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虫瘿是某些昆虫通过产卵或取食刺激,诱导寄主植物细胞异常增生而形成的具有特定结构的增生组织[1-4],是昆虫与寄主植物长期协同进化的结果[5-9]。五倍子是一些蚜虫寄生在漆树科盐肤木属(Rhus)植物叶片上所形成的虫瘿,其主要成分为单宁酸,广泛应用于医药、化工和电子等行业,是我国传统的林特产品[10-13]。角倍是五倍子中分布最为广泛、产量最大的种类,由角倍蚜(Schlechtendalia chinensis Bell)寄生在盐肤木(Rhus chinensis Mill)上形成[14-16]。角倍蚜干母对盐肤木的取食部位有着严格的选择[17]。每年春季,角倍蚜干母在盐肤木细嫩的复叶叶翅上取食,刺激叶组织细胞增生膨大,形成具有不规则突起的虫瘿,干母在封闭的瘿内生长和繁殖,个体数量可达到数千至上万头,秋季虫瘿成熟后自然爆裂,蚜虫迁飞回到藓类寄主上越冬[18]。角倍蚜虫瘿作为具有较大生物量的增生组织,悬挂于盐肤木的复叶上,支撑着瘿内成千上万头蚜虫的生长,数量众多的虫瘿和蚜虫是否会对盐肤木枝叶的生长发育产生不利影响,目前尚无相关报道[19]。
本研究从角倍核心分布区的盐肤木林中取样,统计自然迁飞和人工挂袋两种接种方式下,盐肤木单株虫瘿数量、分枝数、复叶数等数量指标,同时测定虫瘿体积、小叶面积和树木地径等性状指标,分析虫瘿对盐肤木生长的影响。研究结果有助于深入理解蚜虫与寄主植物的寄生关系,也可为盐肤木林的管理和五倍子高产培育提供参考。
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在角倍蚜自然迁飞和人工挂袋接种方式下,有虫瘿与无虫瘿盐肤木的生长指标均有显著差异(表 1),其中自然迁飞条件下,有虫瘿盐肤木的平均侧枝数为2.39±0.11个·株-1、梢头数为4.88±0.36个·株-1、复叶数为79.51±6.00个·株-1,显著高于无虫瘿盐肤木的平均侧枝数1.86±0.07个·株-1、梢头数为1.90±0.09个·株-1、复叶数为36.10±2.29个·株-1;在人工挂袋条件下,有虫瘿盐肤木的平均侧枝数为2.59±0.09个·株-1、梢头数为4.95±0.32个·株-1、复叶数为75.67±5.49个·株-1,显著高于无虫瘿盐肤木的平均侧枝数2.06±0.09个·株-1、梢头数为2.48±0.18个·株-1、复叶数为54.83±5.74个·株-1,且两种接种方式之间的无虫瘿盐肤木的生长指标差异不显著。表明两种接种方式下,相比无虫瘿盐肤木,有虫瘿盐肤木的侧枝数、梢头数和复叶数均增多,显示虫瘿对盐肤木的生长有促进作用。
表 1 接种方式和虫瘿对盐肤木枝叶生长的影响分析
Table 1. Analysis on the influence of host plant foliage growth in two inoculation methods
类别
Category地径
Ground diameter/cm侧枝数
Number of lateral branches梢头数
Tips of branches有虫瘿梢
头数
Galled tips of
branches有虫瘿
梢头比
Galled ratio
tips of branches复叶数Number of
compound leaves有虫瘿复叶数
Galled number
of compound
leaves有虫瘿复叶比
Galled ratio of
compound
leaves虫瘿总数
Total number
of galls有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled自然迁飞
Natural migration4.43±
0.14aA3.81±
0.13aB2.39±
0.11aA1.86±
0.07aB4.88±
0.36aA1.90±
0.09aB2.00±
0.11a0.50±
0.04a79.51±
6.00aA36.10±
2.29aB7.67±
0.82a0.10±
0.01a32.80±
3.85a人工挂袋
Artificial aphid bags4.85±
0.21aA4.21±
0.29aA2.59±
0.09aA2.06±
0.09aB4.95±
0.32aA2.48±
0.18bB2.06±
0.11a0.51±
0.03a75.67±
5.49aA54.83±
5.74bB10.02±
1.16a0.15±
0.01b32.73±
4.29at 1.696 1.237 1.406 1.672 0.15 2.968 0.39 0.072 -0.47 3.032 1.56 3.004 -0.013 P 0.093 0.219 0.162 0.097 0.881 0.004 0.697 0.943 0.639 0.003 0.122 0.003 0.99 n 117 115 117 115 117 115 117 117 117 115 117 117 117 注:小写字母和t值表示自然迁飞与人工挂袋差异(纵向比较),大写字母表示有虫瘿与无虫瘿差异(横向比较),n为调查株数。
Note: lower-case letters and t value represent natural migration and artificial pouch difference (vertical comparison), capital letters represent galled and non-galled difference (horizontal comparison), n is the number of Samples investigated. -
比较两种接种方式下的有虫瘿盐肤木,仅单株虫瘿总体积、单片复叶的小叶总面积、有虫瘿复叶比和单位体积虫瘿所需小叶面积有显著差异,其余指标如有虫瘿梢头数、复叶数和虫瘿总数等均无显著差异(表 1、表 2),其中人工挂袋接种时,单株虫瘿总体积为170.92±14.85 cm3、单片复叶的小叶总面积616.26±32.73 cm2,显著高于自然迁飞时的85.82±9.40 cm3和482.81±28.51 cm2,可以看出,人工挂袋的虫瘿的总体积约为自然迁飞的2倍,由于虫瘿体积与重量呈正相关,即产量也约为后者的2倍;同时,人工挂袋的小叶总面积约为自然迁飞的1.3倍,这可能与人工挂袋时虫瘿体积的增加有关。即人工挂袋接种比自然迁飞更能增加虫瘿体积、倍子产量和小叶面积。
表 2 接种方式对虫瘿和盐肤木叶片生长的影响分析
Table 2. Analysis on influences of gall and foliage growth in two inoculation methods
类别Category 小叶数
Number of leaflets/pcs小叶总面积
Total areas of leaflets/cm2复叶虫瘿数
Number of galled on
the compound leave/pc虫瘿总体积
Total volume of galls
/cm3单位体积虫瘿所需小叶面积
Leaf area per unit volume
of gall/cm2有虫瘿Galled 无虫瘿Non-galled 有虫瘿Galled 无虫瘿Non-galled 自然迁飞Natural migration 9.10±0.23aA 8.90±0.33aA 482.81±28.51aA 438.38±35.36aA 6.03±1.04a 85.82±9.40a 8.72±1.33a 人工挂袋Artificial aphid bags 9.60±0.22aA 9.07±0.28aA 616.26±32.73bA 465.29±24.78aB 7.17±0.59a 170.92±14.85b 4.67±0.61b t 1.591 0.383 3.074 0.623 0.947 4.842 -2.767 P 0.117 0.703 0.003 0.535 0.347 0.000 0.008 注:n=60,小写字母和t值表示自然迁飞与人工挂袋差异,大写字母表示有虫瘿与无虫瘿差异。
Note: n=60, lower-case letters and t value represent natural migration and artificial pouch difference, capital letters represent galled and non-galled difference. -
Pearson相关分析结果:在人工挂袋和自然迁飞接种条件下,虫瘿数与梢头数、有虫瘿梢头数、复叶数和有虫瘿复叶数均分别呈极显著正相关(P < 0.01),与地径和侧枝数的相关性不显著(表 3)。反映出两种接种方式下虫瘿数与盐肤木生长指标的关系具有相似的规律,但唯一的区别在于,自然迁飞条件下,虫瘿数与盐肤木地径呈显著正相关(P < 0.05),而人工挂袋条件下,二者的相关性不显著。表明自然迁飞时,地径越大的盐肤木,虫瘿越多,这与前面角倍蚜优先选择地径较大的盐肤木寄生致瘿的结果相一致。
表 3 两种接种方式下盐肤木生长指标与虫瘿数的相关关系
Table 3. Correlations between foliage growth indexes of Rhus chinensis and gall number in two inoculation methods
类别Category 侧枝数
Number of lateral branches梢头数
Tips of branches有虫瘿梢头数
Galled tips of branches复叶数
Number of compound leaves有虫瘿复叶数
Galled number of compound leaves虫瘿数
Number of galls人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration地径Ground diameter 0.335** 0.439** 0.528** 0.705** -0.036 0.109 0.397** 0.746** 0.225 0.240 0.117 0.300* 侧枝数
Number of lateral branches1 1 0.407** 0.364** 0.485** 0.566** 0.427** 0.296* 0.321** 0.221 0.221 0.184 梢头数Tips of branches 1 1 0.295* 0.405** 0.951** 0.952** 0.576** 0.632** 0.413** 0.584** 有虫瘿梢头数
Galled tips of branches1 1 0.366** 0.335* 0.575** 0.622** 0.520** 0.560** 复叶数Number of compound leaves 1 1 0.585** 0.643** 0.414** 0.608** 有虫瘿复叶数Galled number of compound leaves 1 1 0.910** 0.911** 注:人工挂袋n=66,自然迁飞n=51。
Note: artificial pouch, n=66; natural migration, n=51. -
Pearson相关分析结果:在人工挂袋与自然迁飞条件下,复叶虫瘿体积与小叶数、复叶虫瘿数呈极显著正相关(P < 0.01),单位虫瘿体积所需小叶面积与复叶虫瘿数、复叶虫瘿体积呈极显著负相关(P < 0.01)(表 4)。但不同的是,自然迁飞条件下,单位虫瘿体积所需小叶面积与小叶总面积呈极显著正相关(P < 0.01),而人工挂袋时,二者的相关性不显著。表明无论是人工挂袋还是自然迁飞,复叶虫瘿数越多,则虫瘿总体积越大。
表 4 两种接种方式盐肤木叶片与虫瘿生长指标的相关关系
Table 4. Correlations between leaf growth indexes of Rhus chinensis and gall growth indexes in two inoculation methods
类别Category 小叶总面积
Total areas of leaflets复叶虫瘿数
Number of galled on the compound leave复叶虫瘿总体积
Total volume of galls单位虫瘿体积所需小叶面积
Leaf area per unit volume of gall人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration小叶数Number of leaflets 0.411* 0.244 0.296 0.384* 0.483** 0.503** -0.171 -0.358 小叶总面积Total areas of leaflets 1 1 0.278 0.083 0.191 -0.097 0.226 0.475** 复叶虫瘿数
Number of galled on the compound leave1 1 0.783** 0.719** -0.552** -0.457* 复叶虫瘿总体积Total volume of galls 1 1 -0.695** -0.733** 注:n=60。
Note: n=60.人工挂袋和自然迁飞接种方式致瘿的复叶上虫瘿数量与虫瘿个体体积存在显著负相关关系(P < 0.05)。表明单位复叶上虫瘿的平均体积随着虫瘿数量的增加而减少(图 1)。
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自然迁飞条件下:第一轴特征值为4.07,占特征值总和58.21%,累积贡献率为58.21%;第二轴特征值为1.29,占特征值总和18.43%,前两轴累积贡献率为76.65%。提取前两轴就能够比较好的解释原有变量所包含的信息;第一轴反映梢头数和复叶数是影响虫瘿产量的主要因素,可以归纳为“枝叶因子”;第二轴可以明显区分出盐肤木植物性状与虫瘿性状的排序,其中地径具有较高的特征值。
人工挂袋条件下:第一轴特征值为3.70,占特征值总和52.80%,累积贡献率为52.80%;第二轴特征值为1.38,占特征值总和19.65%,前两轴累积贡献率为72.45%,提取前两轴就能够比较好的解释原有变量所包含的信息,PCA排序能较好的反映盐肤木与虫瘿产量之间的相互关系(表 5),第一主轴为有虫瘿复叶数、梢头数和复叶数的变化梯度,反映了虫瘿产量与有虫瘿复叶数、复叶数、梢头数关系的变化梯度;第二轴为地径的变化梯度,与盐肤木的生长性状密切相关。
表 5 两种接种方式的主成分分数和特征值的系数矩阵
Table 5. Coefficient matrix of composition scores and eigenvalue of principal in two inoculation methods
类别Category 自然迁飞
Natural migration人工挂袋
Artificial aphid bags轴Axis 1 轴Axis 2 轴Axis 1 轴Axis 2 地径Ground diameter 0.33 -0.59 0.24 0.59 侧枝数
Number of lateral branches0.26 -0.07 0.31 0.12 梢头数Tips of branches 0.45 -0.25 0.44 0.34 有虫瘿梢头数
Galled tips of branches0.32 0.47 0.33 -0.43 复叶数
Number of compound leaves0.44 -0.29 0.44 0.26 有虫瘿复叶数Galled number of compound leaves 0.41 0.39 0.45 -0.30 虫瘿数Number of galls 0.40 0.35 0.39 -0.42 特征值Characteristic value 4.07 1.29 3.70 1.38 方差的Variance % 58.21 18.43 52.80 19.65 累积Cumulation % 58.21 76.65 52.80 72.45 综合分析表明,两种接种方式下,与虫瘿产量相关的3个主要因子分别是:梢头数、复叶数和有虫瘿复叶数。
两种接种方式下角倍蚜虫瘿对寄主盐肤木枝叶生长的影响
Effects of Aphid Schlechtendalia chinensis Galls on the Foliage Growth of Host Rhus chinensis under Two Inoculation Methods
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摘要:
目的 明确两种接种方式(自然迁飞和人工挂袋)下角倍蚜虫瘿的数量和分布及其对寄主盐肤木枝叶生长的影响。 方法 在自然迁飞和人工挂袋形成虫瘿的盐肤木林内,按随机区组设计设置样地,在样地内随机选取有虫瘿树和无虫瘿树,分别测量和统计盐肤木及虫瘿的生长指标,采用方差分析虫瘿对盐肤木生长的影响。 结果 在自然迁飞和人工挂袋两种接种方式下,盐肤木单株虫瘿数没有显著差异。但人工挂袋时,单片复叶的平均虫瘿总体积为170.92±14.85 cm3、小叶总面积为616.26±32.73 cm2,显著高于自然迁飞形成虫瘿的总体积85.82±9.40 cm3和小叶总面积482.81±28.51 cm2。表明人工挂袋接种可以增加虫瘿的总体积,从而提高角倍产量;同时,较大体积的虫瘿还促进了盐肤木小叶的生长,使小叶面积增大。 结论 角倍蚜虫瘿对寄主盐肤木的枝叶生长具有促进作用,且这种生长促进与虫瘿的体积呈正相关。与自然迁飞接种相比,人工挂袋接种更能发挥盐肤木的生长和结倍潜力,提高角倍产量。 Abstract:Objective Illustrating the aphid gall number and distribution of Schlechtendalia chinensis under two inoculation methods (natural migration and artificial aphid bags) and their effects on the foliage growth of host Rhus chinensis. Method The sample sites and host plants were selected by random group setting in the fields which inoculated by natural migration and hanging aphid bags. The growth indexes and traits of the galls and their host trees were measured or counted. The host plant growths effected by aphid galls were analyzed using variance analysis. Result No significant difference was observed in the gall numbers per tree of the two inoculation methods. However, the average of total gall volume per compound leaf on the trees inoculated by aphid bags was 170.92±14.85 cm3 which was almost twice the average gall volume (85.82±9.40 cm3) on the trees inoculated by natural migration, and the total area of leaflets per compound leaf on the trees inoculated by artificial pouch was 616.26±32.73 cm2 which was greater than the total area of leaflets (482.81±28.51 cm2) on the trees inoculated by natural migration. It was indicated that the total gall volume increased by inoculation of aphid bags, thereby increasing the yield. Moreover, greater galls may cause extra growths of the leaflets and increase the leaflet areas. Conclusion The aphid galls of S. chinensis promote foliage growth to its host plant R. chinensis. And this growth is positively correlated with the gall sizes. Compared to natural migration, the aphid bag inoculation may increase the gall yield and improve the potential ability of gall formation. -
Key words:
- Schlechtendalia chinensis
- / gall
- / Rhus chinensis
- / foliage growth
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表 1 接种方式和虫瘿对盐肤木枝叶生长的影响分析
Table 1. Analysis on the influence of host plant foliage growth in two inoculation methods
类别
Category地径
Ground diameter/cm侧枝数
Number of lateral branches梢头数
Tips of branches有虫瘿梢
头数
Galled tips of
branches有虫瘿
梢头比
Galled ratio
tips of branches复叶数Number of
compound leaves有虫瘿复叶数
Galled number
of compound
leaves有虫瘿复叶比
Galled ratio of
compound
leaves虫瘿总数
Total number
of galls有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled有虫瘿
Galled无虫瘿
Non-galled自然迁飞
Natural migration4.43±
0.14aA3.81±
0.13aB2.39±
0.11aA1.86±
0.07aB4.88±
0.36aA1.90±
0.09aB2.00±
0.11a0.50±
0.04a79.51±
6.00aA36.10±
2.29aB7.67±
0.82a0.10±
0.01a32.80±
3.85a人工挂袋
Artificial aphid bags4.85±
0.21aA4.21±
0.29aA2.59±
0.09aA2.06±
0.09aB4.95±
0.32aA2.48±
0.18bB2.06±
0.11a0.51±
0.03a75.67±
5.49aA54.83±
5.74bB10.02±
1.16a0.15±
0.01b32.73±
4.29at 1.696 1.237 1.406 1.672 0.15 2.968 0.39 0.072 -0.47 3.032 1.56 3.004 -0.013 P 0.093 0.219 0.162 0.097 0.881 0.004 0.697 0.943 0.639 0.003 0.122 0.003 0.99 n 117 115 117 115 117 115 117 117 117 115 117 117 117 注:小写字母和t值表示自然迁飞与人工挂袋差异(纵向比较),大写字母表示有虫瘿与无虫瘿差异(横向比较),n为调查株数。
Note: lower-case letters and t value represent natural migration and artificial pouch difference (vertical comparison), capital letters represent galled and non-galled difference (horizontal comparison), n is the number of Samples investigated.表 2 接种方式对虫瘿和盐肤木叶片生长的影响分析
Table 2. Analysis on influences of gall and foliage growth in two inoculation methods
类别Category 小叶数
Number of leaflets/pcs小叶总面积
Total areas of leaflets/cm2复叶虫瘿数
Number of galled on
the compound leave/pc虫瘿总体积
Total volume of galls
/cm3单位体积虫瘿所需小叶面积
Leaf area per unit volume
of gall/cm2有虫瘿Galled 无虫瘿Non-galled 有虫瘿Galled 无虫瘿Non-galled 自然迁飞Natural migration 9.10±0.23aA 8.90±0.33aA 482.81±28.51aA 438.38±35.36aA 6.03±1.04a 85.82±9.40a 8.72±1.33a 人工挂袋Artificial aphid bags 9.60±0.22aA 9.07±0.28aA 616.26±32.73bA 465.29±24.78aB 7.17±0.59a 170.92±14.85b 4.67±0.61b t 1.591 0.383 3.074 0.623 0.947 4.842 -2.767 P 0.117 0.703 0.003 0.535 0.347 0.000 0.008 注:n=60,小写字母和t值表示自然迁飞与人工挂袋差异,大写字母表示有虫瘿与无虫瘿差异。
Note: n=60, lower-case letters and t value represent natural migration and artificial pouch difference, capital letters represent galled and non-galled difference.表 3 两种接种方式下盐肤木生长指标与虫瘿数的相关关系
Table 3. Correlations between foliage growth indexes of Rhus chinensis and gall number in two inoculation methods
类别Category 侧枝数
Number of lateral branches梢头数
Tips of branches有虫瘿梢头数
Galled tips of branches复叶数
Number of compound leaves有虫瘿复叶数
Galled number of compound leaves虫瘿数
Number of galls人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration地径Ground diameter 0.335** 0.439** 0.528** 0.705** -0.036 0.109 0.397** 0.746** 0.225 0.240 0.117 0.300* 侧枝数
Number of lateral branches1 1 0.407** 0.364** 0.485** 0.566** 0.427** 0.296* 0.321** 0.221 0.221 0.184 梢头数Tips of branches 1 1 0.295* 0.405** 0.951** 0.952** 0.576** 0.632** 0.413** 0.584** 有虫瘿梢头数
Galled tips of branches1 1 0.366** 0.335* 0.575** 0.622** 0.520** 0.560** 复叶数Number of compound leaves 1 1 0.585** 0.643** 0.414** 0.608** 有虫瘿复叶数Galled number of compound leaves 1 1 0.910** 0.911** 注:人工挂袋n=66,自然迁飞n=51。
Note: artificial pouch, n=66; natural migration, n=51.表 4 两种接种方式盐肤木叶片与虫瘿生长指标的相关关系
Table 4. Correlations between leaf growth indexes of Rhus chinensis and gall growth indexes in two inoculation methods
类别Category 小叶总面积
Total areas of leaflets复叶虫瘿数
Number of galled on the compound leave复叶虫瘿总体积
Total volume of galls单位虫瘿体积所需小叶面积
Leaf area per unit volume of gall人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration人工挂袋
Artificial aphid bags自然迁飞
Natural migration小叶数Number of leaflets 0.411* 0.244 0.296 0.384* 0.483** 0.503** -0.171 -0.358 小叶总面积Total areas of leaflets 1 1 0.278 0.083 0.191 -0.097 0.226 0.475** 复叶虫瘿数
Number of galled on the compound leave1 1 0.783** 0.719** -0.552** -0.457* 复叶虫瘿总体积Total volume of galls 1 1 -0.695** -0.733** 注:n=60。
Note: n=60.表 5 两种接种方式的主成分分数和特征值的系数矩阵
Table 5. Coefficient matrix of composition scores and eigenvalue of principal in two inoculation methods
类别Category 自然迁飞
Natural migration人工挂袋
Artificial aphid bags轴Axis 1 轴Axis 2 轴Axis 1 轴Axis 2 地径Ground diameter 0.33 -0.59 0.24 0.59 侧枝数
Number of lateral branches0.26 -0.07 0.31 0.12 梢头数Tips of branches 0.45 -0.25 0.44 0.34 有虫瘿梢头数
Galled tips of branches0.32 0.47 0.33 -0.43 复叶数
Number of compound leaves0.44 -0.29 0.44 0.26 有虫瘿复叶数Galled number of compound leaves 0.41 0.39 0.45 -0.30 虫瘿数Number of galls 0.40 0.35 0.39 -0.42 特征值Characteristic value 4.07 1.29 3.70 1.38 方差的Variance % 58.21 18.43 52.80 19.65 累积Cumulation % 58.21 76.65 52.80 72.45 -
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