title: Toward understanding the functional role of Ss-RIOK-1, a RIO protein kinase-encoding gene of Strongyloides stercoralis.
abstract: BACKGROUND
  Some studies of Saccharomyces cerevisiae and mammals have shown that RIO protein kinases (RIOKs) are involved in ribosome biogenesis, cell cycle progression and development. However, there is a paucity of information on their functions in parasitic nematodes. We aimed to investigate the function of RIOK-1 encoding gene from Strongyloides stercoralis, a nematode parasitizing humans and dogs.
  
  
  RESULTS
  The RIOK-1 protein-encoding gene Ss-riok-1 was characterized from S. stercoralis. The full-length cDNA, gDNA and putative promoter region of Ss-riok-1 were isolated and sequenced. The cDNA comprises 1,828 bp, including a 377 bp 5'-UTR, a 17 bp 3'-UTR and a 1,434 bp ORF encoding a protein of 477 amino acids containing a RIOK-1 signature motif. The genomic sequence of the Ss-riok-1 coding region is 1,636 bp in length and has three exons and two introns. The putative promoter region comprises 4,280 bp and contains conserved promoter elements, including four CAAT boxes, 12 GATA boxes, eight E-boxes (CANNTG) and 38 TATA boxes. The Ss-riok-1 gene is transcribed throughout all developmental stages with the highest transcript abundance in the infective third-stage larva (iL3). Recombinant Ss-RIOK-1 is an active kinase, capable of both phosphorylation and auto-phosphorylation. Patterns of transcriptional reporter expression in transgenic S. stercoralis larvae indicated that Ss-RIOK-1 is expressed in neurons of the head, body and tail as well as in pharynx and hypodermis.
  
  
  CONCLUSIONS
  The characterization of the molecular and the temporal and spatial expression patterns of the encoding gene provide first clues as to functions of RIOKs in the biological processes of parasitic nematodes.
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Wang
authors.initials: W
authors.lastname: Yuan
authors.affiliation: Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America.
authors.forename: James B
authors.initials: JB
authors.lastname: Lok
authors.affiliation: Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America.
authors.forename: Jonathan D
authors.initials: JD
authors.lastname: Stoltzfus
authors.affiliation: Faculty of Veterinary Science, The University of Melbourne, Parkville, Victoria, Australia.
authors.forename: Robin B
authors.initials: RB
authors.lastname: Gasser
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Fang
authors.initials: F
authors.lastname: Fang
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Wei-Qiang
authors.initials: WQ
authors.lastname: Lei
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Rui
authors.initials: R
authors.lastname: Fang
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Yan-Qin
authors.initials: YQ
authors.lastname: Zhou
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Jun-Long
authors.initials: JL
authors.lastname: Zhao
authors.affiliation: State Key Laboratory of Agricultural Microbiology, Key Laboratory of Development of Veterinary Diagnostic Products, Ministry of Agriculture, College of Veterinary Medicine, Huazhong Agricultural University, Wuhan, China.
authors.forename: Min
authors.initials: M
authors.lastname: Hu
mesh_terms: D000818:Animals; D004285:Dogs; D006801:Humans; D011401:Promoter Regions, Genetic; D017346:Protein Serine-Threonine Kinases / Q000235:genetics* / Q000502:physiology; D017171:Strongyloides stercoralis / Q000201:enzymology / Q000235:genetics*; D014158:Transcription, Genetic
chemical_list: D017346:Protein Serine-Threonine Kinases
references.citation: Dev Biol. 1977 Mar;56(1):110-56
references.pmid: 838129
references.citation: Structure. 2004 Sep;12(9):1585-94
references.pmid: 15341724
references.citation: Cold Spring Harb Symp Quant Biol. 1983;48 Pt 2:633-40
references.pmid: 6586380
references.citation: Dev Biol. 1983 Nov;100(1):64-119
references.pmid: 6684600
references.citation: Mol Microbiol. 2002 Apr;44(2):309-24
references.pmid: 11972772
references.citation: J Biol Chem. 2000 Jun 23;275(25):18845-50
references.pmid: 10858445
references.citation: BMC Bioinformatics. 2013 Aug 08;14:241
references.pmid: 23924163
references.citation: Science. 2002 Dec 6;298(5600):1912-34
references.pmid: 12471243
references.citation: PLoS Pathog. 2009 Apr;5(4):e1000370
references.pmid: 19360119
references.citation: Dev Growth Differ. 2005 Dec;47(9):581-9
references.pmid: 16316403
references.citation: Gene. 2005 Aug 15;356:1-10
references.pmid: 16014321
references.citation: Bioinformatics. 2007 Nov 1;23(21):2947-8
references.pmid: 17846036
references.citation: Methods Cell Biol. 1995;48:451-82
references.pmid: 8531738
references.citation: Mech Dev. 2007 May;124(5):377-89
references.pmid: 17369030
references.citation: WormBook. 2007 May 23;:1-15
references.pmid: 18050491
references.citation: Nat Biotechnol. 2013 Jan;31(1):46-53
references.pmid: 23222703
references.citation: Mol Biochem Parasitol. 1999 Oct 15;103(2):171-81
references.pmid: 10551361
references.citation: Exp Biol Med (Maywood). 2003 Dec;228(11):1282-90
references.pmid: 14681544
references.citation: Parasit Vectors. 2008 Sep 22;1(1):34
references.pmid: 18808669
references.citation: Biotechnol Adv. 2011 May-Jun;29(3):338-50
references.pmid: 21262337
references.citation: Science. 1988 Jul 1;241(4861):42-52
references.pmid: 3291115
references.citation: Mol Biol Evol. 2013 Apr;30(4):772-80
references.pmid: 23329690
references.citation: Nature. 2000 Nov 16;408(6810):325-30
references.pmid: 11099033
references.citation: J Neurosci Res. 1985;13(1-2):75-88
references.pmid: 3973936
references.citation: Rev Infect Dis. 1981 May-Jun;3(3):397-407
references.pmid: 7025145
references.citation: FEBS J. 2007 Sep;274(17):4654-67
references.pmid: 17725716
references.citation: Int J Parasitol. 2003 Nov;33(13):1537-44
references.pmid: 14572516
references.citation: Cold Spring Harb Symp Quant Biol. 1983;48 Pt 2:745-59
references.pmid: 6586384
references.citation: PLoS Biol. 2003 Oct;1(1):E12
references.pmid: 14551910
references.citation: Nature. 2003 Jan 16;421(6920):268-72
references.pmid: 12529643
references.citation: Dev Cell. 2012 May 15;22(5):1101-8
references.pmid: 22560298
references.citation: Nat Protoc. 2012 Mar 01;7(3):562-78
references.pmid: 22383036
references.citation: Mol Cell Biol. 2003 Mar;23(6):2083-95
references.pmid: 12612080
references.citation: Exp Parasitol. 1984 Jun;57(3):287-96
references.pmid: 6723900
references.citation: J Biol Chem. 2011 Jan 21;286(3):1976-86
references.pmid: 21081503
references.citation: Int J Parasitol. 2013 Jun;43(7):515-20
references.pmid: 23500073
references.citation: Mol Biochem Parasitol. 2002 Feb;119(2):279-84
references.pmid: 11814580
references.citation: Mol Cell Biol. 2000 Jan;20(2):429-40
references.pmid: 10611221
references.citation: Nature. 2005 Mar 24;434(7032):462-9
references.pmid: 15791247
references.citation: Nucleic Acids Res. 2001 Aug 15;29(16):3377-84
references.pmid: 11504875
references.citation: Int J Parasitol. 2006 May 31;36(6):671-9
references.pmid: 16500658
references.citation: J Comp Neurol. 1997 Dec 15;389(2):212-23
references.pmid: 9416917
references.citation: WormBook. 2005 Dec 13;:1-19
references.pmid: 18050405
references.citation: WormBook. 2007 Feb 17;:1-18
references.pmid: 18050500
references.citation: Mol Biol Evol. 2011 Oct;28(10):2731-9
references.pmid: 21546353
references.citation: Proc Natl Acad Sci U S A. 1997 May 13;94(10):5172-6
references.pmid: 9144210
references.citation: Biochim Biophys Acta. 2005 Dec 30;1754(1-2):14-24
references.pmid: 16182620
references.citation: J Neurosci. 1988 Jan;8(1):176-84
references.pmid: 3339407
references.citation: J Cell Physiol. 2005 Oct;205(1):1-9
references.pmid: 15887235
references.citation: J Comp Neurol. 1975 Apr 1;160(3):313-37
references.pmid: 1112927
references.citation: EMBO J. 1998 Mar 2;17(5):1476-86
references.pmid: 9482744
references.citation: Philos Trans R Soc Lond B Biol Sci. 1986 Nov 12;314(1165):1-340
references.pmid: 22462104
references.citation: Nat Methods. 2012 Mar 04;9(4):357-9
references.pmid: 22388286
references.citation: Mol Biol Cell. 2012 Jan;23(1):22-35
references.pmid: 22072790
references.citation: Nucleic Acids Res. 2000 Jan 1;28(1):263-6
references.pmid: 10592242
references.citation: Exp Cell Res. 2005 Jun 10;306(2):343-8
references.pmid: 15925590
references.citation: J Neurosci. 1986 Feb;6(2):412-23
references.pmid: 3950704
references.citation: Nucleic Acids Res. 1993 May 25;21(10):2525-6
references.pmid: 8506152
references.citation: J Mol Biol. 1990 Apr 20;212(4):563-78
references.pmid: 2329577
references.citation: Nat Methods. 2008 Jul;5(7):621-8
references.pmid: 18516045
references.citation: FEBS Lett. 2002 Jul 31;524(1-3):31-6
references.pmid: 12135737
references.citation: Development. 1993 Mar;117(3):1071-87
references.pmid: 8325236
references.citation: Mol Neurobiol. 1988 Winter;2(4):227-61
references.pmid: 3078112
references.citation: J Mol Evol. 2000 Feb;50(2):103-15
references.pmid: 10684344
references.citation: PLoS Biol. 2013 Oct;11(10):e1001680
references.pmid: 24143133
references.citation: Nucleic Acids Res. 1993 Jul 1;21(13):3097-103
references.pmid: 8332530
references.citation: PLoS Negl Trop Dis. 2012;6(10):e1854
references.pmid: 23145190
references.citation: PLoS One. 2012;7(6):e38587
references.pmid: 22701676
references.citation: Nucleic Acids Res. 1997 Sep 1;25(17):3389-402
references.pmid: 9254694
references.citation: Genome Res. 2004 Oct;14(10B):2162-8
references.pmid: 15489339
references.citation: EMBO J. 2001 Aug 1;20(15):4204-13
references.pmid: 11483523
references.citation: FEBS J. 2005 Jul;272(14):3698-713
references.pmid: 16008568
references.citation: Genome Biol. 2013 Apr 25;14(4):R36
references.pmid: 23618408
references.citation: J Biol Chem. 2011 Oct 21;286(42):36352-60
references.pmid: 21880710
references.citation: EMBO J. 2000 Dec 15;19(24):6870-81
references.pmid: 11118222
references.citation: Nature. 1998 Mar 5;392(6671):71-5
references.pmid: 9510248
references.citation: J Infect Dis. 1993 Dec;168(6):1479-84
references.pmid: 8245532
references.citation: Exp Parasitol. 2008 Feb;118(2):253-65
references.pmid: 17945217
references.citation: Philos Trans R Soc Lond B Biol Sci. 1976 Aug 10;275(938):327-48
references.pmid: 8806
references.citation: J Biol Chem. 2005 Nov 11;280(45):37297-300
references.pmid: 16183636
references.citation: J Comp Neurol. 1995 Jun 26;357(2):281-95
references.pmid: 7665730
references.citation: Cardiovasc Res. 2004 Aug 1;63(2):196-207
references.pmid: 15249177