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Published on December 3, 2004; 10.1104/pp.104.052951

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Received September 7, 2004
Returned for revision October 15, 2004
Accepted October 15, 2004

Identification of the Arabidopsis Palmitoyl-Monogalactosyldiacylglycerol {Delta}7-Desaturase Gene FAD5, and Effects of Plastidial Retargeting of Arabidopsis Desaturases on the fad5 Mutant Phenotype

Ingo Heilmann , Sergei Mekhedov , Barbara King , John Browse , and John Shanklin *

Department of Biology, Brookhaven National Laboratory, Upton, New York 11973
Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824
Institute of Biological Chemistry, Washington State University, Pullman, Washington 99164

* Corresponding author; email: shanklin{at}bnl.gov.

Hexadeca 7,10,13-trienoic acid (16:3{Delta}7,10,13) is one of the most abundant fatty acids in Arabidopsis (Arabidopsis thaliana) and a functional component of thylakoid membranes, where it is found as an sn-2 ester of monogalactosyldiacylglycerol. The Arabidopsis fad5 mutant lacks activity of the plastidial palmitoyl-monogalactosyldiacylglycerol {Delta}7-desaturase FAD5, and is characterized biochemically by the absence of 16:3{Delta}7,10,13 and physiologically by reduced chlorophyll content and a reduced recovery rate after photoinhibition. While the fad5 mutation has been mapped, the FAD5 gene was not unambiguously identified, and a formal functional characterization by complementation of fad5 mutant phenotypes has not been reported. Two candidate genes (At3g15850 and At3g15870) predicted to encode plastid-targeted desaturases at the fad5 chromosomal locus were cloned from fad5 plants and sequenced. A nonsense mutation changing codon TGG (Trp-98) into TGA (stop) was identified in At3g15850 (ADS3), whereas the fad5 At3g15870 allele was identical to wild type (after correction of a sequencing error in the published wild-type genomic At3g15870 sequence). Expression of a genomic clone or cDNA for wild-type At3g15850 conferred on fad5 plants the ability to synthesize 16:3{Delta}7,10,13 and restored leaf chlorophyll content. Arabidopsis carrying a T-DNA insertion in At3g15870 had wild-type levels of both 16:3{Delta}7,10,13 and chlorophyll. Together, these data formally prove that At3g15850 is FAD5. Interestingly, the fad5 phenotype was partially complemented when extraplastidial {Delta}9-desaturases of the Arabidopsis desaturase (ADS) family were expressed as fusions with a plastidial transit peptide. Tight correlation between leaf 16:3{Delta}7,10,13 levels and chlorophyll content suggests a role for plastidial fatty acid desaturases in thylakoid formation.




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