title: STARCH-EXCESS4 is a laforin-like Phosphoglucan phosphatase required for starch degradation in Arabidopsis thaliana.
abstract: Starch is the major storage carbohydrate in plants. It is comprised of glucans that form semicrystalline granules. Glucan phosphorylation is a prerequisite for normal starch breakdown, but phosphoglucan metabolism is not understood. A putative protein phosphatase encoded at the Starch Excess 4 (SEX4) locus of Arabidopsis thaliana was recently shown to be required for normal starch breakdown. Here, we show that SEX4 is a phosphoglucan phosphatase in vivo and define its role within the starch degradation pathway. SEX4 dephosphorylates both the starch granule surface and soluble phosphoglucans in vitro, and sex4 null mutants accumulate phosphorylated intermediates of starch breakdown. These compounds are linear alpha-1,4-glucans esterified with one or two phosphate groups. They are released from starch granules by the glucan hydrolases alpha-amylase and isoamylase. In vitro experiments show that the rate of starch granule degradation is increased upon simultaneous phosphorylation and dephosphorylation of starch. We propose that glucan phosphorylating enzymes and phosphoglucan phosphatases work in synergy with glucan hydrolases to mediate efficient starch catabolism.
authors.affiliation: Institute of Plant Sciences, ETH Zurich, 8092 Zurich, Switzerland. koliver@ethz.ch
authors.forename: Oliver
authors.initials: O
authors.lastname: Kötting
authors.forename: Diana
authors.initials: D
authors.lastname: Santelia
authors.forename: Christoph
authors.initials: C
authors.lastname: Edner
authors.forename: Simona
authors.initials: S
authors.lastname: Eicke
authors.forename: Tina
authors.initials: T
authors.lastname: Marthaler
authors.forename: Matthew S
authors.initials: MS
authors.lastname: Gentry
authors.forename: Sylviane
authors.initials: S
authors.lastname: Comparot-Moss
authors.forename: Jychian
authors.initials: J
authors.lastname: Chen
authors.forename: Alison M
authors.initials: AM
authors.lastname: Smith
authors.forename: Martin
authors.initials: M
authors.lastname: Steup
authors.forename: Gerhard
authors.initials: G
authors.lastname: Ritte
authors.forename: Samuel C
authors.initials: SC
authors.lastname: Zeeman
mesh_terms: D017360:Arabidopsis / Q000201:enzymology* / Q000235:genetics; D029681:Arabidopsis Proteins / Q000235:genetics / Q000378:metabolism*; D050260:Carbohydrate Metabolism; D004269:DNA, Bacterial / Q000235:genetics; D005936:Glucans / Q000378:metabolism; D016254:Mutagenesis, Insertional; D010766:Phosphorylation; D017027:Protein Tyrosine Phosphatases / Q000235:genetics / Q000378:metabolism*; D011994:Recombinant Proteins / Q000235:genetics / Q000378:metabolism; D013213:Starch / Q000378:metabolism*
chemical_list: D029681:Arabidopsis Proteins; D004269:DNA, Bacterial; D005936:Glucans; D011994:Recombinant Proteins; C036483:T-DNA; D013213:Starch; D017027:Protein Tyrosine Phosphatases