Costs of induced defenses for the invasive plant houndstongue (<Emphasis Type="Italic">Cynoglossum officinale</Emphasis> L.) and the potential importance for weed biocontrol

Inducible plant defenses—those produced in response to herbivore feeding—are thought to have evolved as a cost-saving tactic that allows plants to enact defenses only when needed. The costs of defense can be significant, and loss of plant fitness due to commitment of resources to induced defenses could affect plant populations and play a role in determining the success or failure of weed biocontrol. We used methyl jasmonate (MeJA) to experimentally induce defenses without herbivores in invasive houndstongue plants (Cynoglossum officinale L.) in the field and measured resulting growth and fitness (plant size, seed number, and seed weight). MeJA-treated plants emitted large amounts of plant volatiles and produced leaves with twice as many trichomes as untreated plants. Plants with activated defenses had fewer leaves, were smaller, and produced nutlets that weighed less than plants not investing in defenses. These data indicate that herbivore-induced defenses are costly for houndstongue plants in their invaded range and represent significant indirect costs of herbivory beyond direct feeding damage (e.g., loss of photosynthetic tissue). Notably, the magnitude of defenses elicited upon feeding varies greatly by herbivore species and a better understanding of the costs of defense could help us predict which potential biocontrol herbivores are most likely to be effective.     

Asymmetric plant-mediated cross-effects between a herbivorous insect and a phytopathogenic fungus

• 1 Cross‐effects between a herbivorous insect and a phytopathogenic fungus on their common host plant were examined. Specifically, we addressed the questions whether (i) infection of Chinese cabbage leaves by the fungus Alternaria brassicae affects the development and host selection behaviour of the leaf beetle Phaedon cochleariae and (ii) whether herbivory influences host suitability of Chinese cabbage for A. brassicae.
• 2 Feeding on fungus‐infected leaves prolonged larval development and reduced pupal weight of P. cochleariae. Adult beetles avoided feeding and egg deposition on fungus‐infected leaves. In contrast to these local effects, no systemic effect of phytopathogenic infection on the herbivore was detected.
• 3 Herbivory did not influence fungal growth neither locally nor systemically.
• 4 Thus, our results demonstrate an asymmetric relationship between herbivore and fungus. Whereas herbivory had no visible impact on fungal growth, fungal infection of the plant induced local resistance against P. cochleariae.

     

Effects of genetic structure of Lupinus arboreus and previous herbivory on Platyprepia virginalis caterpillars

Two leaf-feeding caterpillars, western tussock moth (Orgyia vetusta) and ranchman's tiger moth (Platyprepia virginalis) are abundant on Lupinus arboreus along the California coast. Previous experiments and observations suggested that feeding caused by either of these two folivores could reduce the performance and possibly the abundance and distribution of the other species. Previous common garden experiments also indicated that genetically determined characteristics of the host plants were important for O. vetusta. Here we examined the effects of familial origin of the host plant, and previous damage caused by O. vetusta on the abundance of P. virginalis. Plants with parents from one of three locations had higher numbers of P. virginalis than plants with parents from the other two locations. However, this effect of plant origin depended on the statistical analysis and was not as strong as the effect of prior damage by O. vetusta on numbers of P. virginalis. Counter to our expectation, bushes that supported higher levels of damage by O. vetusta in the previous summer had more P. virginalis caterpillars. This strong effect could result by both moth species selecting bushes with the same traits or as the result of herbivory by O. vetusta enhancing the susceptibility of bushes for P. virginalis. Received: 28 October 1998 / Accepted: 14 March 1999     

Influence of release size on establishment and impact of a root weevil for the biocontrol of houndstongue (Cynoglossum officinale)

The root-boring weevil, Mogulones cruciger, was introduced into Canada to control the weed, houndstongue (Cynoglossum officinale). To optimise its use as a biocontrol agent, a 2-year study was performed in British Columbia, Canada to test if the number of M. cruciger released at sites predicted subsequent declines in weed populations. No, 100, 200, 300 or 400 weevils were released in 1999 at field sites (five replicates) corresponding to discrete populations of houndstongue separated by distances of 0.3–3 km. The sites were subsequently monitored for weevil establishment, population change, and host attack, and houndstongue population change. By 2001, M. cruciger had established at all 20 release sites and was present in low numbers in three of five control sites. The year following release, release size was positively correlated with number of adult weevils collected, their damage to host plants, and with subsequent numbers of larvae per plant. In contrast, houndstongue populations were reduced at the same rate and amount, regardless of the experimental release size, within 2 years of release. Significant release treatment×time interactions indicated that factors other than M. cruciger contributed to houndstongue reductions (e.g. drought). However, overall the addition of weevils accelerated the reductions relative to sites with no weevils added. Our study demonstrated that the lowest number within a range of release sizes typically used in weed biocontrol programmes (i.e. 100) was as effective as 200–400 weevils in achieving a consistent amount and rate of houndstongue reduction, and thus, could be implemented to optimise weevil use and achieve predictable biocontrol.     

Effects of an early-season folivorous moth on the success of a later-season species,mediated by a change in the quality of the shared host,Lupinus arboreus Sims

Summary Larvae of Platyprepia virginalis (Lep.: Arctiidae) and Orgyia vetusta (Lep.: Lymantriidae) feed on the foliage of bush lupine (Lupinus arboreus Sims) at Bodega Bay, California, USA, in February–April and May–July, respectively. Female O. vetusta attained lower pupal weights and produced fewer eggs on branches of L. arboreus which had experimentally received P. virginalis damage earlier in the same year, compared to those reared on control branches.Growth rates of first-instar O. vetusta were lower on undamaged leaves of previously damaged L. arboreus than on leaves of undamaged controls. This was true for lupine bushes damaged by P. virginalis in the field; and also for greenhouse-grown lupine plants damaged by Estigmene acrea (Lep.: Arctiidae) when excised foliage was used in the assay. However, fifth (last) instar female O. vetusta larvae showed no preference for damaged versus undamaged greenhouse-grown L. arboreus in their feeding or choice of a site for pupation.These experiments demonstrate the potential for competition-like interactions between herbivores mediated by changes in quality of the shared host.     

Effect of carnivory on larvae and adults of a detritivorous caddisfly, Lepidostoma complicatum: a laboratory experiment

Lepidostomatid caddisfly larvae are typical detritivores, but they occasionally eat small dead animals. A laboratory feeding experiment was conducted with Lepidostoma complicatum (Kobayashi) larvae using two different feeding treatments: leaves or leaves and dead chironomids. L. complicatum larvae showed significantly higher growth rates and adult emergence weight and a significantly earlier emergence for the leaves and dead chironomids treatment than for the leaves alone treatment. However, the adult emergence rate was not different between the two feeding treatments. Thus, it is apparent that ingestion of dead chironomids by detritivorous L. complicatum larvae positively influences larval growth rates, adult emergence weight, and larval development time.     

Within‐plant distribution of 1,4‐benzoxazin‐3‐ones contributes to herbivore niche differentiation in maize

Plant defences vary in space and time, which may translate into specific herbivore‐foraging patterns and feeding niche differentiation. To date, little is known about the effect of secondary metabolite patterning on within‐plant herbivore foraging. We investigated how variation in the major maize secondary metabolites, 1,4‐benzoxazin‐3‐one derivatives (BXDs), affects the foraging behaviour of two leaf‐chewing herbivores. BXD levels varied substantially within plants. Older leaves had higher levels of constitutive BXDs while younger leaves were consistently more inducible. These differences were observed independently of plant age, even though the concentrations of most BXDs declined markedly in older plants. Larvae of the well‐adapted maize pest Spodoptera frugiperda preferred and grew better on young inducible leaves irrespective of plant age, while larvae of the generalist Spodoptera littoralis preferred and tended to grow better on old leaves. In BXD‐free mutants, the differences in herbivore weight gain between old and young leaves were absent for both species, and leaf preferences of S. frugiperda were attenuated. In contrast, S. littoralis foraging patterns were not affected. In summary, our study shows that plant secondary metabolites differentially affect performance and foraging of adapted and non‐adapted herbivores and thereby likely contribute to feeding niche differentiation.     

Preliminary studies on Ramularia crupinae sp. nov. as a potential biological control agent for common crupina (Crupina vulgaris) in the USA

Common crupina, Crupina vulgaris (Asteraceae), is an alien weed of pastures and disturbed noncrop lands in the western USA. Research is underway in Europe to search for the natural enemies to control the weed. Among various natural enemies attacking C vulgaris, a new fungal species, Ramularia crupinae, has been found to cause serious damage to the weed. The fungus has been grown on various media and a modified V8 juice agar has proved to be suitable for both growth and sporulation. Greenhouse and field studies have shown that R. crupinae can kill infected leaves, thus reducing the plant growth, and in severe cases can kill the young plants. During preliminary host-range tests using various members of the Asteraceae, including artichoke and safflower, the fungus attacked only C. vulgaris.     

Chlorogenic acid as an antiherbivore defence of willows against leaf beetles

In the leaves of 13 Finnish willow species, the content of a phenolic, chlorogenic acid, was found to vary from 0 up to 18 mg g^–1 D.W. Effects of pure chlorogenic acid on insect feeding behaviour were tested using four common leaf beetle species which are in the field mainly found on willows with low-chlorogenic acid leaves. One species, Lochmaea capreae L., was invariably deterred by pure chlorogenic acid applied in naturally occurring concentrations on the willow leaves. Accordingly, in 2-choice laboratory feeding trials L. capreae was found to prefer low-chlorogenic acid leaves of four willow species over high-chlorogenic acid leaves of Salix pentandra L. and S. myrsinifolia Salisb. When presented on the leaves of S. phylicifolia L, pure chlorogenic acid inhibited also the feeding by Phratora polaris Sp.-Schn. Instead, chlorogenic acid had no significant effect on Ph. polaris when it was presented on the leaves of another willow S. cinerea L. In laboratory, Ph. polaris did not show general preference for willow species with low chlorogenic acid content in their leaves. Thus, the response of Ph. polaris to chlorogenic acid seems to depend on the plant species. Apparently variation in other traits such as leaf hairyness may easily override the potential effect of chlorogenic acid content on Ph. polaris. To two other leaf beetle species, Galerucella lineola F. and Plagiodera versicolora Laich., chlorogenic acid is an ineffective deterrent even at unnaturally high concentrations. In laboratory, G. lineola and P. versicolora did not prefer willows with low chlorogenic acid content in their leaves. Thus, among four studied leaf beetle species, only L. capreae seems to be clearly affected by this phenolic. Therefore, overall importance of chlorogenic acid as a defence against willow-feeding leaf beetles appears to be very limited.     

The effect of a foliar disease (rust) on the development of Gastrophysa viridula (Coleoptera: Chrysomelidae)

Abstract.

• 1 Gastrophysa viridula Degeer (Coleoptera: Chrysomelidae) and the pathogenic rust fungus Uromyces rumicis (Schum.) Wint. both occur on leaves of Rumex crispus L. and R.obtusifolius L. Individual stages of beetle development, and egg laying, were compared on healthy and infected leaves of each plant species in the laboratory. Oviposition choice was investigated in the field and laboratory.
• 2 Beetles reared on infected leaves of each species had greater larval mortality and slower development than those reared on healthy leaves. Although larvae feeding on infected leaves consumed up to 2.5 times more dry weight than those reared on healthy leaves, they had a lower relative growth rate and pupated at a lower weight. These changes were consistent with the reduced nutritive quality of rust-infected Rumex leaves.
• 3 Fecundity of beetles reared on infected leaves of both species was considerably reduced. Eggs laid by beetles feeding on infected R.crispus leaves also had a reduced viability.
• 4 The beetle developed consistently poorer on healthy R.crispus than on healthy R.obtusifolius throughout its life-cycle. Differences in larval performance were greater between host species than between infected and healthy leaves.
• 5 Oviposition was similar on infected and healthy R.crispus in both the laboratory and field. However, adults consumed less, and laid fewer eggs on infected than on healthy R.obtusifolius. The pattern of egg laying on different aged leaves was affected by rust infection: a greater proportion of eggs was laid on the older, infected leaves, than on the equivalent aged leaves on the healthy plants. Few larvae survived from eggs laid on rusted leaves in the field.

     

Lack of susceptibility of soil‐inhabiting Platyprepia virginalis caterpillars,a native arctiid,to entomopathogenic nematodes in nature

Entomopathogenic nematodes (EPNs) can kill and regulate populations of soil‐inhabiting insects, but studies evaluating these interactions in native ecosystems are rare. The objective of this study was to examine the effects of EPNs on a non‐agricultural caterpillar, Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae), under natural conditions. Platyprepia virginalis caterpillars live in litter on the soil surface feeding beneath bush lupine during summer, autumn, and winter. Initial laboratory assays revealed that the caterpillars were vulnerable to at least two species of EPNs with which they co‐occur in the coastal prairie in northern California (USA). In contrast to laboratory assays, caterpillars survived exposure to prairie soil containing EPNs under natural conditions in field assays. To better understand the divergence between laboratory and field results for this native caterpillar, we used sentinel insects [Galleria mellonella L. (Lepidoptera: Pyralidae)] to identify particular locations where EPNs were present in the field. Platyprepia virginalis caterpillars were caged at these sites but again showed no evidence of susceptibility to EPNs. Platyprepia virginalis caterpillars reduce their exposure to EPNs by spending their time in and above the litter rather than contacting the soil when given the choice in nature. We conclude that P. virginalis is unlikely to serve as a reservoir for EPNs and that nematodes are unlikely to be important mortality factors for P. virginalis in this natural system.     

The effects of wilt symptom development and peroxidase induction on interactions between vascular wilt bacteria and cucumber beetles

Infection of cucumber (Cucumis sativus L.) with the bacterial pathogen Erwinia tracheiphila E. F. Smith causes vascular wilt disease in leaves, which may alter the suitability of the host plant for insects and other pathogens. In this study, densities of spotted (Diabrotica undecimpunctata howardi Barber) and striped (Acalymma vittata (Fab.) cucumber beetles (Coleoptera: Chrysomelidae) were higher on wilted leaves of mature and seedling field plants inoculated with E. tracheiphila. Bacterial infection or feeding by D. undecimpunctata howardii beetles increased total peroxidase enzyme activity in inoculated or infested leaves of greenhouse seedlings, but only beetle feeding induced higher activities in untreated systemic leaves on the same plants. Neither bacterial infection nor beetle infestation led to the development of systemic acquired resistance (SAR) to the fungal pathogen Colletotrichum orbiculare (Berk & Mont.) Arx. Susceptibility to this fungus was greater on E. tracheiphila-infected plants than on controls. The positive association between leaf wilt symptom development and beetle occurrence thus occurs in the presence of an oxidative but not anti-pathogenic response induced by both the insect and the pathogen.     

How do soil nutrients affect within-plant patterns of herbivory in seedlings of Eucalyptus nitens?

This study assessed how the palatability of leaves of different age classes (young, intermediate and older) of Eucalyptus nitens seedlings varied with plant nutrient status, based on captive feeding trials with two mammalian herbivores, red-bellied pademelons (Thylogale billardierii), and common brushtail possums (Trichosurus vulpecula). Seedlings were grown under three nutrient treatments (low, medium and high), and we determined how palatability was related to chemical and physical characteristics of the leaves. Pademelons ate more older leaves than young and intermediate leaves for all treatments. This pattern was best explained by sideroxylonals (formylated phloroglucinol compounds known to deter herbivory by other marsupials), and/or essential oil compounds that were present in lower concentrations in older leaves. In the low-nutrient treatment, possums also ate more of the older leaves. However, in the medium- and high-nutrient treatments, possums ate more intermediate leaves than older leaves and showed a behavioural preference for young leaves (consuming younger leaves first) over intermediate and older leaves, in spite of high levels of sideroxylonals and essential oils. The young leaves did, however, have the highest nitrogen concentration of all the leaf age classes. Thus, either sideroxylonals and essential oils provided little or no deterrent to possums, or the deterrent was outweighed by other factors such as high nitrogen. This study indicates that mammalian herbivores show different levels of relative use and damage to leaf age classes at varying levels of plant nutrient status and, therefore, their impact on plant fitness may vary with environment.     

Effect of plant age and turfgrass competition on the efficacy of the Sclerotinia minor granular bioherbicide on broadleaf plantain and prostrate knotweed

Broadleaf plantain and prostrate knotweed are important weeds of turfgrass systems. The fungus Sclerotinia minor Jagger (IMI 344141) has been registered as a biological herbicide (Sarritor?) for dandelion (Taraxacum officinale) in Canadian turfgrass habitats. The objective of this study was to evaluate the effect of plant age and turfgrass environment on the efficacy of S. minor against two additional weeds; broadleaf plantain and prostrate knotweed. The turfgrass environment alone exerted significant reduction of above and below ground biomass of broadleaf plantain, to the same magnitude as the S. minor treatment in a grass-free environment. Prostrate knotweed biomass, however, was not reduced to this extent by competition with turfgrass. In the presence of grass, S. minor caused a significant biocontrol effect on all studied variables with more than 90% above ground damage on both weed species. Severe damage occurred on 3–6-week-old plantains with 100% above and below ground reduction, although smaller dry weight reductions were observed on older plantains. Treatment with S. minor reduced the dry matter of 3–5-week-old prostrate knotweed by 65–85%, but less damage occurred on older prostrate knotweed. The bioherbicide fungus is destructive for both species, but variation in area of contact due to different growth forms, growth rates and resource allocation patterns due to different life forms resulted in different biocontrol efficacy on the two species. Control of broadleaf plantain was effective – similar to that previously reported for dandelion – but control of prostrate knotweed was only partial.     

Soil-Behaviour of Phytophthora clandestina

Investigations were undertaken to study the nature and behaviour of P. clandestina in soil. The pathogen was recovered only from soil sievings 250–499 μm and 500 μm–0.99 mm, containing small root fragments. In soil the introduced inoculum of the fungus was incapable of saprophytically and competitively colonizing the dead cotyledons of subterranean clover used as bait material. Exposure of the inoculum to increasing numbers of microbes by adding greater proportions of nonsterile fields, oil to the growth medium of the plant had no significant effect on survival rate and fresh shoot weight of subterranean clover. However, microbes present in the field soil reduced the severity of root rot of subterranean clover. P. clandestina was, able to spread between 15–30 mm through pasteurized soil within a period of 20 days.     

Within-plant variation in induced defence in developing leaves of cotton plants

According to optimal defence theory (ODT) plants should invest in stronger defence in the most valuable parts, such as reproductive or young tissue. Cotton plants are known to express high resistance to herbivore feeding in the developing leaves at the top of the plant after herbivore-damage. Cotton plants also have developing leaves on side shoots from the nodes all along the plant. This allowed us to investigate within-plant distribution of defence between younger tissues at different locations on the plant. We found that all developing leaves showed increased resistance to feeding by larvae of the generalist moth Spodoptera littoralis after herbivore damage to leaves of the plant. An increase in the concentration of secondary compounds, terpenoid aldehydes, was found in developing leaves both at the top of the plant and on the side shoots. However, the resistance was stronger in the top leaves than in the side shoot leaves, showing that there is intra-plant variation in the induced response between different leaves of the same age. After the initial damage, larval feeding mainly occurred on the older, fully developed true leaves. Furthermore, the herbivore-induced resistance in the developing leaves reduced upward movement of feeding larvae on the plant and reduced the feeding on the upper parts of the plant over a period of at least 6 days. The plant thus benefits from defending all developing leaves by directing feeding to older, less valuable leaves and lower parts of the plant. The observed distribution of defence within cotton plants supports ODT.     

Effects of trichomes on the behavior and distribution of Platyprepia virginalis caterpillars

Trichomes are an important physical resistance mechanism of plants, as they reduce insect herbivore movement, feeding, and digestion. However, we know little about how trichomes influence herbivore distributions and populations. We conducted laboratory and field experiments to evaluate the preferences of Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae) caterpillars to natural and manipulated densities of trichomes on their primary food, Lupinus arboreus Sims (Fabaceae). We then conducted field surveys to determine whether variation in trichome density among lupine bushes affected caterpillar spatial distribution on the landscape. Platyprepia virginalis caterpillars preferred lupine leaves with fewer trichomes in choice and no‐choice experiments. In the field, caterpillar feeding damage was found more often on leaves with fewer trichomes. These preferences scaled up to the level of bushes in the landscape such that more caterpillars were found on bushes with lower trichome densities than on bushes with higher trichome densities. This is one of few studies to show the potential for trichome density to influence herbivore population size and distribution in a natural system at a landscape level. The results are consistent with trichomes functioning as a resistance mechanism with consequences for herbivore choice, performance, and distribution.     

Two Phytotoxins Isolated from the Pathogenic Fungus of the Invasive Weed Xanthium italicum

Alternariol and altenuisol were isolated as the major phytotoxins produced by an Alternaria sp. pathogenic fungus of the invasive weed Xanthium italicum. Altenuisol exhibited stronger phytotoxic effect compared with alternariol. At 10 μg/mL, alternariol and altenuisol promoted root growth of the monocot plant Pennisetum alopecuroides by 11.1 % and 75.2 %, respectively, however, inhibitory activity was triggered by the increase of concentration, with root elongation being suppressed by 35.5 % and 52.0 % with alternariol and altenuisol at 1000 μg/mL, respectively. Alternariol slightly inhibited root length of the dicot plant Medicago sativa at 10–1000 μg/mL, whereas altenuisol stimulated root growth by 51.0 % at 10 μg/mL and inhibited root length by 43.4 % at 200 μg/mL. Alternariol and altenuisol did not exert strong regulatory activity on another dicot plant, Amaranthus retroflexus, when tested concentration was low, however, when the concentration reached 1000 μg/mL, they reduced root length by 68.1 % and 51.0 %, respectively. Alternariol and altenuisol exerted similar effect on shoot growth of three tested plants but to a lesser extent. It is noteworthy to mention that this is the first report on the phytotoxicity of altenuisol.     

Interactions between the diamondback moth, Plutella xylostella L. and glasshouse and outdoor-grown cabbage cultivars

Interactions between Brassica cultivars (cvs) and adult and larval stages of Plutella xylostella L. were examined. In six-way choice oviposition experiments, glasshouse-grown Chinese cabbage cv. Tip Top was preferred to five outdoor-grown temperate cabbage cultivars but was similar to four out of five tropical cultivars. Bionomic studies with neonate larvae on temperate cabbage cultivars showed that mean larval duration was longer and percentage survival less on older compared with younger plants but that plant age or cultivar had little effect on pupal weight. The fecundity of P. xylostella reared on outdoor-grown cabbage cultivars varied ten-fold between the poorest and the best cultivar but plant age had little effect on P. xylostella reared on Chinese cabbage. In a glasshouse experiment, survival of P. xylostella on Chinese cabbage was significantly greater compared with field-grown glossy-leaved, normal bloom green cabbage and red cabbage. In bioassays with neonate larvae, when the leaves were placed vertically instead of horizontally a significant level of intrinsic resistance was revealed for glasshouse and field-grown glossy-leaved cabbage but not for Chinese cabbage. Observed differences in host status to P. xylostella are discussed in relation to the potential for partial plant resistance in control programmes.     

Influence of endophyte infection, plant age and harvest interval on Rhopalosiphum padi survival and its relation to quantity of N-formyl and N-acetyl loline in tall fescue

Rhopalosiphum padi L. (Homoptera: Aphididae) is sensitive to loline alkaloids present in tall fescue, Festuca arundinacea Shreb., infected with the endophytic fungus, Acremonium coenophialum Morgan-Jones & Gams. Aphid survival was higher on endophyte-free plants regardless of plant age after germination or age of regrowth tissue after clipping. Survival of aphids on endophyte-infected grass was lower on young tissue but increased as plants aged, although it never reached the same level on endophyte-free plants. Both N-formyl and N-acetyl loline increased as uncut or regrowth tissue aged; however, this was influenced by the age of the plant at the initial cut and the clipping frequency. Although even small amounts of loline cause high aphid mortality, the aphids are able to survive on endophyte-infected plants if the tillers have senescing leaves which contain lower amounts of loline. Preference for senescing leaves may help R. padi avoid plant parts containing high amounts of toxic allelochemicals, thus contributing to higher numbers of aphids on older, endophyte-infected plants.