Trends in Plant Science
PerspectivesA guide to the Lhc genes and their relatives in Arabidopsis
Section snippets
EST clones originating from the Lhc super-gene family
The Arabidopsis genome project has already produced an immense amount of data and the full genomic sequence will soon be available. Many of the EST clones currently available (30 000 EST sequences are accessible in the public databases and the corresponding clones can be ordered from the Arabidopsis Stock Center) correspond to Lhc genes, both from the ten that had previously been characterized (five Lhcb1 and one each of Lhca1, Lhca2, Lhca3, Lhca4 and Lhcb4) and from additional members of the
Lhca genes
Lhca1–4 genes encode the polypeptides of light-harvesting complex I (LHC I) associated with PSI. LHC I is composed of dimers of Lhca proteins but the numbers and exact positions of these dimers have not been determined. An Arabidopsis Lhca1 gene was the first to be sequenced6 and Lhca3 (Ref. 7) and Lhca4 (Ref. 8) cDNAs have subsequently been analysed. An Lhca2 sequence has also been reported9 but the encoded protein is different from other Lhca2 gene products. There is no EST clone
Lhcb1–3 genes
The Lhcb1, Lhcb2 and Lhcb3 genes encode the polypeptides of trimeric LHC II. Lhcb1 homotrimers and Lhcb1/Lhcb2 heterotrimers function as antenna for both photosystems, whereas trimers containing Lhcb3 (probably 2Lhcb1/Lhcb3) appear to be confined to PSII.
Five Lhcb1 genes have been characterized previously from Arabidopsis11, 12 and all Lhcb1 EST clones appear to originate from these genes, indicating that no additional Lhcb1 genes are present. There are large differences in the expression
Lhcb4–6 genes
The Lhcb4, Lhcb5 and Lhcb6 proteins (often called CP29, CP26 and CP24, respectively) are probably monomeric proteins that are present in one copy per PSII unit. One Lhcb4 gene has been sequenced13, but two additional Lhcb4 genes are present in Arabidopsis. Two genes have about the same level of expression, whereas the third (Lhcb4.3) is expressed only at a low level. The protein encoded by Lhcb4.3 is different from the other two proteins in that it lacks a large part of the C-terminal region
The cousins, PsbS and ELIPs
Two proteins with homology to the LHC proteins, PsbS and early light-inducible proteins (ELIPs), have been characterized from higher plants previously. This similarity is not only at the sequence level, as both have been shown recently to bind chlorophyll and carotenoids15. All Arabidopsis PsbS EST clones appear to originate from a single gene. The ELIPs are also three-helix members of the super-gene family, and probably associated with PSII. The ELIP EST clones fall into two groups, indicating
Novel members of the family
Analysis of EST clones has resulted in the identification of several novel sequences with significant sequence homology to the Lhc genes. A common feature of all these genes is that they encode proteins with one, not two, ‘generic LHC motifs' in their sequences (Fig. 3). The first of these has been used for more extensive studies, and it transpires that the encoded protein is a single-helix thylakoid protein of 7 kDa (S. Jansson et al., unpublished). The closest homologues to this protein in
Concluding remarks
An analysis of the EST clones present in the public databases has revealed that the LHC super-gene family of Arabidopsis contains >30 members, of which some encode proteins that were previously unknown. Some of these might have a role in high-light acclimation – ongoing studies will clarify this. Do these 30 genes represent the complete super-gene family? It is unlikely that there are additional genes encoding proteins in unit stoichiometry in PSI or PSII, but the possibility that there could
Acknowledgements
My thanks to Ulrika Ganeteg for sequencing EST clones, to Iwona Adamska who generously shared unpublished data and to Vaughan Hurry who commented on the manuscript. Financial support was provided by the Swedish Forestry and Agricultural Research Council and the Foundation for Strategic Research. Finally, this work would not have been possible without the excellent services of the Arabidopsis Biological Resource Center (ABRC) in Ohio, USA.
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