Manganese accumulation in rice: implications for photosynthetic functioning.

In order to gain fundamental insights into the nature of the adaptation to Mn excess, the characterisation of the photosynthetic apparatus in Mn-treated rice was carried out in 21-day-old plants. We found 17- and 11-fold increases in Mn in the leaf tissues and in thylakoid, respectively, when the plants were grown hydroponically in nutrient solutions with Mn concentrations between 0.125 and 32 mg l(-1) (2.3 and 582.5 microM). Net photosynthesis and the photosynthetic capacity decreased after the 0.5 and 2 mg l(-1) (9.1 and 36.4 microM) Mn treatment, respectively. The stomatal conductance displayed a similar trend to that of photosynthetic capacity. The levels of basal chlorophyll fluorescence and the ratio between variable and maximum chlorophyll fluorescence did not vary significantly among treatments, but the photochemical quenching and the quantum yield of non-cyclic electron transport increased until the 2 mg l(-1) (36.4 microM) Mn treatment. The lipid matrix of thylakoids revealed a global increase in the proportions of phospholipids, relative to galactolipids. This pattern was coupled with diminishing levels of monogalactosyldiacylglycerol. The relative ratio between total carotenoids and total chlorophylls decreased until the last Mn treatment, yet the levels of carotenes, zeaxanthin, and violaxanthin plus antheraxanthin displayed different patterns. It was further found that the de-epoxidation state involving the components of the xanthophylls cycle increased until the 8 mg l(-1) (145.6 microM) Mn treatment. The levels of the photosynthetic electron carriers displayed different patterns, with plastocyanin and the high and low forms of cytochrome b559 remaining steady, whereas cytochromes b563 and f increased until the 8 mg l(-1) (145.6 microM) Mn treatment and the quinone pool increased until the highest Mn treatment. It was concluded that Mn-mediated inhibition of rice photosynthesis barely implicates stomatal conductance, as well as the distribution of energy within the photosystems. In this context, alterations to the relative proportions of the different acyl lipids and isoprenoids, as well as to the accumulations of the photosynthetic electron carriers, seem to play a major role.