Temporal and spatial variations of carbon isotope signature reveal substantial contribution of bracts and internode assimilates to grain filling of japonica rice

Carbon isotope composition (δ¹³C) of a plant organ is an inherent signature reflecting its physiological property, and thus is used as an integrative index in crop breeding. It is also a non-intrusive method for quantifying the relative contribution of different source organs to grain filling in cereals. Using the samples collected from two-year field and pot experiments with two nitrogen (N) fertilization treatments, we investigated the temporal and spatial variations of δ¹³C in source organs of leaf, sheath, internode, and bracts, and in sink organ grain. Constitutive nature of δ¹³C was uncovered, with an order of leaf (−27.84‰) < grain (−27.82‰) < sheath (−27.24‰) < bracts (−26.81‰) < internode (−25.67‰). For different positions of individual organs within the plant, δ¹³C of the leaf and sheath presented a diminishing trend from the top (flag leaf and its sheath) to the bottom (the last leaf in reverse order and its sheath). No obvious pattern was found for the internode. For temporal variations, δ¹³C of the leaf and sheath had a peak (the most negative) at 10 days after anthesis (DAA), whereas that of the bracts showed a marked increase at the time point of anthesis, implying a transformation from sink to source organ. By comparing the δ¹³C in its natural abundance in the water-soluble fractions of the sheath, internode, and bracts with the δ¹³C in mature grains, the relative contribution of these organs to grain filling was assessed. With reference to the leaf, the internode accounted for as high as 32.64% and 42.56% at 10 DAA and 20 DAA, respectively. Meanwhile, bracts presented a larger contribution than the internode, with superior bracts being higher than inferior bracts. In addition, N topdressing reduced the contribution of the internode and bracts. Our findings clearly proved the actual significance of non-foliar organs of the internode and bracts for rice yield formation, thus extending our basic knowledge of source and sink relations.