魚(yú)類(lèi)血栓細(xì)胞免疫功能的研究現(xiàn)狀

高迎莉, 韓高尚, 姚海靜, 陳香凝, 周謙青

魚(yú)類(lèi)血栓細(xì)胞免疫功能的研究現(xiàn)狀

高迎莉1, 2, 3, 韓高尚1, 姚海靜1, 陳香凝1, 2, 3, 周謙青1

(1. 江蘇海洋大學(xué) 海洋生命與水產(chǎn)學(xué)院, 江蘇 連云港 222000; 2. 江蘇省海洋生物技術(shù)重點(diǎn)實(shí)驗(yàn)室, 江蘇 連云港 222000; 3. 江蘇省海洋生物資源與環(huán)境重點(diǎn)實(shí)驗(yàn)室, 江蘇 連云港 222000)

血細(xì)胞研究是魚(yú)類(lèi)基礎(chǔ)研究的重中之重。魚(yú)類(lèi)血細(xì)胞的前期研究主要聚焦于淋巴細(xì)胞、巨噬細(xì)胞和嗜中性粒細(xì)胞等細(xì)胞的功能特性, 而忽略了血栓細(xì)胞。血栓細(xì)胞是低等脊椎動(dòng)物具有的類(lèi)似于哺乳動(dòng)物血小板的一類(lèi)細(xì)胞, 該細(xì)胞的經(jīng)典功能是參與凝血和血栓形成。近年來(lái)的研究表明, 血栓細(xì)胞是一類(lèi)特殊的免疫細(xì)胞, 在魚(yú)體免疫應(yīng)答中發(fā)揮重要作用。作者將對(duì)魚(yú)類(lèi)血栓細(xì)胞免疫功能相關(guān)的形態(tài)學(xué)、免疫學(xué)功能和研究前景等內(nèi)容進(jìn)行綜述, 為魚(yú)類(lèi)血栓細(xì)胞功能的多樣性提供基礎(chǔ)資料。

形態(tài)學(xué); 免疫功能; 血栓細(xì)胞; 魚(yú)類(lèi)

1882年, Giulio[1]描述血液中一種新的“形態(tài)學(xué)成分”, 命名為血小板(Platelets), 并發(fā)現(xiàn)血小板在機(jī)體出血和血栓形成中發(fā)揮作用, 因此Giulio被稱(chēng)為“血小板之父”。后來(lái)的研究發(fā)現(xiàn), 哺乳動(dòng)物無(wú)核的血小板來(lái)源于骨髓中的巨核細(xì)胞, 在非特異性和特異性免疫反應(yīng)中發(fā)揮重要作用[2]; 低等脊椎動(dòng)物如鳥(niǎo)類(lèi)[3]、爬行類(lèi)[4]、兩棲類(lèi)[5]和魚(yú)類(lèi)[6]中, 類(lèi)似于血小板的細(xì)胞是有核的血栓細(xì)胞。近年來(lái)的研究表明, 血栓細(xì)胞參與魚(yú)類(lèi)的非特異性免疫反應(yīng)以及炎癥反應(yīng)[7-9]。此外, 血栓細(xì)胞參與魚(yú)類(lèi)的特異性免疫應(yīng)答的研究引起眾多研究者的興趣[7-8, 10-12]。魚(yú)類(lèi)的血栓細(xì)胞是否具有功能多樣性, 有待于進(jìn)一步的研究。作者將對(duì)魚(yú)類(lèi)血栓細(xì)胞免疫功能相關(guān)的形態(tài)學(xué)和細(xì)胞發(fā)生、免疫學(xué)功能等研究現(xiàn)狀進(jìn)行綜述, 以全面闡述魚(yú)類(lèi)血細(xì)胞的新功能, 為魚(yú)類(lèi)血栓細(xì)胞功能的多樣性提供基礎(chǔ)資料。

1 血栓細(xì)胞的形態(tài)學(xué)特征

硬骨魚(yú)血液中, 血栓細(xì)胞在數(shù)量上僅次于紅細(xì)胞, 這些數(shù)量龐大的血栓細(xì)胞參與魚(yú)體的血栓形成和凝血過(guò)程[6]。血栓細(xì)胞成群分布, 呈淚滴型、紡錘型、卵圓型或孤核型等多種形態(tài), 具有指狀突起、絲狀偽足、囊泡、液泡、微管系統(tǒng)、顆粒、線粒體、高爾基體以及核糖體等成分[7, 13-23]。Jagadeeswaran[24]等發(fā)現(xiàn)斑馬魚(yú)()血栓細(xì)胞具有開(kāi)放的管狀系統(tǒng), 血栓細(xì)胞活化后, 出現(xiàn)絲狀偽足, 細(xì)胞發(fā)生凝集反應(yīng)。斑點(diǎn)叉尾鮰()血栓細(xì)胞呈梭狀, 胞內(nèi)具有富含吞噬物質(zhì)的液泡, 胞質(zhì)中含有彌漫性的糖原顆粒[25](圖1a)。聚蓋魚(yú)() 血栓細(xì)胞有梭形和球形兩種, 細(xì)胞核有中心常染色質(zhì), 細(xì)胞質(zhì)有液泡、管狀系統(tǒng)以及吞噬體[26](圖1b)。Azevedo[27]等的研究發(fā)現(xiàn), 白紋笛鯛()血栓細(xì)胞被紅細(xì)胞圍著, 可形成多個(gè)胞質(zhì)突起而呈現(xiàn)不規(guī)則細(xì)胞膜, 同時(shí)血栓細(xì)胞通過(guò)投影接近多個(gè)紅細(xì)胞的細(xì)胞膜; 血栓細(xì)胞核內(nèi)的異染色質(zhì)和常染色質(zhì)中包含部分顆粒原纖維團(tuán)塊及部分發(fā)育的液泡, 這些液泡有的位于胞質(zhì)突起的頂端, 有的在細(xì)胞質(zhì)中隨機(jī)分布。最新研究表明, 人類(lèi)血小板的細(xì)胞膜具有偽足和突起, 細(xì)胞內(nèi)部具有液泡、α-顆粒、致密顆粒和溶酶體顆粒、管狀系統(tǒng)、線粒體和自噬體。此外, 人類(lèi)血小板的形態(tài)結(jié)構(gòu)與年齡相關(guān), 年齡增大, 細(xì)胞膜的規(guī)整性降低, 光滑性減少, 偽足和突起增多且更細(xì)長(zhǎng), 細(xì)胞內(nèi)部的α-顆粒顯著減少[28]。綜上所述, 不同種魚(yú)類(lèi)的血栓細(xì)胞在形態(tài)結(jié)構(gòu)上存在差異, 但如突起、偽足、囊泡、溶酶體等跟免疫相關(guān)的形態(tài)結(jié)構(gòu)均出現(xiàn)在血栓細(xì)胞中, 這為血栓細(xì)胞的功能多樣性奠定了結(jié)構(gòu)基礎(chǔ)。而人類(lèi)血小板的形態(tài)結(jié)構(gòu)是伴隨時(shí)間發(fā)生變化, 因此不同魚(yú)類(lèi)血栓細(xì)胞的結(jié)構(gòu)差異是否與魚(yú)體的年齡、性別、健康狀態(tài)等相關(guān), 仍需要進(jìn)一步探究。

圖1 血栓細(xì)胞的超微結(jié)構(gòu)觀察

a. Ultrastructure of thrombocytes in channel catfish. Fusiform thrombocyte with vacuoles, 1 of which contains phagocytosed material, bar=500 nm[25]; b. Ultrastructure of thrombocytes in fat snook. A fusiform thrombocyte to the left with its nucleus (N), to the right containing large phagosomes (fg), bar=2 μm[26]

細(xì)胞化學(xué)染色是以細(xì)胞形態(tài)學(xué)為基礎(chǔ), 根據(jù)某一特定胞內(nèi)組分的化學(xué)反應(yīng)原理, 在顯微鏡下觀察細(xì)胞化學(xué)成分及其變化的顯微研究方法。常用的有酶類(lèi)(酸性磷酸酶ACP、堿性磷酸酶ALP、過(guò)氧化物酶Peroxidase、酯酶等)、脂類(lèi)(蘇丹黑B染色SBB)、糖原(過(guò)碘酸-希夫反應(yīng)PAS)等細(xì)胞化學(xué)染色, 且常聯(lián)合吉姆薩Giemsa染色用于區(qū)別不同細(xì)胞亞群或者用于細(xì)胞功能的研究(圖2)。硬骨魚(yú)類(lèi)種類(lèi)不同, 血栓細(xì)胞的細(xì)胞化學(xué)特性有差別, 如莫瑞鱈魚(yú)(, Mitchell)[29](圖3)等魚(yú)類(lèi)的血栓細(xì)胞呈現(xiàn)不同程度的細(xì)胞化學(xué)特性的相似性與差異性(表1)。除了藍(lán)旗金槍魚(yú)()[30], 所有魚(yú)類(lèi)的血栓細(xì)胞都是PAS陽(yáng)性, 糖原是中性粒細(xì)胞進(jìn)行胞內(nèi)殺傷以及清除外來(lái)物質(zhì)時(shí)所需的能量來(lái)源, 這從形態(tài)學(xué)角度證明魚(yú)類(lèi)血栓細(xì)胞的吞噬潛能。藍(lán)旗金槍魚(yú)是大洋洄游性魚(yú)類(lèi), 其血栓細(xì)胞的功能與其他魚(yú)類(lèi)存在差異, 表明不同生境魚(yú)類(lèi)的血栓細(xì)胞會(huì)進(jìn)化出與其生境或者生理活動(dòng)相匹配的功能, 藍(lán)旗金槍魚(yú)血栓細(xì)胞是否具有吞噬功能需要進(jìn)一步的研究。人類(lèi)血小板的吞噬溶酶體中含有一些酶類(lèi), 如ACP, 這是血小板具有吞噬能力的證據(jù)[31]。由表1可知, 除了莫瑞鱈魚(yú)、花斑裸鯉[32]和虹鱒[33], 其他魚(yú)類(lèi)血栓細(xì)胞都是ACP陽(yáng)性, 表明這些魚(yú)類(lèi)的血栓細(xì)胞具有吞噬活性。但是, 莫瑞鱈魚(yú)、花斑裸鯉和虹鱒的血栓細(xì)胞超微結(jié)果顯示,細(xì)胞表面具有管狀系統(tǒng), 胞內(nèi)有吞噬小泡和吞噬物, 說(shuō)明這3種魚(yú)類(lèi)血栓細(xì)胞的吞噬活性是機(jī)械式而非酶活式。其他的細(xì)胞化學(xué)染色結(jié)果差別較大, 呈現(xiàn)明顯的種屬特異性, 如大蓋具脂鯉和條紋鯪脂鯉的ALP染色結(jié)果相反; 鏟吻油鯰[34]是蘇丹黑B陰性; 熱帶魚(yú)[35]是過(guò)氧化氫陰性, 但黃鲇魚(yú)[36]和大口黑鱸[37]是過(guò)氧化氫陽(yáng)性; 羅非魚(yú)[38]僅僅檢測(cè)了糖原特性。魚(yú)類(lèi)血栓細(xì)胞的形態(tài)學(xué)研究不僅可用于區(qū)分魚(yú)類(lèi)的不同血細(xì)胞類(lèi)群, 而且可以直接觀察血栓細(xì)胞的外部細(xì)胞膜、突起等結(jié)構(gòu)以及內(nèi)部細(xì)胞器等組成, 這為進(jìn)一步研究血栓細(xì)胞的功能特性奠定結(jié)構(gòu)基礎(chǔ)。此外, 細(xì)胞化學(xué)特性為魚(yú)類(lèi)血栓細(xì)胞的種屬差異性、年齡相關(guān)性、健康狀態(tài)相關(guān)性、吞噬活性、免疫功能相關(guān)性等的研究提供直接證據(jù)。

圖2 魚(yú)類(lèi)血栓細(xì)胞的形態(tài)圖

a. The thrombocytes of channel catfish (arrowhead) are smaller than the erythrocytes, with a spindle-shaped nucleus[25]; b. The thrombocyte of southern stingray (arrowhead)[23]; c. The thrombocyte of white sturgeon (arrowhead)[23]

圖3 莫瑞鱈魚(yú)血栓細(xì)胞的細(xì)胞化學(xué)染色

a. 過(guò)氧化氫染色; b. 酸性磷酸酶(ACP)染色; c. 堿性磷酸酶(ALP)染色; d. 氯乙酸AS萘酚酯酶(NCE)染色; e. 乙酸萘酯酶(NAE)染色; f.α-丁酸萘酯酶(NBE)染色; g. 蘇丹黑B(SBB)染色; h. 糖原(PAS)染色; i. β-葡糖醛酸糖苷酶染色。放大倍數(shù)×100, 標(biāo)尺=10 μm[29]

a. peroxidase staining; b. acid phosphatase (ACP) staining; c. alkaline phosphatase (ALP) staining; d. naphthol AS chloroacetate esterase (NCE) staining; e. naphthyl acetate esterase (NAE) staining; f. α-naphthyl butyrate esterase (NBE) staining; g. Sudan black B (SBB) staining; h. periodic acid Shiff”s (PAS) staining; i. β-glucuronidase staining. Magnification × 100, Bar=10 μm[29]

表1 血栓細(xì)胞和血小板的細(xì)胞化學(xué)特性

續(xù)表

注: “+”. 陽(yáng)性; “–”. 陰性; 鯉魚(yú)[8, 10](); 胭脂魚(yú)[17](); 條紋鯪脂鯉[25](); 大蓋具脂鯉[25](); 聚蓋魚(yú)[26](); 莫瑞鱈魚(yú)[29](, Mitchell); 藍(lán)鰭金槍魚(yú)[30](); 花斑裸鯉[32](); 虹鱒[33](); 鏟吻油鯰[34](); 熱帶魚(yú)[35](); 黃鲇魚(yú)[36](); 大口黑鱸[37](); 羅非魚(yú)[38]()

2 血栓細(xì)胞的免疫學(xué)功能

2.1 吞噬作用

吞噬作用是內(nèi)吞作用的特殊形式, 包括顆粒的水泡內(nèi)化, 形成內(nèi)部囊泡即吞噬溶酶體, 是非特異性防御病原微生物的重要機(jī)制[39]。魚(yú)類(lèi)專(zhuān)業(yè)的吞噬細(xì)胞有單核/巨噬細(xì)胞[40]、嗜中性粒細(xì)胞[41]和樹(shù)突狀細(xì)胞[42], 此外, 淋巴細(xì)胞也具有吞噬作用[43-45]。關(guān)于上述各類(lèi)細(xì)胞吞噬功能的研究多采用細(xì)胞系培養(yǎng)或細(xì)胞分離方法, 將胞內(nèi)寄生蟲(chóng)、酵母菌、熒光微球和胞內(nèi)寄生細(xì)菌等作為被吞噬的顆粒物, 應(yīng)用細(xì)胞化學(xué)染色、光學(xué)顯微鏡、熒光顯微鏡、透射電鏡以及流式細(xì)胞術(shù)等技術(shù), 證明細(xì)胞的吞噬作用。

近年來(lái)的研究結(jié)果表明, 魚(yú)類(lèi)血栓細(xì)胞也具有吞噬作用, 但是血栓細(xì)胞在病原菌吞噬和清除上存在爭(zhēng)議, 這可能是實(shí)驗(yàn)技術(shù)的不同造成的。細(xì)胞形態(tài)學(xué)結(jié)果顯示, 血栓細(xì)胞胞內(nèi)存在具有吞噬能力的顆粒[46], 然而, 細(xì)胞化學(xué)分析表明, 血栓細(xì)胞ACP呈陰性[37]。但人類(lèi)血小板吞噬體酶類(lèi)鑒定結(jié)果表示, ACP陽(yáng)性是吞噬體不可或缺的成分。因此, 這種不一致是由于研究方法的局限性, 還是由于魚(yú)類(lèi)血栓細(xì)胞的細(xì)胞化學(xué)特殊性, 有待于進(jìn)一步研究。脊椎動(dòng)物的血小板吞噬作用具有4個(gè)特征, 且在魚(yú)類(lèi)血栓細(xì)胞中也有體現(xiàn): (1) 血栓細(xì)胞與顆粒性物質(zhì)(非生物顆粒、病原體等)相互作用。鯉魚(yú)血栓細(xì)胞可以攝入并殺傷金黃色釀膿葡萄球菌()[46]以及大腸埃希氏菌()[8]; Burrows[47]等發(fā)現(xiàn), 瘤棘鲆()血栓細(xì)胞能夠在體外主動(dòng)吞噬碳顆粒, 具有較強(qiáng)的吞噬能力; 牙鲆()和鯉魚(yú)血栓細(xì)胞(圖4)能夠在體外吞噬熒光微球[8]; (2) 抗原刺激后, 血栓細(xì)胞胞質(zhì)的溶酶體產(chǎn)物被釋放, 導(dǎo)致胞內(nèi)殺菌作用。白紋笛鯛血栓細(xì)胞的超微結(jié)構(gòu)顯示, 抗原刺激后, 胞內(nèi)微管釋放空泡物質(zhì), 而該物質(zhì)是血栓細(xì)胞吞噬作用的產(chǎn)物, 從細(xì)胞結(jié)構(gòu)上印證血栓細(xì)胞的吞噬作用[27]; 鯉魚(yú)血栓細(xì)胞吞噬溶酶體融合實(shí)驗(yàn)表明, 血栓細(xì)胞可以?xún)?nèi)化細(xì)菌且具有殺菌能力[8]; (3)吞噬作用的代謝產(chǎn)物能作為炎癥反應(yīng)的媒介。Ferdous[3]研究了雞血栓細(xì)胞吞噬作用, 證明其代謝產(chǎn)物介導(dǎo)炎癥反應(yīng)。而Nagasawa[12]等通過(guò)研究血栓細(xì)胞吞噬作用與白細(xì)胞分泌的激活因子之間的關(guān)系, 發(fā)現(xiàn)血栓細(xì)胞的吞噬活性會(huì)因白細(xì)胞分泌激活因子的增加而增加, 從而使其代謝產(chǎn)物在急性炎癥反應(yīng)中發(fā)揮作用; (4)利用補(bǔ)體, 通過(guò)調(diào)理素作用增強(qiáng)吞噬消化功能。研究表明, 鯉魚(yú)血清對(duì)血栓細(xì)胞的吞噬活力起到增強(qiáng)的作用[8]。以上關(guān)于魚(yú)類(lèi)血栓細(xì)胞吞噬作用研究主要集中于體外吞噬, 血栓細(xì)胞體內(nèi)吞噬研究涉及較少, 研究方法單一, 缺乏多種方法的聯(lián)合使用以及方法間的相互佐證, 且血栓細(xì)胞吞噬作用介導(dǎo)的胞內(nèi)殺傷作用等仍然不夠明確, 這在很大程度上限制對(duì)魚(yú)類(lèi)血栓細(xì)胞抗菌功能的認(rèn)知與探索。

圖4 鯉魚(yú)血栓細(xì)胞吞噬熒光微球和細(xì)菌

a.b. 血栓細(xì)胞吞噬熒光微球(X為被吞噬的微球); c. 血栓細(xì)胞通過(guò)延伸偽足吞噬細(xì)菌, 箭頭所示偽足, X.細(xì)菌; d. 小囊泡圍繞著被攝入的細(xì)菌, *. 囊泡, 標(biāo)尺=1 μm[8]

a.b. Phagocytosis with fluorescent beads by thrombocytes (Ingested beads are indicated as X); c. Thrombocytes engulfed bacteria (X) via the extension of pseudopods (arrowhead); d. Small vesicles surrounding the internalized bacteria are indicated by asterisks (*), Bar=1 μm[8]

2.2 非特異性免疫反應(yīng)

機(jī)體組織損傷后, 血小板/血栓細(xì)胞會(huì)參與機(jī)體的穩(wěn)態(tài)以及血栓形成, 進(jìn)而參與炎癥反應(yīng)。血小板/血栓細(xì)胞會(huì)在第一時(shí)間聚集于損傷處, 除了聚集和參與炎癥反應(yīng), 血小板/血栓細(xì)胞表達(dá)和釋放有利于傷口組織修復(fù)的蛋白質(zhì)和物質(zhì)[48]。不同動(dòng)物的血栓細(xì)胞分泌不同的細(xì)胞因子參與炎癥反應(yīng)和傷口修復(fù)。鳥(niǎo)類(lèi)血栓細(xì)胞產(chǎn)生一些生物活性物質(zhì), 即趨化因子(巨噬細(xì)胞炎癥蛋白-1β和一氧化氮)以及炎癥反應(yīng)的媒介物質(zhì), 如誘導(dǎo)型一氧化氮合酶(iNOS)等; 雞血栓細(xì)胞能夠表達(dá)抗炎細(xì)胞因子(轉(zhuǎn)化生長(zhǎng)因子TGF和IL-10)和前炎癥因子(IL-1β, IL-6, IL-8和 IL-12)以及利于傷口修復(fù)的細(xì)胞因子[3]。鯉魚(yú)和虹鱒血栓細(xì)胞能表達(dá)IL-1β、iNOS、腫瘤壞死因子TNFα、TGFβ和一些趨化因子[7-8]。

在哺乳動(dòng)物中, 血小板可以表達(dá)一些非特異性免疫相關(guān)分子, 如Toll樣受體(Toll-like receptors, TLR)和抗原呈遞相關(guān)基因。鯉魚(yú)、大西洋鱈魚(yú)()、三刺魚(yú)()和虎河豚()的血栓細(xì)胞可以表達(dá)輔助分子(TLR4 interactor with leucine-rich repeats, TRIL)[9], 這表明魚(yú)類(lèi)血栓細(xì)胞會(huì)像其他白細(xì)胞一樣, 在非特異性免疫反應(yīng)中直接發(fā)揮作用。

2.3 特異性免疫反應(yīng)

抗原呈遞是激活特異性免疫反應(yīng)過(guò)程中的最后一個(gè)步驟, 主要經(jīng)歷: (1) 攝取抗原; (2) 將抗原呈遞給特異性的胞內(nèi)組分; (3) 將抗原肽釋放到抗原結(jié)合位點(diǎn)-主要組織性相容性復(fù)合體Ⅱ分子(MHC II)上; (4) 激活CD4輔助性T細(xì)胞。魚(yú)類(lèi)的單核/巨噬細(xì)胞、樹(shù)突狀細(xì)胞和B淋巴細(xì)胞除了具有直接抗菌作用外, 還表達(dá)Ⅱ型組織相容性復(fù)合體, 且具有抗原處理和呈遞的分子機(jī)制。低等脊椎動(dòng)物血栓細(xì)胞表面具有MHC Ⅱ分子, 如鳥(niǎo)類(lèi)[49]、虹鱒和鯉魚(yú)[7-8, 10-11]的血栓細(xì)胞能夠表達(dá)MHC Ⅱ分子mRNA, 參與抗原呈遞過(guò)程。此外, 吞噬異物的血栓細(xì)胞以及具有抗原承載力的血栓細(xì)胞在鯉魚(yú)的頭腎和脾臟中都有所發(fā)現(xiàn), 也說(shuō)明血栓細(xì)胞具有運(yùn)載抗原到淋巴組織并進(jìn)入特異性免疫階段的能力[8]。以上研究?jī)H局限于定量PCR的方法, 鑒定血栓細(xì)胞中免疫相關(guān)基因的表達(dá)水平變化, 缺少不同層次(基因水平、蛋白水平、細(xì)胞水平)的科學(xué)證據(jù)來(lái)論證血栓細(xì)胞的免疫功能。目前, 魚(yú)類(lèi)血栓細(xì)胞非特異性和特異性免疫反應(yīng)的分子水平研究與哺乳動(dòng)物血小板還存在很大的差距。

3 血栓細(xì)胞的未來(lái)研究方向

魚(yú)類(lèi)血栓細(xì)胞的免疫功能研究具有較大的發(fā)展空間。目前, 魚(yú)類(lèi)血栓細(xì)胞的研究主要集中于魚(yú)類(lèi)血栓細(xì)胞特性研究, 這將仍是魚(yú)類(lèi)血栓細(xì)胞研究的方向。此外, 魚(yú)類(lèi)血栓細(xì)胞特性是否和人類(lèi)血小板類(lèi)似, 與自身因子(年齡)等因素有關(guān)[28], 仍需深入研究。因此, 魚(yú)類(lèi)血栓細(xì)胞特性與外界環(huán)境因素(棲息地、溶氧、pH等)以及內(nèi)部因子(種類(lèi)、年齡、性別、生理階段等)的關(guān)系將會(huì)是研究的重點(diǎn)。血栓細(xì)胞表面特異性標(biāo)記物及受體[49-52]發(fā)掘, 可以研發(fā)血栓細(xì)胞的特異性單克隆抗體, 如虹鱒[7]、斑馬魚(yú)[53]、鯉魚(yú)[54]和牙鲆[55], 結(jié)合免疫磁珠、流式細(xì)胞術(shù)等方法, 提高魚(yú)類(lèi)血栓細(xì)胞的分離純化效率[56-57]。利用分離純化的血栓細(xì)胞, 在前期研究的基礎(chǔ)上[7-12], 研究血栓細(xì)胞與其他免疫細(xì)胞的相互作用, 比如草魚(yú)血栓細(xì)胞與白細(xì)胞的作用[12], 血小板與樹(shù)突狀細(xì)胞的相互作用[58]; 借鑒血小板[59]和雞血栓細(xì)胞[60]的研究方法, 開(kāi)展魚(yú)類(lèi)血栓細(xì)胞的組學(xué)研究和免疫調(diào)控網(wǎng)絡(luò)分析, 將會(huì)是未來(lái)研究的熱點(diǎn)。此外, 樹(shù)蛙()血栓細(xì)胞系已經(jīng)建立[5], 能否建立魚(yú)類(lèi)血栓細(xì)胞的細(xì)胞系, 開(kāi)展胞外囊泡[61]等熱點(diǎn)問(wèn)題研究, 仍需要進(jìn)一步探究。

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Current research on the immune function of thrombocytes in fish

GAO Ying-li1, 2, 3, HAN Gao-shang1, YAO Hai-jing1, CHEN Xiang-ning1, 2, 3, ZHOU Qian-qing1

(1. College of Marine Life and Fisheries, Jiangsu Oean University, Lianyungang, 222000, China; 2. Jiangsu Key Laboratory of Marine Biotechnology, Lianyungang 222000, China; 3. Jiangsu Key Laboratory of Marine Bioresources and Environment, Lianyungang 222000, China)

The research on blood cells is very crucial to the research on fish. Research is basically focused on the immune functions of lymphocytes, macrophages, and neutrophils. There is limited focus on thrombocytes. Platelets in mammals and thrombocytes in lower vertebrates are implicated in thrombotic and hemostatic processes. The thrombocytes are regarded as special immune cells that play important roles in the immune response of fish based on current reports. This paper systemically reviews the immune-associated morphological features, immune functions, and future research perspectives on thrombocytes, with the aim of providing basic insight into the multi- functional nature of thrombocytes.

Morphology; immune function; thrombocyte; fish

Nov. 6, 2019

Q-1

A

1000-3096(2021)01-0120-09

10.11759/hykx20191106001

2019-11-06;

2020-05-25

江蘇省自然科學(xué)基金青年基金項(xiàng)目(BK20170450); 江蘇省省政策引導(dǎo)類(lèi)計(jì)劃-蘇北科技專(zhuān)項(xiàng)項(xiàng)目(SZ-LYG2017020); 江蘇省高等學(xué)校自然科學(xué)研究面上項(xiàng)目(17KJB240001); 江蘇省生物技術(shù)重點(diǎn)實(shí)驗(yàn)室開(kāi)放基金項(xiàng)目(HS2017003)和江蘇省海洋資源與環(huán)境重點(diǎn)實(shí)驗(yàn)室開(kāi)放基金項(xiàng)目(CXKT20180112); 江蘇省大學(xué)生創(chuàng)新創(chuàng)業(yè)項(xiàng)目(SY201811641105003)

[Natural Science Foundation of Jiangsu Province, No. BK20170450; Policy Guidance Program of Jiangsu Province, No. SZ-LYG2017020; Natural Science Foundation of the Jiangsu Higher Education Institutions of China, No. 17KJB240001; The Open Research Fund of Jiangsu Key Laboratory of Marine Biotechnology, No.HS2017003; The Open Research Fund of Jiangsu Key Laboratory of Marine Bioresources and Environment, No. CXKT20180112; Student Innovation and Entrepreneurship Program of Jiangsu Province, No. SY201811641105003]

高迎莉(1988- ), 女, 江蘇徐州人, 講師, 博士, 主要從事水產(chǎn)動(dòng)物病害與免疫學(xué)研究, E-mail: yingligao0127@126.com

(本文編輯: 譚雪靜)