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Principles of Plant Genetics and Breeding

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MUTAGENESIS IN PLANT BREEDING 211<br />

Khanizadeh, S., F. Laurens, Y. Lespinasse, Y. Groleau, O. Carisse, <strong>and</strong> J.R. DeEll. 2004. ‘Reinette Russet®’ <strong>and</strong> ‘Galarina®’: two<br />

new winter hardy disease resistant apple cultivars released through joint collaboration <strong>of</strong> INRA (France) <strong>and</strong> AAFC (Canada).<br />

Acta Hort. 663:887–889.<br />

Knight, R.L., <strong>and</strong> F.H. Alston. 1968. Sources <strong>of</strong> field immunity to mildew (Podosphaera leucotricha). Can. J. Genet. Cytol.<br />

10:294–298.<br />

Labuschagné, I.F. 2004. Budbreak number as selection criterion for breeding apples adapted to mild winter climatic conditions: a<br />

review. Acta Hort. 663:775–777.<br />

Laurens, F. 1999. Review <strong>of</strong> the current apple breeding programmes in the world: objectives for scion cultivar improvement. Acta<br />

Hort. 484:163–170.<br />

Laurens, F., J.M. Audergon, J. Claverie, et al. 2000. Integration <strong>of</strong> architectural types in French programmes <strong>of</strong> ligneous fruit<br />

species genetic improvement. Fruits 55:141–152.<br />

Laurens, F., M. Chevalier, E. Dolega, et al. 2004. Local European cultivars as sources <strong>of</strong> durable scab resistance in apple. Acta<br />

Hort. 663:115–121.<br />

Laurens, F., Y. Lespinasse, <strong>and</strong> A. Fouillet. 2005. A new scab-resistant apple: ‘Ariane’. Hortscience 40(2):484–485.<br />

Laurens, F., <strong>and</strong> C. Pitiot, 2003. French apple breeding program: a new partnership between INRA <strong>and</strong> the nurserymen <strong>of</strong><br />

NOVADI. Acta Hort. 622:575–582.<br />

Nybom, H. 2004. ‘Frida’ <strong>and</strong> ‘Fredrik’, the first scab resistant apple cultivars developed in Sweden. Acta Hort. 663:871–873.<br />

Parisi, L., V. Fouillet, H.J. Schouten, et al. 2004. Variability <strong>of</strong> the pathogenicity <strong>of</strong> Venturia inaequalis in Europe. Acta Hort.<br />

663:107–113.<br />

Parisi, L., Y. Lespinasse, J. Guillaumès, <strong>and</strong> J. Krüger. 1993. A new race <strong>of</strong> Venturia inaequalis virulent to apples with resistance<br />

due to Vf gene. Phytopathology 83(5):533–537.<br />

Roberts, T., <strong>and</strong> I. Crute. 1993. Apple scab resistance from Malus floribunda 821 (V f ) is rendered ineffective by isolates <strong>of</strong> Venturia<br />

inaequalis from Malus floribunda. Abstracts, Third Workshop on Integrated Control <strong>of</strong> Pome Fruit Diseases. L<strong>of</strong>thus (Norway), 43.<br />

Semon, S.F.A., 2004. Developments in breeding <strong>and</strong> techniques within the community plant variety rights fruit sector. Acta Hort.<br />

663:707–711.<br />

Tartarini, S., F. Gennari, D. Pratesi, et al. 2004. Characterization <strong>and</strong> genetic mapping <strong>of</strong> a major scab resistance gene from the<br />

old Italian apple cultivar ‘Durello di Forli’. Acta Hort. 663:129–133.<br />

Way, R.D., H.S. Aldwinckle, R.C. Lamb, et al. 1990. Genetic resources <strong>of</strong> temperate fruit <strong>and</strong> nut. Acta Hort. 290:1–62.<br />

growing tip determines the sectorial patterns. Sectorial<br />

patterns are mostly unstable, eventually producing periclinal<br />

structures. Because <strong>of</strong> this behavior, the breeder<br />

needs to grow several clonal generations successively,<br />

<strong>and</strong> select for the desired mutant as well as vegetative<br />

stability. Vegetative stability is attained when uniformly<br />

periclinal lines <strong>of</strong> subclones have been isolated. In a<br />

vegetative system, the treated plant meristem is called<br />

the M 1 V 1 , the next vegetative plant being M 1 V 2 , etc.<br />

Mutations from tissue culture systems<br />

Useful mutations have arisen spontaneously or are<br />

deliberately induced in culture conditions. Such processes,<br />

somaclonal variation <strong>and</strong> somatic selection,<br />

were discussed in Chapter 11.<br />

Using induced mutants<br />

A plant may be selected for a specific mutant trait.<br />

However, it does not mean that this plant contains only<br />

one genetic change. It is recommended to backcross this<br />

mutant to its original parents to reselect a pure version<br />

<strong>of</strong> the mutant. Mutants may be directly used in cultivar<br />

development. Like the backcross method, an adapted<br />

line can be improved through mutation breeding to<br />

correct a shortcoming. In sexually reproducing species,<br />

the target mutant traits can be fixed <strong>and</strong> isolated in the<br />

M 2 or M 3 generation (compared to F 6 or F 7 generations<br />

in conventional breeding). The mutant lines may be<br />

used in genetic studies as markers, among other uses.<br />

Limitations <strong>of</strong> mutagenesis as a plant<br />

breeding technique<br />

There are a number <strong>of</strong> disadvantages <strong>of</strong> mutageneis<br />

when used in plant breeding.<br />

1 Associated side effects. Mutations induced by<br />

mutagens are very diverse in nature. Invariably, the<br />

mutagen kills some cells outright while surviving<br />

plants display a wide range <strong>of</strong> deformities. Even those<br />

plants with the desirable mutations always inherit<br />

some undesirable side effects (akin to linkage drag,<br />

or is it a “mutation drag”?). Therefore, it is <strong>of</strong>ten<br />

necessary to transfer a new mutant into a stable<br />

genetic background, free <strong>of</strong> some <strong>of</strong> the associated

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