The study aimed to determine how the physiological responses of the sunflower (Helianthus annuus L.) plant were affected by prolonged drought stress, salinity stress, and boron application, as well as to assess the recovery dynamics following re-watering. The experimental design included well-watered (WW 80% watering), drought stress (DS, 20% watering) salinity stress (SS, 0 control and 13 dS m-1), boron toxicity (Na2O5B2O3.10H2O, at different doses of 0 and 8 mg L-1) and re-watering after a long-term period of drought stress (24 days). The well-irrigated (80% WW) treatment, which included all factors as a the non-stressed control treatment during the experiment was carried out with five replications. Morphological, physiological and biochemical analyses of plants were measured at four time points: at the 10th and 24th days after the onset of the drought stress period and after re-watering, at 2nd and 7th days following. The relative membrane permeability was increased and relative water content was decreased because drought and salinity stress limited water availability and caused an imbalance in the water status of the leaves and stem of the plant. Even though high levels of Na+ and Cl- ions interfered with essential nutrient uptake under drought stress and boron application, Ca+2 ion levels in the leaves significantly increased in the leaves of plants in areas treated with drought, salt, and boron after re-watering. Extended or intense drought and salinity conditions harmed the phloem and xylem tissue cells of the stem by changing cell size and density, which in turn disrupted biochemical processes, including the functioning of water channels under challenging circumstances. Particularly under conditions of salt and drought stress, the vascular bundles in the plant stem were observed to either shrink significantly or assume an irregular shape. Long-term drought reduced relative water content (RWC) values, resulting in plant dehydration and increased osmotic pressure (RMP) in leaf cells, further exacerbated by salinity and drought stress. The plant attempted to regain some of its characteristics in response to these severe stress conditions after re-watering. However, 24 days after the long dry period, even if watering was re-applied, the growth power of the plant was reduced due to the disturbance in membrane permeability as a result of excessive cell damage.

Simple Summary The current review article focused on the complex interaction between plant hormones and transcription factors in the response to salt stress, a pressing global issue that has a considerable effect on agricultural productivity. The study unveils the effect of salt stress on the ion equilibrium and triggers a cascade of molecular reactions that have a response to plant growth and development. Transcription factors are essential in regulating gene expression under salt stress. The transcription factors function in collaboration with hormones to regulate environmental stress responses. The study sheds light on the underlying regulatory networks, thus providing crucial knowledge for cultivating salt-tolerant crops through selective breeding and genetic engineering. This review significantly contributes to the establishment of sustainable agricultural techniques, which are essential in addressing the increasing problem of soil salinization and securing global food security.Abstract The negative impacts of soil salinization on ion homeostasis provide a significant global barrier to agricultural production and development. Plant physiology and biochemistry are severely affected by primary and secondary NaCl stress impacts, which damage cellular integrity, impair water uptake, and trigger physiological drought. Determining how transcriptional factors (TFs) and hormone networks are regulated in plants in response to salt stress is necessary for developing crops that tolerate salt. This study investigates the complex mechanisms of several significant TF families that influence plant responses to salt stress, involving AP2/ERF, bZIP, NAC, MYB, and WRKY. It demonstrates how these transcription factors (TFs) help plants respond to the detrimental effects of salinity by modulating gene expression through mechanisms including hormone signaling, osmotic stress pathway activation, and ion homeostasis. Additionally, it explores the hormonal imbalances triggered by salt stress, which entail complex interactions among phytohormones like jasmonic acid (JA), salicylic acid (SA), and abscisic acid (ABA) within the hormonal regulatory networks. This review highlights the regulatory role of key transcription factors in salt-stress response, and their interaction with plant hormones is crucial for developing genome-edited crops that can enhance agricultural sustainability and address global food security challenges.