Drought stress definition: Stress is an altered physiological condition caused by factors that tend to disrupt the equilibrium.

The flexibility of normal metabolism allows the response initiation to the environmental changes, which fluctuate regularly and are predictable over daily and seasonal cycles. Plants are frequently exposed to many stresses such as drought, low temperature, salt, flooding, heat, oxidative stress and heavy metal toxicity, while growing in nature. Drought is a meteorological term and is commonly defined as a period without significant rainfall. Generally drought stress occurs when the available water in the soil is reduced and atmospheric conditions cause continuous loss of water by transpiration or evaporation. Drought stress tolerance is seen in almost all plants but its extent varies from species to species and even within species. Water deficit and salt stresses are global issues to ensure survival of agricultural crops and sustainable food production (Jaleel et al., 2007b-e; Nakayama et al., 2007). Drought stress is considered to be a moderate loss of water, which leads to stomatal closure and limitation of gas exchange. Desiccation is much more extensive loss of water, which can potentially lead to gross disruption of metabolism and cell structure and eventually to the cessation of enzyme catalyzed reactions (Smirnoff, 1993; Jaleel et al., 2007d). (Jaleel et al., 2008a-e; Farooq et al., 2008). In plants, a better understanding of the morpho-anatomical and physiological basis of changes in water stress resistance could be used to select or create new varieties of crops to obtain a better productivity under water stress conditions (Nam et al., 2001; Martinez et al., 2007). The reactions of plants to water stress differ significantly at various organizational levels depending upon intensity and duration of stress as well as plant species and its stage of growth (Chaves et al., 2002; Jaleel et al., 2008b). Understanding plant responses to drought is of great importance and also a fundamental part for making the crops stress tolerant (Reddy et al., 2004; Zhao et al., 2008). Term stress (stress factor, stressor) indicates the event.Term strain indicates the state (stress response, state of adaptation) evoked within an organism. Too little water is available in a suitable thermodynamic state.Demand exceeds the supply of water
In nature, water is usually the most limiting factor for plant growth. This is also the case in home or commercial landscapes. If plants do not receive adequate rainfall or irrigation, the resulting drought stress can reduce growth more than all other environmental stresses combined. Drought can be defined as the absence of rainfall or irrigation for a period of time sufficient to deplete soil moisture and injure plants. Drought stress results when water loss from the plant exceeds the ability of the plant's roots to absorb water and when the plant's water content is reduced enough to interfere with normal plant processes. In Florida, plants may frequently encounter drought stress. Rainfall is very seasonal and periodic drought occurs regularly. Because Florida's soils are typically sandy and have low water holding capacity, many plants may experience drought stress

after only a few days without water. During drought, local governments may place restrictions on landscape irrigation in order to conserve potable water, and landscape plants may become subject to drought stress. The use of drought tolerant plants in the landscape can reduce the likelihood of plant injury due to drought stress.

CAUSES
How Long Before Drought Stress Develops?

The time required for drought injury to occur depends on the water-holding capacity of the soil, environmental conditions, stage of plant growth, and plant species. Plants growing in sandy soils with low water-holding capacity are more susceptible to drought stress than plants growing in clay soils. A limited root system will accelerate the rate at which drought stress develops. A root system may be limited by the presence of competing root systems, by site conditions such as compacted soils or high water tables, or by container size (if growing in a container). A plant with a large mass of leaves in relation to the root system is prone to drought stress because the leaves may lose water faster than the roots can supply it. Newly installed plants and poorly established plants may be especially susceptible to drought stress because of the limited root system or the large mass of stems and leaves in comparison to roots.
How Does Environment Affect Drought Stress?

Aside from the moisture content of the soil, environmental conditions of high light intensity, high temperature, low relative humidity and high wind speed will significantly increase plant water loss. The prior environment of a plant also can influence the development of drought stress. A plant that has been drought stressed previously and has recovered may become more drought resistant. Also, a plant that was well-watered prior to drought will usually survive drought better than a continuously drought-stressed plant.

Reasons: Soil dryness Inadequate water uptake by plants in shallow soils

 Osmotic binding in saline soils High evaporation Drought stress develops slowly.Intensity increases with time Stress level, time scale crucial!

EFFECTS
A plant responds to a lack of water by halting growth and reducing photosynthesis and other plant processes in order to reduce water use. As water loss progresses, leaves of some species may appear to change color -- usually to blue-green. Foliage begins to wilt and, if the plant is not irrigated, leaves will fall off and the plant will eventually die. Drought symptoms resemble salt stress because high concentrations of salts in the root zone cause water loss from roots. Close examination of environmental and cultural conditions should help identify the specific proble

Different survival mechanisms of plants at dry sites: 1) Drought escape 2) Dehydration avoidance 3) Dehydration tolerance Drought escape Drought periods must occur at a predictable time Important strategy for mediterranean and monsoon climates, not efficient for Central Europe i) Temporal: Whole life cycle or physiologically active phase shifted to

periods without stress e.g. winter wheat, winter barley well suited for their place of origin (Iraq, Iran; summer drought) Selection of early-ripening genotypes ii) Spatial: Development of water-storing belowground organs e.g. geophytesDrought (dehydration) avoidance Tissues are sensitive to dehydration must maintain high water potentials as long as possible 2 groups of drought avoiders: i) Water savers Conserve water ii) Water spenders Absorb water so fast as to meet transpirational losses Anatomical and morphological traits help the plant to increase water uptake reduce water spendingMorpho-anatomical traits (A) Water uptake is improved (1) extensive root system with large active surface area (2) shoot/root ratio shifted in favour of the roots (B) Water loss is reduced (1) transpiration reduced (timely stomatal closure) (1a) smaller but more densely distributed stomata (2) thick cuticle

(3) epicuticular waxes (4) leaf colour (yellow, glaucous) (5) white hairs on leaves (6) leaf angle (7) leaf rolling (8) plant senescence (8a) leaf senescence (9) leaf sheddingDrought (dehydration) tole

Drought stress is characterized by reduction of water content, diminished leaf water potential and turgor loss, closure of stomata and decrease in cell enlargement and growth (Fig. 1). Severe water stress may result in the arrest of photosynthesis, disturbance of metabolism and finally the death of plant (Jaleel et al., 2008c). Water stress inhibits cell enlargement more than celldivision. It reduces plant growth by affecting various physiological and biochemical processes, such as photosynthesis, respiration, translocation, ion uptake, carbohydrates, nutrient metabolism and growth promoters. Water stress results in stomatal closure and reduced transpiration rates, a decrease in the water potential of plant tissues, decrease in photosynthesis and growth inhibition, accumulation of abscisic acid (ABA), proline, mannitol, sorbitol, formation of radical scavenging compounds (ascorbate, glutathione, -tocopherol etc.), and synthesis of new proteins and mRJNAs. Besides these physiological responses plants also undergo morphological changes. One of the largest is the adaptation of plants and chloroplasts

to high light (sun) and low light (shade exposure). This sun-type or shade-type chloroplast adaptation is also induced by many other stress factors including drought (Lichtenthaler et al., 1981)

Water stress greatly suppresses cell expansion and cell growth due to the low turgor pressure