Tremendous strides have been made regarding our understanding of how host plant chemistry influences the interactions between herbivores and their natural enemies. While most work has focused on plant chemistry effects on host location and acceptance by natural enemies, an increasing number of studies examine negative effects. The tritrophic role of plant chemistry is central to several aspects of trophic phenomena including top-down versus bottom-up control of herbivores, enemy-free space and host choice, and theories of plant defense. Furthermore, tritrophic effects of plant chemistry are important in assessing the degree of compatibility between biological control and plant resistance approaches to pest control. Additional research is needed to understand the physiological effects of plant chemistry on parasitoids. Explicit tests are required to determine whether natural enemies can act as selective forces on plant defense. Finally, further studies of natural systems are crucial to understanding the evolution of multitrophic relationships.

Plant chemistry and natural enemy fitness: effects on herbivore and natural enemy interactions.

In recent years, induced plant defenses have received widespread attention from biologists in a variety of disciplines. The mechanisms underlying these defenses and the interactions that mediate them appeal not only to plant physiologists, ecologists, and evolutionary biologists but also to those scientists that search for novel strategies in plant protection. Several recent books (Karban and Baldwin, 1997; Agrawal et al., 1999) and reviews (Baldwin, 1994; Karban et al., 1997; Agrawal and Rutter, 1998; Agrawal and Karban, 1999; Baldwin and Preston, 1999; Dicke et al., 2003) have been devoted entirely to the subject of induced plant defenses. Various forces, ranging from abiotic stresses to biotic factors such as pathogens, arthropods, or higher organisms, may trigger different plant defense responses. Yet, the biochemical pathways that are involved appear to show considerable similarities. This is also true for the so-called indirect defenses. The term indirect defense refers to those adaptations that result in the recruitment and sustenance of organisms that protect the plants against herbivorous attackers. The early published examples of indirect defenses involved intimate plant–ant interactions, in which myrmecophilous plants were shown to have evolved a range of adaptations providing ants with shelter (domatia) and various food sources (Belt, 1874; Janzen, 1966). In return, these plants may obtain a range of benefits because ants can provide nutrition (Thomson, 1981) or more commonly, protection against herbivores, pathogens, and competing plants (e.g. Koptur, 1992; Oliveira, 1997). The well-documented fitness benefits of ant attendance in myrmecophilous plants (Rico-Gray and Thien, 1989; Oliveira, 1997), combined with the fact that domatia and food supplements are difficult to reconcile with other functions, are convincing arguments for the interpretation that these adaptations represent examples

/pdf/recruitment-of-predators-and-parasitoids-by-herbivore-2jbfj3xxe7.pdf

Recruitment of predators and parasitoids by herbivore-injured plants

Crop domestication is the process of artificially selecting plants to increase their suitability to human requirements: taste, yield, storage, and cultivation practices. There is increasing evidence that crop domestication can profoundly alter interactions among plants, herbivores, and their natural enemies. Overall, little is known about how these interactions are affected by domestication in the geographical ranges where these crops originate, where they are sympatric with the ancestral plant and share the associated arthropod community. In general, domestication consistently has reduced chemical resistance against herbivorous insects, improving herbivore and natural enemy performance on crop plants. More studies are needed to understand how changes in morphology and resistance-related traits arising from domestication may interact with environmental variation to affect species interactions across multiple scales in agroecosystems and natural ecosystems.

Crop Domestication and Its Impact on Naturally Selected Trophic Interactions

Host plant identity and host plant chemistry have often been shown to influence host finding and acceptance by natural enemies but comparatively less attention has been paid to the tritrophic effects of host plant and host plant chemistry on other natural enemy fitness correlates, such as survivorship, clutch size, body size, and sex ratio. Such studies are central to understanding both the selective impact of plants on natural enemies as well as the potential for reciprocal selective impact of natural enemies on plant traits. We examined the effects of host plant and host plant chemistry in a tritrophic system consisting of three apiaceous plants (Pastinaca sativa, Heracleum sphondylium and H. mantegazzianum), the parsnip webworm (Depressaria pastinacella) and the polyembryonic parasitic wasp Copidosoma sosares. All of these plants produce furanocoumarins, known resistance factors for parsnip webworms. Furanocoumarin concentrations were correlated neither with the presence nor the number of webworms on a given plant. Concentrations of two furanocoumarins were negatively associated with C. sosares fitness correlates: isopimpinellin with the likelihood that a given webworm would be parasitized and xanthotoxin with both within-brood survivorship (of all-male and mixed-sex broods) and clutch size. Brood sex ratio and body sizes of individual wasps were not correlated with furanocoumarin chemistry. Because additive genetic variation exists in P. sativa for furanocoumarin chemical traits, these are subject to selection by webworms through herbivory. Third trophic level selective impacts on furanocoumarin traits may include selection for reduced production of those chemicals that affect parasitoid survivorship yet do not influence host plant choice by the herbivore. That such might be the case is suggested by patterns of furanocoumarin production in populations of P. sativa with different histories of infestation; in the Netherlands, where parasitism rates of webworms by C. sosares are high, plants produce lower levels of all linear furanocoumarins and proportionately less isopimpinellin than do midwestern U.S. populations of P. sativa, where natural enemies of the webworm are effectively absent.

/pdf/host-plant-host-plant-chemistry-and-the-polyembryonic-15er9ojpu2.pdf

Host plant, host plant chemistry and the polyembryonic parasitoid Copidosoma sosares : indirect effects in a tritrophic interaction

Foraging behavior by six fruit fly parasitoids (Hymenoptera: Braconidae) released as single- or multiple-species cohorts in field cages: Influence of fruit location and host density

Naive female Diachasmimorpha longicaudata (Ashmead), a solitary endoparasitoid of tephritid fruit flies, exhibited positive responses toward volatiles of host fruits in olfactometer and wind tunnel bioassays. Although no significant preference for one of the test fruits, mango, Mangifera indica L., or grapefruit, Citrus paradisi Macfaden, was observed, the number of flights in the wind tunnel was higher in the presence of mangos. In the olfactometer trials, parasitoids preferred fly infested over noninfested grapefruits, and infested over noninfested mangos. Reproductive performance bioassays were conducted using Anastrepha ludens (Loew) larvae (Diptera: Tephritidae) reared in grapefruit, orange, mango, or artificial diet, and Anastrepha obliqua (Macquart) larvae reared in mango as parasitoid hosts. Parasitoid performance was compared for 2 Anastrepha species and 3 fruit species. Significant effects of fruit fly species and of the diet of fruit fly larvae on longevity, size, and progeny production of D. longicaudata were observed. Anastrepha ludens reared in grapefruit was the best host in terms of offspring longevity, size, and number of female progeny, but parasitoids that developed in A. ludens reared in mango had higher overall fecundity. Anastrepha ludens reared in mango was a better host than A. obliqua in the same fruit. No correlation between parasitoid size and demographic parameters was found. The results of this laboratory study showed that host preference and offspring performance are partially related.

https://bioone.org/journals/environmental-entomology/volume-29/issue-1/0046-225X-29.1.87/Host-Species-and-Host-Plant-Effects-on-Preference-and-Performance/10.1603/0046-225X-29.1.87.pdf

Host Species and Host Plant Effects on Preference and Performance of Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

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Host Species and Host Plant Effects on Preference and Performance of Diachasmimorpha longicaudata (Hymenoptera: Braconidae)