One sentence summary: Expression of PDX1.2 is regulated by the HSFA1 transcription factor 15family and is an important abiotic stress tolerance strategy in most eudicots that is not utilized by 16 monocots such as grasses.
ABSTRACT 28Plants sense temperature changes and respond by altering growth and metabolic activity to 29 acclimate to the altered environmental conditions. The B vitamins give rise to vital coenzymes that 30 are indispensable for growth and development but their inherent reactive nature renders them prone 31 to destruction especially under stress conditions. Therefore, plant survival strategies would be 32 expected to include mechanisms to sustain B vitamin supply under demanding circumstances. Here, 33 using the example of vitamin B 6 , we investigate the regulation of biosynthesis across eudicot and 34 monocot species under heat stress. Most eudicots carry a pseudoenzyme PDX1.2 that is a non-35 catalytic homolog of the PDX1 subunit of the vitamin B 6 biosynthesis protein machinery, 36 PYRIDOXINE SYNTHASE. Using Arabidopsis and tomato as models, we show that PDX1.2 is 37 transcriptionally regulated by the HSFA1 transcription factor family. Monocots only carry catalytic 38 PDX1 homologs but do not respond to heat stress as demonstrated for rice and maize, suggesting 39 fundamental differences in the regulation of vitamin B 6 biosynthesis across the two lineages. 40Investigation of the molecular mechanism of PDX1.2 transcription reveals two alternative 41 transcriptional start sites one of which is exclusive to heat stress. Further data suggest that PDX1.2 42 leads to stabilization of the catalytic PDX1s under heat stress conditions, which would serve to 43 maintain vitamin B 6 homeostasis in times of need in eudicots that carry this gene. Our analyses 44indicate an important abiotic stress tolerance strategy in several eudicots, which has not been 45 evolutionarily adapted (or is not required) by monocots such as grasses.
INTRODUCTION 47The B vitamins are essential for survival of all organisms, as they provide important coenzymes for 48 numerous cellular proteins and have more recently been implicated in non-coenzyme related 49 activities (Colinas & Fitzpatrick, 2015). Plants are autonomous for these compounds 50 biosynthesizing them de novo from relatively simple precursors and are a major source of 51 micronutrients required by animals, including humans (Fitzpatrick et al., 2012). Being chemically 52 reactive by nature as coenzymes and vital for cellular function, it is important that homeostasis of B 53 vitamins is maintained with supply and demand of these compounds needing to be strictly 54 coordinated. In this context, the devastating developmental and physiological effects that 55 deficiencies in the B vitamin compounds can have on an organism, in particular humans, is 56 thoroughly documented, as well as consequences of oversupply (Kennedy, 2016, Spector & 57 Johanson, 2007. In plants, while there are numerous reports on B vitamin metabolism per se 58 (biosynthesis, transport), th...