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Plant defense against herbivory

Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants can sense being touched, and they can use several strategies to defend against damage caused by herbivores. Many plants produce secondary metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores, or reduce plant digestibility. Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants can sense being touched, and they can use several strategies to defend against damage caused by herbivores. Many plants produce secondary metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores, or reduce plant digestibility. Other defensive strategies used by plants include escaping or avoiding herbivores in any time and/or any place, for example by growing in a location where plants are not easily found or accessed by herbivores, or by changing seasonal growth patterns. Another approach diverts herbivores toward eating non-essential parts, or enhances the ability of a plant to recover from the damage caused by herbivory. Some plants encourage the presence of natural enemies of herbivores, which in turn protect the plant. Each type of defense can be either constitutive (always present in the plant), or induced (produced in reaction to damage or stress caused by herbivores). Historically, insects have been the most significant herbivores, and the evolution of land plants is closely associated with the evolution of insects. While most plant defenses are directed against insects, other defenses have evolved that are aimed at vertebrate herbivores, such as birds and mammals. The study of plant defenses against herbivory is important, not only from an evolutionary view point, but also in the direct impact that these defenses have on agriculture, including human and livestock food sources; as beneficial 'biological control agents' in biological pest control programs; as well as in the search for plants of medical importance. The earliest land plants evolved from aquatic plants around 450 million years ago (Ma) in the Ordovician period. Many plants have adapted to iodine-deficient terrestrial environment by removing iodine from their metabolism, in fact iodine is essential only for animal cells. An important antiparasitic action is caused by the block of the transport of iodide of animal cells inhibiting sodium-iodide symporter (NIS). Many plant pesticides are glycosides (as the cardiac digitoxin) and cyanogenic glycosides which liberate cyanide, which, blocking cytochrome c oxidase and NIS, is poisonous only for a large part of parasites and herbivores and not for the plant cells in which it seems useful in seed dormancy phase. Iodide is not pesticide, but is oxidized, by vegetable peroxidase, to iodine, which is a strong oxidant, it is able to kill bacteria, fungi and protozoa. The Cretaceous period saw the appearance of more plant defense mechanisms. The diversification of flowering plants (angiosperms) at that time is associated with the sudden burst of speciation in insects. This diversification of insects represented a major selective force in plant evolution, and led to selection of plants that had defensive adaptations. Early insect herbivores were mandibulate and bit or chewed vegetation; but the evolution of vascular plants lead to the co-evolution of other forms of herbivory, such as sap-sucking, leaf mining, gall forming and nectar-feeding. The relative abundance of different species of plants in ecological communities including forests and grasslands may be determined in part by the level of defensive compounds in the different species. Since the cost of replacement of damaged leaves is higher in conditions where resources are scarce, it may also be that plants growing in areas where water and nutrients are scarce may invest more resources into anti-herbivore defenses. Our understanding of herbivory in geological time comes from three sources: fossilized plants, which may preserve evidence of defense (such as spines), or herbivory-related damage; the observation of plant debris in fossilised animal faeces; and the construction of herbivore mouthparts. Long thought to be a Mesozoic phenomenon, evidence for herbivory is found almost as soon as fossils which could show it. As previously discussed, the first land plants emerged around 450 million years ago; however, herbivory, and therefore the need for plant defenses, has undoubtedly been around for longer. Herbivory first evolved due to marine organisms within ancient lakes and oceans. Within under 20 million years of the first fossils of sporangia and stems towards the close of the Silurian, around 420 million years ago, there is evidence that they were being consumed. Animals fed on the spores of early Devonian plants, and the Rhynie chert also provides evidence that organisms fed on plants using a 'pierce and suck' technique. Many plants of this time are preserved with spine-like enations, which may have performed a defensive role before being co-opted to develop into leaves. During the ensuing 75 million years, plants evolved a range of more complex organs – from roots to seeds. There was a gap of 50 to 100 million years between each organ evolving, and it being fed upon. Hole feeding and skeletonization are recorded in the early Permian, with surface fluid feeding evolving by the end of that period.

[ "Gene", "Pathogen", "Jasmonic acid synthesis" ]
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