Papain protects papaya trees from herbivorous insects: role of cysteine proteases in latex

Many plants contain latex that exudes when leaves are damaged, and a number of proteins and enzymes have been found in it. The roles of those latex proteins and enzymes are as yet poorly understood. We found that papain, a cysteine protease in latex of the Papaya tree (Carica papaya, Caricaceae), is a crucial factor in the defense of the papaya tree against lepidopteran larvae such as oligophagous Samia ricini (Saturniidae) and two notorious polyphagous pests, Mamestra brassicae (Noctuidae) and Spodoptera litura (Noctuidae). Leaves of a number of laticiferous plants, including papaya and a wild fig, Ficus virgata (Moraceae), showed strong toxicity and growth inhibition against lepidopteran larvae, though no apparent toxic factors from these species have been reported. When the latex was washed off, the leaves of these lactiferous plants lost toxicity. Latexes of both papaya and the wild fig were rich in cysteine-protease activity. E-64, a cysteine protease-specific inhibitor, completely deprived the leaves of toxicity when painted on the surface of papaya and fig leaves. Cysteine proteases, such as papain, ficin, and bromelain, all showed toxicity. The results suggest that plant latex and the proteins in it, cysteine proteases in particular, provide plants with a general defense mechanism against herbivorous insects.     

Mechanisms of resistance of freshwater macrophytes to herbivory by invasive juvenile common carp

1. Herbivory on freshwater macrophytes has been assumed to be insignificant and rare. More recent evidence suggests herbivory is common and the impact of invasive invertebrate herbivores can be substantial. However, little is known about consumption of macrophytes by fish. 2. We performed a series of feeding assays, based on the consumption by common carp (Cyprinus carpio), to determine if any mechanisms of resistance, structural or chemical, were present in five species of macrophytes (Stuckenia pectinata, Typha latifolia, Scirpus validus, Chara aspera and Ceratophyllum demersum). 3. Carp consumed more fresh whole plant tissue of C. aspera than any other macrophyte, suggesting a lack of structural or chemical deterrents. Typha latifolia, S. validus and C. demersum were consumed least as whole plants, but consumption increased when they were offered in pellet form suggesting structural defence. Crude chemical extracts from S. pectinata significantly reduced consumption of pellets by carp. Thus, plant chemistry and structure both deterred feeding by carp. 4. Experiments that focus on theory are common but their application to managing landscapes is substantially lacking. Our results provide a basis for recommending plants to be used in restoring larval habitat refugia with the aim of increasing the probability for long‐term recovery of an endangered species. Thus, this paper is an example of how experiments that tie theory to application are important for practical applications and for continued testing of theory. 5. We suggest that macrophyte–herbivore interactions play an integral part in aquatic food webs and may be as important in freshwater communities as in marine and terrestrial systems.     

Resistance to pathogens and host developmental stage: a multifaceted relationship within the plant kingdom

The induction of resistance to disease during plant development is widespread in the plant kingdom. Resistance appears at different stages of host development, varies with plant age or tissue maturity, may be specific or broad-spectrum and is driven by diverse mechanisms, depending on plantpathogen interactions. Studies of these forms of resistance may help us to evaluate more exhaustively the plethora of levels of regulation during development, the variability of the defense potential of developing hosts and may have practical applications, making it possible to reduce pesticide applications. Here, we review the various types of developmental resistance in plants and current knowledge of the molecular and cellular processes involved in their expression. We discuss the implications of these studies, which provide new knowledge from the molecular to the agrosystem level.     

Structure of the phytophagous insect fauna of Onopordum thistles in the northern Mediterranean basin

Thistles of the genus Onopordum (Asteraceae: Cardueae) have become serious weed problems in parts of Australia and North America following introduction from their native Eurasian ranges, and are the target of a biological control programme in Australia. This paper analyses the results of three separate surveys of insects feeding and breeding on thistles of the genus Onopordum in the Mediterranean region. Of the 129 species found feeding, 74 species also bred on these thistles. Endophages comprised 54% of the breeding insects, with species that fed in the capitula and within the rosette crown/root area predominating. The Coleoptera and Hemiptera were the dominant endophage and ectophage taxa, respectively. Differences in the Onopordum fauna were noted between host species and between geographic regions, but this was largely due to the generalist component of the fauna. Onopordum specialists showed little variability across regions or between different species within the host genus. On comparing the fauna of Onopordum with that of other thistle genera, it was suggested that plant-herbivore interactions between Onopordum and its fauna are less highly evolved, in that there is less host specialization and a lesser degree of niche partitioning. This appears to be related to the low rate of speciation and habitat specialization within the host genus itself. The potential of certain genus-specific insects as biological control agents for Onopordum spp. is discussed. It is argued that the absence of a high degree of niche specialization could favour successful biological control, as the agent would be capable of inflicting damage over a wider range of resource gradients.     

Consequences of simultaneous elevation of carbon dioxide and temperature for plant–herbivore interactions: a metaanalysis

The effects of elevated carbon dioxide on plant–herbivore interactions have been summarized in a number of narrative reviews and metaanalyses, while accompanying elevation of temperature has not received sufficient attention. The goal of our study is to search, by means of metaanalysis, for a general pattern in responses of herbivores, and plant characteristics important for herbivores, to simultaneous experimental increase of carbon dioxide and temperature (ECET) in comparison with both ambient conditions and responses to elevated CO₂ (EC) and temperature (ET) applied separately. Our database includes 42 papers describing studies of 31 plant species and seven herbivore species. Nitrogen concentration and C/N ratio in plants decreased under both EC and ECET treatments, whereas ET had no significant effect. Concentrations of nonstructural carbohydrates and phenolics increased in EC, decreased in ET and did not change in ECET treatments, whereas terpenes did not respond to EC but increased in both ET and ECET; leaf toughness increased in both EC and ECET. Responses of defensive secondary compounds to treatments differed between woody and green tissues as well as between gymnosperm and angiosperm plants. Insect herbivore performance was adversely affected by EC, favoured by ET, and not modified by ECET. Our analysis allowed to distinguish three types of relationships between CO₂ and temperature elevation: (1) responses to EC do not depend on temperature (nitrogen, C/N, leaf toughness, phenolics in angiosperm leaves), (2) responses to EC are mitigated by ET (sugars and starch, terpenes in needles of gymnosperms, insect performance) and (3) effects emerge only under ECET (nitrogen in gymnosperms, and phenolics and terpenes in woody tissues). This result indicates that conclusions of CO₂ elevation studies cannot be directly extrapolated to a more realistic climate change scenario. The predicted negative effects of CO₂ elevation on herbivores are likely to be mitigated by temperature increase.     

Seasonal changes in leaf nutritional quality influence grass miner performance

1. The plant phenological age hypothesis predicts that phytophagous insects should prefer and perform better on phenologically young plants than on old plants because plant nutritional quality decreases with plant phenological age. This hypothesis was tested under field and laboratory conditions with the grass miner Chromatomyia milii on the free‐growth species Holcus lanatus. 2. The field experiment was conducted at four sample sites in Belgium where nutritional quality of H. lanatus leaves and performance of C. milii were monitored throughout the growth season. Foliar nutritional quality was highest early in the season due to high levels of proteins and soluble carbohydrates and low levels of lignin. Offspring performance (pupal size) decreased with plant phenological age, due at least partially to the decreasing foliar nutritional quality. 3. In the laboratory experiment, preference and performance of C. milii were determined on three different age classes of H. lanatus. Multiple‐choice experiments demonstrated that oviposition preference did not differ among age classes. Offspring survival decreased with plant phenological age, while pupal size did not differ among age classes. The relationship between foliar nutritional quality and plant phenological age was equivocal and did not correspond to the predictions of the plant phenological age hypothesis. 4. The results of the field experiment were consistent with the idea that the phenological age hypothesis holds in free‐growth species. The laboratory experiment gave only little support to the plant phenological age hypothesis. Possible causes for the differences between field and laboratory experiments are discussed.     

Weed biological control: applying science to solve seemingly intractable problems

Abstract   Exotic weeds pose a problem of considerable economic and environmental importance to Australia. As a consequence, Australia has developed into a leading centre of research on weed biological control, with over 60 weeds the targets of past and current projects. Using primarily entomological examples, this review highlights the contributions made by Australian scientists to the development of theory and the improvement of practice in weed biological control. It also shows how biological control practitioners have made use of, and contributed to, broader theory and knowledge of plant-herbivore relationships. Finally, it concludes with some reflections on the future direction of biological control in Australia.     

Age-related changes in defensive traits of Acacia tortilis Hayne

The theory of plant defences proposes that investments in physical and chemical defences are driven by the risk of herbivore damage, and limited by the cost of producing the particular defensive trait in terms of resources that could be directed to other sinks, such as growth and reproduction. We sampled twigs of 18 mature Acacia tortilis trees and their cohort of juveniles to test some predictions of this hypothesis. We expected a higher allocation of defensive traits to leaves and twigs in the young plants than in the mature ones as a result of a higher risk of damage by ungulates at the juvenile stage. Our results show that the juvenile plants produce more spines along their twigs, but have lower concentrations of phenolic compounds in their leaves than in the mature ones. We also expected a negative relation between the concentration of foliar nutrients and phenolic compounds, as predicted by the carbon/nutrient hypothesis. Only mature plants showed this pattern. Reproduction (in mature plants) and water stress (in juvenile plants) did not relate to allocation to secondary compounds as predicted by current hypotheses of plant defence.