豌豆蚜生物生態(tài)學(xué)特性

張 麗，袁明龍

(1.草地農(nóng)業(yè)生態(tài)系統(tǒng)國家重點實驗室，蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020;2.草業(yè)科學(xué)國家級實驗教學(xué)示范中心(蘭州大學(xué)),甘肅 蘭州 730020; 3.農(nóng)業(yè)部草牧業(yè)創(chuàng)新重點實驗室，甘肅 蘭州 730020)

豌豆蚜生物生態(tài)學(xué)特性

張 麗1,2,3，袁明龍1,2,3

(1.草地農(nóng)業(yè)生態(tài)系統(tǒng)國家重點實驗室，蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020;2.草業(yè)科學(xué)國家級實驗教學(xué)示范中心(蘭州大學(xué)),甘肅 蘭州 730020; 3.農(nóng)業(yè)部草牧業(yè)創(chuàng)新重點實驗室，甘肅 蘭州 730020)

豌豆蚜(Acyrthosiphonpisum)是許多豆科作物及牧草的重要害蟲，不但直接取食寄主植物，造成嚴(yán)重的經(jīng)濟損失，而且還會傳播多種植物病毒。豌豆蚜作為生態(tài)學(xué)研究的模式昆蟲，具有復(fù)雜的生活周期、多樣的生殖方式、表型可塑性以及與細(xì)菌復(fù)雜的共生關(guān)系等諸多特點，已成為眾多科學(xué)家感興趣的研究對象。本文全面總結(jié)了豌豆蚜生物生態(tài)學(xué)特性的研究成果，重點綜述了豌豆蚜的多型現(xiàn)象和內(nèi)共生菌研究的最新進(jìn)展，提出研究豌豆蚜在全球氣候和作物種植結(jié)構(gòu)變化下生物生態(tài)學(xué)特性演變規(guī)律的必要性，加強其生態(tài)適應(yīng)性及多型現(xiàn)象等遺傳機制研究，以期為建立該蟲的可持續(xù)防控技術(shù)體系提供依據(jù)。

蚜蟲；生物學(xué)特性；表型可塑性；孤雌生殖；生態(tài)適應(yīng)性

蚜蟲是半翅目昆蟲中較大的一個類群，目前全世界已記錄4 700余種，其中我國約有1 000種，主要分布在北半球溫帶地區(qū)和亞熱帶地區(qū)[1-4]。許多蚜蟲是重要的農(nóng)林牧業(yè)害蟲，如麥長管蚜(Macrosiphumavenae)[5]、棉蚜(Aphisgossypii)[6-7]、桃蚜(Myzuspersicae)[8]和豌豆蚜(Acyrthosiphonpisum)[9-12]等。蚜蟲以刺吸式口器吸食植物韌皮部汁液，使植物營養(yǎng)損失，發(fā)育受阻，并出現(xiàn)畸形生長、早衰，甚至死亡的現(xiàn)象[13-14]；同時，許多蚜蟲還是一些重要植物病毒的傳播媒介[15]。此外，蚜蟲具有復(fù)雜的生活史和生殖方式[16]，高度表型可塑性[11-12,17]，這些特點使其成為眾多昆蟲學(xué)家、生態(tài)學(xué)家和進(jìn)化生物學(xué)家最為感興趣的研究對象之一[2,9,11,18]。

豌豆蚜又稱豆蚜、豆無網(wǎng)長管蚜，屬于蚜總科的蚜科(Aphididae)，該蟲在世界各地廣泛分布，是許多豆科作物及牧草的主要害蟲之一[19]。在美國，豌豆蚜的危害可導(dǎo)致苜蓿生產(chǎn)者每年損失大約6 000萬美元[20-21]。在我國西北苜蓿種植區(qū)，豌豆蚜每年可造成苜蓿生產(chǎn)10%～30%的經(jīng)濟損失[13,22]。豌豆蚜危害植物時，其若蟲和成蟲群集在植物的幼嫩部位，以刺吸式口器吸取植物韌皮部汁液，從而影響植物的生長發(fā)育、開花結(jié)實等正常的生命活動，嚴(yán)重危害時甚至造成整塊田間植株死亡[14,23]。此外，豌豆蚜還是苜蓿(Medicagosativa)花葉病毒、豌豆(Pisumsativum)耳突花葉病毒等25種病毒的主要傳播者[15,24-26]。由于豌豆蚜具有生活周期復(fù)雜、生殖方式多樣、表型可塑性以及與細(xì)菌復(fù)雜的共生關(guān)系[12,16-17,27-28]等特點，使其成為國際蚜蟲基因組聯(lián)盟科學(xué)家青睞的研究對象。2010年科學(xué)家采用鳥槍法成功完成了豌豆蚜全基因組序列的測定工作，使其成為首個被破譯基因組的蚜蟲，也是第一個被測序的半翅目昆蟲[29]。近年來，豌豆蚜的相關(guān)研究已逐漸深入到了組學(xué)水平，使其成為遺傳學(xué)、生態(tài)學(xué)及進(jìn)化生物學(xué)研究的模式物種[17,30-33]。

豌豆蚜在我國各地均有發(fā)生，且作為模式昆蟲具有重要的研究價值，明確豌豆蚜的生活史、發(fā)生規(guī)律、危害性及生態(tài)適應(yīng)性等基礎(chǔ)生物生態(tài)學(xué)特性，是有效開展其預(yù)測預(yù)報及防治的前提和基礎(chǔ)性工作。為此，本文系統(tǒng)總結(jié)豌豆蚜生物生態(tài)學(xué)的研究現(xiàn)狀，深入分析已有研究成果，并重點討論今后應(yīng)開展的研究方向，以期為豌豆蚜的深入研究及有效防控提供理論支持。

1 生物學(xué)特性

1.1 世代及年生活史

豌豆蚜屬于漸變態(tài)類昆蟲，包括卵、若蟲和成蟲3個發(fā)育階段。該蟲一年能繁殖十多代，世代重疊現(xiàn)象嚴(yán)重[16]。豌豆蚜的發(fā)育歷期在不同地區(qū)、不同氣候條件下差異較大，一般隨溫度的升高而縮短[16,22]。田間條件下，平均氣溫在12 ℃以上持續(xù)5 d時，豌豆蚜便可開始繁殖[34]；而在溫室栽培的條件下，全年均可繁殖危害[35]。但不論一年發(fā)生幾代，均以卵的形式越冬。豌豆蚜通常產(chǎn)卵于植物根部，卵一般都單個分布，數(shù)量多時也成群成堆。卵大多呈長橢圓形，初生時為黃色，然后逐漸變?yōu)榫G色，最后成為光亮黑色[16]。豌豆蚜在北方地區(qū)一般11月份產(chǎn)生兩性蚜，交尾后產(chǎn)卵于多年生豆科植物上越冬(圖1)，而溫暖地區(qū)或溫室內(nèi)全年不產(chǎn)生兩性蚜[34,36]。確定豌豆蚜的越冬場所并調(diào)查越冬卵基數(shù)，有利于預(yù)測來年豌豆蚜的發(fā)生數(shù)量并指導(dǎo)有效防控。

圖1 豌豆蚜生活史Fig. 1 Life cycle of the pea aphid

1.2 寄主及取食行為

豌豆蚜具有較為廣泛的寄主植物，不同的寄主植物對豌豆蚜的生長發(fā)育和繁殖均有影響。例如，蠶豆(Viciafaba)和豌豆較紅豆草(Onobrychisviciaefolia)更適合豌豆蚜的生長發(fā)育及繁殖[37-38]。豌豆蚜取食蠶豆的不同品種，其在發(fā)育歷期、存活率以及生殖力方面均存在差異[38]。寄主植物不僅影響豌豆蚜的存活率和繁殖力，而且會影響豌豆蚜的種群結(jié)構(gòu)和遺傳多樣性[39]。同時寄主植物的干旱脅迫也能夠影響豌豆蚜的習(xí)性和其在植物上的豐富度[40]。

豌豆蚜擁有強烈的寄主?；訹41-42]，但這種特性因地而異。例如，在紐約豌豆蚜寄主為苜蓿或紅三葉草，在加利福尼亞其寄主則主要為白三葉草(Trifoliumrepens)，我國分布的豌豆蚜則在苜蓿及豆科作物上取食[22,43-44]。研究表明，豌豆蚜的寄主?；杂衅湎鄳?yīng)的遺傳學(xué)基礎(chǔ)。例如，豌豆蚜當(dāng)?shù)睾鸵M(jìn)種群均表現(xiàn)出低水平的核苷酸多樣性和中低水平的連鎖不平衡[45-46]。取食不同植物的豌豆蚜，決定其寄主植物利用和蚜蟲交配選擇的數(shù)量性狀基因緊密連鎖，從而能夠促進(jìn)寄主專化和蚜蟲生殖隔離的共同進(jìn)化[9]。此外，用不同的固醇類和植物飼養(yǎng)的豌豆蚜，其基因表達(dá)存在差異性[47]。通過豌豆蚜唾液腺轉(zhuǎn)錄組基因的表達(dá)差異研究，揭示了豌豆蚜對不同寄主植物適應(yīng)性差異的分子機理[48]。

蚜蟲以刺吸式口器吸取植物韌皮部汁液來獲得營養(yǎng)，而植物韌皮部汁液富含昆蟲喜食的糖類物質(zhì)[49]。通常，豌豆蚜喜在植物莖和頂部嫩葉上取食，對植物的生長發(fā)育具有重要的影響[19]。豌豆蚜取食紫花苜蓿的生長區(qū)時，植物的拔節(jié)率顯著降低，而當(dāng)取食成熟區(qū)時拔節(jié)率則無顯著變化[50]。豌豆蚜對植物的取食作用還影響植物的同化物分配，而豌豆蚜可通過改變自身的營養(yǎng)成分結(jié)構(gòu)來適應(yīng)植物中營養(yǎng)物質(zhì)的變化，從而適應(yīng)新生境[34,50]。

1.3 生殖活動

豌豆蚜是典型的世代交替昆蟲，即在食物充足、環(huán)境良好的情況下營孤雌生殖，而在環(huán)境惡化和食物匱乏時進(jìn)行有性生殖(圖1)。豌豆蚜的越冬卵通常小于1 mm[16]，而其孤雌生殖的胚胎較有性生殖的卵更小[51]。在孤雌生殖的分娩期間，從母體到第一齡若蟲需約15 min[52]。豌豆蚜在適溫下，卵孵化后經(jīng)過約7 d即可發(fā)育成成蟲，體長達(dá)4～9 mm[2,22]。豌豆蚜雌成蟲繁殖能力強，在適溫下單雌產(chǎn)蚜量可高達(dá)138頭，生殖期長達(dá)2～3個月。豌豆蚜在大約3/4的生活史中營無性生殖，易于近親繁殖；但因其具有識別種群中近親的能力，所以并未出現(xiàn)種群衰退現(xiàn)象，然而體色識別并非其避免近親繁殖的有效策略[53]。

豌豆蚜的生殖受光照、溫度、寄主植物狀態(tài)等外界因素影響，外界條件的變化可使其在孤雌生殖和有性生殖之間發(fā)生轉(zhuǎn)換[16,29]。豌豆蚜在夏天采取孤雌生殖和胎生的方式進(jìn)行繁殖，而夏末會進(jìn)行有性繁殖，并在夏季結(jié)束前產(chǎn)生耐寒的越冬卵[54]?？梢姡愣寡猎诠麓粕诚蛴行陨侈D(zhuǎn)變的季節(jié)性調(diào)整中，日長和溫度起到了關(guān)鍵作用，但日長和溫度影響豌豆蚜孤雌生殖和有性生殖之間相互轉(zhuǎn)變的詳細(xì)機制還未被深入研究。

1.4 多型現(xiàn)象

多型現(xiàn)象在昆蟲中廣泛存在，主要有體色多型、翅多型等。豌豆蚜具有體色多態(tài)性，分為紅、綠兩種色型(圖2)，研究報道較多的是綠色型[12,55-56]。1945年，Harrington首先報道了紅色型豌豆蚜[57]。在我國，賀春貴[19]于2004年首次發(fā)現(xiàn)并記載了豌豆蚜的紅色型。在對甘肅蘭州苜蓿田的調(diào)查中發(fā)現(xiàn)，紅色型豌豆蚜所占比例逐年上升，有超過綠色型的趨勢[58]。

豌豆蚜體色變化受自身基因型和外界環(huán)境因素的共同調(diào)控。Caillaud和Losey[28]研究表明，蚜蟲的體色是由單一的等位基因控制，紅色型則由占優(yōu)勢地位的顯性基因控制。進(jìn)一步研究表明，紅色型豌豆蚜擁有合成類胡蘿卜素的相關(guān)基因，該基因具有編碼類胡蘿卜素脫氫酶的功能，而這些基因來源于真菌基因的橫向轉(zhuǎn)移[11]。蚜蟲內(nèi)共生菌同樣也參與豌豆蚜的體色多態(tài)型的變化。研究表明，兼性共生菌立克次氏小體(Rickettsia)侵染可使紅色型豌豆蚜向綠色型轉(zhuǎn)變[18]。雖然共生菌是否通過調(diào)控編碼類胡蘿卜素合成酶的基因來影響豌豆蚜的體色尚不可知，但可以確定的是有某種共生菌參與了豌豆蚜的體色分化。環(huán)境條件可影響蚜蟲的體色，當(dāng)在寒冷的環(huán)境條件下，蚜蟲體色為綠色；在適宜的環(huán)境條件下，為橙色蚜蟲，而當(dāng)蚜蟲種群數(shù)量增多食物資源匱乏時，則出現(xiàn)白色蚜蟲[13,59]。高溫和低溫都可誘導(dǎo)蚜蟲體色變化：豌豆蚜在8 ℃的低溫條件下幾乎轉(zhuǎn)變?yōu)榫G色，而高溫誘導(dǎo)可使麥長管蚜產(chǎn)生紅色型，暗示豌豆蚜體色多態(tài)性可能與溫度以及季節(jié)變化有關(guān)[60-62]。也有研究表明，寄主?；砸部赡苁钦T導(dǎo)豌豆蚜體色多態(tài)性的重要原因之一[63]。

圖2 豌豆蚜紅色型和綠色型Fig. 2 The red and green morphs of the pea aphid

豌豆蚜不同體色的個體在應(yīng)對天敵等外界環(huán)境因素脅迫的能力方面存在差異[64-66]。生態(tài)學(xué)研究表明，捕食性天敵更易捕食在綠色植物上的紅色型豌豆蚜，而寄生性天敵卻優(yōu)先攻擊綠色型[67-68]。兩種色型的豌豆蚜對于紫外線脅迫的反應(yīng)也不盡相同，當(dāng)采用紫外線對豌豆蚜進(jìn)行12 h輻射脅迫時，與綠色型蚜蟲相比紅色型蚜蟲的生長發(fā)育明顯遲緩[69]。當(dāng)用殺蟲劑吡蟲啉處理蚜蟲種群后，紅色型豌豆蚜較綠色型表現(xiàn)出更高的敏感性[70]。此外，在寄主植物適應(yīng)性方面，豌豆蚜不同體色個體間存在差異，與綠色型相比紅色型具有更寬的食物生態(tài)幅，導(dǎo)致其在面對抗性植物時具有較強的生活力[71-72]。不同體色豌豆蚜體內(nèi)主要營養(yǎng)物質(zhì)含量有所不同。例如，紅色型體內(nèi)含有較多的碳水化合物和脂類，而綠色型體內(nèi)蛋白質(zhì)含量較高，所以當(dāng)其均處于惡劣的環(huán)境條件下時，紅色型個體則具有足夠的能量產(chǎn)生有翅蚜進(jìn)行遷飛從而逃避惡劣環(huán)境[64,73]。不同色型間所表現(xiàn)出的不同生物學(xué)特性，是豌豆蚜對整個生活史進(jìn)行權(quán)衡后所呈現(xiàn)的一些適應(yīng)性特征，利于提高其在自然環(huán)境、特別是逆境中的適合度[74]。

豌豆蚜是典型的翅多型昆蟲。豌豆蚜孤雌生殖會產(chǎn)生翅二型后代，該后代可發(fā)育成有翅或無翅成蚜。有翅豌豆蚜的發(fā)生會直接影響到天敵的釋放效率，因此深入研究其翅多型產(chǎn)生的原因?qū)υ撓x的生物防治具有十分重要的作用。豌豆蚜有翅和無翅型的形成是由環(huán)境條件和豌豆蚜自身基因型共同決定的[75]，其中種群密度和天敵是最為關(guān)鍵的兩個外界因素。當(dāng)種群密度大時，產(chǎn)生有利于遷飛到其它寄主植物上取食的有翅蚜，有翅蚜的轉(zhuǎn)移危害有利于物種間的基因交流，同時可形成新的生物型并危害新的寄主植物，進(jìn)而在一定程度上增加了寄主范圍[76]。天敵因素也可誘導(dǎo)有翅蚜的產(chǎn)生[77]。天敵的存在會使豌豆蚜釋放一些可誘導(dǎo)翅型改變的信息素，如警報信息素等[78-80]。有翅蚜的出現(xiàn)同時受溫度和光周期的影響，豌豆蚜在不同的溫度和光照組合下會產(chǎn)生有翅和無翅孤雌蚜、雄蚜和產(chǎn)卵雌蚜[81]。目前，有關(guān)豌豆蚜兩種翅型間轉(zhuǎn)換的分子機制研究已逐漸開展。對豌豆蚜翅多型遺傳基礎(chǔ)的研究發(fā)現(xiàn)，雄性豌豆蚜中與翅型相關(guān)的基因位點是伴隨性別決定來進(jìn)行調(diào)控的[82]。而誘導(dǎo)翅型發(fā)育基因的位置不僅影響雄性個體的翅型，同樣也與雌性翅的非遺傳多型性相關(guān)[76]。繼張傳溪團(tuán)隊揭示胰島素信號通路中的胰島素受體在褐飛虱(Nilaparvatalugens)翅型形成的分子機制中發(fā)揮開關(guān)式的調(diào)節(jié)作用后[83]，Guo等[84]首次對豌豆蚜胰島素信號通路中相關(guān)基因的功能進(jìn)行了研究，發(fā)現(xiàn)胰島素信號通路中的Apirp5參與了翅二型豌豆蚜的胚胎發(fā)育和代謝調(diào)節(jié)。

翅多型現(xiàn)象是豌豆蚜對生存環(huán)境的一種適應(yīng)性表現(xiàn)。例如，豌豆蚜無翅個體的卵期及世代時間明顯短于有翅個體，且擁有更高的繁殖力[75,85]。豌豆蚜的翅型同樣也會影響其交配行為和繁殖率，在一對一的交配競爭中有翅豌豆蚜?xí)@得較多的交配對象，而在無競爭狀態(tài)下，無翅雄性一生中的繁殖成功率比有翅雄性稍高[86]。相對于雌性，有翅和無翅的雄性在形態(tài)學(xué)上并不明顯，這可能是由于雄性很少在生殖和傳播間進(jìn)行權(quán)衡所致[87]。

1.5 內(nèi)共生菌

在長期協(xié)同進(jìn)化過程中，蚜蟲與其體內(nèi)共生菌形成了復(fù)雜的共生關(guān)系，共生菌與蚜蟲之間密切互作、協(xié)同進(jìn)化[88]。幾乎所有的蚜蟲都含有內(nèi)共生菌，這些細(xì)菌為同一屬巴克納氏菌(Buchnera)，稱為專性共生菌(obligate symbiont)。豌豆蚜和Buchnera的共生關(guān)系可追溯到25億年以前[89]。共生菌位于豌豆蚜腹部的含菌胞內(nèi)，這些菌胞很少分裂，但會隨著蚜蟲的生長而逐漸變大[90-91]。共生菌的數(shù)量在豌豆蚜不同生長階段有所不同；電鏡觀察發(fā)現(xiàn)，豌豆蚜Buchnera數(shù)量在胚胎菌胞的形成過程中數(shù)量增加，而在卵的形成過程中數(shù)量減少[92]。內(nèi)共生菌在蚜蟲寄主內(nèi)是通過卵和孤雌胚胎進(jìn)行垂直傳播的[93]；在進(jìn)行有性生殖時，將傳遞給卵細(xì)胞，而孤雌生殖時則傳遞給胚胎[94]。

內(nèi)共生關(guān)系一直是生物學(xué)研究的熱點，而豌豆蚜和內(nèi)共生菌Buchnera的專性共生關(guān)系使其成為內(nèi)共生關(guān)系研究的理想模型。隨著豌豆蚜及其內(nèi)共生菌Buchnera全基因組測序的完成[10,29]，目前研究主要側(cè)重于依賴生理和基因組數(shù)據(jù)的Buchnera-蚜蟲的進(jìn)化關(guān)系以及兩者間的營養(yǎng)代謝關(guān)系[95-96]。盡管有研究稱在低級階元水平上Buchnera-蚜蟲符合平行分化關(guān)系，但在高級階元水平上卻缺乏嚴(yán)格的對應(yīng)關(guān)系[95]。同時又有人推測，Buchnera基因組的進(jìn)化效率和橫向轉(zhuǎn)移可能導(dǎo)致Buchnera-蚜蟲在高低階元間不同的發(fā)育格局[97]。然而關(guān)于兩者的進(jìn)化關(guān)系仍不清楚，大多數(shù)研究關(guān)注Buchnera-蚜蟲的營養(yǎng)代謝關(guān)系。豌豆蚜與其共生菌共同參與某些必需氨基酸的生物合成，且宿主蚜蟲能精確地調(diào)控Buchnera的新陳代謝[98]。專性共生菌Buchnera具有維生素合成通路，可為蚜蟲提供所需維生素和腺嘌呤，并同時利用蚜蟲產(chǎn)生的鳥嘌呤來滿足自身對于核苷酸的需求[99-100]。

豌豆蚜除含有專性內(nèi)共生菌外，其體內(nèi)還含有多種其它細(xì)菌類群,稱為次級共生菌，又稱兼性共生菌(facultative symbiont)。兼性共生菌是指部分蚜蟲體內(nèi)含有而另一部分蚜蟲體內(nèi)不含有的一類共生菌。迄今，有關(guān)蚜蟲兼性共生菌的研究中，豌豆蚜內(nèi)共生菌的研究較為深入，目前已報道7種不同的兼性共生菌[101-104](表1)。豌豆蚜每個個體可感染多達(dá)4種兼性共生菌[67,101]，其中研究較多的是Serratia。研究發(fā)現(xiàn)，次級共生菌存在于菌胞周圍的鞘細(xì)胞和蚜蟲血腔中[94]。通常，次級內(nèi)共生菌通過垂直方式進(jìn)行傳播[103]。但由于不同地理種群的蚜蟲體內(nèi)含有不同種類的次級內(nèi)共生菌，所以能夠引起混合侵染的水平傳遞可能也是其主要傳播方式[110]。兼性共生菌與豌豆蚜的生物生態(tài)學(xué)特性密切相關(guān)[103]。一方面，兼性共生菌可為豌豆蚜提供營養(yǎng)、調(diào)控蚜蟲生殖、參與蚜蟲的體色變化和誘導(dǎo)翅型的產(chǎn)生[18,109,111]。另一方面，共生菌能夠提高蚜蟲抵抗病原菌、高溫、抵御天敵的能力，同時可提高豌豆蚜對寄主植物的適應(yīng)性來促進(jìn)不同寄主?；偷漠a(chǎn)生[72,88,94,103,106-108,112-115]。此外，兼性共生菌在改變豌豆蚜的免疫力過程中亦扮演著重要角色[116]。

表1 已知豌豆蚜兼性共生菌種類總結(jié)Table 1 The summary of facultative symbionts in Acyrthosiphon pisum

2 生態(tài)學(xué)特性

2.1 溫度及光照

溫度是影響豌豆蚜生長發(fā)育及體色多態(tài)性的重要因素。豌豆蚜各個發(fā)育階段的發(fā)育速率隨溫度的升高而加快[117-118]；在24 ℃條件下豌豆蚜的內(nèi)稟增長率最大，在18 ℃時具有最高的種群凈增殖率，在27 ℃時繁殖力顯著下降；在31 ℃下豌豆蚜若蟲蟲體很小，能蛻皮但不能正常發(fā)育為成蚜[119]。在不同溫度下飼養(yǎng)豌豆蚜，發(fā)現(xiàn)當(dāng)溫度高于19.6 ℃時若蚜死亡率最高[120]。這些研究均表明，豌豆蚜不耐高溫。此外，環(huán)境溫度不僅影響豌豆蚜的生長發(fā)育，還可以塑造豌豆蚜之間的個體差異[121]。目前，關(guān)于溫度對豌豆蚜體色的影響主要集中在低溫[60]，但對于其影響豌豆蚜體色的內(nèi)在分子機制尚不清楚。

豌豆蚜是一類對光照較為敏感的昆蟲，光照除了影響蚜蟲的時空分布、生長發(fā)育及取食行為之外，對豌豆蚜的生殖轉(zhuǎn)變、性別分化、翅型及體色分化均造成不同程度的影響[16,81,122-123]。研究還發(fā)現(xiàn)，豌豆蚜因能夠自身合成光合作用的重要色素類胡蘿卜素，從而具備吸收太陽光作為其代謝能量的特殊能力，推測其為目前唯一具備光合能力的動物[60]。

2.2 殺蟲劑

蚜蟲每年能夠造成數(shù)百億的作物損失，殺蟲劑的使用仍然是當(dāng)前控制蚜蟲包括豌豆蚜最重要的手段之一[35]。控制豌豆蚜的殺蟲劑主要有吡蟲啉、阿維菌素、高效氯氰菊酯等[13]。豌豆蚜與其天敵昆蟲均受到亞致死效應(yīng)的影響，亞致死效應(yīng)對于協(xié)調(diào)化學(xué)農(nóng)藥防治蚜蟲具有重要的作用[124-125]。隨著吡蟲啉亞致死劑量的增加，豌豆蚜F0代成蚜壽命縮短，產(chǎn)蚜量減少，且影響F1代種群的生長發(fā)育及繁殖[126]。

隨著豌豆蚜綠色防治研究的不斷深入，生物農(nóng)藥的研究和應(yīng)用越來越受到重視。蛋白酶抑制劑蛋白有防治害蟲的潛質(zhì)，它可以通過抑制昆蟲體內(nèi)蛋白酶和淀粉酶的作用，延遲幼蟲的發(fā)育，導(dǎo)致其體重減少、免疫力和生殖率降低，最后導(dǎo)致昆蟲死亡[14,35]。同時受生物環(huán)境影響的昆蟲激肽類似物也能夠提高豌豆蚜的拒食素活性和誘發(fā)高死亡率[127-128]等。另外，皂苷、芹黃素糖苷在防控豌豆蚜中均有潛在應(yīng)用前景[129-130]。

2.3 天敵昆蟲

天敵昆蟲在豌豆蚜的控制中發(fā)揮著重要的作用，目前已報道的豌豆蚜自然天敵來自5目7科[35,131](表2)。

表2 已報道的豌豆蚜天敵昆蟲種類Table 2 Reported natural enemiesof Acyrthosiphon pisum

捕食性天敵和寄生性天敵在控制豌豆蚜危害方面均發(fā)揮重要作用。豌豆蚜的捕食性天敵主要有瓢蟲、草蛉、蝽和食蚜蠅等，而寄生性天敵研究相對較多的則為蚜繭蜂[132]。田間調(diào)查發(fā)現(xiàn)，多異瓢蟲是甘肅省苜蓿田中主要的捕食性天敵之一，可在一定程度上減輕豌豆蚜的危害[19]。瓢蟲的個體大小和齡期均能夠影響其對豌豆蚜的捕食效率，個體較大的瓢蟲捕食效率較高[133]，成蟲和4齡幼蟲捕食作用較其它齡期強[134-135]。七星瓢蟲(Coccinellaseptempunctata)對豌豆蚜捕食功能反應(yīng)的研究表明，攻擊率最強且處理時間最短的蟲態(tài)為七星瓢蟲的成蟲[136]。同日齡瓢蟲在豌豆蚜不同密度梯度時，捕食量具有顯著差異[136]。異色瓢蟲(Harmoniaaxyridis)在低種群數(shù)量時，對豌豆蚜兩種色型未表現(xiàn)出偏好選擇性，而在較高種群數(shù)量時，對豌豆蚜紅色型表現(xiàn)出一定的偏好選擇性[137]。大灰食蚜蠅(Metasyrphuscorollae)對豌豆蚜具有較強的捕食作用[138]，而一種雜食性椿象(Macrolophuspygmaeus)對豌豆蚜也具有捕食作用[139]。

3 總結(jié)及展望

豌豆蚜不僅具有重要的經(jīng)濟價值，而且還具有重要的生態(tài)學(xué)研究價值。盡管已對豌豆蚜的基礎(chǔ)生物生態(tài)學(xué)進(jìn)行了廣泛研究，但對其生態(tài)適應(yīng)性、為害機理及遺傳進(jìn)化等方面的研究還十分欠缺。在全球氣候變化和當(dāng)下牧草種植結(jié)構(gòu)及作物布局發(fā)生重大變化的大背景下，需進(jìn)一步深入研究豌豆蚜生物生態(tài)學(xué)特性的演變規(guī)律，尤其要加強豌豆蚜在遺傳、生理、行為等方面對環(huán)境變化響應(yīng)的研究，為控制豌豆蚜的危害提供理論依據(jù)。

豌豆蚜極高的種群密度及其危害性，與其豐富的遺傳多樣性密切相關(guān)。研究豌豆蚜的種群遺傳與進(jìn)化，有助于進(jìn)一步深入認(rèn)識豌豆蚜的生物生態(tài)學(xué)特性，揭示種群的演化規(guī)律及生態(tài)進(jìn)化過程，也可為該蟲的防控提供理論指導(dǎo)。我國苜蓿地豌豆蚜自然種群中紅色型與綠色型的種群密度比率一直處于動態(tài)變化當(dāng)中，且近年來紅色型種群數(shù)量呈逐年上升趨勢。與綠色型相比，豌豆蚜紅色型在綠色植物上具有較高的可見度，故紅色型更易遭遇天敵捕食而在進(jìn)化過程中處于不利地位。然而，關(guān)于豌豆蚜兩種體色型的相互轉(zhuǎn)變以及自然種群相對數(shù)量的演替機制，目前尚不清楚。

隨著人們對豌豆蚜生物生態(tài)學(xué)特性研究的不斷深入，對其生態(tài)適應(yīng)性遺傳機制的研究變得尤為重要。翅多型和體色變化是研究其生態(tài)適應(yīng)性的兩個重要切入點。最近的研究[84]發(fā)現(xiàn)，豌豆蚜胰島素信號通路中的Apirp5與其翅多型密切相關(guān)，這與褐飛虱翅型變化的研究結(jié)果類似[83]，但有關(guān)豌豆蚜翅型變化的遺傳機制仍不甚清楚。研究表明，低溫可誘導(dǎo)豌豆蚜體色在紅綠之間轉(zhuǎn)變[11]，而共生菌同樣也參與了豌豆蚜體色的變化[12]。因此，豌豆蚜體色多態(tài)性可能受到諸多環(huán)境因子及自身遺傳的共同作用。深入研究豌豆蚜體色變化的遺傳學(xué)基礎(chǔ)，是今后應(yīng)該重點關(guān)注的研究內(nèi)容。

近年來，豌豆蚜已從苜蓿田的次要害蟲上升為主要害蟲，原本以抗苜蓿斑蚜為主的部分苜蓿品種對豌豆蚜的抗性喪失，抗性的喪失會造成田間蚜蟲的大爆發(fā)[71]。天敵防治和生物藥劑防治均可作為豌豆蚜防治的有效手段[140]，而一些常用農(nóng)事操作也可在一定程度上控制苜蓿田中的害蟲數(shù)量[141-142]。種植抗性品種不但可從根本上控制蚜蟲的危害，同時也可減少化學(xué)農(nóng)藥的使用從而達(dá)到保護(hù)天敵的目的。因此，選育并利用抗蚜品種是防控豌豆蚜的重要途徑。另外，作物布局顯著影響豌豆蚜的種群數(shù)量及季節(jié)性發(fā)生規(guī)律[23,38]。因此，有必要進(jìn)一步探明作物布局、氣候等因素對豌豆蚜種群產(chǎn)生的影響，以便發(fā)展和完善不同地區(qū)不同作物上豌豆蚜的防控技術(shù)體系。綜上所述，應(yīng)在深入研究豌豆蚜生物生態(tài)學(xué)特性和生態(tài)適應(yīng)性機制的基礎(chǔ)之上，建立以利用抗蚜品種為中心，天敵和生物防治為基礎(chǔ)，化學(xué)防治為輔的豌豆蚜綠色防控技術(shù)體系。

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(責(zé)任編輯 王芳)

Biological and ecological characteristics ofAcyrthosiphonpisum

Zhang Li1,2,3, Yuan Ming-long1,2,3

(1.State Key Laboratory of Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China; 2.National Demonstration Center for Experimental Grassland Science Education (Lanzhou University) , Lanzhou 730020, China; 3.Key Laboratory of Grassland Livestock Industry Innovation, Ministry of Agriculture, Lanzhou University, Lanzhou 730020, China)

Acyrthosiphonpisum(Hemiptera, Aphidoidea, Aphididae) is an important insect pest of many crops and forage grasses. This pest not only causes serious losses to alfalfa production, but also is the vector of many plant viruses. As an ecological model insect,A.pisumfascinates many scientists due to its complex life cycle, multiple reproductive types, polymorphism and symbiotic relationship with bacteria. We summarized the research status of biological and ecological characteristics ofA.pisum, especially for phenotypic polymorphisms and endosymbiotic bacteria. It is necessary to study how the biological characteristics ofA.pisumrespond to global climate and cropping pattern changes, and elucidate the genetic mechanisms of ecological adaptation and polymorphisms; this will be helpful for the establishment of sustainable management strategies for this pest.

aphid; biological characteristics; phenotypic plasticity; parthenogernesis; ecological adaptation

Yuan Ming-long E-mail:yuanml@lzu.edu.cn

10.11829/j.issn.1001-0629.2016-0518

2016-10-08 接受日期：2016-12-12

甘肅省自然科學(xué)基金青年科技基金計劃(1506RJZA211)

張麗(1993-)，女，甘肅定西人，在讀碩士生，主要從事草地昆蟲學(xué)研究。E-mail:2516486351@qq.com

袁明龍(1982-)，男，甘肅靖遠(yuǎn)人，副教授，博士，主要從事草地昆蟲學(xué)及分子生態(tài)學(xué)研究。E-mail:yuanml@lzu.edu.cn

Q969.36+7.2

A

1001-0629(2017)08-1727-14

張麗，袁明龍.豌豆蚜生物生態(tài)學(xué)特性.草業(yè)科學(xué),2017,34(8)：1727-1740.

Zhang L,Yuan M L.Biological and ecological characteristics ofAcyrthosiphonpisum.Pratacultural Science，2017,34(8)：1727-1740.