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. CO2 assimilation by leaves is reduced mainly by stomatal closure, membrane damage and disturbed activity of various enzymes, especially those of CO2 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.

https://hal.archives-ouvertes.fr/hal-00886451/document

Plant drought stress: effects, mechanisms and management

Plant gene responses to changing carbohydrate status can vary markedly Some genes are induced, some are repressed, and others are minimally affected As in microorganisms, sugar-sensitive plant genes are part of an ancient system of cellular adjustment to critical nutrient availability However, in multicellular plants, sugar-regulated expression also provides a mechanism for control of resource distribution among tissues and organs Carbohydrate depletion upregulates genes for photosynthesis, remobilization, and export, while decreasing mRNAs for storage and utilization Abundant sugar levels exert opposite effects through a combination of gene repression and induction Long-term changes in metabolic activity, resource partitioning, and plant form result Sensitivity of carbohydrate-responsive gene expression to environmental and developmental signals further enhances its potential to aid acclimation The review addresses the above from molecular to whole-plant levels and considers emerging models for sensing and transducing carbohydrate signals to responsive genes

Carbohydrate-modulated gene expression in plants.

As the result of intensive research and breeding efforts over the last 20 years, the yield potential and yield quality of cereals have been greatly improved. Nowadays, yield safety has gained more importance because of the forecasted climatic changes. Drought and high temperature are especially considered as key stress factors with high potential impact on crop yield. Yield safety can only be improved if future breeding attempts will be based on the valuable new knowledge acquired on the processes determining plant development and its responses to stress. Plant stress responses are very complex. Interactions between plant structure, function and the environment need to be investigated at various phases of plant development at the organismal, cellular as well as molecular levels in order to obtain a full picture. The results achieved so far in this field indicate that various plant organs, in a definite hierarchy and in interaction with each other, are involved in determining crop yield under stress. Here we attempt to summarize the currently available information on cereal reproduction under drought and heat stress and to give an outlook towards potential strategies to improve yield safety in cereals.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2007.01727.x

The effect of drought and heat stress on reproductive processes in cereals.

Drought tolerance improvement in crop plants: An integrated view from breeding to genomics

Cytokinins (CKs) play a crucial role in various phases of plant growth and development, but the basic molecular mechanisms of their biosynthesis and signal transduction only recently became clear. The progress was achieved by identifying a series of key genes encoding enzymes and proteins controlling critical steps in biosynthesis, translocation, and signaling. Basic schemes for CK homeostasis and root/shoot communication at the whole-plant level can now be devised. This review summarizes recent findings on the relationship between CK structural variation and activity, distinct features in CK biosynthesis between higher plants and Agrobacterium infected plants, CK translocation at whole-plant and cellular levels, and CKs as signaling molecules for nutrient status via root-shoot communication.

CYTOKININS: Activity, Biosynthesis, and Translocation

Continuous increase in the yield of maize (Zea mays L.) in the U.S. Corn Belt has involved an interaction with plant density. A number of contributing traits and mechanisms have been suggested. In this study we used a modeling approach to examine whether changes in canopy and/or root system architecture might explain the observed trends. A maize crop model was generalized so that changes in canopy and root system architecture could be examined. A layered, diurnal canopy photosynthesis model was introduced to predict consequences of change in canopy architecture. A two-dimensional root exploration model was introduced to predict consequences of change in root system architecture. Field experiments were conducted to derive model parameters for the base hybrid (Pioneer 3394). Simulation studies for various canopy and root system architectures were undertaken for a range of sites, soils, and densities. Simulated responses to density compared well with those found in field experiments. The analysis indicated that (i) change in root system architecture and water capture had a direct effect on biomass accumulation and historical yield trends; and (ii) change in canopy architecture had little direct effect but likely had important indirect effects via leaf area retention and partitioning of carbohydrate to the ear. The study provided plausible explanations and identified testable hypotheses for future research and crop improvement effort.

Can Changes in Canopy and/or Root System Architecture Explain Historical Maize Yield Trends in the U.S. Corn Belt?

Water deficit during pollination increases the frequency of kernel abortion in maize (Zea mays L.). Much of the kernel loss is attributable to lack of current photosynthate, but a large number of kernels fail to develop on water-deficient plants even when assimilate supply is increased. We examined the possibility that assimilate utilization by developing ovaries might be impaired at low water potential ([Psi]w). Plants were grown in the greenhouse in 20-L pots containing 22 kg of amended soil. Water was withheld on the first day silks emerged, and plants were hand-pollinated 4 d later when leaf [Psi]w decreased to approximately - 1.8 MPa and silk [Psi]w was approximately -1.0 MPa. Plants were rehydrated 2 d after pollination. The brief water deficit inhibited ovary growth (dry matter accumulation) and decreased kernel number per ear by 60%, compared to controls. Inhibition of ovary growth was associated with a decrease in the level of reducing sugars, depletion of starch, a 75-fold increase in sucrose concentration (dry weight basis), and inhibition of acid invertase (EC 3.2.1.26) activity. These results indicate that water deficits during pollination disrupt carbohydrate metabolism in maize ovaries. They suggest that acid invertase activity is important for establishing and maintaining reproductive sink strength during pollination and early kernel development.

Low Water Potential Disrupts Carbohydrate Metabolism in Maize (Zea mays L.) Ovaries.

Cytokinins are hormones that play an essential role in plant growth and development. The irreversible degradation of cytokinins, catalyzed by cytokinin oxidase, is an important mechanism by which plants modulate their cytokinin levels. Cytokinin oxidase has been well characterized biochemically, but its regulation at the molecular level is not well understood. We isolated a cytokinin oxidase open reading frame from maize (Zea mays), called Ckx1, and we used it as a probe in northern and in situ hybridization experiments. We found that the gene is expressed in a developmental manner in the kernel, which correlates with cytokinin levels and cytokinin oxidase activity. In situ hybridization with Ckx1 and transgenic expression of a transcriptional fusion of the Ckx1 promoter to the Escherichia coli beta-glucuronidase reporter gene revealed that the gene is expressed in the vascular bundles of kernels, seedling roots, and coleoptiles. We show that Ckx1 gene expression is inducible in various organs by synthetic and natural cytokinins. Ckx1 is also induced by abscisic acid, which may control cytokinin oxidase expression in the kernel under abiotic stress. We hypothesize that under non-stress conditions, cytokinin oxidase in maize plays a role in controlling growth and development via regulation of cytokinin levels transiting in the xylem. In addition, we suggest that under environmental stress conditions, cytokinin oxidase gene induction by abscisic acid results in aberrant degradation of cytokinins therefore impairing normal development.

Chris Zinselmeier

Papers

Can Changes in Canopy and/or Root System Architecture Explain Historical Maize Yield Trends in the U.S. Corn Belt?

Low Water Potential Disrupts Carbohydrate Metabolism in Maize (Zea mays L.) Ovaries.

Cytokinin Oxidase Gene Expression in Maize Is Localized to the Vasculature, and Is Induced by Cytokinins, Abscisic Acid, and Abiotic Stress

Improving Drought Tolerance in Maize

The use of gene expression profiling to dissect the stress sensitivity of reproductive development in maize