ABSTRACIThe mechanisms priming the production, the movement, and the transient and final storage of the photoassimilated carbon in the maize plant were examined at the metabolic level during the formation of the seed, with the ultimate aim to identify metabolic steps restricting grain yield and explaining the delay of formation of the reserve molecules. Under normal field conditions, we show that maize directly supplies the developing seed with the photoassimilated carbon which undergoes numerous interconversions from the ear leaf to the grain. The proteins, either in the leaf or in the seed, are primarily synthesized from incoming amino acids. Nevertheless, a secondary in situ synthesis of amino acids provides the proteins with new amino acids. The amino acids of this second set, slowly synthesized in the seed from the photosynthetic carbon skeletons, are not detected in their free form but immediately and regularly incorporated into the seed proteins, in such a way that, after 4 days of chase, the proportion of the radioactive labeling of the amino acids of the different storage protein groups corresponds to their amino acid composition. In the leaf, the labeling of proteins also arises from different metabolisms, but mainly from the photosynthetic metabolism. Contrary to the seed proteins, the time course of the labeled leaf proteins implies a rapid turnover. The second labeling of starch and proteins in the ear leaf involves a reassimilation of C02, a process optimizing the carbon uptake in maize.established in maize under physiological conditions. The only data in this area are those by Sodek and Wilson (27) who tried to follow the metabolism of amino acids by introducing 14C leucine and 14C lysine into the shank and endosperm of developing maize seed. However, these partial results, obtained under artificial conditions in the absence ofdata concerning the incoming precursors, did not give reliable and general information about the real in vivo metabolism involved in the seed protein synthesis. The interest of the interconversion studies lies in the peculiar amino acid balance of the maize seed storage proteins (19), compared to leaf functional proteins. As regards the leaf protein synthesis, most results concern the turnover ofthe major proteins (25) and not their formation from photosynthates.Consequently, we focused our attention on the fate of the photoassimilated carbon into leaf and seed proteins in maize plants grown in the field under normal physiological conditions. Our purpose was to understand, at the metabolic level, the mechanisms priming the production, the movement, and the transient and final storage of the photoassimilated carbon in the maize plant during the formation of the seed, with ultimate aim to identify metabolic steps restricting the plant grain yield. We point out the existence ofdifferent simultaneous metabolic pathways with numerous interconnections between carbon and nitrogen metabolism and some limiting steps in seed protein synthesis which might be now considered as bottlenec...
To gain a better understanding of the biochemical basis for partitioning of photosynthetically fixed carbon between leaf and grain, a '4C02 labeling study was conducted with field-grown maize plants 4 weeks after flowering. The carbon flow was monitored by separation and identification of '4C assimilates and '4C storage components within each tissue during the chase period (from 4 to 96 hours) following a 5 minute '4CO2 pulse.In the labeled ear leaf, the radioactivity strongly decreased to reach, at the end of the experiment, about 12% of the total incorporated radioactivity, mostly associated with sucrose and proteins. Nevertheless, an unexpected reincorporation of radioactivity was observed either in leaf starch or proteins, the day following the pulse. Conversely, the radioactivity in the grain increased to attain 66% of the total incorporated '4C after a 96 hour chase. The photosynthates, mostly sucrose, organic and free amino acids, rapidly translocated towards the developing seeds, served as precursors for the synthesis of seed storage compounds, starch, and proteins. They accumulate in free form for 24 hours before being incorporated within polymerized storage components. This delay is interpreted as a necessary prerequisite for interconversions prior to the polycondensations. In the grain, the labeling of the storage molecules, either in starch or in storage protein groups (salt-soluble proteins, zein, and glutelin subgroups), was independent of their chemical nature but dependent on their pool size.
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