Frédéric Verret

AtHMA4 is an Arabidopsis thaliana P1B-ATPase which transports Zn and Cd. Here, we demonstrate that AtHMA4 is localized at the plasma membrane and expressed in tissues surrounding the root vascular vessels. The ectopic overexpression of AtHMA4 improved the root growth in the presence of toxic concentrations of Zn, Cd and Co. A null mutant exhibited a lower translocation of Zn and Cd from the roots to shoot. In contrast, the AtHMA4 overexpressing lines displayed an increase in the zinc and cadmium shoot content. Altogether, these results strongly indicate that AtHMA4 plays a role in metal loading in the xylem.

The Arabidopsis thaliana AtHMA3 protein belongs to the P(1B)-adenosine triphosphatase (ATPase) transporter family, involved in heavy metal transport. Functional expression of AtHMA3 phenotypically complements the Cd/Pb-hypersensitive yeast strain Deltaycf1, but not the Zn-hypersensitive mutant Deltazrc1. AtHMA3-complemented Deltaycf1 cells accumulate the same amount of cadmium as YCF1-complemented Deltaycf1 cells or wild-type cells, suggesting that AtHMA3 carries out an intracellular sequestration of Cd. A mutant of AtHMA3 altered in the P-ATPase phosphorylation domain did not complement Deltaycf1, suggesting that metal transport rather than chelation is involved. The fusion protein AtHMA3::green fluorescent protein (GFP) is localized at the vacuole, consistent with a role in the influx of cadmium into the vacuolar compartment. In A. thaliana, the mRNA of AtHMA3 was detected mainly in roots, old rosette leaves and cauline leaves. The expression levels were not affected by cadmium or zinc treatments.

Summary Much of our current knowledge on the mechanisms by which Ca 2+ signals are generated in photosynthetic eukaryotes comes from studies of a relatively small number of model species, particularly green plants and algae, revealing some common features and notable differences between ‘plant’ and ‘animal’ systems. Physiological studies from a broad range of algal cell types have revealed the occurrence of animal‐like signalling properties, including fast action potentials and fast propagating cytosolic Ca 2+ waves. Genomic studies are beginning to reveal the widespread occurrence of conserved channel types likely to be involved in Ca 2+ signalling. However, certain widespread ‘ancient’ channel types appear to have been lost by certain groups, such as the embryophytes. More recent channel gene loss is also evident from comparisons of more closely related algal species. The underlying processes that have given rise to the current distributions of Ca 2+ channel types include widespread retention of ancient Ca 2+ channel genes, horizontal gene transfer (including symbiotic gene transfer and acquisition of bacterial genes), gene loss and gene expansion within taxa. The assessment of the roles of Ca 2+ channel genes in diverse physiological, developmental and life history processes represents a major challenge for future studies. Contents Summary 23 I. Introduction 24 II. Physiological features of Ca 2+ channel activity ineukaryotic photoautotrophs 25 III. Expansion, loss and horizontal gene transfer of Ca 2+ ‐permeable channels in photoautotrophs 28 IV. Conclusions 36 Acknowledgements 37 References 37