張 博 蔡 爽 葉倩紅 曾祥芳 譙仕彥

(中國農業(yè)大學動物科技學院，北京 100193)

營養(yǎng)物質調節(jié)母豬卵母細胞質量與繁殖性能的研究進展

張 博 蔡 爽 葉倩紅 曾祥芳*譙仕彥

(中國農業(yè)大學動物科技學院，北京 100193)

飼糧中能量、脂肪、粗蛋白質和纖維會影響到卵泡發(fā)育、發(fā)情周期、激素水平、胚胎附植和產仔數。卵母細胞質量是決定哺乳動物繁殖性能的關鍵因素，影響優(yōu)質胚胎的數量和懷孕率。卵母細胞質量受到糖類、氨基酸和脂肪等營養(yǎng)物質的調節(jié)，營養(yǎng)物質代謝異常會降低卵母細胞質量。本文就最新關于營養(yǎng)物質對母豬卵母細胞成熟和早期胚胎發(fā)育及繁殖性能的影響的研究進行綜述，為通過營養(yǎng)調控提高母豬卵母細胞發(fā)育潛能和繁殖性能的研究與實際應用提供參考。

母豬；卵母細胞質量；繁殖性能；能量；氨基酸；脂肪

早期胚胎發(fā)育對哺乳動物的繁殖性能有著深遠影響。不同哺乳動物早期胚胎死亡率為30%～50%[1]。如何促進早期胚胎發(fā)育、提高卵母細胞質量成為提高繁殖性能的關鍵。在卵母細胞成熟和發(fā)育過程中伴隨著物質代謝和能量代謝。糖類、氨基酸和脂肪等營養(yǎng)物質可調節(jié)卵母細胞質量、受精能力和胚胎發(fā)育[2]。營養(yǎng)物質與繁殖性能密切相關，如能量、蛋白質、脂肪和纖維會影響母豬的發(fā)情、妊娠和產仔。因此，通過營養(yǎng)方法提高卵子及胚胎發(fā)育潛能，對提高母豬的繁殖性能具有重要意義。

1 營養(yǎng)物質對母豬繁殖性能的影響

1.1 飼糧能量來源

碳水化合物作為能量來源可促進卵巢上卵泡的發(fā)育。在母豬后備期至發(fā)情期，以淀粉為主要能量來源比脂肪型飼糧更能促進卵母細胞成熟[3]，這可能與淀粉組的血清和卵泡液中胰島素樣生長因子(IGF)含量升高有關，IGF可以促進下丘腦分泌促卵泡素(FSH)和促黃體素(LH)，并且在卵丘復合體中LH受體mRNA表達水平升高，LH升高有利于后期卵泡發(fā)育。初產母豬哺乳時由于采食量不夠，導致機體分解代謝增加，不利于卵泡發(fā)育和激素分泌[4]。為了增加能量攝入，初產母豬的哺乳飼糧中添加蔗糖，有縮短斷奶后發(fā)情間隔的趨勢，顯著提高妊娠率和血漿中孕酮含量，并且顯著增加下一胎次產仔數，而哺乳期間體重損失和采食量不受影響[5]。有研究表明，當初產母豬在哺乳期間體重損失較小時，與添加脂肪相比，飼糧中添加膨化小麥、葡萄糖和蔗糖能夠促進斷奶后發(fā)情和減少能量負平衡，并不影響產仔數[6]。葡萄糖可以顯著提高后備母豬發(fā)情時血清中雌激素、LH和FSH含量，使后備母豬初情期提前[7]。因此飼糧中淀粉、蔗糖和葡萄糖可以被機體利用，減少母體脂肪動員和體重損失，有利于發(fā)情周期中激素水平升高，從而促進卵泡發(fā)育。

脂肪作為重要能量來源，其在飼糧中的含量和類型會影響母豬的繁殖性能。給妊娠期母豬飼喂高脂飼糧，能夠增加母豬體重，從而顯著提高總產仔數、活產仔數和仔豬斷奶體重，且縮短母豬斷奶后發(fā)情的間隔[8]。母豬飼糧還影響仔豬脂肪代謝，在妊娠后期和哺乳期給母豬飼喂高水平玉米油，仔豬斷奶時皮下脂肪組成和母乳相同，參與脂肪酸代謝的硬脂酰輔酶A去飽和酶顯著下降[9]。不同類型的脂肪對母豬和仔豬的影響不同，有些脂肪易引起氧化應激[8]，如魚油會升高母豬血清、初乳和仔豬血清丙二醛的含量，表明魚油會造成母豬和仔豬氧化應激；而橄欖油可使母乳中白細胞介素-6和腫瘤壞死因子含量下降，從而降低仔豬斷奶前死亡率[10]。母豬高脂飼糧對仔豬的繁殖性能造成不利影響。飼喂高脂飼糧會降低抗氧化能力，減少后代母豬初情期的卵泡數量[11]。

1.2 飼糧能量水平

在經產母豬妊娠期的前期、中期和后期，飼喂7 d不同能量水平的飼糧，隨著能量水平的提高，母豬的體重和背膘厚會增加，這有利于減少泌乳期間體重損失，盡管采食量呈下降趨勢；有利于仔豬初生重提高，但不影響仔豬斷奶體重；且母豬下一胎次的繁殖性能不受影響[12]。短期高能飼糧對后備母豬具有促進作用，在后備期至發(fā)情期飼喂高能飼糧，可以顯著增加大卵泡數量和提高卵母細胞成熟率[3]。能量水平過低則會對繁殖造成不利影響，后備母豬發(fā)情周期前期限飼，會顯著降低血液中的孕酮、吻蛋白和IGF水平，大卵泡和黃體數量也顯著減少，同時發(fā)情表現異常[13]。因此母體在發(fā)情周期前期對能量需求升高，飼喂低能量水平飼糧會抑制繁殖性能，而短期高能量水平飼糧則具有促進作用。

1.3 飼糧粗蛋白質水平

研究表明，低氮飼糧會影響哺乳母豬氮利用效率。母豬哺乳飼糧的粗蛋白質水平由16.0%下降為14.3%，同時補充限制氨基酸，不會影響平均日采食量和體重變化；泌乳前期乳中凈蛋白利用效率、酪蛋白和乳真蛋白水平無顯著差異，而泌乳高峰期乳中凈蛋白利用效率和酪蛋白水平顯著升高；且仔豬生長速度有升高趨勢[14]。當飼糧中粗蛋白質水平降低太多，且不補充限制氨基酸時會抑制仔豬生長。母豬妊娠期和哺乳期飼糧粗蛋白質水平下降50%，肌生成抑制蛋白及其激活蛋白——叉頭框蛋白O3(FoxO3)的mRNA表達水平顯著上調，使仔豬的日增重顯著下降和生長遲緩[15]。飼糧粗蛋白質水平太低還會降低后代繁殖性能，母豬妊娠期和哺乳期飼糧中粗蛋白質水平下降50%，仔豬卵巢的細胞色素芳香酶含量增加，而RNA酶Drosha和細胞色素芳香酶microRNA含量降低，不利于卵泡成熟，成熟卵泡數量減少；并增加凋亡因子半胱天冬酶活性，顆粒細胞凋亡增加[16]。

母豬飼糧中粗蛋白質與碳水化合物比值也會影響到母豬及仔豬代謝。低粗蛋白質高碳水化合物的飼糧會升高血糖與胰島素比值，降低胰島素含量；高粗蛋白質低碳水化合物飼糧會增加胰高血糖素及餐后血糖濃度；并且這2種飼糧都會增加仔豬肝臟中磷酸烯醇式丙酮酸羧基酶和葡萄糖-6-磷酸激酶mRNA水平，從而促進仔豬對葡萄糖利用[17]。

1.4 飼糧纖維來源

飼糧中纖維對繁殖性能具有促進作用，以羽扇豆作為纖維來源比小麥麩纖維更好，因為羽扇豆纖維可以提高發(fā)育至MⅡ期卵母細胞數量；配種前短期飼喂羽扇豆纖維還可以提高胚胎存活率[18]。前人研究表明，與不可溶性纖維相比，可溶性纖維雖然不影響母豬排卵，但能夠提高胚胎成活率和增加活胚胎數量[19]。魔芋粉中含有較多的可溶性纖維，可提高中性洗滌纖維消化率，增加母豬哺乳期采食量，且仔豬斷奶重有升高趨勢[20]。飼糧中一定量的粗纖維不會影響繁殖性能，添加13.4%Tifton干草不會影響產仔數和仔豬斷奶重[21]。且用富含纖維的蔬菜部分代替飼糧中的大麥和小麥，不會影響妊娠母豬血糖和胰島素含量，但會提高血清中短鏈脂肪酸和非酯化脂肪酸含量，并且血液中甜菜堿、二甲基砜和鯊肌醇可作為蔬菜的生物標記分子[22]。

2 卵母細胞質量及其營養(yǎng)調節(jié)

卵母細胞質量在一定程度上決定了早期胚胎發(fā)育和附植，質量差的卵母細胞即使能夠完成受精，形成合子后卵裂率、優(yōu)質胚胎數量[23]和胚胎附植率都顯著降低[24]。卵母細胞成熟和早期胚胎發(fā)育過程受到糖類、氨基酸和脂肪調節(jié)，現總結如圖1。

圖1 營養(yǎng)物質對卵母細胞質量調節(jié)

2.1 糖類物質

2.1.1 葡萄糖

葡萄糖是最重要的單糖，其代謝對卵母細胞成熟和發(fā)育起著重要作用。豬卵母細胞體外試驗表明，添加葡萄糖或丙酮酸可增加卵母細胞發(fā)育至第二次減數分裂中期(MⅡ期)數量和提高囊胚率；卵母細胞需要顆粒細胞把葡萄糖轉化成丙酮酸才能利用，因此卵母細胞沒有顆粒細胞時只能利用丙酮酸，不能利用葡萄糖[25]。老化的豬卵母細胞出現葡萄糖轉運下降[26]，所以葡萄糖轉運異常會降低卵母細胞質量。冷凍保存會對豬胚胎造成損傷，降低胚胎存活率，細胞凋亡率增加；與添加丙酮酸相比，成熟液中添加葡萄糖可以顯著提高胚胎冷凍后的囊胚率[27]。卵母細胞對葡萄糖吸收利用受多種物質的調節(jié)，骨形態(tài)生成蛋白可以促進卵丘卵母細胞復合體對葡萄糖的利用[28]，胰島素會通過促進顆粒細胞增殖來提高對葡萄糖的利用[29]，而肉堿會促進卵母細胞脂肪酸β氧化而抑制對葡萄糖的吸收利用[30]。

葡萄糖經過糖酵解轉變?yōu)槿樘遣a生ATP，通過磷酸戊糖途徑產生煙酰胺腺嘌呤二核苷酸磷酸(NADP)和磷酸核糖，這2條途徑在卵母細胞成熟和發(fā)育過程起著重要作用。豬卵母細胞成熟液中添加磷酸戊糖途徑抑制劑和糖酵解抑制劑，會顯著降低卵母細胞內ATP和谷胱甘肽，從而抑制卵母細胞成熟；增加與凋亡相關基因(半胱天冬酶和Bax)表達，使囊胚細胞凋亡率升高[25]。這2條途徑的生理性抑制劑如ADP和NADP并不影響卵母細胞成熟；而藥理性抑制劑如氟化鈉和氨基煙酰胺，通過改變卵母細胞氧化狀態(tài)和線粒體活性，抑制卵母細胞成熟[31]。

2.2 氨基酸

母豬卵泡液中谷氨酰胺、丙氨酸、甘氨酸、谷氨酸和脯氨酸含量最高[32]。豬胚胎細胞在培養(yǎng)過程中伴隨著氨基酸吸收和產生，有些氨基酸的含量一直下降，如谷氨酰胺、蘇氨酸和精氨酸；有些氨基酸的含量一直升高，如谷氨酸和甘氨酸[33]，這表明卵母細胞或胚胎不同發(fā)育階段對氨基酸利用不同。

2.2.1 精氨酸及其代謝物

精氨酸是條件性必需氨基酸，妊娠期時母體對精氨酸需要量升高。精氨酸是內源合成一氧化氮(NO)的唯一前體物質，NO可以調節(jié)細胞內多條信號通路和促進血管生成。飼糧中直接添加精氨酸或氮氨甲酰谷氨酸(可促進精氨酸內源合成)能通過激活mTOR信號通路，促進滋養(yǎng)層生長和胚胎著床，進而提高小鼠和豬的繁殖性能[34-35]。豬卵母細胞體外培養(yǎng)試驗中，添加NO合成酶抑制劑會顯著抑制卵母細胞恢復減數分裂和顆粒細胞擴展[36]。

精氨酸在體內可經過鳥氨酸循環(huán)生成鳥氨酸，而鳥氨酸在脫羧酶作用下生成腐胺，小鼠卵巢中鳥氨酸和腐胺會隨著卵母細胞成熟周期性變化。鳥氨酸脫羧酶和腐胺在老齡小鼠中容易缺乏，通過飲水補充腐胺會提高卵巢中腐胺水平，減少胚胎被吸收和增加活仔數[37]。在老齡小鼠卵母細胞體外培養(yǎng)試驗中，腐胺可以顯著提高囊胚細胞數和優(yōu)質胚胎比例[38]。

2.2.2 支鏈氨基酸

支鏈氨基酸不僅能夠調節(jié)機體肌肉蛋白質合成，還對卵母細胞發(fā)育有調節(jié)作用。在沙鼠胚胎體外培養(yǎng)試驗中，八細胞胚胎對纈氨酸的吸收顯著增加，吸收后的纈氨酸用于氧化供能，同位素標記法表明其碳鏈用于合成脂質[39]，因此纈氨酸可以作為能量來源。但最新研究表明，對于產仔數高于12頭的哺乳母豬，飼糧中添加纈氨酸對母豬代謝、產奶量和仔豬生長并沒有促進作用[40]。

前人研究表明，不同發(fā)育階段的卵母細胞或囊胚的亮氨酸轉運載體不同，卵母細胞主要通過鈉依賴型載體運輸亮氨酸，而發(fā)育至囊胚階段主要通過鈉非依賴性轉運載體[41]。近期發(fā)現不同發(fā)育階段的卵泡對亮氨酸吸收速率不同，小鼠卵泡在無腔卵泡、有腔卵泡和三級卵泡發(fā)育過程中，對亮氨酸吸收會逐漸增加。這表明在卵泡發(fā)育過程中細胞分裂和代謝增強需要更多的亮氨酸，通過顆粒細胞增殖可以增加膜轉運載體和緊密連接，從而提高亮氨酸轉運效率。但排卵前的卵泡對亮氨酸吸收下降，這可能與卵泡內膠原酶和明膠酶增加，減少卵母細胞與顆粒細胞氨基酸轉運有關[42]。

2.2.3 蛋氨酸

表觀遺傳包括DNA修飾和組蛋白修飾。卵母細胞成熟和早期胚胎發(fā)育過程中伴隨著表觀遺傳變化，蛋氨酸和甜菜堿作為重要甲基供體會影響仔豬表觀遺傳。研究表明，母豬妊娠期飼喂甜菜堿，可顯著降低仔豬肝臟中甘油三酯含量，且脂肪合成基因表達水平顯著下調，如乙酰輔酶A、硬脂酰輔酶A去飽和酶和脂肪酸合成酶，這可能與甜菜堿升高肝臟中腺苷甲硫氨酸與腺苷半胱氨酸比值有關。該比值升高可促進脂肪酸合成酶和硬脂酰輔酶A去飽和酶基因啟動子上的DNA超甲基化[43]。同樣母豬妊娠期飼喂甜菜堿可促進仔豬甜菜堿/蛋氨酸代謝和DNA甲基轉移酶的表達，使IGF2基因片段高度甲基化，增加新生仔豬大腦海馬區(qū)中IGF2表達和細胞增殖/抗凋亡[44]。

2.3 脂肪酸

2.3.1 脂肪酸β氧化

脂肪酸代謝可以為卵母細胞提供ATP，脂肪酸需要進入線粒體內才能進行β氧化，但該反應是限速反應，需要肉堿棕櫚酰轉移酶催化。卵母細胞脂滴和線粒體共定位試驗表明，豬卵母細胞成熟過程中脂滴與線粒體逐漸靠近，進而促進脂肪酸β氧化；添加β氧化抑制劑會顯著抑制卵母細胞恢復減數分裂和早期胚胎發(fā)育[45-46]。研究表明，即使在沒有碳水化合物提供能量情況下，添加肉堿可提高豬卵母細胞的卵裂率。且肉堿對冷凍豬胚胎細胞具有保護作用，發(fā)育液中添加肉堿可以提高解凍后胚胎存活率[47]。

2.3.2 不飽和脂肪酸

與飽和脂肪酸相比，不飽和脂肪酸有利于卵母細胞質量。用高分辨率核磁共振對卵泡液進行脂質代謝分析，發(fā)現不能進行卵裂的卵泡液中飽和脂肪酸含量顯著升高，而多不飽和脂肪酸含量則顯著下降，這些差異脂肪酸可以用于判斷卵母細胞在受精后能否正常卵裂[48]。母豬成熟卵母細胞的顆粒細胞中多不飽和脂肪酸含量，顯著高于沒有成熟顆粒細胞中的含量；卵母細胞中的飽和脂肪酸的組成在其成熟過程中無顯著變化[49]，這表明很少飽和脂肪酸參與卵母細胞代謝。

研究表明，給哺乳期的母豬飼喂必需脂肪酸亞麻酸，可促進3胎次以上母豬斷奶后發(fā)情，且下一胎次產仔數有增加趨勢[50]。在母豬卵母細胞體外培養(yǎng)試驗中，添加亞麻酸可顯著提高卵母細胞內谷胱甘肽含量，減少氧化應激，從而加速細胞核成熟、提高囊胚率和增加體細胞核移植數；絲裂原激活蛋白激酶(MAPK)抑制劑不利于卵母細胞成熟和早期胚胎發(fā)育，而亞麻酸可完全緩解MAPK抑制劑的不利影響[51]。共軛亞油酸對卵母細胞同樣具有上述促進作用，且MAPK表達水平顯著升高[52]；共軛亞油酸能夠降低細胞質顏色深度[53]，這表明共軛亞油酸可通過促進細胞質中脂滴分解，加快脂肪酸代謝為卵母細胞提供能量。

3 小 結

飼糧中能量來源和水平、粗蛋白質、脂肪和纖維直接影響著繁殖性能，如發(fā)情周期、激素和產仔數等。糖類、氨基酸和脂肪等營養(yǎng)物質，會通過調控能量供應、表觀遺傳修飾、線粒體功能和氧化平衡，從而影響卵母細胞成熟和早期胚胎發(fā)育。營養(yǎng)物質代謝異常會降低卵母細胞質量和繁殖性能。因此仍存在很多問題有待解決：一是如何找到卵母細胞質量的標記物質；二是如何添加營養(yǎng)物質實現繁殖性能的最大潛能?？傊?，需要更多的研究來探究繁殖性能與營養(yǎng)物質的關系，最終為通過營養(yǎng)方法提高繁殖性能提供理論支持。

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(責任編輯 田艷明)

Abstract: Nutrients like energy, fat, crude protein and fiber all influence follicular development, estrous cycle, hormone levels, embryos implantation and litter size. Oocyte quality is a key factor in determining the reproductive performance of sows, and influences the number of high quality embryos and pregnancy rate. Oocytes quality is regulated by nutrients including carbohydrate, amino acids and fats, so abnormal metabolism of nutrients will decrease the quality of oocytes. This review summarized the recent study of nutrients regulation of oocytes quality and reproductive performance, with the aim to improve oocytes development potential and practical application in commercial industry through nutritional modulation.[ChineseJournalofAnimalNutrition,2017,29(10)：3436-3443]

Keywords: sows; oocytes quality; reproductive performance; energy; amino acids; fat

ResearchProgressinNutrientsRegulationofSowOocyteQualityandReproductivePerformance

ZHANG Bo CAI Shuang YE Qianhong ZENG Xiangfang*QIAO Shiyan

(CollegeofAnimalScienceandTechnology,ChinaAgriculturalUniversity,Beijing100193,China)

S828

A

1006-267X(2017)10-3436-08

10.3969/j.issn.1006-267x.2017.10.002

2017-04-06

國家自然科學基金(2014C17-31301980)

張 博(1991—)，男,河南民權人，碩士研究生，從事豬營養(yǎng)研究。E-mail： 15311508532@163.com

*通信作者:曾祥芳，副教授，碩士生導師，E-mail： zengxf@cau.edu.cn

*Corresponding author, associate professor, E-mail: zengxf@cau.edu.cn