Kisspeptin对鱼类生殖轴的调控机制研究*

王 滨 柳学周 徐永江 史 宝 刘 权



Kisspeptin对鱼类生殖轴的调控机制研究*

王 滨1,2#柳学周1,2,①#徐永江1,2史 宝1,2刘 权1,3

(1. 农业农村部海洋渔业可持续发展重点实验室 中国水产科学研究院黄海水产研究所 青岛 266071； 2. 青岛海洋科学与技术试点国家实验室海洋渔业科学与食物产出过程功能实验室 青岛 266071； 3. 上海海洋大学水产与生命学院 上海 201306)

Kisspeptin (简称Kiss或者Kp)是由基因编码的一种下丘脑神经肽，通过其受体KissR(也称作GPR54)的介导参与了多种生理过程，如抑制肿瘤转移和参与生殖调控。目前，尽管在鲤形目(Cypriniformes)、鲈形目(Perciforms)、鲽形目(Pleuronectiforms)、鲀形目(Tetraodontiforms)、颌针目(Beloniforms)、鲉形目(Scorpaeniformes)、鲑形目(Salmoniformes)及鳕形目(Gadiformes)等多种鱼类中均鉴定出了基因，但Kiss/KissR系统在鱼类生殖调控中的精确作用及其分子机制尚未完全阐明。尤其是在鱼类中存在2种及3种基因，Kiss/KissR系统对鱼类生殖调控的作用方式更加复杂。本文简要总结鱼类Kiss及其受体的研究进展，并对Kiss的生理学功能、信号转导机制以及表达调控研究进行概括讨论，旨在加深对鱼类Kiss/KissR系统的认识和了解，为后续研究指明方向。

鱼类；Kisspeptin；kisspeptin receptor；生殖；信号转导；基因表达调控

下丘脑神经肽kisspeptin及其受体KissR在哺乳动物生殖调控及青春期启动中发挥了重要作用(Roa, 2011; Tena-Sempere, 2010)。迄今，除鸟类外，在其他脊椎动物中均鉴定出了基因。除鸭嘴兽()外，哺乳类只存在基因；两栖类存在及三种基因；爬行类只存在基因；斑马鱼()、青鳉()、金鱼()、欧洲海鲈()、条纹鲈()及鲐鱼()中存在和两种基因。相反，在尼罗罗非鱼()、斜带石斑鱼()、塞内加尔鳎()、半滑舌鳎()以及星点东方鲀()中只鉴定出了基因(Pasquier, 2014; Um, 2010; Wang, 2017b)。目前，已在多种鱼类中鉴定出了Kiss系统，其在鱼类生殖调控中的生理功能研究也日益完善(Akazome, 2010; Mechaly, 2013; Tena- Sempere, 2012)。本文简要总结鱼类Kiss及其受体的研究进展，并对Kiss的生理学功能、信号转导机制以及表达调控研究进行概括讨论，旨在加深对鱼类Kiss/KissR系统的认识和了解，为后续研究奠定基础。

1 Kisspeptin的发现及与生殖的关系

基因最初是从人()黑色素瘤和乳腺癌细胞中分离得到的，因其具有抑制肿瘤生长和转移的功能，Kiss最初被命名为转移抑制素(Metastin) (Lee, 1996、1997)。Lee等(1999)从大鼠()脑中鉴定出了1种新型G蛋白偶联受体，命名为GPR54。2年后，Kiss被认为是孤儿受体GPR54的内源性配体(Kotani, 2001; Muir, 2001; Ohtaki, 2001)。2003年，2个独立研究组发现，突变导致人特发性性腺功能减退(de Roux, 2003; Seminara, 2003)。随后研究发现，基因敲除或者均影响性腺发育及生殖功能(d¢Anglemont de Tassigny, 2007; Seminara, 2003)，说明Kiss/GPR54系统在哺乳类生殖调控中发挥了关键作用。

近几年，kisspeptin在鱼类生殖调控中的作用也有较多研究。如Kiss1直接促进了金鱼垂体细胞黄体生成素(Luteinizing hormone, LH)分泌(Chang, 2012; Yang, 2010)。Kiss2也促进了欧洲海鲈(Espigares, 2015b)和条纹鲈(Zmora, 2015)垂体细胞LH及卵泡刺激素(Follicle-stimulating hormone, FSH)分泌。此外，Kiss1增加了金鱼垂体细胞的表达水平(Yang, 2010)。然而，Kiss1特异性地降低了欧洲鳗鲡垂体细胞的表达水平 (Pasquier, 2011)。腹腔注射Kiss2促进了斑马鱼垂体及的表达水平(Kitahashi, 2009)，而Kiss2特异性地促进了斜带石斑鱼垂体的表达量，对的表达水平无影响(Shi, 2010)。综上所述，kisspeptin参与了鱼类生殖调控，但具体作用机制因物种而异。

2 鱼类kiss基因类型、结构及时空表达特性

由于基因不是很保守，直到2008年才在非哺乳类中鉴定出了其同源基因。van Aerle等(2008)利用全基因组序列及比较共线性方法，首次在斑马鱼和青鳉等5种鱼类中鉴定出了基因。随后，Biran等(2008)和Kanda等(2008)也通过类似方法，分别在斑马鱼和青鳉中获得了基因。2009年，基因首次在斑马鱼、青鳉和欧洲海鲈中被鉴定出来(Felip, 2009; Kitahashi, 2009)。斑马鱼基因编码116个氨基酸的前体多肽，其C末端核心十肽为YNLNSFGLRY (Y-Y形式) (Biran, 2008; van Aerle, 2008)；斑马鱼基因编码125个氨基酸的前体多肽，其C末端核心十肽为FNYNPFGLRF (F-F形式) (Kitahashi, 2009)。与之类似，其他鱼类C末端十肽序列与斑马鱼高度保守，该十肽也是发挥其功能所需的最短序列(Akazome, 2010; Pasquier, 2014)。在哺乳类中，基因由3个外显子和2个内含子组成，其中，外显子1只编码一部分5¢UTR，外显子2编码另一部分5¢UTR及一部分CDS，剩余另一部分CDS及3¢UTR由外显子3编码(Pasquier, 2014)。同样，斑马鱼基因也是由3个外显子和2个内含子组成，而基因由2个外显子和1个内含子组成(Kitahashi, 2009)。塞内加尔鳎基因也是由2个外显子和1个内含子组成，但是，其存在2种剪接变异体：较短亚型编码正常Kiss2前体多肽；较长亚型编码一种缩短形式的无功能多肽(Mechaly, 2011)。

鱼类及的组织分布因物种而异，即使同一物种不同脑区表达也有所差异。斑马鱼主要在间脑和中脑中表达，其次为后脑，在端脑和垂体中表达量较低(Biran, 2008)；在外周组织中，斑马鱼在胰腺和前肠中表达量较高，其次为性腺(Biran, 2008)。与之类似，青鳉(Felip, 2009; Kitahashi, 2009)、欧洲海鲈(Felip, 2009)、金鱼(Li, 2009; Yang, 2010)、鲐鱼(Shahjahan, 2010)等脑和性腺中表达量也较高。也主要在脑和性腺中高表达，如斑马鱼(Kitahashi, 2009)、青鳉(Kitahashi, 2009)、金鱼(Li, 2009)、欧洲海鲈(Felip, 2009)、塞内加尔鳎(Mechaly, 2011)及南亚黑鲮() (Saha, 2016)等。此外，也在肠、肾脏、心脏等其他外周组织有所表达，具体表达模式具有物种特异性。

鱼类基因在不同发育阶段/生殖周期的表达模式也在斑马鱼等几种鱼类中有所报道。雌性斑马鱼脑表达量在孵化后逐渐升高，84 d时达到峰值；而雄性斑马鱼脑表达量在孵化后6周达到峰值，12周时有所下降(Biran, 2008)。此外，斑马鱼表达量在孵化后30 d达到峰值(Kitahashi, 2009)。上述结果显示，可能参与了斑马鱼青春期启动。鲐鱼脑在不同生殖周期的表达模式具有性别二态性，雄性脑表达量随精巢发育逐渐降低，而雌性脑表达量在卵巢发育过程中保持不变；除了分别在卵黄生成早期和精子生成晚期略微增加外，雌雄脑表达量随性腺发育逐渐降低，均在产卵/排精后达到最小值(Selvaraj, 2010)。然而，精巢表达水平随性腺发育逐渐升高，在精子成熟时期达到峰值；卵巢表达水平也随性腺发育逐渐升高，在卵黄生成后期达到峰值(Selvaraj, 2010)。以上结果表明，可能参与了鲐鱼季节性性腺发育。其他鱼类表达水平也随性腺发育而发生波动(Alvarado, 2013; Migaud, 2012; Park, 2016; Saha, 2016; Shahi, 2017)。

3 鱼类kissr基因类型、结构及时空表达特性

通常，哺乳类下丘脑的表达水平在青春期显著性增加(Dungan, 2006)。鱼类的表达模式也与生殖周期有关。鲻鱼脑的表达水平随性腺发育而降低，在青春期前期表达量最高(Nocillado, 2007)。与之类似，军曹鱼、黑头呆鱼及大西洋庸鲽脑的表达量也均在青春期达到峰值(Filby, 2008; Mechaly, 2010; Mohamed, 2007)。斑马鱼脑的表达量在孵化后8周时显著性增加，随后回到本底水平；而的表达量在孵化后6周时显著增加，随后一直保持到12周(Biran, 2008)。鲐鱼脑在不同生殖周期的表达模式具有性别二态性，雄鱼脑及的表达水平不随精巢发育过程而变化；而雌鱼脑及的表达水平均在卵黄生成早期显著增加并达到峰值，继而随卵巢发育过程又回到本底水平(Ohga, 2013)。精巢表达水平随性腺发育逐渐升高，在精子成熟时期达到峰值；而精巢表达水平不随性腺发育过程而变化(Ohga, 2013)。综上所述，可能参与了鱼类青春期启动及季节性性腺发育。

4 Kisspeptin对鱼类生殖调控作用研究

4.1 Kisspeptin对下丘脑促性腺激素释放激素(Gonadotropin-releasing hormone, GnRH)神经元活性以及表达调控的影响

GnRH是垂体促性腺激素合成与分泌的主要促进因子，在每种硬骨鱼类中存在至少2种GnRH多肽(Zohar, 2010; 王滨等, 2017)。Parhar等(2004)首次在罗非鱼中鉴定出了基因，并进一步证实在GnRH1、GnRH2及GnRH3神经元中表达，这表明Kiss2能够直接作用于GnRH神经元，进而影响其活性及表达调控。在青鳉中，通过电生理学研究表明，Kiss1能够促进GnRH3神经元的电活动(Electrical activity)，而河豚毒素或者阻断突触传递均降低了Kiss1诱导的GnRH3神经元的电活动，这表明Kiss1以间接方式通过突触调控进而激活GnRH3神经元的电活动(Zhao, 2012)。

4.2 Kisspeptin对垂体激素合成与分泌的影响

由于鱼类中存在2种Kiss多肽，Kiss对鱼类垂体激素分泌的影响更加复杂。肌肉注射Kiss1和Kiss2均提高了青春期前的欧洲海鲈血清LH水平(Felip, 2009)；腹腔注射Kiss1而非Kiss2也提高了性成熟雌性金鱼血清LH水平 (Li, 2009)。但Kiss1和Kiss2均不影响金鱼垂体细胞LH分泌(Li, 2009)。相反，另有研究表明，Kiss1直接促进了金鱼垂体细胞LH分泌(Chang, 2012; Yang, 2010)。最近研究报道，Kiss2而非Kiss1促进了欧洲海鲈(Espigares, 2015b)和条纹鲈(Zmora, 2015)垂体细胞LH分泌。Kiss1和Kiss2对杂交条纹鲈LH分泌的调控作用与生殖周期相关。在青春前期，肌肉注射Kiss2而非Kiss1增加了血清中LH水平；在性腺复苏期，Kiss1和Kiss2均增加了血清中LH水平(Zmora, 2012)。关于FSH分泌调控，肌肉注射Kiss2提高了青春期前的欧洲海鲈血清FSH水平，但是，Kiss1无影响(Felip, 2009)。同样，Kiss2而非Kiss1促进了欧洲海鲈垂体细胞FSH分泌(Espigares, 2015b)。此外，Kiss1和Kiss2均促进了条纹鲈垂体细胞FSH分泌(Zmora, 2015)。而长期埋植Kiss2显著性地降低了条纹鲈血清FSH水平(Zmora, 2014)。在鱼类中，关于Kiss对GH分泌的影响仅见于金鱼，Kiss1促进了金鱼垂体细胞GH分泌(Chang, 2012; Yang, 2010)。综上所述，Kiss对垂体激素分泌的调控作用因物种、生殖周期和注射途径而异，甚至在同一物种的不同生殖周期Kiss1和Kiss2可能发挥了不同的作用。

4.3 Kisspeptin对性腺发育及类固醇激素分泌的影响

5 鱼类kisspeptin的信号转导机制

在哺乳类中，Kiss能够激活多种细胞内信号通路，例如PLC/IP3/PKC、MAPK以及Ca2+通路等(Castano, 2009; Pasquier, 2014)，而非哺乳类中有关Kiss信号转导机制的研究相对较少。在两栖类中，Moon等(2009)通过CRE-luc(对应AC/PKA通路)和SRE-luc(对应PLC/PKC通路)报告系统表明，Kiss能够激活转染了牛蛙() Kiss2R的非洲绿猴肾纤维细胞系(CV-1 cells)中SRE-luc的活性，但对CRE-luc活性无影响。此外，PKC抑制剂GF109203X预处理CV-1细胞系显著性地降低了Kiss诱导的SRE-luc的活性，而Rho激酶抑制剂Y-27632预处理CV-1细胞系部分阻断了Kiss诱导的SRE-luc的活性，上述结果显示，牛蛙Kiss2R可能主要与PKC通路偶联，部分与Rho激酶通路偶联(Moon, 2009)。同样，非洲爪蟾() 3种KissR也都与PKC通路偶联(Lee, 2009)。

6 鱼类kiss/kissr系统的表达调控研究

6.1 性类固醇激素及甲状腺激素对kiss/kissr系统的调控作用

Kiss/KissR系统也介导了睾酮(Testosterone, T)对生殖轴的反馈调控。一方面，用睾酮处理卵巢切除后的雌性条纹鲈，降低了其脑中、及的表达水平(Klenke, 2011)。另一方面，用睾酮处理精巢切除后的雄性欧洲海鲈，降低了其下丘脑中的表达水平，却不影响、及的表达水平(Alvarado, 2016)。然而，睾酮促进了雄性欧洲海鲈垂体细胞及的表达水平，对的表达水平无影响(Espigares, 2015b)。此外，睾酮也不影响半滑舌鳎下丘脑中及的表达水平(Wang, 2017b)。目前，关于甲状腺激素(Thyroid hormone)对鱼类系统的调控作用仅见于罗非鱼。腹腔注射甲状腺激素，显著地增加了罗非鱼脑的表达水平，但由于甲状腺激素受体不在Kiss2神经元中表达，这表明甲状腺激素是以间接的方式影响的表达(Ogawa, 2013)。综上所述，性类固醇激素及甲状腺激素通过影响系统的表达水平进而影响鱼类生殖调控。

6.2 Kisspeptin等神经肽对kiss/kissr系统的调控作用

促性腺激素抑制激素(Gonadotropin-inhibitory hormone, GnIH)是迄今为止在脊椎动物中鉴定出的唯一具有抑制生殖功能的下丘脑神经肽，通过其受体GnIHR (之前被称作GPR147)介导作用于脑-垂体-性腺轴进而影响动物生殖调控(Tsutsui, 2010; Ubuka, 2016; Wang, 2018)。目前，从鱼类到哺乳类都鉴定出了的同源基因，并且每种鱼类基因编码有2种或者3种成熟多肽，即GnIH-1、GnIH-2及GnIH-3 (Ogawa, 2014; Tsutsui, 2010; Ubuka, 2016; 王滨等, 2016)。GnIH对的表达调控也有少数报道。在半滑舌鳎中，GnIH-1和GnIH-2均不影响下丘脑中的表达水平(刘权等, 2017)。腹腔注射斜带石斑鱼3种GnIH多肽也不影响其下丘脑的表达水平(Wang, 2015)。此外，哺乳类GnIH同源多肽RFRP3也不影响大鼠表达水平(Johnson, 2008)。尽管侧脑室注射欧洲海鲈GnIH-1不影响其脑、及的表达水平，但是，GnIH-2均降低了及的表达水平，这说明在欧洲海鲈中，GnIH-2主要发挥了生殖调控的抑制作用(Paullada-Salmeron, 2016)。

6.3 光照对kiss/kissr系统的调控作用

光照是影响鱼类及其他脊椎动物生殖调控的一个重要环境因子，其作用主要由松果体夜间分泌的褪黑激素介导(Kitahashi, 2013)。目前，关于光照对鱼类系统的表达调控研究相对较少且存在争议。如持续性光照降低了罗非鱼脑的表达水平，表明光照能够以直接或者间接的方式影响的表达(Martinez-Chavez, 2008)。同样，持续性光照导致欧洲海鲈前中脑及的表达量不再随季节变化而变化(Espigares, 2017)。长光照(繁殖状态)条件下，青鳉下丘脑核腹侧结节中Kiss1神经元的数量显著性高于短光照(非繁殖状态) (Kanda, 2008)。而在模拟自然光照(促进生殖)条件和持续性光照(抑制生殖)条件下，大西洋鳕()脑及的表达量无显著性差异，这说明光照不影响大西洋鳕基因表达(Cowan, 2012)。特别是褪黑激素促进了斑马鱼脑及的表达水平(Carnevali, 2011)，却抑制了欧洲海鲈脑及的表达水平(Alvarado, 2015)。综上所示，Kiss/KissR系统可能介导了光照(及褪黑激素)对鱼类生殖调控过程，然而具体作用机制因物种而异，需要进一步深入研究。

6.4 温度对kiss/kissr系统的调控作用

对变温动物而言，温度是影响其生殖调控的一个重要环境因子。水温升高或者降低均能抑制鱼类生殖，但其分子机制仍不清楚。Kiss作为鱼类生殖调控的一个重要因子，温度对基因的表达调控作用也有了初步研究。斑马鱼最适繁殖温度为26℃~ 28℃，低于20℃或者高于30℃均能降低其繁殖能力(Shahjahan, 2013)。将斑马鱼置于低温(15℃)、正常温度(27℃)和高温(35℃) 7 d后研究发现，低温组斑马鱼全脑的表达量显著性增加，高温组的表达量与正常组相比无显著性差异；而低温组和高温组斑马鱼全脑的表达量较正常组均显著性降低(Shahjahan, 2013)。此外，低温也增加了斑马鱼松果体等部分脑区的表达量，然而，低温和高温均降低了斑马鱼下丘脑等部分脑区的表达量(Shahjahan, 2013)。上述结果表明，温度调控斑马鱼及表达的作用机制是不同的，低温激活了系统，而低温和高温均抑制了系统，说明和系统可能参与了斑马鱼不同的生理功能(Shahjahan, 2013)。

同样，将星点东方鲀置于低温(14℃)、正常温度(21℃)和高温(28℃) 7 d后研究发现，低温组和高温组性腺指数GSI显著性降低；低温组和高温组脑/表达量也显著性降低；与此同时，低温和高温组均抑制了脑、垂体及的表达水平(Shahjahan, 2016)。上述结果表明，低温和高温组通过抑制系统，进而阻断星点东方鲀生殖。银汉鱼()的性别决定、分化与温度紧密相关，低温(17℃~19℃)导致100%全雌，高温(29℃)导致100%全雄，而24℃~25°℃导致雌雄比例各半(Tovar Bohorquez, 2017)。在高温条件下，银汉仔鱼整个脑部的表达量在孵化后4周显著性增加；而在低温条件下，脑部的表达量在孵化后8周内保持不变，这表明Kiss2可能在雄性发育过程中性别决定阶段发挥了重要作用(Tovar Bohorquez, 2017)。

6.5 饥饿对kiss/kissr系统的调控作用

营养状况也会影响动物生殖活动。目前，关于Kiss介导的能量平衡与生殖之间关系的研究较少。在哺乳类中，饥饿导致小鼠()下丘脑及表达量降低(Luque, 2007)。同样，饥饿也降低了大鼠下丘脑的表达量，却增加了的表达量(Castellano, 2005)。在鱼类中，饥饿15 d导致塞内加尔鳎体重减少，却增加了下丘脑及的表达水平，但对胃中及的表达水平无影响(Mechaly, 2011)。同样，饥饿也增加了欧洲海鲈下丘脑、、及的表达水平(Escobar, 2016)。综上所述，饥饿对哺乳类和鱼类系统的不同调控作用表明，该系统可能在哺乳类和鱼类能量平衡过程中起着相反的作用。此外，Kiss/KissR系统是否参与了鱼类摄食调控仍不得而知，需要进一步深入研究。

7 小结

Kiss是一种多功能的神经肽，它在下丘脑-垂体-性腺轴多个水平参与了哺乳动物生殖调控。目前，尽管已在多种鱼类中鉴定出了Kiss/KissR系统，但其在鱼类生殖调控中的精确作用需要进一步研究；Kiss调控垂体激素分泌及其基因表达的信号转导机制网络需要进一步完善；Kiss是否参与鱼类摄食调控及其作用机制尚未阐明；Kiss与其他因子(例如GnIH、GnRH等)之间如何互作，在生殖轴各个水平将多种信号整合进而调控生殖等生理过程仍不清楚，只有阐明上述机制才能更好地了解Kiss参与鱼类生殖、生长及代谢的协调过程。该综述总结了鱼类Kiss及其受体的研究进展，并对Kiss的生理学功能、信号转导机制以及表达调控研究进行概括讨论，增加了人们对Kiss/KissR系统参与鱼类生殖调控机制的认识，为后续研究提供参考。

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(编辑 陈严)

Regulatory Mechanisms of Kisspeptin on the Reproductive Axis in Fish

WANG Bin1,2, LIU Xuezhou1,2①, XU Yongjiang1,2, SHI Bao1,2, LIU Quan1,3

(1. Key Laboratory of Sustainable Development of Marine Fisheries, Ministry of Agriculture and Rural Affairs, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Qingdao 266071; 2 Laboratory for Marine Fisheries and Food Production Processes, Pilot National Laboratory for Marine Science and Technology (Qingdao), Qingdao 266071,; 3 College of Fisheries and Life Science, Shanghai Ocean University, Shanghai 201306)

Kisspeptin (Kiss or Kp), a novel physiologically active peptide encoded by thegene, activates its cognate receptor KissR (also known as GPR54) in various target tissues to exert disparate functions, including inhibition of tumor metastasis and control of reproductive function. Thegene was originally isolated from human melanoma and breast cancer cells, and kisspeptin was initially called metastin in consideration of its suppressive effects on tumor growth and metastasis. With the exception of the platypus, a mammalian monotreme, which has two forms of kisspeptin genes (and), there is only one ligand,and its receptor,in placental mammals. However, this situation is different and even complex in non-mammalian species. Three/genes were described in amphibians, while searches in the chicken genome databases failed to identify these paralogous genes. To date, multiple forms of/genes have been identified in many teleosts, including Cypriniformes, Perciforms, Pleuronectiforms, Tetraodontiforms, Beloniforms, Scorpaeniformes, Salmoniformes and Gadiformes. A dual kisspeptin system,11and, have been detected in medaka, zebrafish, goldfish, chub mackerel, striped bass, and European sea bass, while onlywas identified in orange-spotted grouper, grass puffer,,,, and half-smooth tongue sole. In addition, the physiological relevance and functions of the Kiss/KissR system for the neuroendocrine regulation of reproduction remains to be established in fish. It should be noted that the mechanisms underlying the actions of Kiss on the hypothalamo-pituitary-gonadal (HPG) axis are still far from being fully understood. Given the multiple forms ofandgenes obtained in teleosts, the regulation of fish reproduction by the Kiss system is even complex. This review briefly summarized the progress of research on Kiss and its receptors, with special emphasis on the physiological functions of Kiss in fish, the signaling transduction pathways as well as the regulation ofgene expression. We hope that this review will contribute to future studies.

Fish; Kisspeptin; kisspeptin receptor; Reproduction; Signal transduction; Regulation of gene expression

LIU Xuezhou, E-mail: liuxz@ysfri.ac.cn

10.19663/j.issn2095-9869.20170424001

S917; Q575; Q492

A

2095-9869(2018)04-0173-12

* 中国水产科学研究院基本科研业务费(2017HY-XKQ01; 2017GH05; 2018GH17)、中国水产科学研究院黄海水产研究所基本科研业务费(20603022016018)、国家自然科学基金(31602133；31502145)、山东省自然科学基金(ZR2016CB02)和国家海水鱼类产业技术体系(CARS-47)共同资助[This work was supported by Grants from the Central Public-Interest Scientific InstitutionBasal Research Fund, CAFS (2017HY-XKQ01; 2017GH05; 2018GH17), Special Scientific Research Funds for Central Non-Profit Institutes, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences (20603022016018), the National Natural Science Foundation of China (31602133; 31502145), the Natural Science Foundation of Shandong Province (ZR2016CB02), and China Agriculture Research System (CARS-47)]. 王 滨，E-mail: wangbin@ysfri.ac.cn；柳学周，E-mail: liuxz@ysfri.ac.cn

# 共同第一作者

柳学周，研究员，E-mail: liuxz@ysfri.ac.cn

2017-04-24,

2017-05-18

王滨, 柳学周, 徐永江, 史宝, 刘权. Kisspeptin对鱼类生殖轴的调控机制研究. 渔业科学进展, 2018, 39(4): 173–184

Wang B, Liu XZ, Xu YJ, Shi B, Liu Q. Regulatory mechanisms of Kisspeptin on the reproductive axis in fish. Progress in Fishery Sciences, 2018, 39(4): 173–184