Simple Growth Patterns Can Create Complex Trajectories for the Ontogeny of Constitutive Chemical Defences in Seaweeds

All of the theory and most of the data on the ecology and evolution of chemical defences derive from terrestrial plants, which have considerable capacity for internal movement of resources. In contrast, most macroalgae – seaweeds – have no or very limited capacity for resource translocation, meaning that trade-offs between growth and defence, for example, should be localised rather than systemic. This may change the predictions of chemical defence theories for seaweeds. We developed a model that mimicked the simple growth pattern of the red seaweed Asparagopsis armata which is composed of repeating clusters of somatic cells and cells which contain deterrent secondary chemicals (gland cells). To do this we created a distinct growth curve for the somatic cells and another for the gland cells using empirical data. The somatic growth function was linked to the growth function for defence via differential equations modelling, which effectively generated a trade-off between growth and defence as these neighbouring cells develop. By treating growth and defence as separate functions we were also able to model a trade-off in growth of 2–3% under most circumstances. However, we found contrasting evidence for this trade-off in the empirical relationships between growth and defence, depending on the light level under which the alga was cultured. After developing a model that incorporated both branching and cell division rates, we formally demonstrated that positive correlations between growth and defence are predicted in many circumstances and also that allocation costs, if they exist, will be constrained by the intrinsic growth patterns of the seaweed. Growth patterns could therefore explain contrasting evidence for cost of constitutive chemical defence in many studies, highlighting the need to consider the fundamental biology and ontogeny of organisms when assessing the allocation theories for defence.     

Specificity in Mesograzer-Induced Defences in Seagrasses

Grazing-induced plant defences that reduce palatability to herbivores are widespread in terrestrial plants and seaweeds, but they have not yet been reported in seagrasses. We investigated the ability of two seagrass species to induce defences in response to direct grazing by three associated mesograzers. Specifically, we conducted feeding-assayed induction experiments to examine how mesograzer-specific grazing impact affects seagrass induction of defences within the context of the optimal defence theory. We found that the amphipod Gammarus insensibilis and the isopod Idotea chelipes exerted a low-intensity grazing on older blades of the seagrass Cymodocea nodosa, which reflects a weak grazing impact that may explain the lack of inducible defences. The isopod Synischia hectica exerted the strongest grazing impact on C. nodosa via high-intensity feeding on young blades with a higher fitness value. This isopod grazing induced defences in C. nodosa as indicated by a consistently lower consumption of blades previously grazed for 5, 12 and 16 days. The lower consumption was maintained when offered tissues with no plant structure (agar-reconstituted food), but showing a reduced size of the previous grazing effect. This indicates that structural traits act in combination with chemical traits to reduce seagrass palatability to the isopod. Increase in total phenolics but not in C:N ratio and total nitrogen of grazed C. nodosa suggests chemical defences rather than a modified nutritional quality as primarily induced chemical traits. We detected no induction of defences in Zostera noltei, which showed the ability to replace moderate losses of young biomass to mesograzers via compensatory growth. Our study provides the first experimental evidence of induction of defences against meso-herbivory that reduce further consumption in seagrasses. It also emphasizes the relevance of grazer identity in determining the level of grazing impact triggering resistance and compensatory responses of different seagrass species.     

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.     

Native generalist herbivores promote invasion of a chemically defended seaweed via refuge‐mediated apparent competition

Refuge‐mediated apparent competition was recently suggested as a mechanism that enables plant invasions. The refuge characteristics of introduced plants are predicted to enhance impacts of generalist herbivores on native competitors and thereby result in an increased abundance of the invader. However, this prediction has so far not been experimentally verified. This study tested if the invasion of a chemically defended seaweed is promoted by native generalist herbivores via refuge‐mediated apparent competition. The invader was shown to offer herbivores a significantly better refuge against fish predation compared with native seaweeds. Furthermore, in an experimental community, the presence of herbivores decreased the performance of neighbouring native seaweeds, but increased growth and relative abundance of the invader. These results provides the first experimental evidence that native generalist herbivores can shift a community towards a dominance of a well‐defended invader, inferior to native species in direct competitive interactions, by means of refuge‐mediated apparent competition.     

Testing for the induction of anti-herbivory defences in four Portuguese macroalgae by direct and water-borne cues of grazing amphipods

Herbivory is a key factor in regulating plant biomass, thereby driving ecosystem performance. Algae have developed multiple adaptations to cope with grazers, including morphological and chemical defences. In a series of experiments we investigated whether several species of macroalgae possess anti-herbivore defences and whether these could be regulated to demand, i.e. grazing events. The potential of direct grazing on defence induction was assessed for two brown (Dictyopteris membranacea, Fucus vesiculosus) and two red seaweeds (Gelidium sesquipedale, Sphaerococcus coronopifolius) from São Rafael and Ria Formosa, Portugal. Bioassays conducted with live algal pieces and agar-based food containing lipophilic algal extracts were used to detect changes in palatability after exposure to amphipod attacks (=treatment phase). Fucus vesiculosus was the only species significantly reducing palatability in response to direct amphipod-attacks. This pattern was observed in live F. vesiculosus pieces and agar-based food containing a lipophilic extract, suggesting that lipophilic compounds produced during the treatment phase were responsible for the repulsion of grazers. Water-borne cues of grazed F. vesiculosus as well as non-grazing amphipods also reduced palatability of neighbouring conspecifics. However, this effect was only observed in live tissues of F. vesiculosus. This study is the first to show that amphipods, like isopods, are capable to induce anti-herbivory defences in F. vesiculosus and that a seasonally variable effectiveness of chemical defences might serve as a dynamic control in alga–herbivore interactions.     

Trade-offs between growth and defence in two phylogenetically close invasive species

The success of an invasive plant species could be explained by trade-off between growth and defence. The aim of this paper was to explore the responses of two non-native aquatic macrophytes Elodea canadensis and Elodea nuttallii to herbivores in their introduced range. We assessed the palatability of the two phylogenetically close aquatic plant species in field and their responses to gammarid consumption in spring, summer and autumn in a microcosm experiment. We measured the variation of functional traits for each season. The traits selected were those judged most closely related to the allocation of resources to growth or to resistance against herbivores. We clearly established that the strategies of the two species were different and that their consumption rate differed in summer. In summer, E. canadensis allocated more of its resources to structural defence (leaf toughness). The increase in leaf thickness reduced the palatability of E. canadensis, whereas E. nuttallii stimulated its growth. Moreover, a decrease in dry matter content in E. nuttallii was found during the growing season in field. In autumn, both plant species accumulated nitrogen and phosphorus in their tissues. We also demonstrated that neither species induced efficient chemical defences against the herbivores. The different strategies of these two Elodea species could be explained by their different resident times in the introduced area and by an adaptation of the naturalised E. canadensis to herbivores.     

Water-borne cues of a non-indigenous seaweed mediate grazer-deterrent responses in native seaweeds, but not vice versa

Plants optimise their resistance to herbivores by regulating deterrent responses on demand. Induction of anti-herbivory defences can occur directly in grazed plants or from emission of risk cues to the environment, which modifies interactions of adjacent plants with, for instance, their consumers. This study confirmed the induction of anti-herbivory responses by water-borne risk cues between adjoining con-specific seaweeds and firstly examined whether plant-plant signalling also exists among adjacent hetero-specific seaweeds. Furthermore, differential abilities and geographic variation in plant-plant signalling by a non-indigenous seaweed as well as native seaweeds were assessed. Twelve-day induction experiments using the non-indigenous seaweed Sargassum muticum were conducted in the laboratory in Portugal and Germany with one local con-familiar (Portugal: Cystoseira humilis, Germany: Halidrys siliquosa) and hetero-familiar native species (Portugal: Fucus spiralis, Germany: F. vesiculosus). All seaweeds were grazed by a local isopod species (Portugal: Stenosoma nadejda, Germany: Idotea baltica) and were positioned upstream of con- and hetero-specific seaweeds. Grazing-induced modification in seaweed traits were tested in three-day feeding assays between cue-exposed and cue-free (?=?control) pieces of both fresh and reconstituted seaweeds. Both Fucus species reduced their palatability when positioned downstream of isopod-grazed con-specifics. Yet, the palatability of non-indigenous S. muticum remained constant in the presence of upstream grazed con-specifics and native hetero-specifics. In contrast, both con-familiar (but neither hetero-familiar) native species reduced palatability when located downstream of grazed S. muticum. Similar patterns of grazer-deterrent responses to water-borne cues were observed on both European shores, and were almost identical between assays using fresh and reconstituted seaweeds. Hence, seaweeds may use plant-plant signalling to optimise chemical resistance to consumers, though this ability appeared to be species-specific. Furthermore, this study suggests that native species may benefit more than a non-indigenous species from water-borne cue mediated reduction in consumption as only natives responded to signals emitted by hetero-specifics.     

Foliage value,apparency and defence investment in birch seedlings and trees

Summary Two factors determining plant anti-herbivore defence investment fitness loss due to herbivory and the probability of herbivory occurring in the field were quantified for birch seedlings and trees. Fitness loss due to defoliation (assumed to be related to loss of growth increment compared to controls) appeared to be greater in seedlings compared to trees, but the result was equivocal. In contrast, seedling foliage at the field site — a typical habitat for birch — suffered much less natural defoliation than tree foliage, suggesting that seedlings are markedly less apparent to most birch herbivores than trees. This low apparency should result in lower investment in anti-herbivore defences by seedlings compared to trees — and being a strong effect, should outweigh the possibly greater growth loss suffered by seedlings, which in isolation would tend to increase their optimum defence investment compared to trees. This prediction was tested using palatability trials with a wide range of common birch herbivores and by direct quantification of anti-herbivore defences. Problems and assumptions inherent in these approaches are discussed, but it seems that birch seedlings are genuinely unapparent to herbivores, and consequently do not need the degree of defence investment required by trees.     

Mechanisms and ecological consequences of plant defence induction and suppression in herbivore communities

Background Plants are hotbeds for parasites such as arthropod herbivores, which acquire nutrients and energy from their hosts in order to grow and reproduce. Hence plants are selected to evolve resistance, which in turn selects for herbivores that can cope with this resistance. To preserve their fitness when attacked by herbivores, plants can employ complex strategies that include reallocation of resources and the production of defensive metabolites and structures. Plant defences can be either prefabricated or be produced only upon attack. Those that are ready-made are referred to as constitutive defences. Some constitutive defences are operational at any time while others require activation. Defences produced only when herbivores are present are referred to as induced defences. These can be established via de novo biosynthesis of defensive substances or via modifications of prefabricated substances and consequently these are active only when needed. Inducibility of defence may serve to save energy and to prevent self-intoxication but also implies that there is a delay in these defences becoming operational. Induced defences can be characterized by alterations in plant morphology and molecular chemistry and are associated with a decrease in herbivore performance. These alterations are set in motion by signals generated by herbivores. Finally, a subset of induced metabolites are released into the air as volatiles and function as a beacon for foraging natural enemies searching for prey, and this is referred to as induced indirect defence.Scope The objective of this review is to evaluate (1) which strategies plants have evolved to cope with herbivores and (2) which traits herbivores have evolved that enable them to counter these defences. The primary focus is on the induction and suppression of plant defences and the review outlines how the palette of traits that determine induction/suppression of, and resistance/susceptibility of herbivores to, plant defences can give rise to exploitative competition and facilitation within ecological communities “inhabiting” a plant.Conclusions Herbivores have evolved diverse strategies, which are not mutually exclusive, to decrease the negative effects of plant defences in order to maximize the conversion of plant material into offspring. Numerous adaptations have been found in herbivores, enabling them to dismantle or bypass defensive barriers, to avoid tissues with relatively high levels of defensive chemicals or to metabolize these chemicals once ingested. In addition, some herbivores interfere with the onset or completion of induced plant defences, resulting in the plant’s resistance being partly or fully suppressed. The ability to suppress induced plant defences appears to occur across plant parasites from different kingdoms, including herbivorous arthropods, and there is remarkable diversity in suppression mechanisms. Suppression may strongly affect the structure of the food web, because the ability to suppress the activation of defences of a communal host may facilitate competitors, whereas the ability of a herbivore to cope with activated plant defences will not. Further characterization of the mechanisms and traits that give rise to suppression of plant defences will enable us to determine their role in shaping direct and indirect interactions in food webs and the extent to which these determine the coexistence and persistence of species.     

Plant stages with biotic,indirect defences are more palatable and suffer less herbivory than their undefended counterparts

Plants have evolved several anti‐herbivory strategies, including direct defences, such as mechanical and chemical defences, and indirect or biotic defences, such as the recruitment of defending animals. We examined whether the investment plants make in direct defences differs between those which do and do not invest in biotic defences, by comparing standing herbivory and palatability of congeneric species with and without indirect defences at two ontogenetic stages: before and after the onset of indirect defences. We used Cordia alliodora and Croton suberosus as the species with indirect defences and Cordia elaeagnoides and Croton pseudoniveus as the species without indirect defences. We predicted that herbivores would prefer to eat species and stages with indirect defences to those without them. As predicted, we found that herbivores preferred species and ontogenetic stages with indirect defences in all cases. Overall, however, natural levels of herbivory were lower in species with indirect defences. We conclude that indirect defences offer effective protection against herbivores and posit that their recruitment allows plants to reduce investment in other defence mechanisms. Our results support the notion that plants trade‐off between direct and indirect defensive strategies. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 536–543.     

High CO2 enhances the competitive strength of seaweeds over corals

Space competition between corals and seaweeds is an important ecological process underlying coral-reef dynamics. Processes promoting seaweed growth and survival, such as herbivore overfishing and eutrophication, can lead to local reef degradation. Here, we present the case that increasing concentrations of atmospheric CO(2) may be an additional process driving a shift from corals to seaweeds on reefs. Coral (Acropora intermedia) mortality in contact with a common coral-reef seaweed (Lobophora papenfussii) increased two- to threefold between background CO(2) (400 ppm) and highest level projected for late 21st century (1140 ppm). The strong interaction between CO(2) and seaweeds on coral mortality was most likely attributable to a chemical competitive mechanism, as control corals with algal mimics showed no mortality. Our results suggest that coral (Acropora) reefs may become increasingly susceptible to seaweed proliferation under ocean acidification, and processes regulating algal abundance (e.g. herbivory) will play an increasingly important role in maintaining coral abundance.     

EFFECTS OF CLIMATE CHANGE ON GLOBAL SEAWEED COMMUNITIES

Seaweeds are ecologically important primary producers, competitors, and ecosystem engineers that play a central role in coastal habitats ranging from kelp forests to coral reefs. Although seaweeds are known to be vulnerable to physical and chemical changes in the marine environment, the impacts of ongoing and future anthropogenic climate change in seaweed‐dominated ecosystems remain poorly understood. In this review, we describe the ways in which changes in the environment directly affect seaweeds in terms of their physiology, growth, reproduction, and survival. We consider the extent to which seaweed species may be able to respond to these changes via adaptation or migration. We also examine the extensive reshuffling of communities that is occurring as the ecological balance between competing species changes, and as top‐down control by herbivores becomes stronger or weaker. Finally, we delve into some of the ecosystem‐level responses to these changes, including changes in primary productivity, diversity, and resilience. Although there are several key areas in which ecological insight is lacking, we suggest that reasonable climate‐related hypotheses can be developed and tested based on current information. By strategically prioritizing research in the areas of complex environmental variation, multiple stressor effects, evolutionary adaptation, and population, community, and ecosystem‐level responses, we can rapidly build upon our current understanding of seaweed biology and climate change ecology to more effectively conserve and manage coastal ecosystems.     

Cost of chemical defence in the red alga Delisea pulchra

The concept of a cost of defence is fundamental to theories for the evolution of defences against consumers. However, the evidence for a cost of plant chemical defences is mixed, and often indirect. This is particularly true for marine macroalgae (seaweeds), for which inferences of cost to date rely almost exclusively on phenotypic correlations between one class of secondary metabolites (brown algal phlorotannins) and growth (or, in one instance, fecundity). No studies of the cost of seaweed chemical defense have experimentally manipulated the presence of secondary metabolites in a controlled fashion and only one previous study has considered genetic background as a factor. Here we measured the cost of halogenated furanones to the red seaweed Delisea pulchra in three ways: a) phenotypic correlations between concentrations of furanones and fecundity in field collected thalli; b) genetic correlations between concentrations of furanones and growth for clones of thalli grown from tetraspores, and c) by comparing growth rates of thalli for which furanone production was experimentally inhibited (furanone -) vs thalli which produced furanones (furanone +). Two of our three tests-correlations between furanones and fecundity, and the growth of furanone (+) vs furanone (−) thalli-indicated a cost of furanones to D. pulchra but genetic correlations between furanones and growth did not. We suggest that these apparently conflicting results are consistent with the consequences of apical growth in this alga, and may further result from a cost of furanones only being manifested at critical developmental stages or times of tissue differentiation.     

Competitors as accomplices: seaweed competitors hide corals from predatory sea stars

Indirect biotic effects arising from multispecies interactions can alter the structure and function of ecological communities—often in surprising ways that can vary in direction and magnitude. On Pacific coral reefs, predation by the crown-of-thorns sea star, Acanthaster planci, is associated with broad-scale losses of coral cover and increases of macroalgal cover. Macroalgal blooms increase coral–macroalgal competition and can generate further coral decline. However, using a combination of manipulative field experiments and observations, we demonstrate that macroalgae, such as Sargassum polycystum, produce associational refuges for corals and dramatically reduce their consumption by Acanthaster. Thus, as Acanthaster densities increase, macroalgae can become coral mutualists, despite being competitors that significantly suppress coral growth. Field feeding experiments revealed that the protective effects of macroalgae were strong enough to cause Acanthaster to consume low-preference corals instead of high-preference corals surrounded by macroalgae. This highlights the context-dependent nature of coral–algal interactions when consumers are common. Macroalgal creation of associational refuges from Acanthaster predation may have important implications for the structure, function and resilience of reef communities subject to an increasing number of biotic disturbances.     

Sensitivity and Acclimation of Three Canopy-Forming Seaweeds to UVB Radiation and Warming

Canopy-forming seaweeds, as primary producers and foundation species, provide key ecological services. Their responses to multiple stressors associated with climate change could therefore have important knock-on effects on the functioning of coastal ecosystems. We examined interactive effects of UVB radiation and warming on juveniles of three habitat-forming subtidal seaweeds from Western Australia–Ecklonia radiata, Scytothalia dorycarpa and Sargassum sp. Fronds were incubated for 14 days at 16–30°C with or without UVB radiation and growth, health status, photosynthetic performance, and light absorbance measured. Furthermore, we used empirical models from the metabolic theory of ecology to evaluate the sensitivity of these important seaweeds to ocean warming. Results indicated that responses to UVB and warming were species specific, with Sargassum showing highest tolerance to a broad range of temperatures. Scytothalia was most sensitive to elevated temperature based on the reduced maximum quantum yields of PSII; however, Ecklonia was most sensitive, according to the comparison of activation energy calculated from Arrhenius’ model. UVB radiation caused reduction in the growth, physiological responses and thallus health in all three species. Our findings indicate that Scytothalia was capable of acclimating in response to UVB and increasing its light absorption efficiency in the UV bands, probably by up-regulating synthesis of photoprotective compounds. The other two species did not acclimate over the two weeks of exposure to UVB. Overall, UVB and warming would severely inhibit the growth and photosynthesis of these canopy-forming seaweeds and decrease their coverage. Differences in the sensitivity and acclimation of major seaweed species to temperature and UVB may alter the balance between species in future seaweed communities under climate change.     

Effects of competition and herbivory on interactions between a hard coral and a brown alga

Despite widespread acceptance of the negative effects of macroalgae on corals, very few studies have experimentally tested the competitive nature of the interaction, and most have ignored the potential effects of corals on algae. We report the effects of herbivory and competition on the growth of the branching scleractinian coral Porites cylindrica Dana and the creeping foliose brown alga Lobophora variegata (Lamouroux) Womersley, on an inshore fringing reef of the central Great Barrier Reef. L. variegata overgrows branches of P. cylindrica from the base up, forming a distinct boundary between the alga and the coral tissue. The experiment used exclusion cages to test for effects of herbivores, and removal of algae and coral tissue, at their interaction boundary, to test for inhibition of the competitors by each other. Comparisons of coral branches with the algae present or removed showed that the presence and overgrowth of the alga caused significant coral tissue mortality. Comparisons of branches with coral tissue unmanipulated or damaged showed that the coral inhibited the overgrowth by L. variegata, but that the algae were markedly superior competitors. Importantly, reduced herbivory resulted in faster algal growth and consequent overgrowth and mortality of coral tissue, demonstrating the critical importance of herbivory to the outcome of the competitive interaction.     

Life‐history constraints in grassland plant species: a growth‐defence trade‐off is the norm

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade‐off possibilities. The competition‐defence hypothesis posits a trade‐off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth‐defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth‐defence trade‐off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life‐history groups and within the majority of individual sites. Thus, a growth‐defence trade‐off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.     

Leaf Colour as a Signal of Chemical Defence to Insect Herbivores in Wild Cabbage (Brassica oleracea)

Leaf colour has been proposed to signal levels of host defence to insect herbivores, but we lack data on herbivory, leaf colour and levels of defence for wild host populations necessary to test this hypothesis. Such a test requires measurements of leaf spectra as they would be sensed by herbivore visual systems, as well as simultaneous measurements of chemical defences and herbivore responses to leaf colour in natural host-herbivore populations. In a large-scale field survey of wild cabbage (Brassica oleracea) populations, we show that variation in leaf colour and brightness, measured according to herbivore spectral sensitivities, predicts both levels of chemical defences (glucosinolates) and abundance of specialist lepidopteran (Pieris rapae) and hemipteran (Brevicoryne brassicae) herbivores. In subsequent experiments, P. rapae larvae achieved faster growth and greater pupal mass when feeding on plants with bluer leaves, which contained lower levels of aliphatic glucosinolates. Glucosinolate-mediated effects on larval performance may thus contribute to the association between P. rapae herbivory and leaf colour observed in the field. However, preference tests found no evidence that adult butterflies selected host plants based on leaf coloration. In the field, B. brassicae abundance varied with leaf brightness but greenhouse experiments were unable to identify any effects of brightness on aphid preference or performance. Our findings suggest that although leaf colour reflects both levels of host defences and herbivore abundance in the field, the ability of herbivores to respond to colour signals may be limited, even in species where performance is correlated with leaf colour.     

Plant-mediated effects in the Brassicaceae on the performance and behaviour of parasitoids

Direct and indirect plant defences are well studied, particularly in the Brassicaceae. Glucosinolates (GS) are secondary plant compounds characteristic in this plant family. They play an important role in defence against herbivores and pathogens. Insect herbivores that are specialists on brassicaceous plant species have evolved adaptations to excrete or detoxify GS. Other insect herbivores may even sequester GS and employ them as defence against their own antagonists, such as predators. Moreover, high levels of GS in the food plants of non-sequestering herbivores can negatively affect the growth and survival of their parasitoids. In addition to allelochemicals, plants produce volatile chemicals when damaged by herbivores. These herbivore induced plant volatiles (HIPV) have been demonstrated to play an important role in foraging behaviour of insect parasitoids. In addition, biosynthetic pathways involved in the production of HIPV are being unraveled using the model plant Arabidopsis thialiana. However, the majority of studies investigating the attractiveness of HIPV to parasitoids are based on experiments mainly using crop plant species in which defence traits may have changed through artificial selection. Field studies with both cultivated and wild crucifers, the latter in which defence traits are intact, are necessary to reveal the relative importance of direct and indirect plant defence strategies on parasitoid and plant fitness. Future research should also consider the potential conflict between direct and indirect plant defences when studying the evolution of plant defences against insect herbivory.