Mitra Jabbari, Barat Ali Fakheri, Reza Aghnoum, Nafiseh Mahdi Nezhad, Reza Ataei, Zahra Koochakpour


Association mapping has proven to be a powerful approach for dissecting the genetic basis of complex traits. In this study, QTLs controlling flag leaf characteristics under drought stress were detected in a set of 148 modern spring barley cultivars using AM analysis. Flag leaf length (FLL), flag leaf width (FLW), relative water content (RWC), chlorophyll content, and maximum quantum efficiency of PSII (Fv/Fm) which are important in photosynthetic rate, were evaluated under normal irrigation and drought stress conditions at grain filling stage. Population structure was estimated using Structure2.3 and linkage disequilibrium (LD) was estimated by the ‘Full Matrix LD’ using Tassel5.0. Significant marker/trait associations were investigated based on K-Q matrix using Tassel3.0. The analysis of population structure divided the cultivars into two sub-groups. Significant LD values (P < 0.01) between polymorphic sites with regions of high and low LD were observed. A total of 84 significant putative genomic regions were identified, which delineated into 37 QTLs under two water treatments. Two stable QTLs on 2H and 3H were detected for FLL in drought stress treatment. A QTL for FLL were detected on 2H in normal treatment, which alone explained around 11% of phenotypic variance of FLL. This QTL was also associated with the expression of FLW and explained around 7.5% of phenotypic variance. The results suggest that major loci are located on chromosomes 2H, 3H, 4H and 5H involved in the development of flag leaf characteristics and could be used as selection criteria in barley breeding for drought tolerance.

Texto completo:

PDF (English)


(I) Aghnoum, R., Marcel, T.C., Johrde, A., Pecchioni, N., Schweizer, P., Niks, R.E., 2010. Basal host resistance of barley to powdery mildew: connecting quantitative trait loci and candidate genes. Molecular Plant-Microbe Interactions, 23, 91-102.

(II) Agrama, H.A., Eizenga, G.C., Yan, W., 2007. Association mapping of yield and its components in rice cultivars. Molecular Breed, 19, 341-356.

(III) Ahmad, M.A, Khan, S.H, Khan, A.S, Kazi, A.M, Basra, S.M.A., 2014. Identification of QTLs for drought tolerance traits on wheat chromosome 2A using association mapping. International Journal of Agricultural Biology, 16, 862-870.

(IV) Alizade, A., 2002. Soil, Water and Plants Relationship, third ed. Emam Reza University Press, Mashhad, Iran, ISBN, 964-6582-21-4.

(V) Araus, J.I., Bort, J., Ceccarelli, S., Grando, S., 1997. Relationship between leaf structure and carbon isotope discrimination in field grown barley. Plant Physiology and Biochemistry, 35(7), 533-541.

(VI) Arnott, A., Wapling, J., Mueller, I., Ramsland, P.A., Siba, PM., Reeder, J.C., Barry, A.E., 2014. Distinct patterns of diversity, population structure and evolution in the AMA1 genes of sympatric Plasmodium falciparum and Plasmodium vivax populations of Papua New Guinea from an area of similarly high transmission. Malaria Journal, 13, 233.

(VII) Arriagada, O., Mora, F., Quitral, Y., Del Pozo, A., 2017. Identification of QTL underlying agronomic, morphological and physiological traits in barley under rainfed conditions using SNP markers. Acta Scientiarum. Agronomy Maringá, 39(3), 321-329.

(VIII) Berdahl, J.D., Rasmusson, D.C., Moss, D.N., 1972. Effects of leaf area on photosynthetic rate, light penetration, and grain yield in barley. Crop Science, 12, 177-80.

(IX) Bimpong, I.K., Serraj, R., Chin, J.H., Ramos, J., Mendoza, E.M.T., Hernandez, J.E., Mendioro, M.S., Brar, D.S., 2011. Identification of QTLs for Drought-Related Traits in Alien Introgression Lines derived from Crosses of Rice (Oryza sativa cv. IR64) × O. glaberrima under Lowland Moisture Stress. Journal of Plant Biology, 54, 237-250.

(X) Brachi, B., Meyer, C.G., Villoutreix, R., Platt, A., Morton, T.C., Roux, F., Bergelson, J., 2015. Coselected genes determine adaptive variation in herbivore resistance throughout the native range of Arabidopsis thaliana. Proceedings of the National Academy of Sciences USA, 112, 4032-4037.

(XI) Bradbury, P.J., Zhang, Z., Kroon, D.E., Casstevens, T.M., Ramdoss, Y., Buckler, E.S., 2007. TASSEL: software for association mapping of complex traits in diverse samples. Bioinformatics, 23, 2633-2635.

(XII) Breseghello, F., Sorrells, M.E., 2006. Association mapping of kernel size and milling quality in wheat (Triticum aestivum L.) cultivars. Genetics, 172, 1165-1177.

(XIII) Chan, E.K., Rowe, H.C., Kliebenstein, D.J., 2010. Understanding the evolution of defense metabolites in Arabidopsis thaliana using genome-wide association mapping. Genetics, 185(3), 991-1007.

(XIV) Chen, C.X., Li, S.C., Wang, S.Q., Liu, H.N., Deng, Q.M., Zheng, A.P., et al., 2011. Assessment of the Genetic Diversity and Genetic Structure of Rice Core Parent Guichao 2, its Parents and Derivatives. Journal of Plant Science, 6, 66-76.

(XV) Comadran, J., Russell, J.R., Booth, A., Pswarayi, A., Ceccarelli, S., Grando, S., Stanca, A.M., Pecchioni, N., Akar, T., et al., 2011. Mixed-model association scans of multi-environmental trial data reveal major loci controlling yield and yield related traits in Hordeum vulgare in Mediterranean environments. Theoretical and Applied Genetics, 122, 1363-1373.

(XVI) Comadran, J., Thomas, W.T.B., van Eeuwijk, F.A., Ceccarelli, S., Grando, S., Stanca, A.M., et al., 2009. Patterns of genetic diversity and linkage disequilibrium in a highly structured Hordeum vulgare association mapping population for the Mediterranean basin. Theoretical and Applied Genetics, 119, 175-187.

(XVII) Diab, A.A., Teulat-Merah, B., This, D., Ozturk, N.Z., Benscher, D., Sorrells, M.E., 2004. Identification of drought-inducible genes and differentially expressed sequence tags in barley. Theoretical and Applied Genetics, 109, 1417-1425.

(XVIII) Elberse, I.A.M., Vanhala, T.K., Turin, J.H.B., Stam, P., van Damme, J.M.M., van Tienderen, P.H., 2004. Quantitative trait loci affecting growth-related traits in wild barley (Hordeum spontaneum) grown under different levels of nutrient supply. Heredity, 93, 22-33.

(XIX) Fan, Y., Shabala, S., Ma, Y., Xu, R., Zhou, M., 2015. Using QTL mapping to investigate the relationships between abiotic stress tolerance (drought and salinity) and agronomic and physiological traits. BMC Genomics, 16(1).

(XX) Faraloni, C., Cutino, I., Petruccelli, R., Leva, A.R., Lazzeri, S., Torzillo, G., 2011. Chlorophyll fluorescence technique as a rapid tool for in vitro screening of olive cultivars (Olea europaea L.) tolerant to drought stress. Environmental and Experimental Botany, 73, 49-56.

(XXI) Forster, B.P., Ellis, R.P., Moir, J., Talame, V., Sanguineti, M.C., Tuberosa, R., et al., 2004. Genotype and phenotype associations with drought tolerance in barley tested in North Africa. Annual Apply Biology, 144, 157-68.

(XXII) Genty, B., Briantais, J., Baker, N., 1989. The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochimica et Biophysica Acta, 990, 87-92.

(XXIII) González, A., Martín, I., Ayerbe, L., 2008. Yield and osmotic adjustment capacity of barley under terminal water‐stress conditions. Journal of Agronomy Crop Science, 194, 81-91.

(XXIV) Gregoriou, K., Pontikis, K., Vemmos, S., 2007. Effects of reduced irradiance on leaf morphology, photosynthetic capacity, and fruit yield in olive (Olea europaea L.). Photosynthetica, 45(2), 172-181.

(XXV) Guo, P., Baum, M., Varshney, R., Graner, A., Grando, S., Ceccarelli, S., 2008. QTLs for chlorophyll and chlorophyll fluorescence parameters in barley under post-flowering drought. Euphytica, 163(2), 203-214.

(XXVI) Gyenis, L., Yun, S.J., Smith, K.P., Steffenson, B.J., Bossolini, E., Sanguineti, M.C., et al., 2007. Genetic architecture of quantitative trait loci associated with morphological and agronomic trait differences in a wild by cultivated barley cross. Genome, 50, 714-23.

(XXVII) Hasan, M., Friedt, W., Pons-Kuhnemann, J., Freitag, N.M., Link, K., Snowdon, R.J., 2008. Association of gene-linked SSR markers to seed glucosinolate content in oilseed rape (Brassica napus ssp napus). Theoretical and Applied Genetics, 116, 1035-1049.

(XXVIII) Haseneyer, G., Stracke, S., Paul, C., Einfeldt, C., Broda, A., Piepho, H.P., Graner, A., Geiger, H.H., 2010. Population structure and phenotypic variation of a spring barley world collection set up for association studies. Plant Breeding, 129(3), 271-279.

(XXIX) Horsley, R.D., Franckowiak, J.D., Schwarz, P.B., 2009. Barley, in: Carena MJ, editor. Cereals. US: Springer, p. 227-50.

(XXX) Igartua, E., Casas, A.M., Ciudad, F., Montoya, J.L., Romagosa, I., 1999. RFLP markers associated with major genes controlling heading date evaluated in a barley germ plasm pool. Heredity, 83, 551-559.

(XXXI) Jannink, J.L., Walsh, B., 2002. Association mapping in plant populations, in: Kang M.S. (Ed.), Quantitative Genetics, Genomics and Plant Breeding. CABI Publishing Oxon UK, p. 59-68.

(XXXII) Kerwin, R., Feusier, J., Corwin, J., Rubin, M., Lin, C., Muok, A., et al., 2015. Natural genetic variation in Arabidopsis thaliana defence metabolism genes modulates field fitness. eLife, 4, e05604.

(XXXIII) Khaleghi, E., Arzani, K., Moallemi, N., Barzegar, M., 2012. Evaluation of Chlorophyll Content and Chlorophyll Fluorescence Parameters and Relationships between Chlorophyll a, b and Chlorophyll Content Index under Water Stress in Olea europaea cv. Dezful. World Academy of Science, Engineering and Technology, 68.

(XXXIV) Kraakman, A.T.W., Martı’nez, F., Mussiraliev, B., van Eeuwijk, F.A., Niks, R.E., 2006. Linkage disequilibrium mapping of morphological, resistance, and other agronomically relevant traits in modern spring barley cultivars. Molecular Breeding, 17, 41-58.

(XXXV) Kraakman, A.T.W., Niks, R.E., van den Berg, P.M.M.M., Stam, P., Van Eeuwijk, F.A., 2004. Linkage disequilibrium mapping of yield and yield stability in modern spring barley cultivars. Genetics, 168, 435-446.

(XXXVI) Krause, G.H., Weis, E., 1991. Chlorophyll fluorescence and photosynthesis: the basics. Annual review of plant physiology and plant molecular biology, 42, 313-349.

(XXXVII) Lakew, B., Henry, R.J., Ceccarelli S, Grando S, Eglinton J, Baum M., 2013. Genetic analysis and phenotypic associations for drought tolerance in Hordeum spontaneum introgression lines using SSR and SNP markers. Euphytica, 189, 9-29.

(XXXVIII) Liu, L., Sun, G., Ren, X., Li, C.H., Sun, D., 2015. Identification of QTL underlying physiological and morphological traits of flag leaf in barley. BMC Genetics, 16, 29.

(XXXIX) Maccaferri, M., Sanguineti, M.C., Demontis, A., El-Ahmed, A., Garcia del Moral, L., Maalouf, F., et al., 2011. Association mapping in durum wheat grown across a broad range of water regimes. Journal of Experimental Botany, 62(2), 409-438.

(XL) Mariey, S.A., Mohamed, M.N., Khatab, I.A., El-Banna, A.N., Abdel Khalek, A.F., Al-Dinary, M.E., 2013. Genetic Diversity Analysis of Some Barley Genotypes for Salt Tolerance Using SSR Markers. Journal of Agricultural Science, 5(7).

(XLI) Matin, M.A., Brown, J.H., Ferguson, H., 1989. Leaf water potential, relative water content, and diffusive resistance as screening techniques for drought resistance in barley. Agronomy Journal, 81, 100-5.

(XLII) Melgar, J.C., Guidi, L., Remorini, D., Agati, G., Degli’innocenti, E., Castelli, S., et al., 2009. Antioxidant defences and oxidative-damage in salt-treated olive plants under contrasting sunlight irradiance. Tree Physiology, 29, 1187-1198.

(XLIII) Mir, R.R., Zaman-Allah, M., Sreenivasulu, N., Trethowan, R., Varshney, R.K., 2012. Integrated genomics, physiology and breeding approaches for improving drought tolerance in crops. Theoretical Applied Genetics, 125, 625-645.

(XLIV) Mora, F., Quitral, Y.A., Matus, I., Russell, J., Waugh, R., del Pozo, A., 2016. SNP-Based QTL Mapping of 15 Complex Traits in Barley under Rain-Fed and Well-Watered Conditions by a Mixed Modeling Approach. Front Plant Science, 7, 909.

(XLV) Mueller, J.C., 2004. Linkage disequilibrium for different scales and applications. Briefings in Bioinformatics, 5, 355-364.

(XLVI) Nei, M., Li, H., 1979. Mathematical model for studying genetic variation in terms restriction endonucleases. Proceedings of the National Academy of Sciences, 76, 5269-5273.

(XLVII) Oraguzie, N.C., Wilcox, P.L., 2007. An overview of association mapping, in: Oraguzie NC, Rikkerink EHA, Gardiner SE (eds). Association mapping in plants. Springer-Verlag, New York, p. 1-10.

(XLVIII) Pasam, R.K., Sharma, R., Malosetti, M., Eeuwijk, F.A.V., Haseneyer, G., Kilian, B., Graner, A., 2012. Genome-wide association studies for agronomical traits in a worldwide spring barley collection. BMC Plant Biology, 12, 16.

(XLIX) Pritchard, J.K., Wen, X., Falush, D., 2010. Documentation for structure software: Version 2.3.

(L) Roy, J.K., Smith, K.P., Muehlbauer, G.J., Chao, S., Close, T.J., Steffenson, B.J., 2010. Association mapping of spot blotch resistance in wild barley. Molecular Breeding, 26, 243-256.

(LI) Shahraki, H., Fakheri, B.A., 2016. QTLs Mapping of Morpho-Physiological Traits of Flag Leaf in Steptoe × Morex Doubled Haploid Lines of Barley in Normal and Salinity Stress Conditions. International Journal of Farming and Allied Sciences, 5(5), 356-362.

(LII) Somers, D.J., Banks, T., Depauw, R., Fox, S., Clarke, J., Pozniak, C., McCartney, C., 2003. Genome wide linkage disequilibrium analysis in bread wheat and durum wheat. Genome, 50, 557-567.

(LIII) Teng, S., Qian, Q., Zeng, D., Kunihiro, Y., Fujimoto, K., Huang, D., et al., 2004. QTL analysis of leaf photosynthetic rate and related physiological traits in rice (Oryza sativa L). Euphytica, 135, 1-7.

(LIV) Teulat, B., Zoumarou-Wallis, N., Rotter, B., Salem, M.B., Bahri, H., This, D., 2003. QTL for relative water content in field-grown barley and their stability across Mediterranean environments. Theoretical Applied Genetics, 108, 181-8.

(LV) This, D., Borries, C., Souyris, I., Teulat, B., 2000. QTL study of chlorophyll content as a genetic parameter of drought tolerance in barley. Barley Genetics Newsletter, 30, 20-23.

(LVI) Thornsberry, J., Goodman, M., Doebley, J., Kresovich, S., Nielsen, D., 2001. Dwarf8 polymorphisms associate with variation in flowering time. Nature Genetics, 28, 286-289.

(LVII) Tungland, L., Chapko, L.B., Wiersma, J.V., Rasmusson, D.C., 1987. Effect of erect leaf angle on grain yield in barley. Crop Science, 27(1), 37-40.

(LVIII) Wójcik-Jagła, M., Rapacz, M., Tyrka, M., Kościelniak, J., Crissy, K., Żmuda, K., 2013. Comparative QTL analysis of early short-time drought tolerance in Polish fodder and malting spring barleys. Theoretical Applied Genetics, 126, 3021-3034.

(LIX) Xue, D., Chen, M., Zhou, M., Chen, S., Mao, Y., Zhang, G., 2008. QTL analysis of flag leaf in barley (Hordeum vulgare L.) for morphological traits and chlorophyll content. Journal of Zhejiang University Science B, 9, 938-43.

(LX) Yap, T.C., Harvey, B.L., 1972. Relations between grain yield and photosynthetic parts above the flag leaf node in barley. Canadian Journal of Plant Science, 52, 241-246.

(LXI) Yin, X., Kropff, M.J., Stam, P., 1999. The role of ecophysiological models in QTL analysis: the example of specific leaf area in barley. Heredity, 82, 415-421.

(LXII) Yu, J., Pressoir, G., Briggs, W.H., Bi, I.V., Yamasaki, M., Doebley, J.F., et al., 2005. A unified mixed_model method for association mapping that accounts for multiple levels of relatedness. Nature Genetics, 38, 203-208.

(LXIII) Zhao, J., Sun, H., Dai, H., Zhang, G., Wu, F., 2010. Difference in response to drought stress among Tibet wild barley genotypes. Euphytica, 172, 395-403.

(LXIV) Zhu, C., Gore, M., Buckler, E.S., Yu, J., 2008. Status and prospects of association mapping in plants. Plant Genome, 1, 5-20.



  • Não há apontamentos.


Licença Creative Commons
Esta obra está licenciada sob uma licença Creative Commons Atribuição - Não comercial - Compartilhar igual 4.0 Internacional.