Abstract The root hair (RH) cells can elongate to several hundred times their initial size, and are an ideal model system for investigating cell size control. Their development is influenced by both endogenous and external signals, which are combined to form a integrative response. Surprisingly, a low temperature condition of 10°C causes an increased RH growth in Arabidopsis and in several monocots, even when the development of the rest of the root and aerial parts of the plant are halted. Previously, we demonstrated a strong correlation between the growth response and a significant decrease in nutrient availability in the medium under low temperature conditions. However, the molecular basis responsible for receiving and transmitting signals related to the availability of nutrients in the soil, and their relation to plant development, remain largely unknown. We decided to further investigate the intricate molecular processes behind the particular responsiveness of this root cell type at low temperature. In this study, we have discovered a gene regulatory network (GRN) controlling early transcriptome responses to low temperature. This GNR is commanded by specific transcription factors (FTs), namely ROOT HAIR DEFECTIVE 6-LIKE 4 (RSL4), a member of the homeodomain leucine zipper (HD-Zip I) group I 13 (AtHB13), the trihelix TF GT2-LIKE1 (GTL1), and a previously unidentified MYB-like TF (AT2G01060). Furthermore, we have identified four downstream RSL4 targets AtHB16 , AtHB23 , EARLY-RESPONSIVE TO DEHYDRATION 7 (ERD7) and ERD10 suggesting their participation in the regulation of RH development under these conditions. Functional analysis shows that such components of the RSL4-dependent transcriptional cascade influence the subsequent RH growth response to low temperature. These discoveries enhance our comprehension of how plants synchronize the RH growth in response to variations in temperature and nutrient availability at the cellular level.
