Physiological and ecological constraints play key roles in the evolution of plant growth patterns, especially in relation to defenses against herbivores. Phenotypic and life history theories are unified within the growth-differentiation balance (GDB) framework, forming an integrated system of theories explaining and predicting patterns of plant defense and competitive interactions in ecological and evolutionary time. Plant activity at the cellular level can be classified as growth (cell division and enlargement) of differentiation (chemical and morphological changes leading to cell maturation and specialization). The GDB hypothesis of plant defense is premised upon a physiological trade-off between growth and differentiation processes. The trade-off between growth and defense exists because secondary metabolism and structural reinforcement are physiologically constrained in dividing and enlarging cells, and because they divert resources from the production of new leaf area. Hence the dilemma of plants: They must grow fast enough to compete, yet maintain the defenses necessary to survive in the presence of pathogens and herbivores. The physiological trade-off between growth and differentiation processes interacts with herbivory and plant-plant competition to manifest itself as a genetic trade-off between growth and defense in the evolution of plant life history strategies. Evolutionary theories of plant defense are reviewed. We also extend a standard growth rate model by separating its ecological and evolutionary components, andformalizing the role of competition in the evolution of plant defense. We conclude with a conceptual model of the evolution of plant defense in which plant physiological trade-offs interact with the abiotic environment, competition and herbivory. theory ofplant defense has not kept pace. The complexity of the subject has hindered the development of a synthetic framework, because any comprehensive treatment of plant defense theory must transcend both the disciplines and the levels of biological organiza
Since its accidental introduction from Asia, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has killed millions of ash trees in North America. As it continues to spread, it could functionally extirpate ash with devastating economic and ecological impacts. Little was known about EAB when it was first discovered in North America in 2002, but substantial advances in understanding of EAB biology, ecology, and management have occurred since. Ash species indigenous to China are generally resistant to EAB and may eventually provide resistance genes for introgression into North American species. EAB is characterized by stratified dispersal resulting from natural and human-assisted spread, and substantial effort has been devoted to the development of survey methods. Early eradication efforts were abandoned largely because of the difficulty of detecting and delineating infestations. Current management is focused on biological control, insecticide protection of high-value trees, and integrated efforts to slow ash mortality.
Urbanization affects communities of herbivorous arthropods and provides opportunities for dramatic changes in their abundance and richness. Underlying these changes are creation of impervious surfaces; variation in the density, diversity, and complexity of vegetation; and maintenance practices including pulsed inputs of fertilizers, water, and pesticides. A rich body of knowledge provides theoretical underpinnings for predicting and understanding impacts of urbanization on arthropods. However, relatively few studies have elucidated mechanisms that explain patterns of insect and mite abundance and diversity across urbanization gradients. Published accounts suggest that responses to urbanization are often taxon specific, highly variable, and linked to properties of urbanization that weaken top-down and/or bottom-up processes, thereby destabilizing populations of herbivores and their natural enemies. In addition to revealing patterns in diversity and abundance of herbivores across urbanization gradients, a primary objective of this review is to examine mechanisms underlying these patterns and to identify potential hypotheses for future testing.
Recent studies have investigated interspecific variation in resistance of ash (Fraxinus spp.) to the exotic wood-boring beetle, emerald ash borer (EAB, Agrilus planipennis). Manchurian ash (Fraxinus mandshurica) is an Asian species that has coevolved with EAB. It experiences little EAB-induced mortality compared to North American ashes. Host phloem chemistry, both constitutive and induced, might partly explain this interspecific variation in resistance. We analyzed the constitutive phloem chemistry of three ash species: Manchurian ash and North American white (Fraxinus americana) and green (Fraxinus pennsylvanica) ash. Analysis of the crude phloem extracts revealed the presence of an array of phenolic compounds including hydroxycoumarins, a monolignol, lignans, phenylethanoids, and secoiridoids. Both qualitative and quantitative differences were observed among the three ash species. Hydroxycoumarins and the phenylethanoids, calceloariosides A and B, were present only in the phloem of Manchurian ash and might represent a mechanism of resistance against EAB.
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