高繁殖力黄淮山羊类固醇激素分泌特点与卵泡颗粒细胞基因差异表达的研究
|
摘要
本文调查了黄淮山羊高繁殖力母羊不同胎次与产羔数之间的关系,比较了高低繁殖力母羊在发情周期和卵巢摘除初期外周血P4和E2变化的特点,探讨了不同繁殖性状母羊在黄体期早期的卵泡、黄体及其卵泡闭锁的差异。通过抑制消减杂交技术建立起了高繁殖力母羊卵泡发育颗粒细胞差异表达基因消减cDNA文库,旨在揭示高繁殖力母羊卵泡发育调控及生殖内分泌特点,为进一步研究高繁殖力母羊的遗传育种提供一定的参考依据。
1黄淮山羊高繁殖力母羊产羔调查与分析
为了探究高繁殖力母羊多胎繁殖性状的特点,对江淮地区427只具有高繁殖力黄淮山羊母羊(窝产羔数至少一次产羔≧5)的产羔进行了调查和分析。结果表明,随着胎次的增加,高繁殖力母羊的产羔数增加,直至第5胎次,之后产羔数下降。在所有胎次中,高繁殖力母羊在第5胎次的产羔数达到最大,平均窝产羔数3.78±0.06羔,与其他胎次的产羔数相比较,呈极显著差异；相比之下,母羊在第1胎次的产羔数极显著低于其他胎次。根据不同窝产羔数的母羊所占比例分析,在第1胎次,随窝产羔数的增加,其母羊的比例依次降低,但在之后母羊所占的比例中,窝产1羔和2羔的比例急剧下降；而窝产4～7羔的母羊比例呈递增趋势延至第4～5胎次,其中随着窝产羔数的增加,其母羊所占比例的最大值在不同胎次间的分布依次后移。以上结果提示,高繁殖力母羊的多胎性状在不同胎次间具有明显的差异,其母羊性成熟后在初次繁殖时卵巢卵泡成熟与排卵及其妊娠生理等已经具备了极端高繁殖力的生理特点。
2在发情周期中高低繁殖力母羊外周血孕酮和雌二醇浓度变化的比较
选择生殖机能正常、空怀、经产的黄淮山羊高繁殖力(窝产羔数至少一次产羔≧5)和低繁殖力(每次窝产羔数≦2)母羊各4只。以间隔12 d肌注氯前列烯醇0.1 mg/只·次诱导同期发情后,在自然发情周期中,连续两个情期每天以间隔12 h和在黄体中期和发情前期间隔12 min密集性采集颈静脉瘘管血样,测定外周血E2和P4含量。结果显示,在黄体期的大多数时间内,高繁殖力母羊外周血P4量高于低繁殖力母羊,在黄体期早期的P4量上升速度,高繁殖力羊早于低繁殖力羊；两组母羊在P4的平台期,P4量呈显著性差异。在整个发情周期中,高繁殖力母羊平均E2浓度均高于低繁殖力母羊；在黄体期早期,高繁殖力母羊出现2个E2峰值,而低繁殖力母羊仅有1个。高低繁殖力母羊外周血P4和E2的量均是以非典型的脉冲式变化。以上结果表明,在黄体期早期,高繁殖力母羊外周血P4量的上升速度和卵泡波卵泡发育的周转均高于低繁殖力母羊,而在E2含量上,高繁殖力母羊维持在较高水平。
3高低繁殖力母羊在黄体期早期卵巢卵泡和黄体的比较
高低繁殖力母羊各4只,在氯前列烯醇同期发情后的第3个发情周期7～9 d,通过颈静脉瘘管采集羊的外周血液,测定E2和P4含量,并于第7和第9 d应用羊用腹腔内窥镜和腹部手术方法,观察和测定母羊卵泡和黄体大小。结果显示,在发情周期7～9 d的卵泡发育中,高低繁殖力母羊卵巢小卵泡和大卵泡数均处于减少趋势中,但差异均不显著,而中卵泡数以及总卵泡数均增加,其中高繁殖力母羊的中卵泡数平均增加了7.50个/只,低繁殖力母羊平均增加了4.80个/只。高繁殖力母羊外周血P4和E2浓度分别高于低繁殖力母羊,P4浓度在高低繁殖力母羊之间呈显著性差异；在发情周期第9d,高繁殖力母羊卵巢小卵泡数和Ⅱ型黄体数均分别显著高于低繁殖力羊16.43%和27.73%。以上结果表明,在发情周期7～9 d的卵泡发育中,黄淮山羊高低繁殖力母羊卵巢小卵泡、中卵泡和大卵泡数的发育趋势基本处于相同的生理变化。高繁殖力母羊外周血P4浓度显著高于低繁殖力母羊,促进了高繁殖力母羊卵泡的周转；高繁殖力母羊排卵卵泡以体积相对较小的成熟卵泡增多,卵巢卵泡的共优势化明显,Ⅱ型黄体数增加。
4卵巢摘除初期高低繁殖力母羊外周血孕酮和雌二醇浓度变化的比较
高低繁殖力母羊各4只,在发情周期的第9 d,通过腹部外科手术摘除双侧卵巢,并应用颈静脉瘘管连续5 d和卵巢摘除后第1和第5 d密集性采血。结果显示,母羊卵巢在摘除后的E2和P4分泌,均以非典型性的脉冲式释放。在卵巢摘除后的第1d,高繁殖力母羊外周血液P4和E2浓度均分别大于低繁殖力母羊。高低繁殖力母羊在卵巢去除后的1～3 d的P4水平均呈下降趋势；高繁殖力母羊的E2除在第1 d极显著性高于低繁殖力羊外,在其后的2～5 d,E2的平均值总体呈低于低繁殖力母羊的趋势,并在第5 d呈现差著性差异。以上结果表明,母羊非性腺相关组织在分泌E2和P4时呈现非典型的脉冲式。在外周血E2量上,高低繁殖力母羊之间存在呈现显著性差异,非性腺组织所产生的E2可能参与了母羊的生殖过程。
5山羊黄体期闭锁卵泡的组织定位观察
高低繁殖力母羊各4只,在发情周期第9 d通过腹部外科手术摘除双侧卵巢,迅即投入液氮中保存,冷冻切片,TUNEL检测法结合苏木精染色法,观察卵巢闭锁卵泡的组织学特点。结果显示,在高低繁殖力母羊卵巢闭锁的无腔卵泡中,均极少见到凋亡的卵母细胞。有腔卵泡的闭锁,多表现在卵泡腔的壁颗粒细胞层中有大量颗粒细胞凋亡,表现出明显的荧光反应,并多处于近腔颗粒细胞；而部分卵泡在颗粒层中出现凋亡细胞偶有零星存在的现象,但是在高低繁殖力母羊之间均无明显差异。在高繁殖力母羊的卵巢中,仅一膜之隔的相邻卵泡之间闭锁和非闭锁同时显现。以上结果表明,TUNEL检测结合原片苏木精染色的鉴定为卵巢组织凋亡细胞的定位提供一个较好的方法。在闭锁的有腔卵泡中,卵泡多表现出近腔细胞的凋亡特征,高低繁殖力母羊在闭锁卵泡的组织定位上无显著性差异。
6高繁殖力母羊大小卵泡颗粒细胞差异表达基因消减cDNA文库的构建和筛选
以氯前列烯醇肌肉注射2只高繁殖力母羊40 h后剪取卵巢,在20 min内计数卵泡、测量卵泡直径,以组织钝性分离和收集颗粒细胞,并剪取取样后的卵泡部分组织置入液氮内保存。从颗粒细胞中提取总RNA的大卵泡(5～6 mm)和小卵泡(3～4 mm)均未发生凋亡,总RNA经过双链cDNA的合成和扩增,以抑制消减杂交技术(SSH)构建了山羊卵泡颗粒细胞正向和反向cDNA文库。在正反向文库中,分别随机挑选了190个克隆进行斑点杂交。根据斑点杂交信息,在正反向文库中共选择了110个克隆单向测序,获得了109个EST,聚类后经过Blastn比对后发现,共有10个与已知功能基因相似,16个与已知但功能未知基因相似,65个为新的EST。这些差异基因的发现有助于揭示山羊卵泡发育和分化的调控机制,为进一步研究高繁殖力母羊卵泡共优势化的生殖机理提供一定的参考依据。
The present paper investigated relationship between different parities and litter size of Huanghuai goats at high prolificacy. This research also compared peripheral blood concentrations of progesterone and estradiol of Huanghuai goats with poor prolificacy during the estrous cycle and in the early days after ovariectomy. The different prolific goats were also employed to analyse differences in number and size of their ovaries antral follicles and corepus luteum and in follicular atresia. In this study, a subtractive cDNA library of different expression genes in follicular granulosa cells in Huanghuai goats of high prolificacy was constructed by suppression subtractive hybridization. The objective of this study was to reveal the mechanism of follicles development regulation and their feature of reproduction endocrine in goat of high prolificacy, and to provide certain references for identifying the candidate genes and for its heredity and breeding of their high prolificacy.
1. Investigation of Little Size of Huanghuai Goats with High Prolificacy
The little sizes of 427 highly prolific Huanghuai goats (HP,≥5 kids born per litter on at least one occasion) distributed in the Huanghuai plain were investigated to approach characteristic of their little size as following different parities. The results showed that litter size increased up to parity 5 and then decreased. At all parity levels, parity 5 HP does have the highest mean litter size (3.78±0.06) and parity 1 goats have the smallest. Based on analyzing the percentage of females at different little size, the females decreased at parity 1 as little size increased, and the proportion had 1 or 2 kids born per litter decreased rapidly after parity 1.However, the percentage of goats with 4 to 7 little size increased up to parity 4 to 5, furthermore as little size increased, the maximum number of the does delayed in order at different parity. It was concluded that there were obvious differences in little size at different parity in prolific does, and the limit facts of prolificacy probably was their uterus capacity.
2. Comparison of Concentrations of Progesterone and Estradiol in Peripheral Plasma during the Estrous Cycle between the Goats of High and Poor Prolificacy
Eight Huanghuai goats with 3-5 years old, non-lactating, parity 4-5 were selected to study two sequential natural estrous cycles of does with high prolificacy ((?)5 kids born per litter on at least one occasion, n=4) or poor prolificacy (<3 kids born per litter on any occasion, n=4) by initial synchronization achieved using two i. m. of 0.1 mg of a synthetic analog of prostaglandin F2 alpha given 12 days apart. Jugular blood samples were collected twice a day during the natural estrus by jugular syrinx and at 12-minute intervals for 3.2 hours over 2-3 days of proestrus and mid-luteal phase to assay for progesterone and estradiol. The results showed that progesterone concentrations began to rise earlier and were higher in the HP than in the PP goats on most days of the luteal phase but not during the follicular phase of the cycle. There was a significant difference between the two groups in the magnitude of the progesterone plateau. Mean estradiol concentrations in the HP group remained significantly higher than in the PP group during the estrus cycle. There were two estradiol peaks in the HP goats during the early luteal phase, but only one in the PP group. Progesterone and estradiol were secreted in a pulsatile atypia manner in HP and PP goats. It was concluded that the increase of progesterone in peripheral plasma and the turnover of follicles growth of waves in HP goats were higher than in PP group during the estrus cycle, and were the same for the overall mean estradiol concentrations.
3. Comparison of Number and Size of Follicles and Corpora Lutea in Early Days of Luteal Phase in Huanghuai Goats of High or Poor Prolificacy
Four HP and four PP goats were employed to collect Jugular blood samples individually at seven to nine days in third estrus cycle after synchronization achieved using a synthetic analog of prostaglandin F2 alpha and to assay for progesterone and estradiol. Meanwhile, follicles and corpora lutea were observed and measured with laparoscopy and operation on abdominal region at day 7 and day 9 in estrus cycle respectively. The results showed that number of small and large follicles had a decreasing tendency during 7 to 9 days of estrus cycle, but no difference was observed respectively. Both middle and all follicles increased, namely the mean middle follicles did to 7.50 per HP goat and to 4.48 per PP goat. Concentration of P4 and E2 in peripheral blood in HP does were higher than in PP goats, respectively, however there was no difference in P4 between two groups. Small follicles and typeⅡof corepus lutea in HP goats had obviously higher by 16.43%and 27.73%respectively compared with the PP does at day 9 of estrous cycle. It was concluded that there was a nearly same tendency in physiology of small-, middle- and large-follicles development during 7 to 9 days of estrus cycle in the HP and PP goats. The higher concentration of P4 in HP does promoted the turnover in follicles compared wich PP goats. There were more ovulation follicles characterizing smaller follicle in volume when matured in HP goats, and obvious co-dominance of follicles, and moreⅡtype of corepus lutea after ovulation, compared with those of PP does.
4. Comparison of Concentrations of Progesterone and Estradiol in Peripheral Plasma in Early Days after Ovariectomy between the Goats of High and Poor Prolificacy
Four HP and four PP goats were used to make ovariectomy of both ovaries respectively at day 9 of estrus cycle and to collect Jugular blood samples twice daily from 1 to 5 days after ovariectomy and at 12-minute intervals for 3.2 hours at day 1 and 5, in order to assay for progesterone and estradiol. The results showed that progesterone and estradiol were secreted in a pulsatile atypia manner in HP and PP goats. At day 1 after ovariectomy, the E2 and P4 concentrations were significantly higher in the HP than in the PP goats. Then there were a decreasing tendency in P4 lever in two groups during 1 to 3 days. The overall mean E2 concentrations in HP goats were lower than in the PP group from 2 to 5 days, significant differences were found at day 1 and 5 but their different concentrations were opposite. It was concluded that progesterone and estradiol from nonovaries were secreted in a atypia form of pulse. There existed a significant difference in E2 concentration in peripheral blood between HP and PP goats, furthermore the E2 from nonovaries probably involved in reproduction process in female.
5. Observation of Histological Localization of Atretic Follicle in Luteal Stage in the Goat
Four HP and four PP goats were employed to make ovariectomy of both ovaries respectively at day 9 of estrus cycle. The ovaries were preserved into liquefacient nitrogen quickly and made cryostat section to observe the histological characteristics of atretic follicle by TUNEL detecting and haematoxylin staining method. The results showed that very few oocytes with atresia were found in preantral follicles in the HP and PP goats. Apoptosis displayed more in a number of wall granulose cell near follicle antrum characterized appearance of fluorescence response. Few and individual apoptosis cell was found in granulose cell in some follicles, but no difference between HP and PP does. Follicles that were closed together with only a membrane separated, could have different physiological state at same time, one atresia another nonatresia in HP goats. It was concluded that the method of TUNEL assay and haematoxylin staining could provide a correct way for precisely detecting and localizing apoptotic cells in ovarian section. Apoptosis displayed more in a quantity of wall granulose cell near follicle antrum in atretic antrum follicles. There was a strictly physiological barrier between two adjacent follicles in HP goats.
6. Construction and Selection of Subtracted cDNA Library of Granulosa Cell in Small and Large Follicles in Early Days of Heat in Huanghuai Goats of High Prolificacy
Two HP goats were used to collect both ovaries individually 40 hours after synchronization achieved using a synthetic analog of prostaglandin F2 alpha. Counting and measuring follicles and collecting granulose cells by blunt dissection and preserving a fraction of follicle used into liquid nitrogen were within 20 min. Total RNA was extracted from granulose cells of large (5-6 mm) and small (3-4mm) follicles that were not found atresia, and were synthesized and amplified, then a forward- and reverse-subtractive cDNA library were constructed by suppression subtractive hybridization. A total of 190 clones randomly selected from the library were proved to be differentially expressed in the granulose cells, and 110 clones were selected out to be sequenced in one way based on the dot blot analysis. Among of these,109 EST obtained, and 10 exhibited high homology to be known genes,16 exhibited high homology to be found but its function not known genes,65 identified as new EST by Blastn checking after clustering. These different genes could conduce to reveal the regulation mechanism of follicles development in goats, meanwhile provide some reference for identifying the candidate gene of HP goats.
1黄淮山羊高繁殖力母羊产羔调查与分析
为了探究高繁殖力母羊多胎繁殖性状的特点,对江淮地区427只具有高繁殖力黄淮山羊母羊(窝产羔数至少一次产羔≧5)的产羔进行了调查和分析。结果表明,随着胎次的增加,高繁殖力母羊的产羔数增加,直至第5胎次,之后产羔数下降。在所有胎次中,高繁殖力母羊在第5胎次的产羔数达到最大,平均窝产羔数3.78±0.06羔,与其他胎次的产羔数相比较,呈极显著差异；相比之下,母羊在第1胎次的产羔数极显著低于其他胎次。根据不同窝产羔数的母羊所占比例分析,在第1胎次,随窝产羔数的增加,其母羊的比例依次降低,但在之后母羊所占的比例中,窝产1羔和2羔的比例急剧下降；而窝产4～7羔的母羊比例呈递增趋势延至第4～5胎次,其中随着窝产羔数的增加,其母羊所占比例的最大值在不同胎次间的分布依次后移。以上结果提示,高繁殖力母羊的多胎性状在不同胎次间具有明显的差异,其母羊性成熟后在初次繁殖时卵巢卵泡成熟与排卵及其妊娠生理等已经具备了极端高繁殖力的生理特点。
2在发情周期中高低繁殖力母羊外周血孕酮和雌二醇浓度变化的比较
选择生殖机能正常、空怀、经产的黄淮山羊高繁殖力(窝产羔数至少一次产羔≧5)和低繁殖力(每次窝产羔数≦2)母羊各4只。以间隔12 d肌注氯前列烯醇0.1 mg/只·次诱导同期发情后,在自然发情周期中,连续两个情期每天以间隔12 h和在黄体中期和发情前期间隔12 min密集性采集颈静脉瘘管血样,测定外周血E2和P4含量。结果显示,在黄体期的大多数时间内,高繁殖力母羊外周血P4量高于低繁殖力母羊,在黄体期早期的P4量上升速度,高繁殖力羊早于低繁殖力羊；两组母羊在P4的平台期,P4量呈显著性差异。在整个发情周期中,高繁殖力母羊平均E2浓度均高于低繁殖力母羊；在黄体期早期,高繁殖力母羊出现2个E2峰值,而低繁殖力母羊仅有1个。高低繁殖力母羊外周血P4和E2的量均是以非典型的脉冲式变化。以上结果表明,在黄体期早期,高繁殖力母羊外周血P4量的上升速度和卵泡波卵泡发育的周转均高于低繁殖力母羊,而在E2含量上,高繁殖力母羊维持在较高水平。
3高低繁殖力母羊在黄体期早期卵巢卵泡和黄体的比较
高低繁殖力母羊各4只,在氯前列烯醇同期发情后的第3个发情周期7～9 d,通过颈静脉瘘管采集羊的外周血液,测定E2和P4含量,并于第7和第9 d应用羊用腹腔内窥镜和腹部手术方法,观察和测定母羊卵泡和黄体大小。结果显示,在发情周期7～9 d的卵泡发育中,高低繁殖力母羊卵巢小卵泡和大卵泡数均处于减少趋势中,但差异均不显著,而中卵泡数以及总卵泡数均增加,其中高繁殖力母羊的中卵泡数平均增加了7.50个/只,低繁殖力母羊平均增加了4.80个/只。高繁殖力母羊外周血P4和E2浓度分别高于低繁殖力母羊,P4浓度在高低繁殖力母羊之间呈显著性差异；在发情周期第9d,高繁殖力母羊卵巢小卵泡数和Ⅱ型黄体数均分别显著高于低繁殖力羊16.43%和27.73%。以上结果表明,在发情周期7～9 d的卵泡发育中,黄淮山羊高低繁殖力母羊卵巢小卵泡、中卵泡和大卵泡数的发育趋势基本处于相同的生理变化。高繁殖力母羊外周血P4浓度显著高于低繁殖力母羊,促进了高繁殖力母羊卵泡的周转；高繁殖力母羊排卵卵泡以体积相对较小的成熟卵泡增多,卵巢卵泡的共优势化明显,Ⅱ型黄体数增加。
4卵巢摘除初期高低繁殖力母羊外周血孕酮和雌二醇浓度变化的比较
高低繁殖力母羊各4只,在发情周期的第9 d,通过腹部外科手术摘除双侧卵巢,并应用颈静脉瘘管连续5 d和卵巢摘除后第1和第5 d密集性采血。结果显示,母羊卵巢在摘除后的E2和P4分泌,均以非典型性的脉冲式释放。在卵巢摘除后的第1d,高繁殖力母羊外周血液P4和E2浓度均分别大于低繁殖力母羊。高低繁殖力母羊在卵巢去除后的1～3 d的P4水平均呈下降趋势；高繁殖力母羊的E2除在第1 d极显著性高于低繁殖力羊外,在其后的2～5 d,E2的平均值总体呈低于低繁殖力母羊的趋势,并在第5 d呈现差著性差异。以上结果表明,母羊非性腺相关组织在分泌E2和P4时呈现非典型的脉冲式。在外周血E2量上,高低繁殖力母羊之间存在呈现显著性差异,非性腺组织所产生的E2可能参与了母羊的生殖过程。
5山羊黄体期闭锁卵泡的组织定位观察
高低繁殖力母羊各4只,在发情周期第9 d通过腹部外科手术摘除双侧卵巢,迅即投入液氮中保存,冷冻切片,TUNEL检测法结合苏木精染色法,观察卵巢闭锁卵泡的组织学特点。结果显示,在高低繁殖力母羊卵巢闭锁的无腔卵泡中,均极少见到凋亡的卵母细胞。有腔卵泡的闭锁,多表现在卵泡腔的壁颗粒细胞层中有大量颗粒细胞凋亡,表现出明显的荧光反应,并多处于近腔颗粒细胞；而部分卵泡在颗粒层中出现凋亡细胞偶有零星存在的现象,但是在高低繁殖力母羊之间均无明显差异。在高繁殖力母羊的卵巢中,仅一膜之隔的相邻卵泡之间闭锁和非闭锁同时显现。以上结果表明,TUNEL检测结合原片苏木精染色的鉴定为卵巢组织凋亡细胞的定位提供一个较好的方法。在闭锁的有腔卵泡中,卵泡多表现出近腔细胞的凋亡特征,高低繁殖力母羊在闭锁卵泡的组织定位上无显著性差异。
6高繁殖力母羊大小卵泡颗粒细胞差异表达基因消减cDNA文库的构建和筛选
以氯前列烯醇肌肉注射2只高繁殖力母羊40 h后剪取卵巢,在20 min内计数卵泡、测量卵泡直径,以组织钝性分离和收集颗粒细胞,并剪取取样后的卵泡部分组织置入液氮内保存。从颗粒细胞中提取总RNA的大卵泡(5～6 mm)和小卵泡(3～4 mm)均未发生凋亡,总RNA经过双链cDNA的合成和扩增,以抑制消减杂交技术(SSH)构建了山羊卵泡颗粒细胞正向和反向cDNA文库。在正反向文库中,分别随机挑选了190个克隆进行斑点杂交。根据斑点杂交信息,在正反向文库中共选择了110个克隆单向测序,获得了109个EST,聚类后经过Blastn比对后发现,共有10个与已知功能基因相似,16个与已知但功能未知基因相似,65个为新的EST。这些差异基因的发现有助于揭示山羊卵泡发育和分化的调控机制,为进一步研究高繁殖力母羊卵泡共优势化的生殖机理提供一定的参考依据。
The present paper investigated relationship between different parities and litter size of Huanghuai goats at high prolificacy. This research also compared peripheral blood concentrations of progesterone and estradiol of Huanghuai goats with poor prolificacy during the estrous cycle and in the early days after ovariectomy. The different prolific goats were also employed to analyse differences in number and size of their ovaries antral follicles and corepus luteum and in follicular atresia. In this study, a subtractive cDNA library of different expression genes in follicular granulosa cells in Huanghuai goats of high prolificacy was constructed by suppression subtractive hybridization. The objective of this study was to reveal the mechanism of follicles development regulation and their feature of reproduction endocrine in goat of high prolificacy, and to provide certain references for identifying the candidate genes and for its heredity and breeding of their high prolificacy.
1. Investigation of Little Size of Huanghuai Goats with High Prolificacy
The little sizes of 427 highly prolific Huanghuai goats (HP,≥5 kids born per litter on at least one occasion) distributed in the Huanghuai plain were investigated to approach characteristic of their little size as following different parities. The results showed that litter size increased up to parity 5 and then decreased. At all parity levels, parity 5 HP does have the highest mean litter size (3.78±0.06) and parity 1 goats have the smallest. Based on analyzing the percentage of females at different little size, the females decreased at parity 1 as little size increased, and the proportion had 1 or 2 kids born per litter decreased rapidly after parity 1.However, the percentage of goats with 4 to 7 little size increased up to parity 4 to 5, furthermore as little size increased, the maximum number of the does delayed in order at different parity. It was concluded that there were obvious differences in little size at different parity in prolific does, and the limit facts of prolificacy probably was their uterus capacity.
2. Comparison of Concentrations of Progesterone and Estradiol in Peripheral Plasma during the Estrous Cycle between the Goats of High and Poor Prolificacy
Eight Huanghuai goats with 3-5 years old, non-lactating, parity 4-5 were selected to study two sequential natural estrous cycles of does with high prolificacy ((?)5 kids born per litter on at least one occasion, n=4) or poor prolificacy (<3 kids born per litter on any occasion, n=4) by initial synchronization achieved using two i. m. of 0.1 mg of a synthetic analog of prostaglandin F2 alpha given 12 days apart. Jugular blood samples were collected twice a day during the natural estrus by jugular syrinx and at 12-minute intervals for 3.2 hours over 2-3 days of proestrus and mid-luteal phase to assay for progesterone and estradiol. The results showed that progesterone concentrations began to rise earlier and were higher in the HP than in the PP goats on most days of the luteal phase but not during the follicular phase of the cycle. There was a significant difference between the two groups in the magnitude of the progesterone plateau. Mean estradiol concentrations in the HP group remained significantly higher than in the PP group during the estrus cycle. There were two estradiol peaks in the HP goats during the early luteal phase, but only one in the PP group. Progesterone and estradiol were secreted in a pulsatile atypia manner in HP and PP goats. It was concluded that the increase of progesterone in peripheral plasma and the turnover of follicles growth of waves in HP goats were higher than in PP group during the estrus cycle, and were the same for the overall mean estradiol concentrations.
3. Comparison of Number and Size of Follicles and Corpora Lutea in Early Days of Luteal Phase in Huanghuai Goats of High or Poor Prolificacy
Four HP and four PP goats were employed to collect Jugular blood samples individually at seven to nine days in third estrus cycle after synchronization achieved using a synthetic analog of prostaglandin F2 alpha and to assay for progesterone and estradiol. Meanwhile, follicles and corpora lutea were observed and measured with laparoscopy and operation on abdominal region at day 7 and day 9 in estrus cycle respectively. The results showed that number of small and large follicles had a decreasing tendency during 7 to 9 days of estrus cycle, but no difference was observed respectively. Both middle and all follicles increased, namely the mean middle follicles did to 7.50 per HP goat and to 4.48 per PP goat. Concentration of P4 and E2 in peripheral blood in HP does were higher than in PP goats, respectively, however there was no difference in P4 between two groups. Small follicles and typeⅡof corepus lutea in HP goats had obviously higher by 16.43%and 27.73%respectively compared with the PP does at day 9 of estrous cycle. It was concluded that there was a nearly same tendency in physiology of small-, middle- and large-follicles development during 7 to 9 days of estrus cycle in the HP and PP goats. The higher concentration of P4 in HP does promoted the turnover in follicles compared wich PP goats. There were more ovulation follicles characterizing smaller follicle in volume when matured in HP goats, and obvious co-dominance of follicles, and moreⅡtype of corepus lutea after ovulation, compared with those of PP does.
4. Comparison of Concentrations of Progesterone and Estradiol in Peripheral Plasma in Early Days after Ovariectomy between the Goats of High and Poor Prolificacy
Four HP and four PP goats were used to make ovariectomy of both ovaries respectively at day 9 of estrus cycle and to collect Jugular blood samples twice daily from 1 to 5 days after ovariectomy and at 12-minute intervals for 3.2 hours at day 1 and 5, in order to assay for progesterone and estradiol. The results showed that progesterone and estradiol were secreted in a pulsatile atypia manner in HP and PP goats. At day 1 after ovariectomy, the E2 and P4 concentrations were significantly higher in the HP than in the PP goats. Then there were a decreasing tendency in P4 lever in two groups during 1 to 3 days. The overall mean E2 concentrations in HP goats were lower than in the PP group from 2 to 5 days, significant differences were found at day 1 and 5 but their different concentrations were opposite. It was concluded that progesterone and estradiol from nonovaries were secreted in a atypia form of pulse. There existed a significant difference in E2 concentration in peripheral blood between HP and PP goats, furthermore the E2 from nonovaries probably involved in reproduction process in female.
5. Observation of Histological Localization of Atretic Follicle in Luteal Stage in the Goat
Four HP and four PP goats were employed to make ovariectomy of both ovaries respectively at day 9 of estrus cycle. The ovaries were preserved into liquefacient nitrogen quickly and made cryostat section to observe the histological characteristics of atretic follicle by TUNEL detecting and haematoxylin staining method. The results showed that very few oocytes with atresia were found in preantral follicles in the HP and PP goats. Apoptosis displayed more in a number of wall granulose cell near follicle antrum characterized appearance of fluorescence response. Few and individual apoptosis cell was found in granulose cell in some follicles, but no difference between HP and PP does. Follicles that were closed together with only a membrane separated, could have different physiological state at same time, one atresia another nonatresia in HP goats. It was concluded that the method of TUNEL assay and haematoxylin staining could provide a correct way for precisely detecting and localizing apoptotic cells in ovarian section. Apoptosis displayed more in a quantity of wall granulose cell near follicle antrum in atretic antrum follicles. There was a strictly physiological barrier between two adjacent follicles in HP goats.
6. Construction and Selection of Subtracted cDNA Library of Granulosa Cell in Small and Large Follicles in Early Days of Heat in Huanghuai Goats of High Prolificacy
Two HP goats were used to collect both ovaries individually 40 hours after synchronization achieved using a synthetic analog of prostaglandin F2 alpha. Counting and measuring follicles and collecting granulose cells by blunt dissection and preserving a fraction of follicle used into liquid nitrogen were within 20 min. Total RNA was extracted from granulose cells of large (5-6 mm) and small (3-4mm) follicles that were not found atresia, and were synthesized and amplified, then a forward- and reverse-subtractive cDNA library were constructed by suppression subtractive hybridization. A total of 190 clones randomly selected from the library were proved to be differentially expressed in the granulose cells, and 110 clones were selected out to be sequenced in one way based on the dot blot analysis. Among of these,109 EST obtained, and 10 exhibited high homology to be known genes,16 exhibited high homology to be found but its function not known genes,65 identified as new EST by Blastn checking after clustering. These different genes could conduce to reveal the regulation mechanism of follicles development in goats, meanwhile provide some reference for identifying the candidate gene of HP goats.
引文
1. Dickerson, G E.1970.Efficiency of animal production-molding the biological components. J. Anim. Sci.,30:849.
2. Shelton, M.1971. Some factors affecting efficiency of lamb production. Texas Agric. Exp. Sta. Tech. Rept., No.26.
3. Gabinna, D.1989b. Improvement of the reproductive performance of Rasa Aragonesa flocks in frequent lambing systems. Ⅱ. Repeatability and heritability of sexual precocity, fertility and litter size. Livest. Prod. Sci.,22:87.
4.蒋英,陶雍.中国山羊.西安：陕西科学技术出版社,1988,102-104.
5.杨利国,周东蕊.奶牛繁殖力的影响因素及提高措施.中国奶牛,2001,(1)：52-54
6. Van Vleck, L D, E J Pollak, and E A. Oltenacu. Genetics for the Animal Sciences. W. H. Freeman and Co., New York,1987.
7. Lasley, J F. Genetics of Livestock Improvement (4th Ed.). Prentice-Hall, Hemel Hempstead, U. K., 1987.
8. Abdulkhaliq A M, Harvey W R, Parker C F. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. J. Anim. Sci.,1989,67:3250-3257.
9. Fahmy M H. Development of DLS breed of sheep:genetic and phenotypic parameters of date of lambing and number of lambs born alive. Can. J. Anim. Sci.,1990,70:771-778.
l0.Altarriba J, Varona L, Crarcia-Comes L A et al. Bayesian inference of variance components for number of lambs born alive in Rasa Aragonesa sheep. J. Anim. Sci.,1998,76:23-28.
11.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28(5)：418-423.
12. SHUNICHI SHIMASAKI, R. KELLY MOORE, FUMIO OTSUKA, et al. The Bone Morphogenetic Protein System In Mammalian Reproduction. Endocrine Reviews,2004,25:72-101.
13. Souza C J H, C MacDougall, B K Campbell, et al. The Booroola (FecB) phenotype is associated with a mutation in the bone morphogenetic receptor type 1 B (BMPR1B) gene. Journal of Endocrinology,2001,169, R1-R6
14.姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
15. Philippe Monget, Stephane Fabre, Philippe Mulsant, et al. Regulation of ovarian folliculogenesis by IGF and BMP system in domestic animals. Domestic Animal Endocrinology,2002,23:139-154
16. Campbell,B K, D T Baird, C J H Souza, et al. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction,2003,126:101-111
17. Souza C J H, B K Campbell, Baird, D T. et al. Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology, 1997a,138:5333-5340.
18. Piper L R, Bindon B M and Davis G H. The single gene inheritance of the high litter size of theBooroola Merino. Genetics of Reproduction in sheep,1985,38(2):115-125.
19. McNtty K P and Henderson K M. Gonadotrophins, fecundity genes and ovarian follicular function. Journal of steroid Biochemistry,1987,27:365-373.
20. Davis G H, Dodds K G, Wheeler R, et al. Evidence that an imprinted gene on the X chromosome increases ovulation rate in sheep. Biology of Reproduction,2001,64:216-221.
21. Jonmundosson JV and Adalstinsson S. Single genes for fecundity in Icelandic sheep. In:R. B. Land and D. W. Robinson (Ed) Genetics of Reproduction in sheep.1985,159-168.
22. Davis G H, McEwan J C, Fennessy P F, et al. Infertility due to bilateral ovarian hypoplasia in sheep homozygous (FecXI FecXH) for the Inverdale prolificacy gene located on the X chromosome. Biology of Reproduction,1992,46:636-640.
23. Davis GH, McEwan JC, Fennessy PF, et al. Evidence for the presence of a major gene influencing ovulation rate on the X chromosome of sheep. Biology of Reproduction,1991,44:620-640.
24.柳淑芳,姜运良,杜立新BMPR-IB和BMP15基因作为小尾寒羊多胎性能候选基因的研究.遗传学报,2003,30(8)：755-760.
25.王根林,毛鑫智,Davis G H,等.DAN分析发现我国湖羊和小尾寒羊存在Booroola(FecB)多胎基因.南京农业大学学报,2003,26(1)：104-106.
26. Matzuk MM, Lamb DJ. Genetic dissection of mammalian fertility pathways. Nature Cell Biol (Suppl),2002,4:s41-s49
27.桑润滋主编.动物繁殖生物技术(第二版).北京：中国农业出版社,2006年7月,25-42.
28. Markstrom E, Svernsson Ech, Shao R, et al. Survival factors regulating ovarian apoptosis-dependence on follicle differentiation. Reproduction,2002,123:23-30
29. Manabe N, Goto Y, Matsuda-Minehata F, et al. Regualtion mechanism of selective atresia in porcine follicles:regulation of granulose cell apoptosis during atresia. J Reprod Deve,2004,50:493-514
30.蒋英编著.世界养羊科学及生产(第一版).北京：中国农业大学出版社,1999.
31.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
32. Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization:a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA,1996,93(12):6025-6030.
1.姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
2.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28(5)：418-423.
3.谢庄,程瑞禾,韩玉刚.湖羊公羊产羔效应的统计学分析.南京农业大学学报,1998,21(1)：77-81
4. Waldron D F and D L. Thomas. Increased litter size in Rambouillet sheep:I. Estimation of genetic parameters. J. Anim. Sci.1992,70:3333-3344.
5. Shelton M. and J W Menzies. Genetic parameters of some performance characteristics of range fine-wool ewes. J. Anim. Sci,1968,27:1219.
6.中国羊品种志编写组.中国羊品种志.上海：上海科学技术出版社,1989,29-58.
7.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
8.朱东华,徐忠,管生.长江三角洲白山羊繁殖性能调查研究.中国草食动物,2005年第25卷第2期：35-36
9.陈启康,钱鹤良,沙文锋,等.海门山羊多胎繁殖性能和改良利用的研究.南京农专学报,2003,18(1)：35-39.
10.李金林,冉汝俊.鲁北白山羊多胎早熟性能选育研究.中国养羊,1997,4：12-14.
11.徐云华,曹洪防,李强,等.莱芜黑山羊高繁品系培育.山东畜牧兽医,2001,6：13-14.
12.王维春,熊朝瑞,贾旌旗,等.南江黄羊肉用品种选育研究.四川畜牧兽医,2000,27(7)：40-42.
13.Theresa Wilson, Xi-Yang Wu, Jennifer L, et al. Highly Prolific Booroola Sheep Have a Mutation in the Intracellular Kinase Domain of Bone Morphogenetic Protein IB Receptor (ALK-6) That Is Expressed in Both Oocytes and Granulosa Cells. BIOLOGY OF REPRODUCTION,2001,64, 1225-1235.
14. Hanrahan J P. Crossbreeding studies involving Finnish Landrace and Galway sheep. Proc. Working Symp. Breed Evaluation and Crossing Experiments, Zeist,1974. pp 431-444.
15. Land R B, W S Russell and HP Donald. The litter size and fertility of Finnish Landrace and Tasmanian Merino sheep and their reciprocal crosses. Anim. Prod,1974,18:265.
16. Ricordean G, L Tchamitchian, J Thimonier, et al. First survey of results obtained in France on reproductive and maternal performance in sheep,with particular reference to the Romanov breed and crosses with it. Livest. Rod. Sci,1978,5:181.
17. Bradford G E, A Lahlou-Kassi, Y M Berger, et al. Performance of D'man and Sardi breeds of sheep on accelerated lambing. II. Ovulation rate and embryo survival. Small Ruminant Res,1989,2:241.
18. Davis GH, Montgomery GW, Allison AJ, et al. Segregation of a major gene influencing fecundity in progeny of Booroola sheep. NZ J Agric Res,1982,25:525-529
19. McNatty KP, Lun S, Heath DA, et al. Differences in ovarian activity between Booroola X Merino ewes which were homozygous, heterozygous or non-carriers of a fecundity gene influencing their ovulation rate. J Reprod Fertil,1986,77:193-205
20.Souza CJH, Moraes JCF, Chagas LM. Effect of Booroola gene on time of ovulation and ovulatory dynamics. Anim Reprod Sci,1994,37:7-13
21.Driancourt MA, Cahill LP, Bindon BM. Ovarian follicular populations and preovulatory enlargement in Booroola and control Merino ewes. J Reprod Fertil,1985,73:93-107
22.McNatty KP, O'Keefe LE, Henderson KM, et al.125I labelled hCG binding characteristics in theca interna and other tissues from Romney ewes and from Booroola X Romney ewes with and without a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1986c,77: 477-488
23.Henderson KM, McNatty KP, O'Keeffe LE, et al. Differences in gonadotrophin stimulated cyclic-AMP production by granulose cells from Booroola X Merino ewes which were homozygous, heterozygous or non-carriers of a fecundity gene influencing their ovulation rate. J Reprod Fertil. 1987,81:395-402
24. Hanrahan J P. Ovulation rate as the selection criterion for litter size in sheep. Proc. Aust. SOCA. nim. Prod,1980,13:405
25. Waldron D F, and D L Thomas. Increased Litter Size in Rambouillet Sheep: Ⅱ. Expected Responses from Alternative Selection. J. Anim. Sci.,1992.70:3345-3350
26. Altarriba J, L Varona, L A Garcia-Cortes, et al. Bayesian Inference of Variance Components for Litter Size in Rasa Aragonesa Sheep. J. Anim. Sci.,1998.76:23-28
27. Bunge R, D L Thomas and T G Nash. Performance of Hair Breeds and Prolific Wool Breeds of Sheep in Southern Illinois:Lamb Production of F1 Adult Ewes. J. Anim. Sci.,1995.73:1602-1608
28. Cumming I A. Relationships in the sheep of ovulation rate with live weight, breed, season and plane of nutrition. Aust. J. Exp. Agric.,1977,17:234.
29. Dickerson G E, H A Glimp, H J Tuma, et al. Genetic resources for efficient meat production in sheep. Growth and carcass characteristics of ram lambs of seven breeds. J. Anim. Sci.,1972,34:940.
30. Gunn R G., J M Doney, W F Smith, et al. Effects of age and its relationship with body size on reproductive performance in Scottish Blackface ewes. Anim. Prod.,1986,43:279.
31.ANTONIO GONZALEZ-BULNES, CARLOS J H SOUZA, BRUCE K CAMPBELL, et al. Effect of Ageing on Hormone Secretion and Follicular Dynamics in Sheep with and without the Booroola Gene. Endocrinology.,2004,145(6):2858-2864
32. Ginther OJ, Kot K, Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42,987-1001.
33. Castro de T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52,399-411.
34. Tenorio Filho F, MHB Santos, PG Carrazzoni, et al. Follicular dynamics in Anglo-Nubian goats using transrectal and transvaginal ultrasound. Small Ruminant Research,2007,72,51-56.
35. Mohamed S. Medan, Gen Watanabe, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. BIOLOGY OF REPRODUCTION,2003,69,57-63.
36. Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287
37. Driancourt MA, Webb R, Fry RC. Does follicular dominance occur in ewes? J. Reprod. Fertil,1991, 93:63-70.
38. Gonzalez-Bulnes A, C Di az-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245
39. Adams G. P, Kot K, Smith CA, et al. Effect of the dominant follicle on regression of its subordinates. Can. J. Anim. Sci,1993,73,267-275.
40. Cueto M, A Gibbons, R Alberio, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
41. Gonzalez-Bulnes A, Santiago-Moreno J, Garcia-Garcia RM, et al. Origin of the preovulatory follicle in Mouflon sheep (Ovis gmelini musimon) and effect on the growth of remaining follicles during the follicular phase of oestrous cycle. Anim. Reprod. Sci,2001,65,265-272.
42. Gonzalez-Bulnes A, Santiago Moreno J, Gomez-Brunet A, et al. Follicular dynamics during the oestrous cycle in dairy goats. Anim. Sci,1999b,68,547-554.
43. Gonzalez-Bulnes A, Souza CJH, Campbell BK, et al. Systemic and intraovarian effects of dominant follicles on ovine follicular growth. Anim. Reprod. Sci,2004,84,107-119
44. EVANS A C O and J E FORTUNE. Selection of the Dominant Follicle in Cattle Occurs in the Absence of Differences in the Expression of Messenger Ribonucleic Acid for Gonadotropin Receptors. Endocrinology,1997,138:2963-2971
45. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Anim. Reprod. Sci,2003,78,271-287.
46. Mohamed S. Medan, Gen Watanabe, et al. Follicular and Hormonal Dynamics during the Estrous Cycle in Goats. Joumal of Reproduction and Development,2005, Vol.51:455-463.
47. Mann GE, McNeilly AS, Baird DT. Hormone production in vivo and in vitro from follicles at different stages of estrous cycle in sheep. J Endocrinol,1992,132:225-234.
48. Kawate N, Morita N, Tsuji M, et al. Roles of pulsatile release of LH in the development and maintenance of corpus luteum function in the goat. Theriogenology,2000,54:1133-1143.
49. Xia Y, T O'Shea, R Murison, et al. Concentrations of Progesterone, Follistatin, and Follicle-Stimulating Hormone in Peripheral Plasma Across the Estrous Cycle and Pregnancy in Merino Ewes That Are Homozygous or Noncarriers of the Booroola Gene. BIOLOGY OF REPRODUCTION,2003,69,1079-1084.
50. Bindon BM, Piper LR, Thimonier J. Preovulatory LH characteristics and time of ovulation in the prolific Booroola Merino ewe. J Reprod Fertil,1984,71:519-523.
51. Bindon BM, Piper LR, Cummins LJ, et al. Reproductive endocrinology of prolific sheep: studies of the Booroola Merino. In Genetics of Reproduction in Sheep.1985, pp 217-235 Eds RB Land and DW Robinson. Butterworths, London.
52. Bindon BM. Reproductive biology of the Booroola Merino sheep. Australian Journal of Biological Sciences,1984,37:163-189.
53. McNatty KP, Hudson N, Henderson KM, et al. Differences in gonadotrophin concentrations and pituitary responsiveness to GnRH between Booroola ewes which were homozygous (FF), heterozygous (F+) and non-carriers (++) of a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1987,80:577-588.
54. Fry RC, Clarke IJ, Cummins JT, et al. Induction of ovulation in chronically hypophysectomized Booroola ewes. Journal of Reproduction and Fertility,1988,82:711-715.
55. Hudson NL, O'Connell AR, Shaw L, et al. Effect of exogenous FSH on ovulation rate in homozygous carriers or non-carriers of the Booroola FecB gene after hypothalamic—pituitary disconnection or after treatment with a GnRH agonist. Domestic Animal Endocrinology,1999,16: 69-80.
56. Boulton MI, Haley CS, Springbett AJ et al. The effect of the Booroola (FecB) gene on peripheral FSH concentrations and ovulation rates during oestrus, seasonal anoestrus and on FSH concentrations following ovariectomy in Scottish Blackface ewes. Journal of Reproduction and Fertility,1995,103:199-207.
57. Souza CJH, Campbell BK, BairdDT andWebb R Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology, 1997a,138:5333-5340.
58. Campbell B K, D T Baird, C J H, et al. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction.,2003,126,101-111.
59. Adams N R, S Atkinson, R M Hoskinson, et al. Immunization of ovariectomized ewes against progesterone, oestrogen or cortisol to detect effects of adrenal steroids on reproduction. J. Reprod. Fert,1990,89:477-483.
60. Ellison PT. Correlations of basal oestrogens with adrenal androgens and relative weight in normal women. Annls Hum. Bio,1984,11,327-336.
61. Shaikh AA & Shaikh SA. Adrenal and ovarian steroid secretion in the rat estrous cycle temporally related to gonadotropins and steroid levels found in peripheral plasma. Endocrinology,1975,96:44.
62. Martin GB. Factors affecting the secretion of luteinizing hormone in the ewe. Biol. Rev,1984,9: 1-87.
63. Webb R, Baxter G, Preece RD, et al. Control of gonadotrophin release in Scottish Blackface and Finnish Landrace ewes during seasonal anoestrus. J. Reprod. Fert.,1985,73:369-378.
64. Goodman RL, Pickover SM & Karsch FJ. Ovarian feedback control of follicle stimulating hormone in the ewe:evidence for selective suppression. Endocrinology,1981a,108:772-777.
65. Rawlings NC, Jeffcoate IA & Rieger DL. The influence of oestradiol-17 β and progesterone on peripheral serum concentration of luteinizing hormone and follicle stimulating hormone in the ovariectomized ewe. Theriogenology,1984,22,473-488.
66. Barlow DH, Coutts JRT, Mowat J and Macnaughton MC. Hormonal profiles at the menopause. In Coutts, J. R. T. (ed.), The Functional Morphology of the Human Ovary. MTP Press, Lancaster, 1981,pp.223-234.
67. Atkinson S & Adams NR. Adrenal glands alter oestrogen activity in the uterus of ewes. J. Endocr., 1988,118:375-380.
68. Pant HC, Dobson H & Ward WR. Effect of active immunization against oestrogens on plasma gonadotrophins in the ewe and the response to synthetic oestrogen on LH. J. Reprod. Fert.,1978,53, 241-248.
69. Joseph I B J K, W D Currie and N C Rawlings. Effects of time after ovariectomy, season and oestradiol on luteinizing hormone and follicle-stimulating hormone secretion in ovariectomized ewes. J. Reprod. Fert,1992,94,511-523.
70.桑润滋 主编.动物繁殖生物技术(第二版).北京,中国农业出版社,2006年7月,25-42.
71. Campbell BK. The modulation of gonadotrophic hormone action on the ovary by paracrine and autocrine factors. Anat Histol Embryol,1999,28(4):247-51. 72. Blondin P, Sirard MA. Oocyte and follicular morphology as determining characteristics for developmental competence in bovine oocytes. Mol Reprod Dev.,1995,41:54-62.
73. Geshi M, Takenouchi N, Yamauchi N, et al. Effects of sodium private in nonserum maturation medium on maturation, fertilization and s ubsequrnt development of bovine oocytes with or without cumulus cells. Biol Reprod.,2000,63:1130-1134.
74. Armstrong D G and Webb R. Ovarian follicular dominance: the role of intraovarian growth factors and novel proteins. Rev Reprod,1997,2(1):139-146
75. Campbell B K, D T Baird, C J H Souza and R Webb. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction.,2003,126: 101-111
76. McNatty KP, Henderson KM, Lun S, et al. Ovarian activity in Booroola X Romney ewes which have a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1985,73: 109-120.
77. Hsueh AJW, Billig H and Tsafriri A. Ovarian follicle atresia:a hormonally controlled apoptotic process. Endocr Rev,1994,15:707-724
78. Byskov AG. Follicular atresia. In:Jones RE(ed) The Vertebrate Ovary. Plenum Press, New York, 1978, pp533-562
79. Markstrom E, Svernsson Ech, Shao R, et al. Survival factors regulating ovarian apoptosis-dependence on follicle differentiation. Reproduction.,2002,123:23-30
80. Manabe N, Goto Y, Matsuda-Minehata F, et al. Regulation mechanism of selective atresia in porcine follicles:regulation of granulosa cell apoptosis during atresia. J Reprod Dev.,2004,50(5):493-514.
81. Irving-Rodgers HF, van Wezel IL, Mussard ML, et al. Atresia revisited: two basic patterns of atresia of bovine antral follicles. Reproduction.,2001,122:761-775
82. Chun SY, Eisenhauer KM, Minami S, et al. Hormonal regulation of apoptosis in early antral follicles: follicle-stimulating as a major survival factor. Endocrinology,1996,137:1447-1456
83. Hsu SY, Hsuch AJW. Hormonal regulation of apoptosis. An ovarian perspective. Trends Endocrinol Metab,1997,8:207-213.
84. Wang Y, Chan S, Tsang BK. Role and gonadotropin regulation of X-linked inhibitor of apoptosis expression during ovarian follicular development in vitro. Biol Reprod,2003,68:610-619.
85.杨世华,余世九.生殖激素控制卵泡细胞凋亡的研究进展.动物医学进展,2002,23：1-5.
86. Guthrie HD, Garrett WM, Cooper BS. Follicle-stimulating hormone and insulin-like growth factor-1 attenute apoptosis in cultured porcine grnaulosa cell. Biol Reprod,1998,58:390-396
87. Yu YS, Li W, Han ZB, et al. The effect of follicle-stimulating hormone on follicular development granulose cells apoptosis and steroidogenesis and its mediation by insulin-like growth factor-I in the goat ovary. Theriogenology,2003,60:1691-1704.
88. Manabe N, Imai Y, Myoumoto A, et al. Apoptosis occurs in granulose cells but not cumulius cell in the atretic graafian follicles in multiparous pig ovaries. Acta Histochem Cytochem,1997,30:85-92
89. Manabe N, Imai Y, Ohno H, et al. Apoptosis occurs in granulose cells but not cumulus cells in the atretic antral follicles in the pig ovaries. Experientia,1996,52:647-657
90. Wyllie AH, Kerr JF, Currie AR. Cell death: the significance of apoptosis. Int Rev Cytol.,1980,68: 251-306
91. Palumbo A and Yeh J. In situ localization of apoptosis in the rat ovary during follicular atresia. Biol Reprod.,1994,51:888-95
92. Rosales AM, Avalos A, Vergara M, et al. Multiparametric study of atresia in ewe antral follicles: histology, flow cytometry, internucleosomal DNA fragmentation, and lysosomal enzyme activities in granulose cells and follicular fluid. Mol Reprod Dev,2000,55:270-281
93. Jolly PD, Smith FR, Heath DA, et al. Morphological evidence of apoptosis and the prevalence of apoptotic versus mitotic cells in the membrana granulosa of ovarian follicles during spontaneous and induced atresia in ewes. Biol Reprod.,1997,56:837-846
94. Jolly PD, Tisdall DJ, Heath DA et al. Apoptosis in bovine granulose cells in relation to steroid synthesis, cyclic adenosine 3',5'-monophosphate response to follicle-stimulating hormone and luteinizing hormone, and follicular atresia. Biol Reprod,1994,51:934-944
95. Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization: a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA, 1996,93(12):6025-6030.
96.郑邈,刘文励.抑制性消减杂交应用的研究进展.基础医学与临床.2006,Vol 26,No11：1276-1280.
97. Clontech Laboratories, Inc. PCR-SelectTM cDNA Subtraction Kit User Manual. Version No. PR732188.,2007, www.clontech. com
98. Matzuk MM, Lamb DJ. Genetic dissection of mammalian fertility pathways. Nature Cell Biol (Suppl),2002,4:s41-s49
99. Bedard J, Brule S, Price CA, et al. Serine protease inhibitor-E2 (SERPINE2) is differentially expressed in granulose cells of dominant follicle in cattle. Mol Reprod Dev,2003,64:152-165.
100. Silverman GA, Bird PI, Carrell RW, et al. The serpins are an expanding superfamily of structurally of similar but functionally diverse proteins. Evolution, mechanism of inhibition, novel functions, and a revised nomenclature. J Biol Chem,2001,276:33293-33296.
101. Fayad T, Levesque V, Sirois J, et al. Gene expression profiling of differentially expressed genes in granulose cells if bovine dominant follicles using suppression subtractive hybridization. Biol Reprod,2004,70:523-533.
102. Sisco B, Hagemann LJ, Shelling AN, et al. Isolation of genes differentially expressed in dominant and subordinate bovine follicles. Endocrinology,2003,144:3904-3913.
103. Ndiaye K, Fayad T, Silversides DW, et al. Identification of downregualted messenger RNAs in bovine granulose cells of dominant follicles following stimulation with human chorinoic gonadotropin. Biol Reprod,2005,73:324-333.
104. Mihm M, Baker PJ, Fleming LM, et al. Differentiation of the bovine dominant follicle from the cohort upregulates mRNA expression for new tissue development genes. Reproduction,2008, 135:253-265.
105. Zieiak AE, Forde N, Park SDE, et al. Identification of novel genes associated with domiant follicle development in cattle. Reprod Fertilit Develop,2007,19:967-975.
106. Qin Chen, A., Zheng GuangWang, Zi Rong Xu, et al. Analysis of gene expression in granulosa cells of ovine antral growing follicles using suppressive subtractive hybridization. Animal Reproduction Science,2009,115:39-48.
107. Robert C, Gagne D, Bousquet D, et al. Differential display and suppressive subtractive hybridization used to identify granulose cell messenger RNA associated with bovine oocyte developmental competence. Biol Reprod,2001,64:1812-1820.
108. Espey LL, Richards JS. Temporal and spatial patterns of ovarian gene transcription following:an ovulatory dose of gonadotropin in the rat. Biol Reprod,2002,67:1662-1670.
109. Richards JS, Russell DL, Ochsner S, et al. Ovulation: new dimensions and new regulators of the inflammatory-like response. Annu Rev Physiol.,2002,64:69-92
110. Liu SF, Li HB, Song XM, et al. Candidate genes with ovulation by differential display PCR in small tail Han sheep. Asian-Aust J Anim Sci,2006,19:1229.
111. Hourvitz A, Gershon E, Hennebold JD, et al. Ovulation-selective genes: the generation and characterization of an ovulatry-selective cDAN library. J Endocrinol,2006,188:531-548.
112. Edry I, S Sela-Abramovich, and N Dekel. Meiotic arrest of oocytes depends on cell-to-cell communication in the ovarian follicle. Mol Cell Endocrinol,2006, June 27,252(1-2):102-6.
113. Wyllie AH. Apoptosis and the regulation of cell numbers in normal and neoplastic tissues:an overview. Cancer Metastasis Rev,1992,11(2):95-103.
1. Abdulkhaliq A M, W R Harvey, and C F Parker. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. J. Anim. Sci,1989,67:3:250.
2. Dickerson G E. Efficiency of animal production-molding the biological components. J. Anim. Sci., 1970,30:849.
3.赵有璋.羊生产学.北京：中国农业出版社,1999,99-100.
4.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
5. Abdulkhaliq A M. Harvey W R, Parker C F. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. Journal of Animal Science,1989,67:3250-3257.
6.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28：418-423.
7. Waldron D F. and D L Thomas. Increased litter size in Rambouillet sheep:1. Estimation of genetic parameters. J. Anim. Sci.,1992,70:3333-3344.
8. Juan Altarriba, Luis Varona, Luis A Garcia-Cortes, et al. Bayesian Inference of Variance Components for Litter Size in Rasa Aragonesa Sheep. J. Anim. Sci.,1998,76:23-28
9. Bunge R, D L Thomas and T G Nash. Performance of Hair Breeds and Prolific Wool Breeds of Sheep in Southern Illinois:Lamb Production of F1 Adult Ewes. J. Anim. Sci.,1995,73:1602-1608
10. Cumming I A. Relationships in the sheep of ovulation rate with live weight, breed, season and plane of nutrition. Aust. J. Exp. Agric,1977,17:234.
11. Dickerson G E, H A Glimp, H J Tuma, et al. Genetic resources for efficient meat production in sheep. Growth and carcass characteristics of ram lambs of seven breeds. J. Anim. Sci.,1972,34:940.
12. Gunn R G, J M Doney, W F Smith, et al. Effects of age and its relationship with body size on reproductive performance in Scottish Blackface ewes. Anim. Prod.,1986,43:279.
13. ANTONIO GONZALEZ-BULNES, CARLOS J H SOUZA, BRUCE K CAMPBELL, et al. Effect of Ageing on Hormone Secretion and Follicular Dynamics in Sheep with and without the Booroola Gene. Endocrinology.,2004,145(6):2858-2864
14. Waldron D F, and D L Thomas. Increased litter size in Rambouillet sheep: I. Estimation of genetic parameters. J. Anim. Sci,1992,70:3333.
15. Hanrahan J P. Ovulation rate as the selection criterion for litter size in sheep. Proc. Aust. SOCA. Anim. Prod,1980,13:405.
1. Baird, D. T. and B. K. Campbell.1998. Follicle selection in sheep with breed differences in ovulation rate. Mol. Cell. Endocrinol.145:89-95.
2. Nett, T. M., A. M. Turzillo, M. Baratta and L. A. Rispoli.2002.Pituitary effects of steroid hormones on secretion of follicle-stimulating hormone and luteinizing hormone. Domest. Anim. Endocrinol. 23:33-42.
3 Savio, J. D., W. W. Thatcher, L. Badinga, de la Sota, R. L. and D. Wolfenson.1993a. Regulation of dominant follicle turnover during the oestrous cycle in cows. J. Reprod. Fertil.97:197-203.
4. Mann, G. E., A. S. McNeilly and D. T. Baird.1992.Hormone production in vivo and in vitro from follicles at different stages of the estrous cycle in sheep. J. Endocrinol.132:225-234.
5. de Castro, T., E. Rubianes, A. Menchaca and A. Rivero.1999. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology 52:399-411.
6. Martin, G. B., R. J. Scaramuzzi and J. D. Henstridge.1983.Effects of oestradiol, progesterone and androstenedione on the pulsatile secretion of luteinizing hormone in ovariectomized ewes during spring and autumn. J. Endocrinol.96:181-193.
7. Palta, P., N. Bansal, R. S. Manik, R. S. Manik, B. S. Prakash and M. L. Madan.1998. Interrelationships between follicular size, estradiol-17β (beta), progesterone and testosterone concentrations in individual buffalo ovarian follicles. Asian-Aust. J. Anim. Sci.11:293-299.
8. Gibbons, J. R., K. Kot, D. L. Thomas, M. C. Wiltbank and O. J. Ginther.1999. Follicular and FSH dynamics in ewes with a history of high and low ovulation rates. Theriogenology 52:1005-1020.
9. Vinoles, C., M. Forsberg, G. Banchero and E. Rubianes.2002.Ovarian follicular dynamics and endocrine profiles in Polwarth ewes in high and low body condition. Anim. Sci.74:539-545.
10. Rubianes, E. and A. Menchaca.2003.The pattern and manipulation of ovarian follicular growth in goats. Anim. Reprod. Sci.78:271-287.
11. Cueto, M., A. Gibbons, R. Alberio, H. Taddeo and A. Gonzalez-Bulnes.2006. Timing of emergence of ovulatory follicles in polyovulatory goats. Anim. Reprod. Sci.91:275-284.
12. Souza, C. J. H., L. Keenam, M. P. Boland and J. F. Roche.1997. Follicular dynamics and ovarian steroid secretion in sheep during the follicular and early luteal phases of the estrous cycle. Biol. Reprod.56:483-488.
13. Campbell, B. K., D. T. Baird, C. J. H. Souza and R. Webb.2003.The FecB (Booroola) gene acts at the ovary:in vivo evidence. Reproduction 126:101-111.
14. Scaramuzzi, R. J. and H. M. Radford.1983.Factors regulating ovulation rate in the ewe. J. Reprod. Fertil.69:353-367.
15. Souza, C. J. H., B. K. Campbell, D. T. Baird and R. Webb.1997a. Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology 138:5333-5340.
16. Rawlings, N. C., A. C. O. Evans, A. Honaramooz and P. M. Bartlewski.2003.Antral follicle growth and endocrine changes in prepubertal cattle, sheep and goats. Anim. Reprod. Sci.78:259-270.
17. Xia, Y., T. O'Shea, R. Murison and J. R. McFarlane.2003.Concentrations of progesterone, follistatin and follicle-stimulating hormone in peripheral plasma across the estrous cycle and pregnancy in merino ewes that are homozygous or noncarriers of the Booroola gene. Biol. Reprod.69:1079-1084.
18. Bindon, B. M., L. J. Cummins, L. R. Piper and T. O'Shea.1981.Relation between ovulation rate and plasma progesterone in Merinos with natural and induced high fecundity. Proc. Aust. Soc. Reprod. Biol.13:79 (Abstr.).
19. McNatty, K. P., S. Lun, D. A. Heath, K. Ball, P. Smith, N. L. Hudson, J. McDiarmid, M. Gibb and K. M. Henderson.1986. Differences in ovarian activity between BooroolaxMerino ewes which were homozygous, heterozygous and non-carrier of a major gene influencing their ovulation rate. J. Reprod. Fertil.77:193-205.
20. Niswender, G. D., K. P. McNatty, P. Smith, K. D. Niswender, C. E. Farin and H. R. Sawyer. 1990.Numbers of steroidogenic luteal cells in Booroola Merino ewes. J. Reprod. Fertil.90:185-190.
21. Menchaca, A. and E. Rubianes.2002.Relation between progesterone concentrations during the early luteal phase and follicular dynamics in goats. Theriogenology 57:1411-1419.
22. Adams, G. P., R. I. Matteri and O. J. Ginthe.1992.Effect of progesterone on ovarian follicles, emergence of follicular waves and circulating follicle stimulation hormone in heifers. J. Reprod. Fertil.67:403-410.
23. Rubianes, E., T. de Castro and B. Carbajal.1996. Effect of high progesterone levels during the growing phase of the dominant follicle of Wave 1 in ultrasonically monitored ewes. Can. J. Anim. Sci.76:473-475.
24. Camp, J. C., D. E. Wildt, P. K. Howard, L. D. Stuart and P. K. Chakraborty.1983.Ovarian activity during normal and abnormal length estrous cycles in the goat. Biol. Reprod.28:673-681.
25. Souza, C. J. H., J. C. F. Moraes and L. M. Chagas.1994. Effect of the Booroola gene on time of ovulation and ovulatory dynamics. Anim. Reprod. Sci.37:7-13.
26. Ginther, O. J. and K. Kot.1994. Follicular dynamics during the ovulatory season in goats. Theriogenology 42:987-1001.
27. Montgomery, G. W., K. P. McNatty and G. H. Davis.1992.Physiology and molecular genetics of mutations that increase ovulation rate in sheep. Endocr. Rev.13:309-328.
1. Ginther O J, Beg M A, Donadeu F,X, et al Mechanism of follicle deviation in monovular farm species. Animal Reproduction Science,2003,78:239-257
2. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287.
3. Gaskins C T, Snowder G D, Westman M K et al. Influence of body weight, age, and weight gain on fertility and prolificacy in four breeds of ewe lambs. Journal Animal Science,2005,83:1680-1689.
4. Cueto M, Gibbons A, Alberio R, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
5. Camp J C, Wildt D E, Ho Wardl P K, et al. Ovarian Activity During Normal and Abnormal Length Estrous CyCles in the Goat. Biology of reproduction,1983,28:673-681.
6. Scaramuzzi R J and Turnbull K E. Luteal size and function in the Booroola Merino. Socence Reproduction Biology,1981,13:77.
7. Sanchez T, Wehrman ME, Kojima FN, et al. Dosage of synthetic progestin, Norgestomet, influences luteinizing hormone pulse frequency and endogenous secretion of 17_-estradiol in heifers. Biol. Reprod,1995,52:464-469.
8. Johnson SK, Dailey RA, Inskeep EK, et al. Effect of peripheral concentrations of progesterone on follicular growth and fertility in ewes. Dom. Anim. Endocrinol,1996,13:69-79.
9. Alejo Menchaca, Edgardo Rubianes. Relation between progesterone concentrations during the early luteal phase and follicular dynamics in goats. Theriogenology,2002,57:1411-1419
10. Gonzalez-Bulnes A, Di az-Delfa C, Garcia-Garcia R M, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245.
11. Dukelow W R, Jarosz S J, Jewett D A, et al. Laparoscopic examination of the ovaries in goats and primates. Laboratory Animal Science,1971,21:594-597.
12. Mohamed S Medan, Gen Watanabe, Kazuaki Sasaki, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. Biology of Reproduction,2003,69:57-63.
13. Mohamed S. Medan, Gen Watanabe, et al. Follicular and Hormonal Dynamics during the Estrous Cycle in Goats. Jounal of Reproduction and Development,2005,51.
14. Ginther OJ, Kot K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42:987-1001.
15. Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003:78:271-287
16. de Castro T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52:399-411.
17. McNatty KP, Henderson KM, Lun S, et al. Ovarian activity in Booroola x Romney ewes which have a major gene influencing their ovulation rate Journal of Reproduction and Fertility,1985,73: 109-120
18. McNatty KP, Lun S, Heath DA, et al. Differences in ovarian activity between Booroola x Merino ewes which were homozygous, heterozygous and non-carriers of a major gene influencing their ovulation rate Journal of Reproduction and Fertility,1986b,77:193-205
19. Campbell B K, D T Baird, C J H Souza et al. The FecB (Booroola) gene acts at the ovary:in vivo evidence. Reproduction,2003,126:101-111.
20. Bartlewski, PM, Beard AP, Cook SJ, et al. Ovarian antral follicular dynamics and their relationships with endocrine variables throughout the oestrous cycle in breeds of sheep differing in prolificacy. J. Reprod. Fertil.1999:115:111-124.
21. Evans AC, Duffy P, Hynes N, et al. Waves of follicle development during the estrous cycle in sheep. Theriogenology,2000,53:699-715.
22. Orita J, Tanaka T, Kamonae H, et al. Ultrasonography observation of follicular and luteal dynamics during the oestrous cycle in Shiba goats. J. Reprod. Dev,2000,46:31-37.
23. Schrick FN, Surface RA, Pritchard JY, et al. Ovarian structures during the estrous cycle and early pregnancy in ewes. Biol. Reprod,1993:49:1133-1140.
24. Gonzalez-Bulnes A, C Diaz-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245.
1.王桂敏,马春玲,韩云明等.输卵管结礼术对大鼠子宫胞液雌二醇受体和孕酮受体的影响.河北医科大学学报,1996,(01).
2. Garm O A. Study of bovine nymphomania. Acta Endocrinol,1949,3(3).
3. Seungjoon Kim, Tomomi Tanaka and Hideo Kamomae. Different Effects of Subnormal Levels of Progesterone on the Pulsatile and Surge Mode Secretion of Luteinizing Hormone in Ovariectomized Goats. Biology of Reproduction 2003,(69):141-145.
4. Adams N R, S Atkinson, R M Hoskinson, et al. Immunization of ovariectomized ewes against progesterone, oestrogen or cortisol to detect effects of adrenal steroids on reproduction. J. Reprod. Fert,1990,89:477-483.
5.杨利国.动物繁殖学.北京：中国农业出版社,2003：35-67.
6. Shaikh AA & Shaikh SA. Adrenal and ovarian steroid secretion in the rat estrous cycle temporally related to gonadotropins and steroid levels found in peripheral plasma. Endocrinology,1975,96,44.
7. Joseph I B J K, W D Currie and N C Rawlings. Effects of time after ovariectomy, season and oestradiol on luteinizing hormone and follicle-stimulating hormone secretion in ovariectomized ewes. J. Reprod. Fert,1992,94,511-523.
8. Ellison P T. Correlations of basal oestrogens with adrenal androgens and relative weight in normal women. Annls Hum. Biol,1984,11:327-336.
9. Lothar Trgaer. Steroid hormone biosynthese, stoffwechsel, wirkung. Springer-Verlag Berlin Heidelberg New York,1977:92-147.
10.王明运 主编.激素生物化学(第一版).北京：人民卫生出版社,1987年10月：335.
11.王镜岩,朱圣庚,徐长法.生物化学(第三版,上册).北京,高等教育出版社,2002：556-583.
12.King BF, JH Britt, K LEsbenshade,et al. Evidence for a local role of inhibin or inhibinasubunits in compensatory ovarian hypertrophy. Journal of reproduction and fertility,1995,104:291-295.
1.曹贵方,张涌.动物细胞凋亡与卵泡闭锁.动物医学进展,1997,18(4)：5-8.
2.曹攸梅.卵巢的显微结构及不同季节的卵巢活动.动物学研究,1990,1(1)：17-25.
3. Irving-Rodgers HF, van Wezel IL, Mussard ML, et al. Atresia revisited:two basic patterns of atresia of bovine antral follicles. Reproduction,2001,122,761-775
4. Ginther OJ, Kot K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42:987-1001.
5. de Castro T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52:399-411.
6. Tenorio Filho F, MHB Santos, PG Carrazzoni, et al. Follicular dynamics in Anglo-Nubian goats using transrectal and transvaginal ultrasound. Small Ruminant Research,2007,72:51-56.
7. Mohamed S. Medan, Gen Watanabe, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. BIOLOGY OF REPRODUCTION,2003,69:57-63.
8. Ginther O J, Beg M A, Donadeu F X, et al. Mechanism of Follicle Deviation in Monovular Farm Species. Animal Reproduction Science,2003,78:239-257.
9. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287.
10. Cueto M, A Gibbons, R Alberio, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
11. Gonzalez-Bulnes A, C Diaz-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science.,2005,86:237-245.
12. Camp J C, Wildt D E, Ho Wardl P K, et al. Ovarian Activity During Normal and Abnormal Length Estrous CyCles in the Goat. Biology of reproduction,1983,28:673-681.
13.杨世柱.羊的超数排卵及其内分泌研究进展.国外畜牧学-草食家畜,1989,6：13-16.
14.孙青原,谭景和,杨增明,等.山羊卵母细胞闭锁的超微结构研究.兽医大学学报,1993,13(2)：129-132.
15. Billig H. Endocrenology.1994:134-245.
16. Milis, Noborum, Yoshihirok. Ultrastractural changes in granulose cells in porcrine antral follicles undergoing atresia indicate apoptotic cell death. J Reprod Dev,1998,44:7-14.
17. Yuan CHENG, Naoko INOUE, Fuko MATSUDA-MINEHATA, et al. Changes in expression and localization of connexin 43 mRNA and protein in porcine ovary granulosa cells furing follicular atresia. Journal of Reproduction and development,2005, vol.51:627-637.
18.于元松.山羊卵泡发育、闭锁及其调控机理的研究.东北农业大学博士论文,2002.
19. Alngel Palumbo and John Yem. In situ location of apoptosis in the rat ovary during follicular atresia. Journal of cell bioligy. Animal Reproduction Science,1994,51:888-895.
20. Jolly. Apoptosis in bovine granulose cells in relation to steroid synthesis cyclic adenosine 3'-5'-monophosphute response to fouicle stimulating hormone and luteiniting hormone and follicular atresia. Biol Reprod,1994,51:934-944.
1. Ginther, O.J., Kot, K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42,987-1001.
2、Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78,271-287
3、Cueto, M., A. Gibbons, R. Alberio, et al. Gonzalez-Bulnes. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
4、Rrichards JS, Russell DL, Ochsner S, et al. Novel signaling pathways that control ovarian follicular development, ovulation, and luteinization. Recent Progress in Hormone Research,2002,57 195-220.
5、 Souza, C.J.H., C MacDougall, B K Campbell, et al. The Booroola (FecB) phenotype is associated with a mutation in the bone morphogenetic receptor type 1 B (BMPR1B) gene. Journal of Endocrinology,2001,169, R1-R6.
6、姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
7、 Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization:a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA, 1996,93(12):6025-6030.
8、Fayad T, Levesque V, Sirois J, et al. Gene expression profiling of differentially expressed genes in granulose cells if bovine dominant follicles using suppression subtractive hybridization. Biol Reprod, 2004,70:523-533.
9. Leon J Spicer, Pauline Y Aad, Dustin T Allen, et al. Growth Differentiation Factor 9 (GDF9) Stimulates Proliferation and Inhibits Steroidogenesis by Bovine Theca Cells:Influence of Follicle Size on Responses to GDF9. BIOLOGY OF REPRODUCTION,2008,78,243-253.
10. Diatchenko L, Lau Y FC, Campbell AP, et al. Suppression subtractive hybridization:A method for generating differentially regulated or tissue-specific cDNA probes and libraries. Proc. Natl. Acad. Sci,USA.,1996,93:6025-6030.
11. Gurskaya NG, Diatchenko L, Chenchik A, et al. Equalizing cDNA subtraction based on selective suppression of polymerase chain reaction: Cloning of Jurkat cell transcripts induced by phytohemaglutinin and phorbol 12-myristate 13-acetate. Amal. Bilchem,1996,240:90-97.
12. Manabe N, Goto Y, Matsuda-Minehata F, et al.2004. Regualtion mechanism of selective atresia in porcine follicles:regulation of granulose cell apoptosis during atresia. J Reprod Deve.50:493-514.
13. Espey LL, Richards JS. Temporal and spatial patterns of ovarian gene transcription following an ovulatory dose of gonadotropin in the rat. Biol Reprod,2002,67:1662-1670.
14. Qin Chen, A., Zheng GuangWang, Zi Rong Xu, et al. Analysis of gene expression in granulosa cells of ovine antral growing follicles using suppressive subtractive hybridization. Animal Reproduction Science,2009,115:39-48.
15. Ndiaye K, Fayad T, Silversides DW, et al. Identification of downregualted messenger RNAs in bovine granulose cells of dominant follicles following stimulation with human chorinoic gonadotropin. Biol Reprod,2005,73:324-333.
16. Diorf MN, Lefebvre R, Silversides DW, et al. Induction of alpha-caveolin-1 (alpha CAV1) expression in bovine granulosa cells in response to an ovulatory dose of human chorionic gonadotrophins. Mol Reprod,2006,73:1353-1360
17. Clontech Laboratories, Inc. PCR-SelectTM cDNA Subtraction Kit User Manual. Published 16 April 2007:4-8
18. Clontech Laboratories, Inc. Super SMARTTM PCR cDAN Synthesis Kit User Manual. Published 16 April 2007:13-30.
19.何鹏.抑制性消减杂交技术及其应用研究进展.国外医学临床生物化学与检验学分册.2004第25卷第4期：356-358.
20. Willnow TE, Nykjaer A, Herz J. Lipoprotein receptors:new roles for ancient proteins. Nat Cell Biol, 1999,1.-E157-162.
21. Strickland DK, Gonias SL, Argraves WS. Diverse roles for the LDL receptor family. Trends Endocrin Met,2002,13:66-74.
22. Tania Fayad, Rejean Lefebvre, Johannes Nimpf, et al. Low-Density Lipoprotein Receptor-Related Protein 8 (LRP8) Is Upregulated in Granulosa Cells of Bovine Dominant Follicle:Molecular Characterization and Spatio-Temporal Expression Studies. BIOLOGY OF REPRODUCTION,2007, 76,466-475.
23. FLORA RABAHI, SOPHIE BRuLE, JEAN SIROIS, et al. High Expression of Bovine a Glutathione S-Transferase (GSTA1, GSTA2) Subunits Is Mainly Associated with Steroidogenically Active Cells and Regulated by Gonadotropins in Bovine Ovarian Follicles. Endocrinology,1999:140: 3507-3517.
24. Rapoport R, Sklan D, Hanukoglu I. Electron leakage from the adrenal cortex mitochondrial P450scc and P450c11 systems:NADPH and steroid dependence. Arch Biochem Biophys,1995, 317:412-416.
25. Poli G, Dianzani MU, Cheeseman KH, et al. Separation and characterization of the aldehydic products of lipid peroxidation stimulated by carbon tetrachloride or ADP-iron isolated hepatocytes and rat liver microsomal suspensions. Biochem J,1985,227:629-638.
26. Kaur K, Salomon RG, O'Neil J, et al. (Carboxyalkyl) pyrroles in human plasma and oxidized low-density lipoproteins. Chem Res Toxicol,1997,10:1387-1396.
27. Julie Bedard, Sophie Brle, Christopher A. Price, et al. Serine protease inhibitor-E2 (SERPINE2) is differentially expressed in granulosa cells of dominant follicle in cattle. Mol. Reprod. Dev,2003,64: 152-165.
28. Hasan S, Hosseini G, Pricivalle M, et al. Coordinate expression of antivoagulant hepatin sulfate proteoglycans and serine protease inhibitors in the rat ovary:Apotent system of proteolysis control. Biol Peprod,2002,66:144-158.
29. Cao M, Nicola E, Portela VM, et al. Regulation of serine protease inhibitor-E2 and plasminogen activator expression and secretion by follicle stimulating hormone and growth factors in non-luteinizing bovine granulose cells in cells in vitro. Matrix Biol,2006,25:342-356.
30. Harris A L. Emerging issues of connexin channels:biophysics fills the gap. Quart. Rev. Biophys. 2001,34,325-472.
31. Joanne E I, Gittens, Kevin J Barr, et al. Interplay between paracrine signaling and gap junctional communication in ovarian follicles. Journal of Cell Science,2005,118,113-122.
2. Shelton, M.1971. Some factors affecting efficiency of lamb production. Texas Agric. Exp. Sta. Tech. Rept., No.26.
3. Gabinna, D.1989b. Improvement of the reproductive performance of Rasa Aragonesa flocks in frequent lambing systems. Ⅱ. Repeatability and heritability of sexual precocity, fertility and litter size. Livest. Prod. Sci.,22:87.
4.蒋英,陶雍.中国山羊.西安：陕西科学技术出版社,1988,102-104.
5.杨利国,周东蕊.奶牛繁殖力的影响因素及提高措施.中国奶牛,2001,(1)：52-54
6. Van Vleck, L D, E J Pollak, and E A. Oltenacu. Genetics for the Animal Sciences. W. H. Freeman and Co., New York,1987.
7. Lasley, J F. Genetics of Livestock Improvement (4th Ed.). Prentice-Hall, Hemel Hempstead, U. K., 1987.
8. Abdulkhaliq A M, Harvey W R, Parker C F. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. J. Anim. Sci.,1989,67:3250-3257.
9. Fahmy M H. Development of DLS breed of sheep:genetic and phenotypic parameters of date of lambing and number of lambs born alive. Can. J. Anim. Sci.,1990,70:771-778.
l0.Altarriba J, Varona L, Crarcia-Comes L A et al. Bayesian inference of variance components for number of lambs born alive in Rasa Aragonesa sheep. J. Anim. Sci.,1998,76:23-28.
11.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28(5)：418-423.
12. SHUNICHI SHIMASAKI, R. KELLY MOORE, FUMIO OTSUKA, et al. The Bone Morphogenetic Protein System In Mammalian Reproduction. Endocrine Reviews,2004,25:72-101.
13. Souza C J H, C MacDougall, B K Campbell, et al. The Booroola (FecB) phenotype is associated with a mutation in the bone morphogenetic receptor type 1 B (BMPR1B) gene. Journal of Endocrinology,2001,169, R1-R6
14.姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
15. Philippe Monget, Stephane Fabre, Philippe Mulsant, et al. Regulation of ovarian folliculogenesis by IGF and BMP system in domestic animals. Domestic Animal Endocrinology,2002,23:139-154
16. Campbell,B K, D T Baird, C J H Souza, et al. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction,2003,126:101-111
17. Souza C J H, B K Campbell, Baird, D T. et al. Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology, 1997a,138:5333-5340.
18. Piper L R, Bindon B M and Davis G H. The single gene inheritance of the high litter size of theBooroola Merino. Genetics of Reproduction in sheep,1985,38(2):115-125.
19. McNtty K P and Henderson K M. Gonadotrophins, fecundity genes and ovarian follicular function. Journal of steroid Biochemistry,1987,27:365-373.
20. Davis G H, Dodds K G, Wheeler R, et al. Evidence that an imprinted gene on the X chromosome increases ovulation rate in sheep. Biology of Reproduction,2001,64:216-221.
21. Jonmundosson JV and Adalstinsson S. Single genes for fecundity in Icelandic sheep. In:R. B. Land and D. W. Robinson (Ed) Genetics of Reproduction in sheep.1985,159-168.
22. Davis G H, McEwan J C, Fennessy P F, et al. Infertility due to bilateral ovarian hypoplasia in sheep homozygous (FecXI FecXH) for the Inverdale prolificacy gene located on the X chromosome. Biology of Reproduction,1992,46:636-640.
23. Davis GH, McEwan JC, Fennessy PF, et al. Evidence for the presence of a major gene influencing ovulation rate on the X chromosome of sheep. Biology of Reproduction,1991,44:620-640.
24.柳淑芳,姜运良,杜立新BMPR-IB和BMP15基因作为小尾寒羊多胎性能候选基因的研究.遗传学报,2003,30(8)：755-760.
25.王根林,毛鑫智,Davis G H,等.DAN分析发现我国湖羊和小尾寒羊存在Booroola(FecB)多胎基因.南京农业大学学报,2003,26(1)：104-106.
26. Matzuk MM, Lamb DJ. Genetic dissection of mammalian fertility pathways. Nature Cell Biol (Suppl),2002,4:s41-s49
27.桑润滋主编.动物繁殖生物技术(第二版).北京：中国农业出版社,2006年7月,25-42.
28. Markstrom E, Svernsson Ech, Shao R, et al. Survival factors regulating ovarian apoptosis-dependence on follicle differentiation. Reproduction,2002,123:23-30
29. Manabe N, Goto Y, Matsuda-Minehata F, et al. Regualtion mechanism of selective atresia in porcine follicles:regulation of granulose cell apoptosis during atresia. J Reprod Deve,2004,50:493-514
30.蒋英编著.世界养羊科学及生产(第一版).北京：中国农业大学出版社,1999.
31.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
32. Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization:a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA,1996,93(12):6025-6030.
1.姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
2.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28(5)：418-423.
3.谢庄,程瑞禾,韩玉刚.湖羊公羊产羔效应的统计学分析.南京农业大学学报,1998,21(1)：77-81
4. Waldron D F and D L. Thomas. Increased litter size in Rambouillet sheep:I. Estimation of genetic parameters. J. Anim. Sci.1992,70:3333-3344.
5. Shelton M. and J W Menzies. Genetic parameters of some performance characteristics of range fine-wool ewes. J. Anim. Sci,1968,27:1219.
6.中国羊品种志编写组.中国羊品种志.上海：上海科学技术出版社,1989,29-58.
7.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
8.朱东华,徐忠,管生.长江三角洲白山羊繁殖性能调查研究.中国草食动物,2005年第25卷第2期：35-36
9.陈启康,钱鹤良,沙文锋,等.海门山羊多胎繁殖性能和改良利用的研究.南京农专学报,2003,18(1)：35-39.
10.李金林,冉汝俊.鲁北白山羊多胎早熟性能选育研究.中国养羊,1997,4：12-14.
11.徐云华,曹洪防,李强,等.莱芜黑山羊高繁品系培育.山东畜牧兽医,2001,6：13-14.
12.王维春,熊朝瑞,贾旌旗,等.南江黄羊肉用品种选育研究.四川畜牧兽医,2000,27(7)：40-42.
13.Theresa Wilson, Xi-Yang Wu, Jennifer L, et al. Highly Prolific Booroola Sheep Have a Mutation in the Intracellular Kinase Domain of Bone Morphogenetic Protein IB Receptor (ALK-6) That Is Expressed in Both Oocytes and Granulosa Cells. BIOLOGY OF REPRODUCTION,2001,64, 1225-1235.
14. Hanrahan J P. Crossbreeding studies involving Finnish Landrace and Galway sheep. Proc. Working Symp. Breed Evaluation and Crossing Experiments, Zeist,1974. pp 431-444.
15. Land R B, W S Russell and HP Donald. The litter size and fertility of Finnish Landrace and Tasmanian Merino sheep and their reciprocal crosses. Anim. Prod,1974,18:265.
16. Ricordean G, L Tchamitchian, J Thimonier, et al. First survey of results obtained in France on reproductive and maternal performance in sheep,with particular reference to the Romanov breed and crosses with it. Livest. Rod. Sci,1978,5:181.
17. Bradford G E, A Lahlou-Kassi, Y M Berger, et al. Performance of D'man and Sardi breeds of sheep on accelerated lambing. II. Ovulation rate and embryo survival. Small Ruminant Res,1989,2:241.
18. Davis GH, Montgomery GW, Allison AJ, et al. Segregation of a major gene influencing fecundity in progeny of Booroola sheep. NZ J Agric Res,1982,25:525-529
19. McNatty KP, Lun S, Heath DA, et al. Differences in ovarian activity between Booroola X Merino ewes which were homozygous, heterozygous or non-carriers of a fecundity gene influencing their ovulation rate. J Reprod Fertil,1986,77:193-205
20.Souza CJH, Moraes JCF, Chagas LM. Effect of Booroola gene on time of ovulation and ovulatory dynamics. Anim Reprod Sci,1994,37:7-13
21.Driancourt MA, Cahill LP, Bindon BM. Ovarian follicular populations and preovulatory enlargement in Booroola and control Merino ewes. J Reprod Fertil,1985,73:93-107
22.McNatty KP, O'Keefe LE, Henderson KM, et al.125I labelled hCG binding characteristics in theca interna and other tissues from Romney ewes and from Booroola X Romney ewes with and without a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1986c,77: 477-488
23.Henderson KM, McNatty KP, O'Keeffe LE, et al. Differences in gonadotrophin stimulated cyclic-AMP production by granulose cells from Booroola X Merino ewes which were homozygous, heterozygous or non-carriers of a fecundity gene influencing their ovulation rate. J Reprod Fertil. 1987,81:395-402
24. Hanrahan J P. Ovulation rate as the selection criterion for litter size in sheep. Proc. Aust. SOCA. nim. Prod,1980,13:405
25. Waldron D F, and D L Thomas. Increased Litter Size in Rambouillet Sheep: Ⅱ. Expected Responses from Alternative Selection. J. Anim. Sci.,1992.70:3345-3350
26. Altarriba J, L Varona, L A Garcia-Cortes, et al. Bayesian Inference of Variance Components for Litter Size in Rasa Aragonesa Sheep. J. Anim. Sci.,1998.76:23-28
27. Bunge R, D L Thomas and T G Nash. Performance of Hair Breeds and Prolific Wool Breeds of Sheep in Southern Illinois:Lamb Production of F1 Adult Ewes. J. Anim. Sci.,1995.73:1602-1608
28. Cumming I A. Relationships in the sheep of ovulation rate with live weight, breed, season and plane of nutrition. Aust. J. Exp. Agric.,1977,17:234.
29. Dickerson G E, H A Glimp, H J Tuma, et al. Genetic resources for efficient meat production in sheep. Growth and carcass characteristics of ram lambs of seven breeds. J. Anim. Sci.,1972,34:940.
30. Gunn R G., J M Doney, W F Smith, et al. Effects of age and its relationship with body size on reproductive performance in Scottish Blackface ewes. Anim. Prod.,1986,43:279.
31.ANTONIO GONZALEZ-BULNES, CARLOS J H SOUZA, BRUCE K CAMPBELL, et al. Effect of Ageing on Hormone Secretion and Follicular Dynamics in Sheep with and without the Booroola Gene. Endocrinology.,2004,145(6):2858-2864
32. Ginther OJ, Kot K, Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42,987-1001.
33. Castro de T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52,399-411.
34. Tenorio Filho F, MHB Santos, PG Carrazzoni, et al. Follicular dynamics in Anglo-Nubian goats using transrectal and transvaginal ultrasound. Small Ruminant Research,2007,72,51-56.
35. Mohamed S. Medan, Gen Watanabe, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. BIOLOGY OF REPRODUCTION,2003,69,57-63.
36. Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287
37. Driancourt MA, Webb R, Fry RC. Does follicular dominance occur in ewes? J. Reprod. Fertil,1991, 93:63-70.
38. Gonzalez-Bulnes A, C Di az-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245
39. Adams G. P, Kot K, Smith CA, et al. Effect of the dominant follicle on regression of its subordinates. Can. J. Anim. Sci,1993,73,267-275.
40. Cueto M, A Gibbons, R Alberio, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
41. Gonzalez-Bulnes A, Santiago-Moreno J, Garcia-Garcia RM, et al. Origin of the preovulatory follicle in Mouflon sheep (Ovis gmelini musimon) and effect on the growth of remaining follicles during the follicular phase of oestrous cycle. Anim. Reprod. Sci,2001,65,265-272.
42. Gonzalez-Bulnes A, Santiago Moreno J, Gomez-Brunet A, et al. Follicular dynamics during the oestrous cycle in dairy goats. Anim. Sci,1999b,68,547-554.
43. Gonzalez-Bulnes A, Souza CJH, Campbell BK, et al. Systemic and intraovarian effects of dominant follicles on ovine follicular growth. Anim. Reprod. Sci,2004,84,107-119
44. EVANS A C O and J E FORTUNE. Selection of the Dominant Follicle in Cattle Occurs in the Absence of Differences in the Expression of Messenger Ribonucleic Acid for Gonadotropin Receptors. Endocrinology,1997,138:2963-2971
45. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Anim. Reprod. Sci,2003,78,271-287.
46. Mohamed S. Medan, Gen Watanabe, et al. Follicular and Hormonal Dynamics during the Estrous Cycle in Goats. Joumal of Reproduction and Development,2005, Vol.51:455-463.
47. Mann GE, McNeilly AS, Baird DT. Hormone production in vivo and in vitro from follicles at different stages of estrous cycle in sheep. J Endocrinol,1992,132:225-234.
48. Kawate N, Morita N, Tsuji M, et al. Roles of pulsatile release of LH in the development and maintenance of corpus luteum function in the goat. Theriogenology,2000,54:1133-1143.
49. Xia Y, T O'Shea, R Murison, et al. Concentrations of Progesterone, Follistatin, and Follicle-Stimulating Hormone in Peripheral Plasma Across the Estrous Cycle and Pregnancy in Merino Ewes That Are Homozygous or Noncarriers of the Booroola Gene. BIOLOGY OF REPRODUCTION,2003,69,1079-1084.
50. Bindon BM, Piper LR, Thimonier J. Preovulatory LH characteristics and time of ovulation in the prolific Booroola Merino ewe. J Reprod Fertil,1984,71:519-523.
51. Bindon BM, Piper LR, Cummins LJ, et al. Reproductive endocrinology of prolific sheep: studies of the Booroola Merino. In Genetics of Reproduction in Sheep.1985, pp 217-235 Eds RB Land and DW Robinson. Butterworths, London.
52. Bindon BM. Reproductive biology of the Booroola Merino sheep. Australian Journal of Biological Sciences,1984,37:163-189.
53. McNatty KP, Hudson N, Henderson KM, et al. Differences in gonadotrophin concentrations and pituitary responsiveness to GnRH between Booroola ewes which were homozygous (FF), heterozygous (F+) and non-carriers (++) of a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1987,80:577-588.
54. Fry RC, Clarke IJ, Cummins JT, et al. Induction of ovulation in chronically hypophysectomized Booroola ewes. Journal of Reproduction and Fertility,1988,82:711-715.
55. Hudson NL, O'Connell AR, Shaw L, et al. Effect of exogenous FSH on ovulation rate in homozygous carriers or non-carriers of the Booroola FecB gene after hypothalamic—pituitary disconnection or after treatment with a GnRH agonist. Domestic Animal Endocrinology,1999,16: 69-80.
56. Boulton MI, Haley CS, Springbett AJ et al. The effect of the Booroola (FecB) gene on peripheral FSH concentrations and ovulation rates during oestrus, seasonal anoestrus and on FSH concentrations following ovariectomy in Scottish Blackface ewes. Journal of Reproduction and Fertility,1995,103:199-207.
57. Souza CJH, Campbell BK, BairdDT andWebb R Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology, 1997a,138:5333-5340.
58. Campbell B K, D T Baird, C J H, et al. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction.,2003,126,101-111.
59. Adams N R, S Atkinson, R M Hoskinson, et al. Immunization of ovariectomized ewes against progesterone, oestrogen or cortisol to detect effects of adrenal steroids on reproduction. J. Reprod. Fert,1990,89:477-483.
60. Ellison PT. Correlations of basal oestrogens with adrenal androgens and relative weight in normal women. Annls Hum. Bio,1984,11,327-336.
61. Shaikh AA & Shaikh SA. Adrenal and ovarian steroid secretion in the rat estrous cycle temporally related to gonadotropins and steroid levels found in peripheral plasma. Endocrinology,1975,96:44.
62. Martin GB. Factors affecting the secretion of luteinizing hormone in the ewe. Biol. Rev,1984,9: 1-87.
63. Webb R, Baxter G, Preece RD, et al. Control of gonadotrophin release in Scottish Blackface and Finnish Landrace ewes during seasonal anoestrus. J. Reprod. Fert.,1985,73:369-378.
64. Goodman RL, Pickover SM & Karsch FJ. Ovarian feedback control of follicle stimulating hormone in the ewe:evidence for selective suppression. Endocrinology,1981a,108:772-777.
65. Rawlings NC, Jeffcoate IA & Rieger DL. The influence of oestradiol-17 β and progesterone on peripheral serum concentration of luteinizing hormone and follicle stimulating hormone in the ovariectomized ewe. Theriogenology,1984,22,473-488.
66. Barlow DH, Coutts JRT, Mowat J and Macnaughton MC. Hormonal profiles at the menopause. In Coutts, J. R. T. (ed.), The Functional Morphology of the Human Ovary. MTP Press, Lancaster, 1981,pp.223-234.
67. Atkinson S & Adams NR. Adrenal glands alter oestrogen activity in the uterus of ewes. J. Endocr., 1988,118:375-380.
68. Pant HC, Dobson H & Ward WR. Effect of active immunization against oestrogens on plasma gonadotrophins in the ewe and the response to synthetic oestrogen on LH. J. Reprod. Fert.,1978,53, 241-248.
69. Joseph I B J K, W D Currie and N C Rawlings. Effects of time after ovariectomy, season and oestradiol on luteinizing hormone and follicle-stimulating hormone secretion in ovariectomized ewes. J. Reprod. Fert,1992,94,511-523.
70.桑润滋 主编.动物繁殖生物技术(第二版).北京,中国农业出版社,2006年7月,25-42.
71. Campbell BK. The modulation of gonadotrophic hormone action on the ovary by paracrine and autocrine factors. Anat Histol Embryol,1999,28(4):247-51. 72. Blondin P, Sirard MA. Oocyte and follicular morphology as determining characteristics for developmental competence in bovine oocytes. Mol Reprod Dev.,1995,41:54-62.
73. Geshi M, Takenouchi N, Yamauchi N, et al. Effects of sodium private in nonserum maturation medium on maturation, fertilization and s ubsequrnt development of bovine oocytes with or without cumulus cells. Biol Reprod.,2000,63:1130-1134.
74. Armstrong D G and Webb R. Ovarian follicular dominance: the role of intraovarian growth factors and novel proteins. Rev Reprod,1997,2(1):139-146
75. Campbell B K, D T Baird, C J H Souza and R Webb. The FecB (Booroola) gene acts at the ovary: in vivo evidence. Reproduction.,2003,126: 101-111
76. McNatty KP, Henderson KM, Lun S, et al. Ovarian activity in Booroola X Romney ewes which have a major gene influencing their ovulation rate. Journal of Reproduction and Fertility,1985,73: 109-120.
77. Hsueh AJW, Billig H and Tsafriri A. Ovarian follicle atresia:a hormonally controlled apoptotic process. Endocr Rev,1994,15:707-724
78. Byskov AG. Follicular atresia. In:Jones RE(ed) The Vertebrate Ovary. Plenum Press, New York, 1978, pp533-562
79. Markstrom E, Svernsson Ech, Shao R, et al. Survival factors regulating ovarian apoptosis-dependence on follicle differentiation. Reproduction.,2002,123:23-30
80. Manabe N, Goto Y, Matsuda-Minehata F, et al. Regulation mechanism of selective atresia in porcine follicles:regulation of granulosa cell apoptosis during atresia. J Reprod Dev.,2004,50(5):493-514.
81. Irving-Rodgers HF, van Wezel IL, Mussard ML, et al. Atresia revisited: two basic patterns of atresia of bovine antral follicles. Reproduction.,2001,122:761-775
82. Chun SY, Eisenhauer KM, Minami S, et al. Hormonal regulation of apoptosis in early antral follicles: follicle-stimulating as a major survival factor. Endocrinology,1996,137:1447-1456
83. Hsu SY, Hsuch AJW. Hormonal regulation of apoptosis. An ovarian perspective. Trends Endocrinol Metab,1997,8:207-213.
84. Wang Y, Chan S, Tsang BK. Role and gonadotropin regulation of X-linked inhibitor of apoptosis expression during ovarian follicular development in vitro. Biol Reprod,2003,68:610-619.
85.杨世华,余世九.生殖激素控制卵泡细胞凋亡的研究进展.动物医学进展,2002,23：1-5.
86. Guthrie HD, Garrett WM, Cooper BS. Follicle-stimulating hormone and insulin-like growth factor-1 attenute apoptosis in cultured porcine grnaulosa cell. Biol Reprod,1998,58:390-396
87. Yu YS, Li W, Han ZB, et al. The effect of follicle-stimulating hormone on follicular development granulose cells apoptosis and steroidogenesis and its mediation by insulin-like growth factor-I in the goat ovary. Theriogenology,2003,60:1691-1704.
88. Manabe N, Imai Y, Myoumoto A, et al. Apoptosis occurs in granulose cells but not cumulius cell in the atretic graafian follicles in multiparous pig ovaries. Acta Histochem Cytochem,1997,30:85-92
89. Manabe N, Imai Y, Ohno H, et al. Apoptosis occurs in granulose cells but not cumulus cells in the atretic antral follicles in the pig ovaries. Experientia,1996,52:647-657
90. Wyllie AH, Kerr JF, Currie AR. Cell death: the significance of apoptosis. Int Rev Cytol.,1980,68: 251-306
91. Palumbo A and Yeh J. In situ localization of apoptosis in the rat ovary during follicular atresia. Biol Reprod.,1994,51:888-95
92. Rosales AM, Avalos A, Vergara M, et al. Multiparametric study of atresia in ewe antral follicles: histology, flow cytometry, internucleosomal DNA fragmentation, and lysosomal enzyme activities in granulose cells and follicular fluid. Mol Reprod Dev,2000,55:270-281
93. Jolly PD, Smith FR, Heath DA, et al. Morphological evidence of apoptosis and the prevalence of apoptotic versus mitotic cells in the membrana granulosa of ovarian follicles during spontaneous and induced atresia in ewes. Biol Reprod.,1997,56:837-846
94. Jolly PD, Tisdall DJ, Heath DA et al. Apoptosis in bovine granulose cells in relation to steroid synthesis, cyclic adenosine 3',5'-monophosphate response to follicle-stimulating hormone and luteinizing hormone, and follicular atresia. Biol Reprod,1994,51:934-944
95. Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization: a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA, 1996,93(12):6025-6030.
96.郑邈,刘文励.抑制性消减杂交应用的研究进展.基础医学与临床.2006,Vol 26,No11：1276-1280.
97. Clontech Laboratories, Inc. PCR-SelectTM cDNA Subtraction Kit User Manual. Version No. PR732188.,2007, www.clontech. com
98. Matzuk MM, Lamb DJ. Genetic dissection of mammalian fertility pathways. Nature Cell Biol (Suppl),2002,4:s41-s49
99. Bedard J, Brule S, Price CA, et al. Serine protease inhibitor-E2 (SERPINE2) is differentially expressed in granulose cells of dominant follicle in cattle. Mol Reprod Dev,2003,64:152-165.
100. Silverman GA, Bird PI, Carrell RW, et al. The serpins are an expanding superfamily of structurally of similar but functionally diverse proteins. Evolution, mechanism of inhibition, novel functions, and a revised nomenclature. J Biol Chem,2001,276:33293-33296.
101. Fayad T, Levesque V, Sirois J, et al. Gene expression profiling of differentially expressed genes in granulose cells if bovine dominant follicles using suppression subtractive hybridization. Biol Reprod,2004,70:523-533.
102. Sisco B, Hagemann LJ, Shelling AN, et al. Isolation of genes differentially expressed in dominant and subordinate bovine follicles. Endocrinology,2003,144:3904-3913.
103. Ndiaye K, Fayad T, Silversides DW, et al. Identification of downregualted messenger RNAs in bovine granulose cells of dominant follicles following stimulation with human chorinoic gonadotropin. Biol Reprod,2005,73:324-333.
104. Mihm M, Baker PJ, Fleming LM, et al. Differentiation of the bovine dominant follicle from the cohort upregulates mRNA expression for new tissue development genes. Reproduction,2008, 135:253-265.
105. Zieiak AE, Forde N, Park SDE, et al. Identification of novel genes associated with domiant follicle development in cattle. Reprod Fertilit Develop,2007,19:967-975.
106. Qin Chen, A., Zheng GuangWang, Zi Rong Xu, et al. Analysis of gene expression in granulosa cells of ovine antral growing follicles using suppressive subtractive hybridization. Animal Reproduction Science,2009,115:39-48.
107. Robert C, Gagne D, Bousquet D, et al. Differential display and suppressive subtractive hybridization used to identify granulose cell messenger RNA associated with bovine oocyte developmental competence. Biol Reprod,2001,64:1812-1820.
108. Espey LL, Richards JS. Temporal and spatial patterns of ovarian gene transcription following:an ovulatory dose of gonadotropin in the rat. Biol Reprod,2002,67:1662-1670.
109. Richards JS, Russell DL, Ochsner S, et al. Ovulation: new dimensions and new regulators of the inflammatory-like response. Annu Rev Physiol.,2002,64:69-92
110. Liu SF, Li HB, Song XM, et al. Candidate genes with ovulation by differential display PCR in small tail Han sheep. Asian-Aust J Anim Sci,2006,19:1229.
111. Hourvitz A, Gershon E, Hennebold JD, et al. Ovulation-selective genes: the generation and characterization of an ovulatry-selective cDAN library. J Endocrinol,2006,188:531-548.
112. Edry I, S Sela-Abramovich, and N Dekel. Meiotic arrest of oocytes depends on cell-to-cell communication in the ovarian follicle. Mol Cell Endocrinol,2006, June 27,252(1-2):102-6.
113. Wyllie AH. Apoptosis and the regulation of cell numbers in normal and neoplastic tissues:an overview. Cancer Metastasis Rev,1992,11(2):95-103.
1. Abdulkhaliq A M, W R Harvey, and C F Parker. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. J. Anim. Sci,1989,67:3:250.
2. Dickerson G E. Efficiency of animal production-molding the biological components. J. Anim. Sci., 1970,30:849.
3.赵有璋.羊生产学.北京：中国农业出版社,1999,99-100.
4.刘振中.槐山羊产羔性能再观察.河南畜牧兽医,1994,15：30-32.
5. Abdulkhaliq A M. Harvey W R, Parker C F. Genetic parameters for ewe productivity traits in the Columbia, Suffolk and Targhee breeds. Journal of Animal Science,1989,67:3250-3257.
6.储明星,王秀利.湖羊出生类型与产活羔数表型及遗传参数估计.遗传学报,2001,28：418-423.
7. Waldron D F. and D L Thomas. Increased litter size in Rambouillet sheep:1. Estimation of genetic parameters. J. Anim. Sci.,1992,70:3333-3344.
8. Juan Altarriba, Luis Varona, Luis A Garcia-Cortes, et al. Bayesian Inference of Variance Components for Litter Size in Rasa Aragonesa Sheep. J. Anim. Sci.,1998,76:23-28
9. Bunge R, D L Thomas and T G Nash. Performance of Hair Breeds and Prolific Wool Breeds of Sheep in Southern Illinois:Lamb Production of F1 Adult Ewes. J. Anim. Sci.,1995,73:1602-1608
10. Cumming I A. Relationships in the sheep of ovulation rate with live weight, breed, season and plane of nutrition. Aust. J. Exp. Agric,1977,17:234.
11. Dickerson G E, H A Glimp, H J Tuma, et al. Genetic resources for efficient meat production in sheep. Growth and carcass characteristics of ram lambs of seven breeds. J. Anim. Sci.,1972,34:940.
12. Gunn R G, J M Doney, W F Smith, et al. Effects of age and its relationship with body size on reproductive performance in Scottish Blackface ewes. Anim. Prod.,1986,43:279.
13. ANTONIO GONZALEZ-BULNES, CARLOS J H SOUZA, BRUCE K CAMPBELL, et al. Effect of Ageing on Hormone Secretion and Follicular Dynamics in Sheep with and without the Booroola Gene. Endocrinology.,2004,145(6):2858-2864
14. Waldron D F, and D L Thomas. Increased litter size in Rambouillet sheep: I. Estimation of genetic parameters. J. Anim. Sci,1992,70:3333.
15. Hanrahan J P. Ovulation rate as the selection criterion for litter size in sheep. Proc. Aust. SOCA. Anim. Prod,1980,13:405.
1. Baird, D. T. and B. K. Campbell.1998. Follicle selection in sheep with breed differences in ovulation rate. Mol. Cell. Endocrinol.145:89-95.
2. Nett, T. M., A. M. Turzillo, M. Baratta and L. A. Rispoli.2002.Pituitary effects of steroid hormones on secretion of follicle-stimulating hormone and luteinizing hormone. Domest. Anim. Endocrinol. 23:33-42.
3 Savio, J. D., W. W. Thatcher, L. Badinga, de la Sota, R. L. and D. Wolfenson.1993a. Regulation of dominant follicle turnover during the oestrous cycle in cows. J. Reprod. Fertil.97:197-203.
4. Mann, G. E., A. S. McNeilly and D. T. Baird.1992.Hormone production in vivo and in vitro from follicles at different stages of the estrous cycle in sheep. J. Endocrinol.132:225-234.
5. de Castro, T., E. Rubianes, A. Menchaca and A. Rivero.1999. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology 52:399-411.
6. Martin, G. B., R. J. Scaramuzzi and J. D. Henstridge.1983.Effects of oestradiol, progesterone and androstenedione on the pulsatile secretion of luteinizing hormone in ovariectomized ewes during spring and autumn. J. Endocrinol.96:181-193.
7. Palta, P., N. Bansal, R. S. Manik, R. S. Manik, B. S. Prakash and M. L. Madan.1998. Interrelationships between follicular size, estradiol-17β (beta), progesterone and testosterone concentrations in individual buffalo ovarian follicles. Asian-Aust. J. Anim. Sci.11:293-299.
8. Gibbons, J. R., K. Kot, D. L. Thomas, M. C. Wiltbank and O. J. Ginther.1999. Follicular and FSH dynamics in ewes with a history of high and low ovulation rates. Theriogenology 52:1005-1020.
9. Vinoles, C., M. Forsberg, G. Banchero and E. Rubianes.2002.Ovarian follicular dynamics and endocrine profiles in Polwarth ewes in high and low body condition. Anim. Sci.74:539-545.
10. Rubianes, E. and A. Menchaca.2003.The pattern and manipulation of ovarian follicular growth in goats. Anim. Reprod. Sci.78:271-287.
11. Cueto, M., A. Gibbons, R. Alberio, H. Taddeo and A. Gonzalez-Bulnes.2006. Timing of emergence of ovulatory follicles in polyovulatory goats. Anim. Reprod. Sci.91:275-284.
12. Souza, C. J. H., L. Keenam, M. P. Boland and J. F. Roche.1997. Follicular dynamics and ovarian steroid secretion in sheep during the follicular and early luteal phases of the estrous cycle. Biol. Reprod.56:483-488.
13. Campbell, B. K., D. T. Baird, C. J. H. Souza and R. Webb.2003.The FecB (Booroola) gene acts at the ovary:in vivo evidence. Reproduction 126:101-111.
14. Scaramuzzi, R. J. and H. M. Radford.1983.Factors regulating ovulation rate in the ewe. J. Reprod. Fertil.69:353-367.
15. Souza, C. J. H., B. K. Campbell, D. T. Baird and R. Webb.1997a. Secretion of inhibin A and follicular dynamics throughout the oestrous cycle in sheep with and without the Booroola gene (FecB). Endocrinology 138:5333-5340.
16. Rawlings, N. C., A. C. O. Evans, A. Honaramooz and P. M. Bartlewski.2003.Antral follicle growth and endocrine changes in prepubertal cattle, sheep and goats. Anim. Reprod. Sci.78:259-270.
17. Xia, Y., T. O'Shea, R. Murison and J. R. McFarlane.2003.Concentrations of progesterone, follistatin and follicle-stimulating hormone in peripheral plasma across the estrous cycle and pregnancy in merino ewes that are homozygous or noncarriers of the Booroola gene. Biol. Reprod.69:1079-1084.
18. Bindon, B. M., L. J. Cummins, L. R. Piper and T. O'Shea.1981.Relation between ovulation rate and plasma progesterone in Merinos with natural and induced high fecundity. Proc. Aust. Soc. Reprod. Biol.13:79 (Abstr.).
19. McNatty, K. P., S. Lun, D. A. Heath, K. Ball, P. Smith, N. L. Hudson, J. McDiarmid, M. Gibb and K. M. Henderson.1986. Differences in ovarian activity between BooroolaxMerino ewes which were homozygous, heterozygous and non-carrier of a major gene influencing their ovulation rate. J. Reprod. Fertil.77:193-205.
20. Niswender, G. D., K. P. McNatty, P. Smith, K. D. Niswender, C. E. Farin and H. R. Sawyer. 1990.Numbers of steroidogenic luteal cells in Booroola Merino ewes. J. Reprod. Fertil.90:185-190.
21. Menchaca, A. and E. Rubianes.2002.Relation between progesterone concentrations during the early luteal phase and follicular dynamics in goats. Theriogenology 57:1411-1419.
22. Adams, G. P., R. I. Matteri and O. J. Ginthe.1992.Effect of progesterone on ovarian follicles, emergence of follicular waves and circulating follicle stimulation hormone in heifers. J. Reprod. Fertil.67:403-410.
23. Rubianes, E., T. de Castro and B. Carbajal.1996. Effect of high progesterone levels during the growing phase of the dominant follicle of Wave 1 in ultrasonically monitored ewes. Can. J. Anim. Sci.76:473-475.
24. Camp, J. C., D. E. Wildt, P. K. Howard, L. D. Stuart and P. K. Chakraborty.1983.Ovarian activity during normal and abnormal length estrous cycles in the goat. Biol. Reprod.28:673-681.
25. Souza, C. J. H., J. C. F. Moraes and L. M. Chagas.1994. Effect of the Booroola gene on time of ovulation and ovulatory dynamics. Anim. Reprod. Sci.37:7-13.
26. Ginther, O. J. and K. Kot.1994. Follicular dynamics during the ovulatory season in goats. Theriogenology 42:987-1001.
27. Montgomery, G. W., K. P. McNatty and G. H. Davis.1992.Physiology and molecular genetics of mutations that increase ovulation rate in sheep. Endocr. Rev.13:309-328.
1. Ginther O J, Beg M A, Donadeu F,X, et al Mechanism of follicle deviation in monovular farm species. Animal Reproduction Science,2003,78:239-257
2. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287.
3. Gaskins C T, Snowder G D, Westman M K et al. Influence of body weight, age, and weight gain on fertility and prolificacy in four breeds of ewe lambs. Journal Animal Science,2005,83:1680-1689.
4. Cueto M, Gibbons A, Alberio R, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
5. Camp J C, Wildt D E, Ho Wardl P K, et al. Ovarian Activity During Normal and Abnormal Length Estrous CyCles in the Goat. Biology of reproduction,1983,28:673-681.
6. Scaramuzzi R J and Turnbull K E. Luteal size and function in the Booroola Merino. Socence Reproduction Biology,1981,13:77.
7. Sanchez T, Wehrman ME, Kojima FN, et al. Dosage of synthetic progestin, Norgestomet, influences luteinizing hormone pulse frequency and endogenous secretion of 17_-estradiol in heifers. Biol. Reprod,1995,52:464-469.
8. Johnson SK, Dailey RA, Inskeep EK, et al. Effect of peripheral concentrations of progesterone on follicular growth and fertility in ewes. Dom. Anim. Endocrinol,1996,13:69-79.
9. Alejo Menchaca, Edgardo Rubianes. Relation between progesterone concentrations during the early luteal phase and follicular dynamics in goats. Theriogenology,2002,57:1411-1419
10. Gonzalez-Bulnes A, Di az-Delfa C, Garcia-Garcia R M, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245.
11. Dukelow W R, Jarosz S J, Jewett D A, et al. Laparoscopic examination of the ovaries in goats and primates. Laboratory Animal Science,1971,21:594-597.
12. Mohamed S Medan, Gen Watanabe, Kazuaki Sasaki, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. Biology of Reproduction,2003,69:57-63.
13. Mohamed S. Medan, Gen Watanabe, et al. Follicular and Hormonal Dynamics during the Estrous Cycle in Goats. Jounal of Reproduction and Development,2005,51.
14. Ginther OJ, Kot K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42:987-1001.
15. Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003:78:271-287
16. de Castro T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52:399-411.
17. McNatty KP, Henderson KM, Lun S, et al. Ovarian activity in Booroola x Romney ewes which have a major gene influencing their ovulation rate Journal of Reproduction and Fertility,1985,73: 109-120
18. McNatty KP, Lun S, Heath DA, et al. Differences in ovarian activity between Booroola x Merino ewes which were homozygous, heterozygous and non-carriers of a major gene influencing their ovulation rate Journal of Reproduction and Fertility,1986b,77:193-205
19. Campbell B K, D T Baird, C J H Souza et al. The FecB (Booroola) gene acts at the ovary:in vivo evidence. Reproduction,2003,126:101-111.
20. Bartlewski, PM, Beard AP, Cook SJ, et al. Ovarian antral follicular dynamics and their relationships with endocrine variables throughout the oestrous cycle in breeds of sheep differing in prolificacy. J. Reprod. Fertil.1999:115:111-124.
21. Evans AC, Duffy P, Hynes N, et al. Waves of follicle development during the estrous cycle in sheep. Theriogenology,2000,53:699-715.
22. Orita J, Tanaka T, Kamonae H, et al. Ultrasonography observation of follicular and luteal dynamics during the oestrous cycle in Shiba goats. J. Reprod. Dev,2000,46:31-37.
23. Schrick FN, Surface RA, Pritchard JY, et al. Ovarian structures during the estrous cycle and early pregnancy in ewes. Biol. Reprod,1993:49:1133-1140.
24. Gonzalez-Bulnes A, C Diaz-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science,2005,86:237-245.
1.王桂敏,马春玲,韩云明等.输卵管结礼术对大鼠子宫胞液雌二醇受体和孕酮受体的影响.河北医科大学学报,1996,(01).
2. Garm O A. Study of bovine nymphomania. Acta Endocrinol,1949,3(3).
3. Seungjoon Kim, Tomomi Tanaka and Hideo Kamomae. Different Effects of Subnormal Levels of Progesterone on the Pulsatile and Surge Mode Secretion of Luteinizing Hormone in Ovariectomized Goats. Biology of Reproduction 2003,(69):141-145.
4. Adams N R, S Atkinson, R M Hoskinson, et al. Immunization of ovariectomized ewes against progesterone, oestrogen or cortisol to detect effects of adrenal steroids on reproduction. J. Reprod. Fert,1990,89:477-483.
5.杨利国.动物繁殖学.北京：中国农业出版社,2003：35-67.
6. Shaikh AA & Shaikh SA. Adrenal and ovarian steroid secretion in the rat estrous cycle temporally related to gonadotropins and steroid levels found in peripheral plasma. Endocrinology,1975,96,44.
7. Joseph I B J K, W D Currie and N C Rawlings. Effects of time after ovariectomy, season and oestradiol on luteinizing hormone and follicle-stimulating hormone secretion in ovariectomized ewes. J. Reprod. Fert,1992,94,511-523.
8. Ellison P T. Correlations of basal oestrogens with adrenal androgens and relative weight in normal women. Annls Hum. Biol,1984,11:327-336.
9. Lothar Trgaer. Steroid hormone biosynthese, stoffwechsel, wirkung. Springer-Verlag Berlin Heidelberg New York,1977:92-147.
10.王明运 主编.激素生物化学(第一版).北京：人民卫生出版社,1987年10月：335.
11.王镜岩,朱圣庚,徐长法.生物化学(第三版,上册).北京,高等教育出版社,2002：556-583.
12.King BF, JH Britt, K LEsbenshade,et al. Evidence for a local role of inhibin or inhibinasubunits in compensatory ovarian hypertrophy. Journal of reproduction and fertility,1995,104:291-295.
1.曹贵方,张涌.动物细胞凋亡与卵泡闭锁.动物医学进展,1997,18(4)：5-8.
2.曹攸梅.卵巢的显微结构及不同季节的卵巢活动.动物学研究,1990,1(1)：17-25.
3. Irving-Rodgers HF, van Wezel IL, Mussard ML, et al. Atresia revisited:two basic patterns of atresia of bovine antral follicles. Reproduction,2001,122,761-775
4. Ginther OJ, Kot K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42:987-1001.
5. de Castro T, Rubianes E, Menchaca A, et al. Ovarian dynamics, serum estradiol and progesterone concentrations during the interovulatory interval in goats. Theriogenology,1999,52:399-411.
6. Tenorio Filho F, MHB Santos, PG Carrazzoni, et al. Follicular dynamics in Anglo-Nubian goats using transrectal and transvaginal ultrasound. Small Ruminant Research,2007,72:51-56.
7. Mohamed S. Medan, Gen Watanabe, et al. Ovarian Dynamics and Their Associations with Peripheral Concentrations of Gonadotropins, Ovarian Steroids, and Inhibin During the Estrous Cycle in Goats. BIOLOGY OF REPRODUCTION,2003,69:57-63.
8. Ginther O J, Beg M A, Donadeu F X, et al. Mechanism of Follicle Deviation in Monovular Farm Species. Animal Reproduction Science,2003,78:239-257.
9. Rubianes E, Menchaca A. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78:271-287.
10. Cueto M, A Gibbons, R Alberio, et al. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
11. Gonzalez-Bulnes A, C Diaz-Delfa, RM Garcia-Garcia, et al. Origin and fate of preovulatory follicles after induced luteolysis at different stages of the luteal phase of the oestrous cycle in goats. Animal Reproduction Science.,2005,86:237-245.
12. Camp J C, Wildt D E, Ho Wardl P K, et al. Ovarian Activity During Normal and Abnormal Length Estrous CyCles in the Goat. Biology of reproduction,1983,28:673-681.
13.杨世柱.羊的超数排卵及其内分泌研究进展.国外畜牧学-草食家畜,1989,6：13-16.
14.孙青原,谭景和,杨增明,等.山羊卵母细胞闭锁的超微结构研究.兽医大学学报,1993,13(2)：129-132.
15. Billig H. Endocrenology.1994:134-245.
16. Milis, Noborum, Yoshihirok. Ultrastractural changes in granulose cells in porcrine antral follicles undergoing atresia indicate apoptotic cell death. J Reprod Dev,1998,44:7-14.
17. Yuan CHENG, Naoko INOUE, Fuko MATSUDA-MINEHATA, et al. Changes in expression and localization of connexin 43 mRNA and protein in porcine ovary granulosa cells furing follicular atresia. Journal of Reproduction and development,2005, vol.51:627-637.
18.于元松.山羊卵泡发育、闭锁及其调控机理的研究.东北农业大学博士论文,2002.
19. Alngel Palumbo and John Yem. In situ location of apoptosis in the rat ovary during follicular atresia. Journal of cell bioligy. Animal Reproduction Science,1994,51:888-895.
20. Jolly. Apoptosis in bovine granulose cells in relation to steroid synthesis cyclic adenosine 3'-5'-monophosphute response to fouicle stimulating hormone and luteiniting hormone and follicular atresia. Biol Reprod,1994,51:934-944.
1. Ginther, O.J., Kot, K. Follicular dynamics during the ovulatory season in goats. Theriogenology,1994, 42,987-1001.
2、Edgardo Rubianes, Alejo Menchaca. The pattern and manipulation of ovarian follicular growth in goats. Animal Reproduction Science,2003,78,271-287
3、Cueto, M., A. Gibbons, R. Alberio, et al. Gonzalez-Bulnes. Timing of emergence of ovulatory follicles in polyovulatory goats. Animal Reproduction Science,2006,91:275-284.
4、Rrichards JS, Russell DL, Ochsner S, et al. Novel signaling pathways that control ovarian follicular development, ovulation, and luteinization. Recent Progress in Hormone Research,2002,57 195-220.
5、 Souza, C.J.H., C MacDougall, B K Campbell, et al. The Booroola (FecB) phenotype is associated with a mutation in the bone morphogenetic receptor type 1 B (BMPR1B) gene. Journal of Endocrinology,2001,169, R1-R6.
6、姜运良,吴常信Booroola Merino绵羊多胎基因FecB的研究.中国畜牧杂志,1999,35(4)：51-53.
7、 Diatchenko L, Lau YF, Campbell AP, et al. Suppression subtractive hybrization:a method for generating differentiallv regulated or tissue specific cDAN probes and libraries. Proc Natl Acad USA, 1996,93(12):6025-6030.
8、Fayad T, Levesque V, Sirois J, et al. Gene expression profiling of differentially expressed genes in granulose cells if bovine dominant follicles using suppression subtractive hybridization. Biol Reprod, 2004,70:523-533.
9. Leon J Spicer, Pauline Y Aad, Dustin T Allen, et al. Growth Differentiation Factor 9 (GDF9) Stimulates Proliferation and Inhibits Steroidogenesis by Bovine Theca Cells:Influence of Follicle Size on Responses to GDF9. BIOLOGY OF REPRODUCTION,2008,78,243-253.
10. Diatchenko L, Lau Y FC, Campbell AP, et al. Suppression subtractive hybridization:A method for generating differentially regulated or tissue-specific cDNA probes and libraries. Proc. Natl. Acad. Sci,USA.,1996,93:6025-6030.
11. Gurskaya NG, Diatchenko L, Chenchik A, et al. Equalizing cDNA subtraction based on selective suppression of polymerase chain reaction: Cloning of Jurkat cell transcripts induced by phytohemaglutinin and phorbol 12-myristate 13-acetate. Amal. Bilchem,1996,240:90-97.
12. Manabe N, Goto Y, Matsuda-Minehata F, et al.2004. Regualtion mechanism of selective atresia in porcine follicles:regulation of granulose cell apoptosis during atresia. J Reprod Deve.50:493-514.
13. Espey LL, Richards JS. Temporal and spatial patterns of ovarian gene transcription following an ovulatory dose of gonadotropin in the rat. Biol Reprod,2002,67:1662-1670.
14. Qin Chen, A., Zheng GuangWang, Zi Rong Xu, et al. Analysis of gene expression in granulosa cells of ovine antral growing follicles using suppressive subtractive hybridization. Animal Reproduction Science,2009,115:39-48.
15. Ndiaye K, Fayad T, Silversides DW, et al. Identification of downregualted messenger RNAs in bovine granulose cells of dominant follicles following stimulation with human chorinoic gonadotropin. Biol Reprod,2005,73:324-333.
16. Diorf MN, Lefebvre R, Silversides DW, et al. Induction of alpha-caveolin-1 (alpha CAV1) expression in bovine granulosa cells in response to an ovulatory dose of human chorionic gonadotrophins. Mol Reprod,2006,73:1353-1360
17. Clontech Laboratories, Inc. PCR-SelectTM cDNA Subtraction Kit User Manual. Published 16 April 2007:4-8
18. Clontech Laboratories, Inc. Super SMARTTM PCR cDAN Synthesis Kit User Manual. Published 16 April 2007:13-30.
19.何鹏.抑制性消减杂交技术及其应用研究进展.国外医学临床生物化学与检验学分册.2004第25卷第4期：356-358.
20. Willnow TE, Nykjaer A, Herz J. Lipoprotein receptors:new roles for ancient proteins. Nat Cell Biol, 1999,1.-E157-162.
21. Strickland DK, Gonias SL, Argraves WS. Diverse roles for the LDL receptor family. Trends Endocrin Met,2002,13:66-74.
22. Tania Fayad, Rejean Lefebvre, Johannes Nimpf, et al. Low-Density Lipoprotein Receptor-Related Protein 8 (LRP8) Is Upregulated in Granulosa Cells of Bovine Dominant Follicle:Molecular Characterization and Spatio-Temporal Expression Studies. BIOLOGY OF REPRODUCTION,2007, 76,466-475.
23. FLORA RABAHI, SOPHIE BRuLE, JEAN SIROIS, et al. High Expression of Bovine a Glutathione S-Transferase (GSTA1, GSTA2) Subunits Is Mainly Associated with Steroidogenically Active Cells and Regulated by Gonadotropins in Bovine Ovarian Follicles. Endocrinology,1999:140: 3507-3517.
24. Rapoport R, Sklan D, Hanukoglu I. Electron leakage from the adrenal cortex mitochondrial P450scc and P450c11 systems:NADPH and steroid dependence. Arch Biochem Biophys,1995, 317:412-416.
25. Poli G, Dianzani MU, Cheeseman KH, et al. Separation and characterization of the aldehydic products of lipid peroxidation stimulated by carbon tetrachloride or ADP-iron isolated hepatocytes and rat liver microsomal suspensions. Biochem J,1985,227:629-638.
26. Kaur K, Salomon RG, O'Neil J, et al. (Carboxyalkyl) pyrroles in human plasma and oxidized low-density lipoproteins. Chem Res Toxicol,1997,10:1387-1396.
27. Julie Bedard, Sophie Brle, Christopher A. Price, et al. Serine protease inhibitor-E2 (SERPINE2) is differentially expressed in granulosa cells of dominant follicle in cattle. Mol. Reprod. Dev,2003,64: 152-165.
28. Hasan S, Hosseini G, Pricivalle M, et al. Coordinate expression of antivoagulant hepatin sulfate proteoglycans and serine protease inhibitors in the rat ovary:Apotent system of proteolysis control. Biol Peprod,2002,66:144-158.
29. Cao M, Nicola E, Portela VM, et al. Regulation of serine protease inhibitor-E2 and plasminogen activator expression and secretion by follicle stimulating hormone and growth factors in non-luteinizing bovine granulose cells in cells in vitro. Matrix Biol,2006,25:342-356.
30. Harris A L. Emerging issues of connexin channels:biophysics fills the gap. Quart. Rev. Biophys. 2001,34,325-472.
31. Joanne E I, Gittens, Kevin J Barr, et al. Interplay between paracrine signaling and gap junctional communication in ovarian follicles. Journal of Cell Science,2005,118,113-122.
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |