The cytokinins, which are N 6 -substituted adenine derivatives, control key aspects of crop productivity. Cytokinin levels are controlled via biosynthesis by isopentenyl transferase (IPT), destruction by cytokinin oxidase/dehydrogenase (CKX), and inactivation via glucosylation by cytokinin glucosyl transferases (CGTs). While both yield components and tolerance to drought and related abiotic stressors have been positively addressed via manipulation of IPT and/or CKX expression, much less attention has been paid to the CGTs. As naming of the CGTs has been unclear, we suggest COGT, CNGT, CONGT and CNOGT to describe the O-, N-and dual function CGTs. As specific CGT mutants of both rice and arabidopsis showed impacts on yield components, we interrogated the wheat genome database, IWGSC RefSeq v1.0 & v2.0, to investigate wheat CGTs. Besides providing unambiguous names for the 53 wheat CGTs, we show their expression patterns in 70 developmental tissues and their response characteristics to various stress conditions by reviewing more than 1000 RNA-seq data sets. These revealed various patterns of responses and showed expression generally being more limited in reproductive tissues than in vegetative tissues. Multiple cis-regulatory elements are present in the 3 kb upstream of the start codons of the 53 CGTs. Elements associated with abscisic acid, light and methyl jasmonate are particularly over-represented, indicative of the responsiveness of CGTs to the environment. These data sets indicate that CGTs have potential value for wheat improvement and that these could be targeted in TILLING or gene editing wheat breeding programmes.genes controlling the formation of the deactivated cytokinin Oand N-glucosides. After introducing the cytokinins more generally, this article focuses on the cytokinin O-and N-glucosyl transferases and their potential roles in modifying yield in bread wheat, under ambient and stress conditions.
Cytokinin biosynthesis and metabolismNaturally occurring cytokinins, which are adenine derivatives, fall into two groups: those with an isoprenoid side chain and those with an aromatic one. The isoprenoid cytokinins are considered to be the predominant forms (Jaworek et al., 2019;Sakakibara, 2006). The aromatics are not covered in this review as, while not exhaustively looked for in all cereals, they were not detected in either maize leaves (Lacuesta et al., 2018) or in rice (Kudo et al., 2012), although low levels of 6-benzylamino purine were recently reported in wheat spikes (Jabło nski et al., 2020) and low levels of various aromatic metabolites have been detected in germinating seeds of maize and oats (Stirk et al., 2012).