THE Mlo GENE OF BARLEY

Current literature, mlo alleles, varieties and related topics

Last update 24.11.2008

                                                       page written and maintained till 2008 by Erik Schwarzbach                             

   e-mail:   eschwarzbach@iol.cz   

Important notice!
From 2009 on the Mlo page will be maintained
by Ralph Panstruga from the MPIZ Köln, Germany

under the editorship of Ralph, the Mlo page will continue to address molecular, biochemical, physiological and breeding aspects  of
the Mlo gene  and the pathogen properties, interacting with it, in an understandable language, comprehensive to a broad audience, summarise
existing knowledge and new findings, and be open for discussions.
.
I am sure, the Mlo-page will be under Ralph in good hands and be useful to its visitors. Good luck!

Erik Schwarzbach

                         

 General

 The function of the  Mlo  gene is generally understood as regulating a cell wall repair process, based on the growth of an intracellular papilla. which forms at the site of injury, that might be mechanical or  due  to penetration attempts by insects  or  fungi. Mutations  at  different  sites  of  the gene generally lead to its malfunction, resulting in excessive  papilla  growth. This causes a very high level of resistance to powdery mildew, since the papilla grows   usually  faster  than  the  parasite  penetrates.  The  recessive  alleles  of the Mlo gene, symbolized  "mlo", therefore behave as  very  efficient   genes   for  resistance  against  powdery  mildew, reducing the number of  mildew  conidia,  that  succeed  to  produce  a  colony,   approx.  500  times under laboratory conditions.  The absence of the regulatory function of the Mlo gene  is often  expressed  in mlo-mutants by adverse pleiotropic effects like large necrotic flecks on leaves of   adult  plants  and   reduced  yield. These side effects of mlo are considerably influenced by the genetic background and can be almost eliminated  by  breeding,  as  is  demonstrated  by the existence of more than 140 mlo cultivars in Europe.  The actual level of mlo-based resistance  under  field   conditions is only partly understood, depends on environmental factors and is difficult to reproduce. Exposure to elevated temperature or short term relief from water stress  increase the susceptibility of mlo plants to mildew by about one order of magnitude. Presently  the  mlo-gene  is  the only  source of resistance, against  which  highly virulent mildew races so far did not spread in agricultural environments. The mlo gene  is  therefore  the  most  widely  used source of resistance in spring barley varieties.  The mlo-cultivars cover in  central and western Europe, very roughly estimated, over 60% of the spring barley area.  The Mlo gene is cloned and it's  molecular  structure known. Mlo-orthologs have been found also in all  so far studied cereals and other higher plants.The Mlo-orthologs reverse on single cell basis the mlo based resistance of barley.  Genes,  similar  to  Mlo  with almost  identical  structure and protein properties, have been found in very different botanical taxa, including bryophyta, indicating its very ancient origin in evolution. Mlo analogs seem to be common  in  higher   plants.  Attempts  are  done  to  utilise  the  resistance  mechanism of mlo also in other species. Although  the   parasite   has  the  capacity  to  overcome  this kind of resistance and several genes for  partial  virulence  to mlo   have   been   detected    in   the   parasite,   the   evolution   of   the   mildew population towards high mlo-virulence proceeds under field conditions in small steps and slowly. The so far known most adapted mlo-virulent mildew race HL-3, produced in  the  laboratory  about 30 years ago, and an old field isolate from Japan produce about 50 - 100 times more colonies on  leaves  of  mlo-plants  than  non-adapted mildew races. A recent analysis used molecular technique to compare the mlo virulent HL-3 and its near isogenic avirulent parent line GE-3. During appresorium formation and penetration the activity of two novel Blumeria graminis genes was several times higher in HL-3 than in GE3. It was also shown, that the parasite genes responsible for higher mlo-virulence are not identical in the Japanese isolate and HL3. This makes evolution towards higher mlo virulence likely. Recently the level of mlo virulence in  some field pathotypes,  collected  in  Europe,  approached 15%, relative to the fully compatible interaction, if tested on detached primary leaf leaf segments  on agar. But the mlo virulence of these isolates on later leaves and under field conditions is considerably lower. The selection pressure for increased mlo virulence is growing, since the majority of new spring barley cultivars in Europe has mlo and the proportion of mlo  on the spring barley acreage exceeded already 50%, see the table  multiplication areas of mlo-cultivars on this page.

(Summary by the page author, updated 10.1.2006)


Alleles of the Mlo locus
Many data and links by courtesy of Dave Mathews, GrainGene
link to the GrainGene Database: click here
 (still incomplete - your help to improve the table is welcome)

Allele	Mutant (mother variety)	found by	Stock	Reference	Mutations at aminoacid level
Mlo	Wild type		all non-mlo barleys		Molecular structure in all wild-type plants identical
mlo1	M 66 ( Haisa)	Freisleben & Lein 1942	CIho15217	BGN-8-168	Trp162 → Arg
mlo2	H3502 (Vollkorn)	Hänsel 1971	CIho15223	BGN-8-168	?
mlo3	MC20(Malteria Heda)	Favret 1960	CIho15225	BGN-8-168	frame shift aft. Phe395
mlo4	SR1=REFOMA (FOMA)	Wiberg 1973		BGN-8-168	frame shift aft. Trp159
mlo5	Risoe 5678 (Carlsberg II)	Jorgensen1975	CIho15219	BGN-8-168	Met1 → Ileu
mlo6	Risoe 6018 (Carlsberg II)	Jorgensen1975	CIho15220	BGN-8-168	?
mlo7	Risoe 7085 (Carlsberg II)	Jorgensen1975	CIho15622	BGN-8-168	Gly226 → Asp
mlo8	Risoe 7372 (Carlsberg II)	Jorgensen1975	CIho15221	BGN-8-168	Met1 →Val
mlo9	SZ5139b (Diamant)	Schwarzbach 1967	CIho15227	BGN-8-168	Arg10  →Trp
mlo10	SR7 (Foma)	Wiberg 1973		BGN-8-168	missing Phe182 and Thr183
mlo11	Grannenlose Zweizeilige  EP79, L92, L100 & othera)	Collected in Ethiopia 1937-38, reviewed by Hoffmann & Nover 1959	Hor 2937, Hor4408, Hor3028, CI14017 & other	discussed by Jorgensen 1992	inserted repetitive Mlo-fragments in front of intact Mlo
mlo12	Mut. No. 4122 (Elgina)	Hentrich 1977		IAE-77-333	F240  → L
mlo13	Mut. No. 2018 (Plena)	Hentrich 1977		IAE-77-333	Val50  → Glu
mlo14	Mut. No. 2029 (Plena)	Hentrich 1977		IAE-77-333	?
mlo15	Mut. No. 4123 (Plena)	Hentrich 1977		IAE-77-333	?
mlo16	Mut. No. 2267 (Alsa)	Hentrich 1977		Pifanelli et al 2002	nucleotide substitution
mlo17	Mut. No. 2034 (Plena)	Hentrich 1977		IAE-77-333	Ser31   → Phe
mlo18	Mutant ML-3A (Azuma Golden)	Yamagushi & Yamashita 1979		JJB-29-217	?
mlo19	Mutant ML-4F (Fuji Nijou)	Yamagushi & Yamashita 1979		JJB-29-217	?
mlo20	Mutant ML-9F (Fuji Nijou)	Yamagushi & Yamashita 1979		JJB-29-217	?
mlo21	Mutant ML-13F (Fuji Nijou)	Yamagushi & Yamashita 1979		JJB-29-217	?
mlo22	Mutant B1012 (Bomi)	Röbbelen & Heun 1990		MBC-91-93	?
mlo23	Mutant B1101 (Bomi)	Röbbelen & Heun 1990		MBC-91-93	?
mlo24	Mutant B1865 (Bomi)	Röbbelen & Heun 1990		MBC-91-93	?
mlo25	Mutant N105 (Bomi)	Röbbelen & Heun 1990		MBC-91-93	?
mlo26	Do 2118 (Plena)	Habekuss & Hentrich 1988		Büschges et al 1997	Leu270 → Phe
mlo27	Do 2021 and 8 other mutants	Habekuss & Hentrich 1988		Pifanelli et al 2002	G318  → E
mlo28	Do 4228	Habekuss & Hentrich 1988		Pifanelli et al 2002	T222  → I
mlo29	see Pifanelli 2002	see Pifanelli 2002		Pifanelli et al 2002	P334  → L
mlo30	Do 2234, 2235	Habekuss & Hentrich 1988		Pifanelli et al 2002	del. of 6 am.acids
mlo31	URS1 (Ursula)	Molina-Cano et al. 2003		Molina-Cano et al. 2003	?
mlo32	PRU1 (Prudentia)	Molina-Cano et al. 2003		Molina-Cano et al. 2003	?


a) The accessions are possibly of same origin, the differing names may come from drifting through collections and nurseries, see Jorgensen  1992

 
 
 
 

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Molecular structure of the Mlo gene
(summary of important findings)

The wild type Mlo-allele is   1599 bp long, the coded protein contains 533 aminoacids (Büschges et al. 1997).
The deduced Mlo protein has a predicted molecular weight of 60 kDa and is  predicted to be membrane-anchored by seven membrane-spanning helices. (Internat. Patent Publ. No. WO9804586)

 Graphic representation of the deduced 7TM topology of barley Mlo (Devoto et al 1999)
kindly permitted by the Journal of Biological Chemistry



The   lipid plasma membrane bilayer is represented by a grey horizontal bar.
       Circles with letters represent amino acids identified by the single letter amino acid code.
For the exact nucleotide or aminoacid sequence see Büschges et al. 1997 or the  documentation
to the Mlo patent WO9804586

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Genetic maps of the Mlo locus

The Mlo gene is located  in the mid of the long arm of  Chromosome 4H.In the last years mapping concentrated on molecular marker maps, which are much more detailed than morphological marker maps, but not generally understandable. Below you can see thumbnails of a morphological marker map and a RFLP marker map.

 
To see these and some other  maps in full size or to find links to more maps CLICK HERE
Recently P.M. Hayes et al., 2000, (Oregon State Univ.)  assembled a page summarizing QTL and physical locations for all 7 chromosomes, see http://www.css.orst.edu/barley/nabgmp/qtlsum.htm.  To see the chromosome 4 map  (PDF-format) click here

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The Mlo patent

The defined polynucleotide constituing the Mlo gene has been patented under WO9804586
in 1998 for the applicant

     INNES JOHN CENTRE INNOV LTD (GB); PANSTRUGA RALPH (GB); BUESCHGES RAINER (GB); SCHULZE LEFERT PAUL MARIA JOSE (GB)

The patent involves 73 claims, in summary covering the isolated Mlo polynucleotide and all it's possible alterations by mutation, addition, excision etc. or parts of it, all methods and vectors for heterologous incorporation of such a polynucleotide or testing it's presence using compounds reacting with it or it's products, and transgenic plants with incorporated said polynucleotide and their descendents (for details click the link).

The patent description contains also pictures with the exact nucleotide and aminoacid sequences, of morphological, rflp,  aflp and other fine structure maps.

Fortunately, the patent does not preclude the free use of mutant or spontaneous mlo-alleles in conventional breeding or testing the presence of mlo in inoculation experiments with mlo-virulent mildew.

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Barley varieties with mlo-based mildew resistance

Area of mlo-varieties grown in western and central Europe: there are no exact data available. Several national variety offices publish data on cultivar areas grown for seed certification. From these approximately the percentage of  mlo-varieties actually grown can be estimated. Not all sown seed is certified, but this uncertaneity is with both mlo- and non-mlo-cultivars . The following table summarises the data available till now. Data form mediterranean countries  and east baltic states are not included, since mildew is there usually not a problem and these areas are epidemiologically quite different from central and western Europe

1. recently released cultivars added to the main table:

(Provisional. Please help to complete/correct the data)

 To see the complete table with all 143 mlo cultivars click here: MAIN CULTIVAR TABLE
(there are most likely more, since some might be officially listed as having unknown resistance)

Thanks to John Clarkson & Sue Slater, NIAB, Gerardo Arias, EMBRAPA, J.H. Czembor & Anna Tratwal IHAR, A. Dreiseitl, VUO Kroměříž, P. Gymer, Agrifusion, I. Langer, SELGEN, Bill Thomas, SCRI, L. Jestin, INRA, Helge Skinnes, AUN, M. Oberforster, AGES Vienna, D. Jurecka, UKZUZ, Max Baumer, LFL, R. Habgood, Nickerson and B. Schinkel, Lochow-P, Morten Rasmussen, SvalöfWeibull and Vera Cervena, Piestany, for their contributions to the table.

 A database of barley cultivars with many useful information, also on pedigrees, was set up by Adrian Newton The link to the database is http://wwwinternal.scri.sari.ac.uk/cprad/

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Determination of the presence of mlo in barley lines

There are three possible ways (at least).
1. The older way, using artificial infections of approx 10 days old seedlings with partially mlo-virulent mildew cultures. It is  based on the gene-for-gene relationship. The cultures should  be virulent also to possibly present other resistance genes,  which otherwise would mask the presence of mlo. This way is cheap, large numbers of  individuals or breeding lines can  be tested simultaneously,  but the test  takes  a minimum of four weeks. Link to a laboratory offering such service:  CLICK HERE

2. The use of markers closely linked to mlo and detectable biochemically. The method is more expensive, but faster, depending on the stage in which the marker is expressed (seed, germling, seedling). The method is  reliable, so far no (rare) recombination have occurred in the tested genotype between  mlo and the marker.

3.  Molecular techniques, based on direct determination of the nucleotide sequence within the Mlo locus or on  reactions with a compound, reacting with mlo nucleotides or their products. This is the fastest, most precise,  but most expensive solution  and requires the licensed use of the Mlo-patent.

Laboratories able to offer such servises are welcome to communicate their links, which would be published on this page.

Determination of mlo9 and mlo11 alleles by means of molec. markers in barley lines is offered to breeders by
      EpiGene GmbH,
      Biotechnologie   im    Pflanzenschutz                                                                                                                 85354  Freising,  
      Tel: 0049(0)8161-713189                                
      mail:  Friedrich.Felsenstein@EpiLogic.de

Detection of mlo presence using artificial infections with mlo-virulent mildew is available in my laboratory, which is :involved mainly in monitoring  occurrence and frequernce of partially mlo virulent airborne mildew

      Barley mildew laboratory
      Erik Schwarzbach
      CZ-67172 MIROSLAV, Czech Republic
      Phone: +420  515 333 878
      mail:   eschwarzbach@iol.cz
       

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Information on topics related to mlo

A center of Mlo-related studies based on molecular biology exists at the
Max Planck Institute for Breeding Research with the team of P. Schulze Lefert and Ralph Panstruga,
contact:

Ralph Panstruga
Max-Planck-Institute für Züchtungsforschung
Carl-von-Linné-Weg 10
D-50829 Köln
Telefon: +49(221) 5062 672
Fax: +49(221) 5062 674
panstrug@mpiz-koeln.mpg.de

Adaptation of the mildew population in Europe to mlo

So far, only partial virulence to mlo at low frequency was detected in the agricultural environment in Europe. Several isolates were independently found with increased levels of  virulence for mlo, none of them however reached the virulence level of the mildew isolate HL-3, developed in the laboratory  (see Atzema 1998, Schwarzbach 1998 and the UK-Cereal pathogen Virulence Survey 1999).

The UKCPVS 1999 Annual Report summarises:
"MILDEW OF BARLEY.
Virulence frequencies were similar to previous years, with Vra, Vg, V(CP), Va12 and Va7 present at high levels in all populations. Frequencies of Vh, Va7, VLa and V(Ab) varied over the season, reflecting the presence of Mla7 and Mlh in winter cultivars and MlLa and Mla1 in spring cultivars. The trend towards more complex isolates continued and the diverse nature of the barley mildew population was sustained. The population was again capable of infecting the majority of cultivars in NIAB Recommended List trials. Only spring barley cultivars with mlo genes were mildew-resistant, although some isolates continued to give increased levels of infection on the mlo differentials Apex and Riviera in tests. No new resistances were identified, although Vanessa may carry new, unknown resistance. Due to the very low incidence of barley mildew in N. Ireland, no isolates were obtained."
Recently, however, in some field pathotypes collected in the UK and the Czech Republic a level of mlo virulence approaching 15%, relative to the fully compatible interaction, has been found.  Since those isolates have a very broad virulence spectrum, they might spread and threaten to some extent barley growing.

The next  pictures give an impression of the extent of specific mlo-virulence that exists in three mildew cultures in experiments with simultaneously inoculated triplets of primary leaf segments of APEX (mlo),
DIAMANT (Mlo) and APEX (mlo).


 Click a high resolution image  Click a high resolution image    Click a high resolution image

 You are invited to contribute more information (click here)

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Discussions related to mlo, open to everybody
Feel free to comment mlo-topics, ask questions or present suggestions. Just email your ideas, which then will be, if you want, put here onto the mlo page.

To contribute any information concerning mlo, send please a  mail

** Literature on mlo from 1990 to 2005

** Handling of new Literature since 2006

** Searching the web using SCIRUS

1. Papers referred to on the mlo-page and frequently cited older papers

 

Favret E.A.,1960. Induced mutations for resistance to diseases. Genetica agraria,13,1-26
    (The  mlo-mutant MC20  described, allelic with M66. Allele mlo3)

Freisleben, R. und Lein, A.,1942. Ueber die Auffindung einer mehltauresistenten Mutante nach
     Roentgenbestrahlung einer anfälligen reinen Linie von Sommergerste. Naturwiss.30,608.
     (first description of a x-ray induced mlo-mutant M66 carrying the mlo1-allele)

Hänsel H.,1971. Experience with a mildew-resistant mutant (mut.3502) of ´Vollkorn´ barley induced
    in 1952.  In: Mutation breeding for Disease Resistance, IAEA-PL-412/13,125-129.
    (Mutant carries the mlo2-allele)

Habekuss A. and Hentrich W.,1988. Charakterisierung funktionell verschiedener ml-o Mutanten durch
    Primärinfektion, Pustelwachstum, Inkubationszeit und Befallsverlauf.
    Tag.-Ber. Akad. LWW DDR  Berlin, 272:229-237.

Hentrich W.,1977. Mildew-resistant mutants in spring barley
    Induced Mutations Against Plant Diseases  IAEA/FAO Vienna, 1977, 333-341

Hoffmann W. & I. Nover, 1959. Ausgangsmaterial für die Züchtung mehltauresistenter Gersten.
    Z. Pflanzenzüchtung 42,68-78
    (origin of mildew resistant landraces from the german Ethiopia-expedition 1937-38)

Jorgensen H.J.,1975.Identification of powdery mildew resistant barley mutants and their allelic
    relationships.  Barley Genetics III (Proc. sympos. Garching, 7-12 July 1975), 446-455.

Joergensen J.H. and Jensen H.P.,1979: Inter-allelic recombination in the ml-o locus in barley.
    Barley Genet. Newsl. 9:37-39.
     (functional allelism of non identical mlo-mutations within the same locus]

Nover I. and Schwarzbach E.,1971: Inheritance studies with a mildew resistant barley mutant.
    Barley Genetics Newsl. 1:36-37.
    (discovery of alelism between mutant mlo and mlo in an ethiopian landrace]

Schwarzbach E.,1967. Recessive total resistance of barley against mildew (E.graminis D.C. f. sp.
    Marchal as a  mutation induced  by Ethylmethansulfonate (in czech).
    Genetics and Plant Breeding (Prague),3,159-162.
    (Mutant from which ALEXIS  with mlo9 was developed - the mlo-variety with the highest acreage so far)

Schwarzbach E.,1975: The pleiotropic effects of the mlo gene and their implications in breeding.
    Barley Genetics III (proc. symp. Garching 1975):440-445.
    (genetic background may compensate to a large extent unfavorable side effects of mlo.
      See also Bjoernstad & Aastveit 2000)

Schwarzbach E.,1979: Response to selection for virulence against the ml-o based mildew resistance in
    barley,  not fitting the gene-for-gene hypothesis. Barley Genetics Newsl. 9,85-88.
    (development of the partially mlo-virulent culture HL-3 in the laboratory]

Skou, J.P.,1985: On the enhanced callose deposition in barley with ml-o powdery mildew resistance.
    Phytopathol. Z. 112,207-216,1985.
    (formation of papillae  in mlo and non-mlo barley epidermis after mechanical or fungal injury)

Wiberg A.,1973. Mutants of barley with induced resistance to powdery mildew. Hereditas 75,83-100

Yamaguchi I & Yamashita A ,1979.  Induced mutation of two-rowed barley resistant to powdery
    mildew, Erysiphe graminis f.sp. hordei. I. Comparison of effects of gammarays and ethyleneimine in
    induction of resistant mutation. Japanese Journal of Breeding  29, 217- 227

 

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2. Literature on mlo from 1990 to 2005

click  90 - 91 -  92 - 93 - 94 - 95 - 96 - 97 - 98 - 99 - 00 - 01 - 02 - 03  - 04 - 05 or browse through

1990
Bayles C.J., Ghemawal M.S., Aist J.R., 19*90. Inhibition by 2-deoxy-D-glucose of callose formation,
      papilla deposition, and resistance to powdery mildew in ml-o barley mutant. Physiol. Mol. Plant
      Pathol. 36,63-72

Bjoernstad Ĺ. and Aastveit K, 1990: Pleiotropic effects of the ml-o mildew resistance gene in
    barley in different genetical backgrounds. Euphytica 46,217-226.

Brown J.K.M. and Wolfe M.S.,1990: Structure and evolution of a population of Erysiphe
    graminis f.sp. hordei.. Plant Pathology 39,376-390.

Kjaer B., Jensen H.P., Jensen J. and Jorgensen J.H., 1990. Associations between three ml-o powdery
    mildew resistance genes and agronomic traits in barley. Euphytica 46:185-193

Pedersen L.H., 1990. 1,3-ß-glucansynthase activity and callose synthesis in barley mlo mutants and
    mother varieties, Plant Physiol. 79 (1990) 02 (Abstract).

Röbbelen G. and Heun M., 1990. Genetic analysis of partial resistance against powdery mildew in
     induced mutants of barley. In:  Plant Mutation Breeding for Crop Improvement.
     Symposium  IAEA/FAO Vienna, 18-22 June 1990, 93-111

1991
Aist J.R. and Bushnell W.R.,1991: Invasion of plants by powdery mildew fungi and cellular
    mechanisms of resistance. In: Cole G.T., Hoch H.C. (eds.), The spore and disease
    initiation in plants and animals. Plenum Press N.York, 321-345.

Andersen L., 1991: Mlo aggressiveness in European barley powdery mildew. In:
    Joergensen J.H.(Ed.), Integrated Control of Cereal Mildews: Virulence Patterns and their
    Change. Risoe Natl. Lab., Roskilde, 1991.

Carver T.L.W., Robbins M.P., Zeyen R.J., 1991. Effects of two PAL inhibitors on the susceptibility and
     localized autofluorescent  host cell responses of oat leaves attacked by Erysiphe graminis DC,
     Physiol. Mol. Plant Pathol. 39 (1991) 269-287.

Joergensen J.H.,1991: Mechanism of Mlo resistance to barley powdery mildew. Sveriges
    Utsädesförenings Tidskr. 2, 80-84.1992

Hentrich W., and Habekuss A.,1991. Untersuchungen an heteroallelen mehltauresistenten Mutanten
    des mlo-Locus der Sommergerste. Vorträge für Pflanzenzüchtung 19,311-312.

1992
Andersen L. and Joergensen J.H.,1992: Mlo aggressiveness in barley powdery mildew.
    Norweg. J. Agric. Sci. 7:77-87.

Carver,T.L.W., Zeyen R.J., Robbbins M.P., Dearne G.A.,1992: Effects of PAL inhibition on oat,
    barley and wheat cell response to appropriate and inaproppriate formae speciales of
    Erysiphe graminis DC. Physiological and Molecular Plant Pathology 41,397-409.

Joergensen J.H.,1992: Discovery; characterisation and exploitation of Mlo powdery mildew
    resistance in barley. Euphytica 63:141-152,1992.

1993
Felsenstein, F.G.; Fischbeck, G.: Sommergerste: mlo-Mehltauresistenz gefährdet?
    Pflanzenschutz-Praxis 1993, Nr. 2, S. 30-31.

Wolter,M., Hollricher K., Salamini F., Schulze-Lefert P.,1993: The mlo resistance alleles to
    powdery mildew infection in barley trigger a developmentally controlled defence
    mimic phenotype. Mol. Gen. Genet. 239,122-128.

Zeyen R.J., Ahlstrand G.G., Carver T.L.W., 1993: X-ray microanalysis of frozen-hydrated, freeze-dried,
    and critical point dried leaf specimens: determination of soluble and insoluble chemical elements at
    Erysiphe graminis epidermal cell papilla sites in barley isolines containing Ml-o and ml-o alleles.
    Canadian Journal of Botany 71,284-296

1994
Atzema J., Limpert E., and Wolfe M.S.,1994: Testing isolates of barley powdery mildew for mlo
    virulence. In: Limpert E., Finckh M.R. and Wolfe,M.S. (Eds.), Integrated control of cereal
    mildews and rusts: towards coordination of research across Europe. European Commission,
    Directorate-General XII, Brussels, 47-48.

Carver,T.L.W., Zeyen R.J., Bushnell W.R., Robbbins M.P.,1994: Inhibition of phenylalanine
    ammonia lyase and cinnamyl-alcohol dehydrogenase increased quantitative
    susceptibility of barley to powdery mildew (Erysiophe graminis DC). Physiological and
    Molecular Plant Pathology 44,261-272.

Clark T.A., Zeyen R.J., Smith A.G., Carver T.L.W. and Vance C.P.,1994: Phenylalanine ammonia lyase
    mRNA accumulation, enzyme activity and cytoplasmic responses in barley isolines differing at Ml-a
    and Ml-o loci, attacked by Erysiphe graminis f.sp. Hordei.
    Physiological and Molecular Plant Pathology 44,171-185

Joergensen J.H.,1994: Genetics of Powdery Mildew Resistance in Barley
    Critical reviews in plant sciences 13, 97-119.1995

Schulze-Lefert, P., Freialdenhoven, A., Scherag, B. and Görg, R. 1994. Dissection of resistance pathways in
     barley to powdery mildew attack. M.J. Daniels et al. (eds.),
     Advances in Molecular Genetics of Plant-Microbe Interactions 3, Kluwer Academic Publishers: 309-312.

1995
Clark T.A., Zeyen R.J., Carver T.L.W., Smith A.G., Bushnell, W.R.,1995. Epidermal cell cytoplasmic
    events and response gene transcript accumulation during Erysiphe graminis attack in isogenic
    barley lines differing  at the Ml-o locus. Physiological and Molecular Plant Pathology 46,1-16

Limpert  E., Brändle U., Müller K. and Wolfe M., 1995: Epidemiologie und Populationsgenetik
    windverbreiteter  Krankheitserreger am Getreide: Ergebnisse großräumiger Untersuchungen
    und Konsequenzen für die Züchtung.
    Ber. 46. Arbeitstagung der AGS, Gumpenstein, 21.-23. November 1995,155-162

Lyngkjaer M.F., Jensen H.P. and Ostergard H.,1995: A Japanese powdery mildew isolate with
    exceptionally large infection efficiency on Mlo-resistant barley. Plant Pathology 45,786-790.

NEWTON A.C. and ANDRIVON D,m1995. Assumptions and implications of current gene-for-gene
    hypotheses. Plant pathology 44,607-618

Schally H., Zederbauer R. , Zwatz B.,1995. Virulenzanalyse am Beispiel Sommergerste-Mehltau in
    Österreich unter Nutzung der Kollektionssysteme Sporenfalle und Pflanzendeposition.
    Pflanzenschutzberichte 55(1),52-68

Zeyen R.J., Bushnell W.R., Carver T.L.W., Robbins M.P., Clark T.A., Boyles D.A., Vance C.P.,
    1995: Inhibiting phenylalanine ammonia lyase and cinnamylalcohol dehydrogenase
    suppresses Mla1 (HR) but not mlo5 (non-HR) barley powdery mildew resistances.
    Physiological and Molecular Plant Pathology 47,119-140.

1996
Atzema J.L., Finckh M.R., Wolfe M.S.,1996: Genetics of the response of Erysiphe graminis f.
    sp. hordei to ml-o resistance in barley.
    Proc. 9th Eur. Med. Cer. rust & Powd. Mild. Conf. :55-57.

Brown J.K.M., Jones E.R.L., The effect of the mlo mildew resistance gene on Rhynchosporium secalis
     of barley,   UK Cereal Pathogen Virulence Survey 1995, Annual Report, 1996, pp. 47-49.

Finckh M.R., Atzema J.L., Freialdhoven A., Peterhänsel C., Schulze-Lefert P. and Wolfe M.,1996.
    Occurrence and genetics of virulence in barley powdery mildew to genes involved in Mlo resistance.
    Proc. 9th Cereal Rust & Mildew Conf., Lunteren, Sept. 1996.

Freialdenhoven A ., Peterhansel C., Kurth J., Kreuztaler F. and Schulzelefert P.,1996:
    Identification of genes required for the function of non-race-specific mlo resistance to
    powdery mildew in barley. Plant Cell 8,5-14.

Lyngkjaer M.F., Ostergard H.,1996: Powdery mildew infection of barley with the mlo resistance   gene.
    In: Proc. 9th Cereal Rusts and Powdery Mildew Conference, 2.-6th Sept. Lunteren, NL.
    Cereal Rusts and Powdery Mildew Bull. 24,58-60.

Newton A.C., Young I.M.,1996: Temporary partial breakdown of Mlo powdery mildew
    resistance in barley. Euphytica 63,141-152.

 Newton A. C. Young I. M. ,1996. Temporary partial breakdown of Mlo-resistance in spring
    barley by the sudden relief of soil water stress. Plant Pathology 45,970-974

Rashal I., Tueryapina, R.,1996. The test of Latvian powdery mildew isolates for mlo aggressiveness
    In: Limpert E.,Finckh M.R., Wolfe M.S., 1996, Integrated control of cereal mildews and rusts: towards
    coordination of research across Europe. The European Commission, EUR 16884-COST 817, 43-45

1997
Baker B., Zambryski P., Staskawicz B., Dinesh-Kumar S P, 1997: Signaling in plant-
    microbe interactions . Science 276, 726-733

Baker S., Newton A.C. and Gurr S.J.,1997: Temporary breakdown of Mlo-resistance to mildew
    in spring barley. Proc. Conf. Approaches to improving disease resistance to meet future
    needs: Airborne pathogens of wheat and barley, 11.-13.Nov.1997 Prague, 102 (Poster
    abstract ).

Büschges R., Hollricher K., Panstruga R., Simons G., Wolter M., Frijters A., van Daalen R., van
    der Lee T., Diergarde P., Groenendijk J., Töpsch S., Vos P., Salamini F., Schulze-Lefert P.,
    1997: The barley Mlo gene: A novel control element of plant pathogen resistance.
    Cell 88,695-705.

Elliott, C. Devoto, A., Haizel, T., Hartmann, A., Piffanelli, P., and Schulze-Lefert, P.
    STRUCTURAL AND FUNCTIONAL ANALYSIS OF BARLEY MLO. EUCARPIA -
    SECTION CEREALS MEETING - Sept. 22 - 23, 1997, TULLN - Austria

Jarosch Birgit, Kogel Karl-Heinz and Schaffrath Ulrich,1997: NEW LIGHT ON THE BARLEY
    MLO LOCUS: Although highly effective for resistance against powdery mildew,  mlo conditions
    enhanced susceptibility to the Rice Blast Fungus Magnaporthe grisea.
    EUCARPIA -  SECTION CEREALS MEETING - Sept. 22 - 23, 1997, TULLN - Austria

Jensen B., Munk L., Nitrogen-induced changes in colony density and spore production of Erysiphe
    graminis f. sp. hordei on seedlings of six spring barley cultivars, Plant Pathol. 46 (1997) 191-202.

Lyngkjaer M.F., Carver T.L.W. & Zeyen R.J., 1997: Suppression of resistance to Erysiphe
    graminis f.sp. hordei conferred by the mlo5 barley powdery mildew resistance gene.
    Physiological and Molecular Plant Pathology 50,17-36.

MANNINEN ,O. M., T. TURPEINEN & E. NISSILÄ ,1997:Identification of RAPD markers
    closely linked to the mlo-locus in barley - EUCARPIA - SECTION CEREALS
    MEETING - Sept. 22 - 23, 1997, TULLN - Austria

Peterhansel C., Freialdhoven A., Kurth J., Kolsch R., SchulzeLefert P.,1997. Interaction analyses of genes
     required for resistance responses to powdery mildew in barley reveal distinct pathwazs leading
     to leaf  cell death. Plant Cell 9,1397-1409

Simons G., van der Lee T., Diergaarde P., van Daelen R., Groenendijk J., Frijters A., Büschges R., Hollricher K.,     Töpsch S.,Schulze-Lefert P., Salamini F., Zabeau F., and Vos P. (1997) AFLP-based fine mapping of the         Mlo gene to a 30 kb DNA segment of the barley genome. Genomics 44:61-70.

Simons, G. et al. (1997) AFLP-based fine mapping of the Mlo gene to a 30-kb DNA segment
      Genet. 253, 687­694

Yahyaoui A.H., Reinhold M. and Scharen A.L.
    Virulence spectrum in populations of the barley powdery mildew pathogen, Erysiphe graminis f. sp.         hordei  in Tunisia and Morocco in 1992. 1997. Plant Pathology 46,139-146

1998
Atzema J.L.,1998.  Durability of mlo resistance in barley against powdery mildew caused by Eysiphe
    graminis  f.sp. hordei.  Ph.D. thesis, ETH Zurich, pp. 1-93.
    ( dealing with the  expression, epidemiology and genetics of interactions between powdery mildew and the
    mlo-gene, including crosses between patially mlo-virulent genotypes of mildew)

Baker S.J., Newton A.C., Crabb D., Guy D.C., Jefferies R.A., Mackerron D.K.L., Thomas W.T.B.,
    Gurr, S.J., 1998: Temporary partial breakdown of mlo-resistance in spring barley by
    sudden relief of soil water-stress under field conditions: the effects of genetic
    background and mlo allele. Plant Pathology 47,401-410.

Carver T.L.W., Robbins M.P., Thomas B.J., Troth K., Raistrick N. and Zeyen R.J., 1998. Silicon deprivation
    enhances localized autofluorescent responses and phenylalanine ammonia-lyase activity in oat
    attacked by  by Blumeria graminis. Physiological and Molecular Plant Pathology 52,245-257.

Czembor, J.H.,Czembor, H.J.,1998: Powdery mildew resistance in cultivars of spring barley
    from polish register. Plant breeding and seed science, 42(2),87-99

Huckelhofven R. and Kogel K.H., 1998. Tissue specific superoxide generation at interaction sites
    in resistant and susceptible near-isogenic lines attacked by the powdery mildew fungus
    (Erysiphe graminis f.sp. hordei). Molecular Plant-Mircobe Interactions 11,292-300.

Lahaye T, Hartmann S, Töpsch S, Freialdenhoven A and Schulze-Lefert P. (1998) High-resolution
    genetic and physical mapping of the Rar1 locus in barley.
    Theoretical Applied Genetics 96, .

Lyngkjaer M.F., Ostergard H., 1998: Interaction between powdery mildew and barley with
    mlo5 mildew resistance. Plant Pathology 47,252-238.

Panstruga R, Buschges R, Piffanelli P, Schulze-Lefert P. 1998. A contiguous 60 kb genomic stretch from
    barley reveals molecular evidence for gene islands in a monocot genome. Nucleic Acids Research 26,
    1056-62.

Peterhänsel C, Freialdenhoven A, Kurth J, Kolsch R and Schulze-Lefert P (1997) Interaction
    analyses of genes required for resistance responses to powdery mildew in barley reveal
    distinct pathways leading to leaf cell death.
    The Plant Cell 9: 1397-1409.

Ropenack E., Parr A., Schulze-Lefert P. 1998. Structural analyses and dynamics of soluble
    and cell wall-bound phenolics in a broad spectrum resistance to the powdery mildew fungus
    in barley. Journal of Biological Chemistry 273,9013-9022.

Schwarzbach, E., 1998: The mlo based resistance of barley against mildew and the response of
    mildew populations to the use of varieties with the mlo gene.
    Czech Journal of Genetics and Plant Breeding 34,3-10 (reprints still available)

Thomas, WTB.; Baird. E.; Fuller, JD.; Lawrence, P.; Young, CR.; Russell, J.; Ramsay, L.; Waugh,
    R.; Powell, W. (1998) Identification of a QTL decreasing yield in barley linked to Mlo
    powdery mildew resistance. Molecular Breeding. 4(5):381-393.English.

1999
Devoto A., Piffanelli P., Nilsson I., Wallin E., Panstruga R., von Heijne G., Schulze-Lefert P., 1999. Topology,  subcellular localization, and sequence diversity of the Mlo family in plants,
    J. Biol. Chem. 274 (1999) 34993-35004. PDF-file

Hückelhoven R, Fodor J, Preis C, Kogel K-H, 1999.Hypersensitive cell death and papilla formation in barley     attacked by the powdery mildew fungus are associated with H₂O₂ but not with salicylic acid             
    accumulation. Plant Physiol. 119,1251-1260

Jarosch Birgit, Kogel K-H., and Schaffrath U, 1999. The Ambivalence of the Barley Mlo Locus:
    Mutations Conferring Resistance Against Powdery Mildew (Blumeria graminis f. sp. hordei)
    Enhance Susceptibility to the Rice Blast Fungus Magnaporthe grisea.
    Molecular Plant-Microbe Interactions 12, 508-514.

Lyngkjaer M.F., Carver T.L.W., 1999. Modification of mlo5 resistance to Blumeria graminis attack in
    barley as a consequence of  induced accessibility and inaccessibility,
    Physiol. Mol. Plant Pathol. 55, 163-174.

Piffanelli P., Devoto A., Schulze-Lefert P.,1999. Defence signalling pathways in cereals.
    Curr. Opin. Plant Biol. 2, 295-300

 Shirazu K., Nielsen K., Piffanelli P, Oliver R. and Schulze-Lefert P., 1999. Cell-autonomous
    complementation of mlo resistance using a biolistic transient expression system.
    The Plant Journal 17,293-299

 Slater S.E., Clarkson J.D.S., Mildew of barley, U.K. Cereal Pathogen Virulence Survey 1998, Annual
     Report, 1999, pp. 43-51.

2000
Baker SJ, Newton AC, Gurr SJ, 2000. Cellular characteristics of temporary partial breakdown of
    mlo-resistance in barley to powdery mildew. Physiological and Molecular Plant Pathology 56, 1-11.

Collins, N., A. Sadanandom, F. Zhou, K. Shirasu, C. Elliott, J. Kurth. A. Devoto, P. Piffanelli, T. Lahaye, A. Hartmann, and P. Schulze-Lefert. 2000. The genetic and molecular basis of disease resistance to the         
    powdery mildew fungus in barley. In S. Logue (ed.) Barley Genetics VIII, Proc. Eighth Int. Barley Genetics        Symp., Adelaide, South Australia, 2000, Volume I:61-67.

Dreiseitl, A. & Jorgensen J.H.,2000. Powdery mildew resistance in Czech and Slovak barley cultivars.
     Plant  Breeding 119 (2000), 203-209.

Hovmoeller, M. S.,  Caffier V.,  Jalli M.  et al.,2000. The European barley powdery mildew virulence
    survey and disease nursery 1993-1999.  Agronomie 20 (2000) 729-743

Lyngkjaer M.F., Newton A. C., Atzema J.L. and Baker S.J., 2000.The Barley mlo-gene: an important
    powdery mildew resistance source. Agronomie 20 (2000) 745-756

Schulze-Lefert P, Vogel J, 2000. Closing the ranks to attack by powdery mildew
    Trends in Plant Science 5: (8) 343-348

Schweizer P, Pokorny J, Schulze-Lefert P, Dudler R.,2000. Double-stranded RNA interferes with gene
    function at the single-cell level in cereals. Plant Journal. 24(6):895-903.

Stewart K.J. and Gurr Sarah J.,2000. Abiotic stress and the breakdown of mlo-resistance to barley
  powdery mildew. (Abstract #372 of poster at Plant Biology 2000, ASPP).

2001
Collins N.C. Lahaye T., Peterhänsel C., Freialdenhoven A., Corbitt Margaret, and Schulze-Lefert P. 2001.
  Sequence Haplotypes Revealed by Sequence-Tagged Site Fine Mapping of the Ror1 Gene in the
  Centromeric Region of Barley Chromosome 1H 1[w]
  Plant Physiology, Vol. 125, 1236–1247

Frye C.A., Tang D.Z. and Innes, R.W.,2001. Negative regulation of defence responses in plants by a
  conserved MAPKK kinase. Proc. Natl. Acad. Sci USA, 98: 373-378

 Hinze K., Thompson R.D., Ritter E., Salamini F., and Schulze-Lefert P. (1991). RFLP-mediated targeting
   of the  mlo resistance locus in barley (Hordeum vulgare).
   Proc. Natl. Acad. Sci. USA 88: 3691-3695.

Hückelhoven R., Trujillo M. and Kogel K.-H.,2001. Mutations in Ror1 and Ror2 genes cause
  modification  of hydrogen peroxide accumulation in mlo-barley under attack from the powdery
  mildew fungus.   Molecular Plant Pathology 1,287-292.

Huckelhoven R, Dechert C, Trujillo M, Kogel KH, 2001.Differential expression of putative cell death
 regulator genes in near-isogenic, resistant and susceptible barley lines during interaction with the
 powdery mildew fungus. PLANT MOLECULAR BIOLOGY  47: (6) 739-748

 Kumar J., Hückelhoven R,  Beckhove U.,  Nagarajan S., and Kogel K.-H, 2001:   A Compromised Mlo
   Pathway Affects the Response of Barley to the  Necrotrophic Fungus Bipolaris sorokiniana
   (Teleomorph: Cochliobolus sativus)  and Its Toxins.  Phytopathology 91:127-133

Mercer PC, Ruddock A, 2001. Mildew of barley in Northern Ireland. UK Cereal Pathogen Virulence
  Survey 2000.  Annual Report, pp 60-62.

Paris M., Lance R. and  Jones M G.K., 2001. Single nucleotide primer extensions to type SNPs in barley
 Proc. 10th Australian Barley Technical Symposium, 2001

 Paris, M., Potter, R., and Jones, M.,2001. Typing Barley Mlo Alleles by Single Nucleotide
   Polymorphim  Analysis using MALDI-ToF Mass Spectrometry, Plant and Animal Genome IX, P335.

Schwarzbach E., 2001: Heat induced susceptibility of mlo-barley to powdery mildew (Blumeria graminis  D.C.f.sp. hordei Marchal). Czech J. Genetics and Plant Breeding. 37(3):82-87

Slater S.E. &  Clarkson J.D.S., 2001: Reaction of spring barley cultivars carrying the mlo resistance to
  infection by powdery mildew isolates in the UK. Cereal Rusts and Powdery Mildews Bulletin 2001

Slater S.E. & Clarkson J.D.S., 2001. Mildew of barley. UK Cereal Pathogen Virulence Survey 2000 Annual
 Report, pp50-59
 

2002    

Czembor JH & Czembor HJ, 2002. Selections from barley landrace collected in Libya as new sources of effective resistance to powdery mildew (Blumeria graminis f. sp. hordei). Rostlinna Vyroba 48,217-223
    Comment: a spontaneous mlo-allele in a Libyan landrace. Identity with other mlo alleles  not known.

Elliott C,  Zhou F, Spielmeyer W, Panstruga R and  Schulze-Lefert P, 2002: Functional Conservation of Wheat and Rice Mlo Orthologs in Defense Modulation to the Powdery Mildew Fungus  Molecular Plant-Microbe Interactions 15:1069-1077

Kim MC, Panstruga R, Elliott C,  Müller J.,Devoto A.,Yoon H W, Park HC, Cho MJ and Schulze-Lefert P, 2002. Calmodulin interacts with MLO protein to regulate defence  against mildew in barley.
Nature  416: 447-451.

 Kim MC, Lee SH, Kim JK, et al. 2002:  Mlo, a modulator of plant defense and cell death, is a novel calmodulin-binding protein - Isolation and characterization of a rice Mlo homologue
 J BIOL CHEM 277 (22): 19304-19314

Király L, Kumar J, Hückelhoven R, Kogel K-H, 2002. mlo5, a resistance gene effective against a biotrophic pathogen (Blumeria graminis fsp. hordei) confers enhanced susceptibility of barley to the necrotrophic fungus Bipolaris sorokiniana (teleomorph: Cochliobolus sativus). Acta Biologica Szegediensis 46,135-136, 2002

Panstruga R and Schulze-Lefert P, 2002. Live and let live: insight into powdery mildew disease and resistance. Molecular Plant Pathology 3,495-502.

Paris M, Jones MGK, 2002. Microsatellite genotyping by primer extension and MALDI-ToF mass spectrometry.PLANT MOLECULAR BIOLOGY REPORTER 20 (3): 259-263

Piffanelli P., Zhou F., Casais C., Orme J., Schaffrath U., Collins N., Panstruga R., and Schulze-Lefert, P. 2002: The barley MLO modulator of defence and cell death is responsive to biotic and abiotic stress stimuli.  Plant Physiology 129: 1076-1085.

Schultheiss H, Dechert C, Kogel KH, et al., 2002: A small GTP-binding host protein is required for entry of powdery mildew fungus into epidermal cells of barley.  PLANT PHYSIOL 128 (4): 1447-1454

Schwarzbach E., Slater Susan E. and  Clarkson J.D.S., 2002: Occurrence of partially mlo-virulent isolates of  barley powdery mildew in agricultural environments in Europe.   Cereal Rusts and Powdery Mildews  Bulletin 2002, vol. 30,  www.crpmb.org/2001/0208schwarzbach/

Schwarzbach E.,2002.The effects of silver on microbial contamination of agar medium and on
  interactions between mildew and barley leaf segments with and without the mlo gene.
  Czech J. Genetics and Plant Breeding. 38, 82-86

Slater, S E & Clarkson, J D S, 2002:. Mildew of Barley. UK Cereal Pathogen Virulence Survey 2001
 Annual Report, pp 52-60

Stein M, Somerville SC, 2002: MLO, a novel modulator of plant defenses and cell death, binds calmodulin.    TRENDS PLANT SCI7 (9): 379-380

Zeyen RJ, Carver TLW, Lyngkjaer MF. 2002. The formation and role of papillae. In. Belanger RR, Bushnell
WR, Dik AJ, Carver TLW, eds. The Powdery Mildews: A Comprehensive Treatise. St. Paul: APS Press, 107-125.
The following link is to the book at the publisher: http://www.shopapspress.org/powmilcomtre.html
 

2003

Collins N. C., Thordal-Christensen H., Lipka V, Bau S., Kombrink E., Qiu Jin-Long, Hückelhoven R,
Stein Mo nica , Freialdenhoven A., . Somerville Shauna C & Schulze-Lefert P., 2003. SNARE-protein-mediated disease resistance at the plant cell wall. NATURE  425, 973-977

Czembor JH, Frese L, 2003. Powdery mildew resistance in selections from barley landraces collected from Turkey. BODENKULTUR 54 (1): 35-40

Devoto A, Hartmann H A, Piffanelli P, Elliott C, Simmons C, Taramino G, Goh C, Cohen F E, Emerson B C, Schulze-Lefert P, and Panstruga R, 2003: Molecular phylogeny and evolution of the plant-specific seven transmembrane MLO family.   J. Mol. Evol.  56,77-88

Dreiseitl A, 2003. Adaptation of Blumeria graminis f.sp hordei to barley resistance genes in the Czech Republic in 1971-2000. PLANT SOIL AND ENVIRONMENT 49 (6): 241-248

Hückelhoven R, Dechert C, and Kogel K-H, 2003. OVEREXPRESSION OF BARLEY BAX INHIBITOR 1 INDUCES BREAKDOWN OF LO-MEDIATED PENETRATION RESISTANCE TO BLUMERIA GRAMINIS . Proc. Natl. Acad. Sci. USA, 100 (9), 5555-5560 l

Jarosch B., Jansen M, and Schaffrath U.,2003. Acquired Resistance Functions in mlo Barley Which Is Hypersusceptible to  Magnaporthe grisea. Phytopathology 16:107-114.

Li AL, Kong XY, Zhou RH, Ma ZY, Jia JZ, 2003. Isolation and characterization of Mlo and NBS-LRR-like gene sequences in wheat. ACTA BOTANICA SINICA 45 (4): 472-478

Molina-Cano JL, Simiand JP, Sopena A, Perez-Vendrell AM, Dorsch S, Rubiales D, Swanston JS, Jahoor A, 2003. Mildew-resistant mutants induced in North American two- and six-rowed malting barley cultivars. THEORETICAL AND APPLIED GENETICS 107 (7): 1278-1287

Olesen KL, Carver TLW, Lyngkjaer MF, 2003. Fungal suppression of resistance against inappropriate Blumeria graminis formae speciales in barley, oat and wheat. PHYSIOLOGICAL AND MOLECULAR PLANT PATHOLOGY 62 (1): 37-50

Panstruga R. and Schulze-Lefert P., Corruption of host seven-transmembrane proteins by pathogenic microbes: a common theme in animals and plants?  • REVIEW ARTICLE
Microbes and Infection,  5, 429-437

Paris M, Potter RH, Lance RCM, Li CD, Jones MGK, 2003. Typing Mlo alleles for powdery mildew resistance in barley by single nucleotide polymorphism analysis using MALDI-ToF mass spectrometry. AUSTRALIAN JOURNAL OF AGRICULTURAL RESEARCH 54 (11-12): 1343-1349 2003

Schultheiss H, Dechert C, Kogel KH, Huckelhoven R, 2003. Functional analysis of barley RAC/ROP G-protein family members in susceptibility to the powdery mildew fungus. PLANT JOURNAL 36 (5): 589-601, 2003

Schultheiss H, Dechert C, Kiraly L, Fodor J, Michel K, Kogel KH, Huckelhoven R, 2003. Functional assessment of the pathogenesis-related protein PR-1b in barley. PLANT SCIENCE 165 (6): 1275-1280.

Schulze-Lefert P., and Panstruga R. 2003. Estabishment of biotrophy by parasitic fungi and
reprogramming of host cells for disease resistance. Annual Review of Phytopathology 41: 641-667
(PubMed Abstract)

Zeyen RJ, Kruger WM, Lyngkjaer MF, Carver TLW, 2003. Differential effects of D-mannose and 2-deoxym-D-glucose on attempted powdery mildew fungal infection of inappropriate and appropriate Gramineae. PHYSIOLOGICAL AND MOLECULAR PLANT PATHOLOGY 61 (6): 315-323.

2004
Eichmann, R, Schultheiss, H, Kogel, KH, Huckelhoven, R, 2004.
The barley apoptosis suppressor homologue bax inhibitor-1 compromises nonhost penetration resistance of barley to the inappropriate pathogen Blumeria graminis f. sp tritici
MOLECULAR PLANT-MICROBE INTERACTIONS 17, 484-490

Elliott Candace, Müller Judith, Miklis M., Bhat R.A..  Schulze-LefertP. & Panstruga R., 2004.
Conserved extracellular cysteines and cytoplasmic loop-loop interplay are required
for functionality of the heptahelical MLO protein. Biochemical Journal 385: 243-254.

Hein, I, Campbell, EI, Woodhead, M, Hedley, PE, Young, V, Morris, WL, Ramsay, L, Stockhaus, J, Lyon, GD, Newton, AC, Birch, PRJ, 2004. Characterisation of early transcriptional changes involving multiple signalling pathways in the Mla13 barley interaction with powdery mildew (Blumeria graminis f. sp hordei)
PLANTA 218, 803-813

 Jansen, C. / Korell, M. / Eckey, C. / Biedenkopf, D. / Kogel, K.-H., Identification and transcriptional analysis of powdery mildew-induced barley genes. Plant Science 168,  373-380 

Opalski Krystina S., Schultheiss H., Kogel K.-H. and Hückelhoven R, 2005.The receptor-like MLO protein and the RAC/ROP family G-protein RACB modulate actin reorganization in barley attacked by the biotrophic powdery mildew fungus Blumeria graminis f.sp. hordei. The Plant Journal  41, 291–303

Panstruga, R., 2004.  A golden shot: how ballistic single cell transformation boosts the molecular
analysis of  cereal-mildew interactions.
MOLECULAR PLANT PATHOLOGY 5( 2 ): 141-148.

Piffanelli, P, Ramsay, L, Waugh, R, Benabdelmouna, A, D'Hont, A, Hinze K, Jorgensen, JH, Schulze-Lefert, P, Panstruga, R, 2004. The structure and possible origin of the barley mlo-11 mildew resistance allele.
11th International Cereal Rust and Powdery Mildew Conference, Norwich, 22-27 August 2004.

Piffanelli, P, Ramsay, L, Waugh, R, Benabdelmouna, A, D'Hont, A, Hollricher, K, Jorgensen, JH, Schulze-Lefert, P, Panstruga, R, 2004. A barley cultivation-associated polymorphism conveys resistance to powdery mildew. NATURE 430, 887-891

Schulze-Lefert P, Knocking on the heaven’s wall: pathogenesis of and resistance to biotrophic fungi at the cell wall  •Current Opinion in Plant Biology, 7, 377-383
.  

2005

Freialdenhoven, A., J. Orme, T. Lahaye and P. Schulze-Lefert, 2005. Barley Rom1 reveals a potential link between race-specific and nonhost resistance responses to powdery mildew fungi. Molecular Plant-Microbe Interactions 18, (4), 291-299 (2005)

Grell M.N.,  Holm Kirsten B. and Giese Henriette, 2005. Two novel Blumeria graminis f. sp. hordei genes are induced in planta and up-regulated in mlo virulent isolates., 2005. Physiological and Molecular Plant Pathology 66, 79-89

Hückelhoven R, , 2005.Powdery mildew susceptibility and biotrophic infection strategies. (Minireview)

Jarosch B, Collins NC, Zellerhoff N, Schaffrath U., 2005. RAR1, ROR1, and the actin cytoskeleton contribute to basal resistance to Magnaporthe grisea in barley. Mol Plant Microbe Interact. 18(5):397-404.

Lipka V. and Panstruga R, 2005. Dynamic cellular responses in plant–microbe interactions. Cell biology 245, 2005, 9-17

Muller J, Piffanelli P, Devoto A, Miklis M, Elliott C, Ortmann B, Schulze-Lefert P, Panstruga R., 2005. Conserved ERAD-like quality control of a plant polytopic membrane protein. Plant Cell.17(1):149-163

Maor R and Shirasu K., 2005. The arms race continues: battle strategies between plants and fungal pathogens. 
Current Opinion in Microbiology  8, 399-404

Panstruga, R., 2005. Serpentine plant MLO proteins as entry portals for powdery mildew fungi. Biochemical Society Transactions 33, 2, 389-392 

Panstruga, R., J. L. Molina-Cano, A. Reinstadler and J. Mueller, 2005. Molecular characterization of mlo mutants in North American two- and six-rowed malting barley cultivars. MOLECULAR PLANT PATHOLOGY 6, 3, 315-320 (2005)

Peterhansel, C., Lahaye, T., 2005. Be fruitful and multiply: gene amplification inducing pathogen resistance. Trends in Plant Science 10,  257-260

 Important change since 1.1.2006:

    Due to the growing number of  new papers in the scope of the page, it is increasingly difficult  to update properly and fast the web page. Since almost all scientists at universities and research institutes have now access to powerful scientific search engines, it is faster to look for new papers using advanced information technology. Therefore only papers discussed on the Mlo page or suggested by the authors themselves will be in future added to the list.  The most comprehensive and most efficient search engine for scientific papers is the interdisciplinary  Web of Science of the Institute of Sientific Information  (Philadelphia), which is available at most university libraries. A search can be specified by terms, authors, cited authors, topics etc and logical connections. In Europe it is available through http://wos.mimas.ac.uk/. 
    Single internet users may use PASCAL, http://www.cas.org/ONLINE/DBSS/pascalss.html . It is, however, expensive, about 4 Eur per hit.  There are also free scientific search engines on the web. Powerful and highly effective  is SCIRUS, http://www.scirus.com/srsapp/, from which you also may download  a free scientific search toolbar for the MSIE browser, which I personally recommend as the best solution when working from at home. Another engine is SCIENCE DIRECT, http://www.sciencedirect.com/, free for guest users.
    To compare the search engines, I used the search terms  barley AND mlo
I obtained  169 journal articles with SCIRUS, 68 with PASCAL, and 13  with ScienceDirect.

For your convenience, you may use SCIRUS directly from the Mlo web page:

Search only in Agricultural and Biological Sciences

Copyright statement:

The content of this page is intelectual property of  Erik Schwarzbach. It may be used, copied and spread freely for scientific and other non-commercial purposes. If used for publications, it shall be properly cited. Commercial use requires consent of the author. No other rights are reserved.
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