Chloride: not simply a 'cheap osmoticum', but a beneficial plant macronutrient

Chloride is a nutrient that accumulates to millimolar levels in plants under most growth conditions, including in almost all soil-grown plants. These relatively high chloride concentrations (relative to the demand for chloride in photosynthesis) are beneficial to plants including non-halophytes, as the addition of chloride to the growth medium above the micromolar level increases biomass. As chloride is not metabolized and its only known essential function is in the oxygen-evolving complex in PSII, we discuss how chloride could be beneficial, especially in comparison with nitrate. We review the different routes taken by chloride in plants, from uptake and translocation to the shoot, and inside the cell in different organelles, including different transport mechanisms and the proteins identified. As the selectivity of many proteins to chloride and nitrate is not well established, the mechanisms within proteins to achieve selectivity of one anion over the other are explored. We further discuss the role of chloride as an osmoticum, why it might be preferentially used instead of other anions, and how chloride content in plants might beneficially influence nitrogen use efficiency and water-holding capacity.