Kosová_Cereal Resistance to Fusarium Head Blight and Possibilities

8122019 Kosovaacute_Cereal Resistance to Fusarium Head Blight and Possibilities

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dough stage of kernel development Besides wateravailability also temperature aeration and light arecrucial climatic factors influencing the productionand dispersal of inocula growth competition my-cotoxin production and pathogenicity Fusarium species were found to differ in their climatic dis-tribution and in the optimum climatic conditionsrequired for their persistence but most speciesproduce inocula grow best and are more patho-genic to cereal heads at warm temperatures andunder humid conditions (D et al 2003)

Since the control of FHB spread by fungicidescrop rotation and deep ploughing need not alwaysbe an efficient means of crop protection breedingof FHB resistant cultivars (lines) represents themain approach how to eliminate yield losses and

contamination of grain by mycotoxins caused bythis disease

The problem of FHB in small grain cereals hasbeen reviewed by P et al (1995) M(1997) B and S (2004) and others Thesituation in the European wheat gene pool wa sdescribed precisely by M (2003) B- et al (2009) have recently summarizedthe problems of FHB in wheat and published a listof all FHB resistance QTL (quantitative trait loci)detected in the wheat gene pool so far Basic infor-

mation on pathogen biology including the descrip-tion of its life c ycle production of trichothecenemycotoxins and its population biology is given inthe review by G and K (2004)

In this review we would like to focus on the sourcesof FHB resistance and on progress in the breeding ofFHB resistant cultivars of small grain cereals withrespect to the Central European region

Manifestation of the disease and its spreading

The first record of FHB or scab comes fromS (1884) who reported the occurrence ofthis disease in England and attributed it to thefungus Fusisporium culmorum

The symptoms of the disease are quite similarin various small grain cereals the fir st symptomsappear as small patches of discoloration at the baseof the glume of a floret in a spikelet As the infec-tion progresses the patches of discoloration mayspread to the rachis and invade the whole spikeThe infection is visible as patches which first have

a bleached (light) appearance later they becomeorange salmon pink red or tan or they can even

change the infected seeds into hard dark brownobjects (called lsquotomb stonesrsquo) Recently Vet al (2005) have found out that secreted lipaseencoded by the FGL1 gene of Fusarium gramine-arum acts as a v irulence factor in the process ofwheat spike infection

The major potential risk for both humans andanimals caused by the infection is the productionof mycotoxins either oestrogenic zearalenone(ZEA or ZON) which is mainly found in infectedmaize ( Ze a mays ) ( Y 1991) or non-macrocyclic trichothecenes According to theirchemical nature the trichothecenes produced by

F culmorum and F graminearum are divided intotwo types type A toxins (HT-2 toxin T-2 toxin)and type B toxins including the most abundantly

produced mycotoxins in small grain cereals es-pecially deoxynivalenol (DON) its derivatives3-acetyl-DON and 15-acetyl-DON and nivalenol(NIV) and 4-acetyl-NIV called also fusarenone X(FX) Most Fusarium strains can simultaneouslyproduce multiple types of mycotoxins eg DONand ZEA In general both DON and NIV chemo-types were reported in Africa Asia and Europewhile only the DON chemotype was found in theUSA (M et al 1999 L et al 2002) Ourlatest studies performed in the territory of the

Czech Republic (S et al 2008) showed ver y rare occurrence of the NIV chemotype andhigh prevalence of the DON chemotype DONdiffers from NIV only in the absence of a hydroxylgroup at C-4 However this difference may haveimportant consequences for the fitness of theproducing organism as it alters the bioactivityand toxicity of trichothecenes For example thetoxicity of NIV is several times higher than thatof DON (A et al 2000) The molecularbasis of NIV or DON-producing abilities of theisolates lies in the functionality of the Tri13 gene

coding for a cytochrome P-450 enzyme which isfunctional in NIV-producing isolates while non-functional in DON-producing strains (B etal 2002 L et al 2002)

The infection is spread through both the asexualspores (macroconidia) and the sexual ascosporesby wind rain-splash dispersal or by arthropod

vectors such as mites (Siteroptes graminum) clo- ver leaf weevil ( Hypera punctata) grasshoppers( Melanoplus bivittatus) picnic beetles (Glischro-chilus quarisignatus) and even the common house-

fly ( Musca domestica) Crop residues on the soilsurface also represent an important reservoir of

Czech J Genet Plant Breed 45 2009 (3) 87ndash105



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pathogens containing ascospores macroconidiaand hyphal fragments which can serve as inocu-lum The fungus spreads by direct penetration ofglumes palea and rachilla The fungus infects otherspikelets internally through vascular bundles ofthe rachilla and rachis in susceptible wheat Theinfection of plants by other diseases (eg BYDV)can significantly contribute to the spread of FHB(eg L amp B 2005) Ascospores mac-roconidia (asexual spores) chlamydospores andhyphal fragments in the soi l surface debris are themain sources of inoculum

Types and mechanisms of FHB resistance

FHB resistance is polygenic and it s expression ishighly influenced by the environment It becomesobvious that resistance against FHB in small graincereals is determined quantitatively by severalquantitative trait loci (QTL) It is however im-portant that many studies showed common anddurable resistance to different Fusarium spp caus-ing FHB (S et al 1997 H et al 2003M et al 2005) Though the isolates ofdifferent Fusarium spp differed largely in quantita-tive aggressiveness they did not show any evident

qualitative differences in virulence (Met al 2005 A et al 2006 Š et al 2008)Since the pathogenic specialization can be classi-fied as low wheat breeding for general Fusariumresistance may be possible

Five types of resistance to FHB have been dis-tinguished in common wheat the first two havealready been classified by S and C- (1963) as resistance to the initial infection(Type I resistance) and resistance to subsequentspread (Type II resistance) Later resistance tomycotoxin accumulation indicated by mycotoxin

concentrations uncorrelated with Type I or Type IIresistance was recognized (M amp A1986) The following components (types) of headblight resistance were distinguished by M- (1995 2002) I Resistance to invasion II Re-sistance to spreading III Resistance to kernelinfection IV Tolerance V Resistance to toxinaccumulation VI Resistance to late blightingand VII Resistance to head death above infec-tion site B et al (2008) have proposedto divide Type V resistance into two sub-types

Type V-1 resistance which lies in the promotionof mycotoxin degradation or detoxification and

Type V-2 resistance which means the preventionof mycotoxin biosynthesis

Several mechanisms of FHB resistance haveevolved in cereals Mechanisms of resistance tomycotoxin accumulation (Type V resistance) liein the production of enzymes which are aimed atthe detoxification and degradation of mycotoxinsThese enzymes include members from the familyof pathogenesis-related (PR) proteins like PR-2(glucanases) PR-3 (chitinases) PR-4 (chitinasetypes I II) and PR-5 (thaumatin-like proteins) pro-teins The expression of enzymes like peroxidaseslipoxygenase and other ROS scavenging enzymes isalso enhanced It was found out by L et al (2001)that the FHB-resistant cultivar Sumai 3 one of themost worldwide used source of FHB resistance in

wheat breeding programmes accumulates higherlevels of chitinases and β-13-glucanases than itssusceptible mutant A wide range of se condarymetabolites with pro-oxidant properties like phe-nolic compounds carotenoids and linoleic acidhydroperoxides are synthesized in order to act asmodulators of mycotoxin biosynthesis RecentlyB et al (2007) compared changes in geneexpression initiated by the infection in resistantcultivar Ning 7840 and susceptible cultivar Clarkand found the enhanced expression of defence-

related genes (precursor of chitinase II genesencoding PR proteins P450 three genes withunknown function) in Ning 7840 relative to Clarkduring early stages of fungal stress A detailedreview on cereal mechanisms of degradation of

Fusarium mycotoxins has recently been publishedby B et al (2008)

It is also becoming evident that FHB resistance(predominantly Type I and II resistance ie FHBinitial infection and subsequent spread) is associ-ated with alleles determining plant morphologicaland physiological characteristics especially plant

height heading date (P et al 1995) inflo-rescence architecture and traits associated withflowering such as narrow flower opening (G- et al 2005) or anther extrusion (T2004) The FHB resistance associated with plantphysiological and morphological characteristicsis also called passive resistance (M1995) It can be generally summed up that plantswith loose spike branching (loose heads) exhibitbetter resistance to FHB spread than plants withdense heads however the grain yield of these

cultivars is much lower than of those with denseheads naturally Recently it was found by Z




90

et al (1999) in doubled haploid (DH) lines from across between the two-rowed genotypes of barleyGobernadora and CMB643 that all but two QTLfor resistance to FHB coincide with the QTL forinflorescence architecture and plant height Theauthors hypothesized that the architecture of theinflorescence might affect the spread of FHB inbarley however they concluded that they did notfind out whether the coincidence of these two typesof QTL was due to linkage or pleiotropy and theyconcluded that further research was needed inthis area In wheat several FHB resistance QTLsoverlap with QTL for plant height weight of in-fected spikelets or awns when examining crossesbetween unrelated cultivars which differ in thesemorphological traits (eg G et al 2003

S et al 2008a) Quite a lot of pa-pers have also been published which examine theeffect of allelic constitution at Rht-B1 locus (alsocalled Rht1) and Rht-D1 locus (also called Rht2loci determining plant height located on 4BS and4DS respectively) on FHB resistance in wheatS et al (2008b) have shown ona mapping population between Spark (a cultivarrelatively resistant to FHB carrying Rht-D1a a l-lele) and Rialto (FHB susceptible cultivar carrying

Rht-D1b allele) that the level of FHB resistance

correlates with the allele type at Rht-D1 dwarfinglocus ie the presence of mutant Rht-D1b semi-dwarfing allele which is considered beneficial foragronomic purposes results in shorter plants withdecreased FHB type I resistance when comparedto normal Rht-D1a allele Similar results wereobtained by B et al (2008) from theevaluation of FHB resistance in recombinant linesderived from the crosses ArinaForno ArinaRib-and DreamLynx G16-92Hussar RenanReacutecitalSVP-72017Capo and CapoSumai 3 under naturalconditions at different locations in Europe How-

ever V et al (2008) showed on a set of threesegregating populations of winter wheat that thecorrelation between the presence of Rht-D1b alleleand FHB resistance is not absolute ie lines car-rying Rht-1Db allele with a relatively high level ofFHB resistance can be obtained Similarly H- et al (2008) showed on three sets of RILsbetween European winter wheats that the Rht-D1 genomic region is the major source of variation inFHB resistance in the European wheat gene poolAnalogously to V et al (2008) they also ob-

tained semi-dwarf lines with acceptable levels ofFHB resistance due to the effects of other minor

QTLs Rht-B1b allele at Rht-B1 locus seems tohave a less negative effect especially on Type IIresistance (S et al 2009) At both

Rht-B1b and Rht-D1b loci DELLA proteins whichare negative regulators of gibberellin (GA) signal-ling are located Since it is becoming evident that

Fusarium species exploit GA signalling pathwaysduring plant infection it seems very probable thatthe negative effect of Rht-D1 semi-dwarfing alleles(and orthologous Rht-B1 ones) on FHB resistanceis due to pleiotropy rather than to linkage drag(H et al 2008 N et al 2008)Thus other semi-dwarfing genes are becomingimportant in breeding programmes RecentlyP et al (2008) have shown that the intro-gression of another dwarfing gene Rht8 can lead

to the construction of high-yielding semi-dwarflines with a sufficiently high level of FHB resist-ance The authors explained their success by aphysiologically different mode of action of the Rht8 gene in comparison with widely used Rht-B1b and

Rht-D1b dwarfing genes ( Rht8 is a GA-sensitivegene tightly linked to the Ppd-D1a allele confer-ring photoperiod insensitivity while Rht-B1b and

Rht-D1b alleles are GA-insensitive)Another adverse characteristic that is usually as-

sociated with the introgression of Sumai 3-derived

FHB resistance QTL is kernel shattering Sumai 3is susceptible to shattering Z and M(2007) studied the relationship between FHB re-sistance QTL and the QTL determining kernelshattering in a population of RILs between Sumai3 and FHB susceptible cultivar Stoa and found outthat there would exist a tight linkage between threemajor FHB resistance QTL and kernel shatteringQTL Another association of FHB resistance (ormoderate tolerance) QTL and some physiologicalcharacteristics was shown in a study of Get al (2005) who found an association and an

overlap between FHB resistance QTL and floweropening in a population of RILs between Goldfield(a narrow opening wheat cultivar) and Patterson(a wide opening wheat cultivar)

Generally it can be stated that the introgressionof FHB resistance QTL into elite high-yielding cul-tivars often brings about a potential risk of worsen-ing of important morphological and physiologicalcharacteristics Therefore detailed mapping of theFHB resistance QTL as well as the overlapping QTLdetermining important agronomical characteristics

is necessary in order to make significant progressin FHB resistance breeding




91

Methods for evaluation of FHB resistance

in small grain cereals (wheat)

It is very important to find a reliable methodto assess variety resistance to FHB Several ar-tificial inoculation methods have been reported(B 1985 L 1985) Specific inoculationprocedures to evaluate fungal spread in the ear(Type II resistance) involve the introduction ofinoculum into the central spikelet of an ear at earlyanthesis (S amp C 1963 Bamp S 1996) Inoculation methods include ahypodermic syringe a micropipette or a small tuftof cotton soaked in inoculum These techniqueshave proved to be reliable (W amp M 1988)but are labour-intensive and important resistance

mechanisms contributing to field resistance mightnot be detected Spray inoculation is used for theassessment of both Type I and Type II resistanceIt seems that Type I resistance at present cannotbe directly measured only as a difference betweenpoint and spraying inoculation For breeding thespraying methodology is more suitable as it con-siders a much wider genetic background than thepoint inoculation does (M et al 2008)To minimize yearlocation effects on results itoften appears necessary to support the disease

development by irrigation of plotsManagement practices such as preceding cropand tillage system play an important role in naturalinfection of FHB The use of maize as precedingcrop and reduced tillage system with the applica-tion of maize plant debris for resistance evaluationwas described by B et al (1999) Theevaluation of a response to natural (the precedingcrop maize and reduced tillage system) and artificial

Fusarium spp infection is used for the classificationof resistance to FHB in wheat varieties included inthe Official Trials in the Czech Republic Though

DON contents after natural and artificial infectionwere significantly interrelated the obtained resultsmainly indicated the importance of simultaneousevaluation of FHB resistance under both condi-tions An advantage of the evaluation of varietyresponse under conditions of natural infection canlie in a better correspondence with conditions inagricultural practice but also data obtained afterartificial inoculation are evidently needed to dif-ferentiate better between varieties in resistanceto FHB (C et al 2008)

The resistance of studied wheat accessions couldbe described by disease score on head of Fusar-

ium damaged grains (FDG) reduction of kernelweight per head and DON content (Wśet al 2004) Another criter ion of the expression ofresistance to FHB is area under the disease progresscurve ndash AUDPC (S amp F 1977) L- et al (2004) found out that good correlationsbetween AUDPC and DON content are present atlow to intermediate disease levels Both FDG andreduction in thousand grain weight reflecting thekernel infection can be considered valuable traits(Š et al 2002) but FDG is more frequently usedas an indicator of FHB resistance (A et al 1999 I et al 2000 Wś et al 2002L et al 2003 M et al 2005)A new method for the evaluation of FHB infec-tion with respect to the plant phenological stage

called REML (a statistical method using a mixedmodel) was recently published by E et al (2008) in order to avoid an unintended selectionfor late-heading genotypes when evaluating FHBinfection in wheat

Pre-screening for specific partial disease resist-ance components against FHB using rapid in vitrotechniques and morphological characteristics maybe of use in achieving favourable combinations ofresistance components at a greater frequency inFHB resistance breeding In the detached leaf as-

say the partial disease resistance components suchas incubation period (period from inoculation tothe first appearance of symptoms) latent period(period from inoculation to sporulation) and lesionlength are measured (D amp C 1999B amp C 2004 B et al 2006)Significant relationships were found between theFHB and seedling blight resistance in vitro butsuch relationships were generally highly dependenton the cultivar and therefore the in vitro test islikely the best means for measuring componentsof FHB resistance andor genotype-specific resist-

ance components (B et al 2007)

Sources of natural resistance to FHB

Bread wheat (Triticum aestivum L)

Asia Wheats with the highest levels of FHBresistance come from the Far East region Apartfrom Sumai 3 a Chinese spring wheat cultivarused as a source of FHB Type II resistance globally

(eg B et al 2003 I et al 2008)other cultivars with relatively high FHB resistance




92

come from this region eg Chinese cultivars Ning7840 Ning 894037 (S et al 2003a Z etal 2003) Wangshuibai (Z et al 2004 Jet al 2005 M et al 2005 L et al 20042006 M et al 2006) Wuhan Japanese cultivarsNobeokabozu and Nyu Bai (Nyubai) or Koreancultivar Chokwang (Y et al 2005) Sumai 3is a highly FHB resistant Chinese spring wheatcultivar which was derived from a cross betweenItalian wheat Funo and Chinese landrace lineTaiwan Xiaomai Wheat Three major QTL forFHB resistance located on 3BS 6BS and 5ALhave been described in the crosses using thelines derived from Sumai 3 (W et al 1999 A et al 2001 B etal 2002 2003 Y et al 2003) The analysis of

SSR markers in the studies of S et al (2003a)Y et al (2006) and Y et al (2006) showedthat the major QTL on 3BS 6BS and 5AL origi-nate from Taiwan Xiaomai Wheat The QTL on3BS 6BS 5AL and 2DS in Sumai 3 are currentlythe most precisely characterised FHB resistanceQTL in wheat C et al (2006) preciselymapped the QTL on 3BS as a single Mendelianfactor with a bimodal t ype of inheritance andnamed it Fhb1 gene The Fhb1 gene is predomi-nantly responsible for Type II resistance and is

flanked by SSR markers Xgwm533 and Xgwm493 However L et al (2008) showed that inthe wheat line CM-82036 the Fhb1 gene confersdetoxification of DON and NIV Since the authorsproposed different mechanisms of detoxifica-tion for DON and NIV they suggested that Fhb1 may be a cluster of at least two resistance genesrather than a single gene Nevertheless they con-cluded that further investigations were necessaryto resolve the nature of Fhb1 C et al (2007) also finely mapped 6BS QTL conferringfield resistance as a Mendelian factor named

Fhb2 and flanked by SSR markers gwm133 and gwm644 The QTL on 2DS conferring Type IIIresistance was recently characterised by Het al (2008) who exploited collinearity betweenwheat and rice genomes and proposed that theQTL on 2DS can encode the wheat homologueof rice multidrug resistance-associated protein(MRP) The FHB QTL region on 2DS f lanks thereduced height gene Rht8 which might influencethe initial infection of FHB under field conditionsH et al (2008) thus showed a possible strat-

egy how to identify candidate genes underlyingsome FHB resistance QTL in wheat

Detailed description of new sources and resultsof mapping QTL for FHB resistance in the Asianwheat germplasm was provided by Y (2007)

America From the South American wheat genepool Brazilian spring wheat cultivars Frontanaand Encruzilhada are widely used as sources ofpredominantly Type I FHB resistance QTL (S- et al 2004) Relatively high disease incidence(quite low Type II resistance) and high percentageof spikes showing partial sterility (low Type VIIresistance) were however detected in Frontanaafter single floret inoculations (Š et al 2003)In the North American region the soft red winterwheat cultivar Ernie released in 1994 by the Uni-

versity of Missouri Agricultural Experiment Station(MK et al 1995) represents a promising

source of FHB resistance QTL other than Sumai 3(L et al 2005 2007)

Europe Genetic diversity of FHB resistance inEuropean winter wheat was recently evaluatedby Z et al (2008a) who concluded that thisstudy would assist in the selection of parental linesin order to increase the efficiency of breeding ef-forts for FHB resistance

There is an ample evidence available that someolder winter wheat cultivars eg KooperatorkaPraag 8 and Bizel possess relatively high levels

of FHB resistance and they may serve as alterna-tive sources of FHB resistance non-derived fromSumai 3 and other Asian wheats (S 1990Š amp S 1997 B et al 2008)However due to a low agronomic value thesetall materials have not been widely exploited inthe European wheat breeding programmes untilnow M and D (2006)screened 149 Russian and Ukrainian winter wheatcultivars for FHB resistance and found a moder-ate level of FHB resistance in the cultivars Hos-tianum 237 Odesskaya 16 and their derivatives

As already mentioned above the Rht-D1 locusfor semi-dwarf growth represents a major sourceof variation in the level of FHB resistance in theWestern European winter wheat gene pool (H- et al 2008)

Promising sources of moderate resistance toFHB (better adapted to the conditions that areprevalent in Europe) may be the Swiss cultivarArina (P et al 2004) German cultivarsCansas Petrus and Dream (K et al 2007S et al 2005) French cultivars Renan

(G et al 2003) and Apache (Het al 2008) Dutch cultivar Romanus (B et




93

al 2008) and Romanian cultivar Fundulea F201R(I et al 2002 S et al 2003b) Using thecultivars Centenaire Tulsa Tuareg Elegant andArina as sources of moderate type II FHB resist-ance specific hybrid combinations were identifiedwith this resistance type at the level equal to ormore resistant than the cultivar Arina (Zet al (2008b) B et al (2008) designated as resist-ant to FHB also the German cultivars HermannSobi Sokrates and Toras Swiss cutivar Runal andHungarian cultivar M2234 G et al (2007)investigated FHB resistance in 50 cultivars fromthe UK National List of winter wheat cultivarsapproved for sale and they found out that onlythree cultivars (Soissons Rht- B1 b Rht- D1 aSpark Rht-D1a and Vector) exhibited a higher

level of FHB resistance than the FHB-susceptiblecultivar Wizard A moderate level of resistanceto FHB was also found in the currently grownCzech cultivars Simila and Alana (C et al 2007) Sakura (Hčč et al 2007) (Figure 1)and Bakfis which possesses above all Type V FHBresistance (L amp P 2008) These cultivarsare promising sources of FHB resistance withadaptation to European conditions

The potential sources of FHB resistance adaptedto European conditions were traced back using

pedigree analysis (Figure 2) It is obvious fromthis analysis that it is very difficult to identify themost promising crossing types For example thecultivars Estica Hana Virtue and Caribo can beconsidered as parents occurring more frequentlyin pedigrees of moderately resistant germplasmWhile Estica and Hana can be considered moder-

ately or medium resistant to FHB (Hana only to theaccumulation of mycotoxins in grain) accordingto available literature (Vň et al 2001 Š etal 2002) semi-dwarf Virtue and tall Caribo donot evidently possess a desirable resistance level(B et al 1994 H et al 1999) Outof the five registered cultivars tested for resistanceto FHB that are descendants of Estica four cultivars(Simila Sakura Romanus and Centenaire) canbe considered moderately resistant while Drifterhas to be regarded as susceptible The cultivarHana was found to be a parent of four moderatelyresistant Czech varieties (Alana Alka Samantaand Sakura) but also of susceptible Saskia andVlasta and medium resistant Brea and Sulamit Asdocumented by S et al (2008) the Swiss

cultivar Arina was incorporated in the pedigree ofthe resistant line G16-92 which was used for thedevelopment of resistant German advanced breed-ing lines like HUS 690 and HUS 692 (L Hpersonal communication) Many crosses with Arinawere also performed in the Czech Republic butwithout evident success (L B personalcommunication) The non-adapted sources ofhigh resistance (eg Sumai 3) have not yet beensuccessfully used for introducing FHB resistanceinto adapted high-yielding European cultivars

although there is a long-time e vidence of thissource of resistance It can be derived from thisanalysis that the recent progress reached in thefield of FHB resistance was obtained by combiningeffective genes using the heterogeneous germplasmwith variable response to FHB for crossing Zet al (2008b) analyzed the inheritance of type II

Figure 1 Moderate resistance to FHB in the modern Czech winter wheat cultivar Sakura (on the left) followingspraying inoculation with the mixture of Fusarium graminearum isolates on the right response to FHB in the sus-ceptible Darwin




94

FHB resistance in selected sources of moderateresistance and they showed the importance ofboth general and specific combining ability ef-fects in the interaction with environment Specifichybrid combinations of Western European wheat

cultivars identified in the F1 generation exhibitedFHB resistance at a level equal to or even higherthan in the Arina cultivar

Sources of FHB resistance which do not come

from cultivated wheat

A possible source of FHB resistance for cultivatedwheat varieties belonging to both the hexaploidwheat (Triticum aestivum) and tetraploid wheat

(T turgidum L ssp durum) is represented by theirtetraploid relative wild emmer ndash T turgidum ssp

dicoccoides (Koumlrn ex Aschers et Graebn) ellO et al (2002) investigated FHB resistance inLangdon ndash T dicoccoides chromosome substitutionlines and identified a single major QTL on 3AS Qfhsndsu-3AS associated with an SSR marker Xgwm2

Later C et al (2007) proved that this locus wasnot homoeologous to the 3BS resistance locus Qfhsndsu-3BS of Sumai 3 and proposed new markers

Xfcp401 and Xfcp3972 flanking this locus A TypeI FHB resistance QTL was found on 4AS in a DHpopulation derived from T aestivum ssp macha (Dekapr et Menabde) by S et al (2005) isQTL co-segregated with SSR marker Xgwm165

Wheatgrass ( Lophopyrum = Thinopyrum sp) isanother source of FHB resistance that may be ex-ploited in wheat breeding programmes S and

O (2006) identified a QTL for FHB resistance on7EL of the genome of Lophopyrum elongatum and

Figure 2 Pedigrees of European winter wheat varieties showing moderate resistance to Fusarium head blight (in boldletters underlined)

Cappelle Desprez

Hybride-80-3 Etoile de Choisy

Maris Huntsman Durin Moisson Mironovskaya 808

NS-984-1

Carstens VIII Courtot Zenith

Ibis Carstens 854 Tring 5 Arina Cariplus Mex landrace

G16-92 Hussar

Festival Arminda Renan

HUS 690 HUS 692 Lambros

Armada Virtue Talent Flanders

NRPB-84-4233

Axial M Widgeon Obelisk Cebeco 451 Konsul

CB-306Y-70 Hana

M Hobbit Backa

Tower Mercia

Apache Tonic Moulin W-14-70-13-A-B Carimulti Kavkaz

Sturdy Centrum

Weihensteph 565-52 Caribo Spark NIC90-3390A Xanthos Hoeser Zenith

Cato RPB-49-75 Urban

Kronjuwel Hermann Runal

Vuka Beno Ares Marabu Vlasta Pegassos Arminda Viginta Alka Alana

Falke

Nimbus Kormoran Estica

Flair Piko Samanta

Petrus Disponent Orestis Monopol

Simila

Dream Solitaumlr Bakfis Romanus

Sakura

Glaucus




95

derived chromosomal substitution lines combiningchromosome 7E with the genetic background ofNing 7840 carrying the 3BS FHB resistance locusThe researchers reported that the 7E QTL derivedfrom Lophopyrum had a greater effect on the diseaseresistance than the 3BS locus in their experimentLater S and O (2007) used a set of chro-mosomal translocation lines between Thinopyrum

ponticum and the wheat cultivar Thatcher andidentified FHB resistance QTL (Qfhspur-7EL) on7EL in a 7el2 line which was flanked by codominantmarkers XBE445653 and Xcfa2240 which may beuseful for gene pyramiding techniques

Very recently the mammoth wildrye ndash Leymusracemosus (Lam) Tzvelev ndash has turned to containa new source of FHB resistance which can possibly

be utilized in the breeding programmes aimed at theimprovement of FHB resistance in cultivated wheatQ et al (2008) characterised a series of wheat- Leymus chromosomal translocation lines and described a newresistance gene Fhb3 located at 7Lr1S (short armof chromosome 7 originating from Leymus)

QTL for FHB resistance in common wheat

B et al (2009) have summarized the

results from 52 studies dealing with detection ofFHB resistance QTL in different wheat sources46 studies were carried out with hexaploid wheatsfour with tetraploid wheats (secondary gene poolfor common wheat) and two with wheat-relatedspecies ( Lophopyrum Thinopyrum) (tertiary genepool for common wheat) QTL detections are avail-able for the following resistance sourcesSources of Asian origin Sumai 3 Stoa Ning 7840

ND2603 CM-82036 Ning 894037 AlondraHuapei 57-2 Nyu Bai Patterson Wuhan 1 DH-181 AC Foremost W14 CS -SM3-7ADS CJ

9306 Gamenya Wangshuibai 5DL ChokwangSeri82

Sources of American origin Ernie FrontanaSources of European origin (winter wheats with a

moderate level of FHB resistance) Arina CansasDream Forno F201R Goldfield Hussar LynxPatterson Remus Renan Sincron Spark G16-92 NK93604

Barley ( Hordeum vulgare L)

In barley Type I resistance is important (resist-ance to initial infection) Type II resistance (spread

of infection via vascular bundles in rachilla andrachis within the spike) is less important in barleythan in wheat (Z et al 1999)

Two-rowed barley Gobernadora (Z et al 1999) and six-rowed nonmalting barley ChevronCI 4192 (a landrace from China) and Svanhals(a landrace from Sweden) exhibited low levelsof FHB (S 1999) and belong to themost widely used FHB resistant cultivars Chevronwas first introduced in Switzerland in 1914 It isresistant to FHB and kernel discoloration (KD)but its agronomical use is limited due to pooragronomic characteristics namely weak straw tallplants late heading thin kernels and low yieldTherefore great effort has been exerted to derivemore agronomically valuable lines (materials) from

Chevron (R et al 1999)D P et al (1999) identified QTL for

resistance to FHB DON accumulation and kerneldiscoloration (KD) in F47 breeding lines derivedfrom Chevron times M69 cross (M69 is an elite breedingline susceptible to FHB) S and S(2006) reported that the Chevron alleles at the QTLon chromosome 2H reduced FHB by 43 and thealleles at the 6H region by 22 while increasinggrain protein by 14 gkg These authors also men-tioned that marker-assisted selection (MAS) was

then generally used to select lines homozygousfor the resistant marker allele in the F2 generationprior to single seed descent (SSD)

H et al (2005) studied the nature of FHB resist-ance in a set of 125 RILs between Russia 6 (a two-rowed resistant cultivar of Russian origin) and HES4 (a six-rowed susceptible cultivar) ey identifiedthree FHB resistance QTL two of them located on2HL and the third on 5HS One of the QTL on 2HLwas coincident with the vrs1 locus which determinesthe inflorescence row type and the other QTL on2HL was positioned in the vicinity of the cleistogamy

locus (cly1 or Cly2) which determines inflorescenceopening us in accordance with the results pub-lished by Z et al (1999) it becomes evident thatthe FHB resistance QTL are often associated withQTL determining inflorescence characteristics inbarley Whether this association is due to linkageor pleiotropy remains to be elucidated RecentlyS et al (2008) showed on a set of RILs betweenHarbin two-row and other five two-rowed cultivarsthat no FHB resistance QTL was associated with thevrs1 locus but all RIL populations contained FHB

resistance QTL at cly1Cly2 locus (cleistogamylocus) on 2HL




96

B et al (2004) evaluated the level ofFHB resistance in a set of 143 barley lines contain-ing 88 European spring barley lines and cultivars33 accessions from the genebank at IPK Gaterslebenand 22 lines of North American origin The lineswith the lowest FHB severity were two-rowed lan-draces CIho 4196 and PI 566203 Apart from theSwiss six-rowed cultivar Chevron other Europeanspring barley cultivars with moderate resistance(although with 2ndash3 times higher AUDPC valuesthan the most resistant lines) were identifiedeg Hellana Pixel Secura Thuringia One of therelatively highly resistant cultivars was also anold hulless line HOR 1867 which is the old Czechcultivar Boehmische Nackte The researchers founda significant negative correlation (r = ndash055) be-

tween plant height and FHB resistance SimilarlyŠ et al (2004) found a high level of resistanceto an aggressive isolate of Fus arium culmorum in Chevron and CI 4196 as important sourcesof FHB resistance in the barley germplasm andmoderate resistance in the cultivars Jersey Olbramand Scarlett when examining FHB resistance andDON content in spring barley cultivars registeredin the Czech Republic

Most FHB resistant barley cultivars exhibit pooragronomical characteristics have poor malting

quality and are two-rowed (Z et al 1999)

Fusarium infection in oats ( Avena sativa L)

Oats is generally less susceptible to Fusarium infection than wheat or barley due to the paniclestructure of its inflorescence resulting in singlespikelet infections and slower infection spread Sev-eral Fusarium species infect oats ndash besides F culmo-rum and F graminearum the species F avenaceum(predominant in Poland) F langsethiae (commonin Scandinavia T amp N 2004) F poae

and F sporotrichioides (major sources of Fusarium infection in Canada) appear in regions where oatsis the staple cereal crop Infections caused by

F culmorum and F graminearum are prev alentin wet (humid) years while infections by F lang-

sethiae predominate in dry seasons F avenaceum produces predominantly moniliformin (MON)and F langsethiae and F sporotrichioides producehighly toxic type A trichothecene T-2 toxin andits deacetylated form HT-2 toxin Fusarium HeadBlight (FHB) is often difficult to assess in oats

The most reliable approach to the evaluation of Fusarium infection in oats is scoring of infected

seeds after harvest (expressed mostly as percentageof infected seeds) This is usually assessed by aldquofreeze and blotrdquo method (L 1966) Onehundred seeds are imbibed for 24 h on a wet filterpaper in a transparent plastic box and then frozento kill the germ After the freezing treatment boxeswith seeds are placed under the source of UV-light and a fraction of infected seeds is calculatedafter 10ndash12 days of UV-treatment Alternativelythe evaluation of mycotoxin content (especiallyDON) in kernels is carried out These two datado not often correlate The visible symptoms of

Fusarium infection in oats may include brownishnecrotic seeds or pink mycelium on glumes whichis visible only at high air humidity

ere exists a considerable genetic variation in oat

susceptibility to Fusarium infection and mycotoxinaccumulation among various cultivars (sources)Generally early cultivars and hulless (naked) cultivarsare less susceptible to infection Cultivars Bessin andLena seem to be highly susceptible to F graminearum infection during the germination stage (Bamp S 2008) Among Canadian oats nakedoat cultivar Boudrias and hulled cultivar Leggettreveal a relatively high tolerance to Fusarium in-fections (T et al 2008) A promising sourceof Fusarium resistance in the oat gene pool may be

represented by lines (materials) derived from Avena sterilis In 2001 a set of progeny lines from direct orback-crosses to Avena sterilis (accessions originatingin Eritrea or Israel or unknown) from Dr KJ Freyrsquosprogramme at Iowa State University were inoculatedin the field with F culmorum Single-seed descent(SSD) lines were developed from crosses with thecultivars Hurdal (Norwegian early maturing) andBelinda (Swedish late maturing) ey were tested(spray inoculations with F culmorum) in 2007 andrepeatedly in 2008

Fusarium infection in rye (Secale cereale L)

and triticale (Triticosecale)

Rye and triticale are generally less susceptible toFHB than wheat Cultivated rye and triticale varie-ties also exhibit a narrower range of FHB resist-ance than wheat cultivars do (see eg Met al 2001) M and P (1996)and M et al (2001) found twi ce loweramounts of DON in winter rye varieties than inwinter wheats under field conditions M

et al (2001) reported that under the same infec-tion conditions (an experiment carried out in a




97

growth chamber under controlled conditions)rye accumulated nine times lower DON amountsthan wheat and triticale had five times lower DONlevels than wheat after inoculation with the same

Fusarium culmorum isolate The inheritance ofFHB resistance in rye and triticale is determinedmainly by additive genetic effects According tofield tests carried out by M et al (2001)the winter rye variety Amando exhibited a rela-tively high level of FHB resistance

Triticale is generally less susceptible to infec-tions caused by Fusarium culmorum and Fusarium

graminearum than wheat and barley e predomi-nant Fusarium species found on triticale is Fusariumavenaceum (C et al 2000) the mainmycotoxin product of which is moniliformin (MON)

In triticale as well as in wheat and barley there existdifferences in susceptibility to Fusarium infectionsamong various cultivars I and I (2008) founda negative correlation between AUDPC and relativekernel weight expressed as of the control in tentriticale varieties grown in Romania after an artificialinoculation with F culmorum and F graminearum isolates ey concluded that the evaluation of theseparameters might be beneficial in selection of triticalegenotypes with improved resistance to FHB (eg the cultivars Gorun Lotru Plai and Stil of the ten

Romanian cultivars tested in this study)M et al (2004) determined a positivecorrelation between DON content and percentageof Fusarium damaged kernels (FDK) rating in a setof 55 triticale genotypes after an artificial inocu-lation by F culmorum isolate in a test consistingof six location-by-year combinations In repeatedtests the triticale variety Lasko showed a relativelyhigh level of FHB resistance (O amp W2001) O et al (2004) showed that addi-tive genetic effects are of major importance in theinheritance of FHB resistance in triticale

Characteristics of approaches usable

in breeding for FHB resistance

Recently much progress has been made in thebreeding of small grain cereals especially wheatand barley for resistance to FHB In wheat andbarley the search for sources of FHB resistancehas led to detection of several QTL conferring in-creased resistance to FHB In wheat however the

QTL with a strong effect on FHB resistance weredetected in exotic materials of Asian origin such as

Sumai 3 Wangshuibai etc which are geneticallydistant from the European wheat gene pool SeveralQTL affecting FHB resistance moderately havealso been detected and characterized in Europeanwheat cultivars However the effect of QTL fromthe European wheat gene pool is largely depend-ent on a given genetic background in a breedingpopulation Moreover the introduction of FHBresistance QTL originating from an exotic sourceinto an elite breeding material is often associatedwith the introgression of several other QTL whichnegatively affect agronomically important plantcharacteristics such as plant height inflorescencearchitecture heading date and final yield It isoften unresolved whether this coincidence of FHBresistance and adverse agronomical characteristics

is a consequence of pleiotropy (= two differentphenotypic effects of one locus) or linkage drag(= the presence of two different tightly linked lociwith different effects) This problem ndash pleiotropyor linkage drag ndash should be resolved by fine map-ping of FHB resistance QTL and their associationwith tightly linked molecular markers

Rapid progress in techniques of molecular biol-ogy has led to the construction of more convenient(and cheaper) SSR and STS markers than previouslyused RFLP and AFLP markers The association

of FHB resistance QTL with molecular markerscould be employed in marker-assisted selection(MAS) ie selection of those genotypes that carrya given molecular marker which is tightly linked tothe resistance QTL and thus probably also car rythis desirable QTL Since the inheritance of FHBresistance evidently seems to be predominantlygoverned by additive effects the stacking of non-homoeologous QTL from different sources in onebreeding population (so called QTL pyramiding)could lead to the construction of plant materialcarrying different QTL and exhibiting improved

FHB resistance when compared to the originalQTL sources (eg W et al 2007 2008)

A list of FHB resistance QTL identified in thewheat gene pool so far has recently been pub-lished in B et al (2009) Currentlyresearchers are trying to distinguish homoeologousQTL of different origin from non-homoeologousones One approach how to resolve this problemis meta-QTL (MQTL) analysis L et al (2009) analyzed FHB resistance QTL in popula-tions derived from Sumai 3 Wangshuibai and

Arina and they described 19 MQTL (QTL sharedby different genetic sources) located on 12 chro-




98

mosomes on all three genomes (six ten and threeMQTL on A B and D genomes respectively) SomeMQTL coincided with the previously identifiedFHB resistance QTL (eg regions on chromosomes3BS 5A and 6BS) some were newly identified It isimportant that resolving the genetic architecture ofFHB resistance by QTL meta-analysis may enablethe breeders to choose parents not comprising thesame resistance loci or QTL inter vals and exploitappropriate MQTL together with their respectivemarkers in MAS

In barley major QTL for FHB resistance havebeen co-localized to the loci determining rowtype (Vrs locus determining six-row or two-rowtype of barley inflorescence) and flowering type(Cly locus determining open ndash chasmogamous ndash

or closed ndash cleistogamous ndash type of flowering)on 2HL In addition to these major QTL otherQTL with minor effects have been detected on5H 6H etc

Based on several studies (reviewed for wheatin B et al 2009) it should be notedthat even the major QTL are responsible onlyfor a minor part of phenotypic variation in FHBresistance Thus it appears to be necessary inbreeding programmes to combine genotypic se-lection (selection of genotypes containing FHB

resistance QTL via MAS) with phenotypic selec-tion of genotypes re vealing a sufficient level ofFHB resistance in field tests ( W et al 2007L et al 2009) It is evident that the lat-est improvement in FHB resistance was reachedthrough the combination of favourable not yetidentified genes via phenotypic selection Cultivarswith a higher resistance level were the results ofbreeding programmes targeted on FHB resistancesuch as F201 R (I et al 1997) or the resistanceof these cultivars was detected during the breedingprocess It was shown by W et al (2007) that

both phenotypic and marker-assisted selection(MAS) are efficient tools in practical breedingand have a potential to reduce FHB symptomsand grain DON content The large range of vari-ation for FHB resistance was detected within themarker-based variant and selection gain was sub-stantially enhanced when additional phenotypicselection was accomplished in three steps It ishowever evident that the combination of MASand phenotypic selection can be highly expensiveparticularly when recurrent backcrossing is needed

with the use of non-adapted cultivars as resistancesources Phenotypic selection (evaluation of plant

response to Fusarium infection in field conditions)will probably play also a decisive role in the nearfuture which can be deduced from current breed-ing results However it is highly disadvantageousthat the cultivar response to FHB is strongly in-fluenced by environmental conditions Moreoveranother impediment for the breeders is representedby the multi-component quantitative expressionof this trait which often prevents detection ofsimilar results on reaction to FHB in differentexperiments Therefore the importance of usingreliable screening methods (described earlier inthis study) should be emphasized It should alsobe highlighted that these methods could be morefruitful when applied on a multi-environment andmulti-replication basis Seed shortage complicated

assessment of the resistance level and necessityto combine FHB resistance with other desirableagronomic traits like yield capacity quality andadaptation are the main obstacles to screening forFHB resistance since very early hybrid generationsAs demonstrated in detail eg by I et al (2002)for wheat and S and S (2006) forbarley the field evaluation of FHB resistance doesnot usually start earlier than in F5

Transgenosis is another approach how to im-prove the plant genetic background for higher

FHB resistance However the practical use oftransgenic techniques in breeding programmes islimited by the EU strict legislation Neverthelessa few studies on the improvement of FHB resist-ance via transgenosis have already been publishedeg B et al (2007) who introduced maizeantifungal gene b-32 under CaMV 35S promoterinto the wheat cultivar Veery and obtained trans-genic plants with enhanced Type II and Type III resistance and reduced level of kernel shatteringOther work published by C et al already in1999 reported enhanced resistance in the wheat

cultivar Bobwhite expressing a thaumatin-likeprotein from rice

Conclusions for resistance breeding

ndash It is evident from this study that sources ofhigh FHB resistance predominantly come from agermplasm not adapted to European conditionsor from distant wheat relatives which is a seriousimpediment to breeding progress For cultivation

in the Central European region only moderatelyresistant cultivars are available at present These




99

cultivars (breeding lines) were usually obtainedafter multi-step phenotypic selection through thecombination of favourable not yet identified genesderived from heterogeneous sources differing inresponse to FHB

ndash Owing to the existence of different resistancecomponents and quantitative expression of thistrait it is advantageous to perform simultaneousselection on a multi-environment and multi-replication basis both in conditions of artificialinfection and in conditions of natural infectionunder high disease pressure (after maize as apreceding crop and reduced tillage)

ndash Seed shortage multi-component quantita-tive nature high affection of the disease level byinteractions with the environment and necessity

to combine FHB resistance with other desirableagronomic traits prevent breeders from screeningfor this trait in the early hybrid generations on anindividual plant basis On the plant basis indi-rect selection for morphological or physiologicalcharacters associated with FHB resistance (likegreater plant height GA sensitivity earlinessnarrow flower opening loose spike branching)can be performed but it is unlikely that this se-lection will be successful in general

ndash Clear recommendations for breeding practice

how to select prospective hybrid combinationsare difficult to postulate However some studiesindicate that the analysis for a resistance levelin early hybrid generations (F1 or F2) could beadvantageous from these aspects (Z et al 2008b) The selection of resistant parents havingQTL on different chromosomes is now feasiblein wheat due to the use of a QTL meta-analysisapproach

ndash Even though much progress in the develop-ment of FHB moderately resistant cultivars hasbeen achieved until now mainly with the use of

conventional breeding methods (after crossingof adapted materials) new opportunities for theutilization of genetically distant sources possessinghigh FHB resistance may be given by the identifica-tion of QTL or genes underlying this resistance Itwas shown that the marker-based introgression ofresistance QTL in traditional breeding materialsmay result in a substantial increase in resistancewithin the shortest possible time However thisexpectation appears true only when the donor-QTLeffects are large enough (like on chromosomes 3BS

or 5A in wheat) to suppress several adverse effectsof a given genetic background

ndash To exploit the full range of quantitative vari -ation for FHB resistance phenotypic selection inthe field should follow marker-based selectionto bail out genetic variance that is caused by theother resistance genes undetected in QTL map-ping studies Though laborious time-consumingand restricted by the environmentally dependentsymptom expression phenotypic selection stillremains to be the most effective tool and pro-

vides an opportunity to select for other highlyeconomically important traits (like plant heightresistance to leaf diseases yield components etc)which is especially valuable when introgressingnon-adapted germplasm

Acknowledg ements The authors express deep

thanks to Dr M I NARDI Fundulea Romaniafor supplying important data and useful commentsSupported by the Ministry of Agriculture of the CzechRepublic Projects No QH 81293 and No QH 71213

References

A OA B D SS C S (2006) Pathogenic variationof Fusarium isolates associated with head blight of

wheat in Australia Journal of Phytopathology 154 513ndash521A NJ MC SP Z SL

(2000) Phytotoxicity of selected trichothecenes usingChlamydomonas reinhardtii as a model system NaturalToxins 7 265ndash269

A JA S RW L S W BLF AD C C M-S B MF J S QJ C PB F RC(2001) DNA markers for Fusarium head blight resist-ance QTLs in two wheat populations Theoretical andApplied Genetics 102 1164ndash1168

A E F E G T C J(1999) Fusarium head blight reactions and accumula-tion of deoxynivalenol (DON) and some of its deriva-tives in kernels of wheat triticale and rye Journal ofPhytopathology 147 577ndash590

B A E F G RJ L A G- DA S RS (2008) Phenotypic andmarker-assisted evaluation of spring and winter wheatgermplasm for resistance to fusarium head blightEuphytica 164 803ndash819

B G S G (1996) Variation in Fusarium gramine-

arum and cultivar resistance to wheat scab Plant Dis-ease 80 975ndash979




100

B G S G (2004) Management and resistancein wheat and barley to Fusarium Head Blight AnnualReview of Phytopathology 42 135ndash161

B C L C C E T TF F C M L E (2007) Fusar-

ium head blight evaluation in wheat transgenic plantsexpressing the maize b-32 antifugal gene EuropeanJournal of Plant Pathology 117 129ndash140

B H C J W H (1994)Free proline accumulation in wheat seedlings influencedby Fusarium culmorum infection and the pathogen me-tabolite deoxynivalenol Acta Physiologiae Plantarum16 111ndash116

B GT (1985) Head scab screening methods usedat CIMMYT In V RL K AR (eds)Wheat for More Tropical Environments CIMMYTMexico 169ndash173

B A B G G P X K G ACA P (2007) Fusarium graminearum-inducedchanges in gene expression between Fusarium headblight-resistant and susceptible wheat cultivars Func-tional and Integrative Genomics 7 69ndash77

B Aring S H (2008) Resistance to Fusarium infection in oats ( Avena sativa L) Cereal ResearchCommunications 36B 57ndash62

B AL R-F F B C (2008)Natural mechanisms for cereal resistance to the ac-cumulation of Fusarium trichothecenes EuropeanJournal of Plant Pathology 121 411ndash423

B JM L G F B C BMR A F R N P SD T M D FM (2007) Comparisonof commercial European wheat cultivars to Fusarium infection of head and seedling tissue Plant Pathology56 55ndash64

B DW MC SP A NJ P- RH D AE (2002) Inactivation of acytochrome P-450 is a determinant of trichothecenediversity in Fusarium species Fungal Genetics andBiology 36 224ndash233

B RA C BM (2004) Development andevaluation of an in vitro detached leaf assay for pre-screening resistance to Fusarium head blight in wheatEuropean Journal of Plant Pathology 110 91ndash102

B RA B C M F (2006) Com-ponents of partial disease resistance detected using adetached leaf assay in CIMMYT Fusarium head blightresistant wheat germplasm European Journal of PlantPathology 116 199ndash210

B H L M D ML S- M S B W K H L RP (1999) Resistenzzuumlchtung bei Weizen gegenuumlber

Aumlhrenfusariosen (Breeding of wheat for resistance toFusarium head blight) Bericht űber die Arbeitstagung

1999 der Vereinigung oumlsterreichischer PflanzenzűchterGumpenstein Austria 63-67 (in German)

B H L M H L D LS B S M R P(2002) Molecular mapping of QTL for Fusarium head

blight resistance in spring wheat I Resistance to fungalspread (Type II resistance) Theoretical and AppliedGenetics 104 84ndash91

B H S B H L G MA N L D M T SB L M (2003) Molecular mapping of QTLsfor Fusarium head blight resistance in spring wheat IIResistance to fungal penetration and spread Theoreti-cal and Applied Genetics 107 503ndash508

B H L L S B LM (2004) Variation for resistance to Fusarium headblight in spring barley Euphytica 137 279ndash290

B H L M S M Z- G H L M F T M G- NE N P (2008) Multi-environmentevaluation of level and stability of FHB resistanceamong parental lines and selected offspring derivedfrom several European winter wheat mapping popula-tions Plant Breeding 127 325ndash332

B H B T A JA (2009) QTLmapping and marker-assisted selection for Fusarium head blight resistance in wheat a review Plant Breed-ing 128 1ndash26

B LW W AH T TA (1975)Surveys of Fusaria associated with crown rot of wheatin eastern Australia Australian Journal of AgriculturalResearch 26 791ndash799

C J K P T M KM Gń P P A Ś-JA B J (2000) Moniliformin accumula-tion in kernels of triticale accessions inoculated with Fusarium avenaceum in Poland Journal of Phytopa-thology 148 433ndash439

C WP C PD L DJ K R FB V R M S G BS

(1999) Development of wheat scab symptoms is delayedin transgenic wheat plants that constitutively expressa rice thaumatin-like protein gene Theoretical andApplied Genetics 99 755ndash760

C X F JD H J S RW A TE EM K SF C X (2007) Saturationand comparative mapping of a major Fusarium headblight resistance QTL in tetraploid wheat MolecularBreeding 19 113ndash124

C J Š V M983323 E S S (2007)Resistance of winter wheat varieties registered in theCzech Republic to mycotoxin accumulation in grain

following inoculation with Fusarium culmorum CzechJournal of Genetics and Plant Breeding 43 44ndash52




101

C J Š V M983323 E (2008) Resistance ofwinter wheat varieties to Fusarium head blight underthe conditions of natural and artificial infection CerealResearch Communications 36B 87ndash90

C RJ (1981) Fusarium diseases of wheat and other

small grains in North America In N PE T- TA C RJ (eds) Fusarium ndash DiseasesBiology and Taxonomy Pennsylvania State UniversityPress University Park 39ndash52

C PA S DJ T J C SB-B A (2006) Fine mapping Fhb1 a majorgene controlling fusarium head blight resistance inbread wheat (Triticum aestivum L) Theoretical andApplied Genetics 112 1465ndash1472

C PA S DJ B-B A (2007)Mapping of Fhb2 on chromosome 6BS a gene control-

ling Fusarium head blight field resistance in breadwheat (Triticum aestivum L) Theoretical and AppliedGenetics 114 429ndash437

D P RC S KP C F M- GJ G-M M D-M RS DA R DC (1999) Quantitativetrait loci associated with resistance to Fusarium headblight and kernel discoloration in barley Theoreticaland Applied Genetics 99 561ndash569

B IA F RC S RW B- WA K SF (2003) Detection of QTLlinked to Fusarium head blight resistance in Sumai

3-derived North Dakota bread wheat lines Theoreticaland Applied Genetics 106 1027-1031

D H C BM (1999) Towards the develop-ment of a novel in vitro strategy for early screening of Fusarium ear blight resistance in adult winter wheatplants European Journal of Plant Pathology 105363ndash372

D FM B J C BM (2003) Influ-ence of climatic factors on Fusarium species pathogenicto cereals European Journal of Plant Pathology 109755ndash768

E K W F M T P HP (2008)

REML approach for adjusting the Fusarium head blightrating to a phenological date in inoculated selectionexperiments of wheat Theoretical and Applied Genet-ics 117 65ndash73

G L D F M J-Y B VN S B M G C T M (2003)Mapping of quantitative trait loci for field resistanceto Fusarium head blight in a European winter wheatTheoretical and Applied Genetics 106 961ndash970

G J K L S X O H (2005) DNAmarkers associated with low Fusarium head blight inci-

dence and narrow flower opening in wheat Theoreticaland Applied Genetics 110 1218ndash1225

G N B R J P K J N- P (2007) Evaluation and characterization of re-sistance to fusarium head blight caused by Fusariumculmorum in UK winter wheat cultivars Plant Pathol-ogy 56 264ndash276

G RS K HC (2004) Heading for disaster Fusar ium graminearum on cereal crops MolecularPlant Pathology 5 515ndash525

H H N N X D B T (2008) Dissect-ing of the FHB resistance QTL on the short arm ofwheat chromosome 2D using a comparative genomicapproach from QTL to candidate gene MolecularBreeding 22 71ndash84

H AJ J P H TW P DW(1999) Relationship between cultivar height and sever-ity of Fusarium ear blight in wheat Plant Pathology

48 202ndash208H TW R P DJ H (2003)Progress towards wheat varieties with resistance to Fusarium head blight Food Control 14 239ndash244

H J V H-H M T K VH J S G M V Z- G H L (2008) Inheritance of resistanceto Fusarium head blight in three European winterwheat populations Theoretical and Applied Genetics117 1119ndash1128

Hčč P H A C J (2007) Winterwheat Sakura Czech Journal of Genetics and Plant

Breeding 43 149ndash155H K K T S K T K (2005)

QTL analysis of Fusarium head blight resistance usinga high-density linkage map in barley Theoretical andApplied Genetics 111 1661ndash1672

I M I G (2008) Latest in breeding of resist-ance to FHB in Romanian triticale Cereal ResearchCommunications 36B 103ndash105

I M S NN I G (1997) Breeding wheatfor resistance to fusarium head blight in Romania InB HJ A F K WE B SPSMN A (eds) Wheat Prospects for Global Improve-

ment Proc 5th Int Wheat Conf Ankara Kluwer Aca-demic Publishers Dordrecht Publ No 6 87ndash92

I M G-S J K MG P (2000) Deoxynivalenol accumulation andother scab symptoms in six Romanian wheat genotypesinoculated with Fusarium graminearum MycotoxinResearch 16 15ndash22

I M S NN I G M M(2002) Approaches in breeding wheat for resistanceto Fusarium head blight (FHB) in Romania Petria12 67ndash72

I M SAacute NN I G C M (2008)Contributions to make modern Romanian bread winter




102

wheat more resistant to FHB Cereal Research Com-munications 36B 73ndash76

J G C P Q G B G W X W SZ B Z S L D (2005) QTLs for Fusariumhead blight response in a wheat DH population of

WangshuibaiAlondrarsquosrsquo Euphytica 146 183ndash191K A Z G W G M V

(2007) Effects of environment disease progress plantheight and heading date on the detection of QTLs forresistance to Fusarium head blight in a European winterwheat cross Euphytica 154 17ndash28

L P P J (2008) Winter wheat Bakfis Czech Jour-nal of Genetics and Plant Breeding 44 169ndash170

L T H YK K KH Y SH L YW (2002) Tri13 and tri7 determine deoxynivalenol and nivalenolproducing chemotypes of Gibberella zeae Applied and

Environmental Microbiology 68 2148ndash2154L M K R B H J RS R G H R P(2003) Fusarium head blight reactions and accumula-tion of deoxynivalenol moniliformin and zearalenonein wheat grains Cereal Research Communications31 407ndash414

L M B H K R SR G H R P (2004) Theeffect of inoculation treatment and long-term applica-tion of moisture on Fusarium head blight symptomsand deoxynivalenol contamination in wheat grains

European Journal of Plant Pathology 110 299ndash308L M K A S B B H

B F S R M F SW (2008) Investigations on the ability of Fhb1 toprotect wheat against nivalenol and deoxynivalenolIn 3rd Int FHB Symposium Szeged Cereal ResearchCommunications 36(Suppl B) 429ndash435

L WL F JD M S L DJ CPD G BS (2001) Isolation and characterizationof novel cDNA clones of acidic chitinases and β-13-glucanases from wheat spikes infected by Fusar ium graminearum Theoretical and Applied Genetics 102

353ndash362L T (1966) A modified blotter test for seed

health Netherlands Journal of Plant Patology 72 319ndash321

L F K ZX Z HL X SL W JZ TDG W JB Z CQ M ZQ (2004) Map-ping QTL associated with resistance to Fusarium headblight in the Nanda2419 times Wangshuibai population IType II resistance Theoretical and Applied Genetics109 1504ndash1511

L F X SL Z ZZ Z CQ K ZX

Y GQ T DG Z HL L CJ C Y W JBL QY M ZQ (2006) Mapping QTL associated with

resistance to Fusarium head blight in the Nanda2419 timesWangshuibai population II Type I resistance eoreti-cal and Applied Genetics 112 528ndash535

L ZZ (1985) Recent advances in research on wheatscab in China In V RL K AR (eds)

Wheat for More Tropical Environments CIMMYTMexico 174ndash181

L S A ZA M K AL (2005) Inherit-ance of Fusarium head blight resistance in the soft redwinter wheat Ernie Theoretical and Applied Genetics110 454ndash461

L S A ZA L H M T D GL MK- AL (2007) QTL associated with Fusarium headblight resistance in the soft red winter wheat ErnieTheoretical and Applied Genetics 115 417ndash427

L Y B H (2005) Interactions between

Barley yellow dwarf virus and Fusarium spp affectingdevelopment of Fusarium head blight of wheat Euro-pean Journal of Plant Pathology 113 283ndash295

L M S C-C M T (2009) Revealingthe genetic architecture of FHB resistance in hexaploidwheat (Triticum aestivum L) by QTL meta-analysisMolecular Breeding 23 473ndash488

M HX Z KM G L B GH C HGC ZX L WZ (2006) Quantitative trait loci forresistance to fusarium head blight and deoxynivalenolaccumulation in Wangshuibai wheat under field condi-tions Plant Pathology 55 739ndash745

M M B H G B LM M SA N R R P(2005) QTL analysis of resistance to Fusarium headblight in wheat using a lsquoWangshuibairsquo-derived popula-tion Plant Breeding 124 329ndash333

M SP D TV (2006) Genea-logical analysis of resistance to Fusarium head blightin Russian and Ukrainian cultivars of common wheatTriticum aestivum L Russian Journal of Genetics42 905ndash914

MK AL B JE T DN K KD(1995) Registration of ldquoErnierdquo wheat Crop Science35 1513

M Aacute (1995) Types and components of resist-ance to Fusarium head blight of wheat Plant Breeding114 377ndash386

M Aacute (2002) Theory and practice of thebreeding for Fusarium head blight resistance in wheatJournal of Applied Genetics 43A 289ndash302

M Aacute (2003) Breeding wheat for Fusariumhead blight resistance in Europe In L KJB WR (eds) Fusarium Head Blight of Wheatand Barley APS Press 211ndash240

M Aacute B T K G VM T B V J (2005) Common resistance




103

to different Fusarium spp causing Fusarium headblight in wheat European Journal of Plant Pathology112 267ndash281

M Aacute T B B T V M (2008)Breeding strategies against FHB in winter wheat and

their relation to type I resistance Cereal ResearchCommunications 36B 37ndash43

M T (1997) Breeding wheat and rye for resistanceto Fusarium diseases Plant Breeding 116 201ndash220

M T P J (1996) Correlations among Fusarium culmorum head blight resistance fungalcolonization and mycotoxin contents in winter ryePlant Breeding 115 347ndash351

M T R C L U S- M G HH (2001) Effects of genotypeand genotype-environment interaction on deoxyniva-

lenol accumulation and resistance to Fusarium headblight in rye triticale and wheat Plant Breeding120 97ndash105

M T H N S B OG R S R F (2004) Estimation ofdeoxynivalenol (DON) content by symptom rating andexoantigen content for resistance selection in wheatand triticale Euphytica 139 123ndash132

M JD A PG (1986) Degradation of de-oxynivalenol by suspension cultures of the Fusarium head blight-resistant wheat cultivar Frontana CanadianJournal of Plant Pathology 8 147ndash150

M RNG S AMQ L CA (1999)Molecular comparison of Fusarium populations causingeumartii wilt and dry on potato in Brazil FitopatologicaBrasiliensis 24 149ndash155

N P S G N SA C X (2008) Role of phytohormone signallingin resistance of wheat to Fusarium head blight CerealResearch Communications 36B 213ndash216

O G W G (2001) Genotypic and environ-mental variation of resistance to head blight in triticaleinoculated with Fusarium culmorum Plant Breeding120 297ndash300

O G H N M T (2004) Esti-mates of additive and dominance effects for Fusarium head blight resistance of winter triticale Plant Breed-ing 123 525ndash530

O CD K SF E EM S RW JLR (2002) Genetic dissection of a major Fusariumhead blight QTL in tetraploid wheat Plant MolecualBiology 48 625ndash632

P S S T T R M MW M K B S G (2004)QTL analysis of resistance to Fusarium head blight in

Swiss winter wheat (Triticum aestivum L) Theoreticaland Applied Genetics 109 323ndash332

P DW J P ML L (1995) Fusarium ear blight (scab) in small grain cereals - a review PlantPathology 44 207ndash238

P D F J W G K D LV L F B H O F

(2008) Marker-assisted wheat improvement creatingsemi-dwarf phenotypes with superior Fusarium headblight resistance Cereal Research Communications36B 153ndash155

Q LL P MO F B C PD GBS (2008) Molecular cytogenetic characterizationof alien introgressions with gene Fhb3 for resistanceto Fusarium head blight disease of wheat Theoreticaland Applied Genetics 117 1155ndash1166

R DC W RD D-M RS EL W JV (1999) Registration

of MNBrite barley Crop Science 39 290S K H K T K (2008) Detection of Fusar-ium head blight resistance QTLs using five populationsof top-cross progeny derived from two-row times two-rowcrosses in barley Molecular Breeding 22 517ndash526

S M Z G B HS G M T K V EE H L (2005) Molecular mapping of Fusariumhead blight resistance in the winter wheat populationDreamLynx Theoretical and Applied Genetics 111747ndash756

S M Z G S G M-

T K V E E H L (2008)Molecular mapping of quantitative trait loci for fieldresistance to Fusarium head blight in a European winterwheat population Plant Breeding 127 459ndash464

S HW C J J (1963) Factorsaffecting the resistance of wheat to scab caused byGibberella zeae Phytopathology 53 831ndash838

S G F RE (1977) e effect of nitrogenfertilization on the expression of slow-mildewing resist-ance in Knox wheat Phytopathology 67 1051ndash1056

S X O H (2006) Fusarium head blight resistancederived from Lophopyrum elongatum chromosome

7E and its augmentation with Fhb1 in wheat PlantBreeding 125 424ndash429

S X O H (2007) Molecular mapping of Thi-nopyrum-derived Fusarium head blight resistance incommon wheat Molecular Breeding 20 131ndash140

S X Z M L W O H (2003a) Detectionof Fusarium head blight resistance QTL in a wheatpopulation using bulked segregant analysis Theoreticaland Applied Genetics 106 1041ndash1047

S XR I M O HW (2003b) Quantitativetrait loci conditioning resistance to Fusarium head

blight in wheat blight in wheat line F201R Crop Sci-ence 43 850ndash857




104

Š V S E (1997) Evaluation of the resist-ance of winter wheat varieties to artificial infectionwith Fusarium culmorum in field conditions CerealResearch Communications 24 977ndash983

Š V S S S E C J (2002)

e effect of infection with Fusarium culmorum L ondeoxynivalenol content in grain of selected winter wheat varieties Journal of Applied Genetics 43A 319ndash332

Š V C J S S W HC J P J (2003) Evaluation ofwheat resistance to accumulation of Fusarium myco-toxin DON in grain In M C F F SAM (eds) Proc EUCARPIA Cereal Section MeetingNovember 21ndash25 2002 Salsomaggiore 261ndash266

Š V Tů L C J S S LL Kč L O J (2004) Effect of Fusarium

head blight on mycotoxin content in grain of springbarley cultivars Czech Journal of Genetics and PlantBreeding 40 91ndash101

Š V C J S S (2008) Assessing resist-ance to head blight in wheat cultivars inoculated withdifferent Fusarium isolates Czech Journal of Geneticsand Plant Breeding 44 43ndash59

S WG (1884) Diseases of Field and Garden CropsMacMillan and Co London 208ndash213 (cited afterM 1997)

S CHA (1990) The inheritance of resistance tohead blight caused by Fusarium culmorum in winter

wheat Euphytica 50 11ndash18S G N S A F S

B R J P N P (2008a) Map-ping of QTL associated with Fusarium head blight inspring wheat RL4137 Czech Journal of Genetics andPlant Breeding 44 147ndash159

S G N S A SJ L-W M W Y S J N- P (2008b) Susceptibility to Fusarium headblight is associated with the Rht-D1b semi-dwarfingallele in wheat Theoretical and Applied Genetics116 1145ndash1153

S G N S A HTW B R J P N P (2009)Semi-dwarfing Rht-B1 and Rht-D1 loci of wheat differsignificantly in their influence on resistance to Fusariumhead blight Theoretical and Applied Genetics 118695ndash702

S RW F RC C H (1997) Reac-tion of spring wheats incorporating Sumai-3 derivedresistance to inoculation with seven Fusarium speciesCereal Research Communications 25 667ndash671

S A C E T M G

N F S N P (2005) Identification oftype I resistance to Fusarium head blight control-

led by a major gene located on chromosome 4A ofTriticum macha Theoretical and Applied Genetics111 521ndash529

S BJ (1999) Combating Fusarium head blightof barley an emerging threat to malting barley quality

throughout the world EBC Congress 531ndash538S BJ S KP (2006) Breeding barley

for multiple disease resistance in the Upper Midwestregion of the USA Czech Journal of Genetics and PlantBreeding 42 79ndash85

S B L M G M S US J B H (2004) Molecularmapping of resistance to Fusarium head blight in thespring wheat cultivar Frontana Theoretical and Ap-plied Genetics 109 215ndash224

S T C J S S Š V (2008)

The occurrence of nivalenol producing Fusarium iso-lates on wheat in the Czech Republic Journal of PlantPathology 90(S-3) 38

T M (2004) Incorporation of Fusarium headblight resistance into European winter wheat breedingprogrammes In C SM B T WJ W RW (eds) Proc 2nd Int Symp FusariumHead Blight Incorporating the 8th European FusariumSeminar Orlando 195ndash196

T A M F JW R BGS ME (2008) Progress in assessing the impactof Fusarium head blight on oat in western Canada and

screening of Avena germplasm for resistance CerealResearch Communications 36B 49ndash56

T M N HI (2004) Fusarium langsethiae sp nov on cereals in Europe International Journal ofFood Microbiology 95 247ndash256

Vň M Tů L H H (2001) Reac-tion of winter wheat varieties to Fusarium graminearum and F culmorum in field infection trials and the efficacyof fungicides Plant Protection Science 37 66ndash73

V CA S W S S (2005) A secretedlipase of Fusarium graminearum is a virulence factorrequired for infection of cereals The Plant Journal42 364ndash375

V H-H H J H L K V R- F E E C H K HM T (2008) Effect of the Rht-D1 dwarfinglocus on Fusarium head blight rating in three segre-gating populations of winter wheat Plant Breeding127 333ndash339

W BL M-S B A JAS RW F RC (1999) RFLP mappingof QTL for Fusarium head blight resistance in wheatCrop Science 39 805ndash811

W YZ M JD (1988) Effect of Fusarium graminearum metabolites on wheat tussze in relation




105

to Fusarium head blight In K AR (ed) WheatProduction Constraints in Tropical Environments DFCIMMYT Mexico 239ndash250

W F K V E E G HH M- T (2007) Comparison of phenotypic and

marker-based selection for Fusarium head blight re-sistance and DON content in spring wheat MolecularBreeding 19 357ndash370

W F S CC K V E E S- M H L M T (2008) Marker-basedintroduction of three quantitative-trait loci conferringresistance to Fusarium head blight into an independentelite winter wheat breeding population Theoreticaland Applied Genetics 117 29ndash35

Wś H C J P J BśM B J (2002) Components of resistance

against Fusarium culmorum in spring wheat Journalof Applied Genetics 43A 345ndash354Wś H P J K Z (2004)

Scab response and deoxynivalenol accumulation inspring wheat kernels of different geographical originsfollowing inoculation with Fusarium culmorum Journalof Phytopathology 152 613ndash621

Y J B G S GE (2005) Novel quantitativetrait loci (QTL) for Fusarium head blight resistancein wheat cultivar Chokwang Theoretical and AppliedGenetics 111 1571ndash1579

Y ZP G J S DJ F G P-

JD MK IH (2003) Marker assistedselection of Fusarium head blight resistance genes intwo doubled haploid populations of wheat MolecularBreeding 12 309ndash317

Y Z G J P JD (2006) Geneticdiversity of resistance genes controlling Fusariumhead blight with simple sequence repeat markers inthirty-six wheat accessions from East Asian originEuphytica 148 345ndash352

Y T (1991) Natural occurrence of myco-toxins in small grain cereals (wheat barley rye oatssorghum millet rice) In S JE H

RS (eds) Mycotoxins and Animal Foods CRC PressBoca Raton 301ndash324

Y JB (2007) Identification of new sources and mappingQTL for FHB resistance in Asian wheat germplasmA dissertation Kansas State University ManhattanKansas 103pp

Y J-B B G-H C S-B B T (2006) Marker-

assisted characterization of Asian wheat lines for resist-ance to Fusarium head blight Theoretical and AppliedGenetics 113 308ndash320

Z G M M (2007) Molecular mappingof kernel shattering and its association with Fusariumhead blight resistance in a Sumai3 derived populationTheoretical and Applied Genetics 115 757ndash766

Z X Z M R L B G M H SOE G P L W (2004) Molecular characterizationof Fusarium head blight resistance from wheat varietyWangshuibai Euphytica 139 59ndash64

Z W-C K FL B G-H D LL BLK S NJ (2003) Validation of a major QTLfor scab resistance with SSR markers and use of mark-er-assisted selection in wheat Plant Breeding 122 40ndash46

Z H G L H P K A KD L Z P L S B T TV H (1999) Does function follow form Princi-pal QTLs for Fusarium head blight (FHB) resistanceare coincident with QTLs for inflorescence traits andplant height in a doubled-haploid population of barleyTheoretical and Applied Genetics 99 1221ndash1232

Z RS M H V B E R-R I (2008a) Evaluation of genetic diversity of Fusar-ium head blight resistance in European winter wheatTheoretical and Applied Genetics 117 813ndash828

Z RS M H V H J VB E R-R I (2008b) Combin-ing ability analysis of Fusarium head blight resistancein western European wheat lines Euphytica 162449ndash456

Received for publication June 30 2009

Accepted after corrections August 28 2009

Corresponding author

Ing J C CSc Vyacutezkumnyacute uacutestav rostlinneacute vyacuteroby vvi Drnovskaacute 507 161 06 Praha 6-RuzyněČeskaacute republikatel + 420 233 022 378 fax + 420 233 022 286 e-mail chrpovavurvcz




88

dough stage of kernel development Besides wateravailability also temperature aeration and light arecrucial climatic factors influencing the productionand dispersal of inocula growth competition my-cotoxin production and pathogenicity Fusarium species were found to differ in their climatic dis-tribution and in the optimum climatic conditionsrequired for their persistence but most speciesproduce inocula grow best and are more patho-genic to cereal heads at warm temperatures andunder humid conditions (D et al 2003)

Since the control of FHB spread by fungicidescrop rotation and deep ploughing need not alwaysbe an efficient means of crop protection breedingof FHB resistant cultivars (lines) represents themain approach how to eliminate yield losses and

contamination of grain by mycotoxins caused bythis disease

The problem of FHB in small grain cereals hasbeen reviewed by P et al (1995) M(1997) B and S (2004) and others Thesituation in the European wheat gene pool wa sdescribed precisely by M (2003) B- et al (2009) have recently summarizedthe problems of FHB in wheat and published a listof all FHB resistance QTL (quantitative trait loci)detected in the wheat gene pool so far Basic infor-

mation on pathogen biology including the descrip-tion of its life c ycle production of trichothecenemycotoxins and its population biology is given inthe review by G and K (2004)

In this review we would like to focus on the sourcesof FHB resistance and on progress in the breeding ofFHB resistant cultivars of small grain cereals withrespect to the Central European region

Manifestation of the disease and its spreading

The first record of FHB or scab comes fromS (1884) who reported the occurrence ofthis disease in England and attributed it to thefungus Fusisporium culmorum

The symptoms of the disease are quite similarin various small grain cereals the fir st symptomsappear as small patches of discoloration at the baseof the glume of a floret in a spikelet As the infec-tion progresses the patches of discoloration mayspread to the rachis and invade the whole spikeThe infection is visible as patches which first have

a bleached (light) appearance later they becomeorange salmon pink red or tan or they can even

change the infected seeds into hard dark brownobjects (called lsquotomb stonesrsquo) Recently Vet al (2005) have found out that secreted lipaseencoded by the FGL1 gene of Fusarium gramine-arum acts as a v irulence factor in the process ofwheat spike infection

The major potential risk for both humans andanimals caused by the infection is the productionof mycotoxins either oestrogenic zearalenone(ZEA or ZON) which is mainly found in infectedmaize ( Ze a mays ) ( Y 1991) or non-macrocyclic trichothecenes According to theirchemical nature the trichothecenes produced by

F culmorum and F graminearum are divided intotwo types type A toxins (HT-2 toxin T-2 toxin)and type B toxins including the most abundantly

produced mycotoxins in small grain cereals es-pecially deoxynivalenol (DON) its derivatives3-acetyl-DON and 15-acetyl-DON and nivalenol(NIV) and 4-acetyl-NIV called also fusarenone X(FX) Most Fusarium strains can simultaneouslyproduce multiple types of mycotoxins eg DONand ZEA In general both DON and NIV chemo-types were reported in Africa Asia and Europewhile only the DON chemotype was found in theUSA (M et al 1999 L et al 2002) Ourlatest studies performed in the territory of the

Czech Republic (S et al 2008) showed ver y rare occurrence of the NIV chemotype andhigh prevalence of the DON chemotype DONdiffers from NIV only in the absence of a hydroxylgroup at C-4 However this difference may haveimportant consequences for the fitness of theproducing organism as it alters the bioactivityand toxicity of trichothecenes For example thetoxicity of NIV is several times higher than thatof DON (A et al 2000) The molecularbasis of NIV or DON-producing abilities of theisolates lies in the functionality of the Tri13 gene

coding for a cytochrome P-450 enzyme which isfunctional in NIV-producing isolates while non-functional in DON-producing strains (B etal 2002 L et al 2002)

The infection is spread through both the asexualspores (macroconidia) and the sexual ascosporesby wind rain-splash dispersal or by arthropod

vectors such as mites (Siteroptes graminum) clo- ver leaf weevil ( Hypera punctata) grasshoppers( Melanoplus bivittatus) picnic beetles (Glischro-chilus quarisignatus) and even the common house-

fly ( Musca domestica) Crop residues on the soilsurface also represent an important reservoir of




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M Hobbit Backa

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Sturdy Centrum





Falke


Flair Piko Samanta


Simila


Sakura

Glaucus




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M Hobbit Backa

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Falke


Flair Piko Samanta


Simila


Sakura

Glaucus




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Cappelle Desprez



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M Hobbit Backa

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Sturdy Centrum





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Flair Piko Samanta


Simila


Sakura

Glaucus




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Flair Piko Samanta


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Sakura

Glaucus




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Cappelle Desprez



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M Hobbit Backa

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Flair Piko Samanta


Simila


Sakura

Glaucus




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Cappelle Desprez



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M Hobbit Backa

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Sturdy Centrum





Falke


Flair Piko Samanta


Simila


Sakura

Glaucus




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Kosová_Cereal Resistance to Fusarium Head Blight and Possibilities

Documents

Transcript of Kosová_Cereal Resistance to Fusarium Head Blight and Possibilities