Producción in vitro de embriones 313 PRODUCCIÓN IN VITRO DE EMBRIONES BOVINOS

Producción in vitro de embriones 313 PRODUCCIÓN IN VITRO DE EMBRIONES BOVINOS Gustavo A Palma Introducción La producción in vitro de embriones es el procedimiento para producir embriones en 3 etapas, la maduración in vitro (MIV) de los ovocitos, la fecundación in vitro (FIV) y el cultivo in vitro (CIV) de los posibles cigotos, Figura 1. El desarrollo in vitro de los ovocitos está íntimamente relacionado con la aptitud que adquieren durante su etapa folicular previa. Por ello, para una mayor comprensión de los eventos que ocurren in vitro y de los factores que los afectan fue desarrollado el capítulo de desarrollo folicular temprano (ver capítulo en este libro).

314 Palma Bibliografía Abe H and H Hoshi (2003) Evaluation of bovine embryos produced in high performance serum-free media J Reprod Dev 49, 193-202 Abe H, Yamashita S, Itoh T, Satoh T, H Hoshi (1999) Ultrastructure of bovine embryos developed from in vitromatured and -fertilized oocytes: comparative morphological evaluation of embryos cultured either in serum-free medium or in serum-supplemented medium Mol Reprod Dev 53, 325 335 Abbott AL and Y Ducibella (2001) Calcium and the control of mammalian cortical granule exocytosis Front Biosci 6, D792 D806. Abbott AL, Fissore RA and T Ducibella (1999) Incompetence of preovulatory mouse oocytes to undergo cortical granule exocytosis following induced calcium oscillations Dev Biol 207, 38 48 Abbott AL, Fissore RA and T Ducibella (2001) Identification of a translocation deficiency in cortical granule secretion in preovulatory mouse oocytes Biol Reprod 65, 1640 1647 Adams GP, Matteri RL, Kastelic JP, Ko JCH and OJ Ginther (1992) Association between surges of follicle-stimulating hormone and the emergence of follicular waves in heifers J Reprod Fertil 9, 177-188 Adashi E, CE Resnick, AJ D ercole, ME Svoboda, JJ van Vyk (1985) Insulin-like growth factors as interovarian regulators of granulosa cell growth and function Endocr Rev 6, 400-420 Adashi EY, CE Resnick, ER Hernandez, ME Svoboda and JJ Van Wyk (1988) Characteization and regulation of a specific cell membrane receptor for somatomedin C/insulin like growth factor I in culture rat granulosa cells Endocrinol 122, 194-201 Agarwal A, Saleh RA, MA Bedaiwy (2003) Role of reactive oxygen species in the pathophysiology of human reproduction Fertil Steril 79, 829-43 Aitken RJ and JS Clarkson (1988) Significance of reactive oxygen species and antioxidants in defining the efficacy of sperm preparation techniques J Androl 9, 367 376 Aitken RJ, Clarkson JS and S Fishel (1989) Generation of reactive oxygen species, lipid peroxidation, and human sperm function Biol Reprod 41, 183-197 Aitken RJ, Harkiss D and Buckingham DW (1993) Analysis of lipid peroxidation mechanisms in human spermatozoa Molecular Reproduction and Development 35 302 315 Aitken RJ, Irvine DS, FC Wu (1991) Prospective analysis of sperm-oocyte fusion and reactive oxygen species generation as criteria for the diagnosis of infertility Am J Obstet Gynecol 164, 542-551 Ali AA, Bilodeau JF and MA Sirard (2003) Antioxidant requirements for bovine oocytes varies during in vitro aturation,fertilization and development Theriogenology 59, 939-49 Alberio RH, Aller J, Crespo P, GA Palma (1998) Gestation with frozen thawed in vitro produced embryo from ovarioteomized argentine-holland cows Proceedings of the 14th Meeting of European Embryo Transfer Society, 116 Alberio R and G A Palma (1998) Development of bovine matured oocytes matured in a defined medium supplemented with a low concentration of r-hfsh Theriogenology 49,195 Alberio R, Olivier N, GA Palma (1997) Embryo production is not affected by replacement of a CO2 incubator by a simple sealed system during embryo development Proceedings of 13 th Scientific Meeting of European Embryo Transfer Association, 126 Alberio R y V Zakhartchenko (2001) Clonación de animales de interés zootécnico En: Biotecnología de la Reproducción, Gustavo A. Palma (Ed.), Ediciones INTA, Balcarce, Argentina, 353-394 Albertini DF, Combelles CMH, Benecchi E and MJ Carabtsos (2001) cellular basis for paracrine regulation of ovarian follicle development Reproducion 121, 647-653 Avery B y T Greve (1995) Impact of percoll on bovine spermatozoa for in vitro insemination Theriogenology 44, 871-878 Avery B, T Greve (1995) Apparent (ab) normaliies of in vitro produced bovine embryos Proceedings of the XXX International Symposium of Società Italina per il progreso della Zootecnia, Milan, 171-183 Andersen MM, J Kroll, AG Byskov and M Faber (1976) Protein composition in the fluid of individual and pooled bovine follicles J Reprod Fertil 48, 109-118 Armstrong DT (1993) Recent advances in superovulation of cattle Theriogenology 39, 7-24 Armstrong JD and AM Benoit (1996) Paracrine, and endocrine factors that mediate the influence of nutrition on reproduction in cattle and swine: an in vivo, IGF-I perspective J Anim Sci 74 (Suppl 3), 18-35

Producción in vitro de embriones 315 Armstrong DT, Helm P, Irvine B, Petersen BA, Stubbings RB, McLean D, Stevens GF, RF Seamark (1992) Pregnancies and live birth from in vitro fertilization of calf oocytes collected by laparoscopic follicular aspiration Theriogenology 38, 667-678 Armstrong DT, Irvine B, Earl CR, McLean D, Stevens G, RF Seamark (1994) Gonadotropin regimens for follicular aspiration and in vitro embryo production from calves oocytes Theriogenology 42, 1227 1236 Armstrong DT, Kotaras PJ and CR Earl (1997) Advances in production of embryos in vitro from juvenile and prepubertal oocytes from the calf and lamb Reprod Fertil Dev 9, 333-339 Austin CR (1951) Observations on the penetration of the sperm into the mammalian egg Aust J Sci Res 4, 581-596 Austin CR (1952) The capacitation of mammalian spezies Nature 170, 326-329 Austin CR (1961) The Mammalian Egg Ed: Charles C Thomas. Springfield, Illinois Ayoub TMA and AG Hunter (1990) Characteritics of a bovine oocyte maturation inhibitor on bovine follicular fluid Biol Peprod 42 (Suppl 1), 458 Azzazy HM, Mansour MM and RH Christenson (2005) Doping in the recombinant era: strategies and counterstrategies Clin Biochem 38, 959-965 Bae YS, Kang SW, Seo MS, Baines IC, Tekle E, Chock PB, SG Rhee (1997) Epidermal growth factor (EGF)-induced generation of hydrogen peroxide Role in EGF receptor-mediated tyrosine phosphorylation J Biol Chem 272, 217-221 Baker J, Liu J-P, Robertson EJ and A Efstratiadis (1993) Role of IGFs in embryonic and postnatal growth Cell 75, 73-82 Barnes FL and NL First (1991) Embryonic transcription in in vitro cultured bovine embryos Mol Reprod Dev 29, 117-123 Barnett DK and BD Bavister (1996) What is the relationship between the metabolism of preimplantation embryos and their development in vitro? Mol Reprod Devol, 43, 105-133 Batt PA, Gardiner DK, AWN Cameron (1991) Oxygen concentration and protein source affect the development of preimplantation goat embryos in vitro Reprod Fertil Dev 3: 601 607 Betteridge KJ, Smith C, Stubbings RB, Xu KP and King WA (1989) Potential genetic improvement of cattle by fertilization of fetal oocytes in vitro J of Reprod Fert Supp 38, 87 98 Barnes MA, Bierly ST, Halomo RD and DM Henricks (1980) Follicle stimulating hormone, luteinizing hormone and estradiol-17b response in GnRH treated Holstein heifers Biol Reprod 22, 459-465 Bavister BD and R Yanagimachi (1977) The effects of sperm extracts and energy sources on the motility and acrosome reaction of hamster spermatozoa in vitro Biol Reprod 16, 228-237 Bavister BD (1995) Culture of preimplantation embryo: facts and artifacts Hum Repod Update 1, 91-148 Bavister BD and McKiernan SH (1991) Different lots of bovine serum albumin stimulate or inhibit hamster embryo development in vitro Serono Symposium on Preimplantation Embryo Development 52, II-3 Beconi MT, Affranchino M, Schang LM, N Beorlegui (1991) Influence of antioxidants on SOD activity in bovine sperm Biochem Int 3, 545-553 Behboodi E, Anderson GB, BonDurant RH, Cargill SL, Kreuscher BR, Medrano JF and JD Murray (1995) Birth of large calves that developed from in vitro-derived bovine embryos Theriogenology 44, 227-232 Bedford JM (1970) The influence of oestrogen and progesteron on sperm capacitation in the reproductive tract of the female rabbit J Endocrinol 46, 191-197 Bedford JM (1998) Mammalian fertilization misread? Sperm penetration of the eutherian zona pellucida is unlikely to be a lytic Event Biol Reprod 59, 1275-1287 Bedford JM (2006) Why do penetrating sperm create an oblique path in the zona pellucida? Reproduction 131, 23-25 Bellin ME and RL Ax (1987) Chemical characteristics of follicular glycosaminoglicans Adv Exp Med Bio 219, 731-735 Bertolini M, Wallace CR and GB Anderson (2006) Expression profile and protein levels of placental products as indirect measures of placental function in in vitro-derived bovine pregnancies. Reproduction 131, 163-173 Bevers MM, Dieleman SJ, van den Hurk R and F Izadyar (1997) Regulation and modulation of oocyte maturation in the bovine Theriogenology 47, 13-22

316 Palma Bilodeau S, MA Fortier and MA Sirard (1993) Effect of Adenylate-Cyclase Stimulation on Meiotic resumption and Cyclic-AMP Content of Zona-Free and Cumulus-Enclosed Bovine Oocytes in vitro J Reprod and Fert 97, 5-11 Bize I, Santander G, Cabello P, Driscoll D and C Sharpe (1991) Hydrogen peroxide is involved in hamster sperm capacitation in vitro Biol Reprod 44, 398-403 Blanco A (1993) Acido hialurónico En: Química biológica, Editorial Ateneo, Buenos Aires, 80-81 Blondin P and M A Sirard (1995) Oocyte and follicular morphology as determining characteristics for developmental compertence in bovine oocytes Mol Reprod Dev 41, 54-62 Boehm KD, M Daimon, IG Gorodeski, LA Sheean, WH Utian, J Ilan (1990) Expression of the insulinlike and platelet-derived growth factor genes in human uterine tissues Mol Reprod Dev 27, 93-101 Boland MP, Crosby TF and I Gordon (1989) Norphologycal normality of cattle embryos following superovulation using PMSG Theriogenology 10, 175-180 Boland NI, Humperson PG, Leese HJ and RG Gosden (1993) Pattern of lactate production and steroidegenesis during growth and maturation of mouse ovarian follcicles in vitro Biol Reprod 48, 798-806 Bols P, Vandenheende JMM, Van Soom A and A Kruif (1995) Transvaginal ovum pick-up (OPU) in the cow: a new disponsable needle guidance system Theriogenology 43, 677-687 Bols PEJ, Van Soom A, Ysebaert MT, Vandenheede JMM and A Kruif (1996) Effects of aspiration vacuum and needle diameter on cumulus oocyte complex morphology and developmental capacity of bovine oocytes Theriogenology 45, 1001-1014 Bols PEJ, Ysebaert MT, Van Soom A and A Kruif (1997) Effects of needle tip bevel and aspiration procedure on the morphology and developmental capacity of bovine compact cumulus oocytes complexes Theriogenology 47,1221-1236 Bonavera JJ, Tortonese DJ, Doray JM and F Petraglia (1988) Endocrine response to superovulatory treatment in pre-pubertal beef heifers: its lack of ability to induce puberty Anim Prod 46, 163-167 Boone WR and SS Shapiro (1990) Quality control in the in vitro fertilization laboratory Theriogenology 33, 1-3 Bousquet D, Twagiramungu H, Morin N, Brisson C, Carboneau G and J Durocher (1999) In vitro embryo production in teh cow: A affective alternative to the conventional embryo production approach Theriogenology 51, 59-70 Boyers SP, Davis RO and DF Katz (1989) Automated semen analysis Curr Probl Obstet Gynecol Fertil I, 167 200 Brantmeier SA, RR Grummer and RL Ax (1987) Concentration of high density lipoproteins among follicular sizes in the bovine J Dairey Sci 70, 2145-2149 Brem G y GA Palma (2001) Aplicaciones de la producción in vitro de embriones En: Palma (Ed) Biotecnología de la Reproducción, Ediciones INTA, Balcarce, Argentina, 489-495 Brenneisen P, Wenk J, Klotz LO, Wlaschek M, Briviba K, Krieg T, Sies H, Scharffetter-Kochanek K, (1998) Central role of ferrous/ferric iron in the ultraviolet B irradiation- mediated signaling pathway leading to increased interstitial collagenase (matrix-degrading metalloprotease (MMP)-1) and stromelysin-1 (MMP-3) mrna levels in cultured human dermal fibroblasts J Biol Chem 273, 5279 5287 Bringstock DR, RP Heap, KD Brown (1989) Poliypetide growth factors in uterine tissues and secretions J Reprd Fert 85, 747-758 Brogliatti GM and GP Adams (1997) Ultrasound-guided transvaginal oocyte collection in prepubertal caves. Theriogenology 45, 1163-1176 Brum DS, Figueiró GM, Leivas FG, Bernardi ML, Silva CAM, MIB Rubin (1999) Transport-culture of in vitro produced bovine embryos for 24h Arq Fac Vet UFRGS 27, 208 Buccione R, Schroeder AC, JJ Eppig (1990) Interactions between somatic cells and germ cells throughout mammalian oogenesis Biol Reprod 43, 543-547 Burgess AW and D Metcalf (1980) The nature and action of granulocyte-macrophage colony stimulating factors, Blood 56, 947 958 Busa WB and R Nuccitelli (1984) Metabolic regulation via intracellular ph Am J Physiol 246, 409-438 Byskov AGS (1968) Cell kynetic studies of atresia in the mouse ovary J Reprod Fertil 37, 277-285 Bevers MM, SJ Dieleman, R van den Hurk and F Izadyar (1997) regulation and modulation of oocyte maturation in the bovine Theriogenology 47, 13-22 Bevers MM and JF Beckes (1993) Development of a combined new mechanical and enzimatic method for the isolation of intact preantral follcicles from fetal, calf and adult bovine ovaries Theriogenology 40, 789-799

Producción in vitro de embriones 317 Block J, Drost M, Monson RL, Rutledge JJ, Rivera RM, Paula-Lopes FF, Ocon OM, Krininger CE III, Liu J and PJ Hansen (2003) Use of insulin-like growth factor-1 during embryo culture and treatment of recipients with gonadotropin-releasing hormone to increase pregnancy rates following the transfer of in vitro-produced embryos to heat-stressed, lactating cows J Anim Sci 81, 1590-1602 Brackett BG and JB Server (1970) Capacitation of rabbit spermatozoa in the uterus Fert Steril 21, 687-695 Byrne AT, Southgate J, Brison DR, HJ Leese (2002) Regulation of apoptosis in the bovine blastocyst by insulin and the insulin-like growth factor (IGF) superfamily. Mol Reprod Dev 62, 489-495 Caamano JN, Ryoo ZY, CR Youngs (1998) Promotion of development of bovine embryos produced in vitro by addition of cysteine and beta-mercaptoethanol to a chemically defined culture system J Dairy Sci 81, 369-374 Cain L, Chatterjee S and TJ Collings (1995) In vitro folliculogenesis of rat preantral follicles Encrinology 136, 3369-3377G Cameo M and JA Blaquier (1976) Androgen-controlled specific proteins in rat epididymis J Endocrinol 69, 47 55 Carrol J, Whittingham DG, Wood MJ, Telfer E, RG Gosden (1989) Extra-ovarian production of mature viable mouse oocytes from frozen primary follicles J Reprod Fertil 90, 321-328 Carson RS, Z Zhang, LA Hutchinson, AC Herington, JK Findlay (1989) Growth factors in ovarian function J Reprod Fert 85, 735-746 Chang MC (1951) Fertilizing capacity of spermatozoa deposited in Fallopian tubes Nature 168, 997-998 Chang R (1992) Mantenimiento del ph en la sangre En: Química, McGraw-Hill (ed), 699-701 Chegini N, MJ Rossi, BJ Masterson (1992) Platelet-derived growth factor (PDGF), epidermal growth factor (EGF), and EGF and PDGF ß-receptors in human endometrial tissue: localization and in vitro action Endocrinol 130, 2373-2385 Chohan KR and AG Hunter (2003) Effect of reproductive status on in vitro developmental competence of bovine oocytes J Vet Sci 4, 67-72 Chohan KR, Hunter AG (2004) In vitro maturation, fertilization and early cleavage rates of bovine fetal oocytes Theriogenology 61, 373-380 Chow CK (1991) Vitamin E and oxidative stress Free Radic Biol Med 11, 215-232 Cittadini E, Albano C, Cimino C, R Palermo (1992) Assisted reproductive technologies in severe male infertility Acta Eur Fertil 23, 123-30 Corcoran D, Fair T, Park S, Rizos D, Patel OV, Smith GW, Coussens PM, Ireland JJ, Boland MP, Evans AC and P Lonergan (2006) Suppressed expression of genes involved in transcription and translation in in vitro compared with in vivo cultured bovine embryos Reproduction 131, 651 66 Coonrod SA, Naaby-Hansen S, Shetty J, Shibahara H, Chen M, White JM, JC Herr (1999) Treatment of mouse oocytes with PI-PLC releases 70-kDa (pi 5) and 35- to 45-kDa (pi 5.5) protein clusters from the egg surface and inhibits sperm-oolemma binding and fusion Dev Biol 207, 334-349 Cortvindt R RG, Hu Y, Liu J, JE Smitz (1998a) Timed analysis of the nuclear maturation of oocytes in early preantral mouse follicle culture supplemented with recombinatn gonadotropin Fertil Steril 70,1114-25 Cortvindt R RG, Hu Y, JE Smitz (1998b Recombinant luteinizing hormone as a survival and differentiation factor increase oocyte maturation in recombinant follicle stimulating hormone suplemented mouse preantral follicle culture Hum Reprod 13,1292-1302 Coskun SA, A Sanbuissho, YC Lin, Y Rikihisa (1991) Fertilizability and subsequent developmental ability of bovine oocytes matured in medium containing epidermal growth factor (FGF) Theriogenology 36, 485-494 Coticchio G, Sereni E, Serrao L, Mazzone S, Iadarola I and A Borini (2004) What criteria for the definition of oocyte quality? Ann N Y Acad Sci 1034, 132-144 Crosier AA, Farin PW, Dykstra MJ, Alexander JE and Ch Farin (2000) Ultrastructural morphometry of bovine compact murulae produced in vivo or in vitro Biol Reprod 62, 1459-1465 Cox LJ, Larman MG, Saunders CM, Hashimoto K, Swann K, FA Lai (2002) Sperm phospholipase C from humans and cynomolgus monkeys triggers Ca2+ oscillations, activation and development of mouse oocytes. Reproduction 124, 611 623. Dalvit G, Llanes SP, Descalzo A, Insani M, Beconi M and P Cetica (2005) Effect of alpha-tocopherol and ascorbic acid on bovine oocyte in vitro maturation Reprod Dom Anim 40, 93-97

318 Palma Damiani P, Fissore RA and RT Duby (1998) Biochemical and physiological maturation of cow and calf oocytes Theriogenology 49, 310 Daniel SA, Armstrong DT, Wood MJ, Telfer E, RE Gore-Langton (1989) Growth and development of rat oocyte in vitro Gam Res 24,109-121 Davis BK (1981) Timing of fertilization In: Knobil E and J Neill (eds), the Physiology of Reproduction, Raven Press, New York 189-317 Davis RO and RJ Siemers (1995) Derivation and reliability of kinematic measures of sperm motion In: Seventh International Symposium on Spermatology Plenary Papers Reproduction, Fertility and Development 7, 857-869 Dawson KM and Baltz (1997) Organic osmolytes and embryos: substrates of the Gly and ß transport systems protect mouse zygotes against the effects of raised osmolarity Biol Reprod 56, 1550 1558 De Loos FAM, M Jeunken, E Zeinstra and MM Bevers (1992) Structural aspects of bovine maturation in vitro Mol Reprod Dev 31, 208-214 De Loos FAM, E Zeinstra and MM Bevers (1994) Follicular wall maintain meiotic arrest in bovine oocytes cultured in vitro Mol Reprod Dev 39, 162-165 Dodson SE, Mcleod BJ, W Haresign, Peters AR and GE Lamming (1988) Endocrine changes from birth to puberty in the heifer J Reprod Fertil 82, 527-538 Domes U (2003) Untersuchungen über die Bestimmung der Spermaqualität - insbesondere Motilität und Membranintegrität - und Zusammenhänge zur Fertilität von Besamungshengsten Tierärztlichen Fakultät der Ludwig-Maximilians-Universität München Diss, 1-155 Dominiko and First (1991) Maturation of denuded bovine oocytes result in normal fertilization and development Biol Reprod 44 (Supl 1) 353 Donnay I, Faerge I, Grondahl C, Verhaeghe B, Sayoud H, Ponderato N, C Galli and G Lazzari (2004) Effect of prematuration, meiosis activating sterol and enriched maturation medium on the nuclear maturation and competence to development of calf oocytes Theriogenology 62, 1093-1107 Donahue R (1968) Maturation of mouse oocyte in vitro Sequence and timing of nuclear progression J Exp Zool 169, 237-250 Dong YJ, Varisanga MD, Mtango NR, Aono M, Otoi T and T Suzuki (2001) Improvement of the culture conditions for in vitro production of cattle embryos in a portable CO 2 incubator Reprod Domest Anim 36, 313-318 Dostal J and A Pavlok (1996) Isolation and characterization of maturion inhibiting compuond in bovine follicular fluid Reprod Nutr Dev 36, 681-690 Downs SM (1993) Factors affecting the resumption of meiotica maturation in mammalian oocytes Theriogenology 19, 65-79 Driancourt MA (1991) Follicular dynamics in sheep and cattle Theriogenology 35, 55-79 Driancourt MA, Reynaud K, J Smitz (2001) Differences in follicular function of 3-month-old calves and mature cows Reproduction 121, 463-74 Du F, Looney CR and X Yang (1996) Evaluation of bovine embryos produced in vitro vs In vivo by differential staining of inner cell mass and trophectoderm cells Theriogenology 45, 211 Ducibella T, Huneau D, Agelichio E, Xu Z, Schultz RM, Kopf Fissore R, Madoux S and JP Ozil (2002) Egg-to-embryo transition is driven by differential responses to Ca2+ oscillation number Dev Biol 250, 280-291 Ducibella T, Huneau D, Angelichio E, Xu Z, Schultz RM, Kopf GS, Fissore R, Madoux S, Ozil JP (2002) Egg-to-embryo transition is driven by differential responses to Ca(2+) oscillation number Dev Biol 250, 280-291 Ducibella T, Schultz RM, JP Ozil (2006) Role of calcium signals in early development Semin Cell Dev Biol 17, 324-332 Dumollard R, Duchen M, C Sardet (2006) Calcium signals and mitochondria at fertilisation Semin Cell Dev Biol 17, 314-23 Dumollard R, Marangos P, Fitzharris G, Swann K, Duchen M, J Carroll (2004) Sperm-triggered [Ca2+] oscillations and Ca 2+ homeostasis in the mouse egg have an absolute requirement for mitochondrial ATP production Development 131, 3057 3067 Drahorad J, tesarik J, Cechova D, V Vilim (1991) Proteins and glycosaminoglycans in the intercellular matrix of human cumulus oophorus and their effect on conversión of proacrosin to acrosin J Reprod Fertil 93, 253-262 Dröge W (2002) Free radicals in the physiological control of cell function Physiol Rev 82, 47-95

Producción in vitro de embriones 319 Earl CR, Fry RC, Maclellan LJ, Kelly JM and DT Armstrong (1998) In vitro fertilization and developmental potential of prepubertal calves oocytes In: ametes: Development and function (Luria A, Ganofi F, Enne G and I Giannaroli, Eds) Rome: Serono Symposia, pp 115-137 Echternkamp SE, HJ Howard, AJ Roberts, J Grizzle and T Wise (1994) Relationsships among concentrations of steroids, Insulin-like Growth Factor-I, and Insulin-like factor binding proteins in ovarian follicular fluid of beeff cattle Biol Reprod 51, 971-981 Emond V, MacLaren LA, Kimmins S, Arosh JA, Fortier MA, RD Lambert (2004) Expression of cyclooxygenase-2 and granulocyte-macrophage colonystimulating factor in the endometrial epithelium of the cow is up-regulated during early pregnancy and in response to intrauterine infusions of interferon-τ Biol Reprod 70, 54 64 Eliyahu E, Tsaadon A, Shtraizent N and R Shalgi (2005) The involvement of protein kinase C and actin filaments in cortical granules exocytosis in the rat Reproduction 129, 161 170 Eppig JJ (1979) A comparision between oocyte growth in coculture with granulosa cell-oocytes junctional contact maintained in vitro J Exp Zool 209, 345 Eppig JJ and O Brien MJ (1996) Development in vitro of mouse oocytes from primordial follicles Biol Reprod 54,197-207 Eppig JJ, AC Schroeder (1989) Capacity of mouse oocytes from preantral follcicles to undergo embryogenesis and development to live young after growth, maturation and fertilization in vitro Biol Reprod 41, 268-276 Eppig JJ, Wigglesworth K, Pendola F, Y Hirao (1997) Murine oocytes suppress expression of luteinizing hormone receptor messenger ribonucleic acid by granulosa cells Biol Reprod 56, 976 984 Ehrenwald E, RH Foote and JE Parks (1990) Bovine oviductal fluid components and their potential role in sperm cholesterol efflux Mol Reprod Dev 25, 195-204 Ehrenwald E, JE Parks and RH Foote (1988a) Cholesterol efflux from bovine sperm: I Induction of the acrosome reaction with lysophosphatidilcholine after reducing sperm cholesterol Gamete Res 20, 145-157 Ehrenwald E, JE Parks and RH Foote (1988b) Cholesterol efflux from bovine sperm: II Effect of reducing sperm cholesterol on penetration of zone-free hamster and in vitro matured bovine ova Gamete Res 20, 413-420 Evans JP and GS Kopf GS (1998) Molecular mechanisms of sperm-egg interactions and egg activation Andrologia 30, 297 307 Fabian D, Il kova G, Rehak P, Czikkova S, Baran V, J Koppel (2004) Inhibitory effect of IGF-I on induced apoptosis in mouse preimplantation embryos cultured in vitro Theriogenology 61, 745 755 Fair T, P Hyttel and T Greve (1995) Bovine oocyte diameter in relation to maturational comtetence and transcriptional activity Mol Reprod Dev 42, 437-442 Farin PW, Britt JH, Shaw DW, BD Slenning (1995) Agreement among evaluators of bovine embryos produced in vivo or in vitro Theriogenology 44, 339-350 Farin PW, Crosier AE and CE Farin (2001) Influence of in vitro systems on embryo survival and fetal development in cattle Theriogenology 55, 151 170 Farin PW and CE Farin (1995) Transfere of bovine embryos produced in vivo or in vitro: survial and fetal development Bio Reprod, 52, 676-682 Farin PW, Farin CE, Crosier AE, Blondin P, JE Alexander (1999) Effects of in vitro culture and maternal insulin-like growth factors-1 on development of bovine conceptuses Theriogenology 51, 238 Farin PW, Piedrahita JA and CE Farin (2006) Errors in development of fetuses and placentas from in vitro-produced bovine embryos Theriogenology 65, 178-191 Ferry LP, P Mermillod, A Massip and F Dessy (1994) Bovine embryos cultured in serum-poor oviduct-conditioned medium need cooperation to reach the blastocyst stage Theriogenology 42, 445-453 Figueiredo JR, Hulshof SCJ, van den Hurk R, Ectors FJ, Fontes RS, Nusgens B, Bevers MM, JF Beckers (1993) Development of a combined new mechanical and enzimatic method for the isolation of intact preantral follcicles from fetal, calf adn adult bovine ovaries Theriogenology 40, 789-799 Figueiredo JR, Hulshof SCJ, Thirty M, Nusgens, B van den Hurk R, Ectors FJ, Bevers MM and JF Beckers (1994) Sources of bovine ovaries affect significantly the development of isolated preantral follicle in vitro Proceedings 10 th Meeting of European Embryo Transfer Association, 172

320 Palma Fleming TP, Kwong WY, Porter R, Ursell E, Fesenko I, Wilkins A, Miller DJ, Watkins AJ; JJ Eckert (2004) The embryo and its future Biol Reprod 71, 1046-1054 Fouladi Nashta AA, Waddington D, KHS Campbell (1998) Maintenence of bovine oocytes in meiotic arrest and subsequent development in vitro: A comparative evaluation of antral follicle culture with other methods Biol Reprod 59, 255-262 Fraser LR (1992) Requirements for successful mammalian sperm capacitation and fertilization Arch Pathol Lab Med, 116, 345-50 Fraser LR, Adeoya-Osiguwa SA, Baxendale RW, and R Gibbons (2006) Regulation of mammalian sperm capacitation by endogenous molecules Front Biosci 11, 1636-1645 Fraser LR and CA McDermott (1992) Ca(2+)-related changes in the mouse sperm capacitation state: a possible role for Ca(2+)-ATPase J Reprod Fertil 9, 363-377 Freshney I (1987) The culture environment, In: Culture of animal cells, A manual of basic technique AR Liss, New York, 57-84 Fujita T, Umeki H, Shimura H, Kugumiya K and K Shiga (2006) Effect of group culture and embryoculture conditioned medium on development of bovine embryos J Reprod Dev 52, 137-142 Fukui Y, H Ono (1989) Effects of sera hormones and granulosa cells added to de culture medium for in vitro maturation, fertilization, cleavage and development of bovine oocytes J Reprod Fertil 86, 501-506 Fukui Y, T Sonoyama, H Michizuki and H Ono (1990) Effects of heparin dosage and capacitacion time on in vitro fertilization and cleavage of bovine oocytes matured in vitro Theriogenology 34, 579-591 Galli C, Duchi R, Crotti G, Turini P, Ponderato N, Colleoni S, Lagutina I and G Lazzari (2004) Production and quality of bovine oocytes and embryos Vet Res Commun 28, 121-126 Gandolfi F, Milanesi E, Pocar P, Luciano AM, Brevini TAL, Acocella F, Lauria A and DT Armstrong (1998) Comparative analysis of calf and cow oocytes during in vitro maturation Mol Reprod Dev 49, 168 175 Gandolfi F, Pocar P, Milanesi E, Brevini TAL, Luciano, AM DT and Armstrong, (1996) Comparative analysis of metabolic activity and protein synthesis during IVM of calves and cows oocytes Biol. Reprod 54 (Suppl 1), 157 Gardner AJ and JP Evans (2006) Mammalian membrane block to polyspermy: new insights into how mammalian eggs prevent fertilisation by multiple sperm Reprod Fert Devel 18 53 61 Gardner DK and M Lane (1993) Amino acids and ammonium regulate mouse embryo development in culture Biol Reprod 48, 377 385 Gardner DK and M Lane (1996) Alleviation of the "2-cell block" and development to the blastocyst of CF1 mouse embryos: role of amino acids, EDTA and physical parameters Hum Reprod 11, 2703 2712 Gardner DK and M Lane (1998) Culture of viable human blastocysts in defined sequential serum-free media Hum Reprod 13 Suppl 3, 148-159 Gardner DK, Lane M and P Blatt (1993) Uptake and metabolism of pyruvate and glucosa by individual sheep preattachment embryos developed in vivo Mol Repro Dev 36, 313-319 Gardner DK and Lane (2003) Towards a single embryo transfer Reprod Biomed Online 6, 470-481 Gardner DK, Lane M, Spitzer A and PA Batt (1994) Enhanced rates of cleavage and development in the absence of serum and somatic cells: amino acids, vitamins and culturing embryos un groups stimulate development Biol Rreprod 50, 390-400 Georges M and JM Massey (1991) Velogenetics, or the use of marker assisted selection and germ line manipulation Theriogenology 35, 151 159 Geva E, Lessino JB, Lerner-Geva L and A Amit (1998) Free radicals, antioxidants and human spermatozoa: clinical implications Hum Reprod 13, 1422 1424 Gilchrist RB, Ritter LJ and DT Armstrong (2004a) Oocyte-somatic cell interactions during follicle development in mammals Anim Reprod Sci. 82/83, 431-446 Gonçalves PBD, AG Hernandez, CM Schweizer, SF Carambula (1996) Aspects of oocyte maturation and in vitro fertilization in the bovine Arq Fac Vet UFRGS, 24, 98-116 Good NE (1966) Hydrogen in buffers and biological research Biochemistry 5, 467-477 Goodman Gilman A, LS Goodman, TW Rall F Murad (1986) Heparina En: Las bases farmacológicas de la terapéutica, Editorial Panamericana, Buenos Aires, 1273-1277 Gordon I (1994) Laboratory production of cattle embryos In: CAB International (Ed), Biotechnology in Agriculture No 11, 640 pp

Producción in vitro de embriones 321 Gordon I (2003) Laboratory production of cattle embryos In: CAB International Biotechnology in agriculture series, no 27 2nd edition United Kingdom 645 pp Gosden RG (1998) Biology and technology of primordial follicle development In: Lauria A, Gandolfi F, Enne G, L Gianaroli (eds) Gametes developmental and function, Serono Symposia, 85-98 Gosden RG and E Telfer (1987) Number of follicles in mammalian ovaries and the allometric relations J Zool Lond 211,169-175 Goto K, Y Takuma, N Ooe and K Ogawa In vitro developmetn of bovine oocytes collected from ovaries of individual cows after in vitro fertilization Jpn J Anim Reprod 36, 110-113 Grazulbilska AT, Bilski JJ, Jablonkashariff A, Reynolds LP and DA Redmer (1997) Immunhystomchemical localization of the gap junctional proteins Connexin (Cx)-26 and Cx32 in sheep ovaries Biol Reprod 56, 124 Grazulbilska AT, Redmer DA, Bilski JJ, Jablonkashariff A, Doraiswamy V and LP Reynolds (1998) Gap junctional proteins, connexin-26, connexin-32, and connexin-43 in sheep ovaries throughout the estrous-cycle Endocrine 8, 269-279 Gressier B, Cabanis A, Lebeque S, Brunet C, Dine T, Luyckx M, Cazin M, JC Cazin (1994) Decrease of hypochlorous acid and hydroxyl radical generated by stimulating human neutrophils, comparison in vitro of some thiol-containing drugs Methods Exp Clin Pharmacol 16, 9-13 Guille JJP and H Joenje (1991) Biological significance of oxygen toxicity, an introduction In: Vigo- Pelfrey C (ed) Membrane Lipid Oxidation CRC Press Inc, Boca Raton, FL, 1-32 Greve T, Avery B, H Callesen (1993) Viability of in vivo and in vitro produced embryos Rep Dom Anim 28, 164-169 Greenwald GS and KR Shyamal (1994) Follicular development and its control In: Knobil E, Neill JD, Greenwald GS, Markert CL, DW Pfaff (eds) Raven Press, New York, 629-724 Grippo AA, Henault MA, Anderson SH and GJ Killian (1992) Cation concentrations in fluid from the oviduct ampulla and isthmus of cows during the estrous cycle J Dairy Sci, 75, 58-65 Grupen CG, Fung M and DT Armstrong (2006) Effects of milrinone and butyrolactone-i on porcine oocyte meiotic progression and Guerin P, El Mouatassim S and Y Menezo (2001) Oxidative stress and protection against reactive oxygen species in the pre-implantation embryo and its surroundings Hum Reprod Update 7, 175-189 Guyader-Joly C, Durand M, Morel A, Ponchon S, Marquant-Le Guienne B, Guérin B and P Humblot (2000) Sources of variation in blastocyst production in a commercial ovum pickup in vitro embryo production program in dairy cattle Theriogenology 53, 355 Haber F and Y Weiss (1935) The Catalityc decomposition of hydrogen peroxide by ion salts J Proc Roy Soc 147, 332 351 Haddad T and D Yee (2006) Targeting the insulin-like growth factor axis as a cancer therapy Future Oncol 2, 101-110 Hagemann A, AH Nielsen, B Avery, B Svenstrup, J Nielsen and K Poulsen (1997) Relationship between follicular fluid steroids and tissue renin concentrations and secretions rates in bovine ovaries Exp Clin Endocrinol Diabetes 105, 271-276 Hamatani T, Carter MG, Sharov AA and MDH Ko (2004) Dynamics of global gene expression changes during mouse preimplantation development Dev Cell 6, 117 131 Hamilton JA and GP Anderson (2004) GM-CSF Biology Growth Factors 22, 225-231 Haliwell B (1994) Free radicals, antioxidants, and human disease: curiosity, cause, or consequence? Lancet 344, 721 724 Hancock JT, Desikan R, Neill SJ Role of reactive oxygen species in cell signalling pathways Biochem Soc Trans 200129, 345-350 Hanrahan JP, Gregan SM, Mulsant P, Mullen M, Davis GH, Powell R and SM Galloway (2004) Mutations in the genes for oocyte-derived growth factors GDF9 and BMP15 are associated with both increased ovulation rate and sterility in Cambridge and Belclare sheep (Ovis aries) Biol Reprod 70, 900 909 Harper KM and P Kaye (1992) Enhanced bovine oocyte quality after in vitro maturation (IVM) with insulin-like growth factor-i (IGF-I) and gonadotropins Biol Reprod Dev Suppl 1, 67 Harper KM, BG Brackett (1993) Bovine development after in vitro maturation in a defined medium with epidermal growth factor and low concentrations of gonadotrophins Biol Reprod 48, 409-416 Harrington, WN and GC Godman (1980) A slective inhibitor of cell proliferation from normal serum Proc Natl Acad Sci USA 77, 423-427

322 Palma Hartshorn GM, Sargent IL and DH Barlow (1994) Meiotic progression of mouse oocytes throughout follicle growth and ovulation in vitro Hum Reprod 9, 352-359 Hartwig A (2000) Metallionen zwischen Essenzialität und Toxizität Bichemie 34, 224 231 Hartshorne GM (1997) In vitro culture of ovarian follicles Reviews of Reprod 2, 94-104 Harvey AJ, Kind KL and JG Thompson (2002) Redox regulation of early embryo development Reproduction 123, 479-486 Hasler JF (1994) Commercial applications of in vitro fertilization in catttle Food Animal 16, 1062-1073 Hasler JF (1996) Commercial production of in vitro-derived bovine embryos Arq Fac Vet, UFRGS 24, 117-134 Hasler JF (2000) In-vitro production of cattle embryos: problems with pregnancies and parturition Hum Reprod 15, 47-58 HaslerJF, Henderson WB, Hurtgen PJ, Jin ZQ, McCauley AD, Mower SA, Neely B, Shuey LS, Stokes JE and SA Trimer (1995) Production, freezing and transfer of bovine IVF ambryos and subsequent calving results Theriogenology 43, 141-152 Hazeleger NL, Hill DJ, Stubbings RB and JS Watson (1995) Relationship of morphology and follicular fluid environment of bovine oocytes to their developmental potential in vitro Theriogenology 43, 509-522 Henderson KM, McNeeilly AS and IA Swanson (1982) Gonadotrophin and steroid concentration in bovine follicular fluid and their relationship to follicle size J Reprod Fertil 65, 467-473 Henkel RR and Schill WB (2003) Sperm preparation for ART Reprod Biol Endocrinol 1, 108 Herrler A, CA Krusche and HM Beier (1998) Insulin and Insulin-like growth factor-i promote rabbit blastocysts development and prevent apoptosis Biol reprod 59, 1302-1310 Hervé CP and JY le Cotonnec (1994) Pharmacokinetic and phamacodynamic cahracteristics of recombinant human follocle simulating hormone Ass Rep Rev 4, 110-117 Highighat N and van Winkle (1990) Developmental change in follicular cell-enhanced aminoacid uptake into mouse aocytes that depends on intact gap junction and transpor system gly J Exp Zool 253, 71-82 Hill JR, Burghardt RC, Jones K, Long CR, Looney CR, Shin T, Spencer TE, Thompson JA, Winger QA and ME Westhusin (2000) Evidence for placental abnormality as the major cause of mortality in first-trimester somatic cell cloned bovine fetuses Biol Reprod 63, 1787 1794 Hillery FL, JJ Parrish and NL First (1990) Bull specific effect on fertilization and embryo development in vitro Theriogenology 33, 249 Hirao YN (1994) In vitro growth and maturation of pig oocytes J Reprod Fert 100, 333-339 Ho YS, Dey MS, Crapo JD Antioxidant enzyme expression in rat lungs during hypoxia Am J Physiol 1996270, L810-L818 Holm P, Booth PJ and H Callesen (2002) Kinetics of early in vitro development of bovine in vivo- and in vitro-derived zygotes produced and/or cultured in chemically defined or serum-containing media Reproduction 123, 553 565 Hoshi H (2003) In vitro production of bovine embryos and their application for embryo transfer Theriogenology 59, 675-85 Hosoe M and Y Shioya (1997) Distribution of cortical granules in bovine oocytes classified by cumulus complex Zigote 5, 371-376 Howles CM (1996) Expression of human FSH (Gonal-F) by recombinant DNA technology Exc Hum Reprod 2, 183-189 Huet C, Monget P, Pisselet C, Hennequet C, Locatelli A and D Monniaux (1998) Steroidogenic enzymes, connexin-43, insulin-like-growth-factor-ii Mannose-6-phosphate receptor, extracellularmatrix components and matrix metalloproteinases Biol Reprod 58, 175-185 Hulshof SCJ, Figueiredo JR, Beckers JF, Bevers MM, van der Donk JA and R van den Hurk (1995) Effects of fetal bovine serum, FSH and 17ß-estradiol on the culture of preantral follicles Theriogenology 44, 217-226 Hunter RH, Vajta G and P Hyttel (1998) Long-term stability of the block J. Exp. Zool 280, 182-188 Hussein TS, Froiland DA, Amato F, Thompson JG and RB Gilchrist (2005) Oocytes prevent cumulus cell apoptosis by maintaining a morphogenic paracrine gradient of bone morphogenetic proteins J Cell Sci 118, 5257-5268 Hussein TS, Thompson JG and RB Gilchrist (2006) Oocyte-secreted factors enhance oocyte developmental competence Dev Biol 296, 514-521

Producción in vitro de embriones 323 Hyslop LA, Nixon VL, Levasseur M, Chapman F, Chiba K, A McDougall (2004) Ca 2+ -promoted cyclin B1 degradation in mouse oocytes requires the establishment of a metaphase arrest Dev Biol 269, 206 219 Hyttel P, Viuff D, Avery B, Laurincik J, and Greve (1996) Transcription and cell cycle-dependent development of intranuclear bodies and granules in two-cell bovine embryos J Reprod Fertil 108, 263 270 Hyttel P, Fair T, Callesen H and T Greve (1997) Oocyte growth, capacitation and final maturation in cattle Theriogenology 47, 23-32 Hyttel P, Greve T, H Callesen (1988) Ultrastructure of in-vivo fertilization in cattle J Reprod Fertil 82, 1 13 Hyttel P, T Greve and H Callesen (1989) Ultraestructural espects of oocyte maturation and fertilization in cattle J Reprod Fert 38, 35-47 Ikebuchi Y, Masumoto N, Matsuoka T, Yokoi T, Tahara M, Tasaka K, Miyake A, Y Murata (1998) SNAP-25 is essential for cortical granule exocytosis in mouse eggs Am J Physiol 274, 496 500 Imakawa K, Helmer SD, Nephew KP, Meka CS and RK Christenson (1993) A novel role for GM-CSF: enhancement of pregnancy specific interferon production, ovine trophoblast protein-1 Endocrinology 132,1869 1871 Inoue N, Ikawa M, Isotani A and M Okabe (2005) The immunoglobulin superfamily protein Izumo is required for sperm to fuse with eggs Nature 434, 234-238 Iwasaki S, Yoshiba N, Ushijima H, Watanabe S, T Nakahara (1990) Morfology and proportion of inner cell mass of bovine blastocysts followed fertilized in vitro and in vivo J Reprod Fert 90, 279-284 Jacobsen H, Schmidt M, Hom P, Sangild PT, Greve T and H Callesen Theriogenology 54, 147-158 James AM, Smith RA and MP Murphy (2004) Antioxidant and prooxidant properties of mitochondrial Coenzyme Q Arch Biochem Biophys 423, 47-56 Jeong YW, Park SW, Hossein MS, Kim S, Kim JH, Lee SH, Kang SK, Lee BC and WS Hwang (2006) Antiapoptotic and embryotrophic effects of a -tocopherol and L-ascorbic acid on porcine embryos derived from in vitro fertilization and somatic cell nuclear transfer Theriogenology 66, 2104-2112 Jones KT, Soeller C, MB Cannell (1998) The passage of Ca 2+ and fluorescent markers between the sperm and egg after fusion in the mouse Development 125, 4627 35 Jousan FD and PJ Hansen (2004) Insulin-like growth factor-i as a survival factor for the bovine preimplantation embryo exposed to heat shock Biol Reprod 71, 1665-70 Jousan FD and PJ Hansen (2007) Insulin-like growth factor-i promotes resistance of bovine preimplantation embryos to heat shock through actions independent of its anti-apoptotic actions requiring PI3K signaling Mol Reprod Dev 74, 189-196 Kaji K and A Kudo (2004) The mechanism of sperm-oocyte fusion in mammals Reproduction 127, 423-429 Kajihara Y, Blakewood EG, Myers MW, Kometari N, Goto K and RA Godke (1991) In vitro maturation and fertilization of follicular oocytes obtained from calves Theriogenology 35, 220 Kane MT (1987) Culture media and culture of early embryos Theriogenology 27, 49-57 Kane MT (1990) Regulation of embryonic development by environmental factors In: Wolf DP (Ed) In vitro fertilization and embryo culture Plenum Press, 359-387 Kauffold J, Amer HA, Bergfeld U, Weber W and A Sobiraj (2005) The in vitro developmental competence of oocytes from juvenile calves is related to follicular diameter J Reprod Dev 51, 325-32 Keefer CL (1992) Development of in vitro produced bovine embryos cultured individually in a simple medium: effect of EGF and TGFß1 Theriogenology 37, 236 Keefer CL and AM Paprocki (1995) Effect of percoll following sperm separation on in vitro fertilization of bovine oocytes Theriogenology 43, 244 Keefer CL, SL Stice, J Dobrinsky (1993) Effect of FSH and LH during bovine in vitro maturation on development following in vitro fertilization and nuclear transfer Mol Reprod Dev 36, 469-474 Keefer CL, SL Stice, AM Paprocki and P Golueke (1994) In vitro culture of embryos Cooperative interaction among embryos and role of growth factors Theriogenology 41, 1322-1331 Keskintepe L and BG (1996) Brackett. In vitro developmental competence of in vitro-matured bovine oocytes fertilized and cultured in completely defined media. Biol Reprod.55, 333-339 Kim J-H, Niwa K, Lim J-M and K Okuda (1993a) Effects of phosphate, energy substrate, and aminoacids on development of in vitro-matured, in vitro-fertilized bovine oocytes in a chemically defined, protein-free culture medium Biol Reprod 48, 1320-1325

324 Palma Kim JH, Funahashi H, Niwa K, and K Okuda (1993b) Glucose requeriment at different developmental stages of in-vitro fertilized bovine embryos cutlured in semi-defined medium Theriogenology 39, 875-886 King WA, Guay P and L Picard (1987) A citogenetical study of 7-day old bovine embryos of poor morphological quality Genome 29, 160-164 Kishida R, Lee ES and Y Fukui (2004) In vitro maturation of porcine oocytes using a defined medium and developmental capacity after intracytoplasmic sperm injection Theriogenology 62, 1663-1676 Kobayashi M, Lee ES, Y Fukui (2006) Cysteamine or beta-mercaptoethanol added to a defined maturation medium improves blastocyst formation of porcine oocytes after intracytoplasmic sperm injection Theriogenology 165, 1191-1199 Kurzawa R, Glabowski W, Baczkowski T, P Brelik (2002) Evaluation of mouse preimplantation embryos exposed to oxidative stress cultured with insulin-like growth factor I and II, epidermal growth factor, insulin, transferrin and selenium Reprod Biol 2, 143-162 Kölle S, Sinowatz F, Boie G and G Palma (1998) Differential expression of ZPC in the bovine ovary, oocyte, and embryo Mol Reprod Dev 49, 435 443 Knijn, HM, Wrenzycki, C, Hendriksen, PJ, Vos, PL, Zeinstra, EC, van der Weijden, GC, Niemann, H and SJ Dieleman (2005) In vitro and in vivo culture effects on mrna expression of genes involved in metabolism and apoptosis in bovine embryos Reprod Fertil Dev 17, 775 784 Knott JG, Gardner AJ, Madgwick S, Jones KT, Williams CJ, Schultz RM (2006) Calmodulindependent protein kinase II triggers mouse egg activation and embryo development in the absence of Ca 2+ oscillations Dev Biol 296, 388-395 Knott JG, Kurokawa M, Fissore RA, Schultz RM and CJ Williams (2005) Transgenic RNA interference reveals role for mouse sperm phospholipase Czeta in triggering Ca 2+ oscillations during fertilization Biol Reprod 72, 992-996 Kouchi Z, Fukami K, Shikano T, Oda S, Nakamura Y, Takenawa T, and S Miyazaki (2004) Recombinant phospholipase C{zeta} has high Ca21 sensitivity and induces Ca 2+ oscillations in mouse eggs J Biol Chem 279, 10408 10412 Kraüßlich H, Palma GA, G Brem (1997) Techniques of bovine embryo production and their possible consequences for breeding strategies and the future role of practitioners in embryo transfer Proceedings of 13 Scientific Meeting of European Embryo Transfer Association, 115-122 Krefft H (1993) Untersuchungen zur Wirkung von Wachstumsfaktoren bei der Entwicklung von Rinderembryonen in vitro PhD Thesis Ludwig-Maximilian University of Munich, Germany 1-135 Krisher RL, Lane M and BD Bavister (1999) Developmental competence and metabolism of bovine embryos cultured in semi-defined and defined culture media Biol Reprod 60, 1345-1352 Kruip Th AM, Boni R, Würth AA, Roelfsen MWM and MC Pieterse (1994) Potencial use of Ovum Pick-Up for embryo production and breeding in cattle J of Dairy Sci 75,2857-2879 Kruip TAM and JHG den Daas (1997) In vitro produced and cloned embryos: effects on pregnancy, parturition and offspring Theriogenology 47, 43 52 Kurokawa M, Sato K and RA Fissore (2004) Mammalian fertilization: from sperm factor to phospholipase C zeta Biol Cell 96, 37 45 Kurzawa R, Glabowski W, Baczkowski T, Wiszniewska B and M Marchlewicz (2004) Growth factors protect in vitro cultured embryos from the consequences of oxidative stress Zygote 12, 231-40 Lackey BR, SL Gray and DM Henricks (2000) Physiological basis for use insulin-like growth factors in reproductive applications: a review Theriogenology 53, 1147-1156 Lane M and BD Bavister (1999) Regulation of intracellular ph in bovine oocytes and cleavage stage embryos Mol Reprod Dev 54, 396-401 Lane M and DK Gardner (1991) Effect of incubation volume and embryo density on the development and viability of mouse embryos in vitro In: Proc of the Symposium on preimplantation embryo development Newton Massachusetts, USA, 33 Lane M, DK Gardner (1997) Differential regulation of mouse embryo development and viability by amino acids J Reprod Fertil 109, 153 164 Lang F, Busch GL, Ritter M, Völkl H, Waldegger S, Gulbins E, D Häussinger (1998) Functional significance of cell volume regulatory mechanisms Physiol Rev 78, 247 306 Langendonckt AV, Donnay I, Schurbiers N, Auquier P, Carolan C, Massip A, F Dessy (1997) Effect of supplementation with fetal calf serum on development of bovine embryos in synthetic oviduct fluid médium J Reprod Fertil 109, 87 93 Lappas M, Permezel M, and GE Rice (2003) N-acetyl-Cysteine inhibits phospholipid metabolism, proinflammatory cytokine release, protease activity, and nuclear factor-κb deoxyribonucleic acid-

Producción in vitro de embriones 325 binding activity in human fetal membranes in vitro The Journal of Clinical Endocrinology & Metabolism 88, 1723-1729 Larson RC, GG Ignotz, WB Currie (1990) Defined medium containing TGF-beta and bfgf permits development of embryos beyond the 8-cell Block J Reprod Fert 5, 6 Larson RC, GG Ignotz, WB Currie (1992a) Platelet derived growth factor (PDGF) stimulates development of bovine embryos during fourth cell cycle Development 155, 821-826 Larson RC, GG Ignotz and WB Currie (1992b) Transforming growth factor-beta and basic fibroblast growth factor synergically promote early bovine embryo development during the fourth cell cycle Mol Reprod Dev 33, 432-435 Lazzari G, G Crotti, R Duchi and C Galli (1999) Heparin effects on blastocyst formation and cell number of the IVM-IVF bovine embryo Theriogenology 51, 242 Lazzari G and C Galli (1994) The effect of heparin supplementation during IVM on cumulusexpansion and developmental capacity of bovine oocytes 10th Scientific Meeting of European Embryo Transfer Association, 202 Lawrence TH, Beers WH and NB Gilula (1978) Transmission of hormonal stimulation by cell-to-cell comunication Nature 272, 501-506 Leyens G, Knoops B and I Donnay (2004) Expression of Peroxiredoxins in Bovine Oocytes and Embryos Produced In Vitro Molecular reproduction and development 69, 243-251 Liebler DC (1993) The role of metabolism in the antioxidant function of vitamin E Crit Rev Toxicol 23, 147-169 Lima PF, Oliveira MA, Goncalves PB, Montagner MM, Reichenbach HD, Weppert M, Neto CC, Pina VM, MH Santos (2004) Effects of retinol on the in vitro development of Bos indicus embryos to blastocysts in two different culture systems Reprod Domest Anim 39, 356-360 Lima PF, Oliveira MA, Santos MH, Reichenbach HD, Weppert M, Paula-Lopes FF, Neto CC, PB Goncalves (2006) Effect of retinoids and growth factor on in vitro bovine embryos produced under chemically defined conditions Anim Reprod Sci 95, 184-92 Lee ES, Fukui Y, Lee BC, Lim JM, WS Hwang (2004) Promoting effect of amino acids added to a chemically defined medium on blastocyst formation and blastomere proliferation of bovine embryos cultured in vitro Anim Reprod Sci 84, 257-67 Leibfried-Rutledge ML, ES Crister, JJ Parrish and NL First (1989) In vitro maturation and fertilization of bovine oocytes Theriogenology 31, 61-74 Lenhart JA, Downey BR and CA Bagnell (1998) Connexin-43 gap junction protein expression during follicular development in teh porcine ovary Biol Reprod 58, 583-590 Levy R, Smith SD, Chandler K, Sadovsky Y, DM Nelson (2000) Apoptosis in human cultured trophoblasts is enhanced by hypoxia and diminished by epidermal growth factor Am J Physiol Cell Physiol 278, C982-C988 Li R, Norman RJ, Armstrong DT and RB Gilchrist (2000) Oocyte-secreted factor(s) determine functional differences between bovine mural granulosa cells and cumulus cells Biol Reprod 63, 839 845 Lindl T und J Bauer (1994) Zell- und Gewebekultur In: Gustav Fischer (Ed), Suttgart, Jena, New York, 77-79 Loob DK and J Dorrington (1992) Intraovarian regulation of follicular development Anim. Reprod Sci 28, 343-354 Loewenstein WR (1987) The cell-to-cell channel gap junctions Cell 48, 725-726 Loewenstein WR and B Rose (1992) The cell-cell channel in the control of growth Semin Cell Biol 3, 59-79 Lonergan P (1994) The application of in vitro fertilization techniques to the prediction of bull fertility Reprod Dom Anim 29, 12-21 Lonergan P, Dinnyes A, Fair T, Yang X and M Boland (2000) Bovine oocyte and embryo development following meiotic inhibition with butyrolactone I Mol Reprod Dev 57, 204 209 Lonergan P, Rizos D, Gutierrez-Adan A, Fair T and MP Boland (2003) Oocyte and embryo quality: effect of origin culture conditions and gene expression patterns Reprod Domest Anim 38, 259-267 Lonergan P, Rizos D, Gutierrez-Adan A, Fair T and MP Boland (2003) Effect of culture environment on embryo quality and gene expression - experience from animal studies Reprod Biomed Online 7, 657-63 Lonergan P, Gutierrez-Adan A, Pintado B, Fair T, Ward F, Fuente JD, Boland M (2000) Relationship between time of first cleavage and the expression of IGF-I growth factor, its receptor, and two

326 Palma housekeeping genes in bovine two-cell embryos and blastocysts produced in vitro Mol Reprod Dev 57,146-152 López L, Alvarez N, Nuñez I, Montes I, Solano R, Fuentes D, de Armas R, Palma, GA and G Brem (1995) The influence of body condition on the ovarian morphology and quality of bovine oocytes for in vitro production of embryos Proceedings of the 13 th International Congress on Animal Reproduction, 10-15 López Ruiz L, N Alvarez, I Nuñez, I Montes, R Solano, D Fuentes, R Pedroso, GA Palma and G Brem (1996) Effect of body condition on the developmental competence of IVM/IVF bovine oocytes Theriogenology 45, 292 Lorca T, Cruzalegui FH, Fesquet D, Cavadore JC, Mery J, Means A and M Doree (1993) Calmodulindependent kinase II mediates inactivation of MPF and CSF upon fertilization of Xenopus eggs Nature 366, 270 273 Lorenzo PL, MJ Illera, JC Illera and M Illera (1994) Enhancement of cumulus expansion and nuclear maturation during bovine in vitro by the addition of epidermal growth factor and insulin-like growth factor I J Reprod and Fert 101, 697-701 Lorton SF, NL First (1979) Hyaluronidase does not disperse the cumulus oophorus surrounding bovine ova. Biol Reprod 21, 301 308 Lu KH and C Polge (1992) A summary of two years results in large scale in vitro bovine embryo production Proceedings of the 12th International Congress on Animal Reproduction, The hague 3, 1315-1317 Luciano AM, S Modina, F Gandolfi, A Lauria and DT Armstrong (2000) Effect of cell-to-cell contact on In vitro deoxyribonucleic acid synthesis and apoptosis responses of bovine granulosa cells to insulin-like growth factor-i and epidermal growth factor Biol Reprod 63,1580-1585 Lucy MC (2000) Regulation of ovarian follicular growth by somatotropin and insulin-like growth factors in cattle J Dairy Sci 83, 1635-1647 Ludwig G, J Frick, E Rovan (1996) Glaswollfiltration In: Praxis der Spermatologie, Springer, Berlin 111-112 Luria A, Tennenbaum T, Sun QY, Rubinstein S, H Breitbart (2000) Differential localization of conventional protein kinase C isoforms during mouse oocyte development Biol Reprod 62, 1564 1570 Luvoni GC, Keskintepe L and BG (1996) Brackett Improvement in bovine embryo production in vivo by glutathione-containing culture media Mol Reprod Dev 43, 437-443 Madison V, Avery B and T Greve (1992) Selection of immature bovine oocytes for developmental potential in vitro Anim Reprod Sci 27, 1-11 Malcuit C, Kurokawa M and RA Fissore (2006) Calcium oscillations and mammalian egg activation J Cell Physiol 206, 565-573 Manes C, Lai NC Nonmitochondrial oxygen utilization by rabbit blastocysts and surface production of superoxide radicals J Reprod Fertil 1995104, 69-75 Markoulaki S, Matson S and T Ducibella (2004) Fertilization stimulates long-lasting oscillations of CaMKII activity in mouse eggs Dev Biol 272, 15 25 Marquant-Le Guienne B, Gerard B, Solari A and C Thibault (1989) In vitro culture of bovine eggs fertilized either in vivo or in vitro Reprod Nutr Dev 29, 559-568 Marquant-Le Guienne B, Guyader-Joly C, Ponchon S, Delalleau N, Florin B, Ede P, Ponsart C, Guérin B and P Humblot (2001) Results of in vitro production in a commercial ovum pickup program. In: Annual Conference of the, Theriogenology 55, 433 Marquant-Leguienne B and P Humblot (1998) Practical measures to improve in vitro blastocyst production Theriogenology 3-11 Marquant-Leguienne B, P Humblot, M Thibier, C Thibault (1990) Evaluation of bull fertility by homologous in vitro fertilization tests Reprod Nutr Dev 30, 259-266 Martin KL, Hardy K, Winston RM, Leese HJ (1993) Activity of enzymes of energy metabolism in single human preimplantation embryos J Reprod Fertil 99, 259-66 Martinovich PN (1938) The development in vitro of the mammalian gonad-ovary and oogenesis Proc Roy Soc B 125, 232-249 Massip A, Mermillod, A Dinnye (1995) Morphology and biochemistry of in vitro produced bovine embryos: impliations for their cryopreservations Hum Reprod 10, 3004-3011 Massip A, Mermillod P, Van Langendonkt A, Reichenbach HD, Lonergen P, Carolan C, De Roover R and G brem (1996) Calving outcome following transfer of embryos produced in vitro in different conditions Anim Reprod Sci 44, 1-10

Producción in vitro de embriones 327 Mastromonaco GF, Semple E, Robert C, Rho GJ, Betts DH, WA King (2004) Different culture media requirements of IVF and nuclear transfer bovine embryos Reprod. Domest Anim 39, 462 467 McAvey BA, WortzmanGB, Williams CJ and JP Evans (2002) Involvement of calciumsignaling and the actin cytoskeleton in themembrane block to polyspermy in mouse eggs Biol Reprod 67, 1342 1352 Mc Evoy TG, Robinson JJ, Aitken RP, Findlay PA and IS Robertson (1997) Dietary excessif urea influence the viability and metabolism of preimplantation sheep embryos and many affect fetal growth among survivors Anim Reprod Sci 47, 71-90 Mc Evoy TG, Robinson JJ, Carolan C, Staines ME, Broadberet P and KD Sinclair (1999) Ovine fetal development following embryo cutlure in synthetic ovidct fluid with added of amonium Theriogenology 51, 247 Mc Evoy TG, Sinclair KD, Broadbent PJ, Goodhand KL and JJ Robinson (1998) Reprod Fertil Dev 10, 459-464 McKiernan SH and BD Bavister (2000) Culture of one-cell hamster embryos with water soluble vitamins: pantothenate stimulates blastocyst production Hum Reprod 15, 157-164 McNatty KP, Moore LG, Hudson NL, Quirke LD, Lawrence SB, Reader K, Hanrahan JP, Smith P, Groome NP, Laitinen M, Ritvos O, JL Juengel (2004) The oocyte and its role in regulating ovulation rate: a new paradigm in reproductive biology Reproduction 128, 379-386 McNatty KP, Smith DM, Makris A, OsathanondhR, KJ Ryan (1979) The microenvironment of the humn antral follicle: interrelationships among the steroid levels in antral fluid, the popöulation of granulosa cells, and the status of the oocyte in vivo and in vitro J Clin Endocrinol Metab 49, 851-860 McNatty KP, Smith P, Moore LG, Reader K, Lun S, Hanrahan JP, Groome NP, Laitinen M, Ritvos O, JL Juengel (2005) Oocyte-expressed genes affecting ovulation rate Mol Cell Endocrinol 29, 234, 57-66 Memili E and NL First (1999) Control of gene expression at the onset of bovine embryonic development Biol Reprod 61, 1198 1207 Memili E and NL First (2000) Zygotic and embryonic gene expression in cow: a review of timing and mechanisms of early gene expression as compared with other species Zygote 8, 87-96 Miller JK and E Brzezinska-Slebodzinska (1993) Oxidative stress, antioxidants and animal function J Diary Sci 76, 2812-2823 Miyano T, Ozaki S, Matsumura K, Miyake M, Kato S, K Yamamoto (1998) In vitro growth, fertilization and development of bovine oocetes from early antral follicles En: Lauria A, Gandolfi F, Enne G, L Gianaroli (eds) Gametes developmental and function Serono Symposia, 575 Miyazaki S and M Ito (2006) Calcium signals for egg activation in mammals J Pharmacol Sci 100, 545-552 Modina S, Baraldi Scesi L, Mauri L, Ponzini A, Lauria A, F Gandolfi (1998) localization and expression of EGF, TGF-α and their receptors in bovine ovarian follicles In: Lauria A, Gandolfi F, Enne G, L Gianaroli (eds) Gametes developmental and function, Serono Symposia, 576 Moller CC and PM Warssaman (1989) Characterization of a proteinase that cleaves zona pellucida glycoprotein ZP2 following activation of mouse eggs Dev Biol 132, 103 112. Monniaux D, Monget P, Besnard C, Huet C and C Pisselet (1997) Growth factors and antral follicular development in domestic rumiants Theriogenology 47, 3-12 Moreira F, Paula-Lopes FF, Hansen PJ, Badinga L and WW Thatcher (2002) Effects of growth hormone and insulinlike growth factor-i on development of in vitro derived bovine embryos Theriogenology 57, 895 907 Mortimer ST (2000) CASA-Practical Aspects J Androl 21, 515-524 Motlik J, T Nagai y K Kikuchi (1991) Resumption of meiosis in pig oocytes cultured with cumulus parietal granulosa cells: the effect of protein synthesis inhibition J Exp Zool 259, 386-391 Motlik J, Pavlok A, Lapathitis G and M Kubelka (2000) Impact of two-step in vitro culture systems on developmental potency of oocytes Rep Dom Anim 35, 267-271 Nagano M, Katagiri S, Takahashi Y (2006b)ATP content and maturational/developmental ability of bovine oocytes with various cytoplasmic morphologies Zygote. 14, 299-304 Nagano M, Katagiri S and Y Takahashi (2006) Relationship between bovine oocyte morphology and in vitro developmental potential Zygote 14, 53-61 Narula A, Taneja M, SM Totey (2000) Morphological development, cell number, and allocation of cells to trophectoderm and inner cell mass of in vitro fertilized and parthenogenetically developed buffalo embryos: the effect of IGF-I Mol Reprod Dev 44, 343-51

328 Palma Nayudu PL, SM Osborn (1992) Factors influencing the rate of preantral and antral growth of mouse ovarian follicles in vitro J Reprod Fertil 95, 345-396 Nedambale TL, Du F, Yang X, XC Tian (2006) Higher survival rate of vitrified and thawed in vitro produced bovine blastocysts following culture in defined medium supplemented with betamercaptoethanol Anim Reprod Sci 93, 61-75 Noguchi S (2005) The biological function of insulin-like growth factor-i in myogenesis and its therapeutic effect on muscular dystrophy Acta Myol 24, 115-118 Numabe T, Okinawa T, Kikuchi T and T Horiochi (2000) Birth and Birth rate of heavy calves conceived by transfer of in vitro or in vivo produced bovine embryos Anim Reprod Sci 64, 13-20 Oliver JE, TJ Aitman, JF Powell, CA Wilson, RN Clayton (1989) Insulin-like growth factor-i gene expression in the rat ovary is confined to the granulosa cells of developing follicles Endocrinol 124, 2671-2676 Olivier NS, Palma GA y R Alberio (1998) In vitro production of bovine embryos in water bath Theriogenology 49, 211 Olson SE and GE Seidel Jr (2000) Culture of in vitro-produced bovine embryos with vitamin E improves development in vitro and after transfer to recipients Biol Reprod 62, 248-252 O Neill C (1997) Evidence for the requirement of autocrine growth factors for development of mouse preimplantation embryos in vitro Biol of Reprod 56, 229 237 Osaki S, Matsumura K, Yamamoto K, Miyano T, Miyake M, S Kato (1997) Fertilization of bovine oocytes grown in-vitro Reprod Fert Devel 9, 781-787 Otoi T, S Tachikawa, S Kondo and T Suzuki (1993) Effects of different lots of semen from the same bull on in vitro development of bovine oocyte fertilized in vitro Theriogenology 39, 713-718 Otoi T, Yamamoto K, Koyama N, Tachikawa S and T Suzuki T (1997) Bovine oocyte diameter in relation to developmental competence Theriogenology 48, 769 774 Overstöm EW (1996) In vitro assessment of embryo viability Theriogenology 45, 3-16 Oyamada T, Iwayama H and Y Fukui (2004) Additional effect of epidermal growth factor during in vitro maturation for individual bovine oocytes using a chemically defined medium Zygote 12, 143-150 Oyawoye O, Abdel Gadir A, Garner A, Constantinovici N, Perrett C, P Hardiman (2003) Antioxidants and reactive oxygen species in follicular fluid of women undergoing IVF, relationship to outcome Hum Reprod 18, 2270-2274 Palma GA (1994) Effect of FSH and estradiol-17b for maturarion of calf oocytes on the in vitro development to blastocysts Theriogenology 41, 267 Palma GA and R Alberio (1998) Foetal bovine serum in maturation medium has no beneficial effect on in vitro development and cell number of bovine blastocysts Proceedings of 14 th Scientific Meeting of European Embryo Transfer Association, 228 Palma GA, Aller JF and RH Alberio (1999) Pregnancies after transfer of in vitro-produced bovine blastocysts derived from oocytes matured in defined medium Theriogenology 51, 390 Palma GA y G Brem (1994) Are there bulls unsuitable for IVF? Proceedings of the 10th Meeting of European Embryo transfer Society, Lyon, 230 Palma GA and G Brem (1996) Effect of high concentrations of IGF-I on the development of in vitro produced bovine embryos Arch Anim Breed 39, 85 Palma GA, J Braun, R Stolla and G Brem (1996) The abbility to produce embryos in vitro using semen from different bulls with a low non-return rate Theriogenology 45, 308 Palma GA, A Clement-Sengewald, U Berg and G Brem (1992) Role of the embryo number in the development of in-vitro produced bovine embryos Theriogenology 37, 271 Palma GA, A Clement-Sengewald, H Krefft and G Brem (1993) Effect of insulin-like growth factor (IGFI) on development of in vitro produced bovine embryos in two culture densities 9th Scientific Meeting of European Embryo Transfer Society, Lyon, 248 Palma GA, A Clement-Sengewald and H Krefft (1993a) In vitro production of cattle embryos from calf oocytes Theriogenology 39, 278 Palma GA, A Clement-Sengewald, U Berg and G Brem (1992) Role of the embryo number in the development of in-vitro produced bovine embryos Theriogenology 37, 271 Palma GA, A Clement-Sengewald and G Brem (1994) Effect of FSH and Estradiol-17ß for maturation of calf oocytes on the in vitro development to blastocysts Theriogenology 41, 267 Palma GA, Krefft H, Clement-Sengewald A, J Zhu and G Brem (1993b) Results of large scale in vitro embryo production Proceedings of the 9th Meeting of European Embryo transfer Society, Lyon, 250

Producción in vitro de embriones 329 Palma GA, J Mödl and G Brem (1997) The influence of different combinations of bovine LH and bovine FSH on the maturation of cattle oocytes Reprod Dom Anim 32, 81 Palma GA, M Müller and G Brem (1997b) Effect of Insulin-like growth factor I (IGF-I) at high concentration on blastocyst development of bovine embryos produced in vitro J Reprod Fert 110, 347-353 Palma GA, N Olivier, Mödl J, Wenigerkind H, G Brem (1996) Effect of cumulus investment from ultrasound collected oocytes on the development of in vitro produced embryos by elite cows Proceedings of the 12 th Scientific Meeting of the European Embryo Transfer Association, 180 Palma GA, Olivier N, Alberio RH, Aller J, G Brem (1998) Effect of a Bag system on the development capacity and viability of in vitro produce bovine embryos Proceedings of 14 th Scientific Meeting of European Embryo Transfer Association, 228 Palma GA, Mödl J and G Brem (1997a) The influence of diferent combinations of bovine LH and bovine FSH on the maturation of cattle oocytes Reprod Dom Anim 32, 81 Palma GA, Olivier N, Alberio R and G Brem (1997c) Effect of environmental conditions in the incubator on the development of in vitro produced bovine embryo Biol Reprod 56, 530 Palma GA, Olivier N, Alberio RH and G Brem (1998) In vitro development and viability of bovine embryos produced without gassed incubator Theriogenology 49, 213 Palma GA and Sinowatz (2004) Male and female effects on the in vitro production of bovine embryos Anat Histol Embryol 33, 257-262 Palma GA, Tortonese DJ and F Sinowatz (2001) Developmental capacity of in vitro prepubertal oocytes Anat Histol Embryol 30, 1-6 Palma GA, Wennigerkind H, Mödl J and G Brem (1995b) Biotecnologías aplicadas en la reproducción bovina Estado actual y aplicaciones futuras Rev Arg Prod Anim 1, 159-165 Palma GA, V Zakhartchenko and G Brem (1995a) In vitro production of bovine embryos with oocytes from individual slaughtered elite cows: Developmental competence and subsequent calving rates J Anim Sci 73, 221 Pampfer S and I Donnay (1999) Apoptosis at the time of embryo implantation in mouse and rat Cell Death Differ 6, 533-545 Paria BC and SK Dey (1990) Preimplantation embryo development in vitro: cooperative interactions among embryos and role of growth factors Dev Biol 87, 4756-4760 Parks JE, JW Aroin, RH Foote (1987) Lipids of plasma membrane and outer acrosomal membrane from bovine spermatozoa Biol Reprod 37, 1249-1258 Parrish JJ, JL Parrish and NL First (1984) Effect of swim-up separation and heparin pretreatment of frozen-thawed spermatozoa on in vitro fertilization of bovine oocytes Biol Reprod 30 (Suppl 1) 112 Parrish JJ, Kim CI, IH Bae (1992) Current concepts of cell-cycle regulation and its relationship to oocyte maturation, fertilization and embryo development Theriogenology 38, 277-296 Parrish JJ, JL Susko-Parrish, ML Leifried-Rutledge, ES Crister, WH Eyestone and NL First (1986b) Bovine in vitro fertilization with frozen-thawed semen Theriogenology 25, 591-600 Paulenz H, Greve IS, Tverdal A, Hofmo PO and K Anderssen Berg (1995) Precision of the coulter Counter for routine assesment of boar sperm concentration in comparison with the haemocytometer and spectrophotometer Reprod Dom Anim 30, 107-111 Pavlok A, Lucas-Hahn A and H Niemann (1992) Fertilization and developmental competence of bovine oocytes derived from different categories of antral follicles Mol Reprod Develop 31, 63-67 Persson E, H Rodriguez-Martinez, V Dantzer (1992) Immunocytochemical localization of growth factors during porcine placentation Proceedings of the 12th International Congress on Animal Reproduction The Hague 3, 1267-1269 Pierce GB, Parchment RE, AL Lewellyn (1991) Hydrogen peroxide as a mediator of programmed cell death in the blastocyst Differentiation 46, 181-186 Pincus G and EV Enzmann (1935) The comparative behavior of mamalian eggs in vivo and in vitro J Exp Med 62, 665-675 Pinyopummintr T and BD Bavister (1991) In vitro-matured/in vitro-fertilized bovine oocytes can develop into morulae/blastocysts in chemically defined, protein-free culture media Biol Reprod 45, 736-742 Pollard JW and SP Leibo (1993) Comparative cryobiology of in vitro and in vivo derived bovine embryos Theriogenology 39, 287 Pomar FJ, Teerds KJ, Kidson A, Colenbrander B, Tharasanit T, Aguilar B, BA Roelen (2005) Differences in the incidence of apoptosis between in vivo and in vitro produced blastocysts of farm animal species: a comparative study Theriogenology 63, 2254-2268

330 Palma Popper K (1989) Falsifizierbarkeit In: Logik der Forschung In: JCB Mohr (P Siebeck) Tübngen, 47-59 Ptak G, Matsukawa K, Palmieri C, Della Salda L, Scapolo PA and P Loi (2006) Developmental and functional evidence of nuclear immaturity in prepubertal oocytes Hum Reprod 21, 2228-2237 Prather RS and NL First (1993) Cell-to-cell coupling in early -stage bovine embryos: a preliminary report Theriogenology 39, 561-567 Prelle K, Stojkovic M, Boxhammer K, Motlik J, Ewald D, Arnold GJ and E Wolf (2001) Insulin-like growth factor I (IGFI) and long R3 IGF-I differently affect development and messenger ribonucleic acid abundance for IGF-binding proteins and type I IGF receptors in in vitro produced bovine embryos Endocrinology 142, 1309 1316. Presicce GA, Jiang S, Simkin M, Zhang L, Looney CR, Godke RA and X Yang (1997) Age and hormonal dependence of acquisition of oocyte competence for embryogenesis in prepubertalcalves Biol Reprod 56, 386-392 Rabahni F, Monniaux D, Pisselet C, Chupin D and P Durand (1991) Qualitative and quantitative changes in protein synthesis of bovine follicular cells during the preovulatory period Mol Reprod Dev 30, 365-274 Revel F, Mermillod P, Peynot N, Benard JP and Y Heyman (1995) Low developmental capacity of in vitro matured and fertilised oocytes from calves compared to that of cows J Reprod Fertil 103, 115-120 Rhee SG (1999) Redox signaling: hydrogen peroxide as intracellular messenger Exp Mol Med 31, 53-59 Rice A, Parrington J, Jones KT, K Swann (2000) Mammalian sperm contain a Ca 2+ sensitive phospholipase C activity that can generate InsP3 from PIP2 associated with intracellular organelles Dev Biol 227,125 135 Rizos D, Lonergan P, Ward F, Duffy P and MPBoland (2002) Consequences of bovine oocyte maturation, fertilization or early embryo development in vitro versus in vivo: implications for blastocyst yield and blastocyst quality Mol Reprod Dev 61, 234 248 Rizos D, Ward F, Duffy P, Boland MP and P Lonergan (2002) Consequences of bovine oocyte maturation, fertilization or early embryo development in vitro versus in vivo: implications for blastocyst yield and blastocyst quality Mol Reprod Dev 61, 234-248 Robinson RS, Mann GE, Gadd TS, Lamming GE, Wathes DC (2000) The expression of the IGF system in the bovine uterus throughout the oestrous cycle and early pregnancy J Endocrinol 165, 231-43 Rodgers RJ, van Wezel IL, Lavranos TC, Rodgers HF, Irvine CM and M Krupa (1998) Developmental changes in cells and matrix during follcicle growth In: Lauria A, Gandolfi F, Enne G, L Gianaroli (eds) Gametes developmental and function, Serono Symposia, 85-98 Rodriguez-Martinez H (2004) Selection of bull spermatozoa by swim-up, Workshop Communications of the 15 th International Congress on Animal Reproduction Porto Seguro, Brasil, 85 Rosenkrans CFJr and NL First (1994) Effect of aminoacids and vitamins on cleavage and developmental rate of bovine zygotes in vitro J Anim Sci 72, 434-437 Rougier N and Z Werb (2001) Minireview: Parthenogenesis in mammals Mol Reprod Dev 59, 468-74 Roy SK and BJ Treacy (1993) Isolation and long term culture of human preantral follicles Fert Steril 59,783-790 Russell DF, Baqir S, Bordignon J and DH Betts (2006) The impact of oocyte maturation media on early bovine embryonic development Mol Reprod Dev 73, 1255-1270 Rubinstein E, Ziyyat A, Wolf JP, Le Naour F, C Boucheix (2006) The molecular players of sperm-egg fusion in mammals Semin Cell Dev Biol 17, 254-263 Saeki K, M Nagao, M Hoshi and M Nagai (1995) Effects of heparin, sperm concentration and bull variation on in vitro fertilization of bovine oocyte in a protein-free medium Theriogenology 43, 751-759 Sagirkaya H, Misirlioglu M, Kaya A, First NL, Parrish JJ, E Memili (2007) Developmental potential of bovine oocytes cultured in different maturation and culture conditions Anim Reprod Sci Oct 17; (en prensa) PMID: 17052869 doi:10.1016/j.anireprosci.2006.09.016 Salamone DF, Damiani P, Fissore RA, Robl JM, RT Duby (2001) Biochemical and developmental evidence that ooplasmic maturation of prepubertal bovine oocytes is compromised Biol Reprod 64, 1761-1768 Salamone DF, Fissore RA, Robl JM and RT Duby (2000) Assessment of ooplasmic and nuclear maturity in calf oocytes by reciprocal transfer of metaphase II chromosomes or somatic cells Theriogenology 53, 242

Producción in vitro de embriones 331 Sánchez R, Concha M, Ichikawa T, Henkel R and WM Schill (1996) Glass wool filtration reduces reactive oxygen species by elimination of leukocytes in oligozoospermic patients with leukocytospermia J Assisted Reprod Genet 13, 489 494 Sardet C, Dumollard R, A McDougall (2006) Signals and Calcium waves at fertilization Sem Cell Devel Biol 17, 223-225 Sato E, Matsuo M, H Miyamoto (1990) Meiotic maturation of bovine oocytes in vitro: improvement of meiotic competence by dibutyryl ciclic adenosine 3,5 -monophosphate J Anim Sci 68,1182-1187 Saunders CM, Larman MG, Parrington J, Cox LJ, Royse J, Blayney LM, Swann K, FA Lai (2002) PLC: a sperm-specific trigger of Ca2+ oscillation in eggs and embryo development Development 129, 3533 3544 Saunders CM, Larman MG, Parrington J, Cox LJ, Royse J, Blayney LM, Swann K, FA Lai (2002) PLC: a sperm-specific trigger of Ca2+ oscillation in eggs and embryo development Development 129, 3533 3544 Scherntahner W, Schmoll F, Brem G, K Schellender (1997) Storing bovine ovaries for 24 hours between 15 and 21 C does not influence in vitro production of blastocysts Theriogenology 47, 297 Schernthaner W, Wenigerkind H, Stojkovic M, Palma GA, Modl J, Wolf E, G Brem (1999) Pregnancy rate after ultrasound-guided follicle aspiration in nonlactating cows from different breeds Zentralbl Veterinarmed A 46, 33-7 Shimasaki S, Moore RK, Otsuka F, GF Erickson (2004) The bone morphogenetic protein system in mammalian reproduction Endocr Rev 25 72 101 Schultz GA, A Hogan, A Watson, Smith RM and S Heyner (1992) Insulin-like growth factors and glucose transporters temporal patterns of gene expression in early murine and bovine embryos Reprod Fert Dev 4, 361-371 Schultz RM (1987) Molecular aspects of oocyte growth and maturation, In: Experimental approaches to mammalian embryonic development Univ Press, Cambridge Schultz RM (2005) From egg to embryo: a peripatetic journey Reproduction 130, 825-828 Schultze D, B Brunswig, AK Mukhpadhyay (1989) Renin and prorenin-like activities in bovine ovarian follicles Endocrinol 124, 1389-1398 Schwartz J, MR Schneider, JL Rodrigues and H-D Reichenbach (1998) Effect of short-term storage of bovine oocytes in different media and temperatures on the subsequent in vitro embryo development Theriogenology 49, 217 Schweitzer CM, Gonzalvez PBD, Neves JP, Severo NC, morales CN, AG Hernández (1996) Influencia deo percoll na capacitacao espermática para fecundacao in vitro de ooctos bovinos XI Congreso brasilero de reproduçao animal, Belo horizonte, Minas Gerais, 275 Shamsuddin M (1993) Effect of growth factors on bovine blastocyst development in a serum-free medium PhD Thesis, Swedish University of Agricultural Sciences Sweden, pp 1-60 Shamsuddin M and B Larsson (1993) In vitro development of bovine embryos after fertilization using semen from different donors Reprod Dom Anim 28, 77-84 Shamsuddin M, Larsson B, Gustafsson H and H Rodriguez-Martinez (1993) A serum-free, cell culture system for development of bovine one-cell embryos up to blastocist stage with improve viability Doctoral Thesis, Univ Of Uppsala, Sweden V, 1-12 Sharma RK and A (1996) Agarwal Role of reactive oxygen species in male infertility Urology 48, 835-850 Shimasaki S, Moore, RK, Otsuka, F and GF Erickson (2004) The bone morphogenetic protein system in mammalian reproduction Endocr Rev 25, 72 101 Sikka SC (2004) Role of oxidative stress and antioxidants in andrology and assisted reproductive technology J Androl 25, 5-18 Simmen RCM, Y Ko, FA Simmen (1993) Insulin-like growth factors and blastocyst development Theriogenology 39,163-175 Sinclair KD, Broadbent PJ and DF Dolman (1995) In vitro produced embryos as a means of achieving pregnancy and improving productivity in beef cows Anim Sci 60, 55-64 Sirard MA and P Blondin (1996) Oocyte maturation and IVF in cattle An Reprod Sci 42417-426 Sirard MA, Coenen, K (1993) The co-culture of cumulus enclosed bovine oocytes and hemisections of follicles, effects on meiotic resumption Theriogenology 40, 933-942 Sirard MA, Richard F and M Mayes (1998) Controlling meiotic resumption in bovine oocytes: a review Theriogenology 49, 483-497 Sirard MA, Richard F, Blondin P, C Robert (2006) Contribution of the oocyte to embryo quality Theriogenology 7, 126-136

332 Palma Sirisathien S, Hernandez-Fonseca HJ and BG Brackett (2003) Influences of epidermal growth factor and insulin-like growth factor-1 on bovine blastocyst development in vitro Anim Reprod Sci 77, 21-32 Sjoblom C, Roberts CT, Wikland M and SA Robertson (2005) Granulocyte-macrophage colonystimulating factor alleviates adverse consequences of embryo culture on fetal growth trajectory and placental Morphogenesis Endocrinology 146, 2142-253 Slotte H, Akerlof E and A Pousette (1993) Separation of human spermatozoa with hyaluronic acid induces, and Percoll inhibits, the acrosome reaction Int J Androl 16, 349 354 Solano R, de Armas R, Pupo CA y FO Castro (1994) Short term preservation of intrafolicular oocytes at 4 C Theriogenology 41, 299 Spicer LJ, P Alvarez, TM Prado, GL Morgan y TD Hamilton (2000) Effects of intraovarian infusion of insulin-like growth factor-i on ovarian follicular function in cattle Dom Anim Endocrinol 18, 265-278 Spicer LJ, E Alpisar and SE Echterkamp (1993) Effects of insulin-like growth factor I, and gonadotropins on bovine granulosa cell proliferation, progesterone production, estradiol production and (or) insulin-like growth factor I production in vitro J Anim Sci 71, 1232-1241 Spicer LJ and CS Chamberlain (2000) Production of insulin-like growth factor-i by granulosa cells but not thecal cells is hormonally responsive in cattle J Anim Sci 78, 2919-2926 Spicer LJ, SE Echterkamp, DF Canning y JM Hammond (1988) Relationship betwenn concentrations of immunoreactive insulin-like growth factor-i in follicular fluid and various biochemical markers of differentiation in bovine antral follicles Bio Reprod 39, 573-580 Steinhardt RA and JM Alderton (1982) Calmodulin confers calcium sensitivity on secretory exocytosis Nature 295, 154 155 Steeves TE and DK Gardner (1999) Metabolism of glucose, pyruvate, and glutamine during the maturation of oocytes derived from prepubertal and adult cows Mol Reprod Dev 54, 92 101 Stevenson BR and BL Paul (1989) The molecular contituents of intercellular junctions Cur Opin Cell Biol 1, 884-891 Stice SL, Strelchenko NS, Keefer CL, L Matthews (1996) Pluripotent bovine embryonic cell lines direct embryonic development following nuclear transfer Biol Reprod 54, 100 110 Stranzinger G und S Paufler (1968) Vergleichende Untersuchungen über Spermadichten-messungen mit der Thoma Kammer Neu, dem Eppendorf-Photometer und dem Coulter Counter beim Bullenejakulaten Zuchthygiene 3, 191-200 Stoddart NR, Wild AE and TP Fleming (1996) Stimulation of development in vitro by plateletactivating factor receptor ligands released by mouse preimplantation embryos J Reprod Fert 108, 47 53 Stojkovic M, Westesen K, Zakhartchenko V, Stojkovic P, Boxhammer K and E Wolf (1999) Coenzym Q (10) in submicron-sized dispersion improves development, hatching, cell proliferation, and adenosine triphosphate content of in vitro-produced bovine embryos Biol Reprod 61, 541-547 Stokes PJ, Abeydeera LR, HJ Leese (2005) Development of porcine embryos in vivo and in vitro; evidence for embryo 'cross talk' in vitro Dev Biol 284, 62-71 Sun QY (2003) Cellular and molecular mechanisms leading to cortical reaction and polyspermy block in mammalian eggs Microsc Res Tech 61, 342 348 Sun QY and H Schatten (2006) Regulation of dynamic events by microfilaments during oocyte maturation and fertilization Reproduction 131, 193-205 Sun QY, Wang WH, Hosoe M, Taniguchi T, Chen DY and Y Shioya (1997) Activation of protein kinase C induces cortical granule exocytosis in a Ca2+-independent manner, but not the resumption of cell cycle in porcine eggs Dev Growth Differ 39, 523 529 Sunderland SJ, PG Knight, MP Boland, JF Roche, JJ Ireland (1996) Alterations in Intrafollicular Levels of Different Molecular-Mass Forms of Inhibin During Development of Follicular-Phase and Luteal-Phase Dominant Follicles in Heifers Biol of Reprod 54, 453-462 Süss U and V Madison (1993) Morphology and meiosis of bovine oocytes taken from ovaries collected after slaughter Experientia 39, 674 Süss U and G Stranzinger (1987) Chracterization of in vitro matured and fertilized bovine oocytes Symposium on Biotechnology in animal breeding 11-14, Japanese Center, Berlin Süss U, J Kassner, K Wüthurich and G Stranzinger (1990) Cumulus expansion, in vitro fertilization and embryonic development after in vitro maturation of bovine oocytes in the presence of follicle stimulating hormone Reprod Dom Anim 25, 3-13 Süss U, K Wüthurich and G Stranzinger (1988) Chromosome configuration and time sequence of the first meiotic division in bovine oocytes matured in vitro Biol Reprod 38, 871-880