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      Characterization of Novel Sorghum brown midrib Mutants from an EMS-Mutagenized Population

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          Reducing lignin concentration in lignocellulosic biomass can increase forage digestibility for ruminant livestock and saccharification yields of biomass for bioenergy. In sorghum ( Sorghum bicolor (L.) Moench) and several other C4 grasses, brown midrib ( bmr) mutants have been shown to reduce lignin concentration. Putative bmr mutants isolated from an EMS-mutagenized population were characterized and classified based on their leaf midrib phenotype and allelism tests with the previously described sorghum bmr mutants bmr2, bmr6, and bmr12. These tests resulted in the identification of additional alleles of bmr2, bmr6, and bmr12, and, in addition, six bmr mutants were identified that were not allelic to these previously described loci. Further allelism testing among these six bmr mutants showed that they represented four novel bmr loci. Based on this study, the number of bmr loci uncovered in sorghum has doubled. The impact of these lines on agronomic traits and lignocellulosic composition was assessed in a 2-yr field study. Overall, most of the identified bmr lines showed reduced lignin concentration of their biomass relative to wild-type (WT). Effects of the six new bmr mutants on enzymatic saccharification of lignocellulosic materials were determined, but the amount of glucose released from the stover was similar to WT in all cases. Like bmr2, bmr6, and bmr12, these mutants may affect monolignol biosynthesis and may be useful for bioenergy and forage improvement when stacked together or in combination with the three previously described bmr alleles.

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          Most cited references 18

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          Cloning and characterization of irregular xylem4 (irx4): a severely lignin-deficient mutant of Arabidopsis.

          A severe lignin mutant, irx4, has been identified in Arabidopsis thaliana as a result of its collapsed xylem phenotype. In contrast to previously described irx mutants, irx4 plants have 50% less lignin than wild-type plants, whilst the cellulose and hemicellulose content remained unchanged. These alterations in the composition of irx4 secondary cell walls had a dramatic effect on the morphology and architecture of the walls, which expand to fill most of the cell, and also on the physical properties of irx4 stems. Further analysis indicated that the irx4 mutation occurred in a cinnamoyl-CoA reductase (CCR) gene within a highly conserved intron splice site sequence of intron 2. As a result, CCR mRNA transcripts were incorrectly spliced. Transgenic plants expressing an IRX3 promoter-CCR cDNA construct were used to generate a series of plants with varying degrees of lignin content in order to assess the role of lignin content in determining the physical properties of Arabidopsis stems.
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            Characteristics of plant cell walls affecting intake and digestibility of forages by ruminants.

            Even under the intensive concentrate feeding systems of ruminant animal production in the United States, forages continue to represent the single most important feed resource. Cell-wall concentration and digestibility limit the intake potential and energy availability of forage crops in beef and dairy production. Identification of cell-wall characteristics that should be targets of genetic modification is required if plant breeders and molecular biologists are to successfully improve forages for livestock feeding. As the forage plant cell develops, phenolic acids and lignin are deposited in the maturing cell wall in specific structural conformations, and in a strict developmental sequence. Lignin is the key element that limits cell-wall digestibility, but cross-linkage of lignin and wall polysaccharides by ferulic acid bridges may be a prerequisite for lignin to exert its affect. Lignin composition and p-coumaric acid in the wall are less likely to affect digestibility. Voluntary intake of forages is a critical determinant of animal performance and cell-wall concentration is negatively related to intake of ruminants consuming high-forage diets. Cell walls affect intake by contributing to ruminal fill. A simple model of cell-wall digestion and passage in which ruminal fill is a function of rates of digestion and passage, as well as the indigestible fraction of the cell-wall indicates that cell-wall concentration and rate of passage are the most critical parameters determining ruminal fill. Plant factors that affect rate of passage include those that affect particle size reduction by chewing and those that affect particle buoyancy in the rumen. The latter is primarily affected by 1) the ability of the particulate matter to retain gases, which is probably related to plant anatomy and rate of digestion of the plant tissue, and 2) the rate at which the gas is produced, which is affected by the potentially digestible fraction of the particulate matter and the rate of digestion of this fraction. Increasing rate of digestion should increase rate of passage by diminishing the gas produced and increasing density over time. A reduction in the indigestible cell-wall fraction is beneficial because this will decrease fill and increase digestibility. Animal production and economic benefits from reduced cell-wall concentration and increased digestibility are significant. Because of the high cell-wall concentration and large digestible cell-wall fraction of grasses, reduction in cell-wall concentration would probably be of greater value than improving digestibility in these species. Legumes represent the opposite situation and may benefit more from improvements in the digestibility of their cell walls.
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              Applying genotyping (TILLING) and phenotyping analyses to elucidate gene function in a chemically induced sorghum mutant population

              Background Sorghum [Sorghum bicolor (L.) Moench] is ranked as the fifth most important grain crop and serves as a major food staple and fodder resource for much of the world, especially in arid and semi-arid regions. The recent surge in sorghum research is driven by its tolerance to drought/heat stresses and its strong potential as a bioenergy feedstock. Completion of the sorghum genome sequence has opened new avenues for sorghum functional genomics. However, the availability of genetic resources, specifically mutant lines, is limited. Chemical mutagenesis of sorghum germplasm, followed by screening for mutants altered in important agronomic traits, represents a rapid and effective means of addressing this limitation. Induced mutations in novel genes of interest can be efficiently assessed using the technique known as Targeting Induced Local Lesion IN Genomes (TILLING). Results A sorghum mutant population consisting of 1,600 lines was generated from the inbred line BTx623 by treatment with the chemical agent ethyl methanesulfonate (EMS). Numerous phenotypes with altered morphological and agronomic traits were observed from M2 and M3 lines in the field. A subset of 768 mutant lines was analyzed by TILLING using four target genes. A total of five mutations were identified resulting in a calculated mutation density of 1/526 kb. Two of the mutations identified by TILLING and verified by sequencing were detected in the gene encoding caffeic acid O-methyltransferase (COMT) in two independent mutant lines. The two mutant lines segregated for the expected brown midrib (bmr) phenotype, a trait associated with altered lignin content and increased digestibility. Conclusion TILLING as a reverse genetic approach has been successfully applied to sorghum. The diversity of the mutant phenotypes observed in the field, and the density of induced mutations calculated from TILLING indicate that this mutant population represents a useful resource for members of the sorghum research community. Moreover, TILLING has been demonstrated to be applicable for sorghum functional genomics by evaluating a small subset of the EMS-induced mutant lines.

                Author and article information

                G3 (Bethesda)
                G3: Genes, Genomes, Genetics
                G3: Genes, Genomes, Genetics
                G3: Genes, Genomes, Genetics
                G3: Genes|Genomes|Genetics
                Genetics Society of America
                2 September 2014
                November 2014
                : 4
                : 11
                : 2115-2124
                [* ]Grain, Forage and Bioenergy Research Unit, USDA-ARS, Lincoln, Nebraska 68583
                []Department of Agronomy and Horticulture, University of Nebraska–Lincoln, Lincoln, Nebraska 68583
                []Agronomy Department and Genetics Institute, University of Florida, Gainesville, Florida 32610
                [§ ]Plant Stress and Germplasm Development Unit, USDA-ARS, Lubbock, Texas 79415
                [** ]Department of Plant Pathology, University of Nebraska–Lincoln, Lincoln, Nebraska 68583
                [†† ]Department of Microbiology and Cell Science and Genetics Institute, University of Florida, Gainesville, Florida 32610
                Author notes

                Present address: Chromatin Inc., Alachua, FL 32615.

                [2 ]Corresponding author: Grain, Forage, and Bioenergy Research Unit USDA-ARS, 137 Keim Hall, East Campus UNL, Lincoln, NE 68583-0937. E-mail: Scott.Sattler@

                The use of trade, firm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the United States Department of Agriculture, the Agricultural Research Service, or the Northern Plains Area of any product or service to the exclusion of others that may be suitable.


                SAS and all other SAS Institute Inc. product or service names are registered trademarks or trademarks of SAS Institute Inc. in the USA and other countries. Indicates USA registration.

                Copyright © 2014 Sattler et al.

                This is an open-access article distributed under the terms of the Creative Commons Attribution Unported License (, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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