Plant drought stress: effects, mechanisms and management
Scarcity of water is a severe environmental constraint to plant productivity. Drought-induced loss in crop yield probably exceeds losses from all other causes, since both the severity and duration of the stress are critical. Here, we have reviewed the effects of drought stress on the growth, phenology, water and nutrient relations, photosynthesis, assimilate partitioning, and respiration in plants. This article also describes the mechanism of drought resistance in plants on a morphological, physiological and molecular basis. Various management strategies have been proposed to cope with drought stress. Drought stress reduces leaf size, stem extension and root proliferation, disturbs plant water relations and reduces water-use efficiency. Plants display a variety of physiological and biochemical responses at cellular and whole-organism levels towards prevailing drought stress, thus making it a complex phenomenon. CO(2) assimilation by leaves is reduced mainly by stomatal closure, membrane damage and disturbed activity of various enzymes, especially those of CO(2) fixation and adenosine triphosphate synthesis. Enhanced metabolite flux through the photorespiratory pathway increases the oxidative load on the tissues as both processes generate reactive oxygen species. Injury caused by reactive oxygen species to biological macromolecules under drought stress is among the major deterrents to growth. Plants display a range of mechanisms to withstand drought stress. The major mechanisms include curtailed water loss by increased diffusive resistance, enhanced water uptake with prolific and deep root systems and its efficient use, and smaller and succulent leaves to reduce the transpirational loss. Among the nutrients, potassium ions help in osmotic adjustment; silicon increases root endodermal silicification and improves the cell water balance. Low-molecular-weight osmolytes, including glycinebetaine, proline and other amino acids, organic acids, and polyols, are crucial to sustain cellular functions under drought. Plant growth substances such as salicylic acid, auxins, gibberrellins, cytokinin and abscisic acid modulate the plant responses towards drought. Polyamines, citrulline and several enzymes act as antioxidants and reduce the adverse effects of water deficit. At molecular levels several drought-responsive genes and transcription factors have been identified, such as the dehydration-responsive element-binding gene, aquaporin, late embryogenesis abundant proteins and dehydrins. Plant drought tolerance can be managed by adopting strategies such as mass screening and breeding, marker-assisted selection and exogenous application of hormones and osmoprotectants to seed or growing plants, as well as engineering for drought resistance.