Water-extractable humic substances speed up transcriptional response of maize roots to nitrate

Humic substances are known to positively influence plant growth and nutrition. In particular, the water-ex- tractable fraction of humic substances (WEHS) has been shown to enhance nitrate acquisition, increasing the activity of high affinity nitrate uptake system. However, molecular bases of this physiological response are not clarified so far. Thus, in the present work, the physiological effect of WEHS on nitrate acquisition in maize roots was cor- related with changes in the root transcriptomic profile. Results confirmed that WEHS caused a faster induction of a higher capacity to take up nitrate in maize roots. Comparing the root transcriptomic profile of Nitrate- and Nitrate + WEHS-treated plants with Control (-N) ones, more than 2000 transcripts appeared to be modulated only in the presence of WEHS. Among these, genes involved in nitrate transport and assimilation (NRT1s, NRT2s, NAR2.1, NR, GS, GOGAT, CNX, UPM) were strongly modulated by WEHS. Furthermore, also some genes known to be linked to the nitrogen limitation responses were affected by WEHS, as transcripts coding for transcription factors (as LBD37, NIN-like protein, NFY- A, GRF5) and enzymes of hormones’ metabolism. The modulation of these transcripts might play a crucial role in coordinating the induction to nitrate, favouring its uptake and assimilation in WEHS-treated plants. The over- expression of nitrogen assimilatory genes by WEHS might led to an early feedback regulation of the high affinity nitrate transport system, as being operated by N-metabolites. Results of the present work shed further light on the contribution of the organic soil component to the nitrogen use efficiency in crops.