养分影响入侵种喜旱莲子草对专食性天敌的防御

doi:10.17520/biods.2017301

摘要/Abstract

摘要：

竞争力增强进化假说(Evolution of Increased Competitive Ability, EICA)认为入侵植物在入侵地由于天敌逃逸导致对天敌防御投入降低而增加生长和繁殖等投入。然而, EICA假说所预测防御能力的进化主要集中在防御的组成抗性方面, 而从诱导抗性和耐受性的角度进行的验证还很有限, 并且忽视了生境资源可利用性对植物防御策略的影响。本研究采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)原产地(阿根廷)和入侵地(美国)各9个种群, 在高、低养分和有、无专食性天敌取食处理下的防御相关功能性状(比茎长、根冠比)及其对天敌取食的诱导反应, 以及它们与植物适合度相关指标的关系。结果表明: 入侵地种群具有较高的总生物量(+14.7%)、较低的比茎长(防御钻茎化蛹天敌的重要指标, -27.5%), 暗示其个体大小增加而防御投入降低, 符合EICA假说的预测。入侵地种群耐受性与原产地种群没有显著差异。在高养分下, 入侵地种群的比茎长对专食性天敌取食的诱导反应显著增加, 且与总生物量呈显著正相关。这些结果表明, 相比原产地种群, 入侵地种群具有更高的生物量, 对专食性天敌的组成抗性降低, 且在高养分下对专食性天敌的诱导抗性增加, 推测生境的养分富集可能会降低喜旱莲子草的生物防治效果。

关键词: 竞争力增强进化假说, 植物-昆虫关系, 可塑性, 资源可利用性, 诱导抗性

Abstract

Evolution of Increased Competitive Ability (EICA) has suggested that exotic plants may reduce resource allocation to anti-herbivore defenses and increase allocation to growth due to reduced specialist herbivores in introduced ranges. However, the major limitation for the hypothesis may be that, as important defensive strategies, induced resistance and tolerance were seldom tested and little information is known regarding the evolution of defensive strategies as influenced by resource availability. Nine introduced and nine native populations of Alternanthera philoxeroides were compared for fitness and two morphological traits that influence herbivory, specifically specific stem length (SSL, plants with low SSL have hollow stems favoured by a stem-boring specialist insect Agasicles hygrophila) and root/shoot ratio (RSR) under low and high nutrient availability and with or without A. hygrophila. Introduced populations showed greater total biomass (+14.7%) and lower SSL (-27.5%), demonstrating increased growth and decreased levels of constitutive defense. Tolerance to herbivory was similar between introduced and native populations. SSL plasticity to herbivory was greater in introduced populations under high nutrient levels, and was positively correlated with total biomass. These results suggest that although both functional trait values and plasticity contribute to the adaptation of alligator weeds to a wide range of nutrient availabilities and specialist insect herbivory, introduced populations have evolved higher production of biomass, lower levels of constitutive structural defense, and greater plasticity of structural defense in high nutrient conditions, indicating that the present control practice for A. philoxeroides where introducing A. hygrophila will face great challenges in the future.

Key words: EICA, plant-insect interactions, plasticity, resource availability, induced resistance

周方, 张致杰, 刘木, 潘晓云 (2017) 养分影响入侵种喜旱莲子草对专食性天敌的防御. 生物多样性, 25, 1276-1284. DOI: 10.17520/biods.2017301.

Fang Zhou, Zhijie Zhang, Mu Liu, Xiaoyun Pan (2017) Effects of nutrient levels on defense against specialist insects in an invasive alligator weed. Biodiversity Science, 25, 1276-1284. DOI: 10.17520/biods.2017301.

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https://www.biodiversity-science.net/CN/Y2017/V25/I12/1276

图/表 5

表1 喜旱莲子草原产地(阿根廷)和入侵地(美国)种群在两种养分处理(高养分; 低养分)和专食性天敌取食处理(有取食; 无取食)下适合度性状(生物量, 贮藏根生物量)和功能性状(根冠比, 比茎长)的方差分析或协方差分析结果

Table 1 ANOVA or ANCOVA of the effects of continental origin, population (pop), nutrient level and herbivory on total biomass, storage root biomass (SRB), root/shoot ratio (RSR), specific stem length (SSL) of Alternanthera philoxeroides.

参数 Source of variation	d.f	F
参数 Source of variation	d.f	总生物量 Total biomass	贮藏根生物量 SRB	根冠比 RSR	比茎长 SSL
采样地 Origin (O)	1, 16	36.76^***	2.07	3.67	51.71^***
种群 Population (Pop)	16, 264	9.01^***	5.69^***	10.05^***	14.03^***
养分 Nutrient (N)	1, 264	1.56	53.41^***	126.82^***	57.00^***
取食 Herbivory (H)	1, 264	16.84^***	7.23^**	10.46^**	3.86
养分 × 取食 (N × H)	1, 264	37.81^***	25.75^***	46.80^***	0.95
采样地 × 养分 (O × N)	1, 264	0.33	0.74	0.10	0.36
采样地 × 取食 (O × H)	1, 264	2.14	0.05	0.09	0.12
采样地 × 养分 × 取食 (O × N × H)	1, 264	0.40	0.01	0.68	5.15^*
养分 × 种群 (N × Pop)	16, 264	1.69	0.57	1.49	1.24
养分 × 取食 × 种群 (N × H × Pop)	16, 264	1.51	0.95	1.49	1.37
协变量 Covariates	1, 287		45.18^***	2.15	23.74^***

图1 莲草直胸跳甲的取食处理(有取食, 无取食)和养分处理(高养分, 低养分)对原产地(阿根廷)和入侵地(美国)种群总生物量(a)、贮藏根生物量(b)、根冠比(c)与比茎长(d)的影响。误差棒代表+/-标准误。

Fig. 1 The effects of herbivory by Agasicles hygrophila (with, h+, without, h-) and nutrient level (high, N+, low, N-) on plant total biomass (a), storage root biomass (SRB) (b), root/shoot ratio (RSR) (c), and specific stem length (SSL) (d) of Alternanthera philoxeroides from native (Argentina) and introduced (USA) range. Error bars denote +/- SE.

图2 养分处理(高养分, 低养分)对原产地(阿根廷)和入侵地(美国)种群喜旱莲子草总生物量(a)和贮藏根生物量(b)对莲草直胸跳甲取食耐受性的影响。误差棒代表+/-标准误。

Fig. 2 The effects of nutrient level (high, N+, low, N-) on tolerance of total biomass (a) and SRB (b) to herbivory by Agasicles hygrophila of Alternanthera philoxeroides from native (Argentina) and introduced (USA) range. Error bars denote +/- SE.

图3 养分处理(高养分, 低养分)对原产地(阿根廷)和入侵地(美国)种群喜旱莲子草根冠比(a)和比茎长(b)对莲草直胸跳甲取食的诱导反应(即可塑性指数)的影响。误差棒代表+/-标准误。

Fig. 3 The effects of nutrient level (high, N+, low, N-) on the plasticity (RTR) of RSR (a) and SSL (b) in response to herbivory by Agasicles hygrophila of Alternanthera philoxeroides from native (Argentina) and introduced (USA) range. Error bars denote +/-SE.

图4 在两种养分处理下, 原产地(阿根廷)和入侵地(美国)喜旱莲子草种群的适合度(总生物量、贮藏根生物量SRB)及总生物量、贮藏根生物量对取食作用的耐受性(TBiomass、TSRB)与功能性状(根冠比RSR、比茎长SSL)及功能性状对莲草直胸跳甲取食的可塑性指数(RTR for RSR, RTR for SSL)的相关性。实线代表正相关, 虚线代表负相关。粗线代表显著性水平P < 0.01, 细线代表显著性水平P < 0.05。

Fig. 4 The correlations between plant fitness (total biomass, SRB), tolerance of total biomass (TBiomass) and SRB (TSRB) to herbivory and functional traits (root/shoot ratio, RSR; specific stem length, SSL), relative trait range of functional traits (RTR for RSR, RTR for SSL) in response to herbivory by Agasicles hygrophila for Alternanthera philoxeroides from native (Argentina) and introduced (USA) range grown under high and low nutrient levels. The solid lines signify positive correlation, and dashed ones mean negative, the thick lines mean significant at P < 0.01, and thin lines at P < 0.05.

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