小菊花部特征及花冠精油组分与访花昆虫的相关性
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摘要
菊花(Dendranthema morifolium)是异花授粉植物,具有自交不亲和的特性。其自然授粉的主要媒介为蜂类和蝶类。本项研究以南京农业大学中国菊花种质资源保存中心内栽培小菊为对象,进行了小菊化学生态学及传粉生物学研究;分别于2006年6月-2006年10月与2007年6月-2007年10月进行了野外调查,之后进行了室内分析和昆虫生物测定。研究内容包括:访花昆虫的种类、行为及活动规律:花部特征、花冠精油组分的鉴定和分析比较;利用“Y”型嗅觉仪及触角电位技术(EAG)研究了访花昆虫对不同花冠精油与几种主要花香物质的反应;以此来分析访花昆虫与小菊花部特征及花冠精油组分之间的关系。结果表明:
(1)小菊的访花昆虫有32种,多数隶属于膜翅目、鳞翅目、双翅目和半翅目;不同月份访花昆虫的种类不同;不同种类访花昆虫的形态特征、访花目的及访花行为有所不同,在植物传粉中的作用也有很大差异;西方蜜蜂(Apis mellifera)为小菊的主要传粉者,其种群数量大,平均访花频率达16朵·min~(-1);晴天,西方蜜蜂和大红蛱蝶的访花蜂数与温度的变化呈显著正相关(r=0.876,0.887);气温骤降、阴天及降雨会影响访花昆虫的种类和数量;同一天内不同时间,访花昆虫的访花频率有差异,一般每天11:00-13:00的访花频率最高。
(2)黄花色系较吸引西方蜜蜂访花;菊花的花朵繁密度、花冠直径、管状花花盘直径均与访花蜂数有显著相关性,而株高与访花蜂数无相关性。大红蛱蝶(Vanessaindica)青睐红色花;其访花蝶数与花朵繁密度、管状花花盘直径均有显著相关性,而花冠直径和株高与访花蝶数无相关性。
(3)经气相色谱-质谱(GC-MS)分析得知,不同菊花品种(或材料)花冠精油的组分不同,其主要成分均为单萜和倍半萜类及其含氧衍生物;并且不同品种(或材料)含有某些特定的花冠精油组分,其中樟脑萜(r=-0.909,p<0.05)、6,10,14-三甲基-2-十五酮(r=0.882,p<0.05)与访花蜂数具一定的相关性,α-萜品醇(r=0.979,p<0.01)、顺柠檬烯氧化物(r=0.979,p<0.01)、金合欢烯氧化物(r=0.979,p<0.01)、p-mentha-6,8-dien-2-ol,cis-(r=0.973,p<0.01,p-menth-l-en-8-ol(r=0.962,p<0.01)和4-萜品烯醇(r=0.957,p<0.05)与访花蝶数也呈一定相关性。
(4)小菊的自然结实率与访花蜂数呈显著相关性(r=0.534,p<0.05)。
(5)西方蜜蜂对不同小菊花冠精油挥发物的嗅觉反应不同,并且随着浓度的减低,正趋性增加。其中,西方蜜蜂对0.1μL·mL~(-1)的'金陵春梦'和'金陵晚霞'挥发油表现出明显的正趋性(选择系数分别为:0.72和0.63)。西方蜜蜂对不同品种(或材料)小菊花冠精油的EAG反应亦有差异,其中对04-1-29、'金陵双秀'小菊的EAG反应值显著大于其他供试小菊(p<0.05),而对'金陵锦袍'和04-8-8小菊的EAG反应值显著小于其他供试小菊(p<0.05)。
(6)西方蜜蜂对不同小菊花香物质组分的EAG反应存在差异,乙酸冰片酯的EAG反应值显著大于其他供试小菊挥发性花香物质组分(p<0.05)。西方蜜蜂对不同小菊花香物质组分的嗅觉反应亦不同,西方蜜蜂对0.01μL·mL~(-1)的石竹烯氧化物表现明显的正趋性(选择系数为0.60),对1μL·mL~(-1)的樟脑则表现明显的负趋性(选择系数为-0.65)。
Chrysanthemum (Dendranthema morifolium) is a self-incompatible allogamous plant. Its main pollination vectors are bees and butterflies. This paper firstly focused on and evaluated the chemical ecology and pollination characteristics of chrysanthemum with small flower type in Chrysanthemum Germplasm Resource Preserving Center of China. Our aim is to investigate and evaluate species and visiting behavior of flower-visiting insects of Dendranthema morifolium during flowering period, and the effects of flower color, corolla density, corolla diameter, tubular flower diameter, plant height and essential oils on attracting insect visitors. The volatiles from eight cultivars (or materials) of chrysanthemum with small flower type were collected by steam distilltion. Behavioral responses and electroantennogram (EAG) of Apis mellifera to the volatiles and several essential oil components of Chrysanthemum with small flower type were investigated respectively. The results as follows:
(1) 32 species, belonging to Hymenoptera, lepidoptera, Diptera and Hemiptera, were observed visiting flowering D. morifolium,the species of visiting insects were not same among different months, and the effects of pollination were significantly affected by morphology, behavior and motive of flower-visiting insects. Apis mellifera was the main pollinator, which has a large population, and visiting frequency of A. mellifera was 16 flowers per min. During sunny days, visiting frequency of A mellifera and Vanessa indica were significantly positive correlation with temperature (r=0.876, 0.887, respectively), while sudden dropping-temperature, cloud and rain had important influences upon the number and species of flower-visiting insects visiting flower. Results of field observation further showed that each day, the visiting frequencies to D. morifolium inflorescences were different in different periods; generally, visiting frequency during period from 11:00 to 13:00 was the highest.
(2) Bees preferred yellow colors flower to others including orange yellow, red, mauve, orange and pink. The number of visiting bees was significantly correlated with corolla density (r=0.715,p<0.01),indicating that the effect of a colony was more significant than the size of a single flower head. For all tested traits of a single flower head, tubular flower diameter had the most significant influence on attracting bees (r=0.622,p<0.01),followedby corolla diameter (r=-0.502,p<0.05).Butterflies preferred red color flowers, and tubular flower diameter significantly influenced the number of pollinating butterflies (r=0.635, p<0.01).Results suggested that rewards from the flowers were the dominating attractive factors to visitors. The effect of single flower head on number of visiting butterflies was more significant than corolla density (r=0.487,p<0.05). In all tested traits of a single flower head, flower color and tubular flower diameter were the main influencing factors on numbers of visiting butterflies.
(3) The main essential oil components,such as camphor (r_(bees)-0.191,p>0.05; r_(butterflies)=0.507,p>0.05) and borneol (r_(bees)=0.354,p>0.05;r_(butterflies)=-0.387,p>0.05) had little correlation with number of visiting insects, whereas some specific essential oil components, such as camphene (r=-0.909, p<0.05), 2-pentadecanone,6,10,14-trimethyl-(r=0.882,p<0.05) showed significant correlations with number of visiting bees, andα-terpineol (r=0.979, p<0.01),cis-limonene oxide (r=0.979, p<0.01),E-farnesene epoxide (r=0.979, p<0.01),p-mentha-6,8-dien-2-ol, cis-(r=0.973,p<0.01),p-menth-1-en-8-ol (r=0.962,p<0.01) and p-menth-1-en-4-ol,[R]-[-]- (r=0.957,p<0.05) showed significant correlations with number of visiting butterflies.
(4) The number of visiting bees was significantly correlated with seed setting percentage (r=0.534, p<0.05) .
(5) The olfactory responses of Apis mellifera to volatiles from different cultivars (or materials) were different, and the positive response became greater as the concentration decreased. Apis mellifera was significantly attracted by 0.1μL/mL~(-1) 'jin ling chun meng' and 'jin ling wan xia' volatiles (coefficient of selectivity are 0.72 and 0.63 ). Apis mellifera also has different responses to volatiles from different cultivars (or materials) of chrysanthemum with small flower type, the EAG values of Apis mellifera treated by volatiles of 04-1-29 and 'jin ling shuang xiu' were significant higher than the other cultivars (or materials) (p<0.05),while the values treated by volatiles of 'jin ling jin pao'and 04-8-8 were lower than the other cultivars (or materials)(p<0.05).
(6) Apis mellifera has different responses to volatiles from different aromatic constituents of chrysanthemum with small flower type, the EAG values of Apis mellifera treated by bomeol acetate were significant higher than the other components (p<0.05).The olfactory responses of Apis mellifera to different components of volatiles were also different, Apis mellifera manifests obvious positive tropism to 0.01μL·mL~(-1)CaryophyIIene oxide (coefficient of selectivity is 0.60),while negative tropism to 1μL·mL~(-1) Camphor (coefficient of selectivity is -0.65).
(1)小菊的访花昆虫有32种,多数隶属于膜翅目、鳞翅目、双翅目和半翅目;不同月份访花昆虫的种类不同;不同种类访花昆虫的形态特征、访花目的及访花行为有所不同,在植物传粉中的作用也有很大差异;西方蜜蜂(Apis mellifera)为小菊的主要传粉者,其种群数量大,平均访花频率达16朵·min~(-1);晴天,西方蜜蜂和大红蛱蝶的访花蜂数与温度的变化呈显著正相关(r=0.876,0.887);气温骤降、阴天及降雨会影响访花昆虫的种类和数量;同一天内不同时间,访花昆虫的访花频率有差异,一般每天11:00-13:00的访花频率最高。
(2)黄花色系较吸引西方蜜蜂访花;菊花的花朵繁密度、花冠直径、管状花花盘直径均与访花蜂数有显著相关性,而株高与访花蜂数无相关性。大红蛱蝶(Vanessaindica)青睐红色花;其访花蝶数与花朵繁密度、管状花花盘直径均有显著相关性,而花冠直径和株高与访花蝶数无相关性。
(3)经气相色谱-质谱(GC-MS)分析得知,不同菊花品种(或材料)花冠精油的组分不同,其主要成分均为单萜和倍半萜类及其含氧衍生物;并且不同品种(或材料)含有某些特定的花冠精油组分,其中樟脑萜(r=-0.909,p<0.05)、6,10,14-三甲基-2-十五酮(r=0.882,p<0.05)与访花蜂数具一定的相关性,α-萜品醇(r=0.979,p<0.01)、顺柠檬烯氧化物(r=0.979,p<0.01)、金合欢烯氧化物(r=0.979,p<0.01)、p-mentha-6,8-dien-2-ol,cis-(r=0.973,p<0.01,p-menth-l-en-8-ol(r=0.962,p<0.01)和4-萜品烯醇(r=0.957,p<0.05)与访花蝶数也呈一定相关性。
(4)小菊的自然结实率与访花蜂数呈显著相关性(r=0.534,p<0.05)。
(5)西方蜜蜂对不同小菊花冠精油挥发物的嗅觉反应不同,并且随着浓度的减低,正趋性增加。其中,西方蜜蜂对0.1μL·mL~(-1)的'金陵春梦'和'金陵晚霞'挥发油表现出明显的正趋性(选择系数分别为:0.72和0.63)。西方蜜蜂对不同品种(或材料)小菊花冠精油的EAG反应亦有差异,其中对04-1-29、'金陵双秀'小菊的EAG反应值显著大于其他供试小菊(p<0.05),而对'金陵锦袍'和04-8-8小菊的EAG反应值显著小于其他供试小菊(p<0.05)。
(6)西方蜜蜂对不同小菊花香物质组分的EAG反应存在差异,乙酸冰片酯的EAG反应值显著大于其他供试小菊挥发性花香物质组分(p<0.05)。西方蜜蜂对不同小菊花香物质组分的嗅觉反应亦不同,西方蜜蜂对0.01μL·mL~(-1)的石竹烯氧化物表现明显的正趋性(选择系数为0.60),对1μL·mL~(-1)的樟脑则表现明显的负趋性(选择系数为-0.65)。
Chrysanthemum (Dendranthema morifolium) is a self-incompatible allogamous plant. Its main pollination vectors are bees and butterflies. This paper firstly focused on and evaluated the chemical ecology and pollination characteristics of chrysanthemum with small flower type in Chrysanthemum Germplasm Resource Preserving Center of China. Our aim is to investigate and evaluate species and visiting behavior of flower-visiting insects of Dendranthema morifolium during flowering period, and the effects of flower color, corolla density, corolla diameter, tubular flower diameter, plant height and essential oils on attracting insect visitors. The volatiles from eight cultivars (or materials) of chrysanthemum with small flower type were collected by steam distilltion. Behavioral responses and electroantennogram (EAG) of Apis mellifera to the volatiles and several essential oil components of Chrysanthemum with small flower type were investigated respectively. The results as follows:
(1) 32 species, belonging to Hymenoptera, lepidoptera, Diptera and Hemiptera, were observed visiting flowering D. morifolium,the species of visiting insects were not same among different months, and the effects of pollination were significantly affected by morphology, behavior and motive of flower-visiting insects. Apis mellifera was the main pollinator, which has a large population, and visiting frequency of A. mellifera was 16 flowers per min. During sunny days, visiting frequency of A mellifera and Vanessa indica were significantly positive correlation with temperature (r=0.876, 0.887, respectively), while sudden dropping-temperature, cloud and rain had important influences upon the number and species of flower-visiting insects visiting flower. Results of field observation further showed that each day, the visiting frequencies to D. morifolium inflorescences were different in different periods; generally, visiting frequency during period from 11:00 to 13:00 was the highest.
(2) Bees preferred yellow colors flower to others including orange yellow, red, mauve, orange and pink. The number of visiting bees was significantly correlated with corolla density (r=0.715,p<0.01),indicating that the effect of a colony was more significant than the size of a single flower head. For all tested traits of a single flower head, tubular flower diameter had the most significant influence on attracting bees (r=0.622,p<0.01),followedby corolla diameter (r=-0.502,p<0.05).Butterflies preferred red color flowers, and tubular flower diameter significantly influenced the number of pollinating butterflies (r=0.635, p<0.01).Results suggested that rewards from the flowers were the dominating attractive factors to visitors. The effect of single flower head on number of visiting butterflies was more significant than corolla density (r=0.487,p<0.05). In all tested traits of a single flower head, flower color and tubular flower diameter were the main influencing factors on numbers of visiting butterflies.
(3) The main essential oil components,such as camphor (r_(bees)-0.191,p>0.05; r_(butterflies)=0.507,p>0.05) and borneol (r_(bees)=0.354,p>0.05;r_(butterflies)=-0.387,p>0.05) had little correlation with number of visiting insects, whereas some specific essential oil components, such as camphene (r=-0.909, p<0.05), 2-pentadecanone,6,10,14-trimethyl-(r=0.882,p<0.05) showed significant correlations with number of visiting bees, andα-terpineol (r=0.979, p<0.01),cis-limonene oxide (r=0.979, p<0.01),E-farnesene epoxide (r=0.979, p<0.01),p-mentha-6,8-dien-2-ol, cis-(r=0.973,p<0.01),p-menth-1-en-8-ol (r=0.962,p<0.01) and p-menth-1-en-4-ol,[R]-[-]- (r=0.957,p<0.05) showed significant correlations with number of visiting butterflies.
(4) The number of visiting bees was significantly correlated with seed setting percentage (r=0.534, p<0.05) .
(5) The olfactory responses of Apis mellifera to volatiles from different cultivars (or materials) were different, and the positive response became greater as the concentration decreased. Apis mellifera was significantly attracted by 0.1μL/mL~(-1) 'jin ling chun meng' and 'jin ling wan xia' volatiles (coefficient of selectivity are 0.72 and 0.63 ). Apis mellifera also has different responses to volatiles from different cultivars (or materials) of chrysanthemum with small flower type, the EAG values of Apis mellifera treated by volatiles of 04-1-29 and 'jin ling shuang xiu' were significant higher than the other cultivars (or materials) (p<0.05),while the values treated by volatiles of 'jin ling jin pao'and 04-8-8 were lower than the other cultivars (or materials)(p<0.05).
(6) Apis mellifera has different responses to volatiles from different aromatic constituents of chrysanthemum with small flower type, the EAG values of Apis mellifera treated by bomeol acetate were significant higher than the other components (p<0.05).The olfactory responses of Apis mellifera to different components of volatiles were also different, Apis mellifera manifests obvious positive tropism to 0.01μL·mL~(-1)CaryophyIIene oxide (coefficient of selectivity is 0.60),while negative tropism to 1μL·mL~(-1) Camphor (coefficient of selectivity is -0.65).
引文
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