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      Karrikins Identified in Biochars Indicate Post-Fire Chemical Cues Can Influence Community Diversity and Plant Development

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          Abstract

          Background

          Karrikins are smoke-derived compounds that provide strong chemical cues to stimulate seed germination and seedling growth. The recent discovery in Arabidopsis that the karrikin perception system may be present throughout angiosperms implies a fundamental plant function. Here, we identify the most potent karrikin, karrikinolide (KAR 1), in biochars and determine its role in species unique plant responses.

          Methods

          Biochars were prepared by three distinct commercial-scale pyrolysis technologies using systematically selected source material and their chemical properties, including karrikinolide, were quantified. Dose-response assays determined the effects of biochar on seed germination for two model species that require karrikinolide to break dormancy ( Solanum orbiculatum, Brassica tourneforttii) and on seedling growth using two species that display plasticity to karrikins, biochar and phytotoxins ( Lactuca sativa, Lycopersicon esculentum). Multivariate analysis examined relationships between biochar properties and the plant phenotype.

          Findings and Conclusions

          Results showed that karrikin abundant biochars stimulated dormant seed germination and seedling growth via mechanisms analogous to post-fire chemical cues. The individual species response was associated with its sensitivity to karrikinolide and inhibitory compounds within the biochars. These findings are critical for understanding why biochar influences community composition and plant physiology uniquely for different species and reaffirms that future pyrolysis technologies promise by-products that concomitantly sequester carbon and enhance plant growth for ecological and broader plant related applications.

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          Most cited references20

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          A handful of carbon.

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            A review of biochars' potential role in the remediation, revegetation and restoration of contaminated soils.

            Biochars are biological residues combusted under low oxygen conditions, resulting in a porous, low density carbon rich material. Their large surface areas and cation exchange capacities, determined to a large extent by source materials and pyrolysis temperatures, enables enhanced sorption of both organic and inorganic contaminants to their surfaces, reducing pollutant mobility when amending contaminated soils. Liming effects or release of carbon into soil solution may increase arsenic mobility, whilst low capital but enhanced retention of plant nutrients can restrict revegetation on degraded soils amended only with biochars; the combination of composts, manures and other amendments with biochars could be their most effective deployment to soils requiring stabilisation by revegetation. Specific mechanisms of contaminant-biochar retention and release over time and the environmental impact of biochar amendments on soil organisms remain somewhat unclear but must be investigated to ensure that the management of environmental pollution coincides with ecological sustainability. Copyright © 2011 Elsevier Ltd. All rights reserved.
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              Biochar: a synthesis of its agronomic impact beyond carbon sequestration.

              Biochar has been heralded as an amendment to revitalize degraded soils, improve soil carbon sequestration, increase agronomic productivity, and enter into future carbon trading markets. However, scientific and economic technicalties may limit the ability of biochar to consistently deliver on these expectations. Past research has demonstrated that biochar is part of the black carbon continuum with variable properties due to the net result of production (e.g., feedstock and pyrolysis conditions) and postproduction factors (storage or activation). Therefore, biochar is not a single entity but rather spans a wide range of black carbon forms. Biochar is black carbon, but not all black carbon is biochar. Agronomic benefits arising from biochar additions to degraded soils have been emphasized, but negligible and negative agronomic effects have also been reported. Fifty percent of the reviewed studies reported yield increases after black carbon or biochar additions, with the remainder of the studies reporting alarming decreases to no significant differences. Hardwood biochar (black carbon) produced by traditional methods (kilns or soil pits) possessed the most consistent yield increases when added to soils. The universality of this conclusion requires further evaluation due to the highly skewed feedstock preferences within existing studies. With global population expanding while the amount of arable land remains limited, restoring soil quality to nonproductive soils could be key to meeting future global food production, food security, and energy supplies; biochar may play a role in this endeavor. Biochar economics are often marginally viable and are tightly tied to the assumed duration of agronomic benefits. Further research is needed to determine the conditions under which biochar can provide economic and agronomic benefits and to elucidate the fundamental mechanisms responsible for these benefits. Copyright © by the American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Inc.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                18 August 2016
                2016
                : 11
                : 8
                : e0161234
                Affiliations
                [1 ]School of Agriculture and Food Sciences, University of Queensland, Brisbane, Queensland, Australia
                [2 ]School of Chemistry and Biochemistry, University of Western Australia, Crawley, Western Australia, Australia
                [3 ]School of Agriculture and Food Sciences, University of Queensland, Gatton, Queensland, Australia
                University of California Davis, UNITED STATES
                Author notes

                Competing Interests: A Fellowship for JK was financially supported by Horticulture Innovation Australia Ltd** using voluntary contributions from Brisbane City Council and matched funds from the Australian Government (HG10025, MT13042, http://horticulture.com.au/) and a University of Queensland ECR Grant ( https://www.uq.edu.au/). An ARC Future Fellowship for GRF was financially supported by the Australian Research Council (FT110100304, http://www.arc.gov.au/). **Horticulture Innovation Australia Limited (Hort Innovation) is not considered a commercial source. It is a grower owned, not-for-profit Rural Research and Development Corporation that invests levy, tax-payer and co-investment funds in research, development, extension and marketing projects. Hort Innovation is a provider of project and program management and communications services for these investments with no commercial benefit. Any outputs developed through funded projects that are deemed of commercial value are licensed and sold through third party commercial partners. This does not alter the authors' adherence to PLOS ONE policies on sharing data and materials.

                • Conceptualization: JK RLL ATL GRF.

                • Data curation: JK GRF.

                • Formal analysis: JK ATL GRF.

                • Funding acquisition: JK GRF.

                • Investigation: JK RLL GRF.

                • Methodology: JK RLL ATL GRF.

                • Project administration: JK GRF.

                • Resources: JK GRF.

                • Supervision: JK GRF.

                • Validation: JK RLL ATL GRF.

                • Visualization: JK RLL ATL GRF.

                • Writing - original draft: JK GRF.

                • Writing - review & editing: JK RLL ATL GRF.

                ‡ These authors also contributed equally to this work and are Joint Senior Authors.

                Article
                PONE-D-16-16938
                10.1371/journal.pone.0161234
                4990347
                27536995
                3bd10bde-c2ed-463e-b7c7-d158724bde32
                © 2016 Kochanek et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 26 April 2016
                : 2 August 2016
                Page count
                Figures: 7, Tables: 2, Pages: 19
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/501100000981, Horticulture Australia;
                Award ID: HG10025, MT13042
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100001794, University of Queensland;
                Award ID: ECR Grant
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100000923, Australian Research Council;
                Award ID: FT110100304
                Award Recipient :
                This research and a Fellowship for JK was financially supported by Horticulture Innovation Australia Ltd using voluntary contributions from Brisbane City Council and matched funds from the Australian Government (HG10025, MT13042, http://horticulture.com.au/) and a University of Queensland ECR Grant ( https://www.uq.edu.au/). An ARC Future Fellowship for GRF was financially supported by the Australian Research Council (FT110100304, http://www.arc.gov.au/). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Agriculture
                Crop Science
                Crops
                Fruits
                Tomatoes
                Biology and Life Sciences
                Organisms
                Plants
                Fruits
                Tomatoes
                Biology and Life Sciences
                Plant Science
                Plant Physiology
                Plant Reproduction
                Seed Germination
                Biology and Life Sciences
                Organisms
                Plants
                Flowering Plants
                Lettuce
                Biology and Life Sciences
                Agriculture
                Crop Science
                Crops
                Vegetables
                Lettuce
                Biology and Life Sciences
                Organisms
                Plants
                Vegetables
                Lettuce
                Biology and Life Sciences
                Plant Science
                Plant Anatomy
                Seeds
                Biology and Life Sciences
                Organisms
                Plants
                Seedlings
                Biology and Life Sciences
                Developmental Biology
                Plant Growth and Development
                Biology and Life Sciences
                Plant Science
                Plant Growth and Development
                Biology and Life Sciences
                Plant Science
                Plant Anatomy
                Leaves
                Biology and Life Sciences
                Organisms
                Plants
                Flowering Plants
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