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In addition to physical defenses, chemical defenses are the most effective way to protect plants from adverse biotic factors (phytophagous insects, other herbivores and pathogenic fungi). This requires extra effort from plants to produce secondary defense metabolites at the expense of production of primary metabolites directly linked to the growth and development of plants. There are three main groups of defensive compounds (alkaloids, phenolic compounds and terpenoids). All defensive compounds are rich in carbon. Depending on the chemical formula, carbon makes up from about 40% to over 85% of the molecular weight of various defense compounds. It is not possible to calculate the total carbon mass accumulation in all defense compounds. In this paper we discuss the content of defensive compounds and carbon with respect to defense strategy of plants, functional groups of woody species (coniferous and deciduous trees), tree species, tree biomass components (leaves, branches, bark, roots, etc.) and many other internal (age of trees, age of leaves, stage of development, origin, etc.) as well as external factors, related to soil and climatic conditions
Artificial defoliation is often used in studies of induced defense reactions of plants to damage by arthropods. However, little is known about the role of various external factors that may determine the nature of inducible defenses. Here we tested whether light level under which plants grow could affect the natural invasion of two-spotted spider mite (Tetranychus urticae Koch) with a broad range of host plants. For this study we used two host species that differ widely in shade tolerance: shade-adapted Norway maple (Acer platanoides L.) and sun-adapted European mountain ash (Sorbus aucuparia L.). The purpose of this study was to explore mite invasion and to test whether prior simulated defoliation, light conditions and host species differences in secondary defense metabolites (soluble phenolics) play a role in the pattern of invasion. One-year-old maple and ash seedlings growing in pots were placed into two shade houses that produced a treatment with 5% light transmittance, and the second group of seedlings was placed in full sunlight. The experiment was carried out in a location chronically affected by spider mite presence. The defoliation was performed in mid-May by manual removal of 0 (control), 25, 50, 75 and 100% of leaves. Natural two-spotted spider mite invasion took place in mid-July and was observed only on seedlings growing in the shade. Mites were found on leaves of almost all seedlings. However, the intensity of feeding damage to leaves was related to the level of earlier defoliation. Control and slightly to moderately (25–50%) defoliated seedlings of both species were most affected by spider mites. Mite feeding behavior was directly linked to changes in concentration of leaf carbon and phenolic compounds. Carbon content was positively correlated with the percent of seedlings damaged by spider mites in both tree species (R² > 0.80; P <0.05). The highest concentration of phenolic compounds was observed in leaves of seedlings subjected to 50 and 75% defoliation. Our results indicated that two-spotted spider mites preferred plants grown in a shade environment and within these plants favored leaves which are high in carbon and low in phenolic compounds. Both plant species were able to survive early season leaf damage inflicted by spider mites. Inducible phenolic compounds were among defensive secondary metabolites contributing to low spider mite success in plants recovering from artificial spring defoliation.
Carbon assimilated during photosynthesis is allocated to basic needs, such as growth, defense, and storage of nutrients. The aim of this study was to explore potential relationships between carbon allocation and light conditions during growth, including shade tolerance of species. We studied species that represent light-demanding (Sorbus aucuparia, Betula pendula), intermediate (Carpinus betulus, Quercus robur), and shade-tolerant (Acer platanoides, Fagus sylvatica) trees. We exposed seedlings to two light treatments (full sunlight and shade), and explored how these conditions affect plant growth and biomass allocation, as well as the levels of phenolic compounds, nonstructural carbohydrates, carbon, and nitrogen. We hypothesized that light-demanding species invest less carbon in chemical defenses against pathogens and/or herbivores compared to shade-tolerant species. On the other hand, light-demanding species showed the greater part of assimilated carbon allocate to growth processes. As a result, the stem diameter above the root collar, the mass of leaves, stems, coarse, and fine roots were larger under full-sunlight conditions in all species, except for greater height of A. platanoides and Q. robur under shade conditions. Leaves from full light were characterized by lower nitrogen content, higher carbon and phenolic contents, and a higher carbon/nitrogen ratio compared with leaves from seedlings grown in shade. In the case of shade-tolerant species, a trade-off mechanism can be proposed that such species restrict their usual allocation of carbon to defense and radial growth, while instead of investing it in increasing their heights and storage capacities. According to the light-demanding species, it was not possible to identify a trade-off mechanism and how carbon allocationis restricted upon exposure to shade conditions, except for the reduced allocation to the root mass.
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