Retake the Center Stage - New Development of Rat Genetics

Journal of Genetics and Genomics

Volumn 39, Issue 6, 2012, Pages 261-268

  Zheng, Sushuang ^a   Geghman, Kindiya ^a   Shenoy, Sushila ^a   Li, Chenjian ^a,b  

^a MOUNT SINAI SCHOOL OF MEDICINE   (United States)

^b PEKING UNIVERSITY   (China)

Author keywords

BAC; Knockout rat; Mutagenesis; Rat model; Transgenic rat

Indexed keywords

COMPLEMENTARY DNA; ETHYLNITROSOUREA; MESSENGER RNA; NUCLEASE; OCTAMER TRANSCRIPTION FACTOR 4; PROTEIN TALEN; TRANSCRIPTION FACTOR NANOG; TRANSCRIPTION FACTOR SOX2; UNCLASSIFIED DRUG; ZINC FINGER NUCLEASE; ZINC FINGER PROTEIN;

BACTERIAL ARTIFICIAL CHROMOSOME; DNA MODIFICATION; DNA REPAIR; DOUBLE STRANDED DNA BREAK; EMBRYONIC STEM CELL; GENETIC MANIPULATION; HIGH THROUGHPUT SCREENING; MUTAGENESIS; NONHUMAN; OPEN READING FRAME; PHENOTYPE; REVIEW; TRANSGENICS; TRANSPOSON;

ANIMALS; ANIMALS, GENETICALLY MODIFIED; CHROMOSOMES, ARTIFICIAL, BACTERIAL; DISEASE MODELS, ANIMAL; DNA TRANSPOSABLE ELEMENTS; EMBRYONIC STEM CELLS; ETHYLNITROSOUREA; GENE TARGETING; HUMANS; MUTAGENESIS; RATS; RETROELEMENTS; ZINC FINGERS;

BACTERIA (MICROORGANISMS); RATTUS;

EID: 84863097837     PISSN: 16738527     EISSN: 18735533     Source Type: Journal    
DOI: 10.1016/j.jgg.2012.05.003     Document Type: Review

References (86)

1. Alam M., Mayerhofer A., Schmidt W.J. The neurobehavioral changes induced by bilateral rotenone lesion in medial forebrain bundle of rats are reversed by L-DOPA. Behav. Brain Res. 2004, 151:117-124.
2. Augustin M., Sedlmeier R., Peters T., Huffstadt U., Kochmann E., Simon D., Schoniger M., Garke-Mayerthaler S., Laufs J., Mayhaus M., Franke S., Klose M., Graupner A., Kurzmann M., Zinser C., Wolf A., Voelkel M., Kellner M., Kilian M., Seelig S., Koppius A., Teubner A., Korthaus D., Nehls M., Wattler S. Efficient and fast targeted production of murine models based on ENU mutagenesis. Mamm. Genome 2005, 16:405-413.
3. Boch J., Scholze H., Schornack S., Landgraf A., Hahn S., Kay S., Lahaye T., Nickstadt A., Bonas U. Breaking the code of DNA binding specificity of TAL-type III effectors. Science 2009, 326:1509-1512.
4. Bogdanove A.J., Schornack S., Lahaye T. TAL effectors: finding plant genes for disease and defense. Curr. Opin. Plant Biol. 2010, 13:394-401.
5. Bogdanove A.J., Voytas D.F. TAL effectors: customizable proteins for DNA targeting. Science 2011, 333:1843-1846.
6. Buehr M., Meek S., Blair K., Yang J., Ure J., Silva J., McLay R., Hall J., Ying Q.L., Smith A. Capture of authentic embryonic stem cells from rat blastocysts. Cell 2008, 135:1287-1298.
7. Cermak T., Doyle E.L., Christian M., Wang L., Zhang Y., Schmidt C., Baller J.A., Somia N.V., Bogdanove A.J., Voytas D.F. Efficient design and assembly of custom TALEN and other TAL effector-based constructs for DNA targeting. Nucleic Acids Res. 2011, 39:e82.
8. Christian M., Cermak T., Doyle E.L., Schmidt C., Zhang F., Hummel A., Bogdanove A.J., Voytas D.F. Targeting DNA double-strand breaks with TAL effector nucleases. Genetics 2010, 186:757-761.
9. Chu X., Zhang Z., Yabut J., Horwitz S., Levorse J., Li X.Q., Zhu L., Lederman H., Ortiga R., Strauss J., Li X., Owens K.A., Dragovic J., Vogt T., Evers R., Shin M.K. Characterization of multidrug resistance 1a/P-glycoprotein knockout rats generated by zinc finger nucleases. Mol. Pharmacol. 2012, 81:220-227.
10. Cordes S.P. N-ethyl-N-nitrosourea mutagenesis: boarding the mouse mutant express. Microbiol. Mol. Biol. Rev. 2005, 69:426-439.
11. Cui X., Ji D., Fisher D.A., Wu Y., Briner D.M., Weinstein E.J. Targeted integration in rat and mouse embryos with zinc-finger nucleases. Nat. Biotechnol. 2011, 29:64-67.
12. Ding S., Wu X., Li G., Han M., Zhuang Y., Xu T. Efficient transposition of the piggyBac (PB) transposon in mammalian cells and mice. Cell 2005, 122:473-483.
13. Evans M.J., Kaufman M.H. Establishment in culture of pluripotential cells from mouse embryos. Nature 1981, 292:154-156.
14. Feng Q., Moran J.V., Kazazian H.H., Boeke J.D. Human L1 retrotransposon encodes a conserved endonuclease required for retrotransposition. Cell 1996, 87:905-916.
15. Geurts A.M., Cost G.J., Freyvert Y., Zeitler B., Miller J.C., Choi V.M., Jenkins S.S., Wood A., Cui X., Meng X., Vincent A., Lam S., Michalkiewicz M., Schilling R., Foeckler J., Kalloway S., Weiler H., Menoret S., Anegon I., Davis G.D., Zhang L., Rebar E.J., Gregory P.D., Urnov F.D., Jacob H.J., Buelow R. Knockout rats via embryo microinjection of zinc-finger nucleases. Science 2009, 325:433.
16. Geurts A.M., Cost G.J., Remy S., Cui X., Tesson L., Usal C., Menoret S., Jacob H.J., Anegon I., Buelow R. Generation of gene-specific mutated rats using zinc-finger nucleases. Methods Mol. Biol. 2010, 597:211-225.
17. Geurts A.M., Moreno C. Zinc-finger nucleases: new strategies to target the rat genome. Clin. Sci. (Lond.) 2010, 119:303-311.
18. Gong S., Yang X.W., Li C., Heintz N. Highly efficient modification of bacterial artificial chromosomes (BACs) using novel shuttle vectors containing the R6Kγ origin of replication. Genome Res. 2002, 12:1992-1998.
19. Goto Y., Peachey N.S., Ripps H., Naash M.I. Functional abnormalities in transgenic mice expressing a mutant rhodopsin gene. Invest. Ophthalmol. Vis. Sci. 1995, 36:62-71.
20. Hammer R.E., Maika S.D., Richardson J.A., Tang J.P., Taurog J.D. Spontaneous inflammatory disease in transgenic rats expressing HLA-B27 and human beta 2m: an animal model of HLA-B27-associated human disorders. Cell 1990, 63:1099-1112.
21. Hockemeyer D., Wang H., Kiani S., Lai C.S., Gao Q., Cassady J.P., Cost G.J., Zhang L., Santiago Y., Miller J.C., Zeitler B., Cherone J.M., Meng X., Hinkley S.J., Rebar E.J., Gregory P.D., Urnov F.D., Jaenisch R. Genetic engineering of human pluripotent cells using TALE nucleases. Nat. Biotechnol. 2011, 29:731-734.
22. Hodgson J.G., Agopyan N., Gutekunst C.A., Leavitt B.R., LePiane F., Singaraja R., Smith D.J., Bissada N., McCutcheon K., Nasir J., Jamot L., Li X.J., Stevens M.E., Rosemond E., Roder J.C., Phillips A.G., Rubin E.M., Hersch S.M., Hayden M.R. A YAC mouse model for Huntington's disease with full-length mutant huntingtin, cytoplasmic toxicity, and selective striatal neurodegeneration. Neuron 1999, 23:181-192.
23. Homberg J.R., Mul J.D., de Wit E., Cuppen E. Complete knockout of the nociceptin/orphanin FQ receptor in the rat does not induce compensatory changes in μ, δ and κ opioid receptors. Neuroscience 2009, 163:308-315.
24. Homberg J.R., Olivier J.D., Smits B.M., Mul J.D., Mudde J., Verheul M., Nieuwenhuizen O.F., Cools A.R., Ronken E., Cremers T., Schoffelmeer A.N., Ellenbroek B.A., Cuppen E. Characterization of the serotonin transporter knockout rat: a selective change in the functioning of the serotonergic system. Neuroscience 2007, 146:1662-1676.
25. Hong J., He H., Weiss M.L. Derivation and characterization of embryonic stem cells lines derived from transgenic Fischer 344 and Dark Agouti rats. Stem Cells Dev 2011, Nov 22, [Epub ahead of print].
26. Huang G., Ashton C., Kumbhani D.S., Ying Q.L. Genetic manipulations in the rat: progress and prospects. Curr. Opin. Nephrol. Hypertens. 2011, 20:391-399.
27. Huang G., Tong C., Kumbhani D.S., Ashton C., Yan H., Ying Q.L. Beyond knockout rats: new insights into finer genome manipulation in rats. Cell Cycle 2011, 10:1059-1066.
28. Huang P., Xiao A., Zhou M., Zhu Z., Lin S., Zhang B. Heritable gene targeting in zebrafish using customized TALENs. Nat. Biotechnol. 2011, 29:699-700.
29. Ivics Z., Izsvak Z., Chapman K.M., Hamra F.K. Sleeping Beauty transposon mutagenesis of the rat genome in spermatogonial stem cells. Methods 2011, 53:356-365.
30. Ivics Z., Izsvak Z., Medrano G., Chapman K.M., Hamra F.K. Sleeping Beauty transposon mutagenesis in rat spermatogonial stem cells. Nat. Protoc. 2011, 6:1521-1535.
31. Izsvák Z., Frohlich J., Grabundzija I., Shirley J.R., Powell H.M., Chapman K.M., Ivics Z., Hamra F.K. Generating knockout rats by transposon mutagenesis in spermatogonial stem cells. Nat. Methods 2010, 7:443-445.
32. Jacob H.J., Lazar J., Dwinell M.R., Moreno C., Geurts A.M. Gene targeting in the rat: advances and opportunities. Trends Genet. 2010, 26:510-518.
33. Kaufman C.D., Izsvak Z., Katzer A., Ivics Z. Frog Prince transposon-based RNAi vectors mediate efficient gene knockdown in human cells. J. RNAi Gene Silencing 2005, 1:97-104.
34. Kawamata M., Ochiya T. Generation of genetically modified rats from embryonic stem cells. Proc. Natl. Acad. Sci. USA 2010, 107:14223-14228.
35. Kitada K., Keng V.W., Takeda J., Horie K. Generating mutant rats using the Sleeping Beauty transposon system. Methods 2009, 49:236-242.
36. Kuo Y.M., Li Z., Jiao Y., Gaborit N., Pani A.K., Orrison B.M., Bruneau B.G., Giasson B.I., Smeyne R.J., Gershon M.D., Nussbaum R.L. Extensive enteric nervous system abnormalities in mice transgenic for artificial chromosomes containing Parkinson disease-associated α-synuclein gene mutations precede central nervous system changes. Hum. Mol. Genet. 2010, 19:1633-1650.
37. Li P., Tong C., Mehrian-Shai R., Jia L., Wu N., Yan Y., Maxson R.E., Schulze E.N., Song H., Hsieh C.L., Pera M.F., Ying Q.L. Germline competent embryonic stem cells derived from rat blastocysts. Cell 2008, 135:1299-1310.
38. Li Y., Liu W., Oo T.F., Wang L., Tang Y., Jackson-Lewis V., Zhou C., Geghman K., Bogdanov M., Przedborski S., Beal M.F., Burke R.E., Li C. Mutant LRRK2(R1441G) BAC transgenic mice recapitulate cardinal features of Parkinson's disease. Nat. Neurosci. 2009, 12:826-828.
39. Li T., Huang S., Jiang W.Z., Wright D., Spalding M.H., Weeks D.P., Yang B. TAL nucleases (TALNs): hybrid proteins composed of TAL effectors and Fok I DNA-cleavage domain. Nucleic Acids Res. 2011, 39:359-372.
40. Li T., Huang S., Zhao X., Wright D.A., Carpenter S., Spalding M.H., Weeks D.P., Yang B. Modularly assembled designer TAL effector nucleases for targeted gene knockout and gene replacement in eukaryotes. Nucleic Acids Res. 2011, 39:6315-6325.
41. Lieber M.R. The mechanism of human nonhomologous DNA end joining. J. Biol. Chem. 2008, 283:1-5.
42. Lindsey J.R. Historical foundations in the laboratory rat. The Laboratory Rat 1979, 1-36. Academic Press, New York. H.J. Baker, J.R. Lindsey, S.H. Weisbroth (Eds.).
43. Liu G., Geurts A.M., Yae K., Srinivasan A.R., Fahrenkrug S.C., Largaespada D.A., Takeda J., Horie K., Olson W.K., Hackett P.B. Target-site preferences of Sleeping Beauty transposons. J. Mol. Biol. 2005, 346:161-173.
44. Liu L., Orozco I.J., Planel E., Wen Y., Bretteville A., Krishnamurthy P., Wang L., Herman M., Figueroa H., Yu W.H., Arancio O., Duff K. A transgenic rat that develops Alzheimer's disease-like amyloid pathology, deficits in synaptic plasticity and cognitive impairment. Neurobiol. Dis. 2008, 31:46-57.
45. Liu J., Li C., Yu Z., Huang P., Wu H., Wei C., Zhu N., Shen Y., Chen Y., Zhang B., Deng W.M., Jiao R. Efficient and specific modifications of the Drosophila genome by means of an easy TALEN strategy. J. Genet. Genomics 2012, 5:209-215.
46. Lo Bianco C., Schneider B.L., Bauer M., Sajadi A., Brice A., Iwatsubo T., Aebischer P. Lentiviral vector delivery of parkin prevents dopaminergic degeneration in an α-synuclein rat model of Parkinson's disease. Proc. Natl. Acad. Sci. USA 2004, 101:17510-17515.
47. Machida S., Kondo M., Jamison J.A., Khan N.W., Kononen L.T., Sugawara T., Bush R.A., Sieving P.A. P23H rhodopsin transgenic rat: correlation of retinal function with histopathology. Invest. Ophthalmol. Vis. Sci. 2000, 41:3200-3209.
48. Mahfouz M.M., Li L., Shamimuzzaman M., Wibowo A., Fang X., Zhu J.K. De novo-engineered transcription activator-like effector (TALE) hybrid nuclease with novel DNA binding specificity creates double-strand breaks. Proc. Natl. Acad. Sci. USA 2011, 108:2623-2628.
49. Martin G.R. Isolation of a pluripotent cell line from early mouse embryos cultured in medium conditioned by teratocarcinoma stem cells. Proc. Natl. Acad. Sci. USA 1981, 78:7634-7638.
50. Mashimo T., Takizawa A., Voigt B., Yoshimi K., Hiai H., Kuramoto T., Serikawa T. Generation of knockout rats with X-linked severe combined immunodeficiency (X-SCID) using zinc-finger nucleases. PLoS ONE 2010, 5:e8870.
51. Mátés L., Chuah M.K., Belay E., Jerchow B., Manoj N., Acosta-Sanchez A., Grzela D.P., Schmitt A., Becker K., Matrai J., Ma L., Samara-Kuko E., Gysemans C., Pryputniewicz D., Miskey C., Fletcher B., VandenDriessche T., Ivics Z., Izsvak Z. Molecular evolution of a novel hyperactive Sleeping Beauty transposase enables robust stable gene transfer in vertebrates. Nat. Genet. 2009, 41:753-761.
52. Mathias S.L., Scott A.F., Kazazian H.H., Boeke J.D., Gabriel A. Reverse transcriptase encoded by a human transposable element. Science 1991, 254:1808-1810.
53. Ménoret S., Iscache A.L., Tesson L., Remy S., Usal C., Osborn M.J., Cost G.J., Bruggemann M., Buelow R., Anegon I. Characterization of immunoglobulin heavy chain knockout rats. Eur. J. Immunol. 2010, 40:2932-2941.
54. Miller J.C., Tan S., Qiao G., Barlow K.A., Wang J., Xia D.F., Meng X., Paschon D.E., Leung E., Hinkley S.J., Dulay G.P., Hua K.L., Ankoudinova I., Cost G.J., Urnov F.D., Zhang H.S., Holmes M.C., Zhang L., Gregory P.D., Rebar E.J. A TALE nuclease architecture for efficient genome editing. Nat. Biotechnol. 2011, 29:143-148.
55. Morbitzer R., Elsaesser J., Hausner J., Lahaye T. Assembly of custom TALE-type DNA binding domains by modular cloning. Nucleic Acids Res. 2011, 39:5790-5799.
56. Moreno C., Hoffman M., Stodola T.J., Didier D.N., Lazar J., Geurts A.M., North P.E., Jacob H.J., Greene A.S. Creation and characterization of a renin knockout rat. Hypertension 2011, 57:614-619.
57. Moscou M.J., Bogdanove A.J. A simple cipher governs DNA recognition by TAL effectors. Science 2009, 326:1501.
58. Mussolino C., Morbitzer R., Lutge F., Dannemann N., Lahaye T., Cathomen T. A novel TALE nuclease scaffold enables high genome editing activity in combination with low toxicity. Nucleic Acids Res. 2011, 39:9283-9293.
59. Pavletich N.P., Pabo C.O. Zinc finger-DNA recognition: crystal structure of a Zif268-DNA complex at 2.1 A. Science 1991, 252:809-817.
60. Philipeaux J.M. Note sur l'extirpation des capsules surrenales chez les rats albinos. C. R. Acad. Sci. (Paris) 1856, 43:904.
61. Porteus M.H., Carroll D. Gene targeting using zinc finger nucleases. Nat. Biotechnol. 2005, 23:967-973.
62. Reyon D., Tsai S.Q., Khayter C., Foden J.A., Sander J.D., Joung J.K. FLASH assembly of TALENs for high-throughput genome editing. Nat. Biotechnol 2012, 30:460-465.
63. Sander J.D., Cade L., Khayter C., Reyon D., Peterson R.T., Joung J.K., Yeh J.R. Targeted gene disruption in somatic zebrafish cells using engineered TALENs. Nat. Biotechnol. 2011, 29:697-698.
64. Scholze H., Boch J. TAL effectors are remote controls for gene activation. Curr. Opin. Microbiol. 2011, 14:47-53.
65. Shizuya H., Birren B., Kim U.J., Mancino V., Slepak T., Tachiiri Y., Simon M. Cloning and stable maintenance of 300-kilobase-pair fragments of human DNA in Escherichia coli using an F-factor-based vector. Proc. Natl. Acad. Sci. USA 1992, 89:8794-8797.
66. Smits B.M., Mudde J.B., van de Belt J., Verheul M., Olivier J., Homberg J., Guryev V., Cools A.R., Ellenbroek B.A., Plasterk R.H., Cuppen E. Generation of gene knockouts and mutant models in the laboratory rat by ENU-driven target-selected mutagenesis. Pharmacogenet. Genomics 2006, 16:159-169.
67. Takeda J., Keng V.W., Horie K. Germline mutagenesis mediated by Sleeping Beauty transposon system in mice. Genome Biol. 2007, 8(Suppl. 1):S14.
68. Tesson L., Usal C., Menoret S., Leung E., Niles B.J., Remy S., Santiago Y., Vincent A.I., Meng X., Zhang L., Gregory P.D., Anegon I., Cost G.J. Knockout rats generated by embryo microinjection of TALENs. Nat. Biotechnol. 2011, 29:695-696.
69. Tong C., Huang G., Ashton C., Li P., Ying Q.L. Generating gene knockout rats by homologous recombination in embryonic stem cells. Nat. Protoc. 2011, 6:827-844.
70. Tong C., Li P., Wu N.L., Yan Y., Ying Q.L. Production of p53 gene knockout rats by homologous recombination in embryonic stem cells. Nature 2010, 467:211-213.
71. Ueda S., Kawamata M., Teratani T., Shimizu T., Tamai Y., Ogawa H., Hayashi K., Tsuda H., Ochiya T. Establishment of rat embryonic stem cells and making of chimera rats. PLoS ONE 2008, 3:e2800.
72. Urnov F.D., Rebar E.J., Holmes M.C., Zhang H.S., Gregory P.D. Genome editing with engineered zinc finger nucleases. Nat. Rev. Genet. 2010, 11:636-646.
73. van Boxtel R., Gould M.N., Cuppen E., Smits B.M. ENU mutagenesis to generate genetically modified rat models. Methods Mol. Biol. 2010, 597:151-167.
74. Vasireddy V., Chavali V.R., Joseph V.T., Kadam R., Lin J.H., Jamison J.A., Kompella U.B., Reddy G.B., Ayyagari R. Rescue of photoreceptor degeneration by curcumin in transgenic rats with P23H rhodopsin mutation. PLoS ONE 2011, 6:e21193.
75. von Hörsten S., Schmitt I., Nguyen H.P., Holzmann C., Schmidt T., Walther T., Bader M., Pabst R., Kobbe P., Krotova J., Stiller D., Kask A., Vaarmann A., Rathke-Hartlieb S., Schulz J.B., Grasshoff U., Bauer I., Vieira-Saecker A.M., Paul M., Jones L., Lindenberg K.S., Landwehrmeyer B., Bauer A., Li X.J., Riess O. Transgenic rat model of Huntington's disease. Hum. Mol. Genet. 2003, 12:617-624.
76. Wheeler V.C., Auerbach W., White J.K., Srinidhi J., Auerbach A., Ryan A., Duyao M.P., Vrbanac V., Weaver M., Gusella J.F., Joyner A.L., MacDonald M.E. Length-dependent gametic CAG repeat instability in the Huntington's disease knock-in mouse. Hum. Mol. Genet. 1999, 8:115-122.
77. Woltjen K., Michael I.P., Mohseni P., Desai R., Mileikovsky M., Hamalainen R., Cowling R., Wang W., Liu P., Gertsenstein M., Kaji K., Sung H.K., Nagy A. piggyBac transposition reprograms fibroblasts to induced pluripotent stem cells. Nature 2009, 458:766-770.
78. Wood A.J., Lo T.W., Zeitler B., Pickle C.S., Ralston E.J., Lee A.H., Amora R., Miller J.C., Leung E., Meng X., Zhang L., Rebar E.J., Gregory P.D., Urnov F.D., Meyer B.J. Targeted genome editing across species using ZFNs and TALENs. Science 2011, 333:307.
79. Xu Q., Shenoy S., Li C. Mouse models for LRRK2 Parkinson's disease. Parkinsonism. Relat. Disord. 2012, 18(Suppl. 1):S186-S189.
80. Yamada M., Iwatsubo T., Mizuno Y., Mochizuki H. Overexpression of α-synuclein in rat substantia nigra results in loss of dopaminergic neurons, phosphorylation of alpha-synuclein and activation of caspase-9: resemblance to pathogenetic changes in Parkinson's disease. J. Neurochem. 2004, 91:451-461.
81. Yamamoto S., Nakata M., Sasada R., Ooshima Y., Yano T., Shinozawa T., Tsukimi Y., Takeyama M., Matsumoto Y., Hashimoto T. Derivation of rat embryonic stem cells and generation of protease-activated receptor-2 knockout rats. Transgenic Res 2011, 10.1007/s11248-011-9564-0.
82. Yang X.W., Model P., Heintz N. Homologous recombination based modification in Escherichia coli and germline transmission in transgenic mice of a bacterial artificial chromosome. Nat. Biotechnol. 1997, 15:859-865.
83. Ying Q.L., Wray J., Nichols J., Batlle-Morera L., Doble B., Woodgett J., Cohen P., Smith A. The ground state of embryonic stem cell self-renewal. Nature 2008, 453:519-523.
84. Zan Y., Haag J.D., Chen K.S., Shepel L.A., Wigington D., Wang Y.R., Hu R., Lopez-Guajardo C.C., Brose H.L., Porter K.I., Leonard R.A., Hitt A.A., Schommer S.L., Elegbede A.F., Gould M.N. Production of knockout rats using ENU mutagenesis and a yeast-based screening assay. Nat. Biotechnol. 2003, 21:645-651.
85. Zhang F., Cong L., Lodato S., Kosuri S., Church G.M., Arlotta P. Efficient construction of sequence-specific TAL effectors for modulating mammalian transcription. Nat. Biotechnol. 2011, 29:149-153.
86. Zhao X., Lv Z., Liu L., Wang L., Tong M., Zhou Q. Derivation of embryonic stem cells from Brown Norway rats blastocysts. J. Genet. Genomics 2010, 37:467-473.