Plant Nucleotide Metabolism. Hiroshi Ashihara

and NaMN nucleotides of the initial nucleotide products of purine, pyrimidine, and pyridine biosynthesis (see Figure 3.1). In addition, glutamine and glycine are used for purine ring synthesis. The genes and enzymes of nitrogen assimilation and amino acid biosynthesis have been well studied in plants (Coruzzi et al. 2015).

3.10 Summary

De novo biosynthesis of ribonucleoside monophosphate, interconversion of nucleoside phosphates, conversion of nucleoside diphosphates to triphosphates, biosynthesis of deoxyribonucleotides, nucleic acid (DNA and RNA) biosynthesis, and supply of PRPP and amino acids for nucleotide biosynthesis are reviewed.

Figure 3.9 Outline of amino acids biosynthesis in plants. DAHP, 3-deoxy-D-arabino-heptulosonate-7-phosphate; E4P, erythrose-4-phosphate; F6P, fructose-6-phosphate; F1,6BP, fructose-1,6-bisphosphate; 6PG, 6-phosphogluconate; 3PGA, 3-phosphoglycerate; PEP, phosphoenolpyruvate.

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Part II Purine Nucleotide Metabolism

4 Purine Nucleotide Biosynthesis De Novo

      4.1 Introduction

      The de novo biosynthesis of purine nucleotides was elucidated independently by Buchanan and Greenberg in the 1950s (see, Buchanan 1986). The purine ring is assembled from several small molecules, namely, glycine, glutamine, aspartate, 5-phosphoribosyl-1-pyrophosphate (PRPP), 10-formyl tetrahydrofolate, and carbon dioxide. In the early studies pigeon liver was used, because birds synthesize purines from excess amino nitrogen, the purines are then degraded to urate and excreted. Incorporation of individual precursors into different atoms of urate was used to reveal the purine biosynthesis pathway. Similar precursor feeding studies were performed with leaves of coffee and tea which accumulate the caffeine (Anderson and Gibbs 1962; Proiser and Serenkov 1963). In plant purine studies, N₂-fixing root nodules of tropical legumes have been used as experimental material to investigate de novo purine biosynthesis. Substantial amounts of ureides are produced as the major nitrogen-containing compounds derived from purine nucleotides (Schubert and Boland 1990).

      Arabidopsis thaliana is a popular model organism in plant biology and genetics. It was the first plant genome to be sequenced, completed by the Arabidopsis-Genome-Initiative (2000). The most up-to-date version of the A. thaliana genome is maintained by the Arabidopsis Information Resource (TAIR). Although feeding experiments and detailed analysis of native enzymes of A. thaliana are difficult from a practical perspective, nucleotide metabolism research has developed aided by molecular genetic studies with the available sequence information from Arabidopsis and rice genomes, confirming the hypothesis that purine biosynthesis de novo in plants is essentially the same as that operating in microorganisms and animals (van der Graaff et al. 2004; Zrenner and Ashihara 2011). However, based on