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RESUMEN La Fusariosis de la Espiga de Trigo

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1. Lon G Sisterna M Haidukowsky M Rizzo I 2003 Fusarium graminearum and deoxynivalenol contamination in the durum wheat area of Argentina Microbiol Res 158 1 7 Lu ZX Gaudet D Puchalski B Despins T Frick M y Laroche A 2006 Inducers of resistance reduce common bunt infection in wheat seedlings while differentially regulating defense gene expression Physiol Mol Plant Pathol 67 138 148 Maccaferri M Corneti S Ricci A Stefanelli S Braida E Lancioni P Ammar K Prodi A Pisi A Pascale M Lippolis V Massi A y Tuberosa R 2013 Genetic dissection of Fusarium head blight response in a durum wheat elite population En Abstracts International Symposium Genetics and Breeding of Durum Wheat ENEA Rome 161 Mackintosh CA Lewis J Radmer LE Shin S Heinen SJ Smith LA Wyckoff MN Dill Macky R Evans CK Kravchenko S Baldridge GD Zeyen RJ y Muehlbauer GJ 2007 Overexpression of defense response genes in transgenic wheat enhances resistance to Fusarium head blight Plant Cell Rep 26 4 479 488 Madriz K 2002 Mecanismos de defensa en las interacciones planta pat geno Manejo Integrado de Plagas 3 22 32 Malihipour A Gilbert J Piercey Normore M y Cloutier S 2012 Molecular phylogenetic analysis trichothecene chemotype patterns and variation in aggressiveness of Fusarium isolates causing head blight in wheat Plant Dis 96 1016 1025 Marchionatto JB 1934 Enfermedades del trigo poco conocidas y radicadas
2. Fusariosis de la espiga Fusarium graminearum y Fusarium spp del trigo en la provincia de Entre R os A o 2002 En Actualizaci n T cnica Trigo INTA EEA Paran Serie Extensi n 24 30 34 139 Bibliograf a Gabriel DW 1999 Why do pathogens carry avirulence genes Physiol Mol Plant Pathol 55 205 214 Gabri ls SHEJ Takken FLW Vossen JH de Jong CF Liu Q Turk SCHJ Wachowski LK Peters J Witsenboer HMA de Wit PJGM y Joosten MHAJ 2006 cDNA AFLP combined with functional analysis reveals novel genes involved in the hypersensitive response Mol Plant Microbe Interact 19 567 576 Galich MT 1987 Importancia y difusi n de la fusariosis del trigo en Argentina En Taller sobre la fusariosis de la espiga en Am rica del Sur Ed Kohli MM Encarnaci n Paraguay ISBN 9686127372 9789686127379 Galich MT Galich AN Masiero B Galarza C y Gudelj V 1996 Aspectos del manejo del cultivo de trigo que influyen en la infecci n de Fusarium graminearum Informe T cnico 123 EEA INTA M Ju rez Galich MT 1997 Fusarium Head Blight in Argentina En Fusarium Head Scab Global Status and Future Prospects Eds Dubin HJ Gilchrist L Reeves J y McNab A CIMMYT M xico 19 28 Galich M Alberione E Helguera M y Bainotti C 2006 Characterization and development of Argentinean wheat germplasm with resistance against Fusarium head blight En The Global Fusarium Initiative for International Collaboration Eds Ban T L
3. 1989 Outbreak of Trichothecene Mycotoxicosis Associated with Consumption of Mold Damaged Wheat Products in Kashmir Valley India Lancet 1 35 37 Boutigny AL Richard Forget F y Barreau C 2008 Natural mechanisms for cereal resistance to the accumulation of Fusarium trichothecenes Eur J Plant Pathol 121 4 411 423 Brennan JM Fagan B y Van Maanen A 2003 Studies on in vitro growth and pathogenicity of European Fusarium fungi Eur J Plant Pathol 109 577 587 Breyne P y Zabeau M 2001 Genome wide expression analysis of plant cell cycle modulated genes Curr Opin Plant Biol 4 136 142 Browne R y Cooke B 2004 Development and evaluation of an in vitro detached leaf assay for pre screening resistance to FHB in wheat Eur J Plant Pathol 110 91 102 133 Bibliograf a Browne R y Cooke B 2005 Resistances of wheat to Fusarium spp in an in vitro seed germination assay and preliminary investigations into the relationship with Fusarium head blight resistance Euphytica 141 23 32 Browne RA Murphy JP Cooke BM Devaney D Walsh EJ Griffey CA Hancock JA Harrison SA Hart P Kolb FL McKendry AL Milus EA Sneller C y Van Sanford DA 2005 Evaluation of components of Fusarium head blight resistance in soft red winter wheat germ plasm using a detached leaf assay Plant Dis 89 4 404 411 Browne R 2007 Components of resistance to FHB in wheat detected in a seed germination assay with Microdochium majus and the
4. 2004 Genealogical concordance between the mating type locus and seven other nuclear genes supports formal recognition of nine phylogenetically distinct species within the Fusarium graminearum clade Fungal Genet Biol 41 600 623 O Donnell K Ward TJ Aberra D Kistler HC Aoki T Orwig N Kimura M Bj rnstad S y Klemsdal SS 2008 Multilocus genotyping and molecular phylogenetics resolve a novel head blight pathogen within the Fusarium graminearum species complex from Ethiopia Fung Genet Biol 45 1514 1522 Oliver R Stack R Miller J y Cai X 2007 Reaction of wild emmer wheat accessions to Fusarium head blight Crop Sci 47 893 899 Oliver RE Cai X Friesen TL Halley S Stack RW Xu SS 2008 Evaluation of Fusarium head blight resistance in tetraploid wheat Triticum turgidum L Crop Sci 48 213 222 Ori N Eshed Y Paran I Presting G Aviv D Tanksley S Zamir D y Fluhr R 1997 The RC family from the wilt disease resistance locus I2 belongs to the nucleotide binding leucine rich repeat superfamily of plant resistance genes Plant Cell 9 4 521 532 Otto CD Kianian SF Elias EM Stack RW y Joppa LR 2002 Genetic dissection of a major Fusarium head blight QTL in tetraploid wheat Plant Mol Biol 48 625 632 Paillard S Schnurbusch T Tiwari R Messmer M Winzeler M Keller B y Schachermayr G 2004 QTL analysis of resistance to Fusarium head blight in Swiss winter wheat Triticum aestivum L Theor Appl Genet 109
5. Cassman KG y Liska AJ 2007 Food and fuel for all realistic or foolish Biofuels Bioprod Bioref 1 18 23 135 Bibliograf a Chang WC Lee TY Huang HD Huang HY y Pan RL 2008 PlantPAN Plant Promoter Analysis Navigator for identifying combinatorial cis regulatory elements with distance constraint in plant gene group BMC Genomics 9 561 Chee PW Elias EM Anderson JA y Kianian SF 2001 Evaluation of a high grain protein QTL from Triticum turgidum L var dicoccoides in an adapted durum wheat background Crop Sci 41 295 301 Chen XM Line RF y Leung H 1998 Genome scanning for resistance gene analogs in rice barley and wheat by highresolution electrophoresis Theor Appl Genet 97 345 355 Chen WP Chen PD Liu DJ Kynast R Friebe B Velazhahan R Muthukrishnan S y Gill BS 1999 Development of wheat scab symptoms is delayed in transgenic wheat plants that constitutively express a rice thaumatin like protein gene Theor Appl Genet 99 5 755 760 Chen X Faris JD Hu J Stack RW Adhikari T Elias EM Kianian SF y Cai X 2007 Saturation and comparative mapping of a major Fusarium head blight resistance QTL in tetraploid wheat Mol Breed 19 2 113 124 Cho SH Lee J Jung KH Lee YW Park JC y Baek NC 2012 Genome wide analysis of genes induced by Fusarium graminearum infection in the tolerant and susceptible wheat J Plant Biol 55 64 72 Christensen AB Thordal Christensen H Zimmermann G Gjetting T
6. 4 753 764 Stitcher L Mauch Mani B y Metraux JP 1997 Systemic acquired resistance Annual Rev Phytopathol 35 235 270 Strange RN Majer JR y Smith H 1974 The isolation and identification of choline and betaine as two major components in anthers and wheat germ that stimulate Fusarium graminearum in vitro Physiol Plant Pathol 4 277 290 Takegami S 1957 Studies on the resistance of wheat varieties to Giberella zea Schw Petch head blight and is mechanism 1 Varietal differences of the position of florets in the wheat spikelet Sci Rep Fac Agric 10 33 42 Tamburic Ilincic L Schaafsma AW Fedak G y Falk DE 2000 Selecting for FHB resistance in early generations of winter wheat populations In Proceedings of the 2000 National Fusarium Head Blight Forum Erlanger KY United States 284 287 Tanno K y Willcox G 2006 How fast was wild wheat domesticated Science 311 1886 Tekauz A McCallum B y Gilbert J 2000 Review Fusarium head blight of barley in western Canada Can J Plant Pathol 22 9 16 Tekauz A McCallum B Ames N y Fetch JM 2004 Fusarium head blight of oat current status in western Canada Can J Plant Pathol 26 473 479 Thatcher L Anderson J y Singh K 2005 Plant defense responses what have we learnt from Arabidopsis Func Plant Biol 32 1 19 Trail F 2009 For blighted waves of grain Fusarium graminearum in the postgenomics era Plant Physiol 149 103 110 Tsunewaki K 2009 Plasmon anal
7. Genetic mapping of QTL for resistance to Fusarium head blight spread type 2 resistance in a Triticum dicoccoidesx Triticum durum backcross derived population Theor Appl Genet 126 11 2825 2834 Burgess LW Klein TA Bryden WL y Tobin NF 1987 Head Blight of Wheat Caused by Fusarium Graminearum Group 1 in New South Wales Australia in 1983 Australas Plant Pathol 16 72 78 Bushnell WR Hazen BE y Pritsch C 2003 Histology and physiology of Fusarium head blight En Fusarium head blight of wheat and barley Eds Leonard KJ y Bushnell WR St Paul MN APS Press 44 83 Cai X 2013 USWBSI Durum CP Planning Meeting Report Disponible en http scab pw usda gov pdfs DUR CP Mtg Rep 5 13 pdf Cantrell RG y Joppa LR 1991 Genetic analysis of quantitative traits in wild emmer Triticum turgidum L var dicoccoides Crop Sci 31 645 649 Carbajo HL Gualati A Jensen CA y Loewy T 2001 C pitulo I rea de cultivo Trigo candeal Manual T cnico Chara Experimental Integrada INTA Barrow Carmona M 2000 Fusariosis Sintomatolog a epidemiolog a y control Revista Agromercado 191 58 60 Carmona MA Formento AN Luque A y Scandiani MM 2012 Afio Nifio Preocupaci n y alerta por Fusariosis en trigo y cebada Qu tomar en cuenta C mo decidir la aplicaci n Que aplicar Disponible en http www agroconsultasonline com ar Carr JP Lewsey MG y Palukaitis P 2010 Signaling in induced resistance Adv Virus Res 76 57 121
8. Lyngkjaer MF Dudler R y Schweizer P 2004 The germinlike protein GLP4 exhibits superoxide dismutase activity and is an important component of quantitative resistance in wheat and barley Mol Plant Microbe Interact 17 109 117 Collard BC y Mackill DJ 2008 Marker assisted selection an approach for precision plant breeding in the twenty first century Philos Trans R Soc B Biol Sci 363 1491 557 572 136 Bibliograf a Collinge DB Gregersen P y Thordal Christensen H 1994 The induction of gene expression in response to pathogenic microbes En Mechanisms of plant growth and improved productivity Modern approaches Ed Basra AS New York Marcel Dekker 391 433 Conrath U Pieterse CM y Mauch Mani B 2002 Priming in plant pathogen interactions Trends Plant Sci 7 5 210 216 Dalcero A Torres A Etcheverry M Chulze S y Varsavsky E 1997 Occurrence of deoxynivalenol and Fusarium graminearum in Argentinean wheat Food Addit Contam 14 11 14 Debener T Lehnackers H Arnold M y Dangl JL 1991 Identification and molecular mapping of a single Arabidopsis thaliana locus determining resistance to a phytopathogenic Pseudomonas syringae isolate Plant J 1 289 302 Deihimi T Niazi A Ebrahimi M Kajbaf K Fanaee S Bakhtiarizadeh MR y Ebrahimie E 2012 Finding the undiscovered roles of genes an approach using mutual ranking of coexpressed genes and promoter architecture case study dual roles of thaumatin like pr
9. en la regi n oeste de la zona triguera Bolet n del Ministerio de Agricultura de la Naci n 36 293 299 Mardi M Pazouki L Delavar H Kazemi MB Ghareyazie B Steiner B Nolz R Lemmens M y Buerstmayr H 2006 QTL analysis of resistance to Fusarium head blight in wheat using a Frontana derived population Plant Breed 125 313 317 Matus C diz MA Pozniak CJ Hughes GR y Hucl P 2006 Multiplexed PCR to screen for a major QTL carrying Fusarium head blight resistance in Sumai 3 wheat Can J Plant Sci 86 3 711 716 147 Bibliograf a Mauch Mani B y Metraux J 1998 Salicylic acid and systemic acquired resistance to pathogen attack Ann Bot 82 535 540 McCormick S 2003 The role of DON in pathogenicity En Fusarium Head Blight of Wheat and Barley Eds Leonard KJ y Bushnell WR The American Phytopathological Society St Paul Minnesota 165 183 McFadden ES y Sears ER 1944 The artificial synthesis of Triticum spelta Rec Genet Soc Am 13 26 27 McMullen MP Stack RW y Casper H 1994 Vomitoxin levels in durum and hard red spring wheats infected with Fusarium head blight in 1993 Biol Cultiv Tests for Control of Plant Dis 9 116 McMullen M Jones R Gallenberg D 1997 Scab of wheat and barley a reemerging disease of devasting impact Plant Dis 81 1340 1348 McMullen M 2003 Impacts of Fusarium head blight on the North American agriculture community the power of one disease to catapult change Pages 484 503
10. encoding germin like proteins of barley Regulation and function in basal host resistance Plant Physiol 142 181 192 161
11. resistance to manage Fusarium head blight and deoxynivalenol in wheat Plant Dis 96 957 967 Woo EJ Dunwell JM Goodenough PW Marvier AC y Pickersgill RW 2000 Germin is a manganese containing homohexamer with oxalate oxidase and superoxide dismutase activities Nat Struct Biol 7 1036 1040 Wu AB Li HP Zhao CS y Liao YC 2005 Comparative pathogenicity of Fusarium graminearum isolates from China revealed by wheat coleoptile and floret inoculations Mycopathologia 160 75 83 Xiao J Jin X Jia X Wang H Cao A Zhao W Pei H Xue Z He L Chen Q y Wang X 2013 Transcriptome based discovery of pathways and genes related to resistance against Fusarium head blight in wheat landrace Wangshuibai BMC Genomics 14 1 197 Xu X 2003 Effects of environmental conditions on the development of Fusarium ear blight Eur J Plant Pathol 109 683 689 Yamaguchi T Minami E Ueki J y Shibuya N 2005 Elicitor induced activation of phospholipases plays an important role for the induction of defense responses in suspension cultured rice cells Plant Cell Physiol 46 579 587 Yamaguchi Shinozaki K y Shinozaki K 1994 A novel cis acting element in an Arabidopsis gene is involved in responsiveness to drought low temperature or high salt stress Plant Cell 6 251 64 Yamaguchi Shinozaki K y Shinozaki K 2005 Organization of cis acting regulatory elements in osmotic and cold stress responsive promoters Trends Plant Sci 10 88 94 16
12. to F graminearum infection In this Thesis it was developed an in vitro assay through the evaluation of the variables Germination Coleoptile length Coleoptile weight and Root weight using seven varieties of commercial durum durum wheat cv LDN and the derived resistant lines LDN Dic 3A and LDN DGEI and two common wheat genotypes A601 and A601S3 The seedling variables explained between 51 and 74 of the disease severity being Coleoptile length and weight the ones that more effectively predicted the resistance to FHB The introgressed genotypes showed better performance in the seedling assay and relatively lower damage in the spikes in relation to susceptible ones suggesting that this in vitro test can detect FHB resistance in different genetic backgrouds Thus we propose an in vitro assay based on coleoptile variables to perform quick qualified and cost effective evaluation of the FHB resistance level defining the Seedling Resistance Index highly related to severity for each genotype Thus the present Thesis allowed us to postulate that the interaction wheat F graminearum could be classified as gen for gen leading to the expression of defense related genes Concurrently there were obtained genotypes resistant to F graminearum and it was developed an in vitro assay that predicts the genotypes reponse to infection XV BIBLIOGRAF A Bibliograf a Acevedo Vergara R Araya Mourgues F Cabello Holley MC Herrera Barr
13. 0 Bibliograf a Yli Mattila T Gagkaeva T Ward TJ Aoki T Kistler HC y O Donnell K 2009 A novel Asian clade within the Fusarium graminearum species complex includes a newly discovered cereal head blight pathogen from the far east of Russia Mycologia 101 841 852 Yoda H Hiroi Y y Sano H 2006 Polyamine oxidase is one of the key elements for oxidative burst to induce programmed cell death in tobacco cultured cells Plant Physiol 142 193 206 Zhang Z Schwartz S Wagner L y Miller W 2000 A greedy algorithm for aligning DNA sequences J Comput Biol 7 203 214 Zhang L Meakin H y Dickinson M 2003 Isolation of genes expressed during compatible interactions between leaf rust Puccinia triticina and wheat using cDNA AFLP Mol Plant Pathol 4 469 477 Zhang X Zhou MP Ren LJ Bai GH Ma HQ Scholten OE Guo PG y Lu WZ 2004 Molecular characterization of Fusarium head blight resistance from wheat variety Wangshuibai Euphytica 139 59 64 Zhang G y Mergoum M 2007 Molecular mapping of kernel shattering and its association with Fusarium head blight resistance in a Sumai3 derived population Theor Appl Genet 115 757 766 Zhang C Wang Y Wang J Zhai Z Zhang L Zheng W Yu W Zhou J Lu G Shim WB y Wang Z 2013 Functional characterization of Rho family small GTPases in Fusarium graminearum Fungal Genet Biol 61 90 99 Zimmermann G Ba mlein H Mock HP Himmelbach A y Schweizer P 2006 The multigene family
14. 323 332 Palazzini JM Ramirez ML Torres AM y Chulze SN 2007 Potential biocontrol agents for Fusarium head blight and deoxynivalenol production in wheat Crop Prot 26 1702 1710 Parry DW Jenkinson P y McLeod L 1995 Fusarium ear blight scab in small grain cereals a review Plant Pathol 44 207 238 150 Bibliograf a Patnaik D y khurana P 2001 Wheat biotechnology A minireview Electron J Biotechnol ISSN 0717 3458 Paul PA Lipps PE Hershman DE McMullen MP Draper MA y Madden LV 2007 A quantitative review of tebuconazole effect on Fusarium head blight and deoxynivalenol content in wheat Phytopathol 97 211 220 Pearce RB Strange RN y Smith H 1976 Glycinebetaine and choline in wheat Distribution and relation to infection by Fusarium graminearum Phytochem 15 6 953 954 Peng JH Sun D y Nevo E 2011 Domestication evolution genetics and genomics in wheat Mol Breed 28 3 281 301 Peng JH Sun DF Peng YL y Nevo E 2012 Gene discovery in Triticum dicoccoides the direct progenitor of cultivated wheats Cereal Research Communications DOI 10 1556 CRC 2012 0030 Pestka JJ 2010 Deoxynivalenol mechanisms of action human exposure and toxicological relevance Arch Toxicol 84 663 679 Pinheiro JC y Bates DM 2000 Mixed Effects Models in S and S PLUS Springer New York Polesani M Desario F Ferrarini A Zamboni A Pezzotti M Kortekamp A y Polverari A 2008 cDNA AFLP analysis of plant a
15. C Altmann T y Weiler EW 2000 12 Oxophytodienoate reductase 3 OPR3 is the isoenzyme involved in jasmonate biosynthesis Planta 210 6 979 984 Schisler DA Khan NI Boehm MJ Lipps PE Slininger PJ y Zhang S 2006 Selection and evaluation of the potential of choline metabolizing microbial strains to reduce Fusarium head blight Biol Control 39 497 506 Schmolke M Zimmermann G Buerstmayr H Schweizer G Miedaner T Korzun V Ebmeyer E y Hartl L 2005 Molecular mapping of Fusarium head blight resistance in the winter wheat population Dream Lynx Theor Appl Genet 111 747 756 Schmolke M Zimmermann G Schweizer G Miedaner T Korzun V Ebmeyer E y Hartl L 2008 Molecular mapping of quantitative trait loci for field resistance to Fusarium head blight in a European winter wheat population Plant Breed 127 459 464 Schr dter R 2004 Influence of harvest and storage conditions on trichothecenes levels in various cereals Toxicol Lett 153 47 49 Schroeder HW y Christensen JJ 1963 Factors affecting resistance of wheat to scab caused by Gibberella zeae Phytopathol 53 831 838 Schulze Lefert P 2004 Knocking on the heaven s wall pathogenesis of and resistance to biotrophic fungi at the cell wall Curr Opin Plant Biol 7 377 383 Schweiger W Steiner B Ametz C Siegwart G Wiesenberger G Berthiller F Lemmens M Jia H Adam G Muehlbauer GJ Kreil DP y Buerstmayr H 2013 Transcriptomic characterization of two major Fu
16. Fs identificados se hallan prote nas asociadas a la respuesta temprana a la infecci n receptores NBS LRR y receptores quinasa involucrados en el reconocimiento espec fico del determinante de avirulencia del pat geno Fueron identificados adem s TDFs que aunque no pudo asign rseles una prote na o funci n resultaron espec ficos de la respuesta a la inoculaci n La identidad de TDFs con ESTs de genotecas de espiga de materiales de 7 aestivum inoculadas con F graminearum constituye un sustento adicional para esta afirmaci n El mapeo in silico permiti localizar 28 TDFs en el genoma de T aestivum siendo el brazo cromos mico 5BL el m s representado adem s de obtener las regiones gen micas y regulatorias de varios genes A partir de estas regiones pudo determinarse la existencia de mecanismos de regulaci n de la transcripci n en com n entre algunos genes asociados a los TDFs entre ellas las prote nas WRKY implicadas en la regulaci n de los genes asociados con la defensa ante pat genos La integraci n de la informaci n obtenida sugiere que la interacci n trigo F graminearum no ser a una interacci n compatible como generalmente se XI Resumen cree sino que se tratar a de una interacci n gen a gen que finalmente lleva a la expresi n de genes asociados a la defensa Hemos asumido adem s el desaf o de desarrollar cultivares de trigo candeal resistentes La compleja herencia de la resistencia y los efectos amb
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18. Resumen RESUMEN La Fusariosis de la Espiga de Trigo FET causada por Fusarium graminearum genera p rdidas en rendimiento y contaminaci n de granos con micotoxinas Existe escasa variabilidad gen tica a la resistencia en el germoplasma de trigo candeal La l nea recombinante cromos mica endocriada LDN Dic 3A presenta promisorios niveles de resistencia Los objetivos de esta Tesis comprenden i identificar genes implicados en la resistencia a FET en LDN Dic 3A ii transferir el QTL de resistencia de LDN Dic 3A a variedades susceptibles de trigo candeal iii desarrollar un ensayo in vitro en pl ntula para identificar genotipos resistentes y su relaci n con la severidad de la enfermedad La identificaci n de la expresi n diferencial de genes inducida en diferentes tiempos post inoculaci n con F graminearum entre LDN Dic 3A y el parental susceptible LDN se bas principalmente en la t cnica de cDNA AFLP De 500 fragmentos derivados de transcripci n TDF identificados con las distintas combinaciones de cebadores utilizados 85 mostraron expresi n diferencial el 36 y el 19 fueron identificados en LDN Dic 3A y LDN respectivamente mientras que el 45 se indujeron en ambos genotipos Los patrones de TDFs obtenidos mediante cDNA AFLP demostraron ser reproducibles mediante la t cnica de RT PCR dando validez a nuestro sistema experimental La comparaci n con secuencias depositadas en bases de datos mostr que entre los TD
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20. d the identification of several genes and regulatory regions being the 5BL chromosome arm where most TDFs were located The analysis of the regulatory regions revealed the existence of transcription regulation mechanisms shared by some TDFs associated genes such as WRKY proteins implied in the regulation of genes associated to pathogen defence The present results suggest that wheat F graminearum interaction is governed by gene for gene relationships The development of resistant cultivars has been a difficult task due to the complex inheritance of resistance and the influence of environmental factors The resistance QTL Qfhs ndsu 3AS from XIV Abstract LDN Dic 3A was incorporated in the cultivars Buck Emeralda and Buck Candisur through crosses F3 homozygous individuals for the resistance allele were subjected to marker assisted selection using the Xgwm2 microsatellite linked to the mentioned QTL allowing a reduction in the number of individuals included in the following steps of the breeding program In F4 there were selected the individuals that showed equal or better resistance performances compared to the resistant parent evaluated through the spike severity at 21 days post inoculation The breeding program will continue by selfing resistant genotypes to obtain plant materials that possess both stable resistance and suitable agronomic traits Rapid and trustable assays are required for the evaluation of germoplasm response
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30. ientales son los responsables del escaso xito obtenido hasta el momento por los mejoradores en la incorporaci n al mercado de cultivares resistentes En este trabajo por medio de cruzamientos se incorpor el QTL de resistencia Ofhs ndsu 3AS de LDN Dic 3A en los cultivares BESM y BCAN El microsat lite Xgwm2 ligado al QTL de resistencia permiti reducir la cantidad de individuos que continuaron en el programa de mejoramiento En la generaci n F3 se seleccionaron los individuos homocigotos para el alelo de resistencia y en F4 se evalu la severidad en espiga identificando individuos con niveles de resistencia similares o mejores que el parental resistente El programa de mejoramiento continuar con autofecundaciones de genotipos resistentes hasta alcanzar estabilidad en la resistencia junto con la presencia de caracteres agron micos de inter s La evaluaci n del comportamiento ante F graminearum de nuevos materiales requiere de la existencia de ensayos r pidos y confiables Hemos desarrollado un ensayo in vitro a trav s de la evaluaci n de las variables Germinaci n Largo de coleoptilo Peso de coleoptilo Peso de ra ces utilizando siete variedades comerciales de trigo candeal el trigo candeal LDN y las l neas resistentes LDN Dic 3A y LDN DGEI y dos genotipos de trigo pan A601 y A601S3 Las variables de pl ntula explicaron entre el 51 y el 74 de la severidad de la enfermedad siendo Largo y Peso de coleoptilo las m s eficaces
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34. nd pathogen genes expressed in grapevine infected with Plasmopara viticola BMC Genomics 9 142 Pozniack CJ Clarke JM Singh AK Thomas J y Fedak G 2007 Progress in Fusarium head blight resistance breeding in Canadian Durum Wheat In Proceedings of 5th Canadian Workshop on Fusarium Head Blight Winnipeg Manitoba Canad 47 Prandini A Sigolo S Filippi L Battilani P y Piva G 2008 Review of predictive models for Fusarium head blight and related mycotoxin contamination in wheat Food Chem Toxicol 47 927 931 151 Bibliograf a Pritsch C Muehlbauer GJ Bushnell WR Somers DA y Vance CP 2000 Fungal development and induction of defense response genes during early infection of wheat spikes by Fusarium graminearum Mol Plant Microbe Interact 13 2 159 169 Pritsch C Vance CP Bushnell WR Somers DA Hohn TM y Muehlbauer GJ 2001 Systemic expression of defense response genes in wheat spikes as a response to Fusarium graminearum infection Physiol Mol Plant 58 1 1 12 Purahong W Alkadri D Nipoti P Pisi A Lemmens M y Prodi A 2012 Validation of a modified Petri dish test to quantify aggressiveness of Fusarium graminearum in durum wheat Eur J Plant Pathol 132 381 391 Purahong W Nipoti P Pisi A Lemmens M y Prodi A 2014 Aggressiveness of different Fusarium graminearum chemotypes within a population from Northern Central Italy Mycoscience 55 63 69 R Development Core Team 2008 R A language and en
35. onible en http www laarena com ar includes modulos print php id298015 amp seccionid Somers DJ Fedak G y Savard M 2003 Molecular mapping of novel genes controlling Fusarium head blight resistance and deoxynivalenol accumulation in spring wheat Genome 46 555 564 Somers DJ Fedak G Clarke J y Cao WG 2006 Mapping of FHB resistance QTLs in tetraploid wheat Genome 49 12 1586 1593 Somssich IE y Hahlbrock K 1998 Pathogen defence in plants a paradigm of biological complexity Trends Plant Sci 3 86 90 Song WY Wang GL Chen LL Kim HS Pi LY Holsten T Gardner J Wang B Zhai WX Zhu LH Fauquet C y Ronald P 1995 A receptor kinase like protein encoded by the rice disease resistance gene Xa21 Science 270 1804 1806 155 Bibliograf a Srinivasachary Gosman N Steed A Simmonds J Leverington Waite M Wang Y Snape J Nicholson P 2008 Susceptibility to Fusarium head blight is associated with the Rht DIb semi dwarfing allele in wheat Theor Appl Genet 116 1145 1153 Stack RW 1988 Response of durum wheat to head blight Biol Cultiv Tests for Control of Plant Dis 3 39 Stack RW Elias EM y Joppa LR 1999 Fusarium head blight reaction of durum wheat lines conditioned by Triticum dicoccoides chromosome substitutions Phytopathol 89 S74 abstract Stack RW 2000 Return of an old pathogen Fusarium head blight of small grains Plant Management Network International Disponible en http www plantmanag
36. os G Leyton Buccicardi G Toro Torres M Vasquez Toro R y Zeren Zeren M 2008 Estudio de la competitividad de la cadena de trigo candeal Fundaci n Chile Fundaci n para la Innovaci n Agraria Agostinelli AM Clark AJ Brown Guedira G y Van Sanford DA 2012 Optimizing phenotypic and genotypic selection for Fusarium head blight resistance in wheat Euphytica 186 115 126 Alberione E Arburua M Fissore G y Fornero G 2013 Eficacia en el control qu mico de enfermedades foliares en trigo y cebada Disponible en http www engormix com MA agricultura trigo articulos eficacia control quimico enfermedades t4888 998 p0 htm Altschul SF Madden TL Schaffer AA Zhang J Zhang Z Miller W y Lipman DJ 1997 Gapped BLAST and PSI BLAST A new generation of protein database search programs Nucleic Acids Res 25 3389 3402 Anand A Zhou T Trick HN Gill BS Bockus WW y Muthukrishnan S 2003 Greenhouse and field testing of transgenic wheat plants stably expressing genes for thaumatin like protein chitinase and glucanase against Fusarium graminearum J Exp Bot 54 384 1101 1111 Andersen L 1948 The development of Gibberella zeae head blight of wheat Phytopathol 38 595 611 Atanasoff D 1920 Fusarium blight scab of wheat and other cereals J Agricult Res 20 1 32 Bachem CWB Van der Hoeven RS De Bruijn SM Vreugdenhil D Zabeau M y Visser RGF 1996 Visualization of differential gene expression using a novel method of
37. oteins in biotic and abiotic stresses SpringerPlus 1 1 1 10 Dellaporta SL Wood J y Hicks J 1983 A plant DNA minipreparation version II Plant Mol Biol Rep 1 14 19 21 Del Ponte EM Fernandes JMC Pavan W y Baethgen WE 2009 A model based assessment of the impacts of climate variability on Fusarium head blight seasonal risk in southern Brazil J Phytopathol 157 675 681 De Pater S Greco V Pham K Memelink J y Kijne J 1996 Characterization of a zinc dependent transcriptional activator from Arabidopsis Nucleic Acids Res 24 4624 4631 Desjardins AE y Hohn TM 1997 Mycotoxins in plant pathogenesis Mol Plant Microbe Interact 10 147 152 De Wit PJGM 1997 Pathogen avirulence and plant resistance a key role for recognition Trends Plant Sci 2 452 458 137 Bibliograf a De Wolf ED Madden LV y Lipps PE 2003 Risk assessment models for wheat Fusarium head blight epidemics based on within season weather data Phytopathol 93 28 435 Diamond H y Cooke B 1999 Towards the development of a novel in vitro strategy for early screening of Fusarium ear blight resistance in adult winter wheat plants Eur J Plant Pathol 105 363 372 Dill Macky R y Jones RK 2000 The effect of previous crop residues and tillage on Fusarium head blight of wheat Plant Dis 84 71 76 Dill Macky R 2003 Inoculation methods and evaluation of Fusarium head blight En Fusarium Head Blight of Wheat and Barley Eds Leonard KJ y Bushnel
38. para predecir la resistencia a FET Los genotipos introgresados mostraron un buen comportamiento en el ensayo en pl ntula y menor da o en espiga comparados con los susceptibles sugiriendo que la prueba in vitro es efectiva para la determinaci n de la resistencia a FET en diferentes fondos gen ticos Entonces se propone un ensayo in vitro basado en las variables de coleoptilo para evaluar de manera eficaz r pida y econ mica el nivel de resistencia a FET definiendo el ndice de Resistencia en Pl ntula altamente correlacionado con severidad para cada genotipo Los principales hallazgos de esta Tesis pueden compendiarse indicando que se ha establecido ue la interacci n trigo F graminearum ser a una interacci n gen a gen que lleva a la expresi n de genes asociados a la defensa la obtenci n de genotipos de trigo candeal resistente a XII Resumen fusariosis de la espiga y el dearrollo de un ensayo in vitro predictor del comportamiento de los genotipos ante la infecci n XIII Abstract ABSTRACT Fusarium head blight FHB caused by Fusarium graminearum produce yield losses and contamination of grain with mycotoxins Scarce genetic variability for resistance exists in durum wheat germplasm The LDN Dic 3A recombinant inbred chromosome line showed to be resistant to FBH The goals of this Thesis include 7 identification of genes involved in FHB resistance in LDN Dic 3A ii transference from LDN Dic 3A
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40. relationship to FHB disease development and mycotoxin accumulation from Fusarium graminearum infection Plant Pathol 56 65 72 Buerstmayr H Stierschneider M Steiner B Lemmens M Griesser M Nevo E y Fahima T 2003 Variation for resistance to head blight caused by Fusarium graminearum in wild emmer Triticum dicoccoides originating from Israel Euphytica 130 17 23 Buerstmayr H Lemmens M Schmolke M Zimmermann G Hartl L Mascher F Trottet M Gosman ME y Nicholson P 2008 Multi environment evaluation of level and stability of FHB resistance among parental lines and selected offspring derived from several European winter wheat mapping populations Plant Breed 127 325 332 Buerstmayr H Ban T Anderson JA 2009 QTL mapping and marker assisted selection for Fusarium head blight resistance en wheat a review Plant Breed 128 1 26 Buerstmayr M Lemmens M Steiner B y Buerstmayr H 2011 Advanced backcross QTL mapping of resistance to Fusarium head blight and plant morphological traits in a Triticum macha x T aestivum population Theor Appl Genet 123 2 293 306 Buerstmayr M Hube K Heckmann J Steiner B Nelson J y Buerstmayr H 2012 Mapping of QTL for Fusarium head blight resistance and morphological and developmental traits in three backcross populations derived from Triticum dicoccum x Triticum durum Theor Appl Genet 125 1751 1765 134 Bibliograf a Buerstmayr M Alimari A Steiner B y Buerstmayr H 2013
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42. sarium resistance quantitative trait loci QTLs Fhb1 and Ofhs ifa SA identifies novel candidate genes Mol Plant Pathol DOI 10 1111 mpp 12048 Sestili S Polverari A Luongo L Ferrarini A Scotton M Hussain J Delledonne M Ficcadenti N y Belisario A 2011 Distinct colonization patterns and cDNA AFLP transcriptome profiles in compatible and incompatible interactions between melon and different races of Fusarium oxysporum f sp melonis BMC Genomics 12 122 154 Bibliograf a Shang J Tao Y Chen X Zou Y Lei C Wang J Li X Zhao X Zhang M Lu Z Xu J Cheng Z Wan J y Zhu L 2009 Identification of a new rice blast resistance gene Pid3 by genome wide comparison of paired nucleotide binding site leucine rich repeat genes and their pseudogene alleles between the two sequenced rice genomes Genetics 182 1303 1311 Shi C Yan P Li J Wu H Li Q y Guan S 2014 Biocontrol of Fusarium graminearum growth and deoxynivalenol production in wheat kernels with bacterial antagonists Int J Environ Res Publ Health 11 1094 1105 Scott DB De Jager EJH y Van Wyk PS 1988 Head blight of irrigated wheat in South Africa Phytophylactica 20 317 319 Snyders CHA y Van Eeuwijk FA 1991 Genotype by strain interactions for resistance to Fusarium head blight caused by Fusarium culmorum in winter wheat Theor Appl Genet 81 239 244 Sol I 2013 Toxinas en trigos y harinas Toxicolog a y Nutrici n del INTI Agroalimentos Disp
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44. to susceptible durum varieties of the resistance QTL iii development of an in vitro seedling assay to identify wheat resistant genotypes and their relationship with disease severity Analysis of differential gene expression induced at different time points post inoculation with F graminearum between LDN Dic 3A and the susceptible parental LDN was performed by cDNA AFLP technique A total 85 out of the 500 transcript derived fragments TDFs identified with the diverse primer combination used showed to be differentially expressed 3696 and 19 were identified in LDN Dic 3A and LDN respectively whereas 45 were induced in both genotypes The TDF patterns obtained though cDNA AFLP showed to be reproducible by RT PCR supporting the reliability of our experimental system to identify differentially expressed transcripts Comparison with protein databases revealed that among the cloned TDFs several showed identity to proteins associated with early response to infection to NBS LRR and kinases receptors involved in specific recognition of avirulence pathogen determinant However there was a group of TDFs that in spite of being specific of the inoculation response could not be assigned to characterized proteins The identity of these TDFs with ESTs from libraries from T aestivum inoculated with Fusarium graminearum additionally supports this affirmation The availability of T aestivum genome sequences allowed the in silico mapping of 28 TDFs an
45. vironment for statistical computing R Foundation for Statistical Computing Vienna Austria http www R project org Ram rez ML Reynoso MM Farnochi MC y Chulze S 2006 Vegetative compatibility and mycotoxin chemotypes among Fusarium graminearum Gibberella zeae isolates from wheat in Argentina Eur J Plant Pathol 115 139 148 Ram rez V Garc a Andrade J y Vera P 2011 Enhanced disease resistance to Botrytis cinerea in myb46 Arabidopsis plants is associated to an early down regulation of CesA genes Plant Signal Behav 6 6 911 913 Ravensdale M Rocheleau H Wang L Nasmith C Ouellet T y Subramaniam R 2014 Components of priming induced resistance to fusarium head blight in wheat revealed by two distinct mutants of Fusarium graminearum Mol Plant Pathol DOI 10 111 1 mpp 12145 Reis EM y Carmona M 2002 Fusariosis del trigo Biologia epidemiologia y estrategias para su manejo Eds Reis y Carmona Buenops Aires Argentina ISBN 98743 3959 4 152 Bibliograf a Ribichich KF Lopez SE y Vegetti AC 2000 Histopathological spikelet changes produced by Fusarium graminearum in susceptible and resistant wheat cultivars Plant Dis 84 794 802 R der MS Korzun V Wandehake K Planschke J Tixier MH Leroy P y Ganal MW 1998 A microsatellite map of wheat Genetics 149 2007 2023 Ruan Y Comeau A Langevin F Hucl P Clarke JM Brule Babel A y Pozniak CJ 2012 Identification of novel QTL for resistance to Fusa
46. ysis in the Triticum Aegilops complex Breed Sci 59 455 470 Turner JG Ellis C y Devoto A 2002 The jasmonate signal pathway Plant Cell 14 153 S164 157 Bibliograf a Turner NW Subrahmanyam S y Piletsky SA 2009 Analytical methods for determination of mycotoxins a review Anal Chim Acta 632 168 180 Van Eeuwijk FA Mesterh zy A Kling CI Ruckenbauer P Saur L Buerstmayr H Lemmens M Keizer LCP Maurin N y Snijders CHA 1995 Assessing non specificity of resistance in wheat to head blight caused by inoculation with European strains of Fusarium culmorum F graminearum and F nivale using a multiplicative model for interaction Theor Appl Genet 90 2 221 228 Van Loon LC Rep M y Pieterse CMJ 2006 Significance of inducible defense related proteins in infected plants Annu Rev Phytopathol 44 135 162 Van Slageren MW 1994 Wild wheats a monograph of Aegilops L and Amblyopyrum Jaub and Spach Eig Poaceae Agricultural University Wageningen amp International Center for Agricultural Research in Dry Areas Aleppo Syria Vargas PR y Fern ndez JMC 1997 Modelando o curso da antese em trigo Fitopatologia Brasileira 22 317 Velazquez P y Formento AN 2013 Efecto de la fusariosis de la espiga del trigo Fusarium graminearum y Fusarium spp en un cultivar susceptible Campa a 2012 Paran Entre R os Jornada Regional de Cultivos de Invierno ISBN 978 950 698 308 6 Vidhyasekaran P 2008 Fungal pat

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