Associational susceptibility in broccoli: mediated by plant volatiles,impeded by ozone

Plant‐emitted volatile organic compounds (VOCs) mediate interactions within a plant community. Typically, receiving a signal from a damaged neighbour enhances the defensive attributes of a receiver plant. The mechanisms underlying plant–plant interactions may be divided into active and passive processes, both of which involve transit of VOCs between plants and are vulnerable to environmental perturbation. Numerous studies have documented between‐plant interactions, but the specific effects on a receiver plant's interactions with herbivores have received little attention. Moreover, the relative contributions of active and passive processes to plant defence and the effects of environmental pollutants on the processes have been largely unexplored. We used a system comprising Brassica oleracea var. italica (broccoli) and the specialist herbivore Plutella xylostella to test whether plants previously exposed to herbivore‐damaged neighbours differed from nonexposed plants in their susceptibility to oviposition. We then investigated the roles of active and passive mechanisms in our observations and whether differences in susceptibility remained under elevated ozone concentrations. Plants exposed to herbivore‐damaged neighbours were more susceptible to oviposition than plants exposed to undamaged neighbours, which indicates associational susceptibility. Mechanistically, active and passive volatile‐mediated processes occurred in tandem with the passive process – involving adsorption of sesquiterpenes to receiver plants – appearing to structure the oviposition response. Exposure to ozone rapidly degraded the sesquiterpenes and eliminated the associational susceptibility. Plant volatiles have typically been thought to play roles in between‐plant interactions and to promote receiver plant defence. Here, we show that receiver plants may also become more susceptible to oviposition and thus more likely to be damaged. Extensive disruption of volatile‐mediated interactions by an atmospheric pollutant highlights the need to consider the pervading environment and changes therein when assessing their ecological significance.     

Con‐specific neighbours may enhance compensation capacity in an invasive plant

Facilitation, both by inter‐ and intra‐specific neighbours, is known to be an important process in structuring plant communities. However, only a small number of experiments have been reported on facilitation in plant invasions, especially between invasive con‐specific individuals. Here, we focus on how con‐specific neighbours of the invasive alien plant alligator weed affect the tolerance of alligator weed to herbivory by the introduced biological control agent, Agasicles hygrophila. We conducted greenhouse and garden experiments in which invasive plant density and herbivory intensity (artificial clipping and real herbivory) were manipulated. In the greenhouse experiment, artificial clipping significantly reduced plant biomass when plants were grown individually, but when con‐specific neighbours were present in the same pot, biomass was not significantly different from control plants. Similarly, when compared to control plants, plants that were subjected to herbivory by A. hygrophila produced more biomass when grown with two con‐specific neighbours than when grown alone. Real herbivory also resulted in an increased number of vegetative buds, and again when two con‐specific neighbours were present this effect was increased (a 55.3% increase in buds when there was no neighbour, but a 111.6% increase in buds when two con‐specific neighbours were present). In the garden experiment, in which plants were grown at high density (6 plants per pot), alligator weed fully recovered from defoliation caused by insects at levels from 20–30% to 100%. Our results indicate that the con‐specific association may increase the compensatory ability to cope with intense damage in this invasive plant.     

Facilitation and interference in subalpine meadows of the central Caucasus

Abstract. We studied the effects of neighbours on the biomass of seven randomly chosen species in species‐rich sub‐alpine meadows in the central Caucasus Mountains by comparing the performance of plants with neighbours removed experimentally to that of paired plants with their neighbours left intact. In most cases the removal of neighbours led to significant increases in vegetative and total above‐ground biomass implying the species were limited by competition. However, the neighbour removal led also to an increased leaf wilting for target plants, as well as to strong decline in reproductive effort for some species. We hypothesise that competition may be the prevailing type of interaction in species‐rich sub‐alpine meadow communities, but competitive effects on vegetative production may be balanced, if not outweighed, by facilitation, at least for some species. Such a balance may enhance species coexistence in communities.     

Interference among cotton neighbours after differential reproductive damage

In indeterminate plant species, the rate of vegetative growth usually declines during the stage of active reproductive growth. Fruit shedding, as induced by insect herbivores, could counteract this decline. Due to the relative increase in vegetative growth, plants that have suffered reproductive damage could be better able to intercept light and acquire soil resources than undamaged plants. If so, plants with damaged neighbours might grow less than their counterparts with smaller, undamaged neighbours. This hypothesis was tested in high- and low-density cotton crops subjected to three treatments: (i) undamaged controls; (ii) uniformly damaged, in which all plants were damaged; (iii) non-uniformly damaged, in which every second plant was damaged. Damaged plants had their flowerbuds and young fruits manually removed at 85 days after sowing to simulate shedding as induced by Helicoverpa spp. (Lepidoptera) and mirid bugs (Hemiptera). As expected, damaged plants had greater leaf area and more vegetative dry matter than undamaged ones. This was most pronounced at high plant density. Neighbour status did not affect vegetative growth but it had a substantial, asymmetric effect on the reproductive growth of target plants. Damaged targets recovered to the level of undamaged controls in terms of total fruit number but had a large reduction in the mass of mature fruit due to the limited time available for recovery. The effect of neighbour status, if any, on the production of mature fruit in damaged targets was overridden by the limit imposed to recovery by the duration of the growing season. In contrast, neighbour status affected the production of mature fruit of undamaged targets: undamaged targets with damaged neighbours had 34% (low density) and 56% (high density) less mature fruit mass than their counterparts with undamaged neighbours. This was because (i) reproductive allocation and (ii) the proportion of total fruit that reached maturity in target plants declined with increasing neighbour interference. Most studies dealing with changes in competitive relationships among plants subjected to differential herbivory have shown how undamaged plants may benefit from herbivores that feed on their neighbours. This study shows that differential reproductive damage can cause the opposite effect, as undamaged plants may have a significant reduction in productivity due to the influence of neighbours whose vegetative growth was stimulated by the loss of reproductive organs. Received: 2 June 1996 / Accepted: 8 September 1996     

Kin recognition affects plant communication and defence

The ability of many animals to recognize kin has allowed them to evolve diverse cooperative behaviours; such ability is less well studied for plants. Many plants, including Artemisia tridentata, have been found to respond to volatile cues emitted by experimentally wounded neighbours to increase levels of resistance to herbivory. We report that this communication was more effective among A. tridentata plants that were more closely related based on microsatellite markers. Plants in the field that received cues from experimentally clipped close relatives experienced less leaf herbivory over the growing season than those that received cues from clipped neighbours that were more distantly related. These results indicate that plants can respond differently to cues from kin, making it less likely that emitters will aid strangers and making it more likely that receivers will respond to cues from relatives. More effective defence adds to a growing list of favourable consequences of kin recognition for plants.     

Plant water stress and previous herbivore damage affect insect performance

1. Short‐term changes in plant resistance traits can be affected by abiotic factors or damage by herbivores, although how the combined effects of abiotic factors and previous damage affect subsequent insect larval development is not well understood. 2. Complementary glasshouse and field experiments were conducted to evaluate whether plant water stress and previous herbivore damage influenced monarch (Danaus plexippus) larval development on common milkweed, Asclepias syriaca. 3. In the glasshouse, water stress altered a suite of A. syriaca functional traits but did not affect nutrient content, whereas herbivore damage increased leaf nitrogen (N) and reduced the carbon:nitrogen (C:N) ratio. A bioassay experiment showed that monarch larval survival was lower on well‐watered plants that were previously damaged by monarch larva than on damaged and drought‐stressed plants. Bioassay larvae consumed less leaf tissue of previously damaged plants, whereas monarch larval mass was affected additively by water stress and previous damage, after correcting for the amount of leaf tissue consumed. 4. In a 2‐year field experiment, monarch larval performance was higher on previously damaged A. syriaca plants that received experimentally reduced rainfall, relative to plants receiving ambient rainfall. 5. Collectively, these results from glasshouse and field experiments suggest that insect performance was highest on previously damaged plants under water stress and highlight the additive and interactive roles of abiotic and biotic factors on herbivore performance.     

The jasmonate pathway alters herbivore feeding behaviour: consequences for plant defences

The jasmonate pathway is a highly conserved defensive cascade in plants that regulates the induction of resistance against herbivores; however, its role in herbivore feeding behaviour remains unknown. We used a mutant tomato plant (def‐1) deficient in the production of jasmonate‐related defensive proteins to test the hypothesis that genotypes with a reduced ability to induce resistance have a higher and more concentrated pattern of herbivore damage. Wild‐type and def‐1 plants received either damage by Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) caterpillars or no damage. After treatment, we tested for systemic responses by allowing a free roaming S. exigua caterpillar to feed on the undamaged portions of plants. Weight‐gain and leaf consumption of S. exigua were highest on def‐1 plants, regardless of prior herbivore damage. Def‐1 plants also had fewer numbers of leaves and leaflets eaten, and fewer feeding holes, which was indicative of a more concentrated distribution of damage on mutant compared to wild‐type plants. Following these results, we mimicked the amount and distribution of feeding damage that wild‐type or jasmonate‐deficient plants would receive on wild‐type plants to test whether changes in feeding behaviour may feedback to influence the expression of induced resistance. We mimicked the distribution of damage in wild‐type and jasmonate‐deficient plants by allowing caterpillars to feed on either one (leaf 1 or 2) or two leaves (leaf 1 and 2). Increased herbivore damage resulted in higher proteinase inhibitor (PI) activity, a jasmonate‐regulated defensive protein, and lower S. exigua performance on wild‐type but not jasmonate‐deficient plants. Compared to undamaged plants, a concentrated pattern of herbivore damage increased systemic resistance; these induced responses were greater on leaflets with stronger vascular connections to the damaged leaf. A more dispersed pattern of caterpillar damage altered the expression of induced responses, but the outcome depended on the specific pattern of damage. When leaf 1 was damaged and then leaf 2, the undamaged (third) leaf (which is more strongly connected to leaf 1 than 2) expressed reduced the PI activity compared to plants receiving concentrated damage to leaf 1; whereas in plants where leaf 2 was first damaged and then leaf 1, there were no differences in PI activity in leaf 3 compared to plants receiving concentrated damage to leaf 2. Thus, induction of the jasmonate pathway may not only determine the amount and distribution of feeding damage by herbivores, but this may feedback to affect the subsequent expression of plant defence.     

Self-recognition affects plant communication and defense

Animals have the ability to distinguish self from non-self, which has allowed them to evolve immune systems and, in some instances, to act preferentially towards individuals that are genetically identical or related. Self-recognition is less well known for plants, although recent work indicates that physically connected roots recognize self and reduce competitive interactions. Sagebrush uses volatile cues emitted by clipped branches of self or different neighbours to increase resistance to herbivory. Here, we show that plants that received volatile cues from genetically identical cuttings accumulated less natural damage than plants that received cues from non-self cuttings. Volatile communication is required to coordinate systemic processes such as induced resistance and plants respond more effectively to self than non-self cues. This self/non-self discrimination did not require physical contact and is a necessary first step towards possible kin recognition and kin selection.     

Bt rice could provide ecological resistance against nontarget planthoppers

Genetically engineered (GE) rice lines expressing Lepidoptera‐active insecticidal cry genes from the bacterium Bacillus thuringiensis (Bt) have been developed in China. Field surveys indicated that Bt rice harbours fewer rice planthoppers than non‐Bt rice although planthoppers are not sensitive to the produced Bt Cry proteins. The mechanisms underlying this phenomenon remain unknown. Here, we show that the low numbers of planthoppers on Bt rice are associated with reduced caterpillar damage. In laboratory and field‐cage experiments, the rice planthopper Nilapavata lugens had no feeding preference for undamaged Bt or non‐Bt plants but exhibited a strong preference for caterpillar‐damaged plants whether Bt or non‐Bt. Under open‐field conditions, rice planthoppers were more abundant on caterpillar‐damaged non‐Bt rice than on neighbouring healthy Bt rice. GC–MS analyses showed that caterpillar damage induced the release of rice plant volatiles known to be attractive to planthoppers, and metabolome analyses revealed increased amino acid contents and reduced sterol contents known to benefit planthopper development. That Lepidoptera‐resistant Bt rice is less attractive to this important nontarget pest in the field is therefore a first example of ecological resistance of Bt plants to nontarget pests. Our findings suggest that non‐Bt rice refuges established for delaying the development of Bt resistance may also act as a trap crop for N. lugens and possibly other planthoppers.     

Communication between sagebrush and wild tobacco in the field

Communication between plants has not been widely accepted by most ecologists. However, recent field experiments indicated that wild tobacco plants became more resistant to herbivores when grown in close proximity to clipped sagebrush neighbors. Tobacco plants grown within 15 cm of sagebrush that had been either manually clipped with scissors or damaged by herbivores experienced less naturally occurring folivory than tobacco plants with unclipped neighbors. These results were consistent over five field seasons and involved treatments that were randomly assigned and well replicated. Associated with lower levels of herbivory were increased activities of polyphenol oxidase in tobacco foliage near clipped sagebrush neighbors. Experiments that blocked either air or soil contact between sagebrush and tobacco indicated that the communication was airborne rather than soilborne. Alternative explanations involving altered microenvironmental conditions or avoidance of clipped sagebrush by herbivores were not supported.Much remains to be learned about the natural history of this phenomenon. Apparently the plants must be in close proximity for communication to occur. Preliminary results suggest that communication between sagebrush and other plants may also occur. The mechanisms of communication as well as its ecological and evolutionary significance remain unknown.     

Do plants tap SOS signals from their infested neighbours?

Ecologist have not been able to show unambigous evidence for the involvement of plant-to-plant signal transfer in the defence strategies of plants. However, phytopathologists and plant physiologists recently demonstrated that resistance in undamaged plants can be elicited by volatiles of plant origin. Now that empirical evidence is accumulating, there is every reason to ask why plants use the available information on the infestion status of their neighbours and to assess the fitness advantages associated with the tuning of their defence. The debate on the ecological and evolutionary significance of interplant communication needs to be revived.     

Effects of single and multiple herbivory by host and non‐host caterpillars on the attractiveness of herbivore‐induced volatiles of sugarcane to the generalist parasitoid Cotesia flavipes

It is well known that parasitoids are attracted to volatiles emitted by host‐damaged plants; however, this tritrophic interaction may change if plants are attacked by more than one herbivore species. The larval parasitoid Cotesia flavipesCameron (Hymenoptera: Braconidae) has been used intensively in Brazil to control the sugarcane borer, Diatraea saccharalisFabricius (Lepidoptera: Pyralidae) in sugarcane crops, where Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), a non‐stemborer lepidopteran, is also a pest. Here, we investigated the ability of C. flavipes to discriminate between an unsuitable host (S. frugiperda) and a suitable host (D. saccharalis) based on herbivore‐induced plant volatiles (HIPVs) emitted by sugarcane, and whether multiple herbivory (D. saccharalis feeding on stalk + S. frugiperda feeding on leaves) in sugarcane affected the attractiveness of HIPVs to C. flavipes. Olfactometer assays indicated that volatiles of host and non‐host‐damaged plants were attractive to C. flavipes. Even though host‐ and non‐host‐damaged plants emitted considerably different volatile blends, neither naïve nor experienced wasps discriminated suitable and unsuitable hosts by means of HIPVs emitted by sugarcane. With regard to multiple herbivory, wasps innately preferred the odor blend emitted by sugarcane upon non‐host + host herbivory over host‐only damaged plants. Multiple herbivory caused a suppression of some volatiles relative to non‐host‐damaged sugarcane that may have resulted from the unaltered levels of jasmonic acid in host‐damaged plants, or from reduced palatability of host‐damaged plants to S. frugiperda. In conclusion, our study showed that C. flavipes responds to a wide range of plant volatile blends, and does not discriminate host from non‐host and non‐stemborer caterpillars based on HIPVs emitted from sugarcane. Moreover, we showed that multiple herbivory by the sugarcane borer and fall armyworm increases the attractiveness of sugarcane plants to the parasitoids.     

Neighbour tolerance,not suppression,provides competitive advantage to non‐native plants

High competitive ability has often been invoked as a key determinant of invasion success and ecological impacts of non‐native plants. Yet our understanding of the strategies that non‐natives use to gain competitive dominance remains limited. Particularly, it remains unknown whether the two non‐mutually exclusive competitive strategies, neighbour suppression and neighbour tolerance, are equally important for the competitive advantage of non‐native plants. Here, we analyse data from 192 peer‐reviewed studies on pairwise plant competition within a Bayesian multilevel meta‐analytic framework and show that non‐native plants outperform their native counterparts due to high tolerance of competition, as opposed to strong suppressive ability. Competitive tolerance ability of non‐native plants was driven by neighbour's origin and was expressed in response to a heterospecific native but not heterospecific non‐native neighbour. In contrast to natives, non‐native species were not more suppressed by hetero‐ vs. conspecific neighbours, which was partially due to higher intensity of intraspecific competition among non‐natives. Heterogeneity in the data was primarily associated with methodological differences among studies and not with phylogenetic relatedness among species. Altogether, our synthesis demonstrates that non‐native plants are competitively distinct from native plants and challenges the common notion that neighbour suppression is the primary strategy for plant invasion success.     

Plant responsiveness to root-root communication of stress cues

Background and Aims

Phenotypic plasticity is based on the organism''s ability to perceive, integrate and respond to multiple signals and cues informative of environmental opportunities and perils. A growing body of evidence demonstrates that plants are able to adapt to imminent threats by perceiving cues emitted from their damaged neighbours. Here, the hypothesis was tested that unstressed plants are able to perceive and respond to stress cues emitted from their drought- and osmotically stressed neighbours and to induce stress responses in additional unstressed plants.

Methods

Split-root Pisum sativum, Cynodon dactylon, Digitaria sanguinalis and Stenotaphrum secundatum plants were subjected to osmotic stress or drought while sharing one of their rooting volumes with an unstressed neighbour, which in turn shared its other rooting volume with additional unstressed neighbours. Following the kinetics of stomatal aperture allowed testing for stress responses in both the stressed plants and their unstressed neighbours.

Key Results

In both P. sativum plants and the three wild clonal grasses, infliction of osmotic stress or drought caused stomatal closure in both the stressed plants and in their unstressed neighbours. While both continuous osmotic stress and drought induced prolonged stomatal closure and limited acclimation in stressed plants, their unstressed neighbours habituated to the stress cues and opened their stomata 3–24 h after the beginning of stress induction.

Conclusions

The results demonstrate a novel type of plant communication, by which plants might be able to increase their readiness to probable future osmotic and drought stresses. Further work is underway to decipher the identity and mode of operation of the involved communication vectors and to assess the potential ecological costs and benefits of emitting and perceiving drought and osmotic stress cues under various ecological scenarios.     

Silicon-augmented resistance of plants to herbivorous insects: a review

Silicon (Si) is one of the most abundant elements in the earth's crust, although its essentiality in plant growth is not clearly established. However, the importance of Si as an element that is particularly beneficial for plants under a range of abiotic and biotic stresses is now beyond doubt. This paper reviews progress in exploring the benefits at two‐ and three‐trophic levels and the underlying mechanism of Si in enhancing the resistance of host plants to herbivorous insects. Numerous studies have shown an enhanced resistance of plants to insect herbivores including folivores, borers, and phloem and xylem feeders. Silicon may act directly on insect herbivores leading to a reduction in insect performance and plant damage. Various indirect effects may also be caused, for example, by delaying herbivore establishment and thus an increased chance of exposure to natural enemies, adverse weather events or control measures that target exposed insects. A further indirect effect of Si may be to increase tolerance of plants to abiotic stresses, notably water stress, which can in turn lead to a reduction in insect numbers and plant damage. There are two mechanisms by which Si is likely to increase resistance to herbivore feeding. Increased physical resistance (constitutive), based on solid amorphous silica, has long been considered the major mechanism of Si‐mediated defences of plants, although there is recent evidence for induced physical defence. Physical resistance involves reduced digestibility and/or increased hardness and abrasiveness of plant tissues because of silica deposition, mainly as opaline phytoliths, in various tissues, including epidermal silica cells. Further, there is now evidence that soluble Si is involved in induced chemical defences to insect herbivore attack through the enhanced production of defensive enzymes or possibly the enhanced release of plant volatiles. However, only two studies have tested for the effect of Si on an insect herbivore and third trophic level effects on the herbivore's predators and parasitoids. One study showed no effect of Si on natural enemies, but the methods used were not favourable for the detection of semiochemical‐mediated effects. Work recently commenced in Australia is methodologically and conceptually more advanced and an effect of Si on the plants' ability to generate an induced response by acting at the third trophic level was observed. This paper provides the first overview of Si in insect herbivore resistance studies, and highlights novel, recent hypotheses and findings in this area of research. Finally, we make suggestions for future research efforts in the use of Si to enhance plant resistance to insect herbivores.     

Characterization of alfalfa germplasm expressing resistance to silverleaf whitefly, Bemisia argentifolii

Abstract: Thirty‐eight plants were taken from a University of California alfalfa selection nursery for developing resistance to silverleaf whitefly, Bemisia argentifolii Bellows & Perring. Seventeen of the plants had low whitefly infestation and were categorized as ‘potentially resistant’; 21 of the plants had high whitefly infestation and were categorized as ‘presumed susceptible’. Plants were propagated vegetatively so that replicated measurements of whitefly performance could be made on each genotype. Two colonies of silverleaf whiteflies were used: one reared on alfalfa (alfalfa‐experienced whiteflies), and the other on cotton (alfalfa‐naive whiteflies). The effect of variation among alfalfa genotypes on whitefly performance was similar for both whitefly sources, although on all genotypes, the alfalfa‐experienced whiteflies generally performed better than their alfalfa‐naive counterparts. In greenhouse tests, fecundity of newly eclosed adults (over a 5‐day period) on the 17 potentially resistant genotypes was relatively consistent in being lower than fecundity on the presumed susceptible genotypes. However, in nymphal survival tests, the response on the 17 potentially resistant genotypes was not consistent. Nymphal survival (egg to adult) on some of these was very low, as expected, while nymphal survival on others was as high as on the presumed susceptible genotypes. Fecundity and nymphal survival data were not correlated for alfalfa‐naive whiteflies, and were only weakly correlated (r² = 0.13, d.f. = 32, P = 0.04) for alfalfa‐experienced whiteflies. Thirteen genotypes then were examined in the greenhouse in stage‐specific survival tests, where four genotypes demonstrated high resistance (<10% nymphal survival) and three demonstrated moderate resistance (11–34% survival) compared with the three presumed susceptible genotypes that were tested (51–73% survival). Most of the mortality on the resistant genotypes occurred in the first instar, while mortality was more evenly distributed across the life stages on the susceptible genotypes. Interestingly, if nymphs survived to second instar on the resistant genotypes, then their subsequent survival to adult eclosion was similar to survival of second instar to adult on susceptible genotypes. Six of the genotypes used in the greenhouse stage‐specific survival test also were evaluated in the field for nymphal survival, and these results were consistent with the greenhouse tests.     

Prior flea beetle herbivory affects oviposition preference and larval performance of a potato beetle on their shared host plant

Abstract 1. The herbaceous plant Solanum carolinense (L.) (Solanaceae) is host to a number of specialist insects, including the leaf-feeding beetles Epitrix fuscula (Crotch) and Leptinotarsa juncta (Germar) (Coleoptera: Chrysomelidae). Potted individuals of S. carolinense were subjected to one of two treatments: exposure to herbivory by E. fuscula or exclusion of all herbivores. The effects of E. fuscula herbivory on larval performance and oviposition preference of L. juncta were investigated.
2. Although the masses of the L. juncta pupae did not differ between the two treatments, larvae feeding on damaged plants developed more slowly than those feeding on undamaged plants.
3. In both paired leaf choice trials and whole plant choice trials, larvae of L. juncta showed no preference for undamaged versus damaged hosts.
4. In a field transplant experiment, adult L. juncta females showed slight feeding preferences and strong oviposition preferences for undamaged plants versus plants that had been fed on by E. fuscula .
5. The results are discussed with reference to their implications for plant-mediated competition among herbivores and constraints on the evolution of plant resistance.     

Plant age, communication, and resistance to herbivores: young sagebrush plants are better emitters and receivers

Plants progress through a series of distinct stages during development, although the role of plant ontogeny in their defenses against herbivores is poorly understood. Recent work indicates that many plants activate systemic induced resistance after herbivore attack, although the relationship between resistance and ontogeny has not been a focus of this work. In addition, for sagebrush and a few other species, individuals near neighbors that experience simulated herbivory become more resistant to subsequent attack. Volatile, airborne cues are required for both systemic induced resistance among branches and for communication among individuals. We conducted experiments in stands of sagebrush of mixed ages to determine effects of plant age on volatile signaling between branches and individuals. Young and old control plants did not differ in levels of chewing damage that they experienced. Systemic induced resistance among branches was only observed for young plants. Young plants showed strong evidence of systemic resistance only if airflow was permitted among branches; plants with only vascular connections showed no systemic resistance. We also found evidence for volatile communication between individuals. For airborne communication, young plants were more effective emitters of cues as well as more responsive receivers of volatile cues.     

Damage-induced changes in woody plants and their effects on insect herbivore performance: a meta-analysis

We conducted a meta‐analysis of 68 studies published between 1982 and 2000 in which the responses of woody plants to natural or simulated herbivore damage and/or insect herbivore performance on control and damaged plants were measured. Cumulative meta‐analyses revealed dramatic temporal changes in the magnitude and direction of the plant and herbivore responses reported during the last two decades. Studies conducted in the 1980s reported increase in phenolic concentrations, reduction in nutrient concentrations and negative effect on herbivore performance, consistently with the idea of induced resistance. In contrast, in the early 1990s when the idea that some types of plant damage may result in induced susceptibility was generally accepted, studies reported non‐significant results or induced susceptibility, and smaller effects on herbivores. The above changes may reflect paradigm shifts in the theory of induced defenses and/or the differences between study systems used in the early and the more recent studies. Overall, plant growth and carbohydrate concentrations were reduced in damaged plants despite enhanced photosynthetic rates. Damage increased the concentrations of carbon and phenolics, while terpene concentrations tended to decrease after damage; changes in nutrient concentrations after damage varied according to nutrient mobility, inherent plant growth rate, ontogenetic stage and plant type (deciduous/evergreen). Early season damage caused more pronounced changes in plants than late season damage, which is in accordance with the assumption that vigorously growing foliage has a greater capacity to respond to damage. Insect growth rate and female pupal weight decreased on previously damaged plants, while herbivore survival, consumption and male pupal weight were not significantly affected. The magnitude and direction of herbivore responses depended on the type of plant, the type of damage, the time interval between the damage and insect feeding (rapid/delayed induced resistance), and the timing of the damage.     

The timing of induced resistance and induced susceptibility in the soybean-Mexican bean beetle system

Induced plant responses to herbivory have been demonstrated in many systems. It has been suggested that the timing of these responses may influence the impact of induced resistance on herbivore populations, and may affect the evolution of induced defenses. This study used a bioassay to characterize the time course of systemic induced responses to Mexican bean beetle herbivory in four genotypes of soybeans. The results suggest that the time course of induced responses in this system is more complex than most previous studies have indicated. Herbivory provoked both rapid induced resistance and subsequent induced susceptibility to beetle feeding. All four genotypes of soybean induced significant resistance to beetle damage (beetles preferred undamaged to damaged plants) by 3 days after damage. By 15 days after damage, this resistance had decayed (beetles showed no preference for undamaged over damaged plants), and by 20 days after damage, all four genotypes exhibited significant induced susceptibility (beetles preferred previously damaged plants over undamaged plants). The magnitude of induced resistance in each genotype correlated strongly with the magnitude of induced susceptibility in that genotype. Received: 28 September 1997 / Accepted: 1 December 1997