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Septoria and Stagonospora Diseases of Cereals - CIMMYT ...

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their resistance in either a recessive<br />

or dominant fashion (Eyal, 1999).<br />

However, in most cases, resistance<br />

appears dominant, with the F1<br />

from a cross between a resistant<br />

<strong>and</strong> a susceptible parent expressing<br />

an intermediate level <strong>of</strong> resistance<br />

that is more similar to that <strong>of</strong> the<br />

resistant parent. A few genes may<br />

be enough to confer resistance that<br />

will hold up in farmers’ fields<br />

(Dubin <strong>and</strong> Rajaram, 1996). While<br />

heritabilities tend to be only <strong>of</strong><br />

moderate magnitude, progress in<br />

breeding for resistance is evident.<br />

Although the number <strong>of</strong> genes<br />

available may be quite high,<br />

accumulating a few key ones may<br />

be sufficient to achieve resistance.<br />

As has been shown for durable<br />

resistance to leaf rust (Singh et al.,<br />

1991), several <strong>of</strong> the alleged<br />

components <strong>of</strong> partial resistance to<br />

S. tritici may also be controlled by<br />

only one or a just a few genes<br />

(Jlibene <strong>and</strong> El Bouami, 1995). It<br />

would seem that those components<br />

that are genetically different could<br />

be combined into the same genetic<br />

background by crossing. Once<br />

available, molecular markers will<br />

be <strong>of</strong> tremendous use for<br />

accumulating resistance to such<br />

environmentally sensitive diseases.<br />

The expression <strong>of</strong> S. tritici<br />

resistance was studied in T. tauschii<br />

accessions, synthetic wheats<br />

derived from them, plus<br />

derivatives <strong>of</strong> synthetic wheats<br />

crossed to common wheats. The<br />

results obtained all pointed to<br />

inheritance based on one or a few<br />

dominant genes (Appels <strong>and</strong><br />

Lagudah, 1990; May <strong>and</strong><br />

Lagudah, 1992).<br />

Chromosome 5D <strong>of</strong> T. tauschii<br />

contributed a high level <strong>of</strong> resistance<br />

to S. nodorum in a synthetic cross<br />

with a T. dicoccum line <strong>and</strong> in<br />

derived substitution lines<br />

(Nicholson et al., 1993).<br />

Chromosomes 5D, 3D, <strong>and</strong> 7D<br />

contributed resistance in decreasing<br />

order <strong>of</strong> importance. Aegilops<br />

longissima <strong>and</strong> Ae. speltoides were<br />

shown to contribute S. nodorum<br />

resistance based on two to four<br />

partially dominant to overdominant<br />

genes (Ecker et al., 1990a<br />

<strong>and</strong> b). Triticum timopheevii<br />

transmitted one S. nodorum<br />

resistance gene located on<br />

chromosome 3A to its progeny from<br />

a cross with a susceptible durum<br />

parent (Ma <strong>and</strong> Hughes, 1995).<br />

In most published accounts <strong>of</strong><br />

quantitative analyses, additive gene<br />

effects or general combining ability<br />

(GCA) effects contributed more to<br />

resistance than dominance or special<br />

combining ability (SCA) effects (van<br />

Ginkel <strong>and</strong> Scharen, 1987; Bruno<br />

<strong>and</strong> Nelson, 1990; Danon <strong>and</strong> Eyal,<br />

1990; Wilkinson et al., 1990; Jonsson,<br />

1991; Jlibene et al., 1994). However,<br />

significant non-additive effects were<br />

<strong>of</strong>ten identified in the same studies.<br />

SCA effects tend to be an order <strong>of</strong><br />

magnitude weaker than GCA<br />

effects. Indirectly Nelson <strong>and</strong> Fang<br />

(1994) confirmed these conclusions<br />

when they observed an absence <strong>of</strong><br />

heterosis for components <strong>of</strong> S.<br />

nodorum resistance, indicating a<br />

general lack <strong>of</strong> over-dominance<br />

effects that enhance resistance.<br />

Jlibene et al. (1994) found a small<br />

cytoplasmic effect in a few <strong>of</strong><br />

their crosses.<br />

Breeding for Resistance to the <strong>Septoria</strong>/<strong>Stagonospora</strong> Blights <strong>of</strong> Wheat 119<br />

Triticales were found not to be<br />

any more resistant than most<br />

wheats, <strong>and</strong> a range <strong>of</strong> reactions to<br />

both S. tritici <strong>and</strong> S. nodorum was<br />

observed (Scharen et al., 1990).<br />

Tritordeum amphiploids (doubled<br />

derivatives from crosses between<br />

Hordeum chilense <strong>and</strong> Triticum spp.)<br />

expressed the S. tritici resistance <strong>of</strong><br />

the barley parent both as seedlings<br />

in greenhouse tests <strong>and</strong> adult<br />

plants in field trials (Rubiales et<br />

al., 1992).<br />

Tolerance to S. nodorum was<br />

shown to be a mechanism that is<br />

genetically different from (partial)<br />

resistance <strong>and</strong> involves several<br />

chromosomes (Rapilly et al., 1988).<br />

Distinct characters were identified<br />

within each <strong>of</strong> these two defense<br />

mechanisms.<br />

Although the introduction <strong>of</strong><br />

alien cytoplasm from wild wheat<br />

relatives reduced partial resistance<br />

somewhat, it did increase tolerance<br />

as measured by reduced yield<br />

losses (Keane <strong>and</strong> Jones, 1990).<br />

Agronomic Traits<br />

<strong>and</strong> Resistance<br />

After a period in the 1960s <strong>and</strong><br />

1970s <strong>of</strong> seemingly increased levels<br />

<strong>of</strong> disease susceptibility in the<br />

semidwarf wheats (Santiago,<br />

1970), modern semidwarf<br />

germplasm carrying high levels <strong>of</strong><br />

resistance was subsequently<br />

identified. Many are now widely<br />

grown as commercial varieties.<br />

As in previous studies,<br />

Camacho-Casas et al. (1995)<br />

showed that it is possible to breed<br />

normally maturing semidwarf<br />

wheats if proper disease conditions

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