title: Two P-type ATPases are required for copper delivery in Arabidopsis thaliana chloroplasts.
abstract: Copper delivery to the thylakoid lumen protein plastocyanin and the stromal enzyme Cu/Zn superoxide dismutase in chloroplasts is required for photosynthesis and oxidative stress protection. The copper delivery system in chloroplasts was characterized by analyzing the function of copper transporter genes in Arabidopsis thaliana. Two mutant alleles were identified of a previously uncharacterized gene, PAA2 (for P-type ATPase of Arabidopsis), which is required for efficient photosynthetic electron transport. PAA2 encodes a copper-transporting P-type ATPase with sequence similarity to PAA1, which functions in copper transport in chloroplasts. Both proteins localized to the chloroplast, as indicated by fusions to green fluorescent protein. The PAA1 fusions were found in the chloroplast periphery, whereas PAA2 fusions were localized in thylakoid membranes. The phenotypes of paa1 and paa2 mutants indicated that the two transporters have distinct functions: whereas both transporters are required for copper delivery to plastocyanin, copper delivery to the stroma is inhibited only in paa1 but not in paa2. The effects of paa1 and paa2 on superoxide dismutase isoform expression levels suggest that stromal copper levels regulate expression of the nuclear genes IRON SUPEROXIDE DISMUTASE1 and COPPER/ZINC SUPEROXIDE DISMUTASE2. A paa1 paa2 double mutant was seedling-lethal, underscoring the importance of copper to photosynthesis. We propose that PAA1 and PAA2 function sequentially in copper transport over the envelope and thylakoid membrane, respectively.
authors.affiliation: Biology Department, Colorado State University, Fort Collins, Colorado 80523, USA.
authors.forename: Salah E
authors.initials: SE
authors.lastname: Abdel-Ghany
authors.forename: Patricia
authors.initials: P
authors.lastname: Müller-Moulé
authors.forename: Krishna K
authors.initials: KK
authors.lastname: Niyogi
authors.forename: Marinus
authors.initials: M
authors.lastname: Pilon
authors.forename: Toshiharu
authors.initials: T
authors.lastname: Shikanai
mesh_terms: D000595:Amino Acid Sequence; D017360:Arabidopsis / Q000201:enzymology* / Q000235:genetics; D029681:Arabidopsis Proteins / Q000235:genetics / Q000302:isolation & purification / Q000378:metabolism*; D001483:Base Sequence; D002352:Carrier Proteins / Q000235:genetics / Q000302:isolation & purification / Q000378:metabolism*; D025222:Chloroplast Proton-Translocating ATPases / Q000235:genetics / Q000302:isolation & purification / Q000378:metabolism*; D002736:Chloroplasts / Q000201:enzymology* / Q000235:genetics; D003300:Copper / Q000378:metabolism*; D018076:DNA, Complementary; D005804:Genes, Lethal / Q000502:physiology; D008969:Molecular Sequence Data; D009154:Mutation / Q000235:genetics; D010788:Photosynthesis / Q000502:physiology*; D013482:Superoxide Dismutase / Q000378:metabolism; D020524:Thylakoids / Q000201:enzymology / Q000235:genetics
chemical_list: D029681:Arabidopsis Proteins; D002352:Carrier Proteins; D018076:DNA, Complementary; D003300:Copper; D013482:Superoxide Dismutase; D025222:Chloroplast Proton-Translocating ATPases; C477511:PAA1 protein, Arabidopsis; C507153:PAA2 protein, Arabidopsis