張廷偉,陳萬(wàn)斌,劉長(zhǎng)仲,崔少偉,饒福強(qiáng)

(甘肅農(nóng)業(yè)大學(xué)植物保護(hù)學(xué)院 蘭州 730070)

不同光周期條件下綠色型豌豆蚜性蚜分化規(guī)律研究*

張廷偉,陳萬(wàn)斌,劉長(zhǎng)仲**,崔少偉,饒福強(qiáng)

(甘肅農(nóng)業(yè)大學(xué)植物保護(hù)學(xué)院 蘭州 730070)

為了明確豌豆蚜隨光周期變化時(shí)的種群繁殖策略以及性蚜分化規(guī)律,在4個(gè)光周期(8L∶16D、10L∶14D、12L∶12D 和14L∶10D)條件下,研究了綠色型豌豆蚜經(jīng)光周期連續(xù)誘導(dǎo)2代后(G1和G2),后代(G3)中的性蚜分化情況。結(jié)果表明: 蘭州地區(qū)豌豆蚜性雌蚜均為無(wú)翅型,而雄蚜均為有翅型。光周期對(duì)豌豆蚜性蚜分化有顯著影響(P＜0.05)。在14L∶10D光周期條件下,豌豆蚜后代中無(wú)性蚜分化; 在12L∶12D、10L∶14D 和8L∶16D 3個(gè)光周期條件下均有性蚜分化,而且隨著光照時(shí)間的變短,豌豆蚜G3中雄蚜所占百分比逐漸減小,性雌蚜所占百分比逐漸升高。在12L∶12D光周期條件下,雄蚜比例最高(30.39%),而性雌蚜比例最低(55.67%);在10L∶14D光周期條件下,G3中胎生蚜比例最高,而性蚜比例最低; 在8L∶16D光周期條件下,無(wú)雄蚜分化,而性雌蚜比例最高(90.76%)。在同一短光照條件下,G1產(chǎn)仔日齡對(duì)G3中性蚜分化具有顯著影響(P＜0.05),同時(shí)性蚜分化具有一定的時(shí)間順序,即先產(chǎn)性雌蚜,后產(chǎn)雄蚜,由性雌蚜向雄蚜過(guò)渡過(guò)程中伴隨產(chǎn)生胎生蚜。表明綠色型豌豆蚜性蚜產(chǎn)生的臨界光照時(shí)長(zhǎng)介于12～14 h,相對(duì)較長(zhǎng)的光照條件下有利于雄蚜產(chǎn)生,而光照時(shí)間越短性雌蚜比例越高。說(shuō)明光周期變化是豌豆蚜生殖模式轉(zhuǎn)變和性蚜分化的重要因子,而且豌豆蚜親代經(jīng)歷光周期處理時(shí)間的長(zhǎng)短影響后代中性蚜的分化。

豌豆蚜; 綠色型; 光周期; 雄蚜; 性雌蚜; 胎生蚜

表型可塑性作為生物體對(duì)環(huán)境異質(zhì)性(environmental heterogeneity)適應(yīng)的一種反應(yīng)機(jī)制,而成為研究物種生態(tài)和進(jìn)化相互關(guān)系的熱點(diǎn)[1-2]。蚜蟲(chóng)非遺傳性多型現(xiàn)象非常普遍,主要包括體色多型、翅型多型和生殖多型等[3],其中生殖多型現(xiàn)象被認(rèn)為是蚜蟲(chóng)進(jìn)化和適應(yīng)環(huán)境條件的典范[4]。在夏季長(zhǎng)日照條件下,蚜蟲(chóng)營(yíng)無(wú)性孤雌胎生生殖從而使種群快速增長(zhǎng),對(duì)農(nóng)業(yè)生產(chǎn)造成嚴(yán)重影響; 當(dāng)秋末環(huán)境條件惡化如溫度逐漸降低、光周期逐漸變短時(shí),孤雌胎生蚜通過(guò)母代感知環(huán)境條件變化從而隔代分化出有性個(gè)體(性蚜)進(jìn)行有性生殖以滯育卵越冬。因此,蚜蟲(chóng)性蚜的產(chǎn)生不僅有利于蚜蟲(chóng)渡過(guò)不良環(huán)境,而且有利于種群內(nèi)基因的交流和保持豐富的遺傳結(jié)構(gòu),對(duì)蚜蟲(chóng)的生存和進(jìn)化具有重要作用[5-7],同時(shí)利用人工條件誘導(dǎo)蚜蟲(chóng)性蚜產(chǎn)生對(duì)研究蚜蟲(chóng)多型現(xiàn)象和種群遺傳有重要意義。

豌豆蚜(Acyrthosiphon pisum)亦稱(chēng)為豆無(wú)網(wǎng)長(zhǎng)管蚜,是溫帶地區(qū)豆科作物及牧草上的重要害蟲(chóng)[8-9]。因豌豆蚜具有多型現(xiàn)象普遍、周期性孤雌生殖以及同寄主全周期型生活史等特點(diǎn),使其成為發(fā)育生物學(xué)和進(jìn)化生物學(xué)研究領(lǐng)域中的理想材料[4,9]。在年生活史中,豌豆蚜在光周期變短和溫度降低時(shí)產(chǎn)生1代有性蚜(雄蚜和性雌蚜),一般性雌蚜都為無(wú)翅型,而雄蚜具有翅二型現(xiàn)象。Smith和MacKay[10]報(bào)道豌豆蚜雄蚜的分化有只產(chǎn)生有翅型、只產(chǎn)生無(wú)翅型和能產(chǎn)有翅和無(wú)翅型3種類(lèi)型,而且雄蚜翅型分化差異與地理種群有關(guān)。Caillaud等[11]經(jīng)孟德?tīng)栠z傳研究表明,豌豆蚜雄蚜翅型的有無(wú)取決于X染色體上api基因位點(diǎn)的等位基因。Lees[12]和MacKay[13]分別對(duì)英格蘭和加拿大的豌豆蚜種群在不同光周期下性蚜分化規(guī)律就行了研究,但不同地理種群間性蚜分化具有明顯差異。國(guó)內(nèi)學(xué)者利用低溫和短光照對(duì)桃蚜(Myzus persicae)、棉蚜(Aphis gossypii)、大豆蚜(Aphis glycines)的性蚜分化進(jìn)行了誘導(dǎo)研究[14-17],但國(guó)內(nèi)關(guān)于豌豆蚜性蚜分化的研究尚少有報(bào)道。據(jù)調(diào)查,在我國(guó)西北地區(qū)紅色型豌豆蚜數(shù)量雖正逐年上升,但綠色型豌豆蚜仍為優(yōu)勢(shì)種群,而且每年春季最先發(fā)生的豌豆蚜也都為綠色型[18-20]。因此本文以綠色型豌豆蚜為研究對(duì)象,研究了不同光周期條件對(duì)豌豆蚜性蚜分化的影響,旨在明確豌豆蚜在我國(guó)西北地區(qū)性蚜產(chǎn)生的條件和性蚜分化的規(guī)律以及豌豆蚜在光周期變化時(shí)的種群繁殖策略,為進(jìn)一步研究豌豆蚜的種群遺傳奠定基礎(chǔ)。

1 材料與方法

1.1 供試材料

試驗(yàn)所用綠色型豌豆蚜采自甘肅農(nóng)業(yè)大學(xué)苜蓿試驗(yàn)基地(蘭州),飼養(yǎng)在盆栽蠶豆(Vicia fabae)‘臨蠶2’植株上,在溫度(22±1) ℃、光周期16 L∶8 D、相對(duì)濕度(RH) 60%的條件下飼養(yǎng)多代后備用。

1.2 試驗(yàn)方法

性蚜誘導(dǎo)試驗(yàn)在人工氣候箱中進(jìn)行,溫度為15 ℃,設(shè)置8L∶16D、10L∶14D、12L∶12D、14L∶10D等4個(gè)不同光周期。試驗(yàn)方法參照王玲等[17]略有改動(dòng),蚜蟲(chóng)飼養(yǎng)采用離體葉片培養(yǎng)皿單頭飼養(yǎng)法進(jìn)行[21]。選取1.1中備用植株上2 h內(nèi)初產(chǎn)的1齡若蚜(G1)10頭(盡可能為不同克隆),接入培養(yǎng)皿蠶豆葉片上置于相應(yīng)的光照培養(yǎng)箱中進(jìn)行飼養(yǎng),每一光周期條件下連續(xù)誘導(dǎo)2代。當(dāng)?shù)?代(G1)發(fā)育成熟并開(kāi)始產(chǎn)蚜后,分別收集其產(chǎn)仔日齡第2 d、7 d、12 d時(shí)所產(chǎn)若蚜(G2)各10頭,每皿放1頭,繼續(xù)飼養(yǎng),待G2開(kāi)始產(chǎn)蚜后每天收集一次若蚜(G3),直到G2全部死亡為止。將G2所有后代(G3)在新的葉片上繼續(xù)飼養(yǎng)至成蚜,以備區(qū)分各蚜型。蚜型區(qū)分主要通過(guò)解剖和觀察蚜蟲(chóng)的生殖系統(tǒng)或器官來(lái)判斷,G3中體內(nèi)具有紅色眼點(diǎn)胚胎的無(wú)翅或有翅蚜為孤雌胎生蚜,體內(nèi)具黃色卵粒的無(wú)翅蚜為性雌蚜,腹部具有黑色抱握器的有翅或無(wú)翅蚜為雄蚜[11,22],統(tǒng)計(jì)各蚜型的比例。

1.3 數(shù)據(jù)處理

試驗(yàn)所得的所有數(shù)據(jù),均利用 Microsoft Excel 2003 和SPSS 16.0軟件進(jìn)行統(tǒng)計(jì)分析。表中所出現(xiàn)的誤差處理方式均為平均值±標(biāo)準(zhǔn)誤(mean±SE)。方差分析采用SPSS 16.0軟件One-way ANOVA程序進(jìn)行,多重比較采用Duncan法進(jìn)行。

2 結(jié)果與分析

2.1 不同光周期對(duì)豌豆蚜性蚜分化的影響

由表 1可知,不同光周期對(duì)豌豆蚜性蚜分化比例具有顯著影響(P＜0.05)。在 14L∶10D光周期條件下,豌豆蚜G3中全為胎生蚜,無(wú)性蚜產(chǎn)生。在12L∶12D、10L∶14D和8L∶16D 3組光周期條件下,豌豆蚜G3中均有性蚜產(chǎn)生,且性雌蚜均為無(wú)翅蚜,而雄蚜均為有翅蚜,沒(méi)有發(fā)現(xiàn)無(wú)翅雄蚜。在3組短光照處理中,在12L∶12D光周期條件下,G3中雄蚜比例最高,而性雌蚜比例最低; 在8L∶16D光周期條件下,性蚜中無(wú)雄蚜分化,而性雌蚜比例最高; 而在10L∶14D光周期條件下,胎生蚜比例明顯較8L∶16D和12L∶12D光周期條件下高,而性蚜比例明顯比 8L∶16D和 12L∶12D光周期條件下低(P＜0.05)。表明相對(duì)較長(zhǎng)的光周期有利于雄蚜產(chǎn)生,而光周期越短性雌蚜比例越高。

表1 不同光周期處理兩代后豌豆蚜后代(G3)中各蚜型的分化比例Table 1 Percentages of various morphs of offspring (G3) of pea aphid after treatments of different photoperiods for 2 generations %

2.2 G1產(chǎn)仔日齡對(duì)性蚜分化的影響

由表2可知,在同一光周期條件下,G1不同產(chǎn)仔日齡對(duì)后代G3中各蚜型分化比例具有顯著影響,而且同一產(chǎn)仔日齡不同光周期間各蚜型分化比例也不相同(P＜0.05)。在12L∶12D光周期條件下,G1產(chǎn)仔日齡對(duì) G3中胎生蚜比例和性蚜比例沒(méi)有顯著影響(P＞0.05),而對(duì)雄蚜和性雌蚜的比例有顯著影響(P＜0.05),G12 d產(chǎn)仔日齡時(shí)所產(chǎn)后代G3中性雌蚜比例最高,而雄蚜比例最低,與7 d和12 d產(chǎn)仔日齡時(shí)G3中性雌蚜和雄蚜比例存在顯著差異(P＜0.05)。在10L∶14D光周期條件下,G1產(chǎn)仔日齡對(duì) G3中各蚜型比例均有顯著影響(P＜0.05),隨著 G1產(chǎn)仔日齡的增加,G3中雄蚜比例逐漸降低,而性雌蚜比例逐漸升高,但G17 d產(chǎn)仔日齡時(shí)后代G3中胎生蚜比例最高,而性蚜比例最低。在8L∶16D光周期條件下,G3中均無(wú)雄蚜產(chǎn)生,G1產(chǎn)仔日齡對(duì)G3中胎生蚜比例和性雌蚜比例均有顯著影響,從而影響性蚜總比例(P＜0.05),G112 d產(chǎn)仔日齡時(shí),G3中胎生蚜比例最低,而性雌蚜比例最高。表明豌豆蚜親代經(jīng)歷光周期時(shí)間的長(zhǎng)短會(huì)影響后代中性蚜的分化比例。

表2 不同光周期處理下第1代豌豆蚜(G1)產(chǎn)仔日齡對(duì)第3代(G3)各蚜型分化比例的影響Table 2 Effect of reproductive period of the first generation (G1) on the percentages of various morphs of the third generation (G3) of pea aphid under different photoperiods %

2.3 豌豆蚜性蚜分化的時(shí)間順序

根據(jù)觀察,在 15 ℃條件下,豌豆蚜整個(gè)產(chǎn)蚜期為20 d左右。由圖1可知,豌豆蚜的生殖模式、性蚜分化順序與光周期密切相關(guān),G2不同產(chǎn)蚜期內(nèi)后代G3中各蚜型分化比例不同。在14L∶10D光周期條件下整個(gè)G2產(chǎn)蚜期內(nèi),豌豆蚜G3中全為胎生蚜,無(wú)性蚜分化,以孤雌生殖模式進(jìn)行繁殖(見(jiàn)圖1 d),而當(dāng)光周期突然縮短時(shí)豌豆蚜以?xún)尚陨碁橹鳌⒓嬉怨麓粕衬Ｊ竭M(jìn)行繁殖(見(jiàn)圖1a,b,c)。在12L∶12D、10L∶14D和8L∶16D 3組光周期條件下,在G2產(chǎn)蚜期的前10 d內(nèi),豌豆蚜G3后代中均以產(chǎn)性雌蚜為主,而雄蚜在產(chǎn)蚜 10 d后出現(xiàn),并且隨著光周期逐漸縮短,雄蚜出現(xiàn)的天數(shù)逐漸縮短。在8L∶16D光周期條件下,G2第1 d所產(chǎn)的后代均為性雌蚜,而產(chǎn)蚜5 d后逐漸分化出胎生蚜; 在10L∶14D和12L∶12D光周期條件下,在前期分化出性雌蚜和胎生蚜之后,第10 d又分化出雄蚜。根據(jù)3組短光照條件下不同產(chǎn)蚜期內(nèi)各生物型的分化比例,表明豌豆蚜性蚜分化過(guò)程呈現(xiàn)一定的時(shí)間順序: 即先產(chǎn)性雌蚜,后產(chǎn)雄蚜,由性雌蚜向雄蚜過(guò)渡過(guò)程中伴隨產(chǎn)生胎生蚜。

圖1 不同光周期條件下特定產(chǎn)蚜期豌豆蚜各生物型分化比例Fig.1 Percentages of various morphs of pea aphid in G3under different photoperiods with specific reproductive period of G2

3 討論與結(jié)論

自然界中,溫度與光周期作為重要的環(huán)境因子,協(xié)同作用于生物體的生長(zhǎng)發(fā)育、繁殖和分布。昆蟲(chóng)的年生活史、滯育、世代交替和蚜蟲(chóng)的季節(jié)多型現(xiàn)象都與溫度和光周期變動(dòng)密切相關(guān)[23]。在溫帶地區(qū),光周期誘導(dǎo)是大多數(shù)昆蟲(chóng)滯育的主要原因,由于不同地理緯度間光周期的差異是恒定的,因此不同地理分布的同種昆蟲(chóng)滯育的臨界光周期具有地理差異性[24-25]。大豆蚜[17]、棉蚜[26]以及亞洲玉米螟(Ostrinia furnacalis)[27]等昆蟲(chóng)的不同地理種群研究證實(shí),對(duì)于短日照滯育型昆蟲(chóng)而言,滯育誘導(dǎo)的臨界光周期常隨著地理緯度的升高而延長(zhǎng)。據(jù)報(bào)道,在20 ℃條件下加拿大曼尼托巴地區(qū)豌豆蚜在光照時(shí)長(zhǎng)低于15 h時(shí)就有性蚜分化,而且在15L∶9D光周期條件下產(chǎn)生雄蚜最多[13]。Lees[12]對(duì)英格蘭伯克郡地區(qū)豌豆蚜種群研究發(fā)現(xiàn),在15 ℃,光周期為14L∶10D時(shí),性蚜出現(xiàn)而且最有利于雄蚜產(chǎn)生。本研究結(jié)果表明,在15 ℃條件下,蘭州地區(qū)豌豆蚜在14L∶10D光周期條件下無(wú)性蚜分化,而在12L∶12D光周期條件下開(kāi)始出現(xiàn)性蚜分化并且雄蚜數(shù)量最高(30.39%),可能與蘭州地區(qū)緯度較低有關(guān)[28]。在3組短光照處理下,性蚜比例均在70%以上,表明蘭州地區(qū)豌豆蚜性蚜產(chǎn)生的臨界日照時(shí)長(zhǎng)介于12～14 h。

本研究發(fā)現(xiàn),蘭州地區(qū)豌豆蚜性雌蚜均為無(wú)翅型,而雄蚜均為有翅型。在日光照長(zhǎng)度8～12 h范圍內(nèi),隨著光照時(shí)數(shù)的逐漸縮短,豌豆蚜雄蚜數(shù)量逐漸減少,而性雌蚜數(shù)量逐漸升高,同時(shí)性蚜分化過(guò)程呈現(xiàn)一定的時(shí)間順序,即先分化出性雌蚜,后分化出雄蚜,由性雌蚜向雄蚜過(guò)渡過(guò)程中伴隨產(chǎn)生胎生蚜,此結(jié)果與國(guó)外學(xué)者對(duì)豌豆蚜研究結(jié)果一致[12-13,29]。前人對(duì)蚜蟲(chóng)有翅型和無(wú)翅型生物學(xué)研究表明,在相同條件下有翅型蚜蟲(chóng)比無(wú)翅型蚜蟲(chóng)發(fā)育歷期更長(zhǎng),所需消耗營(yíng)養(yǎng)物質(zhì)更多,其原因在于翅型出現(xiàn)對(duì)于個(gè)體發(fā)育來(lái)說(shuō)是種額外投資,個(gè)體延長(zhǎng)發(fā)育歷期從而獲得完成個(gè)體發(fā)育所需的營(yíng)養(yǎng)物質(zhì)[29-31]。但是,在自然條件下,當(dāng)環(huán)境條件不適宜(光周期變短、食物資源短缺)時(shí),蚜蟲(chóng)可能會(huì)調(diào)整后代中生物型的比例,從而利用有限的資源獲得最大的投資回報(bào)率,如在12L∶12D光周期條件下,豌豆蚜后代中性雌蚜比例為55.67%,雄蚜比例為30.39%,而在8L∶16D光周期條件下,后代中性雌蚜比例達(dá)90.76%,而不分化出雄蚜。隨著光周期逐漸變短,性雌蚜數(shù)量增多,意味著越冬代卵越多和分布越廣。但隨著光周期變短豌豆蚜后代中雄蚜數(shù)量越來(lái)越少,甚至不產(chǎn)生雄蚜,一方面可能是因?yàn)榍捌诠庵芷谙鄬?duì)較長(zhǎng)時(shí)已經(jīng)存在足夠數(shù)量用于交配的雄蚜,另一方面可能是由于雄蚜出現(xiàn)在生殖后期且發(fā)育歷期較長(zhǎng)而導(dǎo)致親代減少對(duì)其投資[17,29]。此外,在短光照條件下,豌豆蚜在生殖期后期仍有10%～25%左右的孤雌胎生蚜產(chǎn)生,則表明只要環(huán)境條件適宜或者當(dāng)環(huán)境條件轉(zhuǎn)為適宜時(shí)該蟲(chóng)仍能以孤雌胎生模式進(jìn)行繁殖而繼續(xù)為害。因此,豌豆蚜在光周期變化時(shí)的這種性蚜分化規(guī)律與母體根據(jù)外在環(huán)境變化對(duì)后代性別分配的調(diào)整以及性蚜個(gè)體差異有關(guān)。

本研究還發(fā)現(xiàn),豌豆蚜G1不同產(chǎn)仔日齡對(duì)后代G3中各蚜型分化比例具有顯著影響,而且同一產(chǎn)仔日齡不同光周期間各蚜型分化比例也不相同(P＜0.05)。在12L∶12D光周期條件下,隨著G1產(chǎn)仔日齡的增加,后代中雄蚜比例表現(xiàn)出升高的趨勢(shì),而隨著光周期變短、G1產(chǎn)仔日齡的增加,后代中雄蚜比例逐漸降低甚至后代中不分化出雄蚜。在10L∶14D和8L∶16D光周期條件下隨著產(chǎn)仔日齡的增加,性雌蚜的比例逐漸增加,而孤雌胎生蚜的比例逐漸降低。G17 d產(chǎn)仔日齡前后對(duì)后代性蚜分化有顯著影響,此結(jié)果與棉蚜、大豆蚜的研究結(jié)果相似[15,17,26],究其原因可能與母代效應(yīng)有關(guān)。母代所經(jīng)歷的環(huán)境(光照、溫度和營(yíng)養(yǎng))變化會(huì)對(duì)子代表現(xiàn)型差異產(chǎn)生影響[32]。郭玉玲等[33]研究發(fā)現(xiàn),菜蛾盤(pán)絨繭蜂母代的光周期經(jīng)歷會(huì)影響子代滯育發(fā)生。因此,豌豆蚜G1產(chǎn)仔日齡的大小,意味著母代經(jīng)歷光周期時(shí)間的長(zhǎng)短,從而影響后代性蚜的分化。

本研究表明,蘭州地區(qū)綠色型豌豆蚜性蚜產(chǎn)生的臨界光照時(shí)長(zhǎng)介于12～14 h,不同光周期下性蚜分化比例不同,而且親代產(chǎn)仔日齡也會(huì)影響后代性蚜的分化。相對(duì)較長(zhǎng)的光照時(shí)長(zhǎng)有利于雄蚜的誘導(dǎo)出現(xiàn),而光照時(shí)長(zhǎng)越短,越有利于性雌蚜數(shù)量的增加,并且性蚜分化過(guò)程呈現(xiàn)一定的時(shí)間順序,即先分化出性雌蚜,后分化出雄蚜,由性雌蚜向雄蚜過(guò)渡過(guò)程中伴隨少量孤雌胎生蚜產(chǎn)生。本文僅對(duì)綠色型豌豆蚜性蚜分化規(guī)律進(jìn)行了研究,由于豌豆蚜具有紅、綠兩種色型,而且兩種色型豌豆蚜在相同寄主、溫度、光周期條件下生長(zhǎng)發(fā)育和繁殖方面存在顯著差異[19,21,34],因此紅色型豌豆蚜性蚜分化規(guī)律與綠色型是否相同有待研究。此外,由于自然界中溫度和光周期共同作用于生物體,因此關(guān)于光溫互作對(duì)豌豆蚜性蚜分化的誘導(dǎo)也有待進(jìn)一步研究。

References

[1]Agrawal A A.Phenotypic plasticity in the interactions and evolution of species[J].Science,2001,294(5541): 321–326

[2]陳倩,沈佐銳,王永模.蚜蟲(chóng)的表型可塑性及其遺傳基礎(chǔ)[J].昆蟲(chóng)學(xué)報(bào),2006,49(5): 859–866 Chen Q,Shen Z R,Wang Y M.Phenotypic plasticity of aphids and its genetic basis[J].Acta Entomologica Sinica,2006,49(5): 859–866

[3]Simpson S J,Sword G A,Lo N.Polyphenism in insects[J].Current Biology,2011,21(18): R738–R749

[4]Simon J C,Stoeckel S,Tagu D.Evolutionary and functional insights into reproductive strategies of aphids[J].Comptes Rendus Biologies,2010,333(6/7): 488–496

[5]Liu L J,Zheng H Y,Jiang F,et al.Comparative transcriptional analysis of asexual and sexual morphs reveals possible mechanisms in reproductive polyphenism of the cotton aphid[J].PLoS One,2014,9(6): e99506

[6]Poupoulidou D,Margaritopoulos J T,Kephalogianni T E,et al.Effect of temperature and photoperiod on the life cycle in lineages ofMyzus persicae nicotianaeandMyzus persicaes.str.(Hemiptera: Aphididae)[J].European Journal of Entomology,2006,103(2): 337–346

[7]Saunders D S.Insect photoperiodism: Effects of temperature on the induction of insect diapause and diverse roles for the circadian system in the photoperiodic response[J].Entomological Science,2014,17(1): 25–40

[8]張廷偉,黃純倩,杜軍利,等.阿爾蚜繭蜂對(duì)不同齡期豌豆蚜的寄生及后代適合度研究[J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2015,23(7): 914–918 Zhang T W,Huang C Q,Du J L,et al.Parasitism and offspring fitness ofAphidius erviHaliday in relation to its hostAcyrthosphon pisumHarris at different stages[J].Chinese Journal of Eco-Agriculture,2015,23(7): 914–918

[9]Brisson J A,Davis G K.Pea aphid[M]//Hunter W,Kole C.Genome Mapping and Genomics in Arthropods.Berlin Heidelberg: Springer,2008,1: 59–67

[10]Smith M A H,MacKay P A.Genetic variation in male alarydimorphism in populations of pea aphid,Acyrthosiphon pisum[J].Entomologia Experimentalis et Applicata,1989,51(2): 125–132

[11]Caillaud M C,Boutin M,Braendle C,et al.A sex-linked locus controls wing polymorphism in males of the pea aphid,Acyrthosiphon pisum(Harris)[J].Heredity,2002,89(5): 346–352

[12]Lees A D.Dual photoperiodic timers controlling sex and female morph determination in the pea aphid,Acyrthosiphon pisum[J].Journal of Insect Physiology,1990,36(8): 585–591

[13]MacKay P A.Production of sexual and asexual morphs and changes in reproductive sequence associated with photoperiod in the pea aphid,Acyrthosiphon pisum(Harris)[J].Canadian Journal of Zoology,1987,65(11): 2602–2606

[14]楊效文,張孝羲.溫度和光周期對(duì)煙蚜性蚜的誘導(dǎo)[J].昆蟲(chóng)知識(shí),1999,36(5): 272–274 Yang X W,Zhang X X.Production of sexual morph ofMyzus persicae(Sulzer) in relation to photoperiod and temperature[J].Entomological Knowledge,1999,36(5): 272–274

[15]龔鵬,張孝羲.溫度和光周期對(duì)棉蚜性蚜產(chǎn)生的誘導(dǎo)[J].植物保護(hù)學(xué)報(bào),2001,28(4): 318–324 Gong P,Zhang X X.The inducement of temperature and photoperiod to produce sexuales ofAphis gossypiiglover[J].Acta Phytophylacica Sinica,2001,28(4): 318–324

[16]徐蕾,趙季秋,許國(guó)慶,等.室內(nèi)人工誘導(dǎo)大豆蚜產(chǎn)生有性世代的研究[J].應(yīng)用昆蟲(chóng)學(xué)報(bào),2015,52(6): 1429–1437 Xu L,Zhao J Q,Xu G Q,et al.Artificial induction of sexuales inAphis glycines(Hemiptera: Aphididae)[J].Chinese Journal of Applied Entomology,2015,52(6): 1429–1437

[17]王玲,楊帥,趙奎軍,等.低溫和光周期對(duì)不同地理種群大豆蚜蚜型的影響[J].大豆科學(xué),2014,33(2): 228–231 Wang L,Yang S,Zhao K J,et al.Effect of low temperature and photoperiod on soybean aphid forms from different geographical populations[J].Soybean Science,2014,33(2): 228–231

[18]王小強(qiáng),韓秀楠,曹馨月,等.豌豆品種對(duì)綠色型豌豆蚜種群參數(shù)的影響[J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2013,21(8): 1004–1008 Wang X Q,Han X N,Cao X Y,et al.Effects of different garden pea cultivars on population parameters of green morph of pea aphid (Acyrthosiphon pisum)[J].Chinese Journal of Eco-Agriculture,2013,21(8): 1004–1008

[19]杜軍利,武德功,呂寧,等.不同溫度條件下兩種色型豌豆蚜的種群參數(shù)[J].草業(yè)學(xué)報(bào),2015,24(11): 91–99 Du J L,Wu D G,Lü N,et al.Population parameters of two pea aphid (Acyrthosiphon pisum) color morphs at different temperatures[J].Acta Prataculturae Sinica, 2015,24(11): 91–99

[20]金娟,王森山,賀春貴.西北不同地理種群紅色型豌豆蚜的遺傳多樣性[J].草地學(xué)報(bào),2013,21(2): 406–412 Jin J,Wang S S,He C G.Genetic diversities ofAcyrthosiphon pisum(Harris) (pink form) populations from different geographic regions in the northwest of China[J].Acta Agrestia Sinica, 2013,21(2): 406–412

[21]馬亞玲,劉長(zhǎng)仲.光周期對(duì)兩種色型豌豆蚜生長(zhǎng)發(fā)育的影響[J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2014,22(12): 1476–1483 Ma Y L,Liu C Z.Effect of photoperiod on growth and development of two color morphs of pea aphid (Acyrthosiphon pisum)[J].Chinese Journal of Eco-Agriculture,2014,22(12): 1476–1483

[22]Via S.Inducing the sexual forms and hatching the eggs of pea aphids[J].Entomologia Experimentalis et Applicata,1992,65(2): 119–127

[23]Nijhout H F.Control mechanisms of polyphenic development in insects: In polyphenic development,environmental factors alter some aspects of development in an orderly and predictable way[J].Bioscience,1999,49(3): 181–192

[24]Tauber E,Kyriacou B P.Insect photoperiodism and circadian clocks: Models and mechanisms[J].Journal of Biological Rhythms,2001,16(4): 381–390

[25]Saunders D S.Insect Clocks[M].3rd ed.Amsterdam: Elsevier Science,2002

[26]劉健,吳孔明,趙奎軍,等.不同地理種群棉蚜對(duì)溫度和光周期的生態(tài)適應(yīng)性[J].生態(tài)學(xué)報(bào),2003,23(5): 863–869 Liu J,Wu K M,Zhao K J,et al.The ecological adaptability ofAphis gossypiicollected from different climate zones to temperature and photoperiod[J].Acta Ecologica Sinica,2003,23(5): 863–869

[27]郭建青,張洪剛,王振營(yíng),等.光周期和溫度對(duì)亞洲玉米螟滯育誘導(dǎo)的影響[J].昆蟲(chóng)學(xué)報(bào),2013,56(9): 996–1003 Guo J Q,Zhang H G,Wang Z Y,et al.Effects of photoperiod and temperature on diapause induction inOstrinia furnacalis(Lepidoptera: Crambidae)[J].Acta Entomologica Sinica,2013,56(9): 996–1003

[28]武德功,杜軍利,賀春貴,等.綠色豌豆蚜不同地理種群的遺傳多樣性[J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2015,23(4): 465–472 Wu D G,Du J L,He C G,et al.Genetic diversity ofAcyrthosiphon pisum(green form) from different geographical populations[J].Chinese Journal of Eco-Agriculture,2015,23(4): 465–472

[29]Lamb R J,Pointing P J.Sexual morph determination in the aphid,Acyrthosiphon pisum[J].Journal of Insect Physiology,1972,18(10): 2029–2042

[30]Mackay P A,Wellington W G.A comparison of the reproductive patterns of apterous and alate virginoparousAcyrthosiphon pisum(Homoptera: Aphididae)[J].The Canadian Entomologist,1975,107(11): 1161–1166

[31]龔豪,李川,史勝利,等.有翅型與無(wú)翅型棉蚜發(fā)育及繁殖能力比較研究[J].西南農(nóng)業(yè)學(xué)報(bào),2009,22(6): 1785–1788 Gong H,Li C,Shi S L,et al.Difference of development and reproduction ofAphis gossypiiGlover winged and wingless morph[J].Southwest China Journal of Agricultural Sciences,2009,22(6): 1785–1788

[32]劉柱東,李典謨,葛紹奎,等.昆蟲(chóng)的母代效應(yīng)[J].昆蟲(chóng)學(xué)報(bào),2003,46(1): 108–113 Liu Z D,Li D M,Ge S K,et al.Maternal effects in insects[J].Acta Entomologica Sinica,2003,46(1): 108–113

[33]郭玉玲,龐淑婷,Ahmed U A E,等.母代光照經(jīng)歷對(duì)菜蛾盤(pán)絨繭蜂滯育發(fā)生的影響[J].中國(guó)生物防治,2007,23(1): 1–4 Guo Y L,Pang S T,Ahmed U A E,et al.Effect of maternal photoperiod experience on diapause incidence ofCotesia plutellae[J].Chinese Journal of Biological Control,2007,23(1): 1–4

[34]金娟,梁金,賀春貴.苜蓿品種對(duì) 2種色型豌豆蚜的抗生性[J].草業(yè)學(xué)報(bào),2013,22(6): 335–340 Jin J,Liang J,He C G.Antibiosis of five alfalfa cultivars to two colour morphs ofAcyrthosiphon pisum[J].Acta Prataculturae Sinica,2013,22(6): 335–340

Differentiation regularity of Acyrthosiphon pisum (green form) sexual morphs in relation to photoperiods*

ZHANG Tingwei,CHEN Wanbin,LIU Changzhong**,CUI Shaowei,RAO Fuqiang
(College of Plant Protection,Gansu Agricultural University,Lanzhou 730070,China)

In order to clarify population reproduction strategy and differentiation regularity ofAcyrthosiphon pisum(green form) sexual morph in relation to photoperiods,the pea aphid sexual morphs were observed after 2 continuous generations at different photoperiods (8L∶16D,10L∶14D,12L∶12D and 14L∶10D) in the lab.The results showed that the ovipara of pea aphid in Lanzhou area were wingless,while males were winged.Photoperiod had a significant effect on the differentiation of pea aphid sexual morphs (P＜ 0.05).No sexual morph was produced at 14L∶10D photoperiod,but a lot were differentiated under the other photoperiods (8L∶16D,10L∶14D and 12L∶12D).In the shorter photoperiod conditions for the 3rdgeneration,percent male sexual morphs gradually diminished while percent ovipara gradually increased with decreasingillumination time.Percent male was highest (30.39%) at 12L∶12D photoperiod,while percent ovipara was lowest (55.67%) at that photoperiod.At 10L∶14D photoperiod,percent vivipara was highest while percent sexual morphs was lowest.Then at 8L∶16D photoperiod,no male was produced and percent ovipara was highest (90.76%) in the 3rdgeneration.The reproductive period of the 1stgeneration also had a significant effect on the differentiation of sexual morphs in the 3rdgeneration under similar short illumination conditions (P＜ 0.05).Meanwhile,there was a certain time sequence in which pea aphid sexual morphs were differentiated (first ovipara and last males) with some vivipara transitioning to sexual morphs.Hence the critical photoperiod of green pea aphid was between 12 to 14 hours of illumination.A relatively longer illumination was advantageous to the differentiation of males,while short illumination favored higher proportion of ovipara.The results suggested that change in photoperiod was a critical factor of reproductive plasticity.The reproduction of pea aphid sexual morphs and the illumination length maternal pea aphid experienced influenced sexual differentiation of offsprings.

Acyrthosiphon pisum(Harris); Green form; Photoperiod; Male aphid; Ovipara; Vivipara

S435.29

: A

: 1671-3990(2017)02-0166-06

10.13930/j.cnki.cjea.160663

張廷偉,陳萬(wàn)斌,劉長(zhǎng)仲,崔少偉,饒福強(qiáng).不同光周期條件下綠色型豌豆蚜性蚜分化規(guī)律研究[J].中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào),2017,25(2): 166-171

Zhang T W,Chen W B,Liu C Z,Cui S W,Rao F Q.Differentiation regularity ofAcyrthosiphon pisum(green form) sexual morphs in relation to photoperiods[J].Chinese Journal of Eco-Agriculture,2017,25(2): 166-171

* 國(guó)家自然科學(xué)基金項(xiàng)目(31260433)和甘肅農(nóng)業(yè)大學(xué)盛彤笙創(chuàng)新基金項(xiàng)目(GSAU-STS-1419)資助

** 通訊作者: 劉長(zhǎng)仲,主要從事昆蟲(chóng)生態(tài)及害蟲(chóng)綜合治理研究。E-mail: liuchzh@gsau.edu.cn

張廷偉,研究方向?yàn)樽魑锉Ｗo(hù)。E-mail: zhangtw@gsau.edu.cn

2016-07-28 接受日期: 2016-09-14

* The study was supported by the National Natural Science Foundation of China (31260433) and the SHENG Tongsheng Science and Technology Innovation Foundation of Gansu Agricultural University (GSAU-STS-1419).

** Corresponding author,E-mail: liuchzh@gsau.edu.cn

Received Jul.28,2016; accepted Sep.14,2016