Food Safety, Animal Health and Welfare and Environmental Impact of Animals derived from Cloning by Somatic Cell Nucleus Transfer (SCNT) and their Offspring and Products Obtained from those Animals

EFSA Journal

Volumn 6, Issue 7, 2008, Pages

[No Author Info available]

Author keywords

Animal Cloning; Animal Health; Animal Welfare; ART; Assisted Reproduction Technology; Bovine; Cattle; Clone; Clones; Environmental Impact; Epigenetic Reprogramming; Food Product; Food Safety; Genetic Diversity; Immunocompetence; Livestock; Offspring; Pig; Progeny; Risk Assessment; SCNT; Somatic Cell Nucleus Transfer; Swine

Indexed keywords

EID: 85078397129     PISSN: None     EISSN: 18314732     Source Type: Journal    
DOI: 10.2903/j.efsa.2008.767     Document Type: Article

References (180)

1. ACNFP (Advisory Committee on Novel Foods and Processes) 1998. Toxicological assessment of novel (including GM) foods. HMSO, London, last accessed 2008–01-23.
2. Archer, G. S., Dindot, S., Friend, T. H., Walker, S., Zaunbrecher, G., Lawhorn, B. and Piedrahita, J. A. 2003a. Hierarchical phenotypic and epigenetic variation in cloned swine. Biol Reprod 69 (2): 430–6.
3. Archer, G. S., Friend, T. H., Piedrahita, J., Nevill, C. H. and Walker, S. 2003b. Behavioral variation among cloned pigs. Applied Animal Behaviour Science 82 (2): 151.
4. Archer, G. S., Friend, T. H., Piedrahita, J., Nevill, C. H. and Walker, S. 2003c. Behavioral variation among cloned pigs. Applied Animal Behaviour Science 81 (4): 321.
5. Arnold, D. R., Bordignon, V., Lefebvre, R., Murphy, B. D. and Smith, L. C. 2006. Somatic cell nuclear transfer alters peri-implantation trophoblast differentiation in bovine embryos. Reproduction 132 (2): 279–90.
6. Balbach, S. T., Jauch, A., Bohm-Steuer, B., Cavaleri, F. M., Han, Y. M. and Boiani, M. 2007. Chromosome stability differs in cloned mouse embryos and derivative ES cells. Dev Biol 308 (2): 309–21.
7. Batchelder, C. A., Bertolini, M., Mason, J. B., Moyer, A. L., Hoffert, K. A., Petkov, S. G., Famula, T. R., Angelos, J., George, L. W. and Anderson, G. B. 2007a. Perinatal physiology in cloned and normal calves: physical and clinical characteristics. Cloning Stem Cells 9 (1): 63–82.
8. Batchelder, C. A., Bertolini, M., Mason, J. B., Moyer, A. L., Hoffert, K. A., Petkov, S. G., Famula, T. R., Angelos, J., George, L. W. and Anderson, G. B. 2007b. Perinatal physiology in cloned and normal calves: hematologic and biochemical profiles. Cloning Stem Cells 9 (1): 83–96.
9. Batchelder, C. A., Hoffert, K. A., Bertolini, M., Moyer, A. L., Mason, J. B., Petkov, S. G., Famula, T. R. and Anderson, G. B. 2005. Effect of the nuclear-donor cell lineage, type, and cell donor on development of somatic cell nuclear transfer embryos in cattle. Cloning Stem Cells 7 (4): 238–54.
10. Beaujean, N., Taylor, J., Gardner, J., Wilmut, I., Meehan, R. and Young, L. 2004. Effect of limited DNA methylation reprogramming in the normal sheep embryo on somatic cell nuclear transfer. Biol Reprod 71 (1): 185–93.
11. Belitz, H. D., Grosch, W., Schieberle, P., 2004. Food Chemistry. Editor. Springer-Verlag GmbH, Pages.
12. Betts, D. H., Perrault, S. D., Petrik, J., Lin, L., Favetta, L. A., Keefer, C. L. and King, W. A. 2005. Telomere length analysis in goat clones and their offspring. Mol Reprod Dev 72 (4): 461–70.
13. Bielanski, A. 1997. A review on disease transmission studies in relationship to production of embryos by in vitro fertilization and to related new reproductive technologies. Biotechnol Adv 15 (3–4): 633–56.
14. Booth, P. J., Viuff, D., Tan, S., Holm, P., Greve, T. and Callesen, H. 2003. Numerical chromosome errors in day 7 somatic nuclear transfer bovine blastocysts. Biol Reprod 68 (3): 922–8.
15. Braastad, B. O., Osadchuk, L. V., Lund, G. and Bakken, M. 1998. Effects of prenatal handling stress on adrenal weight and function and behaviour in novel situations in blue fox cubs (Alopex lagopus). Applied Animal Behaviour Science 57 (1–2): 157–169.
16. Brambrink, T., Hochedlinger, K., Bell, G. and Jaenisch, R. 2006. ES cells derived from cloned and fertilized blastocysts are transcriptionally and functionally indistinguishable. Proc Natl Acad Sci U S A 103 (4): 933–8.
17. Caballero, B. 2003. Encyclopedia of Food Sciences and Nutrition. Editor. Elsevier Science Ltd, Pages.
18. Camargo, L. S., Viana, J. H., Sa, W. F., Ferreira, A. M. and Vale Filho, V. R. 2005. Developmental competence of oocytes from prepubertal Bos indicus crossbred cattle. Anim Reprod Sci 85 (1–2): 53–9.
19. Carroll, J. A., Carter, D. B., Korte, S. W. and Prather, R. S. 2005. Evaluation of the acute phase response in cloned pigs following a lipopolysaccharide challenge. Domest Anim Endocrinol 29 (3): 564–72.
20. Casolini, P., Cigliana, G., Alema, G. S., Ruggieri, V., Angelucci, L. and Catalani, A. 1997. Effect of increased maternal corticosterone during lactation on hippocampal corticosteroid receptors, stress response and learning in offspring in the early stages of life. Neuroscience 79 (4): 1005–12.
21. Charlier, C., Segers, K., Karim, L., Shay, T., Gyapay, G., Cockett, N. and Georges, M. 2001. The callipyge mutation enhances the expression of coregulated imprinted genes in cis without affecting their imprinting status. Nat Genet 27 (4): 367–9.
22. Chastant-Maillard, S., Quinton, H., Lauffenburger, J., Cordonnier-Lefort, N., Richard, C., Marchal, J., Mormede, P. and Renard, J. P. 2003. Consequences of transvaginal follicular puncture on well-being in cows. Reproduction 125 (4): 555–63.
23. Chavatte-Palmer, P., de Sousa, N., Laigre, P., Camous, S., Ponter, A. A., Beckers, J. F. and Heyman, Y. 2006. Ultrasound fetal measurements and pregnancy associated glycoprotein secretion in early pregnancy in cattle recipients carrying somatic clones. Theriogenology 66 (4): 829–840.
24. Chavatte-Palmer, P. and Guillomot, M. 2007. Comparative implantation and placentation. Gynecol Obstet Invest 64 (3): 166–74.
25. Chavatte-Palmer, P., Heyman, Y., Richard, C., Monget, P., LeBourhis, D., Kann, G., Chilliard, Y., Vignon, X. and Renard, J. P. 2002. Clinical, hormonal, and hematologic characteristics of bovine calves derived from nuclei from somatic cells. Biol Reprod 66 (6): 1596–603.
26. Chavatte-Palmer, P., Remy, D., Cordonnier, N., Richard, C., Issenman, H., Laigre, P., Heyman, Y. and Mialot, J. P. 2004. Health status of cloned cattle at different ages. Cloning Stem Cells 6 (2): 94–100.
27. Cibelli, J. B., Stice, S. L., Golueke, P. J., Kane, J. J., Jerry, J., Blackwell, C., Ponce de Leon, F. A. and Robl, J. M. 1998. Cloned transgenic calves produced from nonquiescent fetal fibroblasts. Science 280 (5367): 1256–8.
28. Coan, P. M., Burton, G. J. and Ferguson-Smith, A. C. 2005. Imprinted genes in the placenta-a review. Placenta 26 Suppl A: S10–20.
29. Constant, F., Guillomot, M., Heyman, Y., Vignon, X., Laigre, P., Servely, J. L., Renard, J. P. and Chavatte-Palmer, P. 2006. Large offspring or large placenta syndrome? Morphometric analysis of late gestation bovine placentomes from somatic nuclear transfer pregnancies complicated by hydrallantois. Biol Reprod 75 (1): 122–30.
30. Cooney, C. A., Dave, A. A. and Wolff, G. L. 2002. Maternal methyl supplements in mice affect epigenetic variation and DNA methylation of offspring. J Nutr 132 (8 Suppl): 2393–2400S.
31. Coulon, M., Baudoin, C., Depaulis-Carre, M., Heyman, Y., Renard, J. P., Richard, C. and Deputte, B. L. 2007. Dairy cattle exploratory and social behaviors: is there an effect of cloning? Theriogenology 68 (8): 1097–103.
32. De Sousa, P. A., Dobrinsky, J. R., Zhu, J., Archibald, A. L., Ainslie, A., Bosma, W., Bowering, J., Bracken, J., Ferrier, P. M., Fletcher, J., Gasparrini, B., Harkness, L., Johnston, P., Ritchie, M., Ritchie, W. A., Travers, A., Albertini, D., Dinnyes, A., King, T. J. and Wilmut, I. 2002. Somatic cell nuclear transfer in the pig: control of pronuclear formation and integration with improved methods for activation and maintenance of pregnancy. Biol Reprod 66 (3): 642–50.
33. Dean, W., Santos, F., Stojkovic, M., Zakhartchenko, V., Walter, J., Wolf, E. and Reik, W. 2001. Conservation of methylation reprogramming in mammalian development: aberrant reprogramming in cloned embryos. Proc Natl Acad Sci U S A 98 (24): 13734–8.
34. Du, Y., Kragh, P. M., Zhang, Y., Li, J., Schmidt, M., Bogh, I. B., Zhang, X., Purup, S., Jorgensen, A. L., Pedersen, A. M., Villemoes, K., Yang, H., Bolund, L. and Vajta, G. 2007. Piglets born from handmade cloning, an innovative cloning method without micromanipulation. Theriogenology 68 (8): 1104–10.
35. Eggan, K., Akutsu, H., Hochedlinger, K., Rideout, W., 3rd Yanagimachi, R. and Jaenisch, R. 2000. X-Chromosome inactivation in cloned mouse embryos. Science 290 (5496): 1578–81.
36. Enright, B. P., Taneja, M., Schreiber, D., Riesen, J., Tian, X. C., Fortune, J. E. and Yang, X. 2002. Reproductive characteristics of cloned heifers derived from adult somatic cells. Biol Reprod 66 (2): 291–6.
37. Estrada, J., Sommer, J., Collins, B., Mir, B., Martin, A., York, A., Petters, R. M. and Piedrahita, J. A. 2007. Swine generated by somatic cell nuclear transfer have increased incidence of intrauterine growth restriction (IUGR). Cloning Stem Cells 9 (2): 229–36.
38. FAO (Food and Agricultural Organization), 2007. The State of the World's Animal Genetic Resources for Food and Agriculture. FAO. Food and Agriculture Organization, , last accessed 2008–01-23. 1-523.
39. Farin, P. W. and Farin, C. E. 1995. Transfer of bovine embryos produced in vivo or in vitro: survival and fetal development. Biol Reprod 52 (3): 676–82.
40. Farin, P. W., Piedrahita, J. A. and Farin, C. E. 2006. Errors in development of fetuses and placentas from in vitro-produced bovine embryos. Theriogenology 65 (1): 178–91.
41. FDA (Food and Drug Administration), 2008. Animal Cloning: A Risk Assessment. Center for Veterinary Medicine, U.S. Food and Drug Administration, Department of Health and Human Services, Pages 1–968, www.fda.gov/cvm/CloneRiskAssessment_Final.htm, last accessed 2008–06-271.
42. Forsberg, E. J., Strelchenko, N. S., Augenstein, M. L., Betthauser, J. M., Childs, L. A., Eilertsen, K. J., Enos, J. M., Forsythe, T. M., Golueke, P. J., Koppang, R. W., Lange, G., Lesmeister, T. L., Mallon, K. S., Mell, G. D., Misica, P. M., Pace, M. M., Pfister-Genskow, M., Voelker, G. R., Watt, S. R. and Bishop, M. D. 2002. Production of cloned cattle from in vitro systems. Biol Reprod 67 (1): 327–33.
43. Galli, C., Duchi, R., Moor, R. M. and Lazzari, G. 1999. Mammalian leukocytes contain all the genetic information necessary for the development of a new individual. Cloning 1 (3): 161–70.
44. Galli, C., Lagutina, I., Crotti, G., Colleoni, S., Turini, P., Ponderato, N., Duchi, R. and Lazzari, G. 2003. Pregnancy: a cloned horse born to its dam twin. Nature 424 (6949): 635.
45. Gardner, D. K. and Lane, M. 2005. Ex vivo early embryo development and effects on gene expression and imprinting. Reprod Fertil Dev 17 (3): 361–70.
46. Gluckman, P. D., Hanson, M. A. and Beedle, A. S. 2007a. Early life events and their consequences for later disease: a life history and evolutionary perspective. Am J Hum Biol 19 (1): 1–19.
47. Gluckman, P. D., Hanson, M. A. and Beedle, A. S. 2007b. Non-genomic transgenerational inheritance of disease risk. Bioessays 29 (2): 145–54.
48. Grimshaw, G. M., Sitarenios, G. and Finegan, J. A. 1995. Mental rotation at 7 years: relations with prenatal testosterone levels and spatial play experiences. Brain Cogn 29 (1): 85–100.
49. Grunau, R. V. E., Whitfield, M. F. and Petrie, J. H. 1994a. Pain sensitivity and temperament in extremely low-birth-weight premature toddlers and preterm and full-term controls. Pain 58 (3): 341–346.
50. Grunau, R. V. E., Whitfield, M. F., Petrie, J. H. and Fryer, E. L. 1994b. Early pain experience, child and family factors, as precursors of somatization: a prospective study of extremely premature and fullterm children. Pain 56 (3): 353–359.
51. Gschwind, D., Hassig, M. and Bleul, U. 2003. [Retrospective study of the fertility outlook in cows after caesarean section]. Schweiz Arch Tierheilkd 145 (4): 161–7.
52. Han, Z., Mtango, N. R., Patel, B. G., Sapienza, C. and Latham, K. E. 2008. Hybrid Vigor and Transgenerational Epigenetic Effects on Early Mouse Embryo Phenotype. Biol Reprod:
53. Hanada, H., Takeda, K., Tagami, T., Nirasawa, K., Akagi, S., Adachi, N., Takahashi, S., Izaike, Y., Iwamoto, M., Fuchimoto, D., Miyashita, N., Kubo, M., Onishi, A. and King, W. A. 2005. Chromosomal instability in the cattle clones derived by somatic cell nuclear-transfer. Mol Reprod Dev 71 (1): 36–44.
54. Hashizume, K., Ishiwata, H., Kizaki, K., Yamada, O., Takahashi, T., Imai, K., Patel, O. V., Akagi, S., Shimizu, M., Takahashi, S., Katsuma, S., Shiojima, S., Hirasawa, A., Tsujimoto, G., Todoroki, J. and Izaike, Y. 2002. Implantation and placental development in somatic cell clone recipient cows. Cloning Stem Cells 4 (3): 197–209.
55. Heyman, Y., Chavatte-Palmer, P., Berthelot, V., Fromentin, G., Hocquette, J. F., Martignat, L. and Renard, J. P. 2007a. Assessing the quality of products from cloned cattle: an integrative approach. Theriogenology 67 (1): 134–41.
56. Heyman, Y., Chavatte-Palmer, P., Fromentin, G., Berthelot, V., Jurie, C., Bas, P., Dubarry, M., Mialot, J. P., Remy, D., Richard, C., Martignat, L., Vignon, X. and Renard, J. P. 2007b. Quality and safety of bovine clones and their products. Animal (1): 963–972.
57. Heyman, Y., Chavatte-Palmer, P., LeBourhis, D., Camous, S., Vignon, X. and Renard, J. P. 2002. Frequency and occurrence of late-gestation losses from cattle cloned embryos. Biol Reprod 66 (1): 6–13.
58. Heyman, Y., Richard, C., Rodriguez-Martinez, H., Lazzari, G., Chavatte-Palmer, P., Vignon, X. and Galli, C. 2004. Zootechnical performance of cloned cattle and offspring: preliminary results. Cloning Stem Cells 6 (2): 111–20.
59. Hiendleder, S., Mund, C., Reichenbach, H. D., Wenigerkind, H., Brem, G., Zakhartchenko, V., Lyko, F. and Wolf, E. 2004. Tissue-specific elevated genomic cytosine methylation levels are associated with an overgrowth phenotype of bovine fetuses derived by in vitro techniques. Biol Reprod 71 (1): 217–23.
60. Hiendleder, S., Wirtz, M., Mund, C., Klempt, M., Reichenbach, H. D., Stojkovic, M., Weppert, M., Wenigerkind, H., Elmlinger, M., Lyko, F., Schmitz, O. J. and Wolf, E. 2006. Tissue-specific effects of in vitro fertilization procedures on genomic cytosine methylation levels in overgrown and normal sized bovine fetuses. Biol Reprod 75 (1): 17–23.
61. Hiendleder, S., Zakhartchenko, V. and Wolf, E. 2005. Mitochondria and the success of somatic cell nuclear transfer cloning: from nuclear-mitochondrial interactions to mitochondrial complementation and mitochondrial DNA recombination. Reprod Fertil Dev 17 (1–2): 69–83.
62. Hill, J. and Chavatte-Palmer, P. 2002. Pregnancy and neonatal care of cloned animals. In: Priniples of cloning. J. B. Cibelli, R. P. Lanza, K. Campbell and M. D. West. Academic Press, New york, 247–266.
63. Hill, J. R., Burghardt, R. C., Jones, K., Long, C. R., Looney, C. R., Shin, T., Spencer, T. E., Thompson, J. A., Winger, Q. A. and Westhusin, M. E. 2000. Evidence for placental abnormality as the major cause of mortality in first-trimester somatic cell cloned bovine fetuses. Biol Reprod 63 (6): 1787–94.
64. Hill, J. R., Winger, Q. A., Burghardt, R. C. and Westhusin, M. E. 2001. Bovine nuclear transfer embryo development using cells derived from a cloned fetus. Anim Reprod Sci 67 (1–2): 17–26.
65. Hoffert, K. A., Batchelder, C. A., Bertolini, M., Moyer, A. L., Famula, T. R., Anderson, D. L. and Anderson, G. B. 2005. Measures of maternal-fetal interaction in day-30 bovine pregnancies derived from nuclear transfer. Cloning Stem Cells 7 (4): 289–305.
66. Humpherys, D., Eggan, K., Akutsu, H., Friedman, A., Hochedlinger, K., Yanagimachi, R., Lander, E. S., Golub, T. R. and Jaenisch, R. 2002. Abnormal gene expression in cloned mice derived from embryonic stem cell and cumulus cell nuclei. Proc Natl Acad Sci U S A 99 (20): 12889–94.
67. Inoue, K., Kohda, T., Lee, J., Ogonuki, N., Mochida, K., Noguchi, Y., Tanemura, K., Kaneko-Ishino, T., Ishino, F. and Ogura, A. 2002. Faithful expression of imprinted genes in cloned mice. Science 295 (5553): 297.
68. Jablonka, E. and Lamb, M. J. 2002. The changing concept of epigenetics. Ann N Y Acad Sci 981: 82–96.
69. Jaenisch, R. and Wilmut, I. 2001. Developmental biology. Don't clone humans Science 291 (5513): 2552.
70. Jensen, R. G., Couch, S. C., Bitman, J., Carlson, S. E., Hamosh, M., Newburg, D. S. and Robert, G. J. 1995. Handbook of Milk Composition. Editor. Academic Press, San Diego, Pages.
71. Jeon, H. Y., Hyun, S. H., Lee, G. S., Kim, H. S., Kim, S., Jeong, Y. W., Kang, S. K., Lee, B. C., Han, J. Y., Ahn, C. and Hwang, W. S. 2005. The analysis of telomere length and telomerase activity in cloned pigs and cows. Mol Reprod Dev 71 (3): 315–20.
72. Jiang, L., Carter, D. B., Xu, J., Yang, X., Prather, R. S. and Tian, X. C. 2004. Telomere lengths in cloned transgenic pigs. Biol Reprod 70 (6): 1589–93.
73. Kang, Y.-K., Koo, D.-B., Park, J. S., Choi, Y.-H., Lee, K.-K. and Han, Y.-M. 2001b. Differential inheritance modes of DNA methylation between euchromatic and heterochromatic DNA sequences in ageing fetal bovine fibroblasts. FEBS Letters 498 (1): 1–5.
74. Kang, Y. K., Koo, D. B., Park, J. S., Choi, Y. H., Lee, K. K. and Han, Y. M. 2001a. Influence of oocyte nuclei on demethylation of donor genome in cloned bovine embryos. FEBS Lett 499 (1–2): 55–8.
75. Kasai, K., Sano, F., Miyashita, N., Watanabe, S. and Nagai, T. 2007. Comparison of the growth performances of offspring produced by a pair of cloned cattle and their nuclear donor animals. J Reprod Dev 53 (1): 135–42.
76. Kato, Y., Tani, T., Sotomaru, Y., Kurokawa, K., Kato, J., Doguchi, H., Yasue, H. and Tsunoda, Y. 1998. Eight calves cloned from somatic cells of a single adult. Science 282 (5396): 2095–8.
77. Kato, Y., Tani, T. and Tsunoda, Y. 2000. Cloning of calves from various somatic cell types of male and female adult, newborn and fetal cows. J Reprod Fertil 120 (2): 231–7.
78. Keefer, C. L., Keyston, R., Lazaris, A., Bhatia, B., Begin, I., Bilodeau, A. S., Zhou, F. J., Kafidi, N., Wang, B., Baldassarre, H. and Karatzas, C. N. 2002. Production of cloned goats after nuclear transfer using adult somatic cells. Biol Reprod 66 (1): 199–203.
79. Kishigami, S., Hikichi, T., Van Thuan, N., Ohta, H., Wakayama, S., Bui, H. T., Mizutani, E. and Wakayama, T. 2006. Normal specification of the extraembryonic lineage after somatic nuclear transfer. FEBS Lett 580 (7): 1801–6.
80. Kremenskoy, M., Kremenska, Y., Suzuki, M., Imai, K., Takahashi, S., Hashizume, K., Yagi, S. and Shiota, K. 2006. DNA methylation profiles of donor nuclei cells and tissues of cloned bovine fetuses. J Reprod Dev 52 (2): 259–66.
81. Kruip, T. A. M. and den Daas, J. H. G. 1997. In vitro produced and cloned embryos: Effects on pregnancy, parturition and offspring. Theriogenology 47 (1): 43–52.
82. Laible, G., Brophy, B., Knighton, D. and Wells, D. N. 2007. Compositional analysis of dairy products derived from clones and cloned transgenic cattle. Theriogenology 67 (1): 166–77.
83. Lanza, R. P., Cibelli, J. B., Blackwell, C., Cristofalo, V. J., Francis, M. K., Baerlocher, G. M., Mak, J., Schertzer, M., Chavez, E. A., Sawyer, N., Lansdorp, P. M. and West, M. D. 2000. Extension of cell life-span and telomere length in animals cloned from senescent somatic cells. Science 288 (5466): 665–9.
84. Lay, D. C., Randel, R. D., Friend, T. H., Carroll, J. A., Welsh, T. H., Jenkins, O. C., Neuendorff, D. A., Bushong, D. M. and Kapp, G. M. 1997. Effects of prenatal stress on the fetal calf. 14 (2): 73.
85. Le Neindre, P. 1989. Influence of cattle rearing conditions and breed on social behaviour and activity of cattle in novel environments. Applied Animal Behaviour Sciences 23: 129–140.
86. Lee, R. S., Peterson, A. J., Donnison, M. J., Ravelich, S., Ledgard, A. M., Li, N., Oliver, J. E., Miller, A. L., Tucker, F. C., Breier, B. and Wells, D. N. 2004. Cloned cattle fetuses with the same nuclear genetics are more variable than contemporary half-siblings resulting from artificial insemination and exhibit fetal and placental growth deregulation even in the first trimester. Biol Reprod 70 (1): 1–11.
87. Lee, S. Y., Park, J. Y., Choi, Y. J., Cho, S. K., Ahn, J. D., Kwon, D. N., Hwang, K. C., Kang, S. J., Paik, S. S., Seo, H. G., Lee, H. T. and Kim, J. H. 2007. Comparative proteomic analysis associated with term placental insufficiency in cloned pig. Proteomics 7 (8): 1303–1315.
88. Li, N., Wells, D. N., Peterson, A. J. and Lee, R. S. 2005. Perturbations in the biochemical composition of fetal fluids are apparent in surviving bovine somatic cell nuclear transfer pregnancies in the first half of gestation. Biol Reprod 73 (1): 139–48.
89. Liu, G., Kato, Y. and Tsunoda, Y. 2007. Aging of Recipient Oocytes Reduces the Development of Cloned Embryos Receiving Cumulus Cells. J Reprod Dev 53 (4): 785–90.
90. Lloyd-Thomas, A. R. and Fitzgerald, M. 1996. Do fetuses feel pain? Reflex responses do not necessarily signify pain. Bmj 313 (7060): 797–8.
91. Loi, P., Clinton, M., Vackova, I., Fulka, J., Jr., Feil, R., Palmieri, C., Della Salda, L. and Ptak, G. 2006. Placental abnormalities associated with post-natal mortality in sheep somatic cell clones. Theriogenology 65 (6): 1110–21.
92. Long, J. and Cai, X. 2007. Igf-2r expression regulated by epigenetic modification and the locus of gene imprinting disrupted in cloned cattle. GENE 388 (1–2): 125–134.
93. Lucifero, D., La Salle, S., Bourc'his, D., Martel, J., Bestor, T. H. and Trasler, J. M. 2007. Coordinate regulation of DNA methyltransferase expression during oogenesis. BMC Dev Biol 7: 36.
94. Martin, M., Adams, C. and Wiseman, B. 2004. Pre-weaning performance and health of pigs born to cloned (fetal cell derived) swine versus non-cloned swine. Theriogenology 62 (1–2): 113–22.
95. Martinez-Cerdeno, V., Noctor, S. C. and Kriegstein, A. R. 2006. Estradiol stimulates progenitor cell division in the ventricular and subventricular zones of the embryonic neocortex. Eur J Neurosci 24 (12): 3475–88.
96. Matsuzaki, M. and Shiga, K. 2002. Endocrine characteristics of cloned calves. Cloning Stem Cells 4 (3): 261–7.
97. McConnell, J. M. L. 2006. A mitochondrial component of developmental programming. In: Developmental origins of health and disease. P. D. Gluckman and M. A. Hanson. Cambridge University Press, Cambridge, 75–81.
98. Mellor, D. J. and Gregory, N. G. 2003. Responsiveness, behavioural arousal and awareness in fetal and newborn lambs: experimental, practical and therapeutic implications. N Z Vet J 51 (1): 2–13.
99. Mir, B., Zaunbrecher, G., Archer, G. S., Friend, T. H. and Piedrahita, J. A. 2005. Progeny of somatic cell nuclear transfer (SCNT) pig clones are phenotypically similar to non-cloned pigs. Cloning Stem Cells 7 (2): 119–25.
100. Norman, H. D., Lawlor, T. J., Wright, J. R. and Powell, R. L. 2004b. Performance of Holstein clones in the United States. J Dairy Sci 87 (3): 729–38.
101. Norman, H. D. and Walsh, M. K. 2004a. Performance of dairy cattle clones and evaluation of their milk composition. Cloning Stem Cells 6 (2): 157–64.
102. NZRBCS 2002. Enderby Island Cattle: A New Zealand Rare Breed Society Rescue Project.
103. Ogonuki, N., Inoue, K., Yamamoto, Y., Noguchi, Y., Tanemura, K., Suzuki, O., Nakayama, H., Doi, K., Ohtomo, Y., Satoh, M., Nishida, A. and Ogura, A. 2002. Early death of mice cloned from somatic cells. Nat Genet 30 (3): 253–4.
104. Ohgane, J., Wakayama, T., Kogo, Y., Senda, S., Hattori, N., Tanaka, S., Yanagimachi, R. and Shiota, K. 2001. DNA methylation variation in cloned mice. Genesis 30 (2): 45–50.
105. Onishi, A., Iwamoto, M., Akita, T., Mikawa, S., Takeda, K., Awata, T., Hanada, H. and Perry, A. C. 2000. Pig cloning by microinjection of fetal fibroblast nuclei. Science 289 (5482): 1188–90.
106. Ortegon, H., Betts, D., Lin, L., Coppola, G., Perrault, S., Blondin, P. and King, W. 2007. Genomic stability and physiological assessments of live offspring sired by a bull clone, Starbuck II. Theriogenology 67 (1): 116–126.
107. Palmquist, D. L., Beaulieu, A. D. and Barbano, D. M. 1993. Feed and animal factors influencing milk fat composition. J Dairy Sci 76 (6): 1753–71.
108. Panarace, M., Aguero, J. I., Garrote, M., Jauregui, G., Segovia, A., Cane, L., Gutierrez, J., Marfil, M., Rigali, F., Pugliese, M., Young, S., Lagioia, J., Garnil, C., Forte Pontes, J. E., Ereno Junio, J. C., Mower, S. and Medina, M. 2007. How healthy are clones and their progeny: 5 years of field experience. Theriogenology 67 (1): 142–51.
109. Park, M. R., Cho, S. K., Lee, S. Y., Choi, Y. J., Park, J. Y., Kwon, D. N., Son, W. J., Paik, S. S., Kim, T., Han, Y. M. and Kim, J. H. 2005. A rare and often unrecognized cerebromeningitis and hemodynamic disorder: a major cause of sudden death in somatic cell cloned piglets. Proteomics 5 (7): 1928–39.
110. Peaston, A. E. and Whitelaw, E. 2006. Epigenetics and phenotypic variation in mammals. Mamm Genome 17 (5): 365–74.
111. Petersen, B., Lucas-Hahn, A., Lemme, E., Hornen, N., Hassel, P., Kues, W. A. and Niemann, H. 2007. Preovulatory embryo transfer increases success of porcine somatic cloning. Reproduction, Fertility and Development 19: 155–156.
112. Petyim, S., Bage, R., Madej, A. and Larsson, B. 2007. Ovum pick-up in dairy heifers: does it affect animal well-being? Reprod Domest Anim 42 (6): 623–32.
113. Philpott, M. 1993. The dangers of disease transmission by artificial insemination and embryo transfer. Br Vet J 149 (4): 339–69.
114. Rassoulzadegan, M., Grandjean, V., Gounon, P. and Cuzin, F. 2007. Inheritance of an epigenetic change in the mouse: a new role for RNA. Biochem Soc Trans 35 (Pt 3): 623–5.
115. Rassoulzadegan, M., Grandjean, V., Gounon, P., Vincent, S., Gillot, I. and Cuzin, F. 2006. RNA-mediated non-mendelian inheritance of an epigenetic change in the mouse. Nature 441 (7092): 469–74.
116. Renard, J. P., Maruotti, J., Jouneau, A. and Vignon, X. 2007. Nuclear reprogramming and pluripotency of embryonic cells: Application to the isolation of embryonic stem cells in farm animals. Theriogenology 68, Suppl 1: S196–205.
117. Roemer, I., Reik, W., Dean, W. and Klose, J. 1997. Epigenetic inheritance in the mouse. Curr Biol 7 (4): 277–80.
118. Roselli, C. E., Stadelman, H., Reeve, R., Bishop, C. V. and Stormshak, F. 2007. The ovine sexually dimorphic nucleus of the medial preoptic area is organized prenatally by testosterone. Endocrinology 148 (9): 4450–7.
119. Roussel, S., Boissy, A., Montigny, D., Hemsworth, P. H. and Duvaux-Ponter, C. 2005. Gender-specific effects of prenatal stress on emotional reactivity and stress physiology of goat kids. Hormones and Behaviour 47 (3): 256–266.
120. Savage, A. F., Maull, J., Tian, X. C., Taneja, M., Katz, L., Darre, M. and Yang, X. 2003. Behavioral observations of adolescent Holstein heifers cloned from adult somatic cells. Theriogenology 60 (6): 1097–110.
121. Schaetzlein, S. and Rudolph, K. L. 2005. Telomere length regulation during cloning, embryogenesis and ageing. Reprod Fertil Dev 17 (1–2): 85–96.
122. Senda, S., Wakayama, T., Arai, Y., Yamazaki, Y., Ohgane, J., Tanaka, S., Hattori, N., Yanagimachi, R. and Shiota, K. 2007. DNA methylation errors in cloned mice disappear with advancement of aging. Cloning Stem Cells 9 (3): 293–302.
123. Senda, S., Wakayama, T., Yamazaki, Y., Ohgane, J., Hattori, N., Tanaka, S., Yanagimachi, R. and Shiota, K. 2004. Skewed X-inactivation in cloned mice. Biochem Biophys Res Commun 321 (1): 38–44.
124. Shibata, M., Otake, M., Tsuchiya, S., Chikyu, M., Horiuchi, A. and Kawarasaki, T. 2006. Reproductive and growth performance in Jin Hua pigs cloned from somatic cell nuclei and the meat quality of their offspring. J Reprod Dev 52 (5): 583–90.
125. Shiels, P. G., Kind, A. J., Campbell, K. H., Wilmut, I., Waddington, D., Colman, A. and Schnieke, A. E. 1999. Analysis of telomere length in Dolly, a sheep derived by nuclear transfer. Cloning 1 (2): 119–25.
126. Shiga, K., Umeki, H., Shimura, H., Fujita, T., Watanabe, S. and Nagai, T. 2005. Growth and fertility of bulls cloned from the somatic cells of an aged and infertile bull. Theriogenology 64 (2): 334–43.
127. Shiota, K. and Yanagimachi, R. 2002. Epigenetics by DNA methylation for development of normal and cloned animals. Differentiation 69 (4–5): 162–6.
128. Shoubridge, E. A. and Wai, T. 2007. Mitochondrial DNA and the mammalian oocyte. Curr Top Dev Biol 77: 87–111.
129. Shutler, D., Weary, D., McLellan, N. 2005. The clones need to return: A comment on Archer et al. (2003). Applied Animal Behaviour Science 91 (3–4): 363–365.
130. Sikich, L. and Todd, R. D. 1988. Are the neurodevelopmental effects of gonadal hormones related to sex differences in psychiatric illnesses? Psychiatr Dev 6 (4): 277–309.
131. Sinclair, K. D., McEvoy, T. G., Maxfield, E. K., Maltin, C. A., Young, L. E., Wilmut, I., Broadbent, P. J. and Robinson, J. J. 1999. Aberrant fetal growth and development after in vitro culture of sheep zygotes. J Reprod Fertil 116 (1): 177–86.
132. Slater, R., Cantarella, A., Gallella, S., Worley, A., Boyd, S., Meek, J. and Fitzgerald, M. 2006. Cortical pain responses in human infants. J Neurosci 26 (14): 3662–6.
133. Smith, R. D. 2005. Veterinary Clinical Epidemiology. Editor. CRC Press, Boca Raton, FL, Pages.
134. Smythe, J. W., McCormick, C. M., Rochford, J. and Meaney, M. J. 1994. The interaction between prenatal stress and neonatal handling on nociceptive response latencies in male and female rats. Physiology & Behavior 55 (5): 971–974.
135. Steinborn, R., Schinogl, P., Zakhartchenko, V., Achmann, R., Schernthaner, W., Stojkovic, M., Wolf, E., Muller, M. and Brem, G. 2000. Mitochondrial DNA heteroplasmy in cloned cattle produced by fetal and adult cell cloning. Nat Genet 25 (3): 255–7.
136. Stringfellow, D. A., Givens, M. D. and Waldrop, J. G. 2004. Biosecurity issues associated with current and emerging embryo technologies. Reprod Fertil Dev 16 (2): 93–102.
137. Suemizu, H., Aiba, K., Yoshikawa, T., Sharov, A. A., Shimozawa, N., Tamaoki, N. and Ko, M. S. 2003. Expression profiling of placentomegaly associated with nuclear transplantation of mouse ES cells. Dev Biol 253 (1): 36–53.
138. Sullivan, E. J., Kasinathan, S., Kasinathan, P., Robl, J. M. and Collas, P. 2004. Cloned calves from chromatin remodeled in vitro. Biol Reprod 70 (1): 146–53.
139. Takahashi, S. and Ito, Y. 2004. Evaluation of meat products from cloned cattle: biological and biochemical properties. Cloning Stem Cells 6 (2): 165–71.
140. Tamashiro, K. L., Sakai, R. R., Yamazaki, Y., Wakayama, T. and Yanagimachi, R. 2007. Developmental, behavioral, and physiological phenotype of cloned mice. Adv Exp Med Biol 591: 72–83.
141. Tamashiro, K. L., Wakayama, T., Akutsu, H., Yamazaki, Y., Lachey, J. L., Wortman, M. D., Seeley, R. J., D'Alessio, D. A., Woods, S. C., Yanagimachi, R. and Sakai, R. R. 2002. Cloned mice have an obese phenotype not transmitted to their offspring. Nat Med 8 (3): 262–7.
142. Tamashiro, K. L., Wakayama, T., Blanchard, R. J., Blanchard, D. C. and Yanagimachi, R. 2000. Postnatal growth and behavioral development of mice cloned from adult cumulus cells. Biol Reprod 63 (1): 328–34.
143. Tamashiro, K. L., Wakayama, T., Yamazaki, Y., Akutsu, H., Woods, S. C., Kondo, S., Yanagimachi, R. and Sakai, R. R. 2003. Phenotype of cloned mice: development, behavior, and physiology. Exp Biol Med (Maywood) 228 (10): 1193–200.
144. Tanaka, S., Miyazawa, K., Watanabe, K., Ohwada, S., Aso, H., Yonai, M., Saito, N. and Yamaguchi, T. 2006. Comparison of T cell subsets between somatic cloned and normal cow. AMERICAN JOURNAL OF REPRODUCTIVE IMMUNOLOGY 55 (1): 28–35.
145. Tanaka, S., Oda, M., Toyoshima, Y., Wakayama, T., Tanaka, M., Yoshida, N., Hattori, N., Ohgane, J., Yanagimachi, R. and Shiota, K. 2001. Placentomegaly in cloned mouse concepti caused by expansion of the spongiotrophoblast layer. Biol Reprod 65 (6): 1813–21.
146. Tecirlioglu, R. T. and Trounson, A. O. 2007. Embryonic stem cells in companion animals (horses, dogs and cats): present status and future prospects. Reprod Fertil Dev 19 (6): 740–7.
147. Tenhagen, B. A., Helmbold, A. and Heuwieser, W. 2007. Effect of various degrees of dystocia in dairy cattle on calf viability, milk production, fertility and culling. J Vet Med A Physiol Pathol Clin Med 54 (2): 98–102.
148. Tian, X. C., Kubota, C., Sakashita, K., Izaike, Y., Okano, R., Tabara, N., Curchoe, C., Jacob, L., Zhang, Y., Smith, S., Bormann, C., Xu, J., Sato, M., Andrew, S. and Yang, X. 2005. Meat and milk compositions of bovine clones. Proc Natl Acad Sci U S A 102 (18): 6261–6.
149. Tian, X. C., Xu, J. and Yang, X. 2000. Normal telomere lengths found in cloned cattle. Nat Genet 26 (3): 272–3.
150. Tome, D., Dubarry, M. and Fromentin, G. 2004. Nutritional value of milk and meat products derived from cloning. Cloning Stem Cells 6 (2): 172–7.
151. Van Laere, A. S., Nguyen, M., Braunschweig, M., Nezer, C., Collette, C., Moreau, L., Archibald, A. L., Haley, C. S., Buys, N., Tally, M., Andersson, G., Georges, M. and Andersson, L. 2003. A regulatory mutation in IGF2 causes a major QTL effect on muscle growth in the pig. Nature 425 (6960): 832–6.
152. Veissier, I., Gesmier, V., Le Neindre, P., Gautier, J. Y. and Bertrand, G. 1994. The effects of rearing in individual crates on subsequent social behaviour of veal calves. Applied Animal Behaviour Science 41 (3–4): 199–210.
153. Wakayama, T., Shinkai, Y., Tamashiro, K. L., Niida, H., Blanchard, D. C., Blanchard, R. J., Ogura, A., Tanemura, K., Tachibana, M., Perry, A. C., Colgan, D. F., Mombaerts, P. and Yanagimachi, R. 2000. Cloning of mice to six generations. Nature 407 (6802): 318–9.
154. Wakayama, T. and Yanagimachi, R. 1999. Cloning the laboratory mouse. Semin Cell Dev Biol 10 (3): 253–8.
155. Walker, S., Christenson, R., Ruiz, R., Reeves, D., Pratt, S., Arenivas, F., Williams, N., Bruner, B. and Polejaeva, I. 2007. Comparison of meat composition from offspring of cloned and conventionally produced boars. Theriogenology 67 (1): 178–184.
156. Walker, S. C., Shin, T., Zaunbrecher, G. M., Romano, J. E., Johnson, G. A., Bazer, F. W. and Piedrahita, J. A. 2002. A highly efficient method for porcine cloning by nuclear transfer using in vitro-matured oocytes. Cloning Stem Cells 4 (2): 105–12.
157. Walker, S. K., Hartwich, K.M., Seamark, R.F. 1996. The production of unusually large offspring following embryo manipulation: Concepts and challenges. Theriogenology, 45 (1): 111–120.
158. Walsh, M. K., Lucey, J. A., Govindasamy-Lucey, S., Pace, M. M. and Bishop, M. D. 2003. Comparison of milk produced by cows cloned by nuclear transfer with milk from non-cloned cows. Cloning Stem Cells 5 (3): 213–9.
159. Ward, I. L. and Weisz, J. 1980. Maternal stress alters plasma testosterone in fetal males. Science 207 (4428): 328–9.
160. Watanabe, S. and Nagai, T. 2008. Health status and productive performance of somatic cell cloned cattle and their offspring produced in Japan. Accepted for publication. The Journal of reproduction and development 54 (2):
161. Wells, D. N. 2005. Animal cloning: problems and prospects. Rev Sci Tech 24 (1): 251–64.
162. Wells, D. N., Forsyth, J. T., McMillan, V. and Oback, B. 2004. The health of somatic cell cloned cattle and their offspring. Cloning Stem Cells 6 (2): 101–10.
163. Wells, D. N., Laible, G., Tucker, F. C., Miller, A. L., Oliver, J. E., Xiang, T., Forsyth, J. T., Berg, M. C., Cockrem, K., L'Huillier, P. J., Tervit, H. R. and Oback, B. 2003. Coordination between donor cell type and cell cycle stage improves nuclear cloning efficiency in cattle. Theriogenology 59 (1): 45–59.
164. Wells, D. N., Misica, P. M. and Tervit, H. R. 1999. Production of cloned calves following nuclear transfer with cultured adult mural granulosa cells. Biol Reprod 60 (4): 996–1005.
165. WHO (World Health Organization) 1990. Principles for the toxicological assessment of pesticide residues in food / published under the joint sponsorship of the United Nations Environment Programme, the International Labour Organisation, and the World Health Organization. World Health Organistation, International Programme on Chemical Safety, Geneva. 1–117.
166. WHO/FAO (World Health Organisation/Food and Agricultural Organization) 2001. Evaluation of allergenicity of genetically modified foods. www.fao.org/docrep/007/y0820e/y0820e00.htm, last accessed 2008–01-23.
167. Williams, N. E., Walker, S. C., Reeves, D. E., Sherrer, E., Galvin, J. M., Polejaeva, I., Rampacek, G., Benyshek, L., Christenson, R. K., Graves, W. M. and Pratt, S. L. 2006. A comparison of reproductive characteristics of boars generated by somatic cell nuclear transfer to highly related conventionally produced boars. Cloning Stem Cells 8 (3): 130–9.
168. Wilmut, I., Beaujean, N., de Sousa, P. A., Dinnyes, A., King, T. J., Paterson, L. A., Wells, D. N. and Young, L. E. 2002. Somatic cell nuclear transfer. Nature 419 (6907): 583–6.
169. Wilmut, I., Schnieke, A. E., McWhir, J., Kind, A. J. and Campbell, K. H. 1997. Viable offspring derived from fetal and adult mammalian cells. Nature 385 (6619): 810–3.
170. Wolff, G. L., Kodell, R. L., Moore, S. R. and Cooney, C. A. 1998. Maternal epigenetics and methyl supplements affect agouti gene expression in Avy/a mice. Faseb J 12 (11): 949–57.
171. World Organisation for Animal Health (OIE) 2007. Terrestrial Animal Health Code. Appendix 3.3.2. In vitro fertilised bovine embryos/in vitro maturing oocytes.
172. Wrenzycki, C., Herrmann, D., Lucas-Hahn, A., Korsawe, K., Lemme, E. and Niemann, H. 2005. Messenger RNA expression patterns in bovine embryos derived from in vitro procedures and their implications for development. Reprod Fertil Dev 17 (1–2): 23–35.
173. Wrenzycki, C., Lucas-Hahn, A., Herrmann, D., Lemme, E., Korsawe, K. and Niemann, H. 2002. In vitro production and nuclear transfer affect dosage compensation of the X-linked gene transcripts G6PD, PGK, and Xist in preimplantation bovine embryos. Biol Reprod 66 (1): 127–34.
174. Yamaguchi, M., Ito, Y. and Takahashi, S. 2007. Fourteen-week feeding test of meat and milk derived from cloned cattle in the rat. Theriogenology 67 (1): 152–165.
175. Yamaguchi, M., Itoh M., Ito, Y. and Watatabe, S. 2008. A 12-month feedign study of reproduction/development in rats fed meat/milk powder supplemented diets derived from the progeny of cloned cattle produced by somatic cell nuclear transfer. J Reprod Dev, Advance Publication 1-44.
176. Yang, L., Chavatte-Palmer, P., Kubota, C., O'Neill, M., Hoagland, T., Renard, J. P., Taneja, M., Yang, X. and Tian, X. C. 2005. Expression of imprinted genes is aberrant in deceased newborn cloned calves and relatively normal in surviving adult clones. Mol Reprod Dev 71 (4): 431–8.
177. Yang, X., Smith, S. L., Tian, X. C., Lewin, H. A., Renard, J. P. and Wakayama, T. 2007a. Nuclear reprogramming of cloned embryos and its implications for therapeutic cloning. Nat Genet 39 (3): 295–302.
178. Yang, X., Tian, X. C., Kubota, C., Page, R., Xu, J., Cibelli, J. and Seidel, G., Jr. 2007b. Risk assessment of meat and milk from cloned animals. Nat Biotechnol 25 (1): 77–83.
179. Yonai, M., Kaneyama, K., Miyashita, N., Kobayashi, S., Goto, Y., Bettpu, T. and Nagai, T. 2005. Growth, reproduction, and lactation in somatic cell cloned cows with short telomeres. J Dairy Sci 88 (11): 4097–110.
180. Young, L. E. and Fairburn, H. R. 2000. Improving the safety of embryo technologies: possible role of genomic imprinting. Theriogenology 53 (2): 627–48.