龍?jiān)婍?，姜冬梅，陳咨余，管成，易治鑫，康?/p>

（四川農(nóng)業(yè)大學(xué)動(dòng)物科技學(xué)院/畜禽遺傳資源發(fā)掘與創(chuàng)新利用四川省重點(diǎn)實(shí)驗(yàn)室，成都611130）

外源性亞精胺對(duì)鼠卵巢生殖激素受體基因表達(dá)的影響

龍?jiān)婍?，姜冬梅，陳咨余，管成，易治鑫，康?

（四川農(nóng)業(yè)大學(xué)動(dòng)物科技學(xué)院/畜禽遺傳資源發(fā)掘與創(chuàng)新利用四川省重點(diǎn)實(shí)驗(yàn)室，成都611130）

為探討外源性亞精胺對(duì)鼠卵巢生殖激素受體基因表達(dá)的影響，采用不同質(zhì)量分?jǐn)?shù)（0.05、0.10和0.15 mg/g）亞精胺腹腔注射6周齡昆明鼠，24 h后采集卵巢組織樣品，應(yīng)用實(shí)時(shí)熒光定量聚合酶鏈?zhǔn)椒磻?yīng)檢測(cè)卵巢組織中促卵泡激素受體、促黃體生成素受體、雌激素受體1、雌激素受體2和雄激素受體基因表達(dá)量。結(jié)果顯示：與對(duì)照組相比，0.05 mg/g亞精胺處理組卵巢促黃體生成素受體表達(dá)量顯著高于對(duì)照組（P＜0.05），0.15 mg/g組促黃體生成素受體表達(dá)量顯著低于對(duì)照組（P＜0.05）；隨亞精胺處理量的增加，卵巢組織中促卵泡激素受體、雌激素受體1、雌激素受體2和雄激素受體表達(dá)量均呈逐漸增加的趨勢(shì)，且均在0.15 mg/g處理組中的表達(dá)量顯著高于對(duì)照組（P＜0.05）。綜上表明，亞精胺可通過(guò)介導(dǎo)鼠卵巢組織生殖激素受體基因表達(dá)來(lái)參與調(diào)控鼠卵巢功能。

亞精胺；卵巢；激素受體；基因表達(dá)

SummaryThe polyamines spermidine and spermine and their precursor putrescine are a class of compounds containing two or more amino groups and participate in the regulation of various physiological and pathological processes by interacting with negatively-charged molecules,such as DNA,RNA and proteins.Recently,increasing studies suggest that polyamines also play important roles in follicular development,ovulation and steroidogenesis in female animals.The spermidine exists ubiquitously in species ranging from yeast to mammals.Recently,studies from our and other laboratories indicated that spermidine is involved in regulating animal reproduction through mediating spermatogenesis,oogenesis,follicular development and ovulation.However, to the best of our knowledge,the effect of the spermidine on the expression levels of hormone receptor genes associated with reproduction is unknown.

To demonstrate the effect of the spermidine on the transcription of reproductive hormone receptor genes in mouse ovaries, Kunming mice were administrated with 0.05,0.10 and 0.15 mg/g(body mass)spermidine by intraperitoneal injection.Ovarian tissue samples were collected after 24 h of spermidine administration.The levels of follicle-stimulating hormone receptor (FSHR),luteinizing hormone receptor(LHR),estrogen receptor 1(ER1),ER2and androgen receptor(AR)mRNA expression in mouse ovaries were measured using quantitative real-time polymerase chain reaction.

The results showed that the level ofLHRmRNA expression in ovaries of mice administrated with 0.05 mg/g spermidine was significantly higher than the untreated control group(P＜0.05).The amount ofLHRmRNA expression in ovaries of mice administrated with 0.15 mg/g spermidine was significantly lower than the untreated control group(P＜0.05).With increasing spermidine administration,a gradually increasing trend was observed in the change ofFSHR,ER1,ER2andARexpression levels.The amount ofFSHR,ER1,ER2andARgenes in ovaries of mice administrated with 0.15 mg/g spermidine was significantly high,compared with the untreated control group,respectively(P＜0.05).

In conclusion,our present results demonstrate that exogenous spermidine mediates the transcription of genes associated with reproductive hormone receptors in ovaries of mice.It is indicated that spermidine might play an important role in regulating the responsiveness of the ovary to reproductive hormones through an unknown mechanism.Further research is needed to elucidate the mechanism of action of spermidine mediating ovarian functions in animals.

亞精胺是廣泛存在于動(dòng)物、植物和微生物體內(nèi)的一種多胺，可通過(guò)介導(dǎo)雄性動(dòng)物精子發(fā)生，以及雌性動(dòng)物卵泡發(fā)育、卵子發(fā)生和排卵活動(dòng)，進(jìn)而參與調(diào)控動(dòng)物的繁殖過(guò)程[1-2]。多胺能通過(guò)與多種生殖激素建立復(fù)雜的反饋聯(lián)系，進(jìn)而參與調(diào)控動(dòng)物繁殖[3]。多胺在維持卵巢類(lèi)固醇激素水平和生成過(guò)程中具有重要作用[4]。雄性動(dòng)物體內(nèi)的多胺合成受到雄激素的調(diào)節(jié)[5]。FASHE等[6]研究發(fā)現(xiàn)，多胺合成受阻時(shí)，卵巢性激素含量顯著降低。趙越超[7]研究發(fā)現(xiàn)，性激素可通過(guò)其受體途徑影響鼠子宮內(nèi)多胺代謝。大量研究證實(shí)，雄激素在治療男性不育癥上具有重要作用，亞精胺在不育男性精漿中的含量明顯比正常男性低[8-10]，推測(cè)亞精胺可能通過(guò)作用于雄激素及其受體來(lái)參與調(diào)控雄性動(dòng)物的繁殖功能。而在雌性動(dòng)物中，亞精胺是否與雄激素受體（androgen receptor,AR）存在直接或間接的作用還未有研究報(bào)道。

促卵泡激素（follicle-stimulating hormone,FSH）和促黃體激素（luteinizing hormone,LH）分別與促卵泡激素受體（follicle-stimulating hormone receptor, FSHR）和促黃體生成素受體（luteinizing hormone receptor,LHR）作用，共同參與調(diào)控動(dòng)物卵泡發(fā)育和卵泡顆粒細(xì)胞增殖[11]。LH和FSH還可參與調(diào)控卵巢組織中雌激素的合成，而雌激素通過(guò)負(fù)反饋調(diào)節(jié)機(jī)制影響垂體LH和FSH的分泌[12-13]。雌激素受體（estrogen receptor,ER）包含ER1和ER2這2種亞型，雙敲除ER1和ER2的雌性鼠不能排卵，說(shuō)明ER對(duì)雌性動(dòng)物排卵具有至關(guān)重要的調(diào)控作用[14]。研究表明，多胺可通過(guò)介導(dǎo)哺乳動(dòng)物生殖激素合成與分泌來(lái)參與調(diào)控動(dòng)物繁殖功能[4]。然而，目前有關(guān)亞精胺調(diào)控雌性動(dòng)物卵巢功能的研究相對(duì)較少，而且尚未見(jiàn)有關(guān)亞精胺調(diào)控卵巢生殖激素受體基因表達(dá)影響的報(bào)道。因此，本試驗(yàn)研究了不同質(zhì)量分?jǐn)?shù)亞精胺對(duì)鼠卵巢組織中FSHR、LHR、ER1、ER2和AR基因表達(dá)的影響，以期為闡明多胺調(diào)控哺乳動(dòng)物卵巢功能的作用機(jī)制提供理論依據(jù)。

1 材料與方法

1.1 試驗(yàn)動(dòng)物及樣品采集

選取雌性健康同胞昆明鼠（SPF級(jí)試驗(yàn)用昆明系鼠，購(gòu)于成都達(dá)碩公司），常規(guī)分籠飼養(yǎng)，鼠自由采食；在試驗(yàn)過(guò)程中對(duì)動(dòng)物處置符合動(dòng)物倫理學(xué)標(biāo)準(zhǔn)。6周齡時(shí)，將鼠隨機(jī)分為4組，每組8只，分別腹腔注射生理鹽水和0.05、0.10和0.15 mg/g（體質(zhì)量）亞精胺。注射24 h后頸部脫臼處死，迅速采集鼠卵巢組織，用生理鹽水洗凈，濾紙吸干后，置于－80℃冰箱保存，備用。

1.2 引物設(shè)計(jì)和實(shí)時(shí)熒光定量聚合酶鏈?zhǔn)椒磻?yīng)

按照RNA提取試劑盒（RNAiso Plus kit，TaKaRa公司，大連）說(shuō)明書(shū)提取鼠卵巢組織總RNA，然后參照反轉(zhuǎn)錄試劑盒說(shuō)明書(shū)（TaKaRa公司，大連）將總RNA樣品反轉(zhuǎn)錄成cDNA模板，置于－20℃冰箱中保存，備用?；贕enBank數(shù)據(jù)庫(kù)，采用Primer Premier 5.0和Oligo 6.0軟件設(shè)計(jì)特異性引物，并委托華大基因有限公司合成。引物序列信息見(jiàn)表1。

實(shí)時(shí)熒光定量反應(yīng)體系（10 μL）：實(shí)時(shí)熒光定量聚合酶鏈?zhǔn)椒磻?yīng)試劑盒（iQTMSYBR Green Supermix kit，Bio-Rad公司，北京）5.0 μL，上、下游引物（10 μmol/L）各0.2 μL，cDNA模板0.5 μL，用無(wú)RNA酶水補(bǔ)足至10 μL。反應(yīng)條件：95℃預(yù)變性3 min；95℃變性10 s，57～63℃退火30 s，72℃延伸30 s（采集熒光），40個(gè)循環(huán)；95℃保持10 s，繪制溶解曲線。用3-磷酸甘油醛脫氫酶（glyceraldehyde-3-phosphate dehydrogenase,GAPDH）基因作為內(nèi)參基因。每個(gè)樣品設(shè)3個(gè)重復(fù)。

表1 實(shí)時(shí)熒光定量聚合酶鏈?zhǔn)椒磻?yīng)引物序列Table 1 Primer sequences used in quantitative real-time polymerase chain reaction(PCR)

1.3 數(shù)據(jù)處理及統(tǒng)計(jì)分析

采用2－ΔΔCT法處理數(shù)據(jù)[7]，用GAPDH作為內(nèi)參基因，以空白組基因的表達(dá)量作為對(duì)照組，計(jì)算不同質(zhì)量分?jǐn)?shù)亞精胺處理組卵巢組織FSHR、LHR、ER1、ER2、AR基因的相對(duì)表達(dá)量。應(yīng)用SAS 9.2統(tǒng)計(jì)分析軟件中的MEANS過(guò)程進(jìn)行描述性統(tǒng)計(jì)分析，并進(jìn)行方差分析和鄧肯多重比較，結(jié)果用平均值±標(biāo)準(zhǔn)差表示，P＜0.05表示差異有統(tǒng)計(jì)學(xué)意義。

2 結(jié)果

2.1 亞精胺對(duì)鼠LHR和FSHR基因表達(dá)的影響

由圖1可知：腹腔注射0.05 mg/g亞精胺時(shí)，鼠卵巢組織中LHR相對(duì)表達(dá)量顯著高于其他各組，是對(duì)照組的2.07倍（P＜0.05）；腹腔注射0.15 mg/g亞精胺組LHR表達(dá)量顯著低于對(duì)照組，為對(duì)照組的9%（P＜0.05）。腹腔注射0.15 mg/g亞精胺時(shí)，F(xiàn)SHR表達(dá)量顯著高于對(duì)照組，是對(duì)照組的31.67倍（P＜0.05）；而腹腔注射0.05和0.10 mg/g亞精胺時(shí)，F(xiàn)SHR基因表達(dá)量與對(duì)照組相比差異無(wú)統(tǒng)計(jì)學(xué)意義（P＞0.05）。

圖1 亞精胺對(duì)鼠卵巢LHR和FSHR表達(dá)的影響Fig.1 Effect of spermidine onLHRandFSHRmRNA expression in mouse ovaries

2.2 亞精胺對(duì)ERER1 1和ERER2 2基因表達(dá)的影響

腹腔注射0.15 mg/g亞精胺處理組中，ER1和ER2表達(dá)量均顯著高于對(duì)照組，分別為對(duì)照組的1.95倍和6.43倍（P＜0.05）；而注射0.05和0.10 mg/g組中，ER1和ER2與對(duì)照組間差異均無(wú)統(tǒng)計(jì)學(xué)意義（P＞0.05）（圖2）。

圖2 亞精胺對(duì)鼠卵巢ER ER1 1和ER ER2 2表達(dá)的影響Fig.2 Effect of spermidine onER1andER2mRNA expression in mouse ovaries

2.3 亞精胺對(duì)AR基因表達(dá)的影響

如圖3所示：當(dāng)腹腔注射0.15 mg/g亞精胺時(shí)，卵巢組織中AR相對(duì)表達(dá)量顯著高于對(duì)照組和其他處理組，是對(duì)照組的4.31倍（P＜0.05）；而注射0.05和0.10 mg/g組中，AR表達(dá)量與對(duì)照組間差異無(wú)統(tǒng)計(jì)學(xué)意義（P＞0.05）。

圖3 亞精胺對(duì)鼠卵巢AR表達(dá)的影響Fig.3 Effect of spermidine onARmRNA expression in mouse ovaries

3 討論

生殖激素及其特異性受體在調(diào)控動(dòng)物繁殖過(guò)程中具有關(guān)鍵作用，而FSH和LH在調(diào)控動(dòng)物繁殖過(guò)程中具有核心作用[15-16]。在卵巢顆粒細(xì)胞中LHR表達(dá)量降低會(huì)抑制排卵功能[17]；而FSHR表達(dá)量升高能增強(qiáng)卵巢對(duì)外源性促性腺激素的應(yīng)答能力。THYSSEN等[18]研究發(fā)現(xiàn)，外源性腐胺和亞精胺能抑制下丘腦和垂體組織中LH和FSH的分泌；用二氟甲基鳥(niǎo)氨酸（一種腐胺生物合成的不可逆抑制劑）處理后FSH分泌增加：表明多胺可參與調(diào)控垂體LH和FSH分泌。本研究發(fā)現(xiàn)，腹腔注射亞精胺顯著提高了卵巢組織中FSHR和LHR表達(dá)量：說(shuō)明亞精胺也可通過(guò)介導(dǎo)卵巢FSHR和LHR表達(dá)來(lái)發(fā)揮其對(duì)雌性動(dòng)物繁殖功能的調(diào)控作用。表皮生長(zhǎng)因子（epidermal growth factor,EGF）作為一種卵巢旁分泌因子，通過(guò)介導(dǎo)卵泡發(fā)育和卵母細(xì)胞生成來(lái)調(diào)節(jié)卵巢功能。研究表明，EGF可下調(diào)哺乳動(dòng)物卵巢FSHR表達(dá)[19]。BLACHOWSKI等[20]發(fā)現(xiàn)，多胺能參與EGF信號(hào)轉(zhuǎn)導(dǎo)通路，提示多胺與EGF參與的生殖調(diào)控具有相關(guān)性。本研究發(fā)現(xiàn)，注射0.15 mg/g亞精胺顯著增加了卵巢組織中FSHR表達(dá)量，推測(cè)亞精胺促進(jìn)FSHR表達(dá)的作用可能與EGF有關(guān)，有待進(jìn)一步研究證實(shí)。此外，本研究還發(fā)現(xiàn)，低質(zhì)量分?jǐn)?shù)亞精胺（0.05 mg/g）能促進(jìn)卵巢LHR表達(dá)，但對(duì)FSHR表達(dá)影響不顯著；而高質(zhì)量分?jǐn)?shù)亞精胺（0.15 mg/g）抑制LHR表達(dá)，但促進(jìn)了FSHR表達(dá)：表明亞精胺影響FSHR和LHR表達(dá)的效應(yīng)具有劑量差異。

雌激素受體基因包括ER1和ER2，在鼠卵巢組織中均有表達(dá)，但其表達(dá)量和生理作用各不相同。在未成熟和成熟的嚙齒類(lèi)動(dòng)物和人卵巢中，ER2表達(dá)量遠(yuǎn)高于ER1[21]。研究表明，在卵巢組織中ER2的表達(dá)和分布與鼠的生殖周期存在密切聯(lián)系，尤其是在發(fā)情期的鼠卵巢組織中有高度表達(dá)，其表達(dá)規(guī)律與小鼠體內(nèi)雌激素水平相適應(yīng)[22]。本研究發(fā)現(xiàn)，0.15 mg/g亞精胺處理可顯著增加鼠卵巢組織中ER1和ER2表達(dá)，提示高質(zhì)量分?jǐn)?shù)亞精胺可通過(guò)提高ER表達(dá)進(jìn)而參與調(diào)控鼠卵巢功能。

AR在調(diào)節(jié)卵巢功能和維持女性生育能力方面具有重要作用[23]。KIMURA等[24]發(fā)現(xiàn)，在卵泡發(fā)育早期即有AR的表達(dá)，AR缺陷型鼠易出現(xiàn)卵泡發(fā)育障礙和卵巢早衰。AR和雄激素水平與哺乳動(dòng)物在卵巢發(fā)育過(guò)程中FSHR表達(dá)密切相關(guān)[25]。此外，雄激素還參與調(diào)控卵巢癌的發(fā)生和發(fā)展過(guò)程，AR水平及其活性增強(qiáng)會(huì)使卵巢癌細(xì)胞對(duì)雄激素的敏感性增大，從而促進(jìn)卵巢癌變[26]。本研究結(jié)果表明，0.15 mg/g亞精胺處理組的鼠卵巢組織中AR表達(dá)顯著高于對(duì)照組：表明高質(zhì)量分?jǐn)?shù)亞精胺可通過(guò)促進(jìn)AR表達(dá)進(jìn)而參與調(diào)控鼠卵巢功能。

綜上所述，在本試驗(yàn)中，腹腔注射不同質(zhì)量分?jǐn)?shù)亞精胺后，鼠卵巢組織中LHR、FSHR、ER1、ER2和AR表達(dá)量相較于對(duì)照組均發(fā)生了顯著變化。低質(zhì)量分?jǐn)?shù)的亞精胺促進(jìn)LHR表達(dá)；高質(zhì)量分?jǐn)?shù)的亞精胺促進(jìn)FSHR、ER1、ER2和AR表達(dá)，同時(shí)抑制LHR的表達(dá)，且具有劑量依賴(lài)性。說(shuō)明亞精胺可通過(guò)介導(dǎo)鼠卵巢組織生殖激素受體基因表達(dá)來(lái)參與調(diào)控鼠卵巢功能。

[1]向睿,何琿,康波.多胺調(diào)控動(dòng)物繁殖的作用及其機(jī)制.動(dòng)物營(yíng)養(yǎng)學(xué)報(bào),2014,26(11):3251-3255. XIANG R,HE H,KANG B.Effects of polyamine regulation on animal reproduction and its mechanism.Chinese Journal of Animal Nutrition,2014,26(11):3251-3255.(in Chinese with English abstract)

[2]PEGG A E,CASERO R A.Current status of the polyamine research field.Polyamines:Methods in Molecular Biology,2011, 720:3-35.

[3]VIJAYANATHAN V,THOMAS T J,NAIR S K,et al.Bending of the estrogen response element by polyamines and estrogen receptors alpha and beta:A fluorescence resonance energy transfer study.The International Journal of Biochemistry and Cell Biology,2006,38(7):1191-1206.

[4]LEFEVRE P L C,PALIN M F,MURPHY B D.Polyamines on the reproductive landscape.Endocrine Reviews,2011,32(5):694-712.

[5]GONZALEZ-MONTELONGO M C,MARIN R,PEREZ J A,et al.Polyamines transduce the nongenomic,androgen-induced calcium sensitization in intestinal smooth muscle.Molecular Endocrinology,2013,27(10):1603-1616.

[6]FASHE T M,KEINANEN T A,GRIGORENKO N A,et al.Cutaneous application of alpha-methylspermidine activates the growth of resting hair follicles in mice.Amino Acids,2010,38(2): 583-590.

[7]趙越超.小鼠圍著床期子宮中多胺相關(guān)基因的表達(dá)、調(diào)節(jié)與功能.哈爾濱:東北農(nóng)業(yè)大學(xué),2008:61-67. ZHAOYC.Expression,regulation and function ofpolyamie-related genes in mouse uterus during peri-implantation period.Harbin: Northeast Agricultural University,2008:61-67.(in Chinese with Englishabstract)

[8]MILLER-FLEMING L,OLIN-SANDOVAL V,CAMPBELL K,et al.Remaining mysteries of molecular biology:The role of polyamines in the cell.Journal of Molecular Biology,2015,427 (21):3389-3406.

[9]CALANDRA R S,RULLI S B,FRUNGIERI M B,et al.Polyamines in the male reproductive system.Acta physiologica, pharmacologica et therapeutica latinoamericana:órgano de la Asociación Latinoamericana de Ciencias Fisiológicas y[de]la Asociación Latinoamericana de Farmacología,1996,46(4):209-222.

[10]MELENDREZ C S,RUTTLE J L,HALLFORD D M,et al.Polyamines in ejaculated ram spermatozoa and their relationship with sperm motility.Journal of Andrology,1992,13(4):293-296.

[11]KANG B,GUO J R,YANG H M,et al.Differential expression profiling of ovarian genes in prelaying and laying geese.Poultry Science,2009,88(9):1975-1983.

[12]NATARAJA S G,YU H N,PALMER S S.Discovery and development of small molecule allosteric modulators of glycoprotein hormone receptors.Frontiers in Endocrinology,2015, 6:142.

[13]MEHL N S,KHALID M,SRISUWATANASAGUL S,et al.GnRH-agonist implantation of prepubertal male cats affects their reproductive performance and testicular LH receptor and FSH receptor expression.Theriogenology,2016,85(5):841-848.

[14]LAU K M,TO K F.Importance of estrogenic signaling and its mediated receptors in prostate cancer.International Journal of Molecular Sciences,2016,17(9):1434.

[15]AYOUB M A,YVINEC R,JEGOT G,et al.Profiling of FSHR negative allosteric modulators on LH/CGR reveals biased antagonism with implications in steroidogenesis.Molecular and Cellular Endocrinology,2016,436:10-22.

[16]LóPEZ-DOVAL S,SALGADO R,LAFUENTE A.The expression of several reproductive hormone receptors can be modified by perfluorooctane sulfonate(PFOS)in adult male rats.Chemosphere,2016,155:488-497.

[17]SAMARDZIJA D,POGRMIC-MAJKIC K,FA S,et al.Atrazine blocks ovulation via suppression ofLhrandCyp19a1mRNA and estradiol secretion in immature gonadotropin-treated rats.Reproductive Toxicology,2016,61:10-18.

[18]THYSSEN S M,HOCKL P F,CHAMSON A,et al.Effects of polyamines on the release of gonadotropin-releasing hormone and gonadotropins in developing female rats.Experimental Biology and Medicine,2002,227(4):276-281.

[19]LIU K C,GE W.Differential regulation of gonadotropin receptors (fshrandlhcgr)by epidermal growth factor(EGF)in the zebrafish ovary.General and Comparative Endocrinology,2013,181:288-294.

[20]BLACHOWSKI S,MOTYL T,GRZELKOWSKA K,et al.Involvement of polyamines in epidermal growth factor(EGF), transforming growth factor(TGF)-alpha and-beta 1 action on culture of L6 and fetal bovine myoblasts.International Journal of Biochemistry,1994,26(7):891-897.

[21]SAUNDERS P T K,MILLARMR,WILLIAMS K,etal.Differential expression of estrogen receptor-alpha and-beta and androgen receptor in the ovaries of marmosets and humans.Biology of Reproduction,2000,63(4):1098-1105.

[22]鄭可佳,楊彩榮,張巖,等.ERβ及p-ERβ在小鼠發(fā)情周期卵巢內(nèi)的表達(dá).東北農(nóng)業(yè)大學(xué)學(xué)報(bào),2009,40(7):85-89. ZHENG K J,YANG C R,ZHANG Y,et al.Study on the expression of ERβandp-ERβon mouse ovary during the oestrous cycle.Journal of Northeast Agricultural University,2009,40(7): 85-89.(in Chinese with English abstract)

[23]WALTERS K A.Role of androgens in normal and pathological ovarian function.Reproduction,2015,149(4):193-218.

[24]KIMURA S,MATSUMOTO T,MATSUYAMA R,et al.Androgen receptor function in folliculogenesis and its clinical implication in premature ovarian failure.Trends in Endocrinology and Metabolism,2007,18(5):183-189.

[25]DU X,LI Q Q,PAN Z X,et al.Androgen receptor and miRNA-126*axis controls follicle-stimulating hormone receptor expression in porcine ovarian granulosa cells.Reproduction,2016, 152(2):161-169.

[26]ZHU T Y,YUAN J,XIE Y D,et al.Association of androgen receptor CAG repeat polymorphism and risk of epithelial ovarian cancer.Gene,2016,575(2):743-746.

Effect of exogenous spermidine on expression profile of reproductive hormone receptors in mouse ovaries.

LONG Shiyun,JIANG Dongmei,CHEN Ziyu,GUAN Cheng,YI Zhixin,KANG Bo*(Farm Animal Genetic Resources Exploration and Innovation Key Laboratory of Sichuan Province/College of Animal Science and Technology,Sichuan Agricultural University, Chengdu 611130,China)

spermidine;ovary;hormone receptor;gene expression

Q 492

A

10.3785/j.issn.1008-9209.2016.09.191

Journal of Zhejiang University(Agric.&Life Sci.),2017,43(2):247-252

國(guó)家自然科學(xué)基金（31201798）；高等學(xué)校博士學(xué)科點(diǎn)專(zhuān)項(xiàng)科研基金（20105103120003）。

康波（http://orcid.org/0000-0003-2811-0642），E-mail:bokang@sicau.edu.cn

（First author）：龍?jiān)婍崳╤ttp://orcid.org/0000-0002-5735-2731），E-mail:longsy210@163.com

2016-09-19；接受日期(Accepted):2016-12-06