Indirect cost of a defensive trait: variation in trichome type affects the natural enemies of herbivorous insects on <Emphasis Type="Italic">Datura wrightii</Emphasis>

The costs and benefits of defensive traits in plants can have an ecological component that arises from the effect of defenses on the natural enemies of herbivores. We tested if glandular trichomes in Datura wrightii, a trait that confers resistance to several species of herbivorous insects, impose an ecological cost by decreasing rates of predation by the natural enemies of herbivores. For two common herbivores of D. wrightii, Lema daturaphila and Tupiocoris notatus, several generalized species of natural enemies exhibited lower rates of predation on glandular compared to non-glandular plants. Lower rates of predation were associated with reductions in the residence time and foraging efficiency of natural enemies on plants with glandular trichomes, but not with direct toxic effects of glandular exudate. Our results suggest that the benefit of resistance to herbivores conferred by glandular trichomes might be offset by the detrimental effect of this trait on the natural enemies of herbivores, and that the fitness consequences of this trichome defense might depend on the composition and abundance of the natural-enemy community.     

Good mothers,bad mothers,and the nature of resistance to herbivory in <Emphasis Type="Italic">Solidago altissima</Emphasis>

Evidence of poor correspondence between an insect herbivore’s oviposition preferences and the performance of its offspring has generally been attributed either to maladaptive behavior of the insect mother or inadequate measurement by the researcher. In contrast, we hypothesize that many cases of “bad mothers” in herbivores may be a byproduct of the hierarchical way natural selection works on resistance in host plants. Epistatic selection on the components of resistance (i.e., antixenosis and antibiosis) may generate negative genetic correlations between the resistance components, which could counteract the efforts of herbivores to oviposit on the best hosts for the performance of their offspring. In common garden and greenhouse experiments, we measured aspects of antixenosis and antibiosis resistance in 26 genets of tall goldenrod, Solidago altissima, against two common herbivores: the gall-inducing fly Eurosta solidaginis and the spittlebug Philaenus spumarius. Goldenrod antixenosis and antibiosis were positively correlated against E. solidaginis and negatively correlated against P. spumarius. Analogously, population-wide preference–performance correlations were positive for the gall flies and negative for the spittlebugs. Several natural history differences between the two insects could make gall flies better mothers, including better synchrony of the phenologies of the flies and the host plant, the much narrower host range of the gall flies than the spittlebugs, and the more sedentary lifestyle of the gall fly larvae than the spittlebug nymphs. If these results are typical in nature, then negative genetic correlations in antixenosis and antibiosis in plants may often result in zero or negative population-wide correlations between preference and performance in herbivores, and thus may be an important reason why herbivorous insects often appear to be bad mothers.     

Midgut cysteine protease-inhibiting activity in Trichoplusia ni protects the peritrophic membrane from degradation by plant cysteine proteases

The action of plant cysteine proteases on the midgut peritrophic membrane (PM) of a polyphagous herbivorous lepidopteran, Trichoplusia ni, was studied. Proteins in PMs isolated from T. ni larvae were confirmed to be highly resistant to the serine proteinases trypsin and chymotrypsin, but were susceptible to degradation by plant cysteine proteases, which is consistent with the known molecular and biochemical characteristics of the T. ni PM proteins. However, the PM proteins were not degraded by plant cysteine proteases in larvae or in the presence of larval midgut fluid in vitro. With further biochemical analysis, cysteine protease-inhibiting activity was identified in the midgut fluid of T. ni larvae. The cysteine protease-inhibiting activity was heat resistant and active in the tested pH range from 6.0 to 10.0, but could be suppressed by thiol reducing reagents or reduced by treatment with catalase. In addition to T. ni, cysteine protease-inhibiting activity was also identified from two other polyphagous Lepidoptera species, Helicoverpa zea and Heliothis virescens. In conclusion, results from this study uncovered that herbivorous insects may counteract the attack of plant cysteine proteases on the PM by inhibiting the potentially insecticidal cysteine proteases from plants in the digestive tract. However, the biochemical identity of the cysteine protease-inhibiting activity in midgut fluid has yet to be identified.     

Fitness of Bt‐resistant cabbage loopers on Bt cotton plants

Development of resistance to the insecticidal toxins from Bacillus thuringiensis (Bt) in insects is the major threat to the continued success of transgenic Bt crops in agriculture. The fitness of Bt‐resistant insects on Bt and non‐Bt plants is a key parameter that determines the development of Bt resistance in insect populations. In this study, a comprehensive analysis of the fitness of Bt‐resistant Trichoplusia ni strains on Bt cotton leaves was conducted. The Bt‐resistant T. ni strains carried two genetically independent mechanisms of resistance to Bt toxins Cry1Ac and Cry2Ab. The effects of the two resistance mechanisms, individually and in combination, on the fitness of the T. ni strains on conventional non‐Bt cotton and on transgenic Bt cotton leaves expressing a single‐toxin Cry1Ac (Bollgard I) or two Bt toxins Cry1Ac and Cry2Ab (Bollgard II) were examined. The presence of Bt toxins in plants reduced the fitness of resistant insects, indicated by decreased net reproductive rate (R₀) and intrinsic rate of increase (r). The reduction in fitness in resistant T. ni on Bollgard II leaves was greater than that on Bollgard I leaves. A 12.4‐day asynchrony of adult emergence between the susceptible T. ni grown on non‐Bt cotton leaves and the dual‐toxin‐resistant T. ni on Bollgard II leaves was observed. Therefore, multitoxin Bt plants not only reduce the probability for T. ni to develop resistance but also strongly reduce the fitness of resistant insects feeding on the plants.     

Eucalypt responses to fertilization and reduced herbivory

Summary We manipulated soil fertility and insect attack for two species of Eucalyptus in natural stands of subalpine woodland on shallow, infertile granitic soils. E. pauciflora and E. stellulata responded in similar ways to simultaneous insecticide and fertilizer treatments. Eliminating herbivorous insects produced the largest changes — improved plant growth, increased leaf N and P, and reduced leaf specific density. Fertilizer regime modified some leaf properties, but had little effect on tree growth. E. stellulata trees were initially shorter than E. pauciflora, but grew faster without herbivores; by the end of the experiment both species were the same size when herbivores were removed. Foliage N and P levels increased most in trees with the most balanced fertilizer addition (NPK), and increased in all trees protected from insects, regardless of fertilizer regime. In this system, herbivorous insects exacerbated the effects of nutrientpoor soils, and may affect dominance of Eucalyptus species in mature forests.     

Host plant species affects the quality of the generalist Trichoplusia ni as a host for the polyembryonic parasitoid Copidosoma floridanum

Diet of herbivorous insects can influence both the herbivores and their natural enemies. We examined the direct and indirect effects of diet on the interactions between the polyphagous herbivore Trichoplusia ni Hübner (Lepidoptera: Noctuidae) and its polyembryonic parasitoid Copidosoma floridanum Ashmead (Hymenoptera: Encyrtidae). To determine how host plant species and host plant iridoid glycoside content affect host caterpillars and their parasitoids, parasitized and unparasitized T. ni were given leaves of either Plantago lanceolata L., which contains the iridoid glycosides aucubin and catalpol, Plantago major L. (Plantaginaceae), which contains only aucubin, or Taraxacum officinale F.H. Wigg (Asteraceae), which contains neither. Survival of unparasitized T. ni was much lower when fed P. major compared with the other two host plants, whereas pupae were smallest when fed T. officinale and developed most slowly when fed P. lanceolata as larvae. Neither aucubin nor catalpol were detected in intact Plantago‐fed T. ni larvae or their hemolymph, and only trace amounts of aucubin were detected in frass, suggesting that these compounds are mostly metabolized in the midgut and are not encountered by the parasitoid. Copidosoma floridanum clutch size was almost doubled when reared from P. lanceolata‐fed T. ni compared with T. officinale‐fed larvae and tripled compared with P. major‐fed larvae, although the percent of parasitoids surviving to adulthood was uniformly high regardless of host diet. The observed variation in C. floridanum fitness among host diets is likely mediated by the effect of the diets on host quality, which in turn may be influenced more by other factors in the host plants than their iridoid glycoside profiles. Interactions between plant metabolites, generalist herbivores like T. ni, and their parasitoids may be predominantly indirect.     

Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.     

Parasitoids follow herbivorous insects to a novel host plant,generalist predators less so

The ‘enemy‐free space’ hypothesis predicts that herbivorous insects can escape their natural enemies by switching to a novel host plant, with consequences for the evolution of host plant specialisation. However, if natural enemies follow herbivores to their novel host plants, enemy‐free space may only be temporary. We tested this by studying the colonisation of the introduced tree Eucalyptus grandis (Hill) Maiden (Myrtaceae) by insects in Brazil, where various species of herbivores have added eucalyptus to their host plant range, which consists of native myrtaceous species such as guava. Some herbivores, for example, Thyrinteina leucoceraea Ringe (Lepidoptera: Geometridae), cause outbreaks in eucalyptus plantations but not on guava, possibly because eucalyptus offers enemy‐free space. We sampled herbivores (mainly Lepidoptera species) and natural enemies on eucalyptus and guava and assessed parasitism of Lepidoptera larvae on both host plant species during ca. 2 years. Overall, predators were encountered more frequently on guava than on eucalyptus. In contrast, parasitoids were encountered equally and parasitism rates of Lepidoptera larvae were similar on both host plants. This indicates that herbivores may escape some enemies by moving to a novel host plant. However, this escape may be temporary and may vary with time. We argue that studying temporal and spatial patterns of enemy‐free space and the response of natural enemies to host use changes of their herbivorous prey is essential for understanding the role of natural enemies in the evolution of host plant use by herbivorous arthropods.     

Differential induction of plant chemical defenses by parasitized and unparasitized herbivores: consequences for reciprocal,multitrophic interactions

Insect parasitoids can play ecologically important roles in virtually all terrestrial plant–insect herbivore interactions, yet whether parasitoids alter the defensive traits that underlie interactions between plants and their herbivores remains a largely unexplored question. Here, we examined the reciprocal trophic interactions among populations of the wild cabbage Brassica oleracea that vary greatly in their production of defensive secondary compounds – glucosinolates (GSs), a generalist herbivore, Trichoplusia ni, and its polyembryonic parasitoid Copidosoma floridanum. In a greenhouse environment, plants were exposed to either healthy (unparasitized), parasitized, or no herbivores. Feeding damage by herbivores induced higher levels of the indole GSs, glucobrassicin and neoglucobrassicin, but not any of the other measured GSs. Herbivores parasitized by C. floridanum induced cabbage plants to produce 1.5 times more indole GSs than levels induced by healthy T. ni and five times more than uninduced plants. As a gregarious endoparasitoid, C. floridanum causes its host T. ni to feed more than unparasitized herbivores resulting in increased induction of indole GSs. In turn, herbivore fitness parameters (including differential effects on male and female contributions to lifetime fecundity in the herbivore) were negatively correlated with the aliphatic GSs, sinigrin and gluconapin, whereas parasitoid fitness parameters were negatively correlated with the indole GSs, glucobrassicin and neoglucobrassicin. That herbivores and their parasitoids appear to be affected by different sets of GSs was unexpected given the intimate developmental associations between host and parasitoid. This study is the first to demonstrate that parasitoids, through increasing feeding by their herbivorous hosts, can induce higher levels of non‐volatile plant chemical defenses. While parasitoids are widely recognized to be ubiquitous in most terrestrial insect herbivore communities, their role in influencing plant–insect herbivore relationships is still vastly underappreciated.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

Leaf shape variation and herbivore consumption and performance: a case study with Ipomoea hederacea and three generalists

The effect of leaf shape variation on plant-herbivore interactions has primarily been studied from the perspective of host seeking behavior. Yet for leaf shape to affect plant-herbivore coevolution, there must be reciprocal effects of leaf shape variation on herbivore consumption and performance. We investigated whether alternative leaf morphs affected the performance of three generalist insect herbivores by taking advantage of a genetic polymorphism and developmental plasticity in leaf shape in the Ivyleaf morning glory, Ipomoea hederacea. Across four experiments, we found variable support for an effect of leaf shape genotype on insects. For cabbage loopers (Trichoplusia ni) and corn earworms (Helicoverpa zea) we found opposing, non-significant trends: T. ni gained more biomass on lobed genotypes, while H. zea gained more biomass on heart-shaped genotypes. For army beetworms (Spodoptera exigua), the effects of leaf shape genotype differed depending on the age of the plants and photoperiod of growing conditions. Caterpillars feeding on tissue from older plants (95 days) grown under long day photoperiods had significantly greater consumption, dry biomass, and digestive efficiency on lobed genotypes. In contrast, there were no significant differences between heart-shaped and lobed genotypes for caterpillars feeding on tissue from younger plants (50 days) grown under short day photoperiods. For plants grown under short days, we found that S. exigua consumed significantly less leaf area when feeding on mature leaves than juvenile leaves, regardless of leaf shape genotype. Taken together, our results suggest that the effects of leaf shape variation on insect performance are likely to vary between insect species, growth conditions of the plant, and the developmental stage and age of leaves sampled. Handling editor: May Berenbaum.     

Sulfhydryl-reagent alteration of soybean resistance to the cabbage looper, Trichoplusia ni

The hypothesis that the sulfhydryl reagent, N-ethylmaleimide, would function as an elicitor of alterable resistance in soybean (Glycine max) plants to Trichoplusia ni herbivory was tested experimentally under greenhouse conditions. This elicitory chemical, which allows receptor thiols to add across its carbon-carbon double bond, altered the resistance in one or more leaves of plants at one or more intervals after treatment; and thus yielded results supporting the hypothesis. Leaf dipping and soil application were both effective methods of treatment. Results support the interpretation that an elicitor may function in intact plants by altering the integrity of sulfhydryl groups in receptor macromolecules which are also involved in signaling a change in the plant's biosynthesis of characteristic defensive compounds such as phenylpropanoids including antifeedant and antibiotic flavonoids. Induced feeding non-preference by T. ni was highly correlated positively with the amount of glyceollins in the leaf.     

Effect of plant patch shape on the distribution and abundance of three lepidopteran species associated with Brassica oleracea

• 1 Plant patch shape may affect the abundance of herbivorous insects. Patches of the same size but longer or irregular have a higher perimeter/area relationship (P/A) than square or regular ones, which may determine the immigration, emigration and abundance of individuals in the patch.
• 2 Only specialist species should be affected by plant patch shape. Those species that are more abundant in smaller patches should be more abundant in patches with higher P/A, whereas those that are more abundant in larger patches should be more abundant in patches with lower P/A.
• 3 We studied the density of eggs, larvae and pupae of Pieris brassicae, Plutella xylostella and Trichoplusia ni in square (low P/A) and I‐shaped (high P/A) patches of 144 plants of Brassica oleracea. We also estimated their immigration to these patches, and the final plant weight.
• 4 Plant patch shape affected the abundance, but not the distribution, of the two specialist species. Whereas P. brassicae was denser in I‐shaped patches, P. xylostella was more abundant in square patches. The generalist T. ni was not affected by patch shape. Immigration of P. brassicae was higher in I‐shaped patches, but immigration of P. xylostella and T. ni was not affected by patch shape. Plants were heavier in the centre of square patches.
• 5 Our results suggest that plant patch shape affects the density of herbivorous insects and should be considered independently from other plant patch variables when studying the population dynamics of these organisms.

     

EVOLUTIONARY ECOLOGY OF DATURA STRAMONIUM L. IN CENTRAL MEXICO: NATURAL SELECTION FOR RESISTANCE TO HERBIVOROUS INSECTS

It has been assumed that herbivores constitute a selective agent for the evolution of plant resistance. However, few studies have tested this hypothesis. In this study, we look at the annual weed Datura stramonium for evidence of current natural selection for resistance to herbivorous insects. Paternal half-sib families obtained through controlled crosses were exposed to herbivores under natural conditions. The plants were damaged by two folivorous insects: the tobacco flea beetle Epitrix parvula and the grasshopper Sphenarium purpurascens. Selection was estimated using a multiple-regression analysis of plant size and of damage by the two herbivores on plant fitness measured as fruit production for both individual phenotypes and family breeding values (genetic analysis). Directional phenotypic selection was detected for both larger plant size and lower resistance to the flea beetles, whereas stabilizing phenotypic selection was revealed for resistance to S. purpurascens. However, performing the same analyses on the breeding values of the characters revealed directional and stabilizing selection only for plant size. Thus, no agreement existed between the results of the two types of analyses, nor was there any detectable potential for genetic change in the studied population because of selection on herbivore resistance. The narrow-sense heritability of every trait studied was small (all <0.1) and not different from zero. The potential for evolutionary response to natural selection for higher resistance to herbivores in the studied population of D. stramonium is probably limited by lack of genetic variation. Natural selection acts on phenotypes, and the detection of phenotypic selection on resistance to herbivores confirms their ecological importance in determining plant fitness. However, evolutionary inferences based solely on phenotypic selection analyses must be interpreted with caution.     

Induced resistance in intertidal macroalgae modifies feeding behaviour of herbivorous snails

Herbivory in terrestrial and marine systems can induce changes in plant chemistry affecting the foraging behaviour of herbivores. A model based on terrestrial plant-herbivore interactions predicts herbivory-induced changes in leaf chemistry to be manifested in (1) increased herbivore mobility, (2) increased feeding dispersal and (3) reduced tissue consumption by herbivores. This study is the first to demonstrate that herbivory-induced changes in the tissue chemistry of the brown seaweed Ascophyllum nodosum elicit the same response in the feeding behaviour of the gastropod Littorina obtusata as predicted for herbivorous insects, providing good evidence for the models validity across different ecosystems. The potential benefit of increased feeding dispersal to terrestrial plants as suggested by the model is the prevention of concentrated damage to apical tissues thereby preserving the plants ability to compete for light; A. nodosum does not conform to these predictions. Increased dispersal of feeding damage on A. nodosum away from primary frond tissues would reduce the likelihood of frond breakage implying a fitness benefit of induced resistance.     

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.     

Effects of nonhost and host plants on insect herbivory covarying with plant size in the cruciferous plant <Emphasis Type="Italic">Turritis glabra</Emphasis>

Correlation between plant size and reproductive output may be modified by herbivory in accordance with host plant density and the presence of nonhost plants. To elucidate the effects of nonhost plant density and host plant density on the intensity of herbivory and reproductive output of the host plant in relation to plant size under natural conditions, we investigated the abundance of three lepidopteran insects, Plutella maculipennis, Anthocharis scolymus, and Pieris rapae the intensity of herbivory, and fruit set of their host plant, Turritis glabra (Cruciferae). To elucidate the effects of nonhost and host plant density, we selected four categories of plots under natural conditions: low density of nonhost and high density of host plants; low density of both nonhost and host plants; high density of both nonhost and host plants; and high density of nonhost and low density of host plants. The plant size indicated by stem diameter was a good predictor of the abundance of all herbivorous species. The effects of density of nonhost and host plants on the abundance of insects varied among species and stages of insects. As the abundance of insects affected the intensity of herbivory, herbivory was more apparent on larger host plants in plots with low density of both nonhost and host plants. Consequently, the correlation between plant size and the number of fruits disappeared in low plots with density of both nonhost and host plants. In this T. glabra– herbivorous insect system, the density of nonhost plants and host plants plays an important role in modifying the relationship between plants and herbivores under natural conditions. Received: July 19, 1999 / Accepted: June 15, 2000     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demonstrated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the secondary metabolites accumulation and is referred to as a copperinducible elicitor (CIE). Finally, we discuss how copperinducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

THE EVOLUTION OF RESISTANCE TO HERBIVORY IN IPOMOEA PURPUREA. II. NATURAL SELECTION BY INSECTS AND COSTS OF RESISTANCE

An important component of the process of coevolution between plants and their insect herbivores is the imposition of selection on plants by insects. Although such selection has been inferred from indirect evidence, little direct evidence for it exists. One goal of this study was to seek direct evidence by determining, for a single plant-herbivore system, whether insect herbivores impose selection on their host plants. A second goal was to determine whether costs are associated with genotypes that confer resistance to herbivores, as has been commonly postulated. The annual morning glory, Ipomoea purpurea, exhibits genetic variation in resistance to four different types of insects. For three of these types, most of the genetic variation is additive. Removal of insect herbivores increased the number of seeds produced by I. purpurea by 20% and eliminated additive genetic variation for seed number (fitness). This result implies that herbivores impose selection on some trait(s) of their host plants. Coupled with selection for decreased damage by corn earworms, as revealed by a negative additive genetic covariance between damage and fitness, this result suggests that insect herbivores impose selection on resistance to corn earworms in I. purpurea. Two types of cost of resistance to herbivores were sought in I. purpurea: 1) internal trade-offs in allocation of resources and 2) ecological trade-offs between resistances to different insects. No costs of either type were detected. This result suggests that cost-benefit arguments that attempt to predict the evolution of levels of resistance to herbivores are not applicable to I. purpurea.