人类遗传性多囊肾相关致病基因功能的研究
详细信息    本馆镜像全文|  推荐本文 |  |   获取CNKI官网全文
摘要
多囊肾疾病(polycystic kidney disease, PKD)是一组常见的以双肾形成多个进行性增大囊肿为主要特征的单基因遗传病。根据遗传特征不同,可以分为常染色体显性多囊肾(Autosomal dominant polycystic kidney disease, ADPKD)和常染色体隐性多囊肾(Autosomal recessive polycystic kidney disease, ARPKD)。ADPKD主要由PKD1和PKD2两个基因突变引起,基因产物分别为Polycystin-1 (PC1)和Polycystin-2 (PC2); ARPKD则由PKHD1基因突变引起,基因产物为Fibrocystin/Polyductin (FPO)。PKD1、PKD2和PKHD1的小鼠同源基因PMd1、Pkd2和Pkhd1都已被克隆,两者之间具有高度的保守性,这些基因的突变也能导致小鼠多囊肾。
     本研究首先建立了一个新型的Pkhd1基因敲除小鼠模型,该模型能很好的模拟人类ARPKD。结合实验室前期已经建立的Pkd2基因敲除小鼠,发现Pkhd1和Pkd2的双突变小鼠比单个基因突变表现出更为严重的症状,提示ARPKD和ADPKD这两类具有不同遗传特征的疾病可能具有分子水平的相互关系。进一步研究发现,FPC和PC2在体内能形成一个分子复合体,这一复合体的形成是通过FPC的胞内羧基末端和PC2胞内氨基末端直接相互作用实现的。FPC的缺失能引起PC2蛋白表达的下降,并能显著降低PC2的离子通道活性,然而PC2的缺失并不影响FPC的表达。这一研究表明,FPC和PC2在体内形成一个分子复合物,并于同一分子通路参与肾内管道结构的形成。
     除了PKD1, PKD2和PKHD1,目前已在人和其他动物体内克隆了至少20个多囊肾相关基因。Bicc1,果蝇双尾C (Bic-C)基因的同源基因,是一个小鼠多囊肾致病基因。Bpk (BALB/c polycystic kidneys)和Jcpk (Juvenile congenital polycystic kidney),这两个分别通过自发突变和人工诱变产生的多囊肾动物模型,都由该基因突变引起。Biccl普遍存在于从线虫到人类的各个物种,并且其序列高度保守。在果蝇等低等动物的研究发现,Bic-C是一个具有RNA结合活性的转录后基因表达调控因子。尽管Biccl的多囊肾动物模型为其体内功能研究提供了一些线索,然而Biccl的细胞生物学功能还完全未知。本研究通过建立IMCD细胞的稳定干涉细胞株,发现Biccl为IMCD细胞在三维培养条件下形成管状分支结构所必须,抑制Biccl的表达导致以E-cadherin为基础的细胞连接异常以及纤毛发生障碍,同时细胞的增殖、凋亡和细胞骨架也出现异常。这一研究初步揭示了Biccl的细胞生物学功能,为进一步探寻该基因在体内的功能及多囊肾的发病机制奠定了基础。
Polycystic kidney disease (PKD) is a group of monogenic disorders that result in renal cyst development. Based on different patterns of inheritance, there are two kinds of PKD, Autosomal dominant PKD(ADPKD) and autosomal recessive PKD(ARPKD). Approximately 85% of ADPKD patients have mutations in the PKD1 gene, which encodes polycystin-1 (PC-1). About 15% of ADPKD cases have mutations in the PKD2 gene, which encodes polycystin-2 (PC-2). Fibrocystin or polyductin(FPC), the poduct of PKHD1 gene, is the only identified ARPKD disease gene. Pkdl, Pkd2 and Pkhdl is the mouse homolog of PKD1, PKD2 and PKHD1 respectively, whose mutation also cause mouse polycystic kidney disease.
     In the current study, we generate a new mouse model by disrupting Pkhdl, which perfectly recapitulates the human ARPKD phenotype. Transmutant mice for Pkhd1 and Pkd2 displayed a significantly more severe renal cystic phenotype than single mutant mice, indicating a molecular relationship between ADPKD and ARPKD. More detailed research found that FPC physically interacts with PC2 and that lack of FPC destabilizes PC2 expression in vivo but not vice versa, suggesting that PC2 functions immediately downstream of FPC. In addition, inhibition of FPC expression reduces Pkd2-channel activity, we conclude that a functional and molecular interaction exists between FPC and PC2 in vivo.
     Except PKD1/2 and PKHD1, more than 20 genes have currently been identified to be able to induce PKD phenotypes in human or various animal mutant models. Biccl is a mouse homologue of Drosophila Bicaudctl-C (dBic-C) and the ortholog of dBic-C in many other species (from C.elegans to humans) can be found to be very conserved. In Drosophila, the gene product of dBic-C is currently considered as an RNA-binding molecule and believed to function in regualtion of mRNA post-transcription. Although the mutant mouse models provide some knowledge on functional roles of Biccl in mammalian development, its cellular functions still remain mostly unknown. To this end, we established stable Biccl-silenced MCD cell lines and characterized its functions in cell biology. Our results indicate that the expression of normal Biccl is required to sustain renal tubulogenesis in 3-D cultures of IMCD cells. Lack of this protein resulted in obvious abnormalities in E-cadherin-based cell-cell contact, which is essential for epithelial polarization and regulation of tubulomorphagenesis. These in vitro findings indicate that Biccl functional roles in renal epithelial differentiation and renal tubule development. The observations made in vitro may illuminate Biccl cellular functions in vivo.
引文
Bacallao RL, McNeill H. Cystic kidney diseases and planar cell polarity signaling. Clin Genet.2009 Feb;75(2):107-17
    Barr MM, Sternberg PW. A polycystic kidney-disease gene homologue required for male mating behaviour in C. elegans. Nature.1999 Sep 23;401(6751):386-9.
    Benzing T, Simons M, Walz G. Wnt signaling in polycystic kidney disease. J Am Soc Nephrol.2007 May; 18(5):1389-98.
    Bleyer AJ, Hart TC. Medullary cystic kidney disease type 2. Am J Kidney Dis.2004 Jun;43(6).1142; author reply 1142-3.
    Bukanov NO, Smith LA, Klinger KW, et al. Long-lasting arrest of murine polycystic kidney disease with CDK inhibitor roscovitine. Nature 2006.444:949-52
    Fick GM, Gabow PA. Hereditary and acquired cystic disease of the kidney. Kidney Int. 1994 Oct;46(4):951-64.
    Fischer E, Legue E, Doyen A, et al. Defective planar cell polarity in polycystic kidney disease. Nat Genet.2006 Jan;38(1):21-3.
    Gattone VH 2nd, Wang X, Harris PC, et al.2003. Inhibition of renal cystic disease development and progression by a vasopressin V2 receptor antagonist. Nat. Med. 9:1323-26
    Harris PC, Torres VE. Polycystic Kidney Disease. Annu Rev Med.2008 Oct 23.
    Hildebrandt F, Attanasio M, Otto E. Nephronophthisis:disease mechanisms of a ciliopathy. J Am Soc Nephrol.2009 Jan;20(1):23-35.
    Hildebrandt F, Otto E. Molecular genetics of nephronophthisis and medullary cystic kidney J Am Soc Nephrol.2000 Sep; 11(9):1753-61.
    Ibraghimov-Beskrovnaya O, Bukanov N. Polycystic kidney diseases:from molecular discoveries to targeted therapeutic strategies. Cell Mol Life Sci.2008 Feb;65(4):605-19.
    Leuenroth SJ, Okuhara D, Shotwell JD, et al. Triptolide is a traditional Chinese medicine-derived inhibitor of polycystic kidney disease. Proc. Natl. Acad. Sci. USA 2007.104:4389-94。
    Masyuk TV, Masyuk Al, Torres VE, et al. Octreotide inhibits hepatic cystogenesis in a rodent model of polycystic liver disease by reducing cholangiocyte adenosine 3',5'-cyclic monophosphate. Gastroenterology 2007.132:1104-16
    Park TJ, Haigo SL, Wallingford JB. Ciliogenesis defects in embryos lacking inturned or fuzzy function are associated with failure of planar cell polarity and Hedgehog signaling. Nat Genet.2006 Mar;38(3):303-11.
    Polgar K, Burrow CR, Hyink DP, et al. Disruption of polycystin-1 function interferes with branching morphogenesis of the ureteric bud in developing mouse kidneys. Dev Biol.2005 Oct 1;286(1):16-30.
    Pretorius DH, Reznik V. Senior-Loken syndrome. J Ultrasound Med.2007 Mar;26(3):418.
    Robert A, Margall-Ducos G, Guidotti JE, et al. The intraflagellar transport component IFT88/polaris is a centrosomal protein regulating G1-S transition in non-ciliated cells. J Cell Sci.2007 Feb 15;120(Pt 4):628-37.
    Saburi S, Hester I, Fischer E, et al. Loss of Fat4 disrupts PCP signaling and oriented cell division and leads to cystic kidney disease. Nat Genet.2008 Aug;40(8):1010-5.
    Sharp AM, Messiaen LM, Page G, et al. Comprehensive genomic analysis of PKHD1 mutations in ARPKD cohorts. J Med Genet.2005 Apr;42(4):336-49
    Shillingford JM, Murcia NS, Larson CH, et al. The mTOR pathway is regulated by polycystin-1, and its inhibition reverses renal cystogenesis in polycystic kidney disease. Proc. Natl. Acad. Sci. USA 2006.103:5466-71
    Simons M, Gloy J, Ganner A, et al. Inversin, the gene product mutated in nephronophthisis type Ⅱ, functions as a molecular switch between Wnt signaling pathways. Nat Genet.2005 May;37(5):537-43.
    SweeneyWEJr, von Vigier RO, Frost P, et al. Src inhibition ameliorates polycystic kidney disease. J. Am. Soc. Nephrol.2008.19:1331-41
    Torres VE, Harris PC, Pirson Y. Autosomal dominant polycystic kidney disease. Lancet. 2007 Apr 14;369(9569):1287-301
    Torres VE, Harris PC. Mechanisms of Disease:autosomal dominant and recessive polycystic kidney diseases. Nat Clin Pract Nephrol.2006 Jan;2(1):40-55
    Wilson PD, Goilav B. Cystic disease of the kidney. Annu Rev Pathol.2007;2:341-68.
    Yang B, Sonawane ND, Zhao D, et al. Small-molecule CFTR inhibitors slow cyst growth in polycystic kidney disease. J. Am. Soc. Nephrol.2008.19:1300-10
    Yoder BK. Role of primary cilia in the pathogenesis of polycystic kidney disease. J Am Soc Nephrol.2007 May; 18(5):1381-8.
    Bergmann C, Senderek J, Schneider F, et al., PKHD1 mutations in families requesting prenatal diagnosis for autosomal recessive polycystic kidney disease (ARPKD). Hum Mutat. 200423,487-495.
    Calvet, J. P., and Grantham, J. J. The genetics and physiology of polycystic kidney disease. Semin Nephrol.2001 21,107-123.
    Capisonda, R., Phan, V, Traubuci, J., et al., Autosomal recessive polycystic kidney disease: outcomes from a single-center experience. Pediatr Nephrol.2003 18,119-126.
    Chauvet V, Qian F, Boute N, et al. Expression of PKD1 and PKD2 transcripts and proteins in human embryo and during normal kidney development. Am J Pathol.2002 Mar;160(3):973-83.
    Chevalier RL, Garland TA, Buschi AJ:The neonate with adult-type autosomal dominant polycystic kidney disease. Int J Pediatr Nephrol.1981 2:73-77,
    European PKD1, Consortium. The polycystic kidney disease 1 gene encodes a 14 kb transcript and lies within a duplicated region on chromosme 16. Cell.1994 77,881-894.
    Fischer E, Legue E, Doyen A, et al. Defective planar cell polarity in polycystic kidney disease. Nat Genet.2006 Jan;38(1):21-3.
    Gallagher AR, Esquivel EL, Briere TS, et al. Biliary and pancreatic dysgenesis in mice harboring a mutation in Pkhdl. Am J Pathol.2008 Feb;172(2):417-29.
    Garcia-Gonzalez MA, Menezes LF, Piontek KB, et al. Genetic interaction studies link autosomal dominant and recessive polycystic kidney disease in a common pathway. Hum Mol Genet.2007 16(16):1940-50.
    Gattone VH 2nd, Wang X, Harris PC, Torres VE:Inhibition of renal cystic disease development and progression by a vasopressin V2 receptor antagonist. Nat Med.2003 9: 1323-1326,
    Harris PC, Torres VE. Polycystic Kidney Disease. Annu Rev Med.2008 Oct 23.
    Harris, P. C., Bae, K. T., Rossetti, S., et al. Cyst number but not the rate of cystic growth is associated with the mutated gene in autosomal dominant polycystic kidney disease. J Am Soc Nephrol.2006 17,3013-3019.
    Igarashi P, Somlo S. Genetics and pathogenesis of polycystic kidney disease. J Am Soc Nephrol.2002 13,2384-2398.
    Ibraghimov-Beskrovnaya O, Bukanov N. Polycystic kidney diseases:from molecular discoveries to targeted therapeutic strategies. Cell Mol Life Sci.2008 Feb;65(4):605-19.
    Koulen P, Cai Y, Geng L, et al. Polycystin-2 is an intracellular calcium release channel. Nat Cell Biol.20024:191-197.
    Kucherlapati R, Edelmann W, Somlo S. Cardiac defects and renal failure in mice with targeted mutations in Pkd2. Nat Genet.200024:75-78.
    Lal M, Song X, Pluznick JL, et al. Polycystin-1 C-terminal tail associates with beta-catenin and inhibits canonical Wnt signaling. Hum Mol Genet.2008 Oct 15,17(20):3105-17.
    Lilova MI, Petkov DL. Intracranial aneurysms in a child with autosomal recessive polycystic kidney disease. Pediatr Nephrol.2001 16,1030-1032.
    Lin F, Hiesberger T, Cordes K, et al. Kidney-specific inactivation of the KIF3A subunit of kinesin-Ⅱ inhibits renal ciliogenesis and produces polycystic kidney disease.Proc Natl Acad Sci U S A.2003 100:5286-5291.
    Lonergan, G. J., Rice, R. R., and Suarez, E. S. Autosomal recessive polycystic kidney disease: radiologic-pathologic correlation. Radiographics.2000 20,837-855.
    Lucon, M, Ianhez, L. E., Lucon, A. M, et al. Bilateral nephrectomy of huge polycystic kidneys associated with a rectus abdominis diastasis and umbilical hernia. Clinics.2006 61, 529-534.
    Lumiaho, A., Ikaheimo, R., Miettinen, R., et al. Mitral valve prolapse and mitral regurgitation are common in patients with polycystic kidney disease type 1. Am J Kidney Dis.2001 38, 1208-1216.
    Mai W, Chen D, Ding T, Kim I, Park S, Cho SY, Chu JS, Liang D, Wang N, Wu D, Li S, Zhao P, Zent R, Wu G, et al. Inhibition of Pkhdl Impairs Tubulomorphogenesis of Cultured IMCD Cells Molecular Biology of the Cell.2005 16,4398-4409.
    Masyuk TV, Huang BQ, Ward CJ, et al. Defects in cholangiocyte fibrocystin expression and ciliary structure in the PCK rat. Gastroenterology.2003 125:1303-1310.
    McGrath J, Somlo S, Makova S, Tian X, et al. Two populations of node monocilia initiate left-right asymmetry in the mouse. Cell.2003 114:61-73.
    Menezes LF, Cai Y, Nagasawa Y et al. Polyductin, the PKHD1 gene product, comprises isoforms expressed in plasma membrane, primary cilium, and cytoplasm. Kidney Int. 200466:1345-1355.
    Mochizuki, T, Wu, G, Hayashi, T., et al. PKD2, a gene for polycystic kidney disease that encodes an integral membrane protein. Science.1996 272,1339-1342.
    Montell, C., Birnbaumer, L., and Flockerzi, V. The TRP channels, a remarkably functional family. Cell.2002 108,595-598.
    Morris-Stiff G, Coles G, Moore R et al. Abdominal wall hernia in autosomal dominant polycystic kidney disease. Br J Surg.1997 84,615-617.
    Moser M, Matthiesen S, Kirfel J, et al. A mouse model for cystic biliary dysgenesis in autosomal recessive polycystic kidney disease (ARPKD). Hepatology.2005 May;41(5):1113-21.
    Nauli SM, Alenghat FJ, Luo Y, et al. Polycystins 1 and 2 mediate mechanosensation in the primary cilium of kidney cells. Nat Genet.2003 33:129-137,
    Neumann HP, Krumme B, van Velthoven V, et al. Multiple intracranial aneurysms in a patient with autosomal recessive polycystic kidney disease. Nephrol Dial Transplant.1999 Apr;14(4):936-9.
    Newby LJ, Streets AJ, Zhao Y, et al. Identification, characterization, and localization of a novel kidney polycystin-l-polycystin-2 complex. J Biol Chem.2002 Jun 7;277(23):20763-73.
    Oflaz H, Alisir S, Buyukaydin B, et al. Biventricular diastolic dysfunction in patients with autosomal-dominant polycystic kidney disease. Kidney Int.2005 68,2244-2249.
    Onuchic LF, Furu L, Nagasawa Y, et al. PKHD1, the polycystic kidney and hepatic disease 1 gene, encodes a novel large protein containing multiple immunoglobulin-like plexin-transcription-factor domains and parallel beta-helix 1 repeats. Am J Hum Genet.2002 70, 1305-1317.
    Pennekamp P, Karcher C, Fischer A, et al. The ion channel polycystin-2 is required for left-right axis determination in mice. Curr Biol.2002 12:938-943,
    Qian CN, Knol J, Igarashi P, et al. Cystic renal neoplasia following conditional inactivation of apc in mouse renal tubular epithelium. J Biol Chem.2005 Feb 4;280(5):3938-45.
    Qian F, Germino FJ, Cai Y, et al. PKD1 interacts with PKD2 through a probable coiled-coil domain. Nat Genet.1997 16:179-183.
    Qian F, Watnick TJ, Onuchic LF, et al. The molecular basis of focal cyst formation in human autosomal dominant polycystic kidney disease type Ⅰ. Cell.199687:979-987,
    Rodova M, Islam MR, Maser RL, Calvet JP. et al. The polycystic kidney disease-1 promoter is a target of the beta-catenin/T-cell factor pathway. J Biol Chem.2002 Aug 16;277(33):29577-83.
    Saadi-Kheddouci S, Berrebi D, Romagnolo B, et al. Early development of polycystic kidney disease in transgenic mice expressing an activated mutant of the beta-catenin gene. Oncogene.2001 Sep 20;20(42):5972-81.
    Shannon MB, Patton BL, Harvey SJ, et al. A hypomorphic mutation in the mouse laminin alpha5 gene causes polycystic kidney disease. J Am Soc Nephrol.200617: 1913-1922.
    Sharp AM, Messiaen LM, Page G, et al. Comprehensive genomic analysis of PKHD1 mutations in ARPKD cohorts. J Med Genet.200542,336-349.
    Sutters M, Germino GG. Autosomal dominant polycystic kidney disease:molecular genetics and pathophysiology. J Lab Clin Med.2003 Feb;141(2):91-101.
    Torres VE, Wang X, Qian Q, et al. Effective treatment of an orthologous model of autosomal dominant polycystic kidney disease. Nat Med.2004 10:363-364.
    Trudel M, D'Agati V, Costantini F. C-myc as an inducer of polycystic kidney disease in transgenic mice. Kidney Int.1991 Apr;39(4):665-71.
    Wang S, Luo Y, Wilson PD, et al. The autosomal recessive polycystic kidney disease protein is localized to primary cilia, with concentration in the basal body area. J Am Soc Nephrol.2004 15:592-602.
    Wang S, Zhang J, Nauli SM, et al. Fibrocystin/polyductin, found in the same protein complex with polycystin-2, regulates calcium responses in kidney epithelia. Mol Cell Biol.200727:3241-3252,.
    Ward CJ, Yuan D, Masyuk TV, et al. Cellular and subcellular localization of the ARPKD protein; fibrocystin is expressed on primary cilia. Hum Mol Genet 12:2703-2710,2003 Watnick T, Germino G:From cilia to cyst. Nat Genet.200334:355-356.
    Watnick T, Germino GG. Molecular basis of autosomal dominant polycystic kidney disease. Semin Nephrol.1999 Jul;19(4):327-43.
    Williams SS, Cobo-Stark P, James LR, et al. Kidney cysts, pancreatic cysts, and biliary disease in a mouse model of autosomal recessive polycystic kidney disease.Pediatr Nephrol.2008 May;23(5):733-41.
    Wilson P. Polycystic kidney disease. N Engl J Med.2004 350,151-164.
    Woollard JR, Punyashtiti R, Richardson S, et al. A mouse model of autosomal recessive polycystic kidney disease with biliary duct and proximal tubule dilatation. Kidney Int. 2007 Aug;72(3):328-36.
    Wu G, Tian X, Nishimura S, et al. Trans-heterozygous Pkdl and Pkd2 mutations modify expression of polycystic kidney disease. Human Molecular Genetics.2002 16,1845-1854.
    Wu Y, Dai XQ, Li Q, et al. Kinesin-2 mediates physical and functional interactions between polycystin-2 and fibrocystin. Hum Mol Genet.2006 15:3280-3292,
    Xiong H, Chen Y, Wu G. et al. A Novel Gene Encoding a TIG Multiple Domain Protein Is a Positional Candidate for Autosomal Recessive Polycystic Kidney Disease. Genomics.200280, 96-104.
    Xu C, Rossetti S, Jiang L, et al. Human ADPKD primary cyst epithelial cells with a novel, single codon deletion in the PKDl gene exhibit defective ciliary polycystin localization and loss of flow-induced Ca2+ signaling. Am J Physiol Renal Physiol.2007 Mar;292(3):F930-45.
    Xu GM, Gonzalez-Perrett S, Essafi M, et al. Polycystin-1 activates and stabilizes the polycystin-2 channel. J Biol Chem.2003 Jan 17;278(3):1457-62.
    Yoder BK, Hou X, Guay-Woodford LM. The polycystic kidney disease proteins, polycystin-1, polycystin-2, polaris, and cystin, are co-localized in renal cilia. J Am Soc Nephrol.2002 13(10):2508-16.
    Yonemura K, Yasuda H, Fujigaki Y Adrenal insufficiency due to isolated adrenocorticotropin deficiency complicated by autosomal recessive polycystic kidney disease. Ren Fail.2003 25,485-492.
    Zerres K, Rudnik-Schoneborn S, Mucher G. Autosomal recessive polycystic kidney disease. clinical features and genetics. Adv Nephrol Necker Hosp.199625,147-157.
    Zerres K, Rudnik-Schoneborn S, Senderek J, et al. Autosomal recessive polycystic kidney disease (ARPKD). J Nephrol.2003 16,453-458.
    Zerres K, Rudnik-Schoneborn S, Steinkamm C, et al. Autosomal recessive polycystic kidney disease. J Mol Med.199876,303-309.
    Zhang MZ, Mai W, Li C, et al. PKHD1 protein encoded by the gene for autosomal recessive polycystic kidney disease associates with basal bodies and primary cilia in renal epithelial cells. Proc Natl Acad Sci U S A.2004 Feb;101(8):2311-6.
    Balda MS, Matter K. Epithelial cell adhesion and the regulation of gene expression. Trends Cell Biol 2003; 13:310-318.
    Behm-Ansmant I, Rehwinkel J, Doerks T, et al. mRNA degradation by miRNAs and GW182 requires both CCR4:NOT deadenylase and DCP1:DCP2 decapping complexes. Genes Dev.2006 Jul 15;20(14):1885-98.
    Berridge MV, Tan AS, McCoy KD, et al. THe biochemical and cellular basis of cell proliferation assays that use tetrazolium salts. Biochemica 1996; 4:12-20.
    Boletta A, Qian F, Onuchic LF et al. Polycystin-1, the gene product of PKD1, induces resistance to apoptosis and spontaneous tubulogenesis in MDCK cells. Mol Cell 2000; 6 (5):1267-1273.
    Bouvrette DJ, Price SJ, Bryda EC. K homology domains of the mouse polycystic kidney disease-related protein, Bicaudal-C (Biccl), mediate RNA binding in vitro. Nephron Exp Nephrol 2008; 108 (1):e27-34.
    Chen D, Roberts R, Pohl M, et al. Differential expression of collagen and laminin binding integrins mediate ureteric bud and inner medullary collecting duct cell tubulogenesis. Am J Physiol Renal Physiol 2004; 287.F602-611.
    Chen T, Damaj BB, Herrera C, et al. Self-association of the single-KH-domain family members Sam68, GRP33, GLD-1, and Qkl:role of the KH domain.Mol Cell Biol 1997; 17 (10):5707-5718.
    Cogswell C, Price SJ, Hou X et al. Positional cloning of jcpk/bpk locus of the mouse. Mamm Genome 2003; 14 (4):242-249.
    Dai B, Liang Z, Fu Y, et al. Generatoin of an anti-Biccl antibody and characterization of its subcellular localization. Chin J Cell Mol Immunol 2008; 24:1106-1109.
    Eckmann CR, Crittenden SL, Suh N, et al. GLD-3 and control of the mitosis/meiosis decision in the germline of Caenorhabditis elegans. Genetics.2004 Sep; 168(1).147-60.
    Eulalio A, Behm-Ansmant I, Izaurralde E. P bodies:at the crossroads of post-transcriptional pathways. Nat Rev Mol Cell Biol.2007 Jan;8(1):9-22.
    Flaherty L, Bryda EC, Collins D, Rudofsky U, Montogomery JC. New mouse model for polycystic kidney disease with both recessive and dominant gene effects. Kidney Int 1995;47(2):552-558.
    Gibson TJ, Thompson JD, Heringa J. The KH domain occurs in a diverse set of RNA-binding proteins that include the antiterminator NusA and is probably involved in binding to nucleic acid. FEBS Lett.1993 Jun 21;324(3):361-6.
    Grimm DH, Karihaloo A, Cai Y et al. Polycystin-2 regulates proliferation and branching morphogenesis in kidney epithelial cells. J Biol Chem 2006; 281 (1):137-144.
    Higashiyama S, Iwamoto R, Goishi K, et al. The membrane protein CD9/DRAP 27 potentiates the juxtacrine growth factoractivity of the membrane-anchored heparin-binding EGF-like growth factor. J Cell Biol 1995; 128:929-938.
    Koch CA, Anderson D, Moran MF, et al. SH2 and SH3 domains:elements that control interactions of cytoplasmic signaling proteins. Science.1991 May 3;252(5006):668-74.
    Lin F, Hiesberger T, Cordes K et al. Kidney-specific inactivation of the KIF3A Mahone M, Saffman EE, Lasko PF. Localized Bicaudal-C RNA encodes a protein containing a KH domain, the RNA binding motif of FMR1. EMBO J 1995; 14 (9):2043-2055.
    Mai W, Chen D, Ding T, et al. Inhibition of Pkhdl Impairs Tubulomorphogenesis of Cultured IMCD Cells Mol Biol Cell 2005; 16:4398-4409.
    Matter K, Balda MS. Signalling to and from tight junctions. Nat Rev Mol Cell Biol 2003; 4:225-236.
    Nauta J, Sweeney WE, Rutledge JC, et al. Biliary epithelial cells from mice with congenital polycystic kidney disease are hyperresponsive to epidermal growth factor. Pediatr Res 1995; 37 (6):755-763.
    Nishio S, Hatano M, Nagata M, et al. Pkdl regulates immortalized proliferation of renal tubular epithelial cells through p53 induction and JNK activation. J Clin Invest.2005 Apr;115(4):910-8.
    Schultz J, Ponting CP, Hofmann K, et al. SAM as a protein interaction domain involved in developmental regulation. Protein Sci 1997; 6(1):249-253.
    Stagner EE, Bouvrette DJ, Cheng J, et al. The polycystic kidney disease-related proteins Biccl and SamCystin interact. Biochem Biophys Res Commun.2009 May 22;383(1):16-21.
    Torres VE, Harris PC. Polycystic kidney disease:genes, proteins, animal models, disease mechanisms and therapeutic opportunities. J Intern Med.2007 Jan;261(1):17-31.
    Tran U, Pickney LM, Ozpolat BD, et al. Xenopus Bicaudal-C is required for the differentiation of the amphibian pronephros. Dev Biol 2007; 307 (1):152-164.
    Wegierski T, Steffl D, Kopp C, et al. TRPP2 channels regulate apoptosis through the Ca(2+) concentration in the endoplasmic reticulum. EMBO J.2009 Mar 4;28(5):490-9
    Wessely O, De Robertis EM. The Xenopus homologue of Bicaudal-C is a localized maternal mRNA that can induce endoderm formation. Development 2000; 127(10):2053-2062.
    Wessely O, Tran U, Zakin L, et al. Identification and expression of the mammalian homologue of Bicaudal-C. Mech Dev 2001; 101 (1-2):267-270.
    Wilson PD, Goilav B. Cystic disease of the kidney. Annu Rev Pathol 2007; 2:341.
    Woo D. Apoptosis and loss of renal tissue in polycystic kidney diseases. N Engl J Med 1995; 333(1):18-25.
    Yoder BK, Tousson A, Millican L, et al.Polaris, a protein disrupted in orpk mutant mice, is required for assembly of renal cilium. Am J Physiol Renal Physiol 2002; 282:F541-552.
    Zegers MM, O'Brien LE, Yu W, Datta A, Mostov KE. Epithelial polarity and tubulogenesis in vitro. Trends Cell Biol 2003; 13:169-176.
    Zhou L, Dai B, Wu G. Molecular Characteristics and Functions of Bicaudal-C. Int J Genet 2008 (31):432.
    Bergmann C, Senderek J, Windelen E, et al. Clinical consequences of PKHD1 mutations in 164 patients with autosomal-recessive polycystic kidney disease (ARPKD). Kidney Int. 2005 Mar;67(3):829-48.
    Christopher J. Ward, Marie C. Hogan, Sandro Rossettil, et al. The gene mutated in autosomal recessive polycystic kidney disease encodes a large, receptor-like protein. Nat Genet.2002 Mar;30(3):259-69.
    Fischer E, Legue E, Doyen A, et al. Defective planar cell polarity in polycystic kidney disease. Nat Genet.2006 Jan;38(1):21-3.
    Gallagher AR, Esquivel EL, Briere TS, et al. Biliary and pancreatic dysgenesis in mice harboring a mutation in Pkhdl Am J Pathol.2008 Feb;172(2):417-29.
    Garcia-Gonzalez MA, Menezes LF, Piontek KB, et al. Genetic interaction studies link autosomal dominant and recessive polycystic kidney disease in a common pathway Hum Mol Genet.2007 Aug 15; 16(16):1940-50.
    Germino GG. Linking cilia to Wnts. Nat Genet.2005 May;37(5):537-43.
    Goilav, B., Norton, K. I., Satlin, L. M., et al. Predominant extrahepatic biliary disease in autosomal recessive polycystic kidney disease:a new association. Pediatr Transplant 2006,10,294-298.
    Guay-Woodford LM, Muecher G, Hopkins SD, et al. The severe perinatal form of autosomal recessive polycystic kidney disease maps to chromosome 6p21.1-p12: implications for genetic counseling. Am. J. Hum. Genet.56,1101-1107 (1995).
    Guay-Woodford, L. M., Desmond, R. A. Autosomal recessive polycystic kidney disease: the clinical experience in North America. Pediatrics 2003,111,1072-1080.
    Harris PC, Torres VE. Polycystic Kidney Disease. Annu Rev Med.2008 Oct 23.
    Hogan MC, Griffin MD, Rossetti S, et al. PKHDL1, a homolog of the autosomal recessive polycystic kidney disease gene, encodes a receptor with inducible T lymphocyte expression. Hum Mol Genet.2003 Mar 15;12(6):685-98.
    Kim Ia, Fu Y, Hui K, et al. Fibrocystin/polyductin modulates renal tubular formation by regulating polycystin-2 expression and function.J Am Soc Nephrol.2008 Mar;19(3):455-68.
    Kim Ib, Li C, Liang D,et al. Polycystin-2 expression is regulated by a PC2-binding domain of intracellular portion of fibrocystin.J Biol Chem.2008 Nov 14;283(46):31559-66.
    Lager DJ, Qian Q, Bengal RJ, et al. The pck rat:a new model that resembles human autosomal dominant polycystic kidney and liver disease. Kidney Int.2001 Jan;59(1):126-36.
    Lens XM, Onuchic LF, Wu Q et al. An integrated genetic and physical map of the autosomal recessive polycystic kidney disease region. Genomics 41,463-466 (1997).
    Lilova MI, Petkov DL. Intracranial aneurysms in a child with autosomal recessive polycystic kidney disease. Pediatr Nephrol 2001,16,1030-1032.
    Lonergan, G J., Rice, R. R., Suarez, E. S. Autosomal recessive polycystic kidney disease: radiologic-pathologic correlation. Radiographics,2000,20,837-855.
    Mai W, Chen D, Ding T, et al. Inhibition of Pkhdl impairs tubulomorphogenesis of cultured IMCD cells. Mol Biol Cell.2005 Sep;16(9):4398-409.
    Menezes LF, Cai Y, Nagasawa Y, et al. Polyductin, the PKHD1 gene product, comprises isoforms expressed in plasma membrane, primary cilium, and cytoplasm. Kidney Int. 2004 Oct;66(4):1345-55.
    Moser M, Matthiesen S, Kirfel J, et al. A mouse model for cystic biliary dysgenesis in autosomal recessive polycystic kidney disease (ARPKD). Hepatology 2005, 41:1113-1121.
    Mucher G, Becker J, Knapp M, et al. Fine mapping of the autosomal recessive polycystic kidney disease locus (PKHD1) and the genes MUT, RDS, CSNK2β, and GSTA1 at 6p21.2-pl2. Genomics 48,40-45 (1998).
    Nagasawa Y, Matthiesen S, Onuchic LF, et al. Identification and characterization of Pkhdl, the mouse orthologue of the human ARPKD gene. J Am Soc Nephrol.2002 Sep;13(9):2246-58.
    Neumann, H. P., Krumme, B., van Velthoven, V, et al. Multiple intracranial aneurysms in a patient with autosomal recessive polycystic kidney disease. Nephrol Dial Transplant,1999,14,936-939.
    Onuchic LF, Furu L, Nagasawa Y, et al. PKHD1, the polycystic kidney and hepatic disease 1 gene, encodes a novel large protein containing multiple immunoglobulin-like plexin-transcription-factor domains and parallel beta-helix 1 repeats. Am J Hum Genet. 2002 May;70(5):1305-17.
    Sharp AM, Messiaen LM, Page G, et al. Comprehensive genomic analysis of PKHD1 mutations in ARPKD cohorts. J Med Genet.2005 Apr;42(4):336-49
    Sjoblom T, Jones S, Wood LD, et al. The consensus coding sequences of human breast and colorectal cancers. Science.2006 Oct 13;314(5797):268-74.
    Sweeney WE Jr, Avner ED. Molecular and cellular pathophysiology of autosomal recessive polycystic kidney disease (ARPKD). Cell Tissue Res.2006 Dec;326(3):671-85.
    Torres VE, Harris PC. Mechanisms of Disease:autosomal dominant and recessive polycystic kidney diseases. Nat Clin Pract Nephrol.2006 Jan;2(1):40-55.
    Wang S, Zhang J, Nauli SM, et al. Fibrocystin/polyductin, found in the same protein complex with polycystin-2, regulates calcium responses in kidney epithelia.Mol Cell Biol.2007 Apr;27(8):3241-52.
    Ward CJ, Yuan D, Masyuk TV, et al. Cellular and subcellular localization of the ARPKD protein; fibrocystin is expressed on primary cilia.Hum Mol Genet.2003 Oct 15;12(20):2703-10.
    Williams SS, Cobo-Stark P, James LR, et al. Kidney cysts, pancreatic cysts, and biliary disease in a mouse model of autosomal recessive polycystic kidney disease. Pediatr Nephrol.2008 May;23(5):733-41.
    Woollard JR, Punyashtiti R, Richardson S, et al. A mouse model of autosomal recessive polycystic kidney disease with biliary duct and proximal tubule dilatation. Kidney Int. 2007 Aug;72(3):328-36.
    Xiong H, Chen Y, Yi Y, et al. A novel gene encoding a TIG multiple domain protein is a positional candidate for autosomal recessive polycystic kidney disease. Genomics.2002 Jul;80(1):96-104.
    Yang J, Zhang S, Zhou Q, et al. PKHD1 gene silencing may cause cell abnormal proliferation through modulation of intracellular calcium in autosomal recessive polycystic kidney disease. J Biochem Mol Biol.2007 Jul 31;40(4):467-74.
    Yonemura, K., Yasuda, H., Fujigaki, Y Adrenal insufficiency due to isolated adrenocorticotropin deficiency complicated by autosomal recessive polycystic kidney disease. Ren Fail 2003,25,485-492.
    Zerres K, Mucher G, Bachner L, et al. Mapping of the gene for autosomal recessive polycystic kidney disease (ARPKD) to chromosome 6p21-cen. Nat Genet.1994 Jul;7(3):429-32.
    Zerres K, Mucher G, Becker J. et al. Prenatal diagnosis of autosomal recessive polycystic kidney disease (ARPKD):molecular genetics, clinical experience, and fetal morphology. Am. J. Med. Genet.1998,76,137-144.
    Zerres, K., Rudnik-Schoneborn, S., Mucher, G Autosomal recessive polycystic kidney disease:clinical features and genetics. Adv Nephrol Necker Hosp 1996,25,147-157.
    Zerres, K., Rudnik-Schoneborn, S., Senderek, J., et al. Autosomal recessive polycystic kidney disease (ARPKD). J Nephrol 2003,16,453-458.
    Zhang MZ, Mai W, Li C, et al. PKHD1 protein encoded by the gene for autosomal recessive polycystic kidney disease associates with basal bodies and primary cilia in renal epithelial cells. Proc Natl Acad Sci U S A.2004 Feb 24;101(8):2311-6.