How To Know What Happens When An Ovocito Is Killed

Transcription

1 Punción folicular 289 PUNCIÓN FOLICULAR (OVUM PICK-UP, OPU) EN LA VACA Peter Bols Introducción Aunque un objetivo de interés es conocer los factores que afectan la recolección post-mortem del ovocito, la punción de los folículos ováricos inmaduros es particularmente interesante desde el punto de vista biotecnológico, cuando se lleva a cabo en animales donantes vivos. En ese sentido, los ovocitos obtenidos son destinados a la producción in vitro de embriones durante un prolongado período. Lo cual obviamente no es posible con donantes faenadas o sacrificadas. La recolección de ovocitos por medio de la punción folicular de hembras vivas también permite la modulación endocrina de la actividad ovárica de la donante para influir sobre la cantidad y calidad de los complejos ovocito cumulus (COC) recuperados. Mientras la recuperación de los ovocitos, después de la muerte, se hace simplemente aspirando los folículos ováricos con una jeringa y una aguja, es diferente cuando se aspiran los folículos inmaduros directamente del ovario en una donante viva. La literatura es bastante escasa sobre los efectos del procedimiento de aspiración sobre el número, la morfología y la capacidad de desarrollo subsecuente del complejo recolectado. Existen diferencias hipotéticas básicas entre la aspiración post-mortem e in vivo. En primer lugar, la manipulación transrectal del ovario es necesaria para la recuperación del ovocito en la hembra donante viva, por medio de la visualización del folículo en forma indirecta, a través de laparoscopia, conocida también como imágen de ultrasonido (imaging ultrasound), ver más abajo. En contraste con ello, cuando se puncionan los folículos de ovarios de matadero, el operador puede examinar el ovario directa y completamente. Así, un folículo específico puede escogerse y puede puncionarse bajo control visual directo. Segundo, se puede esperar un efecto mecánico diferente que puede desempeñar un rol importante, mientras el folíuclo es puncionado in vitro con una jeringa y aguja, comparado con la aspiración folicular in vivo, con una presión de vacío de aspiración ajustable, como fue demostrado por Hashimoto y col. (1999a). Finalmente, el uso de hembras donantes vivas permite obtener información adicional sobre el momento de recolección del ovocito, como también conocer los valores endocrinos de la sangre, para ganar más información sobre el estado metabólico de la donante y su posible efecto sobre la calidad del ovocito. Los factores principales que controlan el resultado, con respecto al número y calidad de los ovocitos recuperados al puncionar los folículos inmaduros por medio de OPU en la vaca, pueden ser divididos en 2 categorías principales (Bols, 1997, 2005). La primera categoría incluye los aspectos más técnicos: el procedimiento de aspiración, tipo de aguja y diámetros, más la posible influencia de la presión de vacío de aspiración sobre la morfología de los complejos (COCs). Para documentar este aspecto técnico, se llevaron a cabo varios estudios, examinando el desarrollo y probando la aguja del dispositivo de OPU, que se comentará más adelante. Una segunda categoría puede describirse como aquellos factores biológicos, que incluyen, entre muchos otros, el estímulo hormonal previo a la punción folícular, el momento en que se lleva a cabo el procedimiento en relación con el estado puntual del ciclo estral, la edad, la raza y la condición fisiológica de la donante. De hecho, la recolección de los ovocytos de donantes vivas debe ser considerada como un proceso dinámico. Cuando los ovocitos son recolectados luego de un estímulo hormonal externo, debe esperarse una influencia consecuente sobre la recolección y la calidad de los ovocitos. El capítulo presentará una descripción actualizada de la metodología de la aspiración folicular, el equipamiento necesario y los resultados de la recolección de los ovocitos en hembras donantes vivas.

2 290 Bols Bibliografía Aerts JMJ, Leroy JLMR, Bols PEJ (2004) The use of an endovaginal ultrasound micro-convex array transducer adapted for transvaginal oocyte retrieval in the cow Proceedings of the ICAR meeting, Porto Seguro, Brazil, Vol 2, 435 Aerts JMJ, Oste M, PEJ Bols (2005) Development and practical applications of a method for repeated transvaginal, ultrasound-guided biopsy collection of the bovine ovary Theriogenology, Arlotto TM, Leibfried-Rutledge ML, NL First Size (1990) distribution and meiotic competence of bovine primary oocytes from two locations in the ovary Theriogenology, 188 Arlotto T, Schwartz JL, First NL, ML Leibfried-Rutledge (1995) Aspects of follicle and oocyte stage that affect in vitro maturation and development of bovine oocytes Theriogenology, Armstrong DT, Irvine BJ, Earl CR, McLean D, RF Seamark (1994) Gonadotrophin stimulation regimens for follicular aspiration and in vitro embryo production from calf oocytes Theriogenology, Argov N, Arav A, D Sklan (2004) Number of oocytes obtained from cows by OPU in aerly, but not late lactation increased with plasma insulin and estradiol concentrations and expression of mrna of the FSH receptor in granulosa cells Theriogenology, Assey RJ, Hyttel P, Greve T, B Purwantara (1994) Oocyte morphology in dominant and subordinate follicles Mol Reprod Dev, Baltussen RMWJ, Vos PLAM, Pieterse MC, de Loos FAM, Bevers MM, SJ Dieleman (1992) Transvaginal ultrasound-guided follicle puncture in PMSG/PG treated cows: a comparison between three puncture needles Proceedings of the 12th International Congress on Animal Reproduction, Baracaldo MI, Martinez MF, Adams GP, RJ Mapletoft (2000) Superovulatory response following transvaginal follicle ablation in cattle Theriogenology, Baruselli PS, de Sà Filho MF, Martins CM, Nasser LF, Nogueira MFG, Barros CM, GA Bo (2005) Superovulation and embryotransfer in Bos indicus cattle Proceddings of the XIX Reunião Annual da Sociedade Brasileira de Tecnologia de Embriões (SBTE), august, Brazil Becker F, Kanitz W, Nürnberg G, Kurth J, M Spitschak (1996) Comparison of repeated transvaginal ovum pick-up in heifers by ultrasonographic and endoscopic instruments Theriogenology, Bergfelt DR, Lightfoot KC, GP Adams (1994) Ovarian dynamics following ultrasound-guided transvaginal follicle ablation in cyclic heifers Theriogenology, 161 Blondin P, Sirard MA (1995) Oocyte and follicular morphology as determining characteristics for developmental competence in bovine oocytes Mol Reprod Dev, Bols PEJ, Vandenheede JMM, Van Soom A, A de Kruif (1995) Transvaginal ovum pick-up (OPU) in the cow: a new disposable needle guidance system Theriogenology, Bols PEJ, Van Soom A, Ysebaert MT, Vandenheede JMM, de A (1996a) Kruif Effects of aspiration vacuum and needle diameter on cumulus oocyte complex morphology and developmental capacity of bovine oocytes Theriogenology, Bols PEJ, Van Soom A, A de Kruif (1996b) Gebruik van de transvaginale Ovum Pick-Up (OPU) techniek: geboorte van de eerste OPU kalveren in België (Use of transvaginal oocyte pick-up: first OPU calves bom in Belgium) Vlaams Diergeneeskundig Tijdschrift 65, Bols PEJ (1997) Transvaginal Ovum Pick-Up in the Cow: Technical and Biological Modifications PhD thesis University of Ghent, Ghent, Belgium Bols PEJ, Ysebaert MT, Van Soom A, de Kruif A (1997),Effects of needle tip bevel and aspiration procedure on tbc morphology and developmental capacity of bovine compact cumulus oocyte complexes Theriogenology

3 Punción folicular 291 Bols PEJ, Ysebaert MT, Lein A, Coryn M, Van Soom A, de A Kruif (1998) Effects of long term treatment with bovine somatotropin on follicular dynamics and subsequent oocyte and blastocyst yield during an OPU-IVF program Theriogenology, Bols PEJ, Taneja M, Van de Velde A, Riesen J, Schreiber D, Echelard Y, Ziomek C, X Yang (1999), Pregnancies from prepubertal heifers following repeated oocyte collection and IVF between 6 to 12 months of age Theriogenology 298 Bols PEJ, Van Holder T, Leroy JLMR, Aerts JMJ, A Van Soom (2004a) Ultrasound-guided transvaginal injection of a low dose of FSH-LH into the bovine ovary as an alternative way to stimulate follicular growth: preliminary results Reprod Fert Dev, Bols PEJ, Leroy JLMR, Vanholder T, A Van Soom A (2004b) comparison of a mechanical sector and a linear array transducer for ultrasound-guided transvaginal oocyte retrieval (OPU) in the cow Theriogenology, Bols PEJ (2005) Puncture of immature ovarian follicles in bovine assisted reproduction Verhandelingen van de Koninklijke Academie voor Geneeskunde van België, LXVII, 3, Bordignon V, Morin N, Durocher J, Bousquet D, LC Smith (1997) GnRH improves the recovery rate and the in vitro developmental competence of oocytes obtained by transvaginal follicular aspiration from superstimulated heifers Theriogenology, Bousquet D, Milovanov C, Bell JC, Durocher J, LC Smith (1995) Nuclear and cytoplasmic maturation of oocytes aspirated from large follicles in superovulated heifers Theriogenology, 172 Bousquet D, Twagiramungu H, Morin N, Brisson C, Carboneau G, J Durocher (1999) In vitro embryo production in the cow: an effective alternative to the conventional embryo production approach Theriogenology, Boyd JS, SN Omran (1991) Diagnostic ultrasonography of the bovine female reproductive tract Vet Rec Brackett RG, Bousquet D, Boice ML, Donawick WJ, Evans JF, MA Dressel (1982) Normal development following in vitro fertilization in the cow Biol Reprod, Brogliatti GM, Swan CD, GP Adams (1995)Transvaginal ultrasound-guided oocyte collection in 10 to 16 weeks of age calves Theriogenology, 177 Bungartz L, Lucas-Hahn A, Rath D, H Niemann (1995)Collection of oocytes from cattle via follicular aspiration aided by ultrasound with or without gonadotropin pretreatment and in different reproductive stages Theriogenology, Carlin SD, Garst AS, Tarraf CG, Bailey TL, McGilliard ML, Gibbons JR, Ahmadzadeh A, FC Gwazdauskas (1999) Effects of ultrasound guided OPU on oocyte recovery and hormonal profiles before and after GnRH treatment Theriogenology, Carolan C, Monaghan P, Gallagher M, I Gordon (1993) Developmental capacity of unselected bovine oocytes recovered from individual pairs of ovaries by surface dissection Proceedings of the 9th Reunion AETE Lyon, 180 Callesen H, Greve T, F Christensen (1987) Ultrasonically guided aspiration of bovine follicular oocytes Theriogenology, 217 Chaubal SA, Molina JA, Ohlrichs CL, Ferre LB, Faber DC, Bols PEJ, Riesen JW, Tian X, X Yang (2006) Comparison of different transvaginal ovum pick-up protocols to optimize oocyte retrieval and embryo production over a 10-week period in cows Theriogenology, Chian RC, K Niwa (1994)Effect of cumulus cells present during different periods of culture on maturation in vitro of bovine oocytes Theriogenology, 176 Chohan KR, AG Hunter (2004) In vitro maturation, fertilization and early cleavage rates of bovine fetal oocytes Theriogenology,

4 292 Bols Cohen J, Debache C, Pez JP, Junca AM, P Cohen-Bacrie (1986) Transvaginal sonographically controlled ovarian puncture for oocyte retrieval for in vitro fertilisation J In Vitro Fertil Embryo Transfer, Cox JF, Hormazabal J, A Santa Maria (1993) Effect of the cumulus on in vitro fertilization of bovine matured oocytes Theriogenology, de Armas R, Solano R, Pupo CA, Aguilar A, Aguirre A, Riego E, FO Castro (1994) Effect of the donor oocyte breed on in vitro fertilization results in cattle Theriogenology, 186 De La Sota RL, Lucy MC, Staples CR, WW Thatcher (1993) Effects of recombinant bovine somatotropin (Sometribove) on ovarian function in lactating and non-lactating dairy cows J Dairy Sci, Dellenbach P, Nisand I, Moreau L, Feger B, Plumere C, Gerlinger P, Brun B, Y Rumpler (1984) Transvaginal sonographically controlled ovarian follicle puncture for egg retrieval Lancet, 1467 de Loos F, Van Vliet C, Van Maurik P, ThAM Kruip (1989) Morphology of immature bovine oocytes Gamete Research,, de Loos F (1990) Morphology of the bovine oocyte in relation to its maturation PhD Thesis, University of Utrecht, The Netherlands de Loos FAM, Bevers MM, Dieleman SJ, ThAM Kruip (1991a) Follicular and oocyte maturation in cows treated for superovulation Theriogenology, de Loos FAM, Bevers MM, Dieleman SJ, ThAM Kruip (1991b) Morphology of preovulatory bovine follicles as related to oocyte maturation Theriogenology, de Loos FAM, Maurik P, van Beneden T, ThAM Kruip (1992)Structural aspects of bovine oocyte maturation in vitro Mol Reprod Dev, Donnay I, De Roover R, Van Langendonck A, Massip A, F Dessy (1997)Overall efficiency of an experimental OPU programme in cattle Theriogenology, 156 De Roover R, Genicot G, Leonard S, Bols P, F Dessy (2005a)Ovum Pick-up and in vitro embryo production in cows superstimulated with an individually adapted superstimulation protocol Anim Reprod Sci, De Roover R, Bols PEJ, Genicot G, Ch Hanzen (2005b )Characterisation of low, medium and high responders following FSH stimulation prior to ultrasound-guided transvaginal oocyte retrieval in cows Theriogenology, De Ruigh L, Mullaart E, AM van Wagtendonk-de Leeuw (2000) The effect of FSH stimulation prior to ovum pick-up on oocyte and embryo yield Theriogenology, 349 Dominguez MM (1995) Effects of body condition, reproductive status and breed on follicular population and oocyte quality in cows Theriogenology, Duby RT, Damiani P, Looney CR, Fissore RA, JM Robl (1996)Prepuberal calves as oocyte donors: promises and problems Theriogenology, Edmondson AJ, Fissore RA, Pashen RL, RH Bondurant (1986) The use of ultrasonography for the study of the bovine reproductive tract: I Normal and pathological ovarian structures Anim Reprod Sci, Fair T, Hyttel P, T Greve (1995) Bovine oocyte diameter in relation to maturational competence and transcriptional activity Mol Reprod Dev, Feichtinger W, P Kemeter (1986) Transvaginal sector scan sonography for needle guided transvaginal follicle aspiration and other applications in gynecologic routine and research Fertil Steril, Fortune JE, J Sirois (1989) The use of ultrasonography to study the regulation of follicular development in cattle and horses In: Follicular Development and the Ovulatory Response pp Eds A Tsafriri and N Dekel Serono Symposia, Rome Fry RC, Simpson TL, Squires TI, Parr RA, RM Damanik (1994) Factors affecting transvaginal oocyte pick-up in heifers Theriogenology, 197

5 Punción folicular 293 Fukui Y, Y Sakuma (1980) Maturation of bovine oocytes cultured in vitro: relation to ovarian activity, follicular size and the presence or absence of cumulus cells Biol Reprod, Galli C, Crotti G, Notari C, Turini P, Duchi R, G Lazzari (2001) Embryo production by ovum pick up from live donors Theriogenology, Garcia A and M Salaheddine (1998)Effects of repeated ultrasound-guided transvaginal follicular aspiration on bovine oocyte recovery and subsequent follicular development Theriogenology, Gibbons JR, Beal WE, Krisher RL, Faber EG, Pearson RE, FC Gwazdauskas (1994) Effects of once versus twice weekly transvaginal follicular aspiration on bovine oocyte recovery and embryo development Theriogenology, 206 Gong JG, Bramley T, R Webb (1991) The effect of recombinant bovine somatotropin on ovarian function in heifers: follicular populations and peripheral hormones Biol Reprod, Gong JG, Bramley TA, R Webb (1993) The effect of recombinant bovine somatotropin on ovarian follicular growth and development in heifers J Reprod Fert, Goodhand KL, Watt RG, Staines ME, Hutchinson JSM, PJ Broadbent (1999) In vivo oocyte recovery and in vitro embryo production from bovine donors aspirated at different frequencies or following FSH treatment Theriogenology, Goodhand KL, Staines ME, Hutchinson JSM, PJ Broadbent (2000) In vivo oocyte recovery and in vitro embryo production from bovine oocyte donors treated with progestagen, oestradiol and FSH Anim Reprod Sci, Gordon I (2003) Laboratory Production of Cattle Embryos, Second Edition CAB International University Press, Cambridge, Goto K, Kajihara Y, Kosaka S, Koba M, Nakanishi Y, K Ogawa (1988) Pregnancies after co-culture of cumulus cells with bovine embryos derived from in vitro fertilization of in vitro matured follicular oocytes J Reprod Fertil, Greve T, Callesen H, P Hyttel (1989) Follicular correlates with in vitro fertilization in cattle J Reprod Fert, Hamano S, M Kuwayama (1993) In vitro fertilization and development of bovine oocytes recovered from the ovaries of individual donors: a comparison between the cutting and aspiration method Theriogenology, Hanada A, Enya Y, T Suzuki (1986) Birth of calves by non-surgical transfer of in vitro fertilized embryos obtained from oocytes matured in vitro Jap Journal of Anim Reprod, 208 Hashimoto S, Takakura R, Kishi M, Sudo T, Minami N, M Yamada (1999a) Ultrasound-guided follicle aspiration: the collection of bovine cumulus-oocyte complexes from ovaries of slaughtered or live cows Theriogenology, Hashimoto S, Takakura R, Minami N, M Yamada (1999b) Ultrasound-guided follicle aspiration: effect of the frequency of a linear transvaginal probe on the collection of bovine oocytes Theriogenology, Hasler JF (1992) Current status and potential of embryo transfer and reproductive technology in dairy cattle J Dairy Sci, Hasler JF, Henderson WB, Hurtgen PJ, Jin ZO, McCauly AD, Mower SA, Neely B, Shuey LS, Stokes JE, SA Trimmer (1995) Production, freezing and transfer of bovine IVF embryos and subsequent calving results Theriogenology, Hasler JF (1998) The current status of oocyte recovery, in vitro embryo production, and embryo transfer in domestic animals, with an emphasis on the bovine J Anim Sci, Hawk HW, RJ Wall (1994) Improved yield of bovine blastocysts from in-vitro produced oocytes 1 Selection of oocytes and zygotes Theriogenology,

6 294 Bols Hazeleger NL, Hill DJ, Stubbings RB, JS Walton (1995) Relationship of morphology and follicular fluid environment of bovine oocytes to their developmental potential in vitro Theriogenology, Hendriksen PJM, Steenweg WNM, Harkema JC, Merton JS, Bevers MM, Vos PLAM, SJ Dieleman (2004) Effect of different stages of the follicular wave on in vitro developmental competence of bovine oocytes Theriogenology, Hinrichs K, Kenney DF, RM Kenney (1990) Aspiration of oocytes from mature and immature preovulatory follicles in the mare Theriogenology, Holland EJ, Bindon BM, Piper LR, Thimonier J, Cornish KA, HM Radford (1981) Endoscopy in cattle: techniques for ovarian examination by the paralumbar and midventral routes Anim Reprod Sci, Imai K, Kobayashi S, Tsujino T, Shin-noh M, Goto Y, Kaneyama K, T Kojima (2000) Effect of the frequency of ovum pick-up intervals on follicle number, oocyte recoverey and embryo production rates in cattle Theriogenology, 359 Izadyar F, Colenbrander B, MM Bevers (1996) Growth hormone stimulates in vitro bovine oocyte maturation and subsequent embryonic development Theriogenology, 279 Jain GC, Das GK, Solanki VS, VN Tripathi (1995) Comparative efficiency of three different collection techniques for oocyte retrieval in buffaloes Theriogenology, 240 Katska L (1984a) Comparison of two methods for recovery of ovarian oocytes from slaughter cattle Anim Reprod Sci, Katska L, Z Smorag (1984) Number and quality of oocytes in relation to age of cattle Anim Reprod Sci, Konishi M, Aoyagi Y, Takedomi T, Itakura H, Itoh T, S Yazawa (1996) Presence of granulosa cells during oocyte maturation improved in vitro development of IVM-IVF bovine oocytes that were collected by ultrasound-guided transvaginal aspiration Theriogenology, Kot K, Anderson LE, Wiltbank MC, OJ Ginther (1999) Transvaginal, ultrasound-guided biopsy of the corpus luteum in cattle Theriogenology 52, Kruip ThAM, SJ Dieleman (1982) Macroscopic classification of bovine follicles and its validation by micromorphological and steroid biochemical procedures Reprod Nutr Dev, Kruip ThAM, SJ Dieleman (1985) Steroid hormone concentration in the fluid of bovine follicles relative to size, quality and stage of the oestrus cycle Theriogenology, Kruip ThAM, Boni R, Wurth YA, Roelofsen MWM, MC Pieterse (1994) Potential use of ovum pick-up for embryoproduction and breeding in cattle Theriogenology, Lambert RD, Bernard C Rioux JE, Béland R, D Amours D, A Montreuil (1983) Endoscopy in cattle by the paralumbar route: technique for ovarian examination and follicular aspiration Theriogenology, Laurincik J, Kroslak P, Hyttel P, Pivko J, AV Sirotkin (1992) Bovine cumulus expansion and coronaoocyte disconnection during culture in vitro Reprod Nutr Dev, Lenz S, Leeton J, P Renou (1987) Transvaginal recovery of oocytes for in vitro fertilization using vaginal ultrasound J In Vitro Fertil Embryo Transfer, Leroy JLMR, Vanholder T, Delanghe JR, Opsomer G, Van Soom A, Bols PEJ, Dewulf J, A de Kruif (2004) Metabolic changes in follicular fluid of the dominant follicle in high-yielding dairy cows early post partum Theriogenology, Ligtvoet CH, Bom N, WJ Gussenhove (1989) Technical principles of ultrasound in: Taverne, MAM and Willemse, AH (eds) Diagnostic Ultrasound and Animal Reproduction, Kluwer Medical Publishers, Dordrecht, 1-9 Lonergan P, Monaghan P, Rizos D, Boland MP, I Gordon (1994) Effect of follicle size on bovine oocyte quality and developmental competence following maturation, fertilization and culture in vitro Mol Reprod Dev, 48-53

7 Punción folicular 295 Looney CR, Lindsey BR, Gonseth CL, Johnson DL Commercial aspects of oocyte retrieval and in vitro fertilization (IVF) for embryo production in problem cows Theriogenology 1994(), Lopata A, Johnston IWH, Leeton JF, Muchnicki D, McTalbot J, C Wood (1974) Collection of human oocytes at laparoscopy and laparotomy Fertil Steril, Lopez Ruiz L, Alvarez N, Nunez I, Montes I, Solano R, Fuentes D, Pedroso R, Palma GA, G Brem (1996) Effect of body condition on the developmental competence of IVM/IVF bovine oocytes Theriogenology, 292 Lu KH, Gordon 1, Chen HB, H McGovern (1987) In vitro culture of early bovine embryos derived from in vitro fertilization of follicular oocytes matured in vitro Proceedings of the 3th Reunion AETE, Lyon, 70 Lu KH (1990) Studies related to the in vitro maturation and fertilization of ovarian oocytes in cattle PhD Thesis, National University of Ireland, Dublin Lu KB, Shi DS, Jiang HS, Goulding D, Boland MP, JF Roche (1991) Comparison of the developmental capacity of bovine oocytes from superovulated and non stimulated heifers Theriogenology, 234 Lu KH, 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), Machatkova M, Jokesova E, Petilikova J, V Dvoracek (1995) Developmental competence of bovine embryos derived from oocytes collected at various stages of the estrous cycle Theriogenology, Machatkova M, Krausova K, Jokesova E, M Tomanek (2004) Developmental competence of bovine oocytes: effects of follicle size and the phase of follicular wave on in vitro embryo production Theriogenology, Madison V, Avery B, T Greve (1992) Selection of immature bovine oocytes for developmental potential in vitro Anim Reprod Sci, 1-11 McEvoy TG, Thompson H, Dolman DF, Watt RG, Reis A, ME Stains (2001) Effects of epidural injection and transvaginal aspiration of ovarian follicles in heifers used repeatedly for ultrasoundguided retrieval of ova and embryo production Vet Rec, Meintjens M, Bellow MS, Broussard JR, Paul JB, RA Godke (1995) Transvaginal aspiration of oocytes from hormone-treated pregnant beef cattle for in vitro fertilization J Anim Sci, Merton JS, de Roos APW, Mullaart E, de Ruigh L, Kaal L, Vos PLAM, SJ Dieleman (2003) Factors affecting oocyte quality and quantity in commercial application of embryo technologies in the cattle breeding industry Theriogenology, Moreno JF, Flores-Foxworth G, Westhusin M, DC Kraemer (1993) Influence of pregnancy and presence of a CL on quantity and quality of bovine oocytes obtained from ovarian follicles aspirated post mortem Theriogenology, 271 Motlik J, Fulka J Factors affecting meiotic competence in pig oocytes Theriogenology (1986) 25, Murakami M, Perez O, Ferguson E, Behboodi E, Denniston RS, RA Godke (2003) Use of in vivorecovered oocytes and adult somatic cells from the same donor for nuclear transfer in cattle Vet Rec, Oropeza A, Wrenzycki C, Herrmann D, Hadeler KG, H Niemann (2004) Improvement of the developmental capacity of oocytes from prepubertal cattle by intraovarian insulin-like growth factor- I application Biol Reprod 70, Palma GA, Tortonese DJ, F Sinowatz (2001) Developmental capacity in vitro of prepubertal oocytes Anat Histol Embryol 30, Paul JB, Looney CR, Lindsay BR, RA Godke (1995) Gonadotropin stimulation of cattle donors at estrus for transvaginal oocyte collection Theriogenology, 294

8 296 Bols Pavlok A, Lucas-Hahn A, H Niemann (1992) Fertilization and developmental competence of bovine oocytes derived from different categories of antral follicles Mol Reprod Dev, Pavlok A, Koutecka L, Krejci P, Slavik T, Cerman J, Slaba J, D Dom (1996) Effect of recombinant bovine somatotropin on follicular growth and quality of oocytes in cattle Anim Reprod Sci, Petyim S, Båge R, Forsberg M, Rodriguez-Martinez H, B Larsson (2000) The effect of repeated follicular puncture on ovarian function in dairy heifers J Vet Med A, Petyim S, Båge R, Forsberg M, Rodriguez-Martinez H, B Larsson (2001) Effects of repeated follicular punctures on ovarian morphology and endocrine parameters in dairy heifers J Vet Med A, Petyim S (2002) Ovum Pick-Up in Dairy Heifers: effects on ovary, ovarian function and animal wellbeing Doctoral thesis, Swedish University of Agricultural Sciences, Uppsala Pierson RA, OJ Ginther (1987) Ultrasonographic appearance of the bovine uterus during the estrous cycle J Am Med Vet Assoc, Pieterse MC, Kappen KA, Kruip ThAM, MAM Taverne (1988) Aspiration of bovine oocytes during transvaginal ultrasound scanning of the ovaries Theriogenology, Pieterse MC, Vos PLAM, Kruip ThAM, Wurth YA, van Beneden ThH, Willemse AH, MAM Taveme (199la) Transvaginal Ultrasound guided follicular aspiration of bovine oocytes Theriogenology, Pieterse MC, Vos PLAM, Kruip ThAM, Willemse AH, MAM Taverne (1991b) Characteristics of bovine estrous cycles during repeated transvaginal, ultrasound-guided puncturing of follicles for ovum pick-up Theriogenology, Pieterse MC, Vos PLAM, Kruip ThAM, Wurth YA, van Beneden ThH, Willemse AH, MAM (1992) Taverne Repeated transvaginal ultrasound-guided ovum pick-up in ECG-treated cows Theriogenology, 273 Price CA, Carrière PD, Bhatia B, NP Groome (1995) Comparison of hormonal and histological changes during follicular growth, as measured by ultrasonography, in cattle J Reprod Fertil, Rath D (1993) Current status of ultrasound-guided retrieval of bovine oocytes Embryo Transfer Newsl, Reeves IJ, Rantanen NW, M Hauser (1984) Transrectal real-time ultrasound scanning of the cow reproductive tract Theriogenology, Reichenbach MD, Wiebke NH, Mödl J, Zhu J, G Brem (1994) Laparoscopy through the vaginal fornix of cows for the repeated aspiration of follicular oocytes Vet Rec, Reinders JMC, AM Van Wagtendonck-de Leeuw (1996) Improvement of a MOET program by addition of in vitro production of embryos after ovum pick-up from pregnant donor heifers Theriogenology, 354 Revel F, Mermillod P, Peynot N, Renard JP, Y Heyman (1995) Low developmental capacity of in vitro matured and fertilized oocytes from calves compared with that of cows J Reprod Fertil, Rick G, Hadeler KG, Lemme E, Lucas-Hahn A, Rath D, Schindler L, H Niemann (1996) Long-term ultrasound guided ovum pick-up in heifers from 6 to 15 months of age Theriogenology, 356 Ryan DP, Blakewood EG, Swanson WF, Rodriguez H, RA Godke (1993) Using hormone-treated pregnant cows as a potential source of oocytes for in vitro fertilization Theriogenology, Santl B, Wenigerkind H, Schernthaner W, Mödl J, Stojkovic M, Prelle K, Holtz W, Brem G, E Wolf (1998) Comparison of ultrasound-guided vs laparoscopic transvaginal ovum pick-up (OPU) in simmental heifers Theriogenology, Sasamoto Y, Sakaguchi M, Katagiri S, Yamada Y, H Takahashi (2003) The effects of twisting and type of aspiration needle on the efficiency of transvaginal ultrasound guided ovum pick up in cattle J Vet Med Sci65, Schellander K, Fayrer-Hosken R, Keefer C, Brown L, Malter H, Mcbride C, B Brackett (1989) In vitro fertilisation of bovine follicular oocytes recovered by laparoscopy Theriogenology,

9 Punción folicular 297 Scott CA, Robertson L, de Moura RTD, Paterson C, JS Boyd (1994) Technical aspects of transvaginal ultrasound-guided follicular aspiration in cows Vet Rec, Seneda MM, Esper CS, Garcia JM, de Oliveira JA, R Vantini (2001) Relationship between follicle size and ultrasound-guided transvaginal oocyte recovery Anim Reprod Sci, Sheldon IM, H Dobson (2003) Reproductive challenges facing the cattle industry at the beginning of the 21 st century Reproduction Supplement, 1-13 Shioya Y, Kuwayama M, Fukushima M, Iwasaki S, A Hanada (1988) In vitro fertilization and cleavage capability of bovine follicular oocytes classified by cumulus cells and matured in vitro Theriogenology, Simon L, Bungartz L, Rath D, H Niemann (1993) Repeated bovine oocyte collection by means of a permanently rinsed ultrasound guided aspiration unit Theriogenology, 312 Singh J, Adams GP, RA Pierson (2003) Promise of new imaging technologies for assessing ovarian function Anim Rep Sci, Sirois J, JE Fortune (1988) Ovarian follicular dynamics during the estrous cycle in heifers monitored by real time ultrasonography Biol Reprod, Sosnowski J, Switonski M, Lechniak D, K Molinski (1996) Cytogenetic evaluation of in vitro matured bovine oocytes collected from ovaries of individual donors Theriogenology, Staigmiller RB, RM Moor (1984)Effect of follicle cells on the maturation and developmental competence of ovine oocytes matured outside the follicle Gamete Research, Stubbings RB, JS Walton (1995) Effect of ultrasonically-guided follicle aspiration on estrous cycle and follicular dynamics in Holstein cows Theriogenology, Takagi Y, Mori K, Takahashi T, Sugawara S, Masaki J Differences in development of bovine oocytes recovered by aspiration or by mincing J Anim Sci 1992(), Taneja M, Bols PEJ, Van de Velde A, Ju J-C, Schreiber D, Tripp MW, Levine H, Echelard Y, Riesen J, X Yang (2000) Developmental competence of juvenile calf oocytes in vitro and in vivo: influence of donor animal variation and repeated gonadotropin stimulation Biol Reprod, Thonon F, Ectors FJ, Delval A, Fontes RS, Touati K, JF Beckers (1993) In vitro maturation, fertilization and development rates of bovine oocytes connected with the reproductive status of the donor Theriogenology, 330 Van Arendonck JAM, A-E Liinamo (2003) Dairy cattle production in Europe Theriogenology, van der Schans A, van der Westerlaken CAJ, de Wit AAC, Eyestone WH, HA de Boer (1991) Ultrasound-guided transvaginal collection of oocytes in the cow Theriogenology, 288 Van Soom A, Van Vlaenderen, Mahmoudzadeh AR, Ysebaert MT, A de Kruif (1994a) Salvage of oocytes from sterile genetically valuable cows, resulting in the birth of a calf Anim Reprod Sci, Van Soom A, Mijten P, Van Vlaenderen I, Van den Branden J, Mahmoudzadeh AR, A de Kruif (1994b) Birth of double-muscled Belgian Blue calves after transfer of in vitro produced embryos into dairy cattle Theriogenology, Van Soom A, Bols PEJ, A de Kruif (1995) Improved results in IVF-treatment of sterility patients by careful selection of bulls and by stimulation of cows with FSH before slaughter Reprod Dom Anim, 67 Van Wagtendonck-de Leeuw AM, Mullaart E, de Roos APW, Merton JS, den Daas JHG, Kemp B, L de Ruigh (2000) Effects of different reproduction techniques: AI, MOET or IVP, on health and welfare of bovine offspring Theriogenology, Vergos E In vitro fertilization and embryo culture in cattle PhD Thesis, 1990, National University of Ireland, Dublin Vos PLAM, de Loos FAM, Pieterse MC, Bevers MM, Taverne MAM, SJ Dieleman (1994) Evaluation of transvaginal ultrasound-guided follicle puncture to collect oocytes and follicular fluids at

10 298 Bols consecutive times relative to the preovulatory LH surge in ecg/pg treated cows Theriogenology, Ward FA, Lonergan P, Enright BP, MP Boland (2000) Factors affecting recoverey and quality of oocytes for bovine embryo production in vitro using ovum pick-up technology Theriogenology, Wikland M, Enk L, Hammarberg K, L Nilsson (1987) Use of vaginal transducer for oocyte retrieval in an IVF/ET program J Clin Ultrasound, Wurth YA, ThAM Kruip (1992) Bovine embryo production in vitro after selection of the follicles and oocytes Proceedings of the 12th International Congress on Animal Reproduction (The Hague), Zhang L, Jiang S, Wozniak PJ, Yang X, RA Godke (1995) Cumulus cell function during bovine oocyte maturation, fertilization and embryo development in vitro Mol Reprod Dev,