ENSRNOP00000026194
RefSeq Status:
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2 Ensembl976,705,602 - 76,708,855 (+)Ensembl
Rnor_6.0 Ensembl982,411,013 - 82,414,240 (+)Ensembl
RefSeq Acc Id: ENSRNOT00000111780   ⟹   ENSRNOP00000077839
RefSeq Status:
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2 Ensembl976,705,619 - 76,708,855 (+)Ensembl
RefSeq Acc Id: NM_173142   ⟹   NP_775165
RefSeq Status: PROVISIONAL
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2976,705,619 - 76,708,850 (+)NCBI
Rnor_6.0982,411,013 - 82,414,244 (+)NCBI
Rnor_5.0982,180,271 - 82,183,510 (+)NCBI
RGSC_v3.4974,491,917 - 74,495,148 (+)RGD
Celera974,276,032 - 74,279,263 (+)RGD
Sequence:
RefSeq Acc Id: XM_006245160   ⟹   XP_006245222
RefSeq Status:
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2976,705,591 - 76,708,859 (+)NCBI
Rnor_6.0982,411,010 - 82,414,249 (+)NCBI
Rnor_5.0982,180,271 - 82,183,510 (+)NCBI
Sequence:
RefSeq Acc Id: XM_039083101   ⟹   XP_038939029
RefSeq Status:
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2976,705,591 - 76,708,252 (+)NCBI
RefSeq Acc Id: XM_039083102   ⟹   XP_038939030
RefSeq Status:
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2976,705,591 - 76,707,841 (+)NCBI
Reference Sequences
RefSeq Acc Id: NP_775165   ⟸   NM_173142
- UniProtKB: P57760 (UniProtKB/Swiss-Prot)
- Sequence:
RefSeq Acc Id: XP_006245222   ⟸   XM_006245160
- Peptide Label: isoform X1
- Sequence:
RefSeq Acc Id: ENSRNOP00000026194   ⟸   ENSRNOT00000026194
RefSeq Acc Id: XP_038939029   ⟸   XM_039083101
- Peptide Label: isoform X2
RefSeq Acc Id: XP_038939030   ⟸   XM_039083102
- Peptide Label: isoform X3
RefSeq Acc Id: ENSRNOP00000077839   ⟸   ENSRNOT00000111780
Protein Domains
Protein kinase

Protein Structures
Name Modeler Protein Id AA Range Protein Structure
AF-P57760-F1-model_v2 AlphaFold P57760 1-305 view protein structure

Transcriptome

eQTL   View at Phenogen
WGCNA   View at Phenogen
Tissue/Strain Expression   View at Phenogen

Promoters
RGD ID:13696774
Promoter ID:EPDNEW_R7298
Type:initiation region
Name:Stk16_1
Description:serine/threonine kinase 16
SO ACC ID:SO:0000170
Source:EPDNEW (Eukaryotic Promoter Database, http://epd.vital-it.ch/)
Experiment Methods:Single-end sequencing.
Position:
Rat AssemblyChrPosition (strand)Source
Rnor_6.0982,411,018 - 82,411,078EPDNEW

Additional Information

Database Acc Id Source(s)
AGR Gene RGD:629474 AgrOrtholog
BioCyc Gene G2FUF-27089 BioCyc
Ensembl Genes ENSRNOG00000019294 Ensembl, ENTREZGENE, UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
Ensembl Protein ENSRNOP00000026194 ENTREZGENE, UniProtKB/Swiss-Prot
  ENSRNOP00000077839.1 UniProtKB/TrEMBL
Ensembl Transcript ENSRNOT00000026194 ENTREZGENE, UniProtKB/Swiss-Prot
  ENSRNOT00000111780.1 UniProtKB/TrEMBL
InterPro Kinase-like_dom_sf UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  Prot_kinase_dom UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  Ser/Thr_kinase_AS UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
KEGG Report rno:286927 UniProtKB/Swiss-Prot
NCBI Gene 286927 ENTREZGENE
Pfam Pkinase UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
PhenoGen Stk16 PhenoGen
PROSITE PROTEIN_KINASE_DOM UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  PROTEIN_KINASE_ST UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
SMART S_TKc UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
Superfamily-SCOP SSF56112 UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
UniProt A0A8I5Y7Y0_RAT UniProtKB/TrEMBL
  P57760 ENTREZGENE, UniProtKB/Swiss-Prot


Nomenclature History
Date Current Symbol Current Name Previous Symbol Previous Name Description Reference Status
2004-09-10 Stk16  serine/threonine kinase 16      Symbol and Name status set to approved 1299863 APPROVED
2003-04-06 Stk16  serine/threonine kinase 16      Symbol and Name status set to provisional 70820 PROVISIONAL

RGD Curation Notes
Note Type Note Reference
gene_process activates the promoter for transforming growth factor-beta (Tgfb) 727297
gene_process may function in both transcriptional regulation and protein phosphorylation 727297