Aboveground herbivory increases soil nematode abundance of an invasive plant

入侵植物往往可以影响土壤线虫的群落结构。然而,入侵植物在入侵地并未完全逃逸地上植食性昆虫的取食危害,而我们对昆虫取食是否会调节入侵植物对土壤线虫群落的影响还知之甚少。在本研究中,我们探讨了地上植食性昆虫取食危害如何调节入侵植物空心莲子草(Alternanthera philoxeroides)对土壤线虫群落的影响,并研究了昆虫调节作用的时间效应。我们开展温室实验,对盆栽的空心莲子草施加三种不同的昆虫取食处理[无昆虫取食、专食性昆虫莲草直胸跳甲(Agasicles hygrophila)取食、广食性昆虫虾钳菜披龟甲(Cassida piperata)取食],并在移除所有植食性昆虫后的第1、10和20天测定土壤线虫的群落结构。此外,我们还测定了空心莲子草根系及根系分泌物的碳含量。研究发现,在植食性昆虫被移除的第1天,昆虫取食处理显著提高了空心莲子草根围土壤线虫的丰度,特别是植食性线虫。然而,随着时间推移,这种影响逐渐减退,并在后期(第10和20天)完全消失。此外,专食性昆虫莲草直胸跳甲和广食性昆虫虾钳菜披龟甲的取食危害对土壤线虫丰度的影响并无显著性差异。综上所述,本研究表明地上植食性昆虫的取食危害可以改变入侵植物对土壤线虫群落的影响,但昆虫的调节作用是短暂的。此外,我们的研究强调了在评估入侵植物的影响时,需要同时考虑地上和地下生物群落。     

养分影响入侵种喜旱莲子草对专食性天敌的防御

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。     

论昆虫与植物的相互作用和进化的关系

昆虫与植物是陆地生物群落中最为重要的组成部分,二者间的相互作用是多方面的,其中最为重要的是昆虫选择植物作为食物和生长场所、昆虫为植物传授花粉两方面。该文集中讨论这两方面的相互作用有哪些因素与进化有密切的关系。植食性昆虫根据其寄主植物范围,通常分为专食性（寄主范围窄）和广食性（寄主范围广）。从生态关系来看,广食性的取食行为比专食性的更为有利,但实际情况却与此相反,统观植食性昆虫的取食行为,有向专食性演化更为普遍的倾向。专食性发展有利于提高昆虫对寄主植物的选择效率,还可缓和天敌作用所造成的压力。根据昆虫与植物相互作用的特点,目前已提出很多昆虫与植物的进化理论,包括成对的协同进化、弥散的协同进化、群落的协同进化以及顺序进化。在昆虫对寄主植物的选择中,以植物对昆虫的影响较昆虫对植物的影响更为重要,称为顺序进化是适宜的;昆虫为被子植物传授花粉造成互惠共生,其中的进化关系应称为协同进化。     

昆虫与植物的协同进化:寄主植物-铃夜蛾-寄生蜂相互作用

近数10年内,Ehrlich和Raven于1964年提出的协同进化理论及Jermy于1976年提出的顺序进化理论极大地促进了对昆虫与植物相互作用的研究。文章首先简要介绍有关理论,对植食性昆虫与植物关系研究的若干核心问题进行评述。主要问题包括(1)植食性昆虫如何选择寄主植物?(2)植物次生物质是否作为植物防御昆虫取食的重要屏障?(3)昆虫能否适应植物的化学防御?(4)植食性昆虫寄主范围是否是从广到专演化的?随之,作者结合对铃夜蛾Helicoverpa系统研究取得的结果,对上述问题做了进一步的论证和阐述。最后,在继承协同进化、顺序进化等理论精髓的基础上,根据当今三营养级相互作用领域的研究新进展,提出一个新的假说,即多营养级协同进化假说。该假说肯定植物次生物质在植物防御和昆虫识别寄主植物上的重要作用,同时把其他营养级并列放入交互作用的系统,特别强调第三营养级在昆虫与植物关系演化过程中的参与和寄主转移与昆虫食性专化和广化的联系。     

植物与植食性昆虫相互作用的分子机制

在长期的协同进化中,植物建立起应对昆虫取食为害的精密而又复杂的防御机制,植物转录组调控中防御应答基因的表达及防御物质的合成因不同的昆虫取食方式而异。一般来说,咀嚼式口器昆虫取食时造成大面积组织伤害,可诱导植物产生伤害反应;而刺吸式口器昆虫因其特殊的口针取食,诱导植物激活病原体相关的防御途径。不同的防御途径激活不同的识别机制和信号途径。本文从信号识别和转导上综述了不同食性的昆虫取食植物时所引发的防御反应,分析了昆虫-植物相互作用关系的分子机制。     

专食性-广食性天敌比例影响入侵植物喜旱莲子草生长防御策略

外来植物从原产地到入侵地通常会经历植食性天敌选择压力的变化,其生长防御性状的快速适应性进化是成功入侵的重要机制之一。植食性天敌按食性专一性分为专食性天敌与广食性天敌,并对植物生长防御性状产生不同的选择压力。然而,在自然群落中两类植食性天敌的相对比例可能会随时间和空间的改变而改变,这些变化对入侵植物生长防御进化的影响尚不清楚。本研究以喜旱莲子草(Alternanthera philoxeroides)为研究对象,在同质园条件下比较了原产地(阿根廷)和入侵地(中国)种群在不同专食性-广食性天敌比例处理下生长防御性状的差异。结果显示:专食性-广食性天敌比例对喜旱莲子草生长防御性状的影响无显著差异,但其与来源地之间的交互作用对生长性状中的贮藏根生物量与根冠比具有显著影响。其中入侵地种群的贮藏根生物量与根冠比随着专食性天敌比例的增加而增加,而原产地种群正好相反。通过对比原产地与入侵地种群生长防御相关性状发现,入侵地种群的总生物量(–21.4%, P=0.027)、地上生物量(–22.6%, P=0.026)、生长速率(–17.5%, P <0.001)和黄酮含量(–38.4%, P=0.0...     

Effects of latitude and soil microbes on the resistance of invasive Solidago canadensis to its co-evolved insect herbivore Corythucha marmorata

纬度和土壤微生物对入侵植物加拿大一枝黄花对共同进化的专性食草昆虫抗性的影响在全球化的背景下,入侵植物再次暴露在来自其原产地的专性食草动物面前的可能性越来越大。然而,入侵植物对曾共同进化的专性食草动物的抗性是否存在纬度趋势以及土壤微生物如何影响入侵植物对这类食草动物的抗性还鲜有研究。我们假设入侵植物加拿大一枝黄花(Solidago canadensis)对其曾共 同进化的来自原产地的食草昆虫菊方翅网蝽(Corythucha marmorata)的抗性随纬度的增加而增加,而加拿 大一枝黄花的局部根际微生物可以促进加其对菊方翅网蝽的抗性。为了验证上述假设,我们首先通过野外调查和温室实验,研究了菊方翅网蝽在中国加拿大一枝黄花种群中的分布丰度,以及对加拿大一枝黄花造成的危害是否存在纬度分布规律。其次,我们通过温室实验,检测了加拿大一枝黄花的本地根际土壤微生物是否能促进加拿大一枝黄花对菊方翅网蝽的抗性。研究结果表明,菊方翅网蝽在加拿大一枝黄花中的分布密度和对加拿大一枝黄花的危害程度与纬度以及加拿大一枝黄花的生长呈现正相关,说明加拿大一枝黄花对菊方翅网蝽的抗性具有纬度趋势。然而,在温室实验中,来自不同纬度的加拿大一枝黄花受到菊方翅网蝽的伤害程度没有显著差异。此外,当根际土壤和加拿大一枝黄花的根状茎同时来自于野外被轻度危害的加拿大一枝黄花或同时来自于野外被重度危害的加拿大一枝黄花时,加拿大一枝黄花的抗性最强。这一结果表明,加拿大一枝黄花的局部根际土壤微生物促进了加拿大一枝黄花对菊方翅网蝽的抗性。     

寄主植物叶片物理性状对潜叶昆虫的影响

潜叶昆虫广泛分布于鳞翅目、双翅目、鞘翅目和膜翅目中,其幼虫潜入叶片内部生活和取食,是一类用于研究植物-昆虫-天敌种间关系和协同进化的重要模式生物。有些潜叶昆虫是重要农林害虫。相比外食性昆虫,在叶内取食的潜叶昆虫幼虫更易受到叶片物理性状的直接影响。叶片的着生位置、朝向、大小、颜色和表皮毛等直接决定潜叶虫成虫的取食和产卵选择,从而影响幼虫的空间分布和寄主适应。叶片的某些物理性状也会直接影响幼虫的取食行为、生长发育和被寄生率。研究叶片物理性状的防御作用以及潜叶昆虫对这些防御的适应,有助于了解潜叶昆虫-寄主植物的协同进化。另一方面,外界环境和遗传育种都有可能改变植物叶片的物理特性,而对潜叶害虫产生抗性,从而实现潜叶害虫的可持续生态控制。     

蚜虫诱导的植物免疫反应

在植物与昆虫长期的互作过程中,植物建立起一系列精密而又复杂的防御机制以应对昆虫取食为害,并且能够识别不同取食类型昆虫的效应因子作出不同的防御应答。最近研究揭示了许多植物与蚜虫之间相互抗争的分子机制,这不仅包括植物激素介导的诱导防御途径、植物先天免疫系统和基于gene-for-gene的R抗性识别和作用机制,而且还包括蚜虫在取食过程中分泌的唾液成分,它有助于蚜虫取食韧皮部组织,抑制植物病原相关分子模式激活的免疫反应(Pathogen-associated molecular patterns triggered immunity,PTI)防御,以及被植物核苷酸结合位点区-亮氨酸重复序列区(NBS-LRR)膜受体识别激活效应因子免疫反应(ETI)防御等方面。本文综述了蚜虫诱导的植物防御途径、蚜虫诱导的植物免疫反应、蚜虫效应因子的鉴定与功能分析三方面的最近研究进展,提出了未来发展的研究方向。这些基于病原微生物提出的"zig-zag"模型为进一步理解植物先天免疫、诱导防御系统和蚜虫唾液腺组分的互作提供新理论支撑,为揭示了植物与蚜虫抗性互作的分子机制及有效安全地防治害虫提供了新思路。     

蚜虫诱导的植物免疫反应

在植物与昆虫长期的互作过程中,植物建立起一系列精密而又复杂的防御机制以应对昆虫取食为害,并且能够识别不同取食类型昆虫的效应因子作出不同的防御应答。最近研究揭示了许多植物与蚜虫之间相互抗争的分子机制,这不仅包括植物激素介导的诱导防御途径、植物先天免疫系统和基于gene-for-gene的R抗性识别和作用机制,而且还包括蚜虫在取食过程中分泌的唾液成分,它有助于蚜虫取食韧皮部组织,抑制植物病原相关分子模式激活的免疫反应(Pathogen-associated molecular patterns triggered immunity,PTI)防御,以及被植物核苷酸结合位点区-亮氨酸重复序列区(NBS-LRR)膜受体识别激活效应因子免疫反应(ETI)防御等方面。本文综述了蚜虫诱导的植物防御途径、蚜虫诱导的植物免疫反应、蚜虫效应因子的鉴定与功能分析三方面的最近研究进展,提出了未来发展的研究方向。这些基于病原微生物提出的"zig-zag"模型为进一步理解植物先天免疫、诱导防御系统和蚜虫唾液腺组分的互作提供新理论支撑,为揭示了植物与蚜虫抗性互作的分子机制及有效安全地防治害虫提供了新思路。     

How leaf domatia and induced plant resistance affect herbivores, natural enemies and plant performance

Predators and plant resistance may act together to control herbivorous arthropod populations or antagonistically, which would reduce the control of pest populations. In a field experiment we enhanced predation by adding simulated leaf domatia to plants. Leaf domatia are small structures that often harbor predaceous arthropods that are potentially beneficial to the plant. We also manipulated host plant quality by inducing resistance with controlled, early season exposure of seedlings to spider mite herbivory.
Our manipulations had profound consequences for the natural community of arthropods that inhabited the plants. Leaf domatia had a direct positive effect on abundances of two species of bugs and one species of thrips, all of which are largely predators of herbivores. On leaves with domatia, each of the predators was found inside the domatia two to three times more often than outside the domatia. Eggs of predaceous bugs inside leaf domatia were protected from parasitism compared to eggs outside the domatia. The positive effects of leaf domatia on predator abundances were associated with reduced populations of herbivorous spider mites, aphids, and whiteflies. Plants with experimental leaf domatia showed significantly enhanced reproductive performance.
Induced resistance also affected the community of arthropods. Of the abundant predators, all of which also fed on the plant, only minute pirate bugs were negatively affected by induced resistance. Populations of herbivorous spider mites and whiteflies were directly and negatively affected by induction. In contrast, aphid populations were higher on plants with induced resistance compared to uninduced plants. Effects of induced resistance and domatia were additive for each of the predators and for aphids. However, spider mite and whitefly populations were not suppressed further by employing both induced resistance and domatia compared to each strategy alone. Our manipulations suggest that plant defense strategies can have positive effects on some species and negative effects on others. Negative effects of “resistance traits” on predators and positive effects on some herbivores may reduce the benefits of constitutive expression of resistance traits and may favor inducible defense strategies. Multiple plant strategies such as inducible resistance and morphological traits that aid in the recruitment of predators of herbivores may act together to maximize plant defenses, although they may also be redundant and not act additively.     

Effects of generalist herbivory on resistance and resource allocation by the invasive plant,Phytolacca americana

Successful invasions by exotic plants are often attributed to a loss of co‐evolved specialists and a re‐allocation of resources from defense to growth and reproduction. However, invasive plants are rarely completely released from insect herbivory because they are frequently attacked by generalists in their introduced ranges. The novel generalist community may also affect the invasive plant's defensive strategies and resource allocation. Here, we tested this hypothesis using American pokeweed (Phytolacca americana L.), a species that has become invasive in China, which is native to North America. We examined resistance, tolerance, growth and reproduction of plant populations from both China and the USA when plants were exposed to natural generalist herbivores in China. We found that leaf damage was greater for invasive populations than for native populations, indicating that plants from invasive ranges had lower resistance to herbivory than those from native ranges. A regression of the percentage of leaf damage against mass showed that there was no significant difference in tolerance between invasive and native populations, even though the shoot, root, fruit and total mass were larger for invasive populations than for native populations. These results suggest that generalist herbivores are important drivers mediating the defensive strategies and resource allocation of the invasive American pokeweed.     

For or against: the importance of variation in growth rate for testing the EICA hypothesis

The evolution of increased competitive ability (EICA) hypothesis proposes that invasive species evolve decreased defense and increased competitive ability following natural enemy release. Previous studies have found evidence both for and against EICA. The resource-enemy release hypothesis (R-ERH) suggests that fast-growing species may experience stronger enemy release than slow-growing species. On the basis of R-ERH, the prediction of EICA will be held true for slow-growing genotypes, i.e., the slow-growing genotypes from the introduced range will be less resistant to herbivory and grow faster than those from the home range; while the EICA will not be held for fast-growing genotypes, i.e., there will be no significant differences in growth and defense traits between the introduced and native fast-growing genotypes. We tested these predictions preliminarily using five populations of the invasive plant Alternanthera philoxeroides. This species has two varieties in its home range, which showed a distinct growth-defense strategy: the northern A. p. var. acutifolia (Apa) had higher growth rate but lower resistance, while the southern A. p. var. obtusifolia (Apo) had lower growth rate but higher resistance level. Our results suggest that the EICA hypothesis is consistent with the slow-growing Apo, but not with the fast-growing Apa. We suggest that evolutionary changes in growth or resistance following enemy release are influenced by variation in growth rate within an invasive alien plant. These findings have important implications for the EICA hypothesis, and may partially explain why previous studies have found evidence both for and against EICA.     

Enemy release and plant invasion: patterns of defensive traits and leaf damage in Hawaii

Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.     

Extracts of the invasive shrub Lonicera maackii increase mortality and alter behavior of amphibian larvae

Water-soluble phytochemicals produced by invasive plants may represent novel elements of invaded ecosystems that can precipitate a variety of direct and indirect effects on native organisms. Phenolic compounds in particular are a common plant defense, and these compounds may have disproportionate impacts on amphibians compared to other taxa. We coupled an exploration of invasive plant extract effects on larvae of four amphibian species (the salamander Ambystoma maculatum, the toad Anaxyrus americanus, and the frogs Hyla sp. and Lithobates blairi) with behavioral observations designed to determine whether behavior can ameliorate the negative effects of exposure to invasive plant extracts. Larvae were reared in extracts of the widespread invasive Amur honeysuckle (Lonicera maackii), mixed native leaf litter, and a water control. Anaxyrus americanus tadpoles reared in L. maackii extracts were more likely to die than tadpoles reared in native extracts, but differences in mortality following rearing in native and exotic extracts were not significant for the other three species. Anaxyrus americanus and L. blairi tadpoles made more trips to the surface in L. maackii extracts than in native extracts, consistent with the hypothesis that exotic extracts may interfere with respiratory physiology and suggesting that L. blairi can behaviorally ameliorate the negative effects of L. maackii extracts. Our study highlights both a direct and indirect pathway by which invasive plant extracts may alter the ecological dynamics of native fauna.     

Invasive belowground mutualists of woody plants

Most plants require mutualistic associations to survive, which can be an important limitation on their ability to become invasive. There are four strategies that permit plants to become invasive without being limited by a lack of mutualists. One is to not be dependent on mutualists. The other three strategies are to form novel mutualisms, form associations with cosmopolitan species, or co-invade with mutualists from their native range. Historically there has been a bias to study mutualisms from a plant perspective, with little consideration of soil biota as invasive species in their own right. Here we address this by reviewing the literature on belowground invasive mutualists of woody plants. We focus on woody invaders as ecosystem-transforming plants that frequently have a high dependence on belowground mutualists. We found that co-invasions are common, with many ectomycorrhizal plant species and N-fixing species co-invading with their mutualists. Other groups, such as arbuscular mycorrhizal plants, tend to associate with cosmopolitan fungal species or to form novel associations in their exotic range. Only limited evidence exists of direct negative effects of co-invading mutualists on native mutualist communities, and effects on native plants appear to be largely driven by altered environmental conditions rather than direct interactions. Mutualists that introduce novel ecosystem functions have effects greater than would be predicted based solely on their biomass. Focusing on the belowground aspects of plant invasions provides novel insights into the impacts, processes and management of invasions of both soil organisms and woody plant species.     

No evolution of reduced resistance and compensation for psyllid herbivory by the invasive <Emphasis Type="Italic">Genista monspessulana</Emphasis>

The evolution of redirecting resources from plant defense to growth or reproduction may explain why some exotic species are successful invaders in new environments. For example, the evolution of increased competitive ability hypothesis posits that escape from herbivores by invasive plants results in the selection of more vigorous genotypes that reduce their allocation of resources to defense. In addition, understanding the defense strategy of an invasive plant may help forecast the likely impact of herbivory. We tested the prediction of reduced defense (i.e., resistance) in Genista monspessulana, measured indirectly as the performance of a specialist psyllid herbivore, by comparing five native and introduced plant populations. We also examined the ability of G. monspessulana to compensate for herbivory in the presence and the absence of psyllids for a single plant population from the native and introduced regions. Plant origin (native or introduced) did not influence the psyllid’s abundance and population growth rate, suggesting no change in resistance to herbivory for introduced plants. Similarly, we found no overall difference in plant performance between individuals in the presence and the absence of psyllid herbivory, suggesting that G. monspessulana was able to fully compensate for herbivory. Damaged plants compensated by changing the pattern of branching, which also resulted in greater dry leaf biomass. We conclude that evolution of reduced defenses does not explain the success of G. monspessulana as an invader and that compensation for herbivory may limit the efficacy of the psyllid as a biological control agent.     

Meta‐analysis reveals evolution in invasive plant species but little support for Evolution of Increased Competitive Ability (EICA)

Ecological explanations for the success and persistence of invasive species vastly outnumber evolutionary hypotheses, yet evolution is a fundamental process in the success of any species. The Evolution of Increased Competitive Ability (EICA) hypothesis (Blossey and Nötzold 1995) proposes that evolutionary change in response to release from coevolved herbivores is responsible for the success of many invasive plant species. Studies that evaluate this hypothesis have used different approaches to test whether invasive populations allocate fewer resources to defense and more to growth and competitive ability than do source populations, with mixed results. We conducted a meta‐analysis of experimental tests of evolutionary change in the context of EICA. In contrast to previous reviews, there was no support across invasive species for EICA's predictions regarding defense or competitive ability, although invasive populations were more productive than conspecific native populations under noncompetitive conditions. We found broad support for genetically based changes in defense and competitive plant traits after introduction into new ranges, but not in the manner suggested by EICA. This review suggests that evolution occurs as a result of plant introduction and population expansion in invasive plant species, and may contribute to the invasiveness and persistence of some introduced species.     

Defensive tradeoffs are not prerequisites to plant diversity in a two species model

Trait‐based theories of biodiversity consider interspecific tradeoffs among species‐specific traits as prerequisites to maintaining community evenness, a component of species diversity. Such tradeoffs are commonly observed in plant communities, particularly in relation to traits associated with resistance to herbivory. Indeed, global experiments show that interspecific tradeoffs are common between plant defense and growth or competitive ability; however, the positive effects of herbivory on plant diversity predicted by theories with trait‐based tradeoffs are far less commonly observed. Moreover, both the overall and relative importance of these tradeoffs in promoting plant diversity are not well known. To disentangle the relationships among growth, competition, and defense in relation to plant community evenness, we built a model that describes the effects of a shared herbivore on two plant species with the potential to differ in each of these traits. While tradeoffs between plant defense and growth or competitive ability can increase plant diversity via evenness, this is not always the case nor is it a requirement for increased diversity. Herbivores may increase plant diversity even in the absence of defensive tradeoffs, preferentially consuming apparently maladapted species, by limiting the negative effects of interspecific interactions. Therefore, the importance of defensive tradeoffs in increasing diversity may not be as important, or as straightforward, as previously hypothesized.     

COSTS OF INDUCED RESPONSES AND TOLERANCE TO HERBIVORY IN MALE AND FEMALE FITNESS COMPONENTS OF WILD RADISH

Theory predicts that plant defensive traits are costly due to trade-offs between allocation to defense and growth and reproduction. Most previous studies of costs of plant defense focused on female fitness costs of constitutively expressed defenses. Consideration of alternative plant strategies, such as induced defenses and tolerance to herbivory, and multiple types of costs, including allocation to male reproductive function, may increase our ability to detect costs of plant defense against herbivores. In this study we measured male and female reproductive costs associated with induced responses and tolerance to herbivory in annual wild radish plants (Raphanus raphanistrum). We induced resistance in the plants by subjecting them to herbivory by Pieris rapae caterpillars. We also induced resistance in plants without leaf tissue removal using a natural chemical elicitor, jasmonic acid; in addition, we removed leaf tissue without inducing plant responses using manual clipping. Induced responses included increased concentrations of indole glucosinolates, which are putative defense compounds. Induced responses, in the absence of leaf tissue removal, reduced plant fitness when five fitness components were considered together; costs of induction were individually detected for time to first flower and number of pollen grains produced per flower. In this system, induced responses appear to impose a cost, although this cost may not have been detected had we only quantified the traditionally measured fitness components, growth and seed production. In the absence of induced responses, 50% leaf tissue removal, reduced plant fitness in three out of the five fitness components measured. Induced responses to herbivory and leaf tissue removal had additive effects on plant fitness. Although plant sibships varied greatly (49–136%) in their level of tolerance to herbivory, costs of tolerance were not detected, as we did not find a negative association between the ability to compensate for damage and plant fitness in the absence of damage. We suggest that consideration of alternative plant defense strategies and multiple costs will result in a broader understanding of the evolutionary ecology of plant defense.