妊娠家兔生殖轴系<i>KISS1</i>和<i>TrkA</i>的分布特征和表达规律

doi:10.7668/hbnxb.20193038

摘要/Abstract

摘要：

为明确吻素(KISS1)和酪氨酸蛋白激酶A(TrkA)在诱导排卵动物生殖轴组织中的表达模式和生理功能,解析KISS1和TrkA对诱导排卵动物生殖的调控机制,以家兔为诱导排卵动物模型,采集妊娠中期(14~18 d)家兔生殖轴系组织(下丘脑、垂体、卵巢和子宫),通过苏木精-伊红染色(H&E)、免疫组织化学染色(IHC)、半定量PCR(RT-PCR)、实时荧光定量PCR(qRT-PCR)和免疫印迹(WB)分析妊娠家兔生殖轴组织中KISS1和TrkA的表达规律和分布特征。IHC结果表明,家兔垂体嗜碱性细胞和卵子透明带中KISS1蛋白的阳性反应较强;TrkA蛋白在下丘脑神经内分泌小细胞、垂体嗜碱性细胞、卵巢颗粒黄体细胞、子宫固有层呈阳性表达。RT-PCR结果显示,妊娠家兔生殖轴组织中均有KISS1表达,且两两比较组间差异显著;下丘脑、垂体、卵巢各组织中TrkA mRNA表达丰度组间差异极显著。KISS1和TrkA 基因qRT-PCR结果与RT-PCR结果表达趋势一致。WB结果发现,KISS1蛋白仅在垂体组织表达;TrkA蛋白在生殖轴组织中均有表达,且表达量在子宫、卵巢、垂体、下丘脑各组织依次呈下降趋势。结果说明,KISS1和TrkA参与调控家兔妊娠过程,且TrkA的作用范围更为广泛。

关键词: 家兔, 生殖轴, 生殖激素, 酪氨酸蛋白激酶A(TrkA), 吻素(KISS1)

Abstract:

In order to clarify the expression pattern and physiological function of Kiss element (KISS1)and Tyrosine protein kinase A(TrkA) in the reproductive axis of ovulation inducing animals,and analyze the regulatory mechanism of KISS1 and TrkA on the reproduction of ovulation inducing animals.Domesticated rabbits were taken as an ovulation-inducing model.The reproductive axis tissues including hypothalamus,pituitary,ovary and uterus were collected from the pregnant Oryctolagus cuniculus f. domestica(Gestation lasts about 14—18 days).The expression patterns and distribution of KISS1 and TrkA mRNA and proteins was detected using hematoxylin-eosin(H&E)and immunohistochemistry(IHC)staining,semi-quantitative interpretation(RT-PCR),Real-time fluorescent quantitative PCR (qRT-PCR)and Western Blot(WB)in the HPG tissues.The results of IHC showed strongly positive reaction of KISS1 protein was presented in the pituitary basophils and zona pellucida of ovary while the positive expression of TrkA protein presented in the neuroendocrine cells of hypothalamus,basophils of pituitary,granular lutein cell of ovary and lamina propria of uterus.The results of RT-PCR KISS1 was expressed in the reproductive axis of pregnant rabbits,and there was significant difference between the two groups.The expression abundance of TrkA mRNA in hypothalamus,pituitary and ovary was significantly different between the two groups.The qRT-PCR results of KISS1 and TrkA genes were consistent with the RT-PCR results.The results of WB suggested that the relative expression levels of KISS1 protein was only observed in pituitary tissue while the relative expression levels of TrkA protein was presented in the HPG tissues and with a down-regulated trend.The results indicating that KISS1 and TrkA are participated in the regulation of pregnancy in Oryctolagus cuniculus f. domestica,and TrkA has a wider scope of action.

Keywords: Pregnant Oryctolagus cuniculus f. domestica;Reproductive axis;Reproductive hormone;Tyrosine protein kinase A(TrkA);Kiss element(KISS1)

安笑笑, 张博皓, 戴丽君, 白旭, 石军, 张霞, 张勇, 张全伟, 赵兴绪. 妊娠家兔生殖轴系KISS1和TrkA的分布特征和表达规律[J]. 华北农学报, 2022, 37(5): 213-221. doi: 10.7668/hbnxb.20193038.

AN Xiaoxiao, ZHANG Bohao, DAI Lijun, BAI Xu, SHI Jun, ZHANG Xia, ZHANG Yong, ZHANG Quanwei, ZHAO Xingxu. The Distribution and Expression Patterns of KISS1 and TrkA in Reproductive Axis of Pregnant Oryctolagus cuniculus f. domestica[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(5): 213-221. doi: 10.7668/hbnxb.20193038.

http://www.hbnxb.net/CN/Y2022/V37/I5/213

图/表 7

表1 半定量PCR引物序列及参数

Tab.1 The primer sequences and parameter of RT-PCR

基因 Gene	引物序列(5'—3') Primer sequence(5'—3')	产物长度/bp Fragment lengths	退火温度/℃ Tm	基因登录号 Gene entry number
KISS1	F: TGCCCGAAGGGTCAGCC	154	58.5	XM_008268539.2
	R: GCTGCTGTTCCTCTGTGCCA
TrkA	F: AATGTGGACGGAGGAACAAG	461	58.5	XM_008264290.2
	R: GTGCCGCATGTACTCAAAGA
GAPDH	F: GGTCGGAGTGAACGGATTTG	150	58.5	NM_001082253.1
	R: TTGCCGTGGGTGGAATCATA

表2 实时荧光定量PCR引物序列及参数

Tab.2 The primer sequences and parameter of qRT-PCR

基因 Gene	引物序列(5'—3') Primer sequence(5'—3')	产物长度/bp Fragment lengths	退火温度/℃ Tm	基因登录号 Gene entry number
KISS1	F: CCGGACCCACAGATCAGC	200	60	XM_008268539.2
	R: GTTGTAGGCGGACGAATCC
TrkA	F: GGACAACCCTTTCGAGTTCA	157	60	XM_008264290.2
	R: GAAAAGGCAGGCAAAGACAG
GAPDH	F: GGTCGGAGTGAACGGATTTG	150	60	NM_001082253.1
	R: TTGCCGTGGGTGGAATCATA

图1 妊娠家兔生殖轴组织形态学观察 A.下丘脑;B.垂体;C和D.卵巢;E.黄体;F.子宫;A~F放大倍数均为400× 。 MNEC.神经内分泌大细胞;PNEC.神经内分泌小细胞;CC.嫌色细胞;BC.嗜碱性细胞;EC.嗜酸性细胞;PO.初级卵母细胞;ZP.透明带;FC.卵泡细胞;PF.初级卵泡;IC.间质细胞;FA.卵泡腔;SG.颗粒层;TFI.卵泡膜内层;TFE.卵泡膜外层;SO.次级卵母细胞;GLC.颗粒黄体细胞;TLC.膜黄体细胞;UG.子宫腺;LP.固有层。图2同。

Fig.1 Morphological observation of HPG axis tissues in pregnant Oryctolagus cuniculus f. domestica using H&E staining A.Hypothalamus;B.Pituitary;C and D.Ovary;E.Corpus luteum;F.Uterus;The magnification of A—F was 400×.MNEC.Magnocellular neuroendocrine cell;PNEC.Parvocellular neuroendocrine cell;CC.Chromophobe;BC.Basophilic;EC.Eosinophil;PO.Primary oocyte;ZP.Zona pellucida;FC.Follicular cell;PF.Primary follicle;IC.Interstitial cells;FA.Follicular cavity;SG.Stratum granulosum;TFI.Inner layer of follicular membrane;TFE.Outer layer of follicular membrane;SO.Secondary oocyte;GLC.Granular lutein cells;TLC.Theca lutein cells;UG.Uterine glands;LP.Proper layer.The same as Fig.2.

图2 妊娠家兔生殖轴组织KISS1和TrkA蛋白的表达分布 A.KISS1蛋白IHC染色;B.TrkA蛋白IHC染色;C.阴性对照(A~C放大倍数均为400×)。SCE.单层柱状上皮。

Fig.2 Expression distribution of KISS1 and TrkA proteins in HPG axis tissues of pregnant Oryctolagus cuniculus f. domestica A.KISS1 protein was stained by immunohistochemically;B.TrkA protein was stained by immunohistochemically;C.Negative control (The magnification of A—C was 400×).SCE.Simple columnar epithelium.

图3 半定量PCR分析妊娠家兔生殖轴组织中KISS1和TrkA mRNA表达规律 A'.琼脂糖凝胶电泳检测妊娠家兔生殖轴组织中KISS1和TrkA mRNA表达丰度。 B'. KISS1 mRNA表达水平。C'.TrkA mRNA表达水平。不同小写字母代表组间差异显著(P<0.05);不同大写字母代表组间差异极显著(P<0.01)。图4—5同。

Fig.3 Relative expression levels of KISS1 and TrkA mRNA in the HPG axis of pregnant Oryctolagus cuniculus f. domestica with RT-PCR A'.The mRNA expression levels of KISS1 and TrkA in HPG axis tissues of pregnant rabbits were detected by agarose gel electrophoresis.B'.KISS1 mRNA expression level.C'.TrkA mRNA expression level.Different lowercase letters indicated significant difference between groups(P<0.05);different capital letters indicated significant difference between groups (P<0.01).The same as Fig.4—5.

图4 qRT-PCR分析妊娠家兔生殖轴组织中KISS1和TrkA mRNA表达规律

Fig.4 Relative expression levels of KISS1 and TrkA mRNA in the HPG axis of pregnant Oryctolagus cuniculus f. domestica with qRT-PCR

图5 妊娠家兔生殖轴组织中KISS1和TrkA蛋白表达规律分析 A.聚丙酰胺凝胶电泳检测妊娠家兔生殖轴组织中KISS1、TrkA和β-actin蛋白表达丰度。

Fig.5 Relative expression levels of KISS1 and TrkA protein in the HPG axis of pregnant Oryctolagus cuniculus f. domestica A.The protein expression levels of KISS1,TrkA and β-actin in HPG axis tissues of pregnant rabbits were detected by polyproacrylamide gel electrophoresis.

参考文献 35

[1]	Umaru M A, Bello A. Reproduction in the one humped camel(Camelus dromedarius)in semi arid Nigeria[J]. Scientific Journal of Biological Sciences, 2013, 2(1):1-7.
[2]	王琪. 双峰驼精清诱导排卵分子调控机制的研究[D]. 兰州: 甘肃农业大学, 2020.doi: 10.27025/d.cnki.ggsnu.2020.000050. doi: 10.27025/d.cnki.ggsnu.2020.000050
	Wang Q. Mechanism of molecular regulation was studied of seminal plasma induced-ovulation in the Bactrian camel (Camelus bactrianus)[D]. Lanzhou: Gansu Agricultural University, 2020.
[3]	王志强, 张君, 徐尚荣, 彭巍, 张琳钦. 生殖激素对初情期牦牛子宫生长发育的影响[J]. 青海畜牧兽医杂志, 2018, 48(2):14-18.doi: 10.3969/j.issn.1003-7950.2018.02.006. doi: 10.3969/j.issn.1003-7950.2018.02.006
	Wang Z Q, Zhang J, Xu S R, Peng W, Zhang L Q. Influence of reproductive hormones secretion pattern on uterus development in puberal female yaks[J]. Chinese Qinghai Journal of Animal and Veterinary Sciences, 2018, 48(2):14-18.
[4]	杨淑华, 徐世文, 李金龙, 王金涛. 生殖激素对禽类卵泡细胞凋亡的影响[J]. 吉林畜牧兽医, 2005, 26(10):12-14,17.doi: 10.3969/j.issn.1672-2078.2005.10.006. doi: 10.3969/j.issn.1672-2078.2005.10.006
	Yang S H, Xu S W, Li J L, Wang J T. The effect of reproductive hormone on avian follicular cells apoptosis[J]. Jilin Animal Husbandry and Veterinary Medicine, 2005, 26(10):12-14,17.
[5]	程元, 赵晓钰, 李园园, 赵全庄, 朱建平, 艾山江, 李海英. 新疆伊犁鹅繁殖周期生殖激素变化及卵巢卵泡发育规律的研究[J]. 中国畜牧兽医, 2017, 44(11):3264-3269.doi: 10.16431/j.cnki.1671-7236.2017.11.022. doi: 10.16431/j.cnki.1671-7236.2017.11.022
	Cheng Y, Zhao X Y, Li Y Y, Zhao Q Z, Zhu J P, Ai S J, Li H Y. Study on the changes of reproductive hormone and regulation of follicle development during the reproductive cycle of Yili goose in Xinjiang[J]. China Animal Husbandry & Veterinary Medicine, 2017, 44(11):3264-3269.
[6]	张宏馨. 维生素E对种母鸡繁殖性能的影响及其机理研究[D]. 保定: 河北农业大学, 2013.
	Zhang H X. Study on the influence of vitamin E on the reproductive performance and it's mechanism in breeding hens[D]. Baoding: Hebei Agricultural University, 2013.
[7]	余哲琪, 田佳迎, 李启黉, 陈忠. p38 MAPK抑制剂对热应激雌性雏鸡主要生殖激素分泌的影响[J]. 中国家禽, 2021, 43(1):76-80.doi: 10.16372/j.issn.1004-6364.2021.01.013. doi: 10.16372/j.issn.1004-6364.2021.01.013
	Yu Z Q, Tian J Y, Li Q H, Chen Z. Effects of p38 MAPK inhibitor on reproductive hormones content in heat-stressed female chicks[J]. China Poultry, 2021, 43(1):76-80.
[8]	狄冉, 郭晓飞, 刘秋月, 胡文萍, 王翔宇, 潘章源, 储明星. 生殖激素对绵羊繁殖性能影响的研究进展[J]. 家畜生态学报, 2015, 36(9):1-6.doi: 10.3969/j.issn.1673-1182.2015.09.001. doi: 10.3969/j.issn.1673-1182.2015.09.001
	Di R, Guo X F, Liu Q Y, Hu W P, Wang X Y, Pan Z Y, Chu M X. Research progress in effect of reproductive hormones on ovine reproductive performance[J]. Acta Ecologae Animalis Domastici, 2015, 36(9):1-6.
[9]	田晓, 孙振高. 早孕期雌激素水平预测妊娠结局的研究进展[J]. 医学综述, 2020, 26(5):872-877.doi: 10.3969/j.issn.1006-2084.2020.05.008. doi: 10.3969/j.issn.1006-2084.2020.05.008
	Tian X, Sun Z G. Research progress in prediction of pregnancy outcome by estrogen level in early pregnancy[J]. Medical Recapitulate, 2020, 26(5):872-877.
[10]	聂莉, 赵友波, 赵丹, 龙云, 刘敏, 雷毅, 王翊丞, 张雪芹, 张金虎, 岳利民. 孕激素诱导microRNA调控子宫内膜上皮功能活动的作用研究[C]// 2019中国生理学会学术年会暨张锡钧基金第十五届全国青年优秀生理学学术论文交流会及第十三届全国青年生理学工作者学术会议论文摘要.中国生理学会, 2019, 1:164-165.
	Nie L, Zhao Y B, Zhao D, Long Y, Liu M, Lei Y, Wang Y C, Zhang X Q, Zhang J H, Yue L M. Study on the role of progesterone induced microRNA in regulating endometrial epithelial function[C]// 2019 academic annual meeting of Chinese Physiological Society and the 15th National Youth excellent physiological academic paper exchange meeting of Zhang Xijun foundation and the 13th National Youth physiological workers academic conference.Chinese Association for Physiological Sciences, 2019, 1:164-165.
[11]	Melnick A P, Pereira N, Murphy E M, Rosenwaks Z, Spandorfer S D. How low is too low? Cycle day 28 estradiol levels and pregnancy outcomes[J]. Fertility and Sterility, 2016, 105(4):905-909.doi: 10.1016/j.fertnstert.2015.11.046. doi: 10.1016/j.fertnstert.2015.11.046 pmid: 26707516
[12]	Pinilla L, Aguilar E, Dieguez C, Millar R P, Tena-Sempere M. Kisspeptins and reproduction:Physiological roles and regulatory mechanisms[J]. Physiological Reviews, 2012, 92(3):1235-1316.doi: 10.1152/physrev.00037.2010. doi: 10.1152/physrev.00037.2010 pmid: 22811428
[13]	Han S K, Gottsch M L, Lee K J, Popa S M, Smith J T, Jakawich S K, Clifton D K, Steiner R A, Herbison A E. Activation of gonadotropin-releasing hormone neurons by kisspeptin as a neuroendocrine switch for the onset of puberty[J]. The Journal of Neuroscience, 2005, 25(49):11349-11356.doi: 10.1523/JNEUROSCI.3328-05.2005. doi: 10.1523/JNEUROSCI.3328-05.2005 URL
[14]	Silveira L G, Latronico A C, Seminara S B. Kisspeptin and clinical disorders[J]. Kisspeptin Signaling in Reproductive Biology, 2013:7 84:187-199.doi: 10.1007/978-1-4614-6199-9_9. doi: 10.1007/978-1-4614-6199-9_9
[15]	彭乔, 桂文武. Kisspeptin/KISS1R系统在妊娠和辅助生殖领域中的研究进展和应用前景[J]. 中国计划生育和妇产科, 2021, 13(4):24-28.doi: 10.3969/j.issn.1674-4020.2021.04.07. doi: 10.3969/j.issn.1674-4020.2021.04.07
	Peng Q, Gui W W. Research progress and application prospects of Kisspeptin/KISS1R system in the field of pregnancy and assisted reproduction[J]. Chinese Journal of Family Planning & Gynecotokology, 2021, 13(4):24-28.
[16]	侯晓峰. 神经生长因子对小鼠卵巢颗粒细胞增殖及孕酮分泌的影响[D]. 长春: 吉林大学, 2014.
	Hou X F. Effect of nerve growth factor on proliferation of granulosa cells and secretion of progesterone in mouse ovary[D]. Changchun: Jilin University, 2014.
[17]	Zhao X X, Li X L, Chen B X. Isolation of ovulation-inducing factors in the seminal plasma of bactrian camel(Camelus bactrianus)by DEAE-cellulose chromatography[J]. Reproduction in Domestic Animals, 2001, 36(3/4):177-181. doi: 10.1046/j.1439-0531.2001.d01-33.x URL
[18]	Ratto M H, Leduc Y A, Valderrama X P, van Straaten K E, Delbaere L T J, Pierson R A, Adams G P. The nerve of ovulation-inducing factor in semen[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(37):15042-15047.doi: 10.1073/pnas.1206273109. doi: 10.1073/pnas.1206273109 pmid: 22908303
[19]	陈健楠, 张全伟, 王琪, 王媛媛, 马友记, 张勇, 赵兴绪. MTNR1a和MTNR1b在牦牛不同组织中的表达及其在睾丸中的定位[J]. 兽类学报, 2020, 40(1):64-70.doi: 10.16829/j.slxb.150313. doi: 10.16829/j.slxb.150313
	Chen J N, Zhang Q W, Wang Q, Wang Y Y, Ma Y J, Zhang Y, Zhao X X. Expression of MTNR1a and MTNR1b in different tissues and localization in testes of yak(Bos grunniens)[J]. Acta Theriologica Sinica, 2020, 40(1):64-70.
[20]	赵彩英, 李怡娜, 王琪, 甘泽, 张勇, 马友记, 张全伟, 赵兴绪. 雄性小鼠生殖轴组织中AURKA表达定位及敲低后对双氢睾酮合成代谢的影响[J]. 农业生物技术学报, 2020, 28(9):1614-1622.doi: 10.3969/j.issn.1674-7968.2020.09.010. doi: 10.3969/j.issn.1674-7968.2020.09.010
	Zhao C Y, Li Y N, Wang Q, Gan Z, Zhang Y, Ma Y J, Zhang Q W, Zhao X X. Expression and localization analysis of AURKA in HPGA of male mouse(Mus musculus)and effect of its knockdown on DHT anabolism[J]. Journal of Agricultural Biotechnology, 2020, 28(9):1614-1622.
[21]	钱晨曦, 董杰, 王晓红. Kisspeptin及其受体KISS1R在女性生殖系统中的分布及局部调节作用研究进展[J]. 解放军医学院学报, 2021, 42(10):1114-1117,1129.doi: 10.3969/j.issn.2095-5227.2021.10.021. doi: 10.3969/j.issn.2095-5227.2021.10.021
	Qian C X, Dong J, Wang X H. Distribution and local regulatory effect of Kisspeptin/KISS1R in female reproductive system[J]. Academic Journal of Chinese PLA Medical School, 2021, 42(10):1114-1117,1129.
[22]	Levanti M B, Germanà A, Abbate F, Montalbano G, Vega J A, Germanà G. TrkA and p75NTR in the ovary of adult cow and pig[J]. Journal of Anatomy, 2005, 207(1):93-96.doi: 10.1111/j.1469-7580.2005.00423.x. doi: 10.1111/j.1469-7580.2005.00423.x pmid: 16011548
[23]	黄厚才. 家兔交配前后及妊娠期生殖内分泌变化的研究[J]. 畜牧与兽医, 1996, 28(6):245-246.
	Huang H C. Preliminary analysis on changes of plasma camp and cgmp levels in neonatal ruminants[J]. Animal Husbandry & Veterinary Medicine, 1996, 28(6):245-246.
[24]	Smith C A, Shoemaker C M, Roeszler K N, Queen J, Crews D, Sinclair A H. Cloning and expression of R-Spondin1 in different vertebrates suggests a conserved role in ovarian development[J]. BMC Developmental Biology, 2008, 8:72-78.doi: 10.1186/1471-213X-8-72. doi: 10.1186/1471-213X-8-72 pmid: 18651984
[25]	de Roux N, Genin E, Carel J C, Matsuda F, Chaussain J L, Milgrom E. Hypogonadotropic hypogonadism due to loss of function of the KiSS1-derived peptide receptor GPR54[J]. Proceedings of the National Academy of Sciences of the United States of America, 2003, 100(19):10972-10976.doi: 10.1073/pnas.1834399100. doi: 10.1073/pnas.1834399100 pmid: 12944565
[26]	Harms J F, Welch D R, Miele M E. KISS1 metastasis suppression and emergent pathways[J]. Clinical & Experimental Metastasis, 2003, 20(1):11-18.doi: 10.1023/a:1022530100931. doi: 10.1023/a:1022530100931
[27]	赵丹丹, 郑连文, 楚琪, 徐影. Kisspeptin在维持妊娠及预测妊娠结局方面的研究进展[J]. 中国计划生育和妇产科, 2021, 13(10):17-21.doi: 10.3969/j.issn.1674-4020.2021.10.05. doi: 10.3969/j.issn.1674-4020.2021.10.05
	Zhao D D, Zheng L W, Chu Q, Xu Y. Research progress of Kisspeptin in maintaining pregnancy and predicting pregnancy outcome[J]. Chinese Journal of Family Planning & Gynecotokology, 2021, 13(10):17-21.
[28]	朱晓玲, 李成娇, 侯晓峰, 石德顺, 刘庆友, 周虚. 神经生长因子NGF及其受体TrkA在雌性水牛生殖器官的表达定位[J]. 吉林农业大学学报, 2011, 33(3):315-318.doi: 10.13327/j.jjlau.2011.03.018. doi: 10.13327/j.jjlau.2011.03.018
	Zhu X L, Li C J, Hou X F, Shi D S, Liu Q Y, Zhou X. Immunohistochemical localization of nerve growth factor and its receptors tyrosine kinase A on the genital in female buffalo[J]. Journal of Jilin Agricultural University, 2011, 33(3):315-318.
[29]	马永和. 神经生长因子及其受体在母牛生殖系统中的表达与功能研究[D]. 长春: 吉林大学, 2010.
	Ma Y H. Expression and function of nerve growth factor and its receptor in reproductive system in cow[D]. Changchun: Jilin University, 2010.
[30]	王勇. NGF及其受体对卵泡发育调控的分子机制研究[D]. 合肥: 中国科学技术大学, 2013.
	Wang Y. The role of NGF and its receptor in regulation of mouse follicular development[D]. Hefei: University of Science and Technology of China, 2013.
[31]	章苓, 王林芳, 丁斐. 小鼠生后发育过程中子宫 NGF基因表达的变化[J]. 南通医学院学报, 2001, 21(4):355-356.doi: 10.3969/j.issn.1674-7887.2001.04.012. doi: 10.3969/j.issn.1674-7887.2001.04.012
	Zhang L, Wang L F, Ding F. Gene expression of nerve growth factor in mouse uterus during postnatal development[J]. Journal of Nantong University (Medical Sciences), 2001, 21(4):355-356.
[32]	Ren L Q, Medan M S, Weng Q, Jin W Z, Li C M, Watanabe G, Taya K. Immunolocalization of nerve growth factor(NGF)and its receptors(TrkA and p75LNGFR)in the reproductive organs of Shiba goats[J]. The Journal of Reproduction and Development, 2005, 51(3):399-404.doi: 10.1262/jrd.16082. doi: 10.1262/jrd.16082 URL
[33]	Dissen G A, Hill D F, Costa M E, Les Dees C W, Lara H E, Ojeda S R. A role for TrkA nerve growth factor receptors in mammalian ovulation[J]. Endocrinology, 1996, 137(1):198-209.doi: 10.1210/en.137.1.198. doi: 10.1210/en.137.1.198 pmid: 8536613
[34]	安世钰, 张国敏, 尤佩华, 孟繁星, 刘建玲, 张婷婷, 杨花, 樊懿萱, 王锋. PPARGC1A/NRF1/TFAM通路核心基因在湖羊睾丸发育过程中的表达变化[J]. 南京农业大学学报, 2019, 42(2):345-351.doi: 10.7685/jnau.201806041. doi: 10.7685/jnau.201806041
	An S Y, Zhang G M, You P H, Meng F X, Liu J L, Zhang T T, Yang H, Fan Y X, Wang F. Research on the change of PPARGC1A/NRF1/TFAM pathway genes expression in the development of Hu sheep testis[J]. Journal of Nanjing Agricultural University, 2019, 42(2):345-351.
[35]	刘海龙, 李娟, 蒲雪峰, 龙树林, 滕芸. 神经生长因子、酪氨酸激酶及B7-H4与宫颈癌病情进展、预后的关系研究[J]. 实用医院临床杂志, 2021, 18(5):51-56.doi: 10.3969/j.issn.1672-6170.2021.05.014. doi: 10.3969/j.issn.1672-6170.2021.05.014
	Liu H L, Li J, Pu X F, Long S L, Teng Y. The relationship between the NGF,TrkA and B7-H4 and the progression and prognosis of cervical cancer[J]. Practical Journal of Clinical Medicine, 2021, 18(5):51-56.

选择文件类型/文献管理软件名称

选择包含的内容

妊娠家兔生殖轴系KISS1和TrkA的分布特征和表达规律

The Distribution and Expression Patterns of KISS1 and TrkA in Reproductive Axis of Pregnant Oryctolagus cuniculus f. domestica

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 7

参考文献 35

相关文章 5

编辑推荐

Metrics

本文评价

[1]	肖正龙, 张龚炜, 杨宇, 王杰, 赖松家. 家兔ATG16L1基因多态性与非特异性消化道紊乱易感性的研究[J]. 华北农学报, 2013, 28(4): 234-238.
[2]	王芳, 胡波, 任雪枫, 范志宇, 张则斌, 徐为中, 何孔旺. 新西兰白兔γ-干扰素在昆虫细胞中的表达及其活性测定[J]. 华北农学报, 2010, 25(3): 9-13.
[3]	徐直, 蒋振国, 徐一树, 张冬梅. 黑白花奶牛发情周期血液中促卵泡素、促黄体素、促乳素、孕酮、雌二醇的含量及变化规律[J]. 华北农学报, 1996, 11(3): 121-127.
[4]	宁长申, 刘维忠. 家兔13种球虫卵囊形态观察[J]. 华北农学报, 1988, 3(2): 81-85.
[5]	房海, 陈翠珍. 家兔致病性大肠艾希氏杆菌的血清型鉴定[J]. 华北农学报, 1987, 2(4): 144-144.

模态框（Modal）标题

选择文件类型/文献管理软件名称

选择包含的内容

妊娠家兔生殖轴系KISS1和TrkA的分布特征和表达规律

The Distribution and Expression Patterns of KISS1 and TrkA in Reproductive Axis of Pregnant Oryctolagus cuniculus f. domestica

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 7

参考文献 35

相关文章 5

编辑推荐

Metrics

本文评价