Insights into Salinity Stress-Induced Morpho-Physiological and Molecular Responses and Nanoparticle- and Nanobiochar-Mediated Tolerance Mechanisms During Seed Germination

Abstract

Soil salinity is a major environmental constraint that threatens global food security by significantly inhibiting seed germination and early seedling establishment. Salinity disrupts all three phases of seed germination: Phase I (imbibition), where reduced water absorption capacity reduces seed hydration and delays metabolic reactivation; Phase II (lag phase), where ionic toxicity and oxidative stress impair enzyme activity, reserve mobilization, and cellular metabolism; and Phase III (radicle protrusion), where limited cell division and length prevent radicle emergence and seedling establishment. These disturbances reduce germination percentage, germination rate, germination index, germination energy, and plant vigor, while increasing average germination time. At the morpho-physiological level, salinity impairs water absorption, membrane stability, photosynthetic pigment accumulation, and root–shoot development. Biochemically, excessive accumulation of reactive oxygen species (ROS), hydrogen peroxide (H2O2), and malondialdehyde (MDA) causes cellular damage and metabolic dysfunction. At the molecular level, salinity alters the expression of the aquaporin gene family (PIPs, TIPs, NIPs, and SIPs), suppresses starch mobilization by reducing α-amylase, enhances abscisic acid (ABA) signaling, and inhibits gibberellic acid (GA) biosynthesis, all of which cause inhibition of germination and early growth. As a result, an effective strategy is needed to improve seed germination under saline conditions. Therefore, the second focus of this review is to critically evaluate the potential of nanoparticles (NPs) and nanobiochar (NBC) as emerging tools to mitigate salinity stress during seed germination. Current evidence suggests that NPs and NBC enhance water absorption, maintain membrane strength, improve nutrient availability, promote antioxidant defense systems, and regulate osmotic adjustment in saline environments. Furthermore, these nanomaterials alter key molecular pathways involved in aquaporin expression, hormonal homeostasis, and reserve mobilization, thereby promoting successful germination and seedling establishment. By combining recent advances in physiological, biochemical, and molecular research, this review provides a comprehensive understanding of salinity-induced germination disruption and highlights the potential of NP- and NBC-based approaches to improve crop establishment under saline conditions.