Application Progress of Selenium and Selenoprotein in Aquaculture

WANG Lianshun; LI Zimeng; JIN Xiaomin; HUANG Jiaqiang

doi:10.3969/j.issn.1006-267x.2023.01.009

Chinese Journal of Animal Nutrition >

2023 , Vol. 35 >Issue 1: 77 - 85

DOI: https://doi.org/10.3969/j.issn.1006-267x.2023.01.009

REVIEW

Application Progress of Selenium and Selenoprotein in Aquaculture

WANG Lianshun ^,¹ ,
LI Zimeng ² ,
JIN Xiaomin ² ,
HUANG Jiaqiang ^,¹^,³^,^*

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¹ College of Fisheries and Life, Dalian Ocean University, Dalian 116023, China
² College of Marine Resources and Environment, Hebei Normal University of Science and Technology, Qinhuangdao 066600, China
³ Department of Nutrition and Health, China Agricultural University, Beijing 100083, China

^* associate professor, E-mail: JQHUANG@cau.edu.cn

Received date: 2022-05-26

Online published: 2023-01-11

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Abstract

With the rapid development of aquaculture industry, the intensive scale of aquaculture is expanded and the density of aquaculture animals is increased, which is extremely easy to cause oxidative stress reaction, which forces aquaculture practitioners to find non-toxic, harmless and efficient feed additives to improve the anti-stress ability of aquatic animals. Selenium (SE) is an essential trace element for human and animal growth and breeding, its biological function is mainly expressed through selenoprotein, and adding a certain amount of selenium to feed can enhance the resistance of aquatic animals. This paper summarizes the types of selenium and selenoprotein and the physiological mechanism of selenium on aquatic animals. At the same time, it summarizes the application of selenium in aquaculture, in order to summarize the role of different selenium sources as aquatic feed additives in improving the growth, immunity and meat quality of economic aquatic animals, so as to provide scientific reference for their efficient application in aquaculture.

Key words： selenium; selenoprotein; aquatic animals; mechanism of action

Cite this article

WANG Lianshun , LI Zimeng , JIN Xiaomin , HUANG Jiaqiang . Application Progress of Selenium and Selenoprotein in Aquaculture[J]. Chinese Journal of Animal Nutrition, 2023 , 35(1) : 77 -85 . DOI: 10.3969/j.issn.1006-267x.2023.01.009

硒(selenium,Se)是最早由瑞典科学家Berzelius于1817年发现的一种非金属元素^[1]。在20世纪40年代以前,人们普遍认为硒是一种有毒物质,会威胁人类健康。直到1957年,硒被证实是动物机体必需的微量元素;1973年,世界卫生组织(WHO)正式宣布了硒是动物机体及人类必需的微量元素^[2-3]。随着人类对硒的认识越来越深入,硒在生理机能和应用方面的研究也越来越多元化^[4]。已有研究表明,硒在动物的生长、抗氧化和免疫系统中起着重要作用,其作为饲料添加剂提供给水产动物时,可以极大地改善水产动物的健康状况^[5]。

水产品以其高蛋白质、低脂肪、营养物质均衡及味道鲜美等诸多优势受到广大消费者的青睐。随着我国经济发展与提质增效需求的不断上升,水产养殖规模也在不断扩大。然而,集约化的水产养殖模式导致水产动物抗应激能力逐渐下降,水产病害种类也逐渐增加,严重影响了水产动物生产性能与食品安全,经济效益受到重创的报道也时有发生^[6⇓-8]。因此,寻找无毒无害且高效的饲料添加剂,成为了当前水产饲料行业的研究热点。硒是水产动物生长所必需的微量元素,在集约化水产养殖系统中,养殖鱼类主要从饲料中获取机体所需的硒^[9]。已有研究表明,硒缺乏会导致水产动物肌肉张力降低,生长性能下降,防御机制减弱,进而出现氧化损伤、组织病变,并出现高死亡率,而硒过量会通过各种有害症状对动物产生毒性,并可能导致饲料效率降低、机体生殖衰竭、组织遭到破坏,甚至出现器官畸形^[5,9-10]。可见,硒过量或缺乏都会对鱼体造成不良影响。本文针对不同硒源对水产动物的生理作用,以及硒在水产养殖中的应用效果进行全面综述,以期为硒在水产养殖中的合理应用提供理论基础。

1 硒的存在形式

在自然界中,硒主要以无机硒和有机硒2种形式存在。无机硒一般以硒酸盐和亚硒酸盐的形式存在,早期曾是水产动物饲料中硒源补充剂的主要形式^[11]。由于其毒性较大且不易被吸收利用,故研究重点逐渐向有机硒转变。有机硒主要以硒代蛋氨酸、硒代半胱氨酸和硒代胱氨酸等形式存在,其吸收率高、生物活性强,且毒性较低^[12]。纳米硒是一种新形态的硒,其相比于无机硒和有机硒具有更高的生物利用度、更低的毒性和更强的生物活性^[13],在水产饲料添加剂方面的应用具有很大的潜力。不管是无机硒、有机硒还是纳米硒,其作为饲料添加剂被动物摄入到体内,都会先转化为机体需要的含硒氨基酸或硒蛋白,再发挥生理作用。

2 硒蛋白的种类与功能

目前,硒蛋白组已经在多种生物中进行了分析,在真核生物中已鉴定出29种硒蛋白^[14],并且在人类中已鉴定出25种硒蛋白^[15]。相对于哺乳动物而言,鱼类硒蛋白在合成过程中硒代半胱氨酸的合成效率更高,且鱼类具有更多的硒蛋白^[16-17]。目前已知的鱼类硒蛋白的种类、分布及功能如表1所示。

表1 鱼类硒蛋白的种类、分布及功能

Table 1 Type, distribution and function of selenoproteins of fish species

硒蛋白种类 Selenoprotein types	鱼类 Fish species	分布 Distribution	功能 Function	参考文献 Reference
GPX1	金头鲷、南方蓝鳍金枪鱼、日本鳗鲡	肝脏、脾脏;幽门盲肠道、肝脏; 鳃、皮肤、肌肉、肝脏、脾脏、肾脏、肠道	中和活性氧和改善氧化应激	[18 ⇓-20]
GPX2	尼罗罗非鱼	鳃、肝脏、肾脏、肌肉	抗氧化,与成熟精子的结构和功能有关	[15]
GPX3	黄颡鱼	肝脏、脑、背肌、肠系膜脂肪、前肠、头肾、鳃、心脏、脾脏	抗氧化	[21]
GPX4	金头鲷、南方蓝鳍金枪鱼、日本鳗鲡、黄颡鱼	肝脏、大脑;幽门盲肠、大脑、心脏、肝脏;幽门盲肠、肝脏;鳃、皮肤、肌肉、肝脏、脾脏、肾脏、肠道	影响大脑的发育和功能,维持机体正常嗅觉信号转导,抗氧化	[18 ⇓⇓⇓-22]
DIO1	牙鲆	肠道、鳃、肝脏、肌肉、鳍、眼睛	影响牙鲆变态过程中眼睛的移动	[23]
DIO2	牙鲆	脊髓、肌肉、肠道、胸鳍、眼睛周围、冠状幼鳍	影响牙鲆变态过程中眼睛的移动	[23]
DIO3	牙鲆	鳃、上下颌、眼睛周围、尾鳍、背鳍、臀鳍、腹鳍、胸鳍、肌肉、支鳍骨、体表皮肤、鼻孔、肠道、肌肉	影响牙鲆变态过程中眼睛的移动和侧线发育	[23]
TXNRD1	松江鲈	血液、心脏、肝脏、鳃、胃、肠道、皮肤、肌肉、肾脏、脾脏、脑、卵	参与机体基础代谢和氧化应激过程中的免疫反应	[24]
TXNRD2	黄颡鱼	背肌、鳃、大脑、心脏、脾脏、头肾、肠系膜脂肪、前肠	还原硫氧还蛋白, 保护细胞免受氧化	[21]
TXNRD3	黄颡鱼	肠系膜脂肪、脑、头肾、鳃、背肌、前肠、心脏、肝脏、脾脏	还原硫氧还蛋白, 保护细胞免受氧化	[21]
SELENOJ	斑马鱼	肝脏、肾脏、脾脏、心脏、肌肉、卵巢、肠道、眼睛、睾丸、大脑	抗氧化,与二磷酸腺苷核糖基化有关	[25]
SELENOP	黄颡鱼	心脏、大脑、脾脏、肾脏、肠道、肌肉、肠系膜脂肪、鳃、睾丸、卵巢	转运机体正常代谢所需蛋白	[21]
SELENOF	黄颡鱼、草鱼	肝脏;肝胰腺、肌肉	诱导糖原分解和脂质代谢	[26-27]
SELENOM	黄颡鱼	心脏、脾脏、肠道	调节细胞内内质网应激和钙稳态调节	[21]
硒蛋白种类 Selenoprotein types	鱼类 Fish species	分布 Distribution	功能 Function	参考文献 Reference
SELENOS	尼罗罗非鱼、草鱼	大脑、脾脏、肾脏;鳃、肝脏、肾脏、肌肉	调节细胞内内质网应激和钙稳态调节	[15,27]
SELENON	黄颡鱼	心脏、大脑、肾脏、肠道	调节细胞内内质网应激和钙稳态调节	[26]
SELENOT	黄颡鱼	心脏、大脑、脾脏、肾脏、肠道、肌肉、肠系膜脂肪、鳃、睾丸、卵巢	抗氧化应激并确保内质网稳态	[21]
SELENOE	草鱼	肝胰腺、脂肪组织	参与脂质代谢	[27]
SELENOL	草鱼、尼罗罗非鱼	肝胰腺、肌肉;鳃、肝脏、肾脏、肌肉	参与脂质代谢、氧化还原	[15,27]
SELENOU	草鱼	肝胰腺、肌肉、脂肪组织	参与脂质代谢	[27]
SELENOK	尼罗罗非鱼	鳃、肝脏、肾脏、肌肉	调节鳃中钙离子代谢	[15]
SEPHS2	尼罗罗非鱼	鳃、大脑	参与氧化应激防御	[15]
SCLY	尼罗罗非鱼	鳃、垂体、肾脏	维持矿物质平衡,平衡肾脏中的血浆稳态和肝脏中的血液载体分子平衡	[15]

GPX:谷胱甘肽过氧化物酶 glutathione peroxidase;DIO:碘甲状腺原氨酸脱碘酶 iodothyronine deiodinase;TXNRD:硫氧还蛋白还原酶 thioredoxin reductase;SELENOJ:硒蛋白J selenoprotein J;SELENOP:硒蛋白P selenoprotein P;SELENOM:硒蛋白M selenoprotein M;SELENOS:硒蛋白S selenoprotein S;SELENON:硒蛋白N selenoprotein N;SELENOT:硒蛋白T selenoprotein T;SELENOE:硒蛋白E selenoprotein E;SELENOL:硒蛋白L selenoprotein L;SELENOU:硒蛋白U selenoprotein U;SELENOK:硒蛋白K selenoprotein K;SEPHS2:硒磷酸合成酶 selenophosphate synthetase;SCLY:硒代半胱氨酸裂解酶 selenocysteine lyase。

3 硒对水产动物抗氧化、促生长和免疫调节的作用机制

3.1 抗氧化作用

在鱼类养殖过程中,由于受到拥挤胁迫、缺氧和营养不良等因素的影响,鱼体内会产生过量的氧自由基,从而引起细胞内的氧化-抗氧化系统动态平衡失调,进而诱导机体发生氧化应激^[28]。细胞内调节氧化还原的含硒酶有谷胱甘肽过氧化物酶(GPX)和硫氧还蛋白还原酶(TXNRD)等^[29]。硒是GPX的活性中心,GPX通过催化谷胱甘肽将过氧化氢和脂质过氧化物等有害物还原成水或无毒的羟基化合物,清除机体内的氧化自由基,保护生物膜和生物大分子免受氧化损伤^[30]。TXNRD可以催化还原性辅酶Ⅱ(NADPH)还原硫氧还蛋白(Trx)上的-S2,进而维持Trx的还原型,解除机体的氧化应激反应^[31]。

研究表明,硒通过清除氧化自由基和脂质过氧化物能缓解机体的氧化应激,减轻对细胞膜和线粒体DNA造成的损伤^[32⇓-34]。研究发现,不同浓度及不同形式的硒能够提高机体内GPX、超氧化物歧化酶(SOD)、过氧化氢酶(CAT)等抗氧化酶的活性,增强机体抗氧化能力,缓解因不良条件导致的氧化应激^{[34⇓⇓-37]}。

GPX4能够减少位于磷脂双分子层中的脂质氢过氧化物,在哺乳动物中,GPX4与成熟精子中线粒体包裹的相关结构蛋白有关^[38]。Porter等^[39]推测GPX4减少脂质氢过氧化物的能力表明它可能是大型海洋硬骨鱼(如蓝鳍金枪鱼)中的一种重要抗氧化酶,金枪鱼在身体组织中积累高水平ω-3长链多不饱和脂肪酸(LC-PUFA)。LC-PUFA对氧化损伤高度敏感,因此GPX4可能在减少细胞膜中的LC-PUFA氢过氧化物方面发挥作用,Bain等^[40]推测GPX4在雄性鱼类生育力中的作用可能与哺乳动物一致。研究发现,GPX1、TXNRD2、TXNRD3、硒蛋白P(SELENOP)^[21]、硒蛋白L(SELENOL)^[41]等都具有氧化还原功能。

3.2 促生长作用

鱼类的生长性能取决于几个因素,如日常管理、水质、疫苗接种和温度。除此之外,可以通过搭配营养均衡的水产饲料提高饲料的消化率,进而增强鱼类的生长性能^[42-43]。硒能通过调节某种生长激素——甲状腺激素(thyroxine),间接调控鱼类的生命代谢活动。硒以硒代半胱氨酸的形式存在于甲状腺素脱碘酶(DIO)的活性中心,通过此种方式参与甲状腺激素的代谢调节。DIO的作用是催化四碘甲状腺原氨酸(T₄)转变为三碘甲状腺原氨酸(T₃),进而调控甲状腺激素合成和降解,还可提高胰岛素水平,促进肌肉生长和新陈代谢,从而促进动物的生长发育^[5,44]。

饲料中不添加硒或添加量过少都不能满足鱼类生长需要,可能导致生长迟缓^[43]。研究表明,通过增加饲料中的亚硒酸盐水平,硒表现出胰岛素样作用,体内己糖激酶活性增加,促进了虹鳟(Oncorhynchus mykiss)的生长和新陈代谢^[44]。在草鱼(Grass carp)饲料中添加不同水平的亚硒酸钠,显著提高了草鱼幼鱼的体重,增加了蛋白质合成和血浆三碘甲状腺原氨酸水平,并促进了生长激素的分泌和鱼体生长^[45]。斑马鱼中硒蛋白N(SELENON)消耗导致机体紊乱和整体肌肉结构改变,而小鼠出现类似于SELENON导致的遗传性神经肌肉疾病症状,例如脊柱强直、脊柱后凸,这些结果表明SELENON可能对成年机体肌肉中肌肉祖细胞的增殖和维持至关重要^[46-47]。

3.3 免疫调节作用

鱼类的抗氧化防御系统可以抵抗应激,消除过量的活性氧并保护宿主,过量的活性氧可以杀伤病原体,但也会对正常的机体组织造成损害,并引起机体的过度免疫,进而增强机体的抗病性^[48]。硒以硒蛋白的形式参与机体抗氧化和免疫调节机制,硒蛋白K(SELENOK)和硒蛋白S(SELENOS)是与炎症和免疫相关的2种硒蛋白,并且在导致内质网应激的条件下发挥保护细胞的作用^[49],此外,GPX可以通过保护中性粒细胞免受超氧化物衍生的自由基的伤害,中性粒细胞产生自由基以杀死外来微生物^[50]。硒能促进吞噬细胞发挥吞噬和杀菌作用来增强机体的非特异性免疫功能,可以参与调节T淋巴细胞、B淋巴细胞、自然杀伤性细胞和中性粒细胞的功能,还能増强体液免疫,刺激免疫球蛋白的形成,提高血液中免疫球蛋白含量,从而增强机体的免疫能力^[49]。

在尼罗罗非鱼的研究中发现,硒显著上调了其肝脾组织中促炎因子——肿瘤坏死因子-α(TNF-α)和白细胞介素-1β(IL-1β)表达,其原因是硒作为免疫刺激物可以促进巨噬细胞分泌,增强免疫防御机制并抵抗病原入侵^[51]。饲料中添加硒后,草鱼幼鱼血清中谷草转氨酶(AST)和谷丙转氨酶(ALT)活性增加,体内免疫球蛋白含量增加,说明硒对草鱼幼鱼的肝脏等组织器官具有保护作用,可以增强鱼类的先天性免疫反应^[48]。

4 不同硒源在水产养殖中的应用研究

硒在水产养殖方面的应用,重点在于探讨不同硒源的饲料添加剂对水产动物的繁殖、生长发育、免疫力等方面的作用。

4.1 无机硒在水产养殖中的应用研究

无机硒主要以亚硒酸盐的形式被动扩散进入动物机体内,必须在肠道内先转化为硒化物才能被吸收^[43]。在水产动物饲料中添加无机硒,往往达不到预期的补硒效果,一般存在以下3个问题:第一,无机硒添加量低,在饲料中难以均匀分布,导致饲料质量参差不齐;第二,水产动物很难吸收利用无机硒,利用率较低,而过量添加会造成水产动物中毒的现象;第三,水产动物将未利用的无机硒排泄到水中,会污染周围的水体环境^[52-53]。

在进行亚硒酸钠和乳酸杆菌对虹鳟生长以及免疫反应研究时,结果表明同时使用亚硒酸钠和乳酸杆菌作为饲料添加剂可改善虹鳟鱼的生长和新陈代谢,并且乳酸杆菌对虹鳟鱼天然免疫应答的影响也在亚硒酸盐的作用下得到增强^[44]。姚朝瑞等^[54]通过研究硒对亚硝酸钠导致草鱼肝细胞氧化损伤的保护作用时,发现给草鱼肝细胞补充硒在一定程度上能缓解亚硝酸钠暴露导致的抗氧化系统失衡,抵抗亚硝酸钠暴露带来的氧化应激,降低细胞凋亡率,维持细胞正常生长。

4.2 有机硒在水产养殖中的应用研究

有机硒以主动转运的方式被机体各组织吸收利用,并参与硒蛋白的合成。许多研究表明,在几种水产动物中,有机硒比无机硒安全性能高,更具生物活性,并且具有更高的生物利用度^[13,44],因为无机硒必须先经过自然或人工合成为硒代半胱氨酸或硒代蛋氨酸等硒化合物才可以发挥作用,而有机硒被摄入机体后会直接裂解为硒化物,而硒代蛋氨酸可转化为硒代半胱氨酸^[55]。

Bakhshalinejad等^[56]在肉鸡饲粮中添加亚硒酸钠和2种有机硒富硒酵母、硒蛋氨酸进行喂养,发现与添加无机硒相比,添加有机硒能显著提高肉鸡的平均日增重(ADG)。在研究不同硒源对鲤鱼(Cyprinus arpio)的生长、肌肉硒浓度、肌肉邻近成分、血液酶活性和抗氧化状态的影响时,发现饲喂硒代蛋氨酸较亚硒酸钠的鱼增重更多,表明在基础饲料中添加纳米硒和硒代蛋氨酸可以提高鲤鱼的生长性能;硒在肝脏和肌肉中积累量更多,表明有机硒在肌肉中的生物利用度可能比无机硒更高,因为它与蛋白质结合导致鱼肌肉中的硒含量更高^[57]。在饲料中添加不同浓度的酵母硒分别对异育银鲫(Carassius auratus gibelio)、虹鳟有不同的生理影响,分别会提高水产动物的生长性能、非特异免疫能力以及改善抗氧化能力^[8,58]。

4.3 纳米硒在水产养殖中的应用研究

从代谢和生理功能方面来看,有机硒和无机硒的水溶性、渗透性和生物利用度较低,相比之下,纳米硒则表现出更高的生物利用度、更低的毒性和更强的生物活性^[13,59],部分原因是纳米颗粒微小、吸附性强,可以增加其与肠壁的接触面积,有利于胃肠道的消化与吸收,提高饲料利用率,提高动物生产效率,进而降低饵料系数,促进动物生长^[51],此外,在同样低剂量条件下,纳米硒显示出抑制肿瘤发展和免疫调节的作用^[8]。

在探究纳米硒对氟化钠诱导小鼠肝细胞组织细胞凋亡的影响时,纳米硒改善了氟化钠的毒性,通过提高肝细胞的抗氧化能力,减轻氟诱导的氧化应激,并通过清除氧自由基环节氟暴露引起的细胞损伤以及抑制细胞凋亡信号通路转导,调控相关基因的表达^[60]。Yu等^[37]在研究纳米硒对草鱼抗氧化能力的影响时发现,纳米硒通过促进肝胰腺中核因子E2相关因子2(Nrf2)的mRNA表达,显著提高了GPX1和CAT的mRNA表达,提高了草鱼的抗氧化能力。在研究添加不同浓度的纳米硒对尼罗罗非鱼的影响试验中发现,添加1 mg/kg的纳米硒显著改善了鱼类生长、血液学指标以及免疫参数,表明纳米硒可以改善罗非鱼的营养生理和免疫力^[61]。

5 小结与展望

综上所述,在水产动物饲料中添加不同硒源对水产动物的生长、抗氧化和免疫等方面具有重要的生理作用,而且其在不同水产动物体内的生物利用情况、浓度和形态变化不同,需要继续探寻更具生物利用价值和毒性更小的硒形态,探讨硒和其他生物、非生物因子的相互作用,不同营养元素对硒的作用效果。此外,硒缺乏或过量都会对水产动物造成不利影响,因此需明晰硒对具有重要功能的生物大分子结构与功能的影响,从而为其双重生物效应(有益与毒性)提供明确的机理性解释。继续研究水产动物对不同硒源的摄入量,确定其安全用量范围,重视硒在食物链中传递的规律及生物效应和人体健康风险评估,以探讨不同养殖生物中普遍添加硒后人类食用安全性的问题。根据已有研究发现,硒蛋白是硒作用于机体的关键,水生生物中的硒蛋白种类明显多于人类和哺乳动物,但目前在水产领域中有关单个硒蛋白的研究较少,应继续更新研究方法,利用蛋白质组学等新方法与硒研究相结合,以明确水产动物新型硒蛋白的重要数据。硒在水产养殖生产中具有广阔的应用前景,深入研究硒将为水产动物的健康养殖以及改善经济水产动物生长、免疫、肉质等方面提供科技支撑。

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