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        凡納濱對(duì)蝦鰓組織對(duì)高溫和氨氮脅迫的生理響應(yīng)

        2020-12-11 09:10:33熊大林段亞飛徐敬明詹愛軍陳成勛張家松
        關(guān)鍵詞:高溫

        熊大林 段亞飛 徐敬明 詹愛軍 陳成勛 張家松

        摘要:【目的】明確高溫和氨氮單因素脅迫及二者復(fù)合脅迫對(duì)凡納濱對(duì)蝦(Litopenaeus vannamei)鰓組織的毒性效應(yīng),為開展凡納濱對(duì)蝦健康養(yǎng)殖提供理論依據(jù)?!痉椒ā繉Ⅲw質(zhì)健康的凡納濱對(duì)蝦隨機(jī)分成4組,即對(duì)照(正常海水)組、高溫(33 ℃)脅迫組、氨氮(15 mg/L)脅迫組、高溫與氨氮復(fù)合脅迫(氨氮濃度15 mg/L,水溫33 ℃)組。脅迫72 h后,從每個(gè)平行中隨機(jī)挑取凡納濱對(duì)蝦鰓組織制作石蠟切片觀察其病理變化,并采用試劑盒測(cè)定鰓組織生理功能指標(biāo)的變化情況?!窘Y(jié)果】經(jīng)高溫和氨氮單因素脅迫或二者復(fù)合脅迫后均致使凡納濱對(duì)蝦鰓組織發(fā)生病理性結(jié)構(gòu)變化,表現(xiàn)為角皮層變薄或消失,上皮細(xì)胞逐漸分解,入鰓和出鰓血管變形或裂解,鰓絲連接松散,腔內(nèi)出現(xiàn)液泡等。凡納濱對(duì)蝦鰓組織超氧陰離子自由基(O[-2]·)和過氧化氫(H2O2)含量均在氨氮脅迫后顯著降低(P<0.05,下同),但經(jīng)高溫脅迫和高溫與氨氮復(fù)合脅迫后無顯著變化(P>0.05,下同);總抗氧化能力(T-AOC)和總超氧化物歧化酶(T-SOD)活性在高溫或氨氮脅迫后均顯著升高;谷氨酸脫氫酶(GDH)活性在高溫和氨氮單因素脅迫及二者復(fù)合脅迫后均顯著升高;谷氨酰胺酶(GLS)活性經(jīng)氨氮脅迫后顯著升高,經(jīng)高溫脅迫和高溫與氨氮復(fù)合脅迫后則顯著降低;己糖激酶(HK)活性在高溫脅迫后顯著降低,經(jīng)氨氮脅迫后顯著升高;乳酸脫氫酶(LDH)活性僅在氨氮脅迫后顯著升高;酸性磷酸酶(ACP)活性經(jīng)氨氮脅迫后顯著升高,而在高溫脅迫和高溫與氨氮復(fù)合脅迫后均顯著降低。【結(jié)論】經(jīng)高溫和氨氮單因素脅迫或二者復(fù)合脅迫后,凡納濱對(duì)蝦鰓組織出現(xiàn)明顯的病理性變化,其氧化應(yīng)激、氨氮代謝和能量代謝相關(guān)生理指標(biāo)均發(fā)生顯著變化。氨氮脅迫造成對(duì)蝦鰓組織生理紊亂,且高溫脅迫加重氨氮毒性,而降低對(duì)蝦鰓組織生理功能。因此,對(duì)蝦健康養(yǎng)殖特別是工廠化高密度養(yǎng)殖過程中因避免高溫環(huán)境下氨氮脅迫對(duì)凡納濱對(duì)蝦的毒性效應(yīng)。

        關(guān)鍵詞: 凡納濱對(duì)蝦;鰓組織;高溫;氨氮脅迫;復(fù)合脅迫;生理響應(yīng)

        中圖分類號(hào): S968.22? ? ? ? ? ? ? ? ? ? ? ? ? ? 文獻(xiàn)標(biāo)志碼: A 文章編號(hào):2095-1191(2020)09-2296-08

        Physiological responses in gills of Litopenaeus vannamei exposed to the combined stress of temperature and ammonia

        XIONG Da-lin1,2,? DUAN Ya-fei1,? XU Jing-ming3,? ZHAN Ai-jun4,

        CHEN Cheng-xun2,? ZHANG Jia-song1,3*

        (1South China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences/Key Lab of South China Sea Fishery Resources Exploitation & Utilization, Ministry of Agriculture and Rural Affairs/Guangdong Provincial Key Lab

        of Fishery Ecology Environment, Guangzhou? 510300, China; 2College of Fisheries, Tianjin Agricultural University, Tianjin? 300384, China; 3College of Landscape Architecture and Life Science, Chongqing University of Arts and Sciences, Chongqing? 402160, China; 4Shenzhen Academy of Inspection and Quarantine, Shenzhen,

        Guangdong? 518010, China)

        Abstract:【Objective】This study was to clarify the toxic effects of high temperature and ammonia single factor stress and their combined stress on the gills of Litopenaeus vannamei. Then, it could provide? theoretical basis for healthy bree-ding of L. vannamei. 【Method】The healthy L. vannamei were divided into four groups: the control(seawater) group, the high temperature(33 ℃) stress group, the ammonia(15 mg/L) stress group, and the combined of high temperature and ammonia stress(15 mg/L ammonia and 33 ℃ water) group. After stress for 72 h, the gill tissues of L. vannamei were randomly selected from each parallel to make paraffin sections to observe the pathological changes. And changes of physiolo-gical function indexes of the L. vannamei were determined by kit. 【Result】The single factor of high temperature and ammonia and the combined stress all caused pathological structural changes in the gill tissue, which was manifested as the thinning or disappearance of the cuticle layer, the gradual decomposition of epithelium, and the deformation and lysis of the afferent and efferent vessels of the gills. The gill filaments were loosely connected, and vacuoles appeared in the cavity. Oxidative damage indexes including superoxide anion radical(O[-2]·) and hydrogen peroxide(H2O2) levels were significantly decreased in gill tissue in the ammonia stress group(P<0.05, the same below), but no significant changes in the high temperature stress group and the combined stress group(P>0.05, the same below). The total antioxidant capacity(T-AOC) and total superoxide dismutase(T-SOD) activities were significantly increased in high temperature stress group and the ammonia stress group. The glutamate dehydrogenase(GDH) activity was significantly increased in all the stress groups, glutaminase(GLS) activity was significantly increased in the ammonia stress group, but significantly decreased in the high temperature stress group and the combined stress group. Hexokinase(HK) activity was significantly decreased in the high temperature stress group, but increased in the ammonia stress group;lactate dehydrogenase(LDH) activity was only significantly increased in the ammonia stress group;acid phosphatase(ACP) activity was significantly increased in the ammonia stress group, but significantly decreased in the high temperature stress group and the combined stress group. 【Conclusion】The results show that high temperature and ammonia single factor and their combined stress all have significant effects on the physiological functions of the gill of L. vannamei,the gill tissue shows obvious pathological changes, oxidative stress, and ammonia nitrogen metabolism and energy metabolism-related physiological indicators all change significantly. The ammonia stress can induce the physiological disorder of gill tissue, while the high temperature stress aggravates the ammonia toxicity and reduces the physiological function of the shrimp. Therefore, the process of healthy shrimp farming should pay attention to the toxic effect caused by ammonia stress in high temperature environment, especially in high-density factory farming.

        1. 3 組織形態(tài)觀察

        脅迫72 h后,從每個(gè)平行中隨機(jī)挑取3尾凡納濱對(duì)蝦的鰓組織置于4%多聚甲醛試劑中固定24 h。固定后采用不同濃度梯度的乙醇(70%、80%、90%和100%)進(jìn)行脫水,再經(jīng)二甲苯透明后進(jìn)行石蠟包埋,使用切片機(jī)制作4 μm的組織切片,最后以蘇木精和伊紅染料進(jìn)行染色,于光學(xué)顯微鏡下觀察鰓組織的病理變化,并拍照記錄。

        1. 4 酶活性測(cè)定

        將保存的鰓組織樣品置于冰上解凍,在預(yù)冷PBS中漂洗去除組織液,濾紙拭干;稱重后置于離心管中,加入9倍體積的PBS,使用組織勻漿機(jī)低溫研磨制成10%組織勻漿,然后在4 ℃下2000 r/min離心15 min,取上清液用于酶活性測(cè)定。酸性磷酸酶(ACP)、總超氧化物歧化酶(T-SOD)、谷氨酸脫氫酶(GDH)、谷氨酰胺酶(GLS)、己糖激酶(HK)、乳酸脫氫酶(LDH)活性和總抗氧化能力(T-AOC),以及超氧陰離子自由基(O[-2]·)、過氧化氫(H2O2)和總蛋白含量均采用南京建成生物工程研究所生產(chǎn)的試劑盒進(jìn)行測(cè)定,按照具體說明進(jìn)行操作。

        1. 5 統(tǒng)計(jì)分析

        試驗(yàn)數(shù)據(jù)采用SPSS 19.0進(jìn)行單因素方差分析(One-way ANOVA)和Duncan?s多重比較。

        2 結(jié)果與分析

        2. 1 高溫和氨氮脅迫對(duì)凡納濱對(duì)蝦鰓組織形態(tài)的影響

        與對(duì)照組(圖1-A)相比,高溫(圖1-B)和氨氮(圖1-C)單因素脅迫及二者復(fù)合脅迫(圖1-D)均造成凡納濱對(duì)蝦鰓組織出現(xiàn)一定程度的損傷,包括角皮層變薄或消失,上皮細(xì)胞逐漸分解,入鰓和出鰓血管變形或裂解,血細(xì)胞數(shù)量減少,隔膜變寬,鰓絲連接松散,腔內(nèi)出現(xiàn)液泡等。

        2. 2 高溫和氨氮脅迫對(duì)凡納濱對(duì)蝦鰓組織氧化應(yīng)激指標(biāo)的影響

        由圖2可知,與對(duì)照組相比,凡納濱對(duì)蝦鰓組織O[-2]·和H2O2含量均在氨氮脅迫后顯著降低(P<0.05,下同),在高溫脅迫組和高溫與氨氮復(fù)合脅迫組中無顯著變化(P>0.05,下同),但高溫脅迫組的O[-2]·含量顯著低于高溫與氨氮復(fù)合脅迫組。在抗氧化酶方面,凡納濱對(duì)蝦鰓組織T-AOC經(jīng)高溫脅迫、氨氮脅迫及高溫與氨氮復(fù)合脅迫后均顯著升高,其中又以氨氮脅迫組的T-AOC最高;T-SOD活性在高溫脅迫組和氨氮脅迫組均顯著升高,但在高溫與氨氮復(fù)合脅迫組無顯著變化。

        2. 3 高溫和氨氮脅迫對(duì)凡納濱對(duì)蝦鰓組織氨氮代謝酶活性的影響

        由圖3可看出,與對(duì)照組相比,凡納濱對(duì)蝦鰓組織GDH活性經(jīng)高溫脅迫、氨氮脅迫及高溫與氨氮復(fù)合脅迫后均顯著升高,且在3個(gè)脅迫組中無顯著差異。凡納濱對(duì)蝦鰓組織GLS活性經(jīng)氨氮脅迫后顯著升高,而經(jīng)高溫脅迫和高溫與氨氮復(fù)合脅迫后均顯著降低,且高溫脅迫組的GLS活性顯著低于高溫與氨氮復(fù)合脅迫組。

        2. 4 高溫和氨氮脅迫對(duì)凡納濱對(duì)蝦鰓組織能量代謝酶活性的影響

        由圖4可看出,與對(duì)照組相比,凡納濱對(duì)蝦鰓組織HK活性經(jīng)高溫脅迫后顯著降低,經(jīng)氨氮脅迫后顯著升高,而在高溫與氨氮復(fù)合脅迫組無顯著變化;凡納濱對(duì)蝦鰓組織LDH活性僅在氨氮脅迫后顯著升高,在高溫脅迫組和高溫與氨氮復(fù)合脅迫組中均無顯著變化;凡納濱對(duì)蝦鰓組織ACP活性經(jīng)氨氮脅迫后顯著升高,但經(jīng)高溫脅迫和高溫與氨氮復(fù)合脅迫后均顯著降低,且高溫脅迫組和高溫與氨氮復(fù)合脅迫組間無顯著差異。

        3 討論

        氨氮是對(duì)蝦養(yǎng)殖過程中重要的環(huán)境因子,對(duì)對(duì)蝦的生長、存活和生理功能均有重要影響。對(duì)蝦是排氨動(dòng)物,機(jī)體內(nèi)大部分氨氮經(jīng)代謝后以尿素和尿酸形式通過鰓組織排出體外(Weihrauch et al.,2004)。當(dāng)對(duì)蝦遭受氨氮脅迫時(shí),則通過鰓組織解毒代謝以降低氨氮對(duì)機(jī)體的損傷(Chen,2000)。目前,有關(guān)氨氮脅迫對(duì)甲殼動(dòng)物鰓組織生理功能影響的研究已有較多報(bào)道,如氨氮脅迫會(huì)造成Macrobrachium amazonicum鰓組織血細(xì)胞浸潤,細(xì)胞腫脹、壞死,層狀上皮增厚或裂解損傷,且損傷效應(yīng)隨氨氮濃度升高而加重,在80 mg/L總氨氮脅迫下甚至出現(xiàn)鰓組織水腫(Dutra et al.,2017);但高溫脅迫下氨氮對(duì)甲殼動(dòng)物鰓組織的毒性效應(yīng)至今尚未明確。為此,本研究探討了高溫和氨氮脅迫對(duì)凡納濱對(duì)蝦鰓組織生理功能的影響,結(jié)果顯示凡納濱對(duì)蝦鰓組織在高溫脅迫、氨氮脅迫及高溫與氨氮復(fù)合脅迫后均發(fā)生病理性結(jié)構(gòu)改變,表明高溫和氨氮單因素脅迫或復(fù)合脅迫均會(huì)損傷對(duì)蝦鰓組織的形態(tài)結(jié)構(gòu)。

        氧化應(yīng)激是動(dòng)物機(jī)體對(duì)環(huán)境脅迫的應(yīng)答機(jī)制之一。環(huán)境脅迫能誘導(dǎo)機(jī)體產(chǎn)生大量活性氧,包括 O[-2]·和H2O2(段亞飛等,2014)。對(duì)蝦在進(jìn)化過程中已形成較完善的抗氧化系統(tǒng),用于清除機(jī)體內(nèi)過量的活性氧,維護(hù)機(jī)體穩(wěn)態(tài)(Mathew et al.,2007)。其中,T-AOC可反映機(jī)體抗氧化系統(tǒng)的整體水平(Lewis et al.,1995);SOD是抗氧化系統(tǒng)重要的標(biāo)志酶,在機(jī)體清除活性氧過程中首先被誘導(dǎo),可將O[-2]·分解為H2O2和O2,并將H2O2傳遞給下游抗氧化酶(Mercier et al.,2006)。任海等(2014)研究表明,脊尾白蝦(Exopalaemon carinicauda)血淋巴和肝胰腺中的SOD活性隨氨氮脅迫濃度的增加和脅迫時(shí)間的延長,呈先上升后下降的變化趨勢(shì);朱孟凱和姚翠鸞(2015)研究表明,15 ℃低溫及31和35 ℃高溫脅迫均會(huì)導(dǎo)致凡納濱對(duì)蝦肝胰腺中的O[-2]·含量顯著增加;王蕓等(2017)研究發(fā)現(xiàn),中國對(duì)蝦(Fenneropenaeus chinensis)經(jīng)8 mg/L氨氮脅迫6 h后,其血淋巴的T-AOC顯著升高,但隨后逐漸降低。本研究結(jié)果表明,凡納濱對(duì)蝦鰓組織O[-2]·和H2O2含量經(jīng)氨氮脅迫后均顯著降低,但在高溫脅迫和高溫與氨氮復(fù)合脅后無顯著變化;相應(yīng)的氨氮脅迫組凡納濱對(duì)蝦鰓組織抗氧化酶T-AOC和T-SOD活性均最高,由此推測(cè)高溫和氨氮單因素脅迫或二者復(fù)合脅迫均會(huì)誘導(dǎo)凡納濱對(duì)蝦鰓組織產(chǎn)生氧化應(yīng)激反應(yīng),而抗氧化酶在機(jī)體清除活性氧過程中發(fā)揮了重要作用。

        谷氨酸代謝是甲殼動(dòng)物機(jī)體應(yīng)答氨氮脅迫的重要生理調(diào)節(jié)途徑(Cheng and Chen,2001)。GDH和GLS是谷氨酸生成或分解的關(guān)鍵調(diào)節(jié)酶,在甲殼動(dòng)物氨氮代謝過程中扮演重要角色。其中,GLS可催化機(jī)體多余的氨和谷氨酸生成谷氨酰胺,而GDH可催化谷氨酸發(fā)生氧化脫氨基反應(yīng),從而有效完成機(jī)體氨氮解毒代謝(王鏡巖等,2002;Hong et al.,2007)。劉勝男等(2015)、周發(fā)林等(2016)研究發(fā)現(xiàn),在氨氮脅迫下,三疣梭子蟹(Portunus trituberculatus)和斑節(jié)對(duì)蝦(Penaeus monodon)的鰓組織中GDH基因表達(dá)水平及其酶活性均顯著上調(diào);Zhang等(2019)研究表明,河蜆(Corbicula fluminea)經(jīng)氨氮脅迫后其消化腺中的GLS基因表達(dá)水平顯著上調(diào)。在本研究中,凡納濱對(duì)蝦鰓組織GDH活性經(jīng)高溫脅迫、氨氮脅迫及高溫與氨氮復(fù)合脅迫后均顯著升高,而GLS活性經(jīng)高溫脅迫和高溫與氨氮復(fù)合脅迫后顯著降低,推測(cè)是高溫和氨氮單因素脅迫或二者復(fù)合脅迫均對(duì)凡納濱對(duì)蝦鰓組織氨氮解毒代謝產(chǎn)生顯著影響,尤其是高溫脅迫抑制了氨氮代謝能力。

        動(dòng)物機(jī)體通過改變自身代謝以適應(yīng)逆境環(huán)境。HK是糖酵解過程的重要調(diào)節(jié)酶,其活性變化可反映機(jī)體對(duì)葡萄糖的利用情況(田相利,2001);LDH是無氧糖酵解和糖異生過程的調(diào)節(jié)酶,可催化乳酸轉(zhuǎn)變?yōu)楸幔瑸闄C(jī)體供能(Viru,1994)。郭彪等(2008)研究表明,水溫由17 ℃突增至27 ℃后,凡納濱對(duì)蝦肝胰臟中的HK活性也隨脅迫時(shí)間的延長呈先升高后降低的變化趨勢(shì),于脅迫3 h時(shí)達(dá)最高值,是水溫突變前的2.54倍。Shan等(2019)研究證實(shí),凡納濱對(duì)蝦在1.61 mg/L氨氮濃度脅迫下,其血淋巴HK活性隨脅迫時(shí)間的延長呈先升高后下降的變化趨勢(shì),而LDH活性呈逐漸升高趨勢(shì)。本研究結(jié)果表明,凡納濱對(duì)蝦鰓組織HK和LDH活性經(jīng)氨氮脅迫后均顯著升高,而在高溫與氨氮復(fù)合脅迫后無顯著變化;此外,HK活性在高溫脅迫后顯著降低,由此推斷氨氮脅迫誘導(dǎo)了凡納濱對(duì)蝦鰓組織能量代謝,高溫脅迫降低了對(duì)蝦鰓組織能量代謝,而高溫與氨氮復(fù)合脅迫對(duì)能量代謝過程無明顯影響。ACP是巨噬細(xì)胞內(nèi)重要的磷酸單酯水解酶類,在對(duì)蝦免疫反應(yīng)過程中發(fā)揮重要作用(陳清西等,1996)。艾春香和曾媛媛(2011)研究表明,氨氮脅迫導(dǎo)致擬穴青蟹(Scylla paramamosain)鰓組織ACP活性顯著降低。但本研究發(fā)現(xiàn)凡納濱對(duì)蝦鰓組織ACP活性在氨氮脅迫后顯著升高,而在高溫脅迫和高溫與氨氮復(fù)合脅迫后均顯著降低,即氨氮脅迫誘導(dǎo)了凡納濱對(duì)蝦鰓組織免疫反應(yīng),但與高溫復(fù)合脅迫即抑制機(jī)體免疫反應(yīng)。

        4 結(jié)論

        經(jīng)高溫和氨氮單因素脅迫或二者復(fù)合脅迫后,凡納濱對(duì)蝦鰓組織出現(xiàn)明顯的病理性變化,其氧化應(yīng)激、氨氮代謝和能量代謝相關(guān)生理指標(biāo)均發(fā)生顯著變化。氨氮脅迫造成對(duì)蝦鰓組織生理紊亂,且高溫脅迫加重氨氮毒性,降低對(duì)蝦鰓組織生理功能。因此,對(duì)蝦健康養(yǎng)殖特別是工廠化高密度養(yǎng)殖過程中因避免高溫環(huán)境下氨氮脅迫對(duì)凡納濱對(duì)蝦的毒性效應(yīng)。

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