孟 倩,孟 玲,李保平

南京農(nóng)業(yè)大學(xué)植物保護(hù)學(xué)院,農(nóng)作物生物災(zāi)害綜合治理教育部重點(diǎn)實(shí)驗(yàn)室, 南京 210095

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短時(shí)高溫脅迫對(duì)斑痣懸繭蜂發(fā)育指標(biāo)的影響

孟 倩,孟 玲,李保平*

南京農(nóng)業(yè)大學(xué)植物保護(hù)學(xué)院,農(nóng)作物生物災(zāi)害綜合治理教育部重點(diǎn)實(shí)驗(yàn)室, 南京 210095

全球氣候變化不僅包括平均氣溫上升,而且諸如熱浪的極端天氣事件出現(xiàn)的頻率和程度也增大。寄生蜂雖在寄主體內(nèi)完成生長(zhǎng)發(fā)育,也會(huì)受到極端氣溫的影響。為探究短時(shí)高溫對(duì)寄生性天敵斑痣懸繭蜂(Meteoruspulchricornis)幼蟲(chóng)發(fā)育表現(xiàn)的影響,以斜紋夜蛾(Spodopteralitura)幼蟲(chóng)為寄主,分別對(duì)1—6日齡子代蜂進(jìn)行39℃、持續(xù)4h的高溫脅迫處理,以發(fā)育全程進(jìn)行適溫(晝29℃、夜26℃)處理為對(duì)照,觀察子代存活、發(fā)育歷期、羽化和成蟲(chóng)壽命等發(fā)育指標(biāo)。高溫脅迫處理1日和5日齡子代蜂幼蟲(chóng)使幼蟲(chóng)存活率下降,與對(duì)照相比分別降低36.1%和28.6%；高溫脅迫可延長(zhǎng)子代蜂幼蟲(chóng)發(fā)育歷期,與對(duì)照相比,高溫處理2、3、5日和6日齡子代蜂幼蟲(chóng)使幼蟲(chóng)發(fā)育歷期分別延長(zhǎng)了5.0%,5.2%,7.0%和12.1%；高溫脅迫處理5、6日齡子代蜂幼蟲(chóng)使羽化出的成蟲(chóng)體型(用后足脛節(jié)長(zhǎng)度表示)比對(duì)照分別減小1.8% 和 2.6%。高溫脅迫處理對(duì)子代蜂蛹發(fā)育歷期、羽化率以及成蟲(chóng)壽命等均沒(méi)有顯著影響。研究結(jié)果說(shuō)明,短時(shí)高溫脅迫對(duì)斑痣懸繭蜂高齡幼蟲(chóng)的負(fù)面影響比對(duì)低齡幼蟲(chóng)大。

溫度適應(yīng)；熱激；溫度；生活史特征；寄生蜂

全球氣候變暖是國(guó)際社會(huì)面臨的嚴(yán)峻挑戰(zhàn)[1]。全球氣候變暖不僅包括平均溫度的升高,還包括極端氣候事件(比如熱浪)頻率和強(qiáng)度的增大[2]。因此,高強(qiáng)度、高頻率的熱浪將更加常見(jiàn)[3]。昆蟲(chóng)作為變溫動(dòng)物,其生長(zhǎng)發(fā)育對(duì)氣溫極為敏感。昆蟲(chóng)有其生長(zhǎng)、發(fā)育、繁殖及存活等生命活動(dòng)的適宜溫度范圍,當(dāng)溫度超過(guò)最適范圍后,昆蟲(chóng)的生命活動(dòng)就會(huì)受到嚴(yán)重的影響[4]。了解昆蟲(chóng)在高溫脅迫下行為以及生理上的反應(yīng),可以幫助人們預(yù)測(cè)在氣候變暖情況下昆蟲(chóng)的分布、生態(tài)位的變化以及物種滅跡的可能性[5]。特別是寄生性天敵,其控制害蟲(chóng)的能力大部分取決于它們對(duì)環(huán)境的適應(yīng)能力[6],通常依賴于寄主的生理?xiàng)l件以及寄主植物的生態(tài)環(huán)境,因此對(duì)氣溫的變化更加敏感。對(duì)寄生性天敵昆蟲(chóng)的相關(guān)研究表明,對(duì)燕麥蚜繭蜂(Aphidiusavenae)、阿爾蚜繭蜂(A.ervi)、桃赤蚜繭蜂(A.matricariae)和角釉小蜂(Hemiptarsenusvaricornis)等的幼蟲(chóng)期進(jìn)行短暫的高溫脅迫,可使其存活率下降、發(fā)育歷期延長(zhǎng)以及繁殖力下降等[7- 10]。但這些研究均把幼蟲(chóng)期作為一個(gè)整體蟲(chóng)態(tài)進(jìn)行脅迫處理,未進(jìn)一步區(qū)分其不同階段對(duì)高溫脅迫的反應(yīng)。寄生蜂幼蟲(chóng)期發(fā)育經(jīng)歷著較大的形態(tài)和生理變化過(guò)程[11],其對(duì)脅迫的反應(yīng)可能存在差異。對(duì)此,本研究通過(guò)比較寄生蜂幼蟲(chóng)不同時(shí)期對(duì)高溫脅迫的反應(yīng),旨在找出敏感時(shí)期,從而為預(yù)測(cè)高溫脅迫對(duì)寄生性天敵的影響提供依據(jù)。

斑痣懸繭蜂(Meteoruspulchricornis)為單寄生、容性內(nèi)寄生蜂,是舞毒蛾(Lymantriadispar)[12]、棉鈴蟲(chóng)(Helicoverpaarmigera)[13]、斜紋夜蛾(Spodopteralitura)和甜菜夜蛾(S.exigua)[14]等重要農(nóng)林害蟲(chóng)的優(yōu)勢(shì)種寄生蜂。該蜂活動(dòng)于夏、秋季,偏好寄生幼蟲(chóng)中間幾個(gè)齡期。本研究以斜紋夜蛾幼蟲(chóng)為寄主,于寄主幼蟲(chóng)發(fā)育期內(nèi)不同時(shí)期進(jìn)行短時(shí)高溫處理,觀察和比較子代蜂的生長(zhǎng)發(fā)育表現(xiàn),以揭示寄生蜂幼蟲(chóng)期不同階段對(duì)高溫脅迫的反應(yīng)。

1 材料與方法

1.1 供試?yán)ハx(chóng)

斑痣懸繭蜂從南京農(nóng)業(yè)大學(xué)江浦實(shí)驗(yàn)農(nóng)場(chǎng)大豆試驗(yàn)田采集的斜紋夜蛾幼蟲(chóng)飼養(yǎng)獲得,在室內(nèi)以斜紋夜蛾2—3齡幼蟲(chóng)作為寄主繼代飼養(yǎng)。供試寄主斜紋夜蛾幼蟲(chóng)在室內(nèi)用人工飼料繼代飼養(yǎng)[15]。飼養(yǎng)環(huán)境的溫度為26℃,光周期為14h光/10h暗,相對(duì)濕度為60%+10%。

1.2 實(shí)驗(yàn)方法

取供試寄主斜紋夜蛾2齡末至3齡初的幼蟲(chóng),單頭稱體重后放入試管中,釋放1頭4—6日齡有寄生經(jīng)歷的雌蜂,觀察到寄生蜂完成1次有效針刺(產(chǎn)卵器扎入約5 s,拔出時(shí)拍動(dòng)翅膀)后,取出寄主幼蟲(chóng),單頭放于有人工飼料的培養(yǎng)皿中,置于人工氣候箱內(nèi)進(jìn)行溫度處理：常溫(在23:00—次日6:00時(shí)之間為26℃,其余時(shí)間為29℃)和高溫脅迫(在12:00—16:00時(shí)之間為39℃,其余同對(duì)照),39℃是我國(guó)華東地區(qū)夏季極端高溫日略高于上限氣溫(37—38℃)的溫度(南京農(nóng)業(yè)氣象網(wǎng)頁(yè)http://nq.njqxj.gov.cn/)。設(shè)6個(gè)幼蟲(chóng)日齡處理,即寄生后第1、2、3、4、5或6天接受高溫脅迫處理,將刺扎寄生后一直進(jìn)行常溫處理的幼蟲(chóng)作為對(duì)照。每日觀察2次(8:00 和20:00),直到寄生蜂羽化為成蟲(chóng),觀察和測(cè)量子代蜂存活、羽化、成蟲(chóng)壽命和體型大小(用后足脛節(jié)長(zhǎng)度代表)。每處理重復(fù)≥50次。

1.3 數(shù)據(jù)分析

子代蜂存活的觀測(cè)值為二進(jìn)制數(shù)據(jù),符合二項(xiàng)式分布型,故采用logistic模型擬合；子代蜂幼蟲(chóng)發(fā)育歷期(從雌蜂產(chǎn)卵到子代蜂化蛹的時(shí)間)、蛹?xì)v期(從子代蜂化蛹到羽化的時(shí)間)、子代蜂后足脛節(jié)長(zhǎng)度和成蟲(chóng)壽命觀測(cè)值屬于連續(xù)數(shù)值數(shù)據(jù),符合高斯分布型,故采用一般線性回歸模型擬合。分析中以寄主體重為協(xié)變量,以消除其影響。為確立幼蟲(chóng)期對(duì)高溫脅迫的敏感階段,將各個(gè)幼蟲(chóng)日齡處理與對(duì)照進(jìn)行比較。假說(shuō)檢驗(yàn)的顯著性概率水平為5%。數(shù)據(jù)分析用R統(tǒng)計(jì)軟件[16]。

2 結(jié)果與分析

2.1 高溫脅迫對(duì)斑痣懸繭蜂子代蜂存活的影響

擬合logistic 模型結(jié)果表明,日齡為1—6 d的子代蜂經(jīng)過(guò)高溫脅迫后的存活率在51.1%到80.6%之間。與對(duì)照相比,寄生后第1、5天接受高溫脅迫處理的子代蜂幼蟲(chóng)存活率顯著下降,分別下降36.1%和28.6% (圖1)。高溫脅迫不影響子代蜂蛹至成蟲(chóng)的羽化,羽化率在50%到87%之間(圖1)。

圖1 高溫脅迫對(duì)子代蜂幼蟲(chóng)存活和成蟲(chóng)羽化的影響Fig.1 Effects of the heat stress on larval survival to pupa and pupa-to-adult emergence rates of offspring parasitoids receiving the treatment at different larval ages幼蟲(chóng)各處理水平(1— 6d)以及對(duì)照的存活率觀測(cè)樣本數(shù)分別為45、40、39、44、49、31和30,成蟲(chóng)羽化率觀測(cè)樣本數(shù)分別為23、30、24、24、27、25和28; *代表與對(duì)照間存在顯著差異(P < 0.05)

2.2 高溫脅迫對(duì)斑痣懸繭蜂幼蟲(chóng)發(fā)育歷期和蛹?xì)v期的影響

一般線性模型擬合表明,日齡為1—6 d的子代蜂幼蟲(chóng)經(jīng)過(guò)高溫脅迫后的發(fā)育歷期平均值在7.9 d到8.7 d之間。與對(duì)照相比,寄生后第2、3、5、6天接受高溫脅迫處理的子代蜂幼蟲(chóng)發(fā)育歷期顯著延長(zhǎng),分別延長(zhǎng)5.0%、5.2%、7.0%、12.1%(圖2 A)。高溫脅迫不影響子代蜂蛹的發(fā)育歷期,平均為6.0—6.4 d(圖2 B)。

圖2 高溫脅迫對(duì)子代蜂幼蟲(chóng)發(fā)育歷期和蛹?xì)v期的影響Fig.2 Effects of the heat stress on larval and pupal duration of offspring parasitoids receiving the treatment at different larval ages圖柱上的短柄代表標(biāo)準(zhǔn)誤 Bars are standard errors of the mean

2.3 高溫脅迫對(duì)斑痣懸繭蜂成蟲(chóng)體型大小和壽命的影響

一般線性模型擬合表明,高溫脅迫處理日齡為1—6 d的子代蜂幼蟲(chóng)后,發(fā)育至成蟲(chóng)后的后足脛節(jié)長(zhǎng)度在1.55—1.59 mm之間。與對(duì)照相比,寄生后第5、6天接受高溫脅迫處理的子代蜂成蟲(chóng)體型顯著減小,分別減小1.8%和2.6% (圖3)。高溫脅迫不影響子代蜂成蟲(chóng)壽命,平均為5.2—5.8 d(圖3)。

圖3 高溫脅迫對(duì)子代蜂成蟲(chóng)后足脛節(jié)長(zhǎng)度和壽命的影響Fig.3 Effects of the heat stress on adult hind tibia length and longevity of offspring parasitoids receiving the treatments at different larval ages

3 討論

本研究發(fā)現(xiàn),高溫脅迫斑痣懸繭蜂不同日齡幼蟲(chóng)對(duì)幼蟲(chóng)發(fā)育表現(xiàn)具有不同的影響,對(duì)不同發(fā)育指標(biāo)的影響也不一致：高溫脅迫處理的1日和5日齡子代蜂幼蟲(chóng)的幼蟲(chóng)期存活率顯著低于其他處理的日齡幼蟲(chóng)；但高溫處理晚期幼蟲(chóng)對(duì)幼蟲(chóng)發(fā)育歷期和羽化出蜂體型大小的不良影響大于處理早期幼蟲(chóng)。

一般來(lái)說(shuō),斑痣懸繭蜂幼蟲(chóng)期通常可以劃分為3個(gè)時(shí)期,即卵期(1—2 d)、初齡幼蟲(chóng)期(3—4 d)和高齡幼蟲(chóng)期(5—6 d)。寄生蜂的胚胎發(fā)育是一個(gè)連續(xù)的過(guò)程,根據(jù)發(fā)育特征可分為5個(gè)階段：早期發(fā)育階段,胚體伸長(zhǎng)期,原軀原頭分化階段,器官形成階段,胚胎成熟期[17]。產(chǎn)卵后20—32 h為原腸胚形成及器官發(fā)育期,這一階段胚帶細(xì)胞裂殖旺盛,并形成原腸溝,是卵期生長(zhǎng)發(fā)育的關(guān)鍵[18]。若干研究表明,溫度不僅影響卵特性,而且影響卵孵化后的表現(xiàn)[19]；但有時(shí)后期表現(xiàn)可彌補(bǔ)早期經(jīng)歷的不足[20]。此外,昆蟲(chóng)卵期由于快速熱傳導(dǎo)以及較短的發(fā)育歷期,因此,昆蟲(chóng)卵期可能對(duì)高溫脅迫更加敏感[21]。例如,吳靜[22]對(duì)螟黃赤眼蜂(Trichogrammachilonis)的研究發(fā)現(xiàn),卵期受到的影響大于后續(xù)蟲(chóng)態(tài)。但在某些植食性昆蟲(chóng)中,高齡幼蟲(chóng)受高溫脅迫的影響大于低齡幼蟲(chóng)。例如,家蠶(Bombyxmori)幼蟲(chóng)對(duì)高溫的耐受性順序?yàn)椋?齡>2齡>3齡>4齡>5齡,即高齡幼蟲(chóng)對(duì)高溫脅迫較敏感[23]；對(duì)麥無(wú)網(wǎng)長(zhǎng)管蚜(Metopolophiumdirhodum)進(jìn)行短暫的高溫脅迫后,也發(fā)現(xiàn)老熟幼蟲(chóng)對(duì)高溫具有較差的耐受性[24]。由這些研究可以推測(cè),昆蟲(chóng)受高溫脅迫的影響并非隨發(fā)育進(jìn)程表現(xiàn)出線性變化趨勢(shì)。對(duì)容性寄生蜂來(lái)說(shuō),對(duì)抗寄主強(qiáng)大的免疫系統(tǒng)是在發(fā)育過(guò)程中面對(duì)的挑戰(zhàn)[25]。但迄今很少有關(guān)于溫度變化對(duì)免疫系統(tǒng)影響的研究報(bào)道[26]。

本研究發(fā)現(xiàn),2、3、5日和6日齡子代蜂幼蟲(chóng)經(jīng)過(guò)高溫脅迫后,幼蟲(chóng)發(fā)育歷期延長(zhǎng),說(shuō)明短暫的高溫脅迫可能抑制子代蜂的生長(zhǎng)發(fā)育速度。在高適溫區(qū)范圍內(nèi),昆蟲(chóng)通常隨著溫度的升高,發(fā)育速率反而減慢,使卵期和幼蟲(chóng)期延長(zhǎng)[27]。昆蟲(chóng)各蟲(chóng)態(tài)發(fā)育速率均與外界溫度之間呈“S”型曲線關(guān)系。例如,楊孝龍等[28]研究發(fā)現(xiàn),在19—3l℃溫度范圍內(nèi),紅點(diǎn)唇瓢蟲(chóng)(Chilocoruskuwanae) 各蟲(chóng)態(tài)的發(fā)育歷期會(huì)隨溫度的升高而縮短,但超過(guò)33℃后,其發(fā)育速率受到抑制；林智慧等[29]發(fā)現(xiàn)南美斑潛蠅(Liriomyzahuidobrensis)卵期隨著溫度的升高而延長(zhǎng),40℃時(shí)卵期最長(zhǎng),顯著長(zhǎng)于其他溫度下(31℃和37℃)的卵期,說(shuō)明高溫對(duì)卵的發(fā)育有明顯抑制作用。然而,本研究中1和4日齡子代蜂幼蟲(chóng)發(fā)育歷期并沒(méi)有延長(zhǎng),究其原因,可能與寄生蜂不同時(shí)期對(duì)高溫的忍耐力或敏感性差異有關(guān),具體原因有待進(jìn)一步研究。

一般認(rèn)為,成蟲(chóng)體型大小是其繁殖力和適合度的線性函數(shù),從而成為衡量昆蟲(chóng)繁殖力[30]和適合度[31]的一項(xiàng)重要的適合度相關(guān)特征。本研究發(fā)現(xiàn),5日和6日齡子代蜂幼蟲(chóng)經(jīng)過(guò)高溫脅迫后,成蟲(chóng)體型顯著減小,而日齡為0—4 d的子代蜂幼蟲(chóng)經(jīng)過(guò)高溫脅迫后,成蟲(chóng)體型未有明顯變化。從該結(jié)果可以推測(cè),高溫脅迫高齡幼蟲(chóng)可能影響成蟲(chóng)的繁殖力。對(duì)其他昆蟲(chóng)的研究也有類似的發(fā)現(xiàn)。對(duì)麥無(wú)網(wǎng)長(zhǎng)管蚜高齡幼蟲(chóng)(4齡)進(jìn)行高溫脅迫后,成蟲(chóng)生殖力比低齡幼蟲(chóng)(2、3齡)下降得多[24]；小菜蛾(Plutellaxylostella)高齡幼蟲(chóng)經(jīng)過(guò)40℃高溫脅迫處理后,生殖力比低齡幼蟲(chóng)下降得多[32]；對(duì)褐飛虱(Nilaparvatalugens)的1齡若蟲(chóng)進(jìn)行40℃ 高溫脅迫處理后,發(fā)現(xiàn)成蟲(chóng)的產(chǎn)卵量明顯下降[33]；同樣,4齡異色瓢蟲(chóng)(Harmoniaaxyridis)幼蟲(chóng)經(jīng)過(guò)高溫脅迫后,成蟲(chóng)體型明顯變小,而對(duì)初齡幼蟲(chóng)的高溫處理卻沒(méi)有影響[34]。對(duì)昆蟲(chóng)不同發(fā)育階段進(jìn)行高溫脅迫,其結(jié)果對(duì)成蟲(chóng)的繁殖力是否有影響取決于昆蟲(chóng)是否有足夠的時(shí)間從熱傷害中恢復(fù)有關(guān)[32]。高齡幼蟲(chóng)經(jīng)過(guò)高溫脅迫后,可能沒(méi)有足夠的時(shí)間修復(fù)熱損傷,因此導(dǎo)致成蟲(chóng)的繁殖力下降。本研究未發(fā)現(xiàn)成蟲(chóng)壽命受高溫脅迫處理的不利影響,原因可能在于觀測(cè)時(shí)間間隔太大,導(dǎo)致很多觀測(cè)值重疊而無(wú)法獲得統(tǒng)計(jì)上的顯著性差異。

本研究結(jié)果說(shuō)明,不同日齡的斑痣懸繭蜂子代幼蟲(chóng)對(duì)高溫的耐受性存在明顯差異,總體而言,高齡幼蟲(chóng)更容易受到高溫脅迫的負(fù)面影響,是對(duì)高溫脅迫較敏感的時(shí)期。

致謝：南京農(nóng)業(yè)大學(xué)植物保護(hù)學(xué)院榮星幫助試驗(yàn),特此致謝。

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Effects of short-term heat stress on developmental parameters ofMeteoruspulchricornis(Hymenoptera: Braconidae)

MENG Qian, MENG Ling, LI Baoping*

CollegeofPlantProtection,KeyLaboratoryofMonitoringandManagementforPlantDiseasesandInsectPests,MinistryofAgriculture,NanjingAgriculturalUniversity,Nanjing210095,China

Global warming causes an increase in the average temperature butalso the intensity and frequency of extreme climatic conditions, such as short heat notoly waves. Extreme temperatures can impact parasitoid growth and development in hosts. Therefore, to understand the effect of heat waves on parasitic natural enemies of insect pests, we made laboratory experiments to examine developmental performances of the offspring of the solitary endoparasitoid,Meteoruspulchricornis, usingSpodopteralituralarvae as hosts. Offspring parasitoids in the hosts

the heat shock treatment by 39℃ for 4 hours at the age of 1, 2, 3, 4, 5, or 6 days old after parasitism. Developmental survival, duration, adult body size, and longevity of offspring parasitoids were observed. Larval survival to pupa decreased in the host larvae that received the heat treatment at the age of 1 d and 5 d, by 36.1% and 28.6%, respectively, compared to controls. Larval duration was prolonged by 5.0%, 5.2%, 7.0%, and 12.1%, respectively, in the host larvae that received the heat treatment at the age of 2, 3, 5, and 6 d, respectively, compared to control. Adult hind tibia length decreased by 1.8% and 2.6%, respectively, in the hosts that received the heat treatment at the age of 5 and 6 d. Pupa-to-adult duration, pupa-to-adult emergence rate, and adult longevity were not influenced by the heat shock treatment. The results of the present study suggest that older larvae ofM.pulchricorniswere more vulnerable to heat waves than younger ones.

thermal adaptation; heat shock; temperature; life history traits; parasitoid wasp

國(guó)家自然科學(xué)基金項(xiàng)目(31570389)；國(guó)家科技支撐計(jì)劃項(xiàng)目(2012BAC19B01);國(guó)家公益性行業(yè)(農(nóng)業(yè))科研專項(xiàng)(200903003,201103002)

2015- 11- 14; 網(wǎng)絡(luò)出版日期:2016- 10- 29

10.5846/stxb201511142311

*通訊作者Corresponding author.E-mail: lbp@njau.edu.cn

孟倩,孟玲,李保平.短時(shí)高溫脅迫對(duì)斑痣懸繭蜂發(fā)育指標(biāo)的影響.生態(tài)學(xué)報(bào),2017,37(8):2838- 2843.

Meng Q, Meng L, Li B P.Effects of short-term heat stress on developmental parameters ofMeteoruspulchricornis(Hymenoptera: Braconidae).Acta Ecologica Sinica,2017,37(8):2838- 2843.