Physiological requirements for wheat ideotypes in response to drought threat. Biochemical and physiological studies on drought influence on plants are systematically summarized in review papers (Foyer and Noctor 2. Jogaiah et al. 2. Labudda and Azam 2. Reynolds and Tuberosa 2. Much research was based on Arabidopsis thaliana as a model organism. Very often, the experiments involved drought treatment under conditions that were definitely different from the field conditions (Claeys et al.

Vadez et al. 2. 01. In the case of higher plants, including crops, the phenomena of dehydration tolerance and dehydration avoidance are types of reactions to water shortage in arable soil. These two responses to drought stress are complementary (Levitt 1. The terms: “drought stress avoidance and tolerance” are fundamental and relevant to the physiological status of plants in the conditions of water shortage.

The term “drought resistance” is a common working phrase used, i. Blum 2. 01. 1b). Water shortage induces dysfunction of plant vital processes leading to the inhibition of growth and development and, in case of long- term water deficit, to tissue decay. It is accompanied by the reduction of water potential and cellular activity, turgor and cell volume reduction, inducing increase of osmoprotectant concentration, changes in macromolecule structure and spatial relationship between cellular compartments accompanied by reduced transpiration and photosynthesis. These processes often result in reduced crop yield which is unacceptable from the economic point of view (Kacperska 2. Maseda and Ferniez 2.

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Richards et al. 2. At the biochemical level, dehydration- tolerant plants are characterized by: (1) more effective identification and transduction of signals (including ABA) (Cutler et al. Pantin et al. 2. 01. Grudkowska and Zagda.

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Zhang et al. 2. 00. LEA proteins (abundant late embryogenesis) interacting primarily with trehalose (Iturriaga 2. Caramelo and Iusem 2. Hussain et al. 2. Jaleel et al. 2. 00. Pinheiro and Chaves 2.

Atkin and Macherel 2. Zagda. 2. 01. 0). At the molecular level, drought induces the expression of several genes including those encoding protective metabolites (i. LEA proteins, Hsp (Heat shock proteins), proline, glycine betaine); genes for sugar osmoprotectant (mannitol and trehalose) synthesis; genes for enzymes maintaining the correct cell membrane structure, e. Among genes encoding signal transductors, kinase genes and genes for transcription factors encoding proteins from the following families can be found: DREB (dehydration response elements binding), ERF (ethylene responsive transcription factor), WRKY (transcription factors with the conserved amino acid sequence: WRKYGQK), MYB (MYelo. Blastosis), b. HLH (basic Helix Loop Helix), b. ZIP (basic Leucine Zipper Domain), NAC (transcription factors of Nam, Ataf.

Cuc. 2, genes possessing common domain) and genes encoding zinc finger protein family (Gosal et al. Regulation of drought- induced modifications of gene expression by RNAi was also studied (Bhatnagar- Mathur et al. The genes for phytohormone synthesis and phytohormone- dependent stress defense signaling such as ABA and gibberellin, jasmonic acid and brassinosteroids were also identified to be engaged in drought resistance regulation (Fleury et al.

Hu and Xiong 2. 01. Shinozaki and Yamaguchi- Shinozaki 2. Zhou et al. 2. 01. Despite muli- year studies on plant genomes, proteomes and metabolomes, the data regarding mechanisms of plant tolerance to environmental stresses are still insufficient for a complete understanding of the processes determining stress tolerance and consequently for economically successful genetic manipulations (Atkinson and Urwin 2. Bhatnagar- Mathur et al. Rybka 2. 01. 1; Tardieu 2. Vadez et al. Legitimizations for economically viable genetic manipulation are recent results of studies on DREB1.

A transgenic groundnut (Vadez et al. Biochemical theoretical achievements are reflected in practice by detection of higher ABA level in leaves of drought- tolerant near isogenic lines (NILs) of pearl millet (Kholov. Vadez et al. However, the extraction of genetic components of complex qualitative traits is still laborious and expensive, since the diversity of phenotypes resulting from complex interaction between plant and environment is difficult to describe due to the multitude of variables in the .

This brings a new meaning to phenotypes in field observation as well as in the frames of functional genomics and phenomics studies (Araus and Cairns 2. Chern et al. 2. 01.

Craufurd et al. 2. Schoppach and Sadok 2. Vadez et al. 2. 00. Recently, Vadez and co- workers (2.

Water- capturing process related to root structure and water conservation for growth phases most sensitive to drought are the key factors guaranteeing effective crop production in drought- threatened areas. The mechanisms influencing root water uptake and its usage by shoots are highly complex (Passioura 2. Watch Full Abundant Acreage Available (2017) 2002. Although cultivars with deeper roots with proper architecture are often considered to be more water stress tolerant, it is not a sufficient selection trait to ensure achieving the breeding goal.

Besides positive correlations found between root structure and water extraction, there are some reports showing a poor relation between roots and water uptake among grain crops and legumes (Vadez et al. This controversy partially results from technical difficulties concerning root studies as well as non- uniform water availability in the soil. Until new experimental methods are introduced, the non- destructive measurements of water uptake at consecutive growth phases, using new lysimetric methods, seem to be an easier target for studies on relations between water capture and crop yield (Vadez et al. It is postulated that genotype adaptation to water shortage is beneficial if this genotype is able to take advantage of small amounts of water available at critical stages of growth. The grain filling period was confirmed experimentally as the important one (Christopher et al. Vadez et al. 2. 01.

Wasson et al. Ten millimeters of subsoil water absorbed after anthesis by wheat roots, between depths of 1. Water extraction by roots at later stages of cereal plants growth would be possible, if roots continued to grow till that time, a feature which should be screened in breeding processes. Such an approach to the problem of water uptake by the roots is appropriate only for soil profiles which are sufficiently deep and rich in water. Water extraction at later stages appears to be dependent on water- saving mechanisms at earlier stages of plant growth and on shoot water demand.

Since root and shoot growth are under the same genetic control, faster root growth can cause quick exploration of soil water. For this reason, faster rate of root growth should be an important trait for genotypes extracting all soil- available water prior to maturity in climate conditions of short rainfall duration or in areas with deep soil profiles rich in water in deeper layers (Vadez et al.

Despite the difficulties in testing and correlation with the yield not always being observed, roots seem to be the force of the second green revolution which will increase the crop yield to feed future generations (Gewin 2. Water conservation on plant and canopy level is the second most important factor of field water stress tolerance underlined by Vadez and co- authors (2. The rate of leaf canopy development and the leaf area at a time close to anthesis are crucial components for determining water usage. Fast canopy development reduces soil transpiration and thereby water loss during the vegetation (Blum 2. On the other hand, it was shown that genotypes of legume, such as chickpea, peanut, cowpea as well as sorghum, tolerant to water stress, have smaller leaf canopy at the vegetative stage.

Experiment on maize showed leaf size reduction under water pressure deficit and also under low vapor pressure deficit, documenting hydraulic control of leaf development.