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      Silicon Supplementation of Rescuegrass Reduces Herbivory by a Grasshopper

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          Abstract

          The theory of coevolution suggests that herbivores play an important role in the diversification and composition of plant communities. A prevalent idea holds that grasses and grazing animals participated in an evolutionary “arms race” as grassland ecosystems started spreading across the continents. In this race, besides other things, silicification in the form of phytoliths occurred in the grasses, and the graminivorous herbivores responded through specialized mandibles to feed on plants rich in phytoliths. It is important to understand whether these mandibles equip the herbivores in different environments or the grasses can augment their defense by channelizing their energy in high resource milieu. Here we used rescuegrass ( Bromus catharticus; Family: Poaceae), an alien species of South America, to understand the mechanism of resistance offered by this species against a local insect herbivore ( Oxya grandis; Family: Acrididae), graminivorous grasshopper, in different silicon-rich environments. We used different concentrations of silicon and observed the types of phytoliths formed after Si amendments and studied the effect of phytoliths on mandible wear of the grasshopper. Silicon concentrations increased ca. 12 fold in the highest supplementation treatments. The results reveal that higher foliar silica concentration in Si-rich plants did not result in changing the morphology of the phytoliths; still the leaf tissue consumption was lower in higher Si treatments, perhaps due to mandibular wear of the grasshoppers. The study opens a new dimension of investigating the role of Si amendments in reducing herbivory.

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          Phylogenetic variation in the silicon composition of plants.

          Silicon (Si) in plants provides structural support and improves tolerance to diseases, drought and metal toxicity. Shoot Si concentrations are generally considered to be greater in monocotyledonous than in non-monocot plant species. The phylogenetic variation in the shoot Si concentration of plants reported in the primary literature has been quantified. Studies were identified which reported Si concentrations in leaf or non-woody shoot tissues from at least two plant species growing in the same environment. Each study contained at least one species in common with another study. Meta-analysis of the data revealed that, in general, ferns, gymnosperms and angiosperms accumulated less Si in their shoots than non-vascular plant species and horsetails. Within angiosperms and ferns, differences in shoot Si concentration between species grouped by their higher-level phylogenetic position were identified. Within the angiosperms, species from the commelinoid monocot orders Poales and Arecales accumulated substantially more Si in their shoots than species from other monocot clades. A high shoot Si concentration is not a general feature of monocot species. Information on the phylogenetic variation in shoot Si concentration may provide useful palaeoecological and archaeological information, and inform studies of the biogeochemical cycling of Si and those of the molecular genetics of Si uptake and transport in plants.
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            International code for phytolith nomenclature 1.0.

            Phytoliths (microscopic opal silica particles produced in and between the cells of many plants) are a very resilient, often-preserved type of microfossil and today, phytolith analysis is widely used in palaeoenvironmental studies, botany, geology and archaeology. To date there has been little standardization in the way phytoliths are described and classified. This paper presents the first International Code for Phytolith Nomenclature (ICPN), proposing an easy to follow, internationally accepted protocol to describe and name phytoliths.
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              Physical defences wear you down: progressive and irreversible impacts of silica on insect herbivores.

              1. Silica in the leaves of grasses can act as a defence against both vertebrate and invertebrate herbivores. The mechanisms by which silica affects herbivore performance are not well characterized. Here we expose an insect herbivore Spodoptera exempta to high-silica diets and test two mechanisms by which silica has been proposed to act as a defence. First, that silica reduces the digestibility of leaves and second, that silica causes wear to insect mandibles, both of which could potentially impact on herbivore performance. 2. Silica reduced the efficiency with which S. exempta converted ingested food to body mass and the amount of nitrogen absorbed from their food, leading to reduced insect growth rates. The measure of how efficiently herbivores utilize digested food (ECD) was unaffected by silica. 3. These effects occurred even with short-term exposure to silica-rich diets, but they also increased markedly with the duration of exposure and affected late instars more than early instar larvae. This appears to be due to the progressive impacts of silica with longer exposure times and suggests that herbivores cannot adapt to silica defences, nor do they develop a tolerance for silica with age. 4. Exposure to silica-rich diets caused increased mandible wear in S. exempta. This effect was extremely rapid, occurring within a single instar, further reducing feeding efficiency and growth rates. These effects on insect growth and feeding efficiency are nonreversible, persisting after the herbivore has switched diets. Up to a third of this residual impact can be explained by the degree of mandible wear caused by previous silica-rich diets. 5. The impacts of silica on S. exempta larvae were progressive with exposure time and could not be compensated for, even by switching to a different diet. Thus, herbivores cannot easily adapt to physical defences such as silica, suggesting this defence will have major implications for herbivore fitness.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                24 May 2019
                2019
                : 10
                : 671
                Affiliations
                [1] 1 Department of Botany, University of Kashmir , Srinagar, India
                [2] 2 Division of Entomology, Sher-e-Kashmir University of Agricultural Sciences and Technology of Kashmir , Srinagar, India
                Author notes

                Edited by: Martin John Hodson, Oxford Brookes University, United Kingdom

                Reviewed by: Scott Nicholas Johnson, Western Sydney University, Australia; Paul-andré Calatayud, Institut de recherche pour le développement (IRD), France; Gerald Juma, University of Nairobi, Kenya

                *Correspondence: Irfan Rashid, ecoirfan@ 123456yahoo.co.in

                This article was submitted to Plant Physiology, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2019.00671
                6543128
                b0a897eb-a0c2-47b1-9d1d-e7b16b6a8172
                Copyright © 2019 Mir, Rashid, Hussain, Reshi, Assad and Sofi.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 20 December 2018
                : 03 May 2019
                Page count
                Figures: 6, Tables: 0, Equations: 0, References: 60, Pages: 8, Words: 5472
                Categories
                Plant Science
                Original Research

                Plant science & Botany
                herbivory,phytolith,grass,mandible wear,silicon
                Plant science & Botany
                herbivory, phytolith, grass, mandible wear, silicon

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