史彩華, 胡靜榮, 李傳仁, 張友軍

(1. 長(zhǎng)江大學(xué), 荊州 434025; 2. 中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所, 北京 100081)

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專論與綜述

環(huán)境脅迫下昆蟲的耐寒適應(yīng)機(jī)制研究進(jìn)展

史彩華1,2, 胡靜榮1, 李傳仁1, 張友軍2*

(1. 長(zhǎng)江大學(xué), 荊州 434025; 2. 中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所, 北京 100081)

昆蟲是變溫動(dòng)物,為了安全越冬,昆蟲通常改變自身結(jié)構(gòu)和物質(zhì)構(gòu)成以適應(yīng)低溫的到來(lái),這一適應(yīng)機(jī)制與體內(nèi)特殊的生理生化物質(zhì)緊密相關(guān),如海藻糖、葡萄糖、甘油、山梨醇、脂肪酸和氨基酸等小分子抗凍保護(hù)劑。這些抗凍保護(hù)劑具有穩(wěn)定細(xì)胞膜結(jié)構(gòu)和保護(hù)蛋白質(zhì)功能的作用。雖然昆蟲耐寒性的研究不斷深入,但目前我們?nèi)匀缓茈y確定影響耐寒的關(guān)鍵因素是什么？為什么有些昆蟲在低于-20℃環(huán)境下還能存活？為了弄清這一科學(xué)問(wèn)題,科學(xué)家們利用轉(zhuǎn)錄組、基因組、蛋白質(zhì)組和代謝組等各種組學(xué)剖析低溫脅迫后昆蟲生理反應(yīng)的分子機(jī)制。本文旨在綜述前人對(duì)昆蟲耐寒性的研究,為將來(lái)其他昆蟲或動(dòng)植物的耐寒性研究提供參考依據(jù)。同時(shí),也為新型生物農(nóng)藥的開發(fā)和天敵昆蟲的人工助增提供廣闊思路。

昆蟲; 耐寒性; 抗凍保護(hù)劑; 機(jī)制

昆蟲耐寒性研究最早起源于18世紀(jì)30年代,Bachmetjew發(fā)現(xiàn)了昆蟲的過(guò)冷卻點(diǎn)[1]。直到20世紀(jì)60年代,Salt[2]應(yīng)用簡(jiǎn)單的儀器設(shè)備進(jìn)行了昆蟲耐低溫的生理試驗(yàn)。隨著科學(xué)技術(shù)的發(fā)展,許多高科技的手段被用于揭示低溫生物學(xué)問(wèn)題,比如:氣相色譜、液相色譜、電泳法、差示掃描量熱法、核磁共振光譜和各種組學(xué),包括轉(zhuǎn)錄組、基因組、基因芯片、蛋白質(zhì)組和代謝組等。

影響昆蟲耐寒的因素主要包括生物因素和非生物因素。生物因素主要包括寄主營(yíng)養(yǎng)、發(fā)育歷期及雌雄差異等;非生物因素主要有季節(jié)性氣溫變化、快速冷馴化、環(huán)境濕度、光周期及不同地理位置等。昆蟲耐寒是各種因素相互作用的結(jié)果[3],并非單一的體系,而是由多種抗寒物質(zhì)組成的復(fù)雜系統(tǒng)[4-6]。昆蟲應(yīng)對(duì)低溫耐寒的對(duì)策主要分為兩類:一類是行為對(duì)策,即生態(tài)適應(yīng)對(duì)策,當(dāng)冬季低溫來(lái)臨時(shí),昆蟲通過(guò)行為活動(dòng)尋找合適的躲避場(chǎng)所,比如遷飛、休眠和滯育等;另一類是生理對(duì)策,即改變體內(nèi)生理生化物質(zhì)比例,提高耐寒性。目前,昆蟲耐寒性研究主要圍繞過(guò)冷卻點(diǎn)、小分子抗凍保護(hù)劑、抗凍蛋白和熱休克蛋白等方面進(jìn)行機(jī)理解析[7]。然而,不同昆蟲耐寒機(jī)理不盡相同,本文對(duì)前人的大量研究進(jìn)行了綜述。

1 昆蟲耐低溫的生理生化機(jī)制

昆蟲是變溫動(dòng)物,能否安全越冬直接關(guān)系到昆蟲下一年種群的繁衍。19世紀(jì)60年代,人們開始意識(shí)到昆蟲的耐寒性與昆蟲自身生理調(diào)節(jié)緊密相關(guān)。

1.1 過(guò)冷卻點(diǎn)的降低

溫度低于昆蟲體液冰點(diǎn)而不凝固的現(xiàn)象稱作過(guò)冷卻現(xiàn)象。過(guò)冷卻點(diǎn)是描述昆蟲耐寒性的主要指標(biāo)之一。昆蟲的過(guò)冷卻點(diǎn)在不同個(gè)體和群體之間差異很大,受環(huán)境、自身發(fā)育、機(jī)體代謝等多種因素影響,因此,這些因素也影響昆蟲的耐寒性。進(jìn)入過(guò)冷卻狀態(tài)是北半球溫帶和寒溫帶地區(qū)昆蟲采取的主要越冬策略。蘋淡褐卷蛾(Epiphyaspostvittana)幼蟲經(jīng)冷馴化處理后,過(guò)冷卻點(diǎn)和死亡率均明顯降低[8]。美洲芹鳳蝶(Papiliozelicaon)越冬蛹過(guò)冷卻點(diǎn)為-20.5℃時(shí),可以在-30℃幸存,但經(jīng)春季暖溫暴露后,過(guò)冷卻點(diǎn)升為-17℃,在-20℃僅能存活1 h[9]。由此說(shuō)明低溫冷馴化對(duì)降低昆蟲過(guò)冷卻點(diǎn)至關(guān)重要,也是自然環(huán)境下昆蟲能夠正常越冬必不可少的步驟。

1.2 體內(nèi)冰核物質(zhì)的排除與結(jié)構(gòu)調(diào)整

生物體中常有異源催化物質(zhì),促使水分子重新排列,并在零下溫度結(jié)冰,這種催化物質(zhì)被稱作冰核促成活性因子(ice nucleating active agents,INA),簡(jiǎn)稱為冰核。早期在細(xì)菌的核中得到證實(shí),細(xì)胞外膜上的蛋白、碳水化合物和脂類限制了冰核的形成。同樣,在昆蟲體液中,也存在冰核。除去冰核可以增加昆蟲的耐寒性,提高小分子防凍保護(hù)劑的合成。食物和細(xì)菌在昆蟲腸道內(nèi)被認(rèn)為是潛在的冰核物質(zhì),因此,在低溫下可以通過(guò)停止取食的方式除去這些潛在的冰核物質(zhì)。最近,Worland等[10-11]在彈尾目昆蟲中發(fā)現(xiàn)蛻皮可以降低過(guò)冷卻點(diǎn),可能由于彈尾目昆蟲通過(guò)蛻皮脫掉了潛在的冰核物質(zhì)。近年來(lái)通過(guò)基因芯片技術(shù)證明了蛻皮與過(guò)冷卻點(diǎn)降低存在一定的聯(lián)系。另外,冰核結(jié)構(gòu)也是影響過(guò)冷卻點(diǎn)的因素,不同的冰核結(jié)構(gòu)有助于山楂粉蝶(Aporiacrataegi)耐受-85℃低溫[12]。因此,說(shuō)明不同昆蟲通過(guò)不同的途徑處理冰核來(lái)達(dá)到抵抗低溫的目的。

1.3 小分子抗凍保護(hù)劑的積累

越冬昆蟲體內(nèi)聚集小分子抗凍保護(hù)劑,如甘油、多元醇、糖類等,可增加昆蟲體內(nèi)束縛水的含量;直接與酶及其他蛋白相互作用,提高昆蟲機(jī)體的耐寒性。有研究報(bào)道,長(zhǎng)角血蜱(Haemaphysalislongicornis)滯育卵中山梨醇和甘油的積累是重要的越冬抗凍保護(hù)劑[13];冰川搖蚊(Pseudodiamesabranickii)體內(nèi)主要以葡萄糖和蔗糖作為小分子抗凍保護(hù)劑[14];大螟(Sesamiainferens)體內(nèi)小分子抗凍保護(hù)劑,如甘油、海藻糖、果糖、葡萄糖和肌醇含量的變化與季節(jié)氣溫波動(dòng)完全吻合[15]。因此,說(shuō)明昆蟲抵抗低溫并非單一物質(zhì)行為,而是存在多元抗凍保護(hù)系統(tǒng)。

1.4 脂類的轉(zhuǎn)化

目前,研究與耐寒相關(guān)的脂類主要有軟脂酸、亞油酸、脂肪酸酯和油酸等。脂肪是生物體的三大基礎(chǔ)物質(zhì)之一,具有貯能和保溫等功能,同時(shí)在低溫耐寒方面起著非常重要的作用[16-18]。昆蟲脂肪耐寒主要表現(xiàn)在:其一,脂肪具有貯能、供能的功效,如低溫飼養(yǎng)條件下劍川無(wú)鉤蝠蛾(Hepialusjianchuanensis)幼蟲體內(nèi)脂肪含量明顯高于較高溫度飼養(yǎng)的個(gè)體[19];其二,脂肪可以減少昆蟲體內(nèi)自由水含量,從而增加其耐寒性[20];其三,游離脂肪含量影響昆蟲過(guò)冷卻點(diǎn)的變化。作為脂肪代謝產(chǎn)物的甘油可以降低白蠟窄吉丁(Agrilusplanipennis)的過(guò)冷卻點(diǎn),保護(hù)其組織不受低溫的傷害[21]。

脂肪酸是脂肪合成的底物,因此,它與昆蟲的耐寒性有著緊密的聯(lián)系。一般情況下,隨著溫度的降低,不飽和脂肪酸含量升高,飽和脂肪酸含量降低,增加昆蟲體內(nèi)不飽和脂肪酸含量可以阻止細(xì)胞膜脂在低溫下結(jié)晶。Nieminen等[22]的研究表明,頸利虱蠅(Lipoptenacervi)體內(nèi)不飽和脂肪酸含量在晚秋季節(jié)顯著高于夏季。

脂肪參與昆蟲體內(nèi)各種代謝反應(yīng),直接或間接地影響昆蟲的多種生理反應(yīng)。同時(shí),脂肪含量的變化影響昆蟲耐寒性的機(jī)制也并非單一,因此,需要進(jìn)一步從分子層面深入研究。

1.5 自由氨基酸的合成

溫度脅迫也能觸發(fā)昆蟲體內(nèi)自由氨基酸的代謝反應(yīng)。在低溫條件下,自由氨基酸對(duì)昆蟲機(jī)體具有抗凍保護(hù)的作用。蠋蝽(Armachinensis)經(jīng)快速冷馴化處理后,體內(nèi)自由氨基酸含量發(fā)生較大變化,其中丙氨酸和谷氨酸含量明顯升高,脯氨酸含量明顯降低[23]。相反,植物抗寒性研究中普遍認(rèn)為脯氨酸是一類抗凍保護(hù)劑,其含量隨著低溫脅迫大幅度增加。因此,說(shuō)明自由氨基酸在不同生物體耐寒中的作用機(jī)制并不相同。

1.6 酶激活及離子平衡

無(wú)機(jī)離子(inorganic ions)是生命體不可缺少的一部分,在溶質(zhì)的跨膜運(yùn)動(dòng)中扮演著重要的角色,如K+、Na+和Ca2+等。當(dāng)昆蟲受到低溫脅迫時(shí),體內(nèi)會(huì)積累大量有毒代謝物質(zhì),堿性金屬離子和各種酶類可以幫助調(diào)節(jié)機(jī)體受損,確保正常的新陳代謝。在低溫暴露下,昆蟲血淋巴中K+濃度顯著增加,Na+濃度下降及腸道內(nèi)水分流失[24],這一觀點(diǎn)在黑腹果蠅(Drosophilamelanogaster)中已經(jīng)得到證實(shí)[25]。然而,東亞飛蝗(Locustamigratoria)經(jīng)低溫脅迫后血淋巴中K+濃度并未發(fā)生顯著變化[26]。說(shuō)明堿性金屬離子在不同昆蟲種類耐寒中的作用和變化動(dòng)態(tài)并非一致,如低溫暴露下,Ca2+在亞洲玉米螟(Ostriniafurnacalis)中誘導(dǎo)快速冷馴化,刺激甘油合成[27]。

溫度變化會(huì)通過(guò)一個(gè)信號(hào)傳導(dǎo)系統(tǒng)激活糖原磷酸化酶,糖原和其他冷凍保護(hù)劑隨著溫度的變化相互轉(zhuǎn)化。越冬過(guò)程中,溫度或光周期刺激直接影響昆蟲生長(zhǎng)發(fā)育中的激素,改變合成多元醇的酶活機(jī)制,如保幼激素(JH)和保幼激素類似物(JHA)刺激二化螟幼蟲體內(nèi)甘油聚集,蛻皮激素則降低其甘油濃度[28]。SOD和CAT能夠清除細(xì)胞內(nèi)的活性氧、羥自由基及其他過(guò)氧化物,以免對(duì)細(xì)胞造成毒害[29]。LDH是糖酵解途徑的關(guān)鍵酶之一,若其活性升高會(huì)直接影響昆蟲體內(nèi)能量代謝,促進(jìn)新陳代謝,致使昆蟲不耐寒。Na+,K+-ATP酶有促進(jìn)離子運(yùn)輸?shù)闹匾δ?低溫誘導(dǎo)使離子在細(xì)胞內(nèi)外平衡,啟動(dòng)相關(guān)生理生化反應(yīng)[30]。如異色瓢蟲經(jīng)冷馴化處理后體內(nèi)細(xì)胞保護(hù)酶超氧化物歧化酶(SOD)與過(guò)氧化氫酶(CAT)活性升高,與新陳代謝有關(guān)的乳酸脫氫酶(LDH)及Na+,K+-ATP酶活性降低[31]。

2 昆蟲耐寒性的分子機(jī)制

2.1 抗冷保護(hù)劑基因調(diào)控

2.1.1 脂類

脂肪酸脫飽和酶(fatty acid desaturase, FAD)是一類在脂肪酸鏈上將碳碳單鍵轉(zhuǎn)化為碳碳雙鍵的酶,是合成不飽和脂肪酸的關(guān)鍵酶[32]。昆蟲中FAD保幼激素的種類很多,但主要屬于FAD9(△9-?；o酶脫飽和酶)和FAD11(△11-?；o酶脫飽和酶)[33-34]。在魚類[35]、細(xì)菌[36]、植物[37]和昆蟲[38]的耐低溫研究中均證實(shí)了細(xì)胞膜上飽和脂肪酸轉(zhuǎn)變成不飽和脂肪酸時(shí),FAD9基因起到了非常關(guān)鍵的作用。關(guān)于FAD9基因序列已經(jīng)在黑腹果蠅[39]、粉紋夜蛾(Trichoplusiani)[40]、家蠶(Bombyxmori)[41]和家蠅(Muscadomestica)[42]中得到鑒定,序列在兩端保守性低,但中間部分存在兩個(gè)相對(duì)保守的疏水區(qū),富含3個(gè)極保守的組氨酸基序和4 次跨膜結(jié)構(gòu)域,這3個(gè)極保守的組氨酸基序與酶活性中心的形成相關(guān),主要是螯合金屬離子,結(jié)合氧[43]。

Kayukawa等[38]在蔥地種蠅(Deliaantiqua)中最先證實(shí)了FAD9基因與耐寒相關(guān)。隨后,又有學(xué)者陸續(xù)在白紋伊蚊(Aedesalbopictus)、麻蠅(Sarcophagacrassipalpis)等幾種昆蟲證實(shí)耐寒與FAD9基因有關(guān)[44-45]。當(dāng)溫度降低時(shí)膜脂流動(dòng)性減少,該信號(hào)激活FAD9合成不飽和脂肪酸用來(lái)提高昆蟲耐寒性。FAD9在低溫下有兩種調(diào)節(jié)機(jī)制[35]:其一,當(dāng)溫度開始下降時(shí),激活昆蟲體內(nèi)現(xiàn)有的FAD9基因,增加其活性,但表達(dá)量不變;其二,當(dāng)溫度繼續(xù)下降時(shí),FAD9基因表達(dá)量升高,酶活性也進(jìn)一步增加。關(guān)于FAD調(diào)控脂肪酸代謝的機(jī)制至今研究還不夠深入,有待進(jìn)一步完善。

2.1.2 甘油激酶

在低溫狀態(tài)時(shí),昆蟲合成多元醇等抗凍保護(hù)劑阻止細(xì)胞結(jié)冰,增加其適應(yīng)性和生存能力。甘油是昆蟲血淋巴中主要的抗凍保護(hù)劑之一[46],具有兩個(gè)獨(dú)立的合成途徑:一個(gè)是利用多元醇脫氫酶催化甘油醛和煙酰胺腺嘌呤二核苷磷酸(NADPH+H+)途徑合成甘油[47];另一個(gè)是利用甘油磷酸脫氫酶(GPDH)和葡萄糖激酶(GK)途徑合成甘油[48]。小菜蛾通過(guò)后者完成甘油合成,其基因組注釋中有4個(gè)葡萄糖激酶(GK)基因和1個(gè)甘油磷酸脫氫酶(GPDH)基因[49],經(jīng)RNAi驗(yàn)證GK1與耐寒相關(guān)[50]。GPDH可能在耐寒過(guò)程中起輔助作用,催化合成3-磷酸甘油[51],這一假設(shè)需要在其他昆蟲中進(jìn)一步研究證明。

2.1.3 海藻糖

海藻糖也是昆蟲血淋巴中主要的抗凍保護(hù)劑之一,由2個(gè)葡萄糖殘基通過(guò)糖苷鍵結(jié)合而成[52]。海藻糖的合成途徑有5 種,最主要的是TPS/TPP途徑。這個(gè)途徑中,海藻糖-6-磷酸合成酶起關(guān)鍵作用,即尿苷二磷酸葡萄糖和6-磷酸葡萄糖在海藻糖6-磷酸合成酶的催化作用下合成6-磷酸海藻糖,然后在海藻糖6-磷酸磷酸酯酶作用下生成海藻糖[53]。

隨著季節(jié)變化,昆蟲體內(nèi)海藻糖在血淋巴中的含量也隨之變化,秋季增加,冬季結(jié)束后下降[54]。同樣,這一特性在低溫條件或昆蟲冬滯育中也存在[55]。完全處于冬滯育狀態(tài)的蘋果蠹蛾(Cydiapomonella)幼蟲體內(nèi)海藻糖含量比滯育初期的高3倍,而且海藻糖濃度與蘋果蠹蛾幼蟲過(guò)冷卻能力和低溫生存能力相關(guān)[56]。異色瓢蟲在低溫誘導(dǎo)下,TPS的表達(dá)量隨著溫度的降低而顯著升高,在降溫和升溫處理?xiàng)l件下,TPS的表達(dá)量呈現(xiàn)先升高后下降的表達(dá)趨勢(shì)[57]。進(jìn)一步說(shuō)明TPS基因的表達(dá)與耐寒相關(guān)。這一機(jī)制的研究為明確昆蟲的耐寒途徑提供了新的思路。

2.2 抗凍蛋白和熱滯活性

抗凍蛋白是一類具有熱滯活性,吸附在冰核表面抑制冰核增長(zhǎng)的冰結(jié)合蛋白,能夠?qū)鼋Y(jié)溫度降到熔點(diǎn)之下,這一現(xiàn)象稱作熱滯作用?？箖龅鞍啄軐⒗ハx血淋巴的冰點(diǎn)降低超過(guò)5℃[58]。同時(shí),抗凍蛋白還可改變冰晶形態(tài)、降低過(guò)冷卻點(diǎn)、降低玻璃化和去玻璃化損傷,具有明顯的季節(jié)性[59]。目前,至少有50種昆蟲產(chǎn)生抗凍蛋白[60]。昆蟲抗凍蛋白的分子量一般約為8～9 kD,無(wú)糖基化位點(diǎn)并且含有較多的親水性氨基酸,有40%～50%的氨基酸殘基能形成氫鍵。由于一些昆蟲的抗凍蛋白相比魚類,具有更強(qiáng)的抗凍能力[61],因此,昆蟲抗凍蛋白的研究成為生物技術(shù)的熱門話題。

目前,盡管大部分昆蟲抗凍蛋白研究集中在探索和測(cè)量血淋巴中熱滯活性,但是也有少量昆蟲的抗凍蛋白序列被測(cè)序并純化,如:黃粉蟲(Tenebriomolitor)、赤翅甲(Dendroidescanadensis)、雪跳蚤(Hypogastruraharveyi)、樅色卷蛾(Choristoneurafumiferana)等[62]。比較這4種昆蟲的抗凍蛋白序列,發(fā)現(xiàn)它們之間相似度非常低,表明昆蟲抗凍蛋白之間由不同始祖獨(dú)立進(jìn)化而來(lái),至少包含17個(gè)cf型亞族和13個(gè)D型亞族。同一基因不同的亞型多拷貝現(xiàn)象存在明顯的適應(yīng)性,因此,隨著環(huán)境信號(hào)的改變,這些蛋白可能發(fā)揮不同的功能[63]。將抗凍蛋白轉(zhuǎn)入昆蟲細(xì)胞系中進(jìn)行表達(dá),可以控制神經(jīng)內(nèi)分泌,使生物體對(duì)環(huán)境因素誘導(dǎo)產(chǎn)生不敏感的反應(yīng)[64]。也有研究表明,有些生物體轉(zhuǎn)入抗凍蛋白基因后,提高了熱滯活性,增強(qiáng)了抗凍能力;然而,也有些生物體轉(zhuǎn)入抗凍蛋白基因后,雖然提高了熱滯活性,但是抗凍能力并無(wú)明顯差異[65]。可能存在如下原因[66-67]:(1)抗凍蛋白是多基因家族,熱滯活性不可能區(qū)分不同亞型的抗凍蛋白,不同形式抗凍蛋白亞型混合,產(chǎn)生了耐寒性;(2)細(xì)胞膜的穩(wěn)定性對(duì)抗寒非常重要,即使轉(zhuǎn)基因抗凍蛋白集中在細(xì)胞膜上且能正確翻譯與修飾,但它們不可能無(wú)限制地穩(wěn)定細(xì)胞膜;(3)穩(wěn)定細(xì)胞膜抵抗低溫需要一些協(xié)同作用的物質(zhì),如甘油。因此,昆蟲體內(nèi)抗凍蛋白的產(chǎn)生與抵抗低溫的關(guān)系非常復(fù)雜,需要通過(guò)現(xiàn)代分子生物學(xué)手段進(jìn)行深入研究。

2.3 熱休克蛋白

熱休克蛋白(heat shock proteins, HSPs)是一種抗逆蛋白,通常被認(rèn)為具有保護(hù)細(xì)胞不受傷害的作用。在大部分昆蟲中,冷休克可以誘導(dǎo)熱休克蛋白家族基因的表達(dá),但在果蠅中,不管是冷脅迫還是快速冷馴化,HSP70的表達(dá)幾乎恒定不變[68]。因此,在早期昆蟲耐寒性研究中,黑腹果蠅不是理想的模式昆蟲,HSP70不隨著冷休克而上調(diào),但這并不意味其他熱休克蛋白基因也不變化,特別是小熱休克蛋白基因家族,這一觀念隨著后來(lái)研究的深入得到了證實(shí)[69],如HSP19.5,HSP20.8和HSP21.7在美洲斑潛蠅(Liriomyzasativae)耐寒中起到非常重要的作用,其中HSP20.8對(duì)低溫更敏感[70]。

冬滯育在昆蟲中普遍存在,其研究主要集中在熱休克蛋白基因[71]。與滯育相關(guān)的基因有兩種[72]:一種是直接調(diào)控滯育;另一種是輔助啟動(dòng)滯育機(jī)制。不同物種之間調(diào)控滯育的基因不同,如HSP90在麻蠅(S.crassipalpis)滯育中表達(dá)上調(diào)[73],但在蔥地種蠅(Deliaantiqua)滯育中表達(dá)下調(diào)[74],說(shuō)明不同種類的昆蟲參與耐寒調(diào)控的熱休克蛋白并非一致。

熱休克蛋白基因家族與滯育的關(guān)系非常復(fù)雜,主要受外部環(huán)境的刺激影響。麻蠅(S.crassipalpis)通過(guò)RNAi技術(shù)干擾HSP23和HSP70基因,并不影響其滯育,但影響蛹的耐寒性[75]。因此,HSP23和HSP70被認(rèn)為主要與麻蠅越冬耐寒相關(guān)。

目前,確定與滯育觸發(fā)相關(guān)的基因有兩類[76]。一類是滯育激素,不同物種之間表現(xiàn)并非一致,滯育激素在家蠶中誘導(dǎo)胚胎滯育[77],然而,在煙芽夜蛾(Heliothisvirescens)[78]和棉鈴蟲(Helicoverpaarmigera)[79]中解除滯育。另一類是P38 MAPK的磷酸化作用,在豆長(zhǎng)刺螢葉甲(Atrachyamenetriesi)[80]、家蠶(Bombyxmori)[81]和麻蠅(Sarcophagacrassipalpis)[82]等3種昆蟲的滯育與解除中得到證實(shí)。p38 MAPK的磷酸化作用在誘導(dǎo)麻蠅進(jìn)入快速冷馴化過(guò)程中起著關(guān)鍵作用,溫度在0℃ 左右10 min內(nèi)啟動(dòng)p38 MAPK磷酸化途徑[83]。

2.4 水通道蛋白

隨著分子生物學(xué)高科技手段的發(fā)展與運(yùn)用,生物體中仍然存在一些特殊的基因在耐寒中起著重要作用,如水通道蛋白就是其中之一。水通道蛋白是跨膜蛋白,可以選擇性地讓水或其他小分子物質(zhì)通過(guò)細(xì)胞膜。目前,水通道蛋白在黑腹果蠅[84]、白紋伊蚊(A.albopictus)[85]、岡比亞按蚊(Anophelesgambiae)[86]和Eurostasolidaginis[87]等昆蟲中得到克隆并鑒定。通過(guò)免疫印跡法得知E.solidaginis水通道蛋白與哺乳動(dòng)物同源,水通道蛋白的表達(dá)量與氣候溫度和干燥直接相關(guān),抑制水通道蛋白活性可以明顯降低E.solidaginis脂肪體和中腸細(xì)胞在低溫下的存活[87]。由于水通道蛋白具有控制水分和甘油流動(dòng),增加滲透調(diào)節(jié)和代謝運(yùn)輸?shù)墓δ?因此,可以推測(cè)水通道蛋白與耐寒性存在一定的關(guān)系,但需要進(jìn)一步研究證實(shí)。

3 展望

昆蟲受低溫脅迫時(shí),會(huì)啟動(dòng)一系列抗寒機(jī)制。通過(guò)調(diào)控相關(guān)抗寒基因,合成抗逆物質(zhì),增加細(xì)胞內(nèi)抗凍蛋白和熱休克蛋白等提高抗寒能力,保護(hù)機(jī)體不受低溫的傷害。同時(shí),這一生理生化過(guò)程隨著溫度的恢復(fù)是可逆的。

昆蟲耐寒性研究是昆蟲生態(tài)學(xué)和生物進(jìn)化學(xué)研究的熱點(diǎn),也為害蟲的測(cè)報(bào)和防治提供理論基礎(chǔ)[88],尤其對(duì)新農(nóng)藥的開發(fā)提供了新的思路。

昆蟲脂肪酸脫飽和酶在低溫下促進(jìn)不飽和脂肪酸的形成,減少膜脂流動(dòng)性,提高耐寒性。因此,可以通過(guò)基因工程等相關(guān)手段開發(fā)新的生物制劑,阻斷害蟲體內(nèi)脂肪酸脫飽和酶的活性,降低害蟲耐寒能力,讓其不能在低溫條件下越冬,達(dá)到害蟲防治的效果。同時(shí),對(duì)于有益生物,也可以通過(guò)此方法保護(hù)其耐寒越冬。這一研究為今后的生產(chǎn)實(shí)踐提供了廣闊的思路。

昆蟲體內(nèi)冰核物質(zhì)能夠提高昆蟲過(guò)冷卻點(diǎn),降低昆蟲耐寒性,導(dǎo)致越冬昆蟲大量死亡。若開發(fā)成新型的生物制劑,與其他防治方法配合使用,可能開辟一個(gè)全新的害蟲防治領(lǐng)域,具有潛在的生態(tài)學(xué)意義。

總之,隨著基因芯片、蛋白質(zhì)組學(xué)、代謝組學(xué)和基因組學(xué)等的發(fā)展與應(yīng)用,昆蟲耐寒的生理生化機(jī)制逐漸清晰。盡管果蠅不適合作為耐寒研究的模式昆蟲,但果蠅有完整的基因組序列、成熟的育種系、突變體和細(xì)胞系。根據(jù)果蠅的基因功能,推測(cè)其他昆蟲某類基因的功能,再進(jìn)行試驗(yàn)證明。研究昆蟲的耐寒性機(jī)制,可為農(nóng)業(yè)昆蟲的測(cè)報(bào)提供理論參考,尤其對(duì)天敵昆蟲的低溫儲(chǔ)存具有重要意義,同時(shí),在新農(nóng)藥開發(fā)與利用方面具有較大前景。這些研究有望推動(dòng)和促進(jìn)害蟲農(nóng)業(yè)防治和生物防治事業(yè)的發(fā)展。

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(責(zé)任編輯:田 喆)

Research progress in the cold tolerance mechanism of insects under environmental stress

Shi Caihua1,2, Hu Jingrong1, Li Chuanren1, Zhang Youjun2

(1. Yangtze University, Jingzhou 434025, China; 2. Institute of Vegetables and Flowers,Chinese Academy of Agricultural Sciences, Beijing 100081, China)

As ectothermic animals, insects usually convert its structure and physiological material composition in order to survive safely in winter. This adaptation mechanism is closely related to the special physiological and biochemical substances in the body, such as trehalose, dextrose, glycerol, sorbitol, fatty acids, amino acids and other frost protection agents of small molecules. These antifreeze agents have the function of stabilizing the cell membrane structure and protecting the protein function. Currently, research on the cold tolerance of insects is deepening, but it is still difficult to determine the key factors that affect the cold tolerance. Why some insects can survive below -20℃? In order to clarify the scientific issues, a large number of scholars have used the theory of transcriptomics, genomics, proteomics and metabolomics to analyze the molecular mechanisms of insect physiological responses after low temperature stress. The purpose of this article is to review the cold tolerance aspects of previous studies and provide a reference for future studies of other insects or cold flora and fauna. At the same time, it also provides a broad way for the development of new bio-pesticides and increasing natural enemies of insects artificially.

insect; cold tolerance; frost protection agent; mechanism

Reviews

2016-02-16

2016-05-05

公益性行業(yè)(農(nóng)業(yè))科研專項(xiàng)(2013203027)

Q 965

A

10.3969/j.issn.0529-1542.2016.06.003

* 通信作者 E-mail:zhangyoujun@caas.cn