DELLA-PIF Regulation of Nitrogen Assimilation: from Arabidopsis Model to Long-Term Translation to Crop Efficiency Gains

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The famous cereal 'green revolution' of the 1960s/1970s increased crop yields, averted famine and fed a growing world population. Green Revolution Varieties (GRVs) of rice and wheat were the genetic foundation of the green revolution. GRVs carry mutant growth regulatory genes that confer dwarfism, and this dwarfism increases yield because it reduces loss due to 'lodging' (flattening of plants by wind and rain), hence causing the yield increases of the green revolution. However, the mutant growth regulatory genes also cause GRVs to be less efficient in assimilating the nitrogen (N) supplied to them in the form of fertilizer. As a result, N that is not assimilated by GRVs is dissipated into the wider environment, where it causes severe damage to terrestrial and aquatic ecosystems, together with atmospheric greenhouse-gas pollution that precipitates climate change. Because today's high-yielding crop varieties still depend upon the mutant dwarfing genes for their high yields, it is necessary to find ways of developing new crop varieties that retain the benefits of GRV dwarfism but that are more efficient in their use of N fertilizers (have improved N use efficiency, NUE).Here we propose to exploit the rapid genetics and molecular biology of the genetic model Arabidopsis to make discoveries that will enable future enhancement of GRV NUE. The GRV dwarfing genes cause accumulation of a class of growth inhibitory proteins called DELLAs, and DELLAs also accumulate in the dwarf Arabidopsis GRV mutant model gai. Accumulated DELLAs inhibit the action of another class of regulatory proteins, the PIFs (or Phytochrome Interacting Factors). Our recent preliminary evidence from studies of Arabidopsis suggest that the inhibitory effect of DELLAs on PIFs may explain the reduced NUE of GRVs, and it is this novel and exciting finding that we exploit in this proposal. We will therefore first further test our working hypothesis that interactions between DELLAs and PIFs affect the assimilation of N: that the DELLAs accumulated in GRVs and gai oppose PIF function, thus reducing N assimilation. If this hypothesis is correct, modulation of the DELLA-PIF relationship may provide a novel route towards improving GRV NUE. We have the following objectives: A. Obtain an in-depth understanding of PIF-regulation of Arabidopsis and rice N assimilation - essentially performing genetic tests of the role of PIFs in regulation of N metabolism and assimilation in Arabidopsis and rice. B. Determine how the DELLA-PIF interaction regulates the abundance of mRNA encoding nitrate reductase (NR), a key enzyme in N assimilation - this an exploration of how the DELLA-PIF interaction controls the expression of the gene encoding that enzyme.C. Determine if the DELLA-PIF interaction also directly affects the abundance and/or specific enzymatic activity of the NR enzyme itself.D. Determine if NUE can be increased despite retaining yield-enhancing dwarfism. This is important because it could lead to the development of crops which retain the high yields of current GRVs, but at reduced environmental cost. First, we will determine if increasing PIF activity might confer such benefits. However, because increasing the activity of PIFs themselves in GRVs might have additional unwanted consequences, we will additionally explore other routes (downstream of PIFs) to improving GRV NUE whilst retaining yield-enhancing dwarfism. Inherent in our strategy is initial translation of findings from Arabidopsis model to crop (rice), exploiting our long-standing combined expertise in DELLA biology, model-crop translations, and whole genome sequence analysis. Our long-term aim (future proposals) is to use the fundamental understanding gained here in the development of rice and wheat GRVs having enhanced NUE, thus enhancing global food security and reducing agricultural environmental degradation.