Effects of Drought Stress on Some Anatomical Characteristics of Barley Leaves

Document Type : Research Paper


Assistant Professor, Department of Agriculture, Miandoab Branch, Islamic Azad University, Miandoab, Iran


In this research, effect of drought stress on leaf characteristics was investigated in the experimental field of Miandoab Azad University, Iran using four facultative barley cultivars. Two separate experiments were conducted in pot and field conditions. In both experiments a factorial arrangement was used and the treatments were completely randomized in four replications. In each experiment half of the experimental units were drought stressed and the other half were irrigated normally. For the pot experiment, plants were sampled at fourth leaf stage in order to obtain winter leaves. For the field experiment, when plants reached at late stem elongation stage, penultimate leaves were sampled and used for preparing microscopic slides.
Xylem and phloem diameter and mesophyll, bundle sheath and epidermal cells area were measured in the transverse sections prepared from middle parts of the leaves. Significant differences were observed among genotypes under drought stress in terms of leaf characteristics. Results also showed that drought stress changes the diameter and the surface area of the cells. However, the changes were not the same in the winter and spring leaves. The changes in some cases such as diameter of xylem vessels were considerable. For example, in the cultivars Sahra and Jonob, the winter leaves had bigger xylem vessels under drought stress as compared to the normal condition while in the spring leaves the xylem diameter was smaller under the same condition.


Baik BK and Ullrich SE, 2008. Barley for food: characteristics, improvement and renewed interest. Journal of Cereal Science 48: 233-242.
Bohnert HJ and Jensen RG, 1996. Strategies for engineering water stress tolerance in plants. Trends in Biotechnology 14: 89-97.
Bongi G, Mencuccini M and Fontanazza G, 1987. Photosynthesis of olive leaves: effect of light flux density, leaf age, and temperature and H2O vapor pressure deficit on gas exchange. Journal of the American Society for Horticultural Science 112: 143-148.
Bosabalidis AM and Kofidis G, 2002. Comparative effects of drought stress on leaf anatomy of two olive cultivars. Plant Science 163(2): 375-379.
Chartzoulakis K, Patakas A, Kofidis G, Bosabalidis A and Nastou A, 2002. Water stress affects leaf anatomy, gas exchange, water relations and growth of two avocado cultivars. Scientia Horticulturae 95: 39-50.
Chaves MM and Oliveira MM, 2004. Mechanisms underlying plant resilience to water deficits: prospects for water-saving agriculture. Journal of Experimental Botany 55: 2365-2384.
Cochard H, Nardini A and Coll L, 2004. Hydraulic architecture of leaf blades: where is the main resistance? Plant, Cell and Environment 27: 1257-1267.
Culter JM, Rains DW and Loomis RS, 1977. The importance of cell size in the water relations of plants. Physiologia Plantarum 40: 255-260.
Dias J, Pimenta JA, Medri ME, Boeger MRT and de Freitas CT, 2007. Physiological aspects of sun and shade leaves of Lithraea molleoides (Vell.) Engl. (Anacardiaceae). Brazilian Archives of Biology and Technology 50(1): 91-99.
Djanaguiramana M, Prasada PVV, Boyle DL and Schapaugha WT, 2011. High-temperature stress and soybean leaves: leaf anatomy and photosynthesis. Crop Science 51: 2125-2531.
Drake PL, Froend RH and Franks PJ, 2013. Smaller, faster stomata: scaling of stomatal size, rate of response and stomatal conductance. Journal of Experimental Botany 64: 495-505.
Evans JR, Caemmerer SV, Setchell BA and Hudson GS, 1994. The relationship between CO2 transfer conductance and leaf anatomy in transgenic tobacco with a reduced content of rubisco. Australian Journal of Plant
Physiology 21(4): 475-495.
Fitter A and Hay R, 2002. Environmental Physiology of Plants. Academic Press. 367 pages.
Fricke W and Flowers TJ, 1998. Control of leaf cell elongation in barley. Generation rates of osmotic pressure and growth-associated water potential gradients. Planta 206: 53-65.
Guerfel M, Baccouri O, Boujnah D, Chaidi W and Zarrouk M, 2009. Impact of water stress on gas exchange, water relations, chlorophyll content and leaf structure in the two main Tunisian   olive (Oleaeuropaea L.) cultivars. Scientia Horticulturae 119: 257-263.
Hallik L, Niinemets U and Wright IJ, 2009. Are species shade and drought tolerance reflected in leaf-level structural and functional differentiation in Northern Hemisphere temperate woody flora? New Phytologist 184: 257-274.
Harb A, Krishnan A, Madana M, Ambavaram R and Pereira A, 2010. Molecular and physiological analysis of drought stress in Arabidopsis reveals early responses leading to acclimation in plant growth. Journal of Plant Physiology 154: 1254-1271.
Hu Y, Fromm J and Schmidhalter U, 2005. Effect of salinity on tissue architecture in expanding wheat leaves. Planta 220: 838-848.
Hu Y, Schnyder H and Schmidhalter U, 2000. Carbohydrate accumulation and partitioning in elongating leaves of wheat in response to saline soil conditions. Australian Journal of Plant
Physiology 27: 363- 370.
Japp AP and Newman I, 1987. Morphological and anatomical effects of severe drought on the roots of Lolium perenne  L. New Phytologist 105: 393-402.
Jones MB, Leafe LE and Stiles W, 1980. Water stress in field grown perennial ryegrass. Its effects on leaf water status, stomatal resistance and leaf morphology. Annals of Applied Biology 96: 103-10.
Kazemi Arbat H, 2005. Morphology and Anatomy of Cereal Crops. Volume 2. University of Tabriz Press, Tabriz, Iran (In Persian).
Kiseleva LS and Kaminskaya DA, 2002. Hormonal regulation of assimilate utilization in barley leaves in relation to the development of their source function. Russian Journal of Plant Physiology 49(4): 535-540.
Krcek M, Slamka P, Olsovska K, Brestic M and Bencikova M, 2008. Reduction of drought stress effect in spring barley (Hordeum vulgare L.) by nitrogen fertilization. Plant, Soil and Environment 54(1): 7-13.
Lee KW, Choi GJ, Kim KY, Ji HC, Zaman R and Lee SH, 2011. Identification of drought induced differentially expressed genes in barley leaves using the annealing control-primer-based GeneFishing technique. Australian Journal of Crop Science 5(11): 1364-1369.
Mansoor U, Ashraf M and Rao AR, 2002. Variation in leaf anatomy in wheat germplasm from varying drought-hit habitats. International Journal of Agriculture and Biology 4: 12-16.
McClendon JH, 1992. Photographic survey of the occurrence of bundle sheath extension in deciduous dicots. Plant Physiology 99: 1677-1679.
McKim HL, Walsh JE and Arion DN, 1980. Review of Techniques for Measuring Soil Moisture in Situ. United States Army Corps of Engineers. Cold Regions Research and Engineering Laboratory, Hanover, New Hampshire, USA. Report 80-31.
Mediavilla S, Escudero A and Heilmeier H, 2001. Internal leaf anatomy and photosynthetic resource-use efficiency: interspecific and intraspecific comparisons. Tree Physiology 21: 251-259.
Minchin PEH, Thorpe MR, Farrar JE and Koroleva OA, 2002. Source-sink coupling in young barley plants and control of phloem loading. Journal of Experimental Botany 53(374): 1671-1676.
Niinemets U and Sack L, 2006. Structural determinants of leaf light harvesting capacity and photosynthetic potentials. In: Esser K, Luttge UE, Beyschlag W and Murata J(Eds). Progress in Botany, Vol. 67. Pp. 385-419. Springer-Verlag, Berlin.
Payvandi S, Daly KR, Joes BL, Talboys P and Roose T, 2014. A mathematical model of water and nutrient transport in xylem vessels of a wheat plant. Bulletin of Mathematical Biology 76: 566-596.
Press MC, 1999. The functional significance of leaf structure: a search for generalizations. New Phytologist 143: 213-219.
Rebetzke GJ, Condon AG and Richards RA, 2010. Genomic regions for canopy temperature and their genetic association with stomatal conductance and grain yield in bread wheat (Triticum aestivum L.). Functional Plant Biology 8: 150-154.
Reddy AR, Chiatanya KV and Vivekanandan M, 2004. Drought induced responses of photosynthesis and antioxidant metabolism in higher plants. Journal of Plant Physiology 161: 1189-1202.
Syvertsen JP, Lloyd J, Mcconchiehie C, Kriedemann PE and Farquhar GD, 1995. On the relationship between leaf anatomy and CO2 diffusion through the mesophyll of hypostomatous leaves. Plant, Cell & Environment 18: 149-157.
Talame V, Ozturk NZ, Bohnert HJ and Tuberosa R, 2007. Barley transcript profiles under dehydration shock and drought stress treatments: a comparative analysis. Journal of Experimental Botany 58(2): 229–240.
Vasantha S, Alarmelu S, Hemaprabha G and Shanthi RM, 2005. Evaluation of promising sugarcane genotypes for drought. Journal of Sugar Technology 7(2): 82-83.
Verma V, Foulkes MJ, Worland AJ, Sylvester-Bradley R, Caligari PDS and Snape JW, 2004. Mapping quantitative trait loci for flag leaf senescence as a yield determinant in winter wheat under optimal and drought-stressed environments. Euphytica 135: 255-263.
Zagdanska B and Kozdoj J, 1994. Water stress-induced changes in morphology and anatomy of flag leaf of spring wheat. Acta Societatis Botanicorum Poloniae 63(1): 61–66.
Zagdanska B and Wisniewski K, 1996. Endoproteinase activities in wheat leaves upon water deficit. Acta Biochimica Polonica 43(3): 515–520.