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. Streptomyces and . Suzuki, Nevertheless, Phs and Bms structurally differ from one to another according the presence of a dihydrothiazole ring for Phs instead of a thiazole for Bms, 1979.

. Griggr, Members of the Ph family differ according the structure of a basic group which is different for each Ph molecule, influencing their spectrum of action, These differences in structure will impact on the strength of the link between the antibiotic and the DNA, thus on the degree of resistance, 1984.

. Reiter, They express their cytotoxic action by inducing single-or double-stranded DNA scissions, All these related antibiotics have a strong activity since they are able to kill both prokaryotic and eukaryotic cells, even at low concentrations (Umezawa, 1967.

. Franklin, . Snow, and . Bodet, , 1981.

B. Ascomycota and . Punt, Botrytis cinerea: MIC 15 µg.mL -1, Colletotrichum gloeosporioides: MIC 10 µg.mL -1, p.1, 1989.

(. Pardo, Penicillium chrysogenum: MIC 10 µg.mL -1, 1988.

. Tang, )) seem to be relatively sensitive to Ph and Bm even if the resistance level can differ between species and strains, Schizosaccharomyces pombe: MIC < 1 µg.mL -1, 1995.

A. Nidulans and A. Mic-;-alderton, is 10 µg.mL -1 while the A. oryzae one is ten-fold higher, 100 µg.mL -1 (Punt et al., 1992), Candida glabrata strains: MIC from 3.125 to 6.25 µg, p.1, 2006.

, Nevertheless, differences in the susceptibility to Zeo, Ph and Bm are observed for a same strain

. Suzuki, The same statement was observed with Clitopilus passeckerianus and Coprinopsis cinerea for which Ph 100 µg.mL -1 was able to completely inhibit the fungal growth while with 200 µg.mL -1 Zeo growth inhibition was not complete, Zeo seems to be less effective than Ph and Bm. For example, Aspergillus oryzae growth was inhibited by 200 µg.mL -1 Bm toward 100 µg.mL -1 Ph, 2005.

. Wang, HERIVAUX Anaïs | Les récepteurs histidine kinases : structure et distribution chez les eucaryotes et caractérisation fonctionnelle chez S. apiospermum rencontré dans la mucoviscidose 380 with acceptable concentrations, 2005.

. Laplaza, On the other hand, Ph seems to have a higher activity than Bm as demonstrated in Saccharomyces cerevisiae (Ph MIC = 0.1 µg.mL -1 toward Bm MIC = 5 µg.mL -1 ), While in Aspergillus nidulans and Neurospora crassa, the opposite statement was observed, 1987.

, Different genes, all originated from bacteria, encoding resistance to Ph, Bm and Zeo are available

. Gatignol, According the gene, the resulting proteins can inhibit antibiotic activity through stoichiometric or enzymatic mechanisms. Three resistance genes seem to encode a protein which bind to the antibiotic and thus inhibit reversibly the action of this latter, 1984.

. Gatignol, All these genes are able to confer resistance to both Ph and Bm (Gatignol et al., 1990). Nevertheless, the Shble resistance gene was demonstrated to express a higher resistance to Ph than the Tn5 ble gene in plants, ii) the Shble gene from Streptoalloteichus hindustanus, a tallysomycin-producing bacteria, 1987.

, All the Ph/Bm resistance genes have been demonstrated to be suitable for the genetic disruption of genes in yeast and fungal organisms. The most frequently used is the Shble gene from S. hindustanus in Ascomycota (Mattern and Punt, Drocourt, 1988.

;. Silar and . Santos, , 1995.

. Cardoza, The E. coli ble gene was also reported, at a lower extent, 1988.

. Cheevadhanarak, )) and yeast species (Gaillardin and Ribet, 1987.

. Kudla, The three other resistance genes available are less mentioned in the literature. The ble gene from S. aureus was reported in Candida famata transformation (Dmytruk et al., 2006) and the blm genes from S. verticillus was used in Pichia pastoris and Aspergillus oryzae transformation (Munshi and Lee, 1988.

. Annexes--revue,

, HERIVAUX Anaïs | Les récepteurs histidine kinases : structure et distribution chez les eucaryotes et caractérisation fonctionnelle chez S. apiospermum rencontré dans la mucoviscidose, p.381

. Van-engelenburg, The stability of the integration of the resistance gene in fungi is species dependant. In most cases the Ph/Zeo resistance gene was stably inherited, 1989.

. Cheevadhanarak, Rozman and Komel, 1992; Schuren and Wessels, 1991.

. Kummasook, Only few reports showed a low stability in the resistance gene integration, like in Aspergillus sojae, Phanerochaete chrysosporium, 1989.

. Kolar, According the studies, a low or high background is observed on plates containing the transformed cells, particularly in Basidiomycota transformation (Gaillardin and Ribet, 1987.

. Kim, For example, in the case of Trametes versicolor, the observed background was associated with the presence of osmotic stabilizers and the acidification of the medium by the fungus, 1991.

. Wang, To overcome this problem, research teams constructed codon-modified Ph/Zeo resistance genes consisting in modifying the five CTG codons of the Shble gene by leucine codons as yeasts belonging to the CTG clade will traduce a serine instead of a leucine (CUG, Some subtleties are important to keep in mind prior the use of these different resistance genes in fungal transformation. Indeed, all these genes are not functional in yeast species belonging to the CTG clade, 2003.

. Kilaru, , 2009.

. Kolar, Other important points must be underlined when using Ph or Bm as selection markers. First, the selection of transformed cells is mainly done on minimal medium, 1988.

. Van-engelenburg, , 1989.

K. Rozman and . Santos, , 1992.

. Teunissen, even if some experiments used rich media mostly in yeast transformations, 1987.

, HERIVAUX Anaïs | Les récepteurs histidine kinases : structure et distribution chez les eucaryotes et caractérisation fonctionnelle chez S. apiospermum rencontré dans la mucoviscidose, p.382

A. , , 2002.

Y. , , 2009.

. Lin, varying according the species. That is to say that the optimum medium pH to use Ph and Bm in Candida glabrata, Then, it was shown that Ph activity in Saccharomyces cerevisiae was dependent on aerobic conditions and growth phase, 1970.

, As a conclusion, the phleomycin/bleomycin/zeocin family can be used in numerous fungal species

. Gatignol, One of the advantages of these selection markers is that they do not bring any additional phenotype than the one desired, vol.500, p.bp, 1985.

. Sugiyama, Johansson and Hahn-Hägerdal), even this background seems to be lower than for geneticin (Johansson and Hahn-Hägerdal). In addition, they can induce DNA double-strand breaks thus they must be used with caution because they are highly toxic, 1994.

A. Nidulans and A. Orbach, Gliocladium virens (Ossanna and Mischke, Trichoderma harzianum (Inglis et al., 1999) but also the benA gene from Aspergillus nidulans, 1978.

. Haptmann, Benomyl is usually solubilized in dimethyl sulfoxide (Wood, 1982.

S. Leger, , vol.or absolute ethanol, 1989.

S. ;. Picknett, . Ossanna, and ;. Mischke, Transformed cells should be selected on rich media containing Ben, Annexes -Revue, 1985.

, HERIVAUX Anaïs | Les récepteurs histidine kinases : structure et distribution chez les eucaryotes et caractérisation fonctionnelle chez S. apiospermum rencontré dans la mucoviscidose, p.388

. Churchill, it was reported that low frequencies of transformation are associated with the use of ben R or benA as a selectable marker, particularly in filamentous fungi, Bunkers, 1988.

. Fernandez-larrea, ;. Stahl, and . Goettel, Finally, low concentrations of Ben in the medium are sufficient to select, 1989.

. Upchurch, µg.mL -1 for Cercospora kikuchii, 1990.

. Razanamparany, Gliocladium virens (Ossanna and Mischke, 1990), Penicillium sp. (Picknett and Saunders, Trichoderma harzianum (Inglis et al., 1999), and from 5 to 13 µg.mL -1 for Acremonium chrysogenum, 1985.

. Duarte, , 2006.

. Orbach, thus easy to integrate in a construction. Moreover, benomyl is inexpensive in comparison to other commonly used selectable markers, Ben-resistance genes seems to be convenient selectable marker to transform fungi such as the ben R gene from Neurospora crassa, p.340, 1986.

. Carboxin-;-von-schmeling, this molecule has been widely employed as a systemic fungicide (Snel and Edgington, 1970; Erwin, 1970) to control the spread of phytopathogen fungal species (Powelson and Shaner, Clitopilus passeckerianus MIC = 5 µg.mL -1, 1966.

, HERIVAUX Anaïs | Les récepteurs histidine kinases : structure et distribution chez les eucaryotes et caractérisation fonctionnelle chez S. apiospermum rencontré dans la mucoviscidose, p.390

. Keon, Indeed, in the SdhC a change from a threonine residue to an isoleucine was observed and an asparagine change for glutamine in the SdhD. The first described mutation was different from the one identified in the SdhC from Coprinopsis cinerea, the third cysteine rich cluster subunit of the Ip subunit of the SDH, is able to confer resistance to Cbx in fungi, 1976.

. Shima, it was thus easy to develop theses resistance gene as selection agents in fungal transformation. Nevertheless, these genes can only be used for homologous recombination, Hebeloma cylindrosporum, 2000.

. Irie, Ustilago esculenta, Ustilago maydis, 1972.

. Ruiz-herrera, , 1999.

. Skinner, , 1998.

. Shima, Nevertheless, in some rare cases, heterologous transformation gave rise to carboxin-resistant transformants (e.g. the sdi R gene from Agaricus bisporus in Coprinopsis cinerea, Aspergillus oryzae in Penicillium simplicissium, 2000.

. Shima, the sdi1 R gene from Coprinopsis cinerea in Clitopilus passeckerianus, Aspergillus oryzae in Aspergillus parasticus, 2009.

. Honda, but for a long time it was considered that using these resistance genes impaired the pathogenicity of the transformed strain, thus that Cbx resistance genes cannot be used as selection markers, Cbx resistance genes were successfully used as a dominant selection marker in fungal transformation, 1999.

. Edgington, Concentrations of Cbx used to select transformed cells is generally lower in basidiomycetous (e.g. 60 µg.mL -1 in Clitopilus passeckerianus, pp.40-41, 1966.

. Coprinopsis and . Kilaru, 0.2 µg.mL -1 in Hebeloma cylindrosporum, 2009.

. Irie, 50 µg.mL -1 in Magnaporthe oryzae, µg.mL -1 in Lentinula edodes, 2003.

. Honda, 5 µg.mL -1 in Ustilago esculenta, µg.mL -1 in Pleurotus ostreatus, p.75, 1992.

. Shima, 40 µg.mL -1 in Zymoseptoria tritici, Selection of the transformed cells is usually done on rich media in basidiomycetes, 2009.

. Honda, Finally, in some experiments it was precised that transformation plates were incubated in the dark, 1998.

. Ando, As a conclusion, carboxin is interesting in the transformation of basidiomycetous fungal species as far as in most cases the transformation efficiency obtained is higher or the same as the one with traditional selection markers, the discrimination is better, few studies reported the presence of false-positive transformants. It is more complicate to rule on ascomycetes species transformation because few data are available. Then, this drug is less expensive than hygromycin or geneticin, 2009.

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. Hansen, Jenks and Reines also demonstrated that the insertion of a hydroxyl side-chain at position 523 confers MPA resistance. Candida species show different degrees of resistance toward MPA. Indeed, in liquid and solid complete medium, C. parapsilosis was inhibited with 200 µg.mL -1 and 150 µg.mL -1 in liquid minimal medium, Candida auris was inhibited in minimal solid media containing, p.50, 2001.

. Mpa-(defosse, C. albicans strains showed a more limited resistance to MPA as far as their growth was inhibited between 5 and 10 µg.mL -1 of drug in the medium, 1989.

. Köhler, MPA induces cytokinesis defects in C. albicans morphogenesis and the IMH3 gene, 1997.

. Mpa-(gallagher, What is interesting here concerning the high resistance to MPA is that M. guilliermondii is not a MPA-producing species contrary to P. brevicompactum. Further investigations demonstrated that this yeast possess an additional IMH3.2 gene and the presence of a serine at position 251 is responsible for MPA resistance, 1987.

. Köhler, 2007) selection marker. In these experiments, C. albicans, C. dubliniensis strains were selected with low concentrations of MPA (5, 10 µg.mL -1 ) wile C. parapsilosis necessitated 150 to 200 µg.mL -1 MPA. The IMH3.2 gene from M. guilliermondii was used in heterologous transformation of a huge number of yeasts belonging to the CTG clade such as C. auris and Lodderomyces elongisporus with 50 µg.mL-1 MPA, Transformation experiments using the IMPDH resistance genes were only reported in yeasts. The IMH3 resistance gene from C. albicans was both used as a homologous, 1997.

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