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      金邊鯉皮膚黑色素合成及轉(zhuǎn)運(yùn)相關(guān)基因表達(dá)分析

      2020-04-14 04:59:03韋玲靜呂業(yè)堅(jiān)鄒輝文衍紅甘寶江張桂姣張盛嚴(yán)雪瑜葉香塵
      關(guān)鍵詞:差異表達(dá)基因黑色素

      韋玲靜 呂業(yè)堅(jiān) 鄒輝 文衍紅 甘寶江 張桂姣 張盛 嚴(yán)雪瑜 葉香塵

      摘要:【目的】研究影響金邊鯉皮膚色素沉著的關(guān)鍵基因及信號(hào)通路,揭示其皮膚顏色差異形成的原因,為金邊鯉良種培育提供候選基因信息。【方法】基于突變型金邊鯉黃色皮膚(Y組)和黑色皮膚(B組)及野生型金邊鯉黑色皮膚(W組)的轉(zhuǎn)錄組測(cè)序數(shù)據(jù),篩選分析獲得部分共有差異表達(dá)基因,利用實(shí)時(shí)熒光定量PCR檢測(cè)差異表達(dá)基因和黑色素合成相關(guān)基因的表達(dá)情況,并分析各基因間的相關(guān)性。【結(jié)果】依據(jù)轉(zhuǎn)錄組數(shù)據(jù)篩選獲得ACT3、MYH2、TPM1、TPM2和TPM3共5個(gè)來源于心肌腎上腺素信號(hào)通路及與細(xì)胞動(dòng)力相關(guān)的基因;除ACT3基因外,其余4個(gè)基因(MYH2、TPM1、TPM2和TPM3)在Y組皮膚中的相對(duì)表達(dá)量均顯著低于B組和W組(P<0.05,下同),與轉(zhuǎn)錄組分析的差異基因表達(dá)趨勢(shì)一致。MC-1R、TYR、TYRP1和TYRP2等4個(gè)黑色素合成相關(guān)基因在金邊鯉皮膚中的表達(dá)情況為:B組皮膚中MC-1R基因的相對(duì)表達(dá)量顯著高于Y組和W組;TYR、TYRP1和TYRP2基因在3組皮膚中的相對(duì)表達(dá)量排序均為W組>B組>Y組,其中W組的相對(duì)表達(dá)量顯著高于B組和Y組。相關(guān)性分析結(jié)果顯示,MC-1R和TYR基因與ACT3、MYH2、TPM1、TPM2和TPM3基因呈顯著或極顯著(P<0.01)正相關(guān),而TYRP1和TYRP2基因與這5個(gè)差異表達(dá)基因均無顯著相關(guān)性(P>0.05)。【結(jié)論】ACT3、MYH2、TPM1、TPM2和TPM3基因及其所在的心肌腎上腺素信號(hào)通路可能影響黑色素合成與運(yùn)動(dòng),且與黑色素合成相關(guān)基因存在相關(guān)性,可作為金邊鯉體色變異研究中的標(biāo)記基因。

      關(guān)鍵詞: 金邊鯉;黑色素;差異表達(dá)基因;黑色素合成相關(guān)基因;心肌腎上腺素通路;色素沉著

      中圖分類號(hào): S965.116? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?文獻(xiàn)標(biāo)志碼: A 文章編號(hào):2095-1191(2020)02-0453-08

      Skin melanin synthesis of Jinbian carp and expression analysis of transport related genes

      WEI Ling-jing1, LYU Ye-jian1, ZOU Hui1, WEN Yan-hong2, GAN Bao-jiang1,

      Zhang Gui-jiao3, ZHANG Sheng1, YAN Xue-yu1, YE Xiang-chen1*

      (1Aquatic Species Introduction and Breeding Center of Guangxi, Nanning? 530031, China; 2Liuzhou Aquaculture Technology Extending Station, Liuzhou,Guangxi? 545066, China; 3Aquaculture Technology Extension Station of Rongshui County, Rongshui, Guangxi? 545300, China)

      Abstract:【Objective】The key genes and signaling pathways affecting skin pigmentation of Jinbian carp were stu-died, to reveal the causes of skin color variation mechanism and provide candidate gene information for further cultivating new varieties of Jinbian carp. 【Method】Part of differentially co-expressed genes were screened based on the transcriptome sequencing data of yellow skin(group Y) and black skin(group B) in mutant type, and black skin (group W) in wild type of Jinbian carp, thenreal-time fluorescence quantitative PCR was used to detect the expressions of co-differentially expressed genes and genes related to melanin synthesis, and the correlations among the genes were analyzed. 【Result】A total of five genes,including ACT3, MYH2, TPM1, TPM2 and TPM3, were derived from the adrenergic signaling in cardiomyocytes pathways and associated with cell power were screened according to the transcriptome data.Apart from ACT3, the mRNA expression levels of the other fourgenes(MYH2, TPM1, TPM2 and TPM3) in group Y were all significantly lower than group B and group W(P<0.05, the same below), which was consistent with the trend of the result in transcriptome analysis. The mRNA expression levels of four genesrelating to melanin synthetic, including MC-1R, TYR, TYRP1 and TYRP2 were detected,which showed that the relative expression of MC-1R gene in group B was significantly higher than group Y and group W. The expression of TYR, TYRP1 and TYRP2 in the skin of the three groups showed an order of group W>group B>group Y, in which group W was significantly higher than group B and group Y. Correlation ana-lysis showed that the mRNA expressions levels of MC-1R and TYR gene were significantor extremely(P<0.01) positively correlated with ACT3, MYH2, TPM1, TPM2 and TPM3 genes, while TYRP1 and TYRP2 had no significant correlation with the fivegenes(P>0.05). 【conclusion】ACT3, MYH2, TPM1, TPM2 and TPM3 genes and their adrenergic signaling in cardiomyocytes pathways may affect the synthesis and movement of melanin, which may be related to the skin color variation of the Jinbian carp. These genes can be used as marker genes in the study of skin color variation of Jinbian carp.

      Key words: Jinbian carp; melanin; differentially expressed genes(DEGs); melanin synthesis related genes; adrener-gic signaling in cardiomyocytes pathways; pigmentation

      Foundation item: Special Project of National Staple Freshwater Fish Industry Technology System Construction(CARS-45)

      0 引言

      【研究意義】色素是由色素細(xì)胞合成的生物高分子化合物,在細(xì)胞骨架微管和微絲的作用下,色素顆粒通過聚集或分散作用而影響魚類體色(譚玉文等,2008)。目前已發(fā)現(xiàn)的魚類色素細(xì)胞有5種,包括黑色素細(xì)胞、紅/黃色素細(xì)胞、白色素細(xì)胞、虹彩細(xì)胞和藍(lán)色素細(xì)胞(Kelsh,2010)。其中由神經(jīng)脊細(xì)胞分化而來的黑色素細(xì)胞在魚體中分布最廣泛,能合成分泌具有抗輻射、抗氧化及免疫調(diào)節(jié)等功能的真黑素和褐黑素(Parichy et al.,2007;Geng et al.,2010;鄒宇等,2012)。魚類體色除了反映魚體的生理健康狀況外,還直接影響其市場(chǎng)價(jià)格。因此,研究魚類體色變異的分子機(jī)制,對(duì)提高魚類體色遺傳穩(wěn)定性及其經(jīng)濟(jì)價(jià)值具有重要意義?!厩叭搜芯窟M(jìn)展】大量研究證實(shí),MC-1R(Melanocortin-1 receptor,黑色素皮質(zhì)1受體)、TYR(Tyrosinase,酪氨酸酶)、TYRP1(Tyrosinase-related protein 1,酪氨酸相關(guān)蛋白1)、TYRP2(Tyrosinase-related protein 2,酪氨酸相關(guān)蛋白2)、ASIP(Agouti signaling protein,刺鼠信號(hào)蛋白)、MITF(Microphthalmia-associated transcription factor,小眼輔助轉(zhuǎn)錄因子)和SOX10(SRY-related HMG-box,Y染色體性別決定基因相關(guān)轉(zhuǎn)錄因子10)等基因在黑色素合成過程中發(fā)揮重要作用(Masca-renhas et al.,2010;Miao et al.,2010;李康樂,2014;徐瑩,2014)。相對(duì)于其他脊椎動(dòng)物,魚類擁有更復(fù)雜且多樣化的色素沉積方式,受多個(gè)基因及信號(hào)通路調(diào)控。目前,關(guān)于魚類機(jī)體色素沉積的研究已有較多報(bào)道。Jiang等(2014)基于轉(zhuǎn)錄組分析興國紅鯉和黃河鯉皮膚組織時(shí)發(fā)現(xiàn),鯉魚皮膚色素沉積與黑色素合成、Wnt信號(hào)通路及MAPK信號(hào)通路有關(guān);McCauley等(2004)研究發(fā)現(xiàn)Oca2基因部分缺失及MC-1R基因突變能引起墨西哥麗脂鯉體色變異;肖穎琦(2017)認(rèn)為MAPK信號(hào)通路中的dusp6、hsp27、map2k4和mef2c基因與玫瑰高原鰍體色白化有關(guān)。原肌球蛋白(Tropomyosin,TMs)被認(rèn)為是肌球蛋白(Myosin)的穩(wěn)定蛋白,是一種高度保守和多樣化的肌動(dòng)蛋白家族,可調(diào)節(jié)肌球蛋白的動(dòng)力學(xué)和結(jié)構(gòu)特性,幾乎存在于所有的動(dòng)物機(jī)體內(nèi)(Lehman et al.,2009)。原肌球蛋白的編碼基因(TPM)包含TPM1、TPM2、TPM3和TPM4共4種亞型,TPM基因受損或缺失將導(dǎo)致細(xì)胞惡性轉(zhuǎn)化,形成腫瘤或產(chǎn)生畸形,如食管鱗癌、直腸癌和足內(nèi)翻非綜合征等(趙秀麗,2006;趙月鳴,2015;Weymouth et al.,2016)。肌球蛋白是動(dòng)物肌肉的主要構(gòu)成蛋白,而肌球蛋白重鏈(Myosin heavy chain,MYH)是肌球蛋白的組成單位,在非肌肉細(xì)胞中發(fā)揮細(xì)胞骨架的作用,同時(shí)參與細(xì)胞物質(zhì)運(yùn)輸、能量代謝及信號(hào)傳導(dǎo)等途徑。黑素體從黑色素細(xì)胞轉(zhuǎn)移到樹突,再傳遞給周圍角質(zhì)形成細(xì)胞的過程均依賴于細(xì)胞骨架中肌動(dòng)蛋白(Actin)和肌球蛋白等提供動(dòng)力(Hara et al.,2000)。肌動(dòng)蛋白在黑色素細(xì)胞樹突的遠(yuǎn)端聚集,作為黑素體運(yùn)輸馬達(dá),其中肌球蛋白5a(MYO5A)可與黑素親和素(MLPH)及GTP結(jié)合蛋白(Rab27a)結(jié)合形成黑色素轉(zhuǎn)運(yùn)復(fù)合體,將黑素體轉(zhuǎn)運(yùn)至樹突末端,終止黑素體在樹突中的雙向長距離運(yùn)動(dòng)(Matesic et al.,2001;Fontanesi et al.,2014)。Wu等(1998)研究發(fā)現(xiàn),肌球蛋白基因突變會(huì)導(dǎo)致小鼠毛發(fā)顏色變淺,究其原因是黑素體運(yùn)輸受損而聚集在細(xì)胞核周區(qū)。Hara等(2000)研究表明,細(xì)胞骨架成分和微管相關(guān)動(dòng)力蛋白參與黑色素在角質(zhì)形成細(xì)胞內(nèi)的分布及運(yùn)動(dòng),且動(dòng)力蛋白與黑素體存在共區(qū)域化?!颈狙芯壳腥朦c(diǎn)】金邊鯉是針對(duì)稻田養(yǎng)殖而選育出的鯉魚新選系,其外形特點(diǎn)為背鰭兩側(cè)有能穩(wěn)定遺傳的金色條帶。本課題組前期在金邊鯉皮膚轉(zhuǎn)錄組的研究中富集到與肌球蛋白和肌動(dòng)蛋白有關(guān)的差異表達(dá)基因(Yan et al.,2019),但目前金邊鯉皮膚變異的分子機(jī)制尚未清楚?!緮M解決的關(guān)鍵問題】測(cè)定肌動(dòng)蛋白3基因(ACT3)、肌球蛋白重鏈2基因(MYH2)、原肌球蛋白3個(gè)亞型基因(TPM1、TPM2和TPM3)及黑色素合成相關(guān)基因(MC-1R、TYR、TYRP1和TYRP2)在金邊鯉不同皮膚中的表達(dá)情況,并分析其相關(guān)性,旨在探究金邊鯉皮膚顏色差異形成的原因,為其良種培育提供候選基因信息。

      1 材料與方法

      1. 1 試驗(yàn)材料

      供試鯉魚取自廣西融水縣融榮水產(chǎn)繁殖場(chǎng)(同一養(yǎng)殖池塘),分為有金邊和無金邊的鯉魚,6月齡,體質(zhì)量65.17±12.90 g/尾,體長14.35±0.91 cm。采集8尾突變型金邊鯉魚背部的黃色皮膚和黑色皮膚,分別記為Y組和B組;同時(shí)采集8尾野生型金邊鯉魚背部的黑色皮膚,記為W組(圖1)。樣品存于2 mL的凍存管中,以液氮速凍后轉(zhuǎn)入-80 ℃冰箱保存?zhèn)溆谩?/p>

      1. 2 總RNA提取及cDNA合成

      3. 2 腎上腺素信號(hào)通路在色素沉著中的作用

      色素細(xì)胞膜上通常分布有細(xì)小的肌纖維和神經(jīng)末梢。色素細(xì)胞在肌纖維的收縮牽引下改變形態(tài),促使色素顆粒在驅(qū)力蛋白的作用下運(yùn)動(dòng),進(jìn)而分散或聚集在機(jī)體各部位,是魚類體色變化的主要原因。多巴(Levodopa)是黑色素合成的中間產(chǎn)物,其常見代謝途徑是被TYR多次催化形成多巴醌(Dopaquinone),最終氧化形成黑色素(Hearing,2011)。多巴的另一代謝途徑是形成兒茶酚胺,兒茶酚胺在相應(yīng)的轉(zhuǎn)移酶作用下可轉(zhuǎn)化為多巴胺(Dopamine)、去甲腎上腺素(Noradrenaline,NE)和腎上腺素(Adrena-line,AD)(余鵬程,2017)。已有研究證明,黑色素細(xì)胞上存在β腎上腺素受體,且AD負(fù)責(zé)黑色素顆粒的擴(kuò)散,使動(dòng)物體色變深;NE負(fù)責(zé)黑色素團(tuán)內(nèi)色素聚集而使動(dòng)物體色變淺,均屬于神經(jīng)調(diào)控體色變化范疇(Fujii,2000;Sivamani et al.,2009)。腎上腺素能受體(Adrenergic receptor,AR)屬于G蛋白偶聯(lián)受體,是兒茶酚胺類物質(zhì)作用的受體蛋白,可分為α1、α2、β1和β2等4個(gè)亞型(Gillbro et al.,2004;Li et al.,2004)。β1-AR能增加細(xì)胞內(nèi)的Ca2+濃度,β2-AR可刺激腺苷酸環(huán)化酶(cAMP),與黑色素合成的關(guān)鍵步驟相似。β1-AR拮抗劑會(huì)抑制白色素細(xì)胞色素顆粒擴(kuò)散,但對(duì)黑色素細(xì)胞無影響;β2-AR介導(dǎo)AD抑制黑色素聚集;α2-AR激動(dòng)劑能調(diào)控黑色素細(xì)胞和紅色素細(xì)胞的色素合成,同時(shí)通過介導(dǎo)NE引起黑色素顆粒聚集,且對(duì)黑色素細(xì)胞凋亡有影響(Fujii,2000)。Chou等(1989)認(rèn)為AD誘導(dǎo)黑色素聚集的信號(hào)轉(zhuǎn)導(dǎo)系統(tǒng)第二信使并不是cGMP和cAMP,也不是通過Ca2+信號(hào)通路,而是存在一個(gè)未知的第二信使。余鵬程(2017)認(rèn)為β2-AR通過G蛋白—腺苷酸環(huán)化酶—蛋白激酶A(GS-AC-PKA)信號(hào)通路調(diào)節(jié)TYR活性或增加其相關(guān)基因(TYR、TYRP和TYRP2等)的表達(dá)量,從而影響黑色素生成。

      本研究在金邊鯉皮膚轉(zhuǎn)錄組富集到的ACT3、MYH2、TPM1、TPM2和TPM3等5個(gè)差異表達(dá)基因均來源于腎上腺素信號(hào)通路(Adrenergic signaling in cardiomyocytes)。在心肌腎上腺素信號(hào)通路中,AD與β1-AR或β2-AR受體結(jié)合活化Gs蛋白,激活腺苷酸環(huán)化酶(AC),影響第二信使cAMP濃度,隨后激活蛋白激酶A系統(tǒng)(PKA)和Ca2+信號(hào)通路,最終影響ACT3、TPM和MYH2基因的表達(dá),與余鵬程(2017)認(rèn)為β2-AR通過GS-AC-PKA通路調(diào)節(jié)黑色素生成的結(jié)論一致。孫福亮(2016)在新吉細(xì)毛羊和小尾寒羊皮膚轉(zhuǎn)錄分析中同樣富集到TPM1、TPM2等與心肌收縮和心肌腎上腺素信號(hào)通路相關(guān)的差異表達(dá)基因。本研究富集到的ACT3、MYH和TPM等基因家族均與黑色素運(yùn)動(dòng)的驅(qū)動(dòng)蛋白有關(guān),且除了ACT3基因外,其余4個(gè)基因在金邊鯉黃色皮膚中的表達(dá)量均顯著低于黑色皮膚,且與MC-1R和TYR基因表達(dá)呈顯著或極顯著相關(guān),推測(cè)這些基因是通過心肌腎上腺素信號(hào)通路影響金邊鯉皮膚黑色素合成或轉(zhuǎn)運(yùn)途徑,最終影響金邊鯉皮膚色素沉著。但究竟是TPM、MYH2、Actins等基因表達(dá)差異影響黑色素顆粒運(yùn)動(dòng)而導(dǎo)致金邊鯉皮膚顏色差異,還是金邊鯉金色皮膚中黑色素合成途徑受阻才影響這些基因的表達(dá),尚有待進(jìn)一步探究。

      4 結(jié)論

      MC-1R、TYR、TYRP1和TYRP2基因及其所在信號(hào)通路可能影響金邊鯉金色皮膚形成,但并非主要因素;ACT3、MYH2、TPM1、TPM2和TPM3基因及其所在的心肌腎上腺素信號(hào)通路可能通過影響黑色素合成和運(yùn)動(dòng),是金邊鯉體色變異的主要影響因素。ACT3、MYH2、TPM1、TPM2和TPM3基因與黑色素合成相關(guān)基因存在相關(guān)性,可作為金邊鯉體色變異研究中的標(biāo)記基因。

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