Anda di halaman 1dari 8

Biologa de la reproduccin

43

BIOLOGA DE LA REPRODUCCIN
Gustavo A Palma

Introduccin
La capacidad reproductiva es, obviamente, la condicin indispensable de toda especie para
preservarse. En produccin animal tiene un significado central, dado que la cra de animales de
inters productivo basa su existencia comercial en la fertilidad, la reproduccin peridica y sistemtica
de los individuos. En otras palabras, reproduccin eficiente. Por otra parte la reproduccin sexual
confiere a los organismos el beneficio evolutivo del proceso de recombinacin gentica, a diferencia
de la reproduccin asexuada, porque elimina las mutaciones deletreas de la poblacin, segn la
hiptesis de la epistasis sinrgica o reductora de la carga mutacional. Esto es, elimina a aquellos
genotipos con mutaciones mltiples que tengan menos adecuacin (epistasis negativa), como
tambin permite incorporar caractersticas deseadas.
A fin de comprender mejor las particularidades reproductivas de la mayora de las especies
domsticas y la humana, las posibilidades biotecnolgicas, sus limitaciones y las alteraciones
patolgicas que tienen lugar, es importante considerar la dinmica biolgica poblacional, como
estrategia evolutiva, antes que fuera condicionada por el hombre y el comportamiento social humano.
El crecimiento y tamao de una poblacin est determinado por 2 factores de inters biolgico:
La natalidad (tasa de nacimientos)
La mortalidad (tasa de muertes)
De esta manera la tasa de crecimiento de una poblacin estar determinada por los
nacimientos (b) en relacin con las muertes (d):
r = (b - d), dnde r = la proporcin de crecimiento de poblacin, b = la proporcin del nacimiento
y d = la mortalidad. Los factores que controlan a la poblacin caracterizan ese potencial bitico,
a travs de:
La capacidad de reproducirse (factores intrnsecos) y
la interaccin (o resistencia) al ambiente (factores extrnsecos)
El crecimiento de la poblacin depender de los animales presentes, de la tasa (r, de growth
rate) y la capacidad mxima de crecimiento (k, de carring capacity). Esos son los puntos de partida de
la biologa reproductiva de las poblaciones llamadas k y n, o tambin conocidos como seleccin k y
seleccin n, Fig 1. Las poblaciones n estn constituidas por aquellos individuos de alto potencial
bitico porque que viven en ambientes muy inestables. Con tamao poblacional fluctuante, sufren
grandes oscilaciones de la poblacin, incluso hasta la eliminacin de varias generaciones, aunque
tambin pueden reproducirse con la misma rapidez. Estn ampliamente distribuidos y sacan provecho
rpidamente de las buenas oportunidades ambientales, pudiendo as, reestablecer la poblacin
inicial, lo que los caracteriza de oportunistas. Algunos ejemplos de las poblaciones n son: las
bacterias, los insectos, los peces, el plancton y los roedores.
Por el contrario, la estrategia k de reproduccin corresponde a aquellos individuos que viven en
equilibrio con un ambiente propicio, el cual les ofrece las condiciones ptimas para una vida longeva.
Con un ambiente que asegura a los individuos tantos o ms recursos de los que pueden consumir,
tienen un tamao poblacional estable y responden lentamente a las oportunidades ambientales
(cuadro 1).
Todas las especies animales de inters productivo y la humana pertenecen a la poblacin k. La
estrategia biolgica de la poblacin k establece que la capacidad reproductiva de un individuo no es
una funcin prioritaria y por lo tanto todo lo que atente contra su bienestar, atentar tambin contra la
reproduccin.

44

Palma
Crecimiento poblacional

Nr. de individuos

Tiempo en meses
Fig 1. Poblaciones animales segn sus particularidades reproductivas
Esta estrategia podra brindar una hiptesis biolgica para explicar determinados fenmenos
del comportamiento social humano de porqu, por ejemplo, las sociedades de algunos pases
industrializados con las mejores condiciones de vida, tienen la mayor longevidad, las menores tasas
de mortalidad y una tasa reproductiva negativa. Esto es, la capacidad de la especie humana de
reconocer su condicin de bienestar y proyectarle sus deseos, fantasas y aspiraciones a su futuro,
subordina su reproduccin, disminuyndola.
Cuadro 1. Estrategias reproductivas de las poblaciones k
Extenso tiempo de desarrollo embrionario
Tarda edad de la pubertad
Reproduccin durante la mayor parte de su vida
Extenso perodo de gestacin
Bajo nmero y alto cuidado de la cra de cras
Suponiendo que esta hiptesis fuera aceptable, es posible intuir, entonces, que mientras una
pareja alemana prefiere disfrutar de las buenas condiciones de vida que le ofrece su ambiente sin
reproducirse, para evitar competir luego con su descendencia, un animal sometido a situaciones
ambientales (productivas, reproductivas o biotecnolgicas) que atenten contra su bienestar,
disminuir su eficiencia reproductiva.
No es necesario que las condiciones ambientales pongan en peligro la vida del individuo para
que disminuya significativamente la eficiencia reproductiva. El manejo y seleccin inadecuados de los
reproductores que componen los rodeos lecheros, la aplicacin errnea de las biotecnologas con
tratamientos hormonales para la superovulacin en programas de transferencia de embriones o la
sobre-estimulacin ovrica, en programas de la puncin folicular para la produccin in vitro de
embriones, pueden provocar efectos adversos (ver captulo superovulacin y OPU). Tambin permite
comprender por qu los bovinos de razas europeas, que fueron exportados a pases tropicales en la
dcada del 70 no se reprodujeron como lo hacan bajo sus propias condiciones de origen. La
caracterstica de que respondan lentamente a las condiciones ambientales explicara tambin porque
no sirvieron las cmaras especialmente diseadas en Alemania para adaptar, durante semanas, a
toros antes de ser exportados al trpico. Lentamente puede significar en biologa adaptativa
numerosas generaciones. Tambin explica las significativas diferencias en la eficiencia reproductiva
de la especie bufalina, cuando es producida bajo mejores condiciones (ver captulo biologa de la
reproduccin de la hembra bufalina), como es el caso en Argentina y Brasil frente a India.
Se conoce que la desnutricin afecta a la reproduccin, no obstante, no es necesario que el
animal est desnutrido para que no ovule o lo haga sin presentar celo (celo silente), o ste sea corto.

Biologa de la reproduccin

45

Eso explicara por qu animales en buen estado pero carentes, por ejemplo de selenio en la dieta,
tienen inferior produccin de ovocitos y stos de baja calidad.
Cuando se enfoca la posible explicacin fenomenolgica, desde el punto de vista de la
implacable estrategia poblacional k, muchos de los alcances limitados de la biotecnologa de la
reproduccin deberan ser considerados como grandes xitos.
Actualmente las acciones mdicas en la reproduccin se concentran en las siguientes reas:
Comprensin de los fenmenos biolgicos
Estatus ginecolgico (reconocimiento del ciclo estral)
Estatus androlgico (evaluacin de la calidad espermtica)
Interpretacin del diagnstico
Diagnstico y tratamiento de enfermedades
Control del fenmeno reproductivo grupal con el fin de incrementar la eficiencia reproductiva
(control reproductivo y fertilidad de los rodeos), como los tratamientos hormonales
Control o asistencia individual para incrementar o establecer la capacidad reproductiva, como la
reproduccin asistida en las especies humana y animal, con fines teraputicos y econmicos
(inseminacin artificial intravaginal, intrauterina o intratubrica) produccin in vitro de embriones,
produccin de mellizos idnticos, clonacin a travs de transferencia nuclear, etc.
Control de la reproduccin con fines anticonceptivos, para bloquear la eficiencia reproductiva
femenina y masculina
El objetivo del presente captulo es presentar algunos fenmenos reproductivos que permitan
interpretar ms cabalmente las interacciones neuro-hormonales presentadas por Rivera y Callejas
(ver libros rojo y azul en el CD), como tambin las consecuencias biolgicas de la relacin entre
individuos, de stos con el ambiente y el posible efecto de este ltimo sobre la respuesta a los
tratamientos biotecnolgicos. Se presentar el rol de los esteroides en el desarrollo folicular, el control
hormonal y vascular de la regresin luteal, las prostaglandinas intraluteales, el efecto de la nutricin y
sus seales neuropeptdicas sobre la biologa reproductiva.

46

Palma

Bibliografa
Acosta TJ, Yoshizawa N, Ohtani M and A Miyamoto (2002) Local changes in blood flow within the
early and midcycle corpus luteum after prostaglandin F2 injection in the cow Biol Reprod 66, 651
658
Ahima RS and JS Flier (2000) Adipose tissue as an endocrine organ Trends Endocrinol Metab 11,
327332
Amstalden M, Garcia MR, Williams SW, Stanko RL, Nizielski SE, Morrison CD, Keisler DH, GL
Williams (2000) Leptin gene expression, circulating leptin, and luteinizing hormone pulsatility are
acutely responsive to short-term fasting in prepubertal heifers: Relationships to circulating insulin
and insulin-like growth factor I Biol Reprod 63, 127-133
Asselin E, Goff AK, Bergeron H and MA (1996) Fortier Influence of sex steroids on the production of
prostaglandins F2 and E2 and response to oxytocin in cultured epithelial and stromal cells of the
bovine endometrium Biol Reprod 54, 371379
Bah MM, Acosta TJ, Pilawski W, Deptula K, Okuda K, DJ Skarzynski (2006) Role of intraluteal
prostaglandin F(2alpha), progesterone and oxytocin in basal and pulsatile progesterone release
from developing bovine corpus luteum Prostaglandins Other Lipid Mediat 79, 218-29
Barb CR, Kraeling RR, Rampacek GB, Barnett JB (1999) Neuropeptide Y: a possible link between LH
and GH in the gilt J Anim Sci 77, 223
Beach FA (1976) Sexual attractivity, proceptivity, and receptivity in female mammals Horm Behav 7,
105138
Beata C (2001) Appeasing Pheromones in Mammals 26th WSAVA congress proceedings; Vancouver,
British Columbia, Canada August 8-11
Belund H, Lindstom P and I Savic (2006) Brian reponse to putative pheromones in lesbian women
PNAS 1003, 8269-8274
Bensafi M, Brown WM, Khan R, Levenson B and N Sobel (2004) Sniffing human sex-steroid derived
compounds modulates mood, memory and autonomic nervous system function in specific
behavioral contexts Behav Brain Res 152, 11-22
Bensafi M, Brown WM, Tsutsui T, Mainland JD, Johnson BN, Bremner EA, Young N, Mauss I, Ray B,
Gross J, Richards J, Stappen I, Levenson RW and N Sobel (2003) Sex-Steroid Derived
Compounds Induce Sex-Specific Effects on Autonomic Nervous System Function in Humans
Behav Neurosci 117, 1125-1134
Boland MP, Lonergan lP and D OCallaghanz (200l) Effect of nutrition on endocrine parameters,
ovarian physiology, and oocyte and embryo development Theriogenology 55, 1323-1340
Brown JL and A Eklund (1994) Kin recognition and the major histocompatibility complex: an integrative
review Am Nat 14, 435-461
Christiansen K, Behavioural correlates of testosterone In: E Nieschlag and HM Behre (Ed.),
Testosterone. Action-deficiency-substitution (2nd ed.), Springer, Heidelberg (1998), 107142
Claus R (1994): Pheromone. In: Dcke, F. (Ed.): Veterinrmedizinische Endokrinologie Gustav
Fischer Verlag - Jena, Stuttgart, 691712
Claus R, Over R, M Dehnard (1990) Effect of male odour on LH secretion and the induction of
ovulation in seasonally anoestrous goats Anim Reprod Sci 22, 2738
Cohen-Tanoudji J, Einhorn J, JP Signoret (1994) Ram sexual pheromone: first approach of chemical
identification Physiol Behav 56, 955961
Davis JS, Weakland LL, Weiland DA, Farese RV and LA West (1987) Prostaglandin F2 stimulates
phosphatidylinositol 4,5-bisphosphate hydrolysis and mobilizes intracellular Ca2+ in bovine luteal
cells. Proc Natl Acad Sci84, 37283732
Day ML and LH Anderson (1998) Current concepts on the control of puberty in cattle J Anim Sci 76,
115
Diskin MG, Mackey DR, Roche JF, JM Sreenan (2003) Effects of nutrition and metabolic status on
circulating hormones and ovarian follicle development in cattle 78, 345-70
Diskin MG, Murphy JJ and JM Sreenan (2006) Embryo survival in dairy cows managed under pastoral
conditions Anim Reprod Sci 96, 297-311

Biologa de la reproduccin

47

Drummond AE (2006) The role of steroids in follicular growth Reprod Biol Endocrinol 4, 16
Doving KB and D Trotier (1998) Struture and function of the vomeronasal organ J Exp Biol 201, 29132925
Elias CF, Lee C, Kelly J, Aschkenasi C, Ahima RS, Bjrbk C, Flier JS, Saper CB, JK Elmquist
(1999) Leptin differentially regulates NPY and POMC neurons projecting to the lateral hypothalamic
area Neuron 23, 775-786
ESHRE Capri Workshop Group (2006) Nutrition and reproduction in women Hum Reprod Update 12,
193-207
Evans A and F Mulligan (2006) Nutrition and fertility in dairy cattle (In Press) Available online 3 August
Fortune JE, Rivera GM, MY Yang (2004) Follicular development: the role of the follicular
microenvironment in selection of the dominant follicle Anim Reprod Sci 82-83, 109-126
Fukushima N, Hanada R, Teranishi H, Fukue Y, Tachibana T, Ishikawa H, Takeda S, Takeuchi Y,
Fukumoto S, Kangawa K, Nagata K, M Kojima (2005) Ghrelin directly regulates bone formation.
Bone Miner Res 20, 790798
Gazal OS, Leshin LS, Stanko RL, Thomas MG, Keisler DH, Anderson LL, GL Williams (1998)
Gonadotropin-releasing hormone secretion into third-ventricle cerebrospinal fluid of cattle:
correspondence with the tonic and surge release of luteinizing hormone and its tonic inhibition by
suckling and neuropeptide Y Biol Reprod 59, 676683
Gnanapavan S, Kola B, Bustin SA, Morris DG, McGee P, Fairclough P, Bhattacharya S, Carpenter R,
Grossman AB and M Korbonits (2002) The tissue distribution of the mRNA of ghrelin and subtypes
of its receptor, GHS-R, in humans J Clin Endocrinol Metab 87, 29882991
Gelez H and C Fabre-Nys (2004) The "male effect" in sheep and goats: a review of the respective
roles of the two olfactory systems Horm Behav 46, 257-271
Gelez H, Archer E, Chesneau D, Campan R, C Fabre-Nys (2004) Importance of learning in the
response of ewes to male odor Chem Senses 29, 555563
Gelez H, C Fabre-Nys (2006) Role of the olfactory systems and importance of learning in the ewes'
response to rams or their odors Reprod Nutr Dev 46, 401-415
Goff AK (2004) Steroid hormone modulation of prostaglandin secretion in the ruminant endometrium
during the estrous cycle Biol Reprod 71, 11-16
Gong JG (2002) Influence of metabolic hormones and nutrition on ovarian follicle development in
cattle: practical implications Dom Anim Endocrinol 23, 229241
Grammer K, Fink B, N Neave (2005) Human pheromones and sexual attraction Eur J Obs Gyn
Reprod Biol 118, 135-152
Hampton JH, Manikkam M, Lubahn DB, Smith MF and HA Garverick (2004) Androgen receptor mRNA
expression in the bovine ovary Domest Anim Endocrinol 27, 81-88
Hayashi S and T Kimura (1974) Sex-attractant emitted by female mice Physiol Behav 13, 563567.
Hayashi K, Acosta TJ, Berisha B, Kobayashi S, Ohtani M, Schams D and A Miyamoto (2003) Changes
in prostaglandin secretion by the regressing bovine corpus luteum Prostaglandins Other Lipid
Mediat; 70, 339349
Henson MC and VD.Castracane (2006) Leptin in pregnancy: an update Biol Reprod 74, :218-229
Horie K, Takakura K, Fujiwara H, Suginami H, Liao S and T Mori (1992) Immunohistochemical
localisation of androgen receptor in the human ovary throughout the menstrual cycle in relation to
oestrogen and progesterone receptor expression Hum Reprod 7, 184-190.
Hurst JL, Payne CE, Nevison CM, Marie A D, Humphries RE, Robertson DHL, Cavaggioni A and RJ
Beynon (2001) Individual recognition in mice mediated by major urinary proteins Nature 414, 631634
Iqbal J, Kurose Y, Canny B, IJ Clarke (2006) Effects of central infusion of ghrelin on food intake and
plasma levels of growth hormone, luteinising hormone, prolactin, and cortisol secretion in sheep
Endocrinology 147, 510519
Iwata E, Kikusui T, Takeuchi Y, Mori Y (2003) Substances derived from 4-ethyl octanoic acid account
for primer pheromone activity for the "male effect" in goats J Vet Med Sci 65, 1019-10121
Izard MK and JG Vandenbergh (1982) The effects of bull urine on puberty and calving date in
crossbred beef heifers J Animal Sci 55, 11601168

48

Palma

Jacob S and MK McClintock (2000) Psychological state and mood effects of steroidal chemosignals in
women and men Horm Behav 37, 57-78
Jacob S, Hayreh DJ and MK McClintock (2001) Context-dependent effects of steroid chemosignals on
human physiology and mood Physiol Behav 74, 15-27
Jacob S, McClintock MK, Zelano B and V Ober (2002) Paternally inherited HLA alleles are associated
with women's choice of male odor. Nat Genet 30, 175-179
Jamnongjit M and SR Hammes (2006) Ovarian steroids: the good, the bad, and the signals that raise
them Cell Cycle 5, 1178-1183
Jordan WC and MW Bruford (1998) New perspectives on mate choice and the MHC Heredity 81, 239245
Karlson P and M Lscher (1959) Pheromones: a new term for a class of biologically active
substances Nature 183, 5556
Katongole CB, Naftolin F and RV Short (1971) Relationship between blood levels of luteinizing
hormone, and testosterone in bulls and the effect of sexual stimulation J Endocrinol 50, 457466
Kawamura K, Sato N, Fukuda J, Kodama H, Kumegai J, Tanikawa H, Nakamura A, Honda Y, Sato T,
T Tanaka (2003) Ghrelin inhibits the development of mouse preimplantation embryos in vitro
Endocrinology 144, 26232633
Kershaw EE and JS Flier (2004) Adipose tissue as an endocrine organ J Clin Endocrinol Metab 89,
25482556
Kindahl H, Lindell JO and LE Edqvist (1981) Release of prostaglandin F2 during the oestrous cycle
Acta Vet Scand 77, 143158
Kojima M, Hosoda H, Date Y, Nakazato M, Matsuo H and K Kangawa (1999) Ghrelin is a growthhormone-releasing acylated peptide from stomach Nature 402, 656-660
Korbonits M, Goldstone AP, Gueorguiev M and AB Grossman (2004) Ghrelin a hormone with multiple
functions Frontiers in Neuroendocrinology 25, 27-68
Kumar KR, Archunan G, Jeyaraman R and S Narasimhan (2000) Chemical characterization of bovine
urine with special reference to oestrus Vet Res Comm 24, 445-454
Liberles SD and LB Buck (2006) A second class of chemosensory receptors in the olfactory epithelium
Nature 442, 645-650
Lucy MC (2000) Regulation of ovarian follicular growth by somatotropins and insulin-like growth
factors in cattle J Dairy Sci 83, 16351647
Lucy MC (2003) Mechanisms linking nutrition and reproduction in postpartum cows Reproduction in
Domestic Ruminants V Reproduction Suppl 61, 415417
Lundstrm JN (2005) Psychological and Neurological modulation of a putative human pheromone
Acta Universitatis Upsaliensis Faculty of social Sciencies, Uppsala, 1-73
Maier ChM (2005) Wirkung eines synthetisch hergestellten Pheromon-analogs (PAP) auf das
Wohlbefinden von Mastschweinen beim Transport zum Schlachthof, Dissertation, Mnchen, 3-10
Matchock RL and EJ Susman (2006) Family Composition and Menarcheal Age: Anti-Inbreeding
Strategies Am J Hum Biol 18, 481491
McGlone JJ and DL Anderson (2002) Synthetic maternal pheromone stimulates feeding behavior and
weight gain in weaned pigs J Anim Sci 80, 3179-3183
Schams D and B Berisha (2004) Regulation of corpus luteum function in cattle-an overview Reprod
Dom Anim 39, 241251.
Meidan R, Levy N, Kisliouk T, Podlovny L, Rusiansky M, E Klipper (2005) The yin and yang of corpus
luteum-derived endothelial cells: balancing life and death Domestic Anim Reprod29, 318328.
Meredith M (2001) Human vomeronasal organ function: a critical review of best and worst cases
Chem Senses 26, 433 445
Morrison CD, Daniel JA, Buff PR, DH Keisler (1999) Continuous, intracerebroventricular infusion of
neuropeptide Y induces a prolonged suppression of LH J Anim Sci 77, 216
Mulac-Jericevic B, Mullinax RA, DeMayo FJ, Lydon JP and OM Conneely (2000) Subgroup of
reproductive functions of progesterone mediated by progesterone receptor-B isoform Science 289,
1751-1754

Biologa de la reproduccin

49

Mulac-Jericevic B, Lydon JP, DeMayo FJ and OM Conneely (2003) Defective mammary gland
morphogenesis in mice lacking the progesterone receptor B isoform Proc Natl Acad Sci, 100, 97449749
Myers MG Jr (2004) Leptin receptor signaling and the regulation of mammalian physiology Recent
Prog Horm Res 59, 287-304
Nakahara K, Nakagawa M, Baba Y, Sato M, Toshinai K, Date Y, Nakazato M, Kojima M, Miyazato M,
Kaiya H, Hosoda H, Kangawa K, N Murakami () Maternal ghrelin plays an important role in rat fetal
development during pregnancy Endocrinology 147, 1333-1342
Nelson RJ and S Chiavegatto (2001) Molecular basis of aggression Trends Neurosci 24, 71319
Over R, Cohen-Tannoudji J, Claus M and JP Signoret (1990) Effect of pheromones from male goats
on LH secretion in anestrous ewes Physiol Behav 48, 665668
Pearson Helen (2006) Early embryos can yield stem cells... and survive Nature 442, 858
Penny LA, Armstrong DG, Bramley T,Weeb R, Collins RA,Watson ED (1999) Immune cells and
cytokine production in the bovine corpus luteum throughout the estrous cycle and after induced
luteolysis J Reprod Fertil 115, 8796
Penton-Voak IS, Jones BC, Little AC, Baker S, Tiddeman B, Burt DM and DI Perrett (2001) Symmetry,
sexual dimorphism in facial proportions and male facial attractiveness Proc R Soc Lond B Biol Sci
268, 1617-1623
Petri G, Wysocki CJ, Barhart KT, Sondheimer SJ, JJ Leiden (2003) Male axilar extracts contain
pheromones that effect pulsatile secretion of luteinizing hormone and mood in women recipients
Horm Behav 20, 474-482
Petroff MG, Petroff BK, JL Pate (2001) Mechanisms of cytokine-induced death of cultured bovine
luteal cells Reproduction 121, 753760
Rekwot PI, Ogwu D, Oyedipe EO and VO Sekoni (2001) The role of pheromones and biostimulation in
animal reproduction Anim Reprod Scie 65, 157-170
Rasmussen LE and DR Greenwood (2003) Frontalin: a chemical message of musth in Asian
elephants (Elephas maximus) Chem Senses 28, 433-46
Rekwot PI, Ogwu D, Oyedipe EO and VO Sekoni (2000a) Effects of bull exposure and body growth on
onset of puberty in Bunaji and Friesian x Bunaji heifers Reprod Nutr Dev 40, 19
Rekwot PI, Ogwu D, EO Oyedipe (2000b) The influence of bull biostimulation, season and parity on
resumption of ovarian activity of Zebu (Bos indicus) cattle following parturition Animal Reprod Sci
63, 111
Rekwot PI, Ogwu D, Oyedipe EO and VO Sekoni (2001) The role of pheromones and biostimulation in
animal reproduction Anim Reprod Sci65, 157-70
Roberson MS, Wolfe MW, Stumpf TT, Werth LA, Cupp AS, Kojima ND, Wolfe PL, Kittok RJ, JE Kinder
(1991) Influence of growth rate and exposure to bulls on age at puberty in beef heifers J Animal Sci
69, 292298
Rosa HJD, Juniper DT and MJ Bryant (2000) The effect of exposure to oestrus ewes on rams sexual
behaviour, plasma testosterone concentrations and ability to stimulate ovulation in seasonally
anoestrus ewes Appl Anim Behav Sci 67, 293-305
Rushton JP and TA Bons (2005) Mate choice and friendship in twins: evidence for genetic similarity
Psychol Sci 16: 555559
Salcedo R, Ponce ML, Young HA, Wasserman K, Ward JM, Kleinman HK, Oppenheim JJ, Murphy WJ
(2000) Human endothelial cells express CCR2 and respond to MCP-1: direct role of MCP-1 in
angiogenesis and tumor progression Blood 96, 34-40
Sambrus, 1971 citado por Weitze K-F (2001) Andrologie beim Bullen En: Bush W, A Holzmann (Ed.),
Physiologie und Pathologie der Fortpflanzung bei mnnlichen Tieren, Schattauer Verlag, Stuttgart,
New York, 119-216
Shearer MK, LS Katz (2006) Female-female mounting among goats stimulates sexual performance in
males Hor Beh 50, 33-37
Skarzynski DJ, Jaroszewski JJ, K Okuda (2005) Role of tumor necrosis factor-alpha and nitric oxide in
luteolysis in cattle Domest Anim Endocrinol 29, 340-346

50

Palma

Slomczynska M, Duda M and K Slzak (2001) The expression of androgen receptor, cytochrome P450
aromatase and FSH receptor mRNA in the porcine ovary Folia Histochem Cytobiol 39, 9-13
Smith OB and B Somande (1994) Interaction between nutrition and reproduction in farm animals
Proceedings of a regional seminar held by the International Foundation for Science, Niamey, Niger,
January 17-21, 7-26
Spencer AG, Woods JW, Arakawa T, Singer II and WL Smith (1998) Subcellular localization of
prostaglandin endoperoxide H synthases-1 and -2 by immunoelectron microscopy J Biol Chem
273, 98869893
Steckler T, Wang J, Bartol FF, Roy SK and V Padmanabhan (2005) Fetal programming: prenatal
testosterone treatment causes intrauterine growth retardation, reduces ovarian reserve and
increases ovarian follicular recruitment Endocrinology 146, 3185-3193
Stern K and MK McClintock (1998) Regulation of ovulation by human pheromones Nature 392, 177179
Tarin JJ and V Gomez-Piquer (2002) Do women have a hidden heat period? Hum Reprod 17, 22432248
Towson DH, OConnor CL, JK Pru (2002) Expression of Monocyte Chemoattractant Protein-1 and
Distribution of Immune Cell Populations in the Bovine Corpus Luteum Throughout the Estrous
Cycle Biol Reprod 66, 361366
Vandenbergh JG Acceleration and inhibition of puberty in female mice by pheromones J Reprod Fertil
Suppl 19, 411-419
Vulliemoz NR, Xiao E, Xia-Zhang L, Germond M, Rivier J, M Ferin (2004) Decrease in luteinizing
hormone pulse frequency during a five-hour peripheral ghrelin infusion in the ovariectomized
rhesus monkey J Clin Endocrinol Metab 89, 5718-5723
Wade GN and JE Jones (2004) Neuroendocrinology of nutritional infertility Am J Physiol Regul Integr
Comp Physiol 287, R1277-1296
Walkden-Brown SW, Restall H, H Henniawati (1993) The male effect in the Australian cashmere goat:
1. Ovarian and behavioural response of seasonally anovulatory does following the introduction of
bucks Anim Reprod Sci 32, 4153
Wauters M, Considine RV and van LF Gaal (2000) Human leptin: from an adipocyte hormone to an
endocrine mediator Eur J Endocrinol 143, 293311
Webb R, Garnworthy PC, Gong J-G and DG Armstrong (2004) Control of follicular growth: local
interactions and nutritional influences J Anim Sci 82, E63-E74
Wedekind C, Seebeck T, Bettens F, AJ Paepke (1995) MHC-dependent mate preferences in humans
Proc R Soc Lond B 260, 245-249
Weitze K-F (2001) Andrologie beim Bullen En: Bush W, A Holzmann (Ed.), Physiologie und Pathologie
der Fortpflanzung bei mnnlichen Tieren, Schattauer Verlag, Stuttgart, New York, 119-216
Whitten WK (1959) Occurrence of anoestrus in mice caged in groups J Endocrinol 18, 102107
Wilson OE and WH Bossert (1963) Chemical communication among animals Recent Progr Hormone
Res 19, 673716
Wiltbank MC and JS Ottobre (2003) Regulation of intraluteal production of prostaglandins Reprod Biol
Endocrinol 10 1:91
Wyatt TD (2004) Pheromones and animal behaviour In: Communication by smell and taste,
Cambridge University Press, pp 391
Wysocki JCh and G Preti (2004) Facts, fallacies, fears and frustrations with human pheromones Anat
Rec Part A 281A, 1201-1211
Xiao C, Liu JM, Sirois J, AK Goff (1998) Regulation of cyclooxygenase-2 and prostaglandin F
synthase gene expression by steroid hormones and interferon-t in bovine uterine endometrial cells
Endocrinology 139, 22932299
Yoon H, Enquist LW and C Dulac (2005) Olfactory inputs to hypothalamic neurons controlling
reproduction and fertility Cell 123, 669-682
Zeng X-nong, Leyden JJ, Lawley HJ, Sawano K, Nohara I and G Preti Analysis of characteristic
odours from human male axillae J Chem Ecol 17, 1469-1492

Anda mungkin juga menyukai