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First published online January 24, 2008; 10.1104/pp.107.111476 Plant Physiology 146:1085-1097 (2008) © 2008 American Society of Plant Biologists OPEN ACCESS ARTICLE
Tie-dyed2 Functions with Tie-dyed1 to Promote Carbohydrate Export from Maize Leaves1,[C],[W],[OA]Department of Biology, Pennsylvania State University, University Park, Pennsylvania 16802
Regulation of carbon partitioning is essential for plant growth and development. To gain insight into genes controlling carbon allocation in leaves, we identified mutants that hyperaccumulate carbohydrates. tie-dyed2 (tdy2) is a recessive mutant of maize (Zea mays) with variegated, nonclonal, chlorotic leaf sectors containing excess starch and soluble sugars. Consistent with a defect in carbon export, we found that a by-product of functional chloroplasts, likely a sugar, induces tdy2 phenotypic expression. Based on the phenotypic similarities between tdy2 and two other maize mutants with leaf carbon accumulation defects, tdy1 and sucrose export defective1 (sxd1), we investigated whether Tdy2 functioned in the same pathway as Tdy1 or Sxd1. Cytological and genetic studies demonstrate that Tdy2 and Sxd1 function independently. However, in tdy1/+; tdy2/+ F1 plants, we observed a moderate chlorotic sectored phenotype, suggesting that the two genes are dosage sensitive and have a related function. This type of genetic interaction is referred to as second site noncomplementation and has often, though not exclusively, been found in cases where the two encoded proteins physically interact. Moreover, tdy1; tdy2 double mutants display a synergistic interaction supporting this hypothesis. Additionally, we determined that cell walls of chlorotic leaf tissues in tdy mutants contain increased cellulose; thus, tdy mutants potentially represent enhanced feedstocks for biofuels production. From our phenotypic and genetic characterizations, we propose a model whereby TDY1 and TDY2 function together in a single genetic pathway, possibly in homo- and heteromeric complexes, to promote carbon export from leaves.
1 This work was partially supported by the National Research Initiative of the U.S. Department of Agriculture Cooperative State Research, Education and Extension Service (grant no. 2004–35304–14948 to D.M.B.). The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: David M. Braun (dbraun{at}psu.edu). [C] Some figures in this article are displayed in color online but in black and white in the print edition. [W] The online version of this article contains Web-only data. [OA] Open Access articles can be viewed online without a subscription. www.plantphysiol.org/cgi/doi/10.1104/pp.107.111476 * Corresponding author; e-mail dbraun{at}psu.edu. Received October 22, 2007; accepted January 14, 2008; published January 24, 2008.
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