蘇 睿，秦毅恒，梁 鋮，董 坤，張學(xué)文*

(1.云南省農(nóng)業(yè)科學(xué)院蠶桑蜜蜂研究所，云南蒙自 661101；2.云南農(nóng)業(yè)大學(xué)食品科學(xué)技術(shù)學(xué)院，東方蜜蜂研究所，昆明 650201)

不同濃度復(fù)合液體維生素對(duì)凹紋胡蜂訪(fǎng)問(wèn)偏好的影響

蘇 睿1，秦毅恒1，梁 鋮1，董 坤2，張學(xué)文1*

(1.云南省農(nóng)業(yè)科學(xué)院蠶桑蜜蜂研究所，云南蒙自 661101；2.云南農(nóng)業(yè)大學(xué)食品科學(xué)技術(shù)學(xué)院，東方蜜蜂研究所，昆明 650201)

本研究旨在為配制凹紋胡蜂VespauelutinaaurariaSmith的人工飼料提供基本理論依據(jù)。2015年7月20日至7月24日，室外招引凹紋胡蜂，以分別添加了2‰、6‰和10‰復(fù)合液體維生素的40%蔗糖溶液作為試驗(yàn)組，相同濃度的蔗糖溶液作為對(duì)照，觀(guān)察其對(duì)4組不同食物的訪(fǎng)問(wèn)次數(shù)和訪(fǎng)問(wèn)時(shí)間。結(jié)果表明，凹紋胡蜂對(duì)2‰復(fù)合液體維生素組訪(fǎng)問(wèn)次數(shù)最高，極顯著高于對(duì)照、6‰和10‰維生素組(P<0.01)，而對(duì)含有10‰復(fù)合液體維生素的糖溶液組訪(fǎng)問(wèn)次數(shù)最低。此外，凹紋胡蜂訪(fǎng)問(wèn)2‰維生素組和6‰維生素組的時(shí)間差異不顯著(P>0.05)，但均極顯著高于對(duì)照組和10‰維生素組(P<0.01)。研究結(jié)果表明，凹紋胡蜂對(duì)添加了2‰維生素的蔗糖溶液具有顯著偏好。

凹紋胡蜂；飼料；訪(fǎng)問(wèn)次數(shù)；訪(fǎng)問(wèn)時(shí)間

凹紋胡蜂VespauelutinaaurariaSmith是一種重要的野生經(jīng)濟(jì)昆蟲(chóng)(李鐵生，1985；馮穎等，2001)，隸屬于胡蜂科Polistes胡蜂屬Vespa(李鐵生，1985)。目前國(guó)內(nèi)外有關(guān)胡蜂的報(bào)道主要集中在其生物學(xué)特性、藥用價(jià)值、生物防治及采集行為生態(tài)學(xué)等方面(董大志和王云珍, 1989; 馮穎等, 2001; D'adamo and Lozada, 2010; Desouhantetal., 2010; Schueller and Jeanne, 2012; Lópezetal., 2013; Lozada and D'adamo, 2014; Santoroetal., 2015)，而對(duì)其飼料的研究極其匱乏，僅有個(gè)別報(bào)道探索了日糧蛋白對(duì)胡蜂存活時(shí)間的影響(Soyelu, 2013)。事實(shí)上，關(guān)于蜂類(lèi)飼料的研究也僅集中在蜜蜂和熊蜂方面(吳杰等, 2009; 馮倩倩等, 2011; 劉俊峰等, 2011; 鄭本樂(lè)等, 2012; Pasqualeetal., 2013; 徐希蓮等, 2013; Vanderplancketal., 2014; Hill and Burdett, 2015; 蓋琴寶等, 2015)。

盡管凹紋胡蜂具有較高的生物防治、藥用及食用價(jià)值(李鐵生，1985；陳玉慧等，1997)，然而人們對(duì)其關(guān)注度遠(yuǎn)遠(yuǎn)不夠，僅有個(gè)別報(bào)道初步探索了其生物學(xué)特性、飼養(yǎng)管理及人工飼料等方面(董大志和王云珍，1989；鄭國(guó)權(quán)和譚墾，2008)。凹紋胡蜂嗜食蝗蟲(chóng)、蒼蠅、蜻蜓、蟋蟀及膜翅目和鱗翅目中的多種幼蟲(chóng)以獲得蛋白質(zhì)和維生素等營(yíng)養(yǎng)成分(董大志和王云珍，1989)。據(jù)報(bào)道，含糖量較高的水果(如蘋(píng)果、桃、梨)、糖蜜和低脂肪肉類(lèi)(如兔肉)可作為凹紋胡蜂人工繁育的飼料(董大志和王云珍，1989)。但其研究缺乏有關(guān)凹紋胡蜂繁育各階段所需飼料及不同飼料對(duì)其生長(zhǎng)發(fā)育影響的深入探討，關(guān)于其飼料的配制缺乏理論依據(jù)，難以實(shí)現(xiàn)其商業(yè)化飼養(yǎng)。維生素作為凹紋胡蜂的重要營(yíng)養(yǎng)物質(zhì)，對(duì)維持蜂群的正常生長(zhǎng)、發(fā)育和繁殖有著不可替代的作用。花蜜和水果是凹紋胡蜂獲得維生素的主要途徑，然而當(dāng)外界缺乏攝食來(lái)源時(shí)，配制維生素人工代用飼料極其重要。闡明凹紋胡蜂對(duì)不同濃度維生素的訪(fǎng)問(wèn)偏好是合理配制其維生素飼料的關(guān)鍵。

本研究旨在通過(guò)比較含有不同濃度復(fù)合液體維生素的蔗糖溶液與純蔗糖溶液對(duì)凹紋胡蜂訪(fǎng)問(wèn)次數(shù)及訪(fǎng)問(wèn)時(shí)間的影響，探明凹紋胡蜂對(duì)不同濃度維生素的攝食偏好，以期為凹紋胡蜂的規(guī)模化人工飼養(yǎng)、科學(xué)配制維生素飼料提供基礎(chǔ)數(shù)據(jù)。

1 材料與方法

1.1 試驗(yàn)蜂群

本試驗(yàn)所用蜂群為云南省農(nóng)業(yè)科學(xué)院蠶桑蜜蜂研究所提供的凹紋胡蜂，共計(jì)6群。試驗(yàn)中所用的復(fù)合液體維生素購(gòu)買(mǎi)于上海陽(yáng)農(nóng)生物科技有限公司(主要成分為維生素A、維生素D、維生素E、維生素B1、維生素B2、維生素B6、煙酰胺等多種維生素)。

1.2 方法

1.2.1配制溶液

準(zhǔn)確稱(chēng)取1600 g蔗糖，加入2400 mL蒸餾水充分溶解，配制成濃度為40%(g/100 g)的蔗糖溶液作為母液，并以此母液作為對(duì)照組胡蜂的食物(以下簡(jiǎn)寫(xiě)為蔗糖溶液)；用移液槍依次準(zhǔn)確量取1 mL、3 mL和5 mL的復(fù)合液體維生素，分別加入3個(gè)500 mL的容量瓶中，蔗糖母液定容，配制成維生素濃度分別為2‰、6‰和10‰的復(fù)合液體維生素蔗糖混合溶液(以下分別簡(jiǎn)寫(xiě)為2‰、6‰和10‰維生素溶液)，作為試驗(yàn)組胡蜂的食物。

1.2.2偏好試驗(yàn)

2015年7月20日至7月24日，開(kāi)展偏好試驗(yàn)。試驗(yàn)設(shè)4個(gè)處理，分別在4個(gè)飼喂器中加入10 mL不同處理組的食物。為消除位置效應(yīng)對(duì)其訪(fǎng)問(wèn)偏好的影響，試驗(yàn)設(shè)置4個(gè)取食點(diǎn)(重復(fù))，每個(gè)取食點(diǎn)間隔5 m，試驗(yàn)時(shí)每個(gè)取食點(diǎn)同時(shí)擺放4個(gè)飼喂器，盛以不同處理組的食物，在1個(gè)取食點(diǎn)試驗(yàn)記錄完畢后，隨機(jī)更換取食點(diǎn)和新的飼喂器及其擺放次序；每個(gè)取食點(diǎn)4組飼喂器擺放次序見(jiàn)表1，飼喂器間隔30 cm；每個(gè)取食點(diǎn)記錄時(shí)，具體記錄方法為：每天8 ∶30-11 ∶00和14 ∶30-17 ∶00每隔10 min記錄一次單位時(shí)間內(nèi)(30 min)凹紋胡蜂對(duì)4個(gè)處理組飼喂器的訪(fǎng)問(wèn)次數(shù)及其訪(fǎng)問(wèn)時(shí)間。

表1 飼喂器擺放次序

1.3 數(shù)據(jù)處理

用Excel 2003整理數(shù)據(jù)，繪制圖表；運(yùn)用SPSS 20.0軟件系統(tǒng)對(duì)4種處理下凹紋胡蜂的訪(fǎng)問(wèn)次數(shù)和訪(fǎng)問(wèn)時(shí)間進(jìn)行方差分析，多重比較采用LSD法。

2 結(jié)果與分析

4種處理?xiàng)l件下凹紋胡蜂對(duì)不同濃度維生素的訪(fǎng)問(wèn)次數(shù)如圖1所示。從圖1可以看出，凹紋胡蜂對(duì)2‰維生素的訪(fǎng)問(wèn)次數(shù)高達(dá)22.22±2.33次(n=32)，極顯著高于對(duì)照組(14.41±1.87，n=32，P=0.0020)、6‰維生素組(15.03±1.49，n=32，P=0.0040)和10‰維生素組(8.22±0.98，n=32，P<0.01)；而對(duì)10‰維生素組的訪(fǎng)問(wèn)次數(shù)最低，極顯著低于其余3組(P<0.01)；但凹紋胡蜂對(duì)6‰維生素組的訪(fǎng)問(wèn)次數(shù)與對(duì)照組差異不顯著(n=32，P>0.05)。

圖1 凹紋胡蜂對(duì)不同濃度維生素的訪(fǎng)問(wèn)次數(shù)Fig.1 The foraging frequency of wasp on different concentration vitamin注：CK，40%蔗糖溶液；2‰、6‰和10‰維生素分別代表40%蔗糖溶液中含有2‰、6‰和10‰的復(fù)合液體維生素；小寫(xiě)字母表示0.05水平上不同處理組之間訪(fǎng)問(wèn)次數(shù)的顯著性差異，相同小寫(xiě)字母表示差異不顯著(P>0.05)，不同字母表示差異極顯著(P<0.01)；下同。Note: CK, 40% sucrose solution; 2‰、6‰ and 10‰ vitamin, 2‰、6‰ and 10‰ multiple vitamins added into 40% sucrose solution, respectively; Lowercase letters mean significant difference at 0.05 level foraging frequency of wasp on different treatments; the same small letter mean no significant difference (P>0.05), while the alternate small letter mean extremely significant difference (P<0.01); the same as below.

從凹紋胡蜂對(duì)不同濃度維生素的平均訪(fǎng)問(wèn)時(shí)間來(lái)看(見(jiàn)圖2)，盡管其對(duì)2‰維生素組(27.42±0.38 s，n=397)和6‰維生素組(27.64±0.54 s，n=241)的平均訪(fǎng)問(wèn)時(shí)間差異不顯著(P=0.7340)，但二者均極顯著高于對(duì)照組(24.67±0.50 s，n=212)和10‰維生素組(23.87±0.85 s，n=119)(P<0.01)，這表明與純蔗糖溶液相比凹紋胡蜂更喜歡添加了復(fù)合維生素的蔗糖溶液，但當(dāng)復(fù)合維生素的濃度達(dá)到10‰時(shí)，凹紋胡蜂的平均訪(fǎng)問(wèn)時(shí)間最短。

圖2 凹紋胡蜂對(duì)不同濃度維生素的平均取食時(shí)間Fig.2 The average foraging time of wasp fed with different concentration vitamins

試驗(yàn)中觀(guān)察到，當(dāng)環(huán)境中同時(shí)存在4種不同處理組的食物時(shí)，第一次來(lái)訪(fǎng)的凹紋胡蜂需要通過(guò)在4個(gè)飼喂器間徘徊選擇，才能最終選定其偏好的食物，而一旦選定就會(huì)頻繁返回到其偏好的飼喂器取食。對(duì)于其厭惡的食物，凹紋胡蜂往往只訪(fǎng)問(wèn)1-3 s，隨即重新選擇飼喂器進(jìn)行取食；而對(duì)于其偏愛(ài)的食物，訪(fǎng)問(wèn)時(shí)間長(zhǎng)達(dá)45.7-63 s。由表2可知，與對(duì)照組相比，凹紋胡蜂訪(fǎng)問(wèn)2‰維生素的時(shí)間急劇增加并達(dá)到最大；而隨著維生素濃度增加，凹紋胡蜂訪(fǎng)問(wèn)的最長(zhǎng)時(shí)間和最短時(shí)間均急劇縮短，當(dāng)維生素濃度達(dá)到10‰時(shí)，其最短訪(fǎng)問(wèn)時(shí)間僅為2.6 s。

表2 凹紋胡蜂對(duì)不同濃度維生素的最短和最長(zhǎng)訪(fǎng)問(wèn)時(shí)間

綜合來(lái)看，凹紋胡蜂對(duì)2‰維生素組和6‰維生素組的訪(fǎng)問(wèn)次數(shù)和訪(fǎng)問(wèn)時(shí)間均極顯著高于對(duì)照組，這暗示凹紋胡蜂偏好添加了2‰復(fù)合維生素的糖溶液；但其對(duì)10‰維生素組的訪(fǎng)問(wèn)次數(shù)最低，訪(fǎng)問(wèn)時(shí)間也最短，這可能是因?yàn)楦邼舛鹊膹?fù)合維生素影響了食物的口感，從而導(dǎo)致其不再積極取食。

3 結(jié)論與討論

3.1 凹紋胡蜂食物搜尋定位能力

試驗(yàn)中觀(guān)察到，一旦凹紋胡蜂對(duì)環(huán)境中同時(shí)存在的4組食物(含有0‰、2‰、6‰、10‰維生素的蔗糖溶液)最終做出選擇后，若更換取食點(diǎn)和食物擺放次序，凹紋胡蜂仍然會(huì)迅速找到2‰維生素組；甚至該組食物已被采食完，凹紋胡蜂仍然會(huì)優(yōu)先訪(fǎng)問(wèn)該組食物(2‰維生素組)，只有經(jīng)過(guò)幾次嘗試仍未覓到食物時(shí)，凹紋胡蜂才會(huì)取食其它組食物。這暗示，凹紋胡蜂對(duì)食物具有一定的搜尋定位能力，但若加強(qiáng)干擾(如設(shè)置幾個(gè)不同的取食點(diǎn)，同時(shí)擺放不同排列次序的維生素糖溶液、增加其選擇機(jī)會(huì)；加大飼喂盤(pán)間距等)，其對(duì)食物的定位能力是否受到影響，有待進(jìn)一步研究。據(jù)報(bào)道，群居性胡蜂會(huì)受以往采集經(jīng)驗(yàn)的影響，頻繁回到之前已經(jīng)成功覓食并且能夠重復(fù)取食的同一種食物資源采集點(diǎn)(Sabrinaetal., 2014; Santoroetal., 2015)。本試驗(yàn)也驗(yàn)證了這一點(diǎn)。事實(shí)上，群居性胡蜂是一種典型的雜食蜂并且會(huì)采用各種不同的機(jī)制進(jìn)行定位并篩選所需的食物資源(Lozada and D'adamo, 2011)。然而，群居性的凹紋胡蜂是如何對(duì)食物進(jìn)行定位目前仍尚未可知。

據(jù)報(bào)道，社會(huì)性蜜蜂具有將巢內(nèi)的食物氣味與野外蜜源進(jìn)行聯(lián)想學(xué)習(xí)的能力，這對(duì)于其招募同伴外出采集是必要的(Santoroetal., 2015)，這一點(diǎn)在德國(guó)胡蜂Vespulagermanica和Polybiaoccidentalis上也得到了驗(yàn)證(Schuelleretal., 2010; Tayloretal., 2010; Tayloretal., 2011; Tayloretal., 2012)。最近的研究首次闡明了當(dāng)外界存在可利用的食物資源時(shí)，通過(guò)化學(xué)信號(hào)，普通胡蜂Vespulavulgaris將食物資源信息傳遞給巢內(nèi)同伴，而這種信號(hào)能夠讓沒(méi)有經(jīng)驗(yàn)的胡蜂對(duì)食物資源進(jìn)行定位(Santoroetal., 2015)。此外，蜜蜂利用舞蹈語(yǔ)言向同伴傳遞食物信息并招募同伴外出采集，然而群居性的胡蜂通常不會(huì)招募同伴到有食物的地方(Santoroetal., 2015)。目前，金環(huán)胡蜂Vespamandarinia和普通胡蜂是已知的兩種能夠利用信號(hào)進(jìn)行招募的群居性胡蜂(Santoroetal., 2015)。本試驗(yàn)中，盡管凹紋胡蜂能迅速定位訪(fǎng)問(wèn)過(guò)的飼喂盤(pán)，但來(lái)訪(fǎng)問(wèn)的同一群凹紋胡蜂往往只有幾頭，另外，試驗(yàn)中觀(guān)察到，新加入采集的凹紋胡蜂第一次訪(fǎng)問(wèn)飼喂盤(pán)時(shí)，對(duì)食物沒(méi)有偏好，而是經(jīng)過(guò)幾次徘徊選擇之后才對(duì)食物表現(xiàn)出偏好，這暗示凹紋胡蜂可能并不能就糖類(lèi)食物資源信息進(jìn)行巢內(nèi)交流、招募同伴。

3.2 凹紋胡蜂對(duì)維生素的選擇偏好

本試驗(yàn)結(jié)果表明，凹紋胡蜂能夠識(shí)別并對(duì)環(huán)境中同時(shí)存在的4種不同濃度的維生素糖溶液具有不同的訪(fǎng)問(wèn)偏好。試驗(yàn)中連續(xù)3 d觀(guān)察發(fā)現(xiàn)，凹紋胡蜂對(duì)2‰維生素組的取食量最多，并且對(duì)該組食物的取食表現(xiàn)出非隨機(jī)性的聚集分布(多達(dá)4-6頭)，最先采食完的概率為75%，而對(duì)10‰維生素組的取食量最少，最后采食完。綜合訪(fǎng)問(wèn)次數(shù)和取食量來(lái)看，含有2‰維生素的糖溶液對(duì)凹紋胡蜂具有更強(qiáng)的吸引力。然而，試驗(yàn)中凹紋胡蜂重復(fù)訪(fǎng)問(wèn)同一飼喂盤(pán)，暗示凹紋胡蜂對(duì)維生素糖溶液的選擇偏好似乎受其覓食經(jīng)驗(yàn)的影響。有趣的是，這種覓食經(jīng)驗(yàn)并未傳遞給巢內(nèi)其它同伴，因?yàn)樾录尤氩杉陌技y胡蜂仍然需要通過(guò)多次選擇，才會(huì)最終選擇2‰維生素組。有文獻(xiàn)報(bào)道，經(jīng)過(guò)飼喂訓(xùn)練的德國(guó)胡蜂會(huì)根據(jù)覓食經(jīng)驗(yàn)頻繁訪(fǎng)問(wèn)已經(jīng)采集過(guò)的食物，并且這種記憶能夠保持1 h(Sabrinaetal., 2014)。而凹紋胡蜂對(duì)2‰維生素的糖溶液偏好的記憶能夠保持多久尚不明確。

另外，本試驗(yàn)中，無(wú)論更換取食點(diǎn)還是食物的擺放次序，凹紋胡蜂都能迅速找到添加了2‰維生素的糖溶液，暗示其對(duì)食物味道具有偏好，但對(duì)食物位置沒(méi)有偏好。據(jù)報(bào)道，無(wú)刺蜂Tetragonulabiroi喜歡訪(fǎng)問(wèn)距離巢穴最近(1 m)或者距離地面1 m高的飼喂盤(pán)，這與本試驗(yàn)凹紋胡蜂對(duì)食物位置沒(méi)有偏好的結(jié)果不同，這可能與最優(yōu)采集理論、無(wú)刺蜂對(duì)食物位置的記憶或其同伴離開(kāi)飼喂盤(pán)時(shí)做的化學(xué)標(biāo)記有關(guān)(Ciaretal., 2013)。而本試驗(yàn)中凹紋胡蜂頻繁取食同一食物，是否與其離開(kāi)飼喂盤(pán)時(shí)做了標(biāo)記有關(guān)，有待進(jìn)一步驗(yàn)證。

總體來(lái)看，凹紋胡蜂喜歡添加了維生素的糖溶液，但當(dāng)維生素濃度達(dá)到10‰時(shí)，凹紋胡蜂對(duì)10‰維生素組的訪(fǎng)問(wèn)次數(shù)和取食量都低于對(duì)照組，這可能是因?yàn)樘侨芤褐刑砑?0‰的維生素適口性變差。因此，筆者建議今后在配制凹紋胡蜂的維生素代用飼料時(shí)，維生素的添加水平為2‰。

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Effectofdifferentmultiplevitaminsconcentrationonthevisitingpreferenceofwasp

SU Rui1, QIN Yi-Heng1, LIANG Cheng1, DONG Kun2, ZHANG Xue-Wen1*

(1. Institute of Sericulture and Apiculture, Yunnan Academy of Agricultural Sciences, Mengzi 661101, Yunnan Province, China; 2. Eastern Bee Research Institute, College of Food Science and Technology, Yunnan Agricultural University, Kunming 650201, China)

This experiment was conducted to investigate the effect of different amount of multiple vitamins on the foraging preference of wasps. We hope that our work could provide a theoretical basis for further making artificial diet of wasps. Here, we investigated foraging frequency and foraging time of waspsVespauelutinaaurariaSmith to asses their visiting preference when foraging on 4 food groups of 40% sucrose solutions that were added with 0‰, 2‰, 6‰, 10‰ of multiple vitamins, respectively, from July 20 to July 24, 2015. Four feeders containing these 4 food groups, respectively, were in a line to attract wasp visits. The results demonstrated that the foraging frequency of wasps feeding on the 2‰ multiple vitamins group was very significantly higher than that of wasps feeding on the other groups (sucrose solutions added with 0‰, 6‰ and 10‰ multiple vitamins, respectively) (P<0.01). Among the four groups, there was lowest foraging frequency for wasps feeding on the group of sucrose solution added with 10‰ multiple vitamins. According to the average foraging time, wasps preferred to the two food groups contained 2‰ and 6‰ multiple vitamins, and the foraging time of wasps feeding on the two groups was very significantly longer than the time of wasps feeding on the control group and 10‰ multiple vitamins group (P<0.01), but there was no significant difference between the first two groups (P>0.05). It is concluded that the visiting preferences of wasp vary on different concentrations of multiple vitamins in sucrose solutions. And wasps prefer to the sucrose solution added with 2‰ multiple vitamins, according to a comprehensive comparison for foraging frequency, foraging time and feeding amount of wasps foraging activities on different feeders. Our work sheds light on the foraging preference of wasp for different concentrations of multiple vitamins, which were previously seemed be neglected. However, much work remains to be done in order to clarify the chemical composition of wasp artificial diet for the commercialized breeding.

VespauelutinaaurariaSmith; artificial diet; foraging frequency; foraging time

蘇睿，秦毅恒，梁鋮,等.不同濃度復(fù)合液體維生素對(duì)凹紋胡蜂訪(fǎng)問(wèn)偏好的影響[J].環(huán)境昆蟲(chóng)學(xué)報(bào)，2017,39(5):1142-1147.

Q968.1

A

1674-0858(2017)05-1142-06

國(guó)家蜂產(chǎn)業(yè)技術(shù)體系專(zhuān)項(xiàng)(CARS-45-SYZ17)；國(guó)家自然科學(xué)基金項(xiàng)目(31460577，31572339)

蘇睿，女，1986年生，陜西潼關(guān)人，碩士，研究實(shí)習(xí)員，研究方向?yàn)槊鄯涫诜?，E-mail：surui13572630023@163.com

*通訊作者Author for correspondence, E-mail: zxw216226@163.com

Received: 2016-06-22; 接受日期Accepted: 2016-08-20