N-terminal truncated RHT-1 proteins generated by translational reinitiation cause semi-dwarfing of wheat Green Revolution alleles

The unprecedented wheat yield increases during the Green Revolution were achieved through the introduc-tion of the Reduced height(Rht)-B1b and Rht-D1b semi-dwarfing alleles.These Rht-1 alleles encode growth-repressing DELLA genes containing a stop codon within their open reading frame that confers gibberellin(GA)-insensitive semi-dwarfism.In this study,we successfully took the hurdle of detecting wild-type RHT-1 proteins in different wheat organs and confirmed their degradation in response to GAs.We further demonstrated that Rht-B1b and Rht-D1b produce N-terminal truncated proteins through trans-lational reinitiation.Expression of these N-terminal truncated proteins in transgenic lines and in Rht-D1c,an allele containing multiple Rht-D1b copies,demonstrated their ability to cause strong dwarfism,resulting from their insensitivity to GA-mediated degradation.N-terminal truncated proteins were detected in spikes and nodes,but not in the aleurone layers.Since Rht-B 1b and Rht-D1b alleles cause dwarfism but have wild-type dormancy,this finding suggests that tissue-specific differences in translational reinitiation may explain why the Rht-1 alleles reduce plant height without affecting dormancy.Taken together,our findings not only reveal the molecular mechanism underlying the Green Revolution but also demonstrate that trans-lational reinitiation in the main open reading frame occurs in plants.