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Principles of Plant Genetics and Breeding

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226 CHAPTER 13<br />

Bread wheat<br />

(Triticum aestivum)<br />

2n = 42<br />

Genome = AABBDD<br />

Intercross<br />

<strong>and</strong> select<br />

AABBDDRR<br />

2n = 56<br />

Primary octoploid<br />

Method 1<br />

F 1<br />

ABDR<br />

n = 28<br />

Sterile haploid<br />

Figure 13.6 Steps in the development <strong>of</strong> triticale.<br />

<strong>Breeding</strong> alloploids<br />

Alloploids may be induced by crossing two species with<br />

different genomes, followed by chromosome doubling <strong>of</strong><br />

the hybrid. Compared to autoploids, inducing alloploids<br />

is not commonly done by plant breeders. If successful,<br />

the newly induced amphiploid instantly becomes a new<br />

species (unable to cross to either parent). It also becomes<br />

reproductively isolated from its parents. Success <strong>of</strong><br />

induced alloploids is enhanced by the proper choice<br />

<strong>of</strong> parents. In particular, using parents with low ploidy<br />

levels increases the chance <strong>of</strong> high fertility <strong>and</strong> seed set<br />

in the amphiploid. Commercially successful induced<br />

alloploids are few. The most noted success with induced<br />

alloploidy is the commercially grown amphiploid, triticale<br />

(× Triticosecale), derived from a cross between wheat<br />

(Triticum) <strong>and</strong> rye (Secale) (Figure 13.6). The objective<br />

<strong>of</strong> developing triticale is to obtain a product that com-<br />

×<br />

Rye<br />

(Secale cereale)<br />

2n = 14<br />

Genome = RR<br />

×<br />

AABBDDRR<br />

2n = 56<br />

Fertile<br />

Embryo<br />

culture<br />

Double<br />

×<br />

AABB (RR/DD)<br />

2n = 42<br />

Secondary hexaploid<br />

Method 2<br />

F 1<br />

ABR<br />

n = 21<br />

Sterile haploid<br />

AABBRR<br />

2n = 42<br />

Fertile<br />

Durum wheat<br />

(Triticum turgidum)<br />

2n = 28<br />

Genome = AABB<br />

Intercross<br />

<strong>and</strong> select<br />

AABBRR<br />

2n = 42<br />

Primary hexaploid<br />

bines the qualities <strong>of</strong> wheat with the hardiness <strong>of</strong> rye. In<br />

lieu <strong>of</strong> doubling the F 1 to produce the desired synthetic<br />

product, a wheat × rye cross may be undertaken. The F 1<br />

plant possesses 28 chromosomes <strong>and</strong> exhibits intermediate<br />

traits that favor rye (hairy neck, spike length). All<br />

F 1 s are sterile because <strong>of</strong> the formation <strong>of</strong> univalents<br />

<strong>and</strong> irregular gametogenesis. F 1 s are backcrossed to<br />

wheat to produce progenies containing 42 chromosomes<br />

(seven from rye <strong>and</strong> the rest from wheat). The<br />

wheat chromosomes form bivalents at meiosis, while<br />

the rye chromosomes form univalents. The bivalent<br />

wheat chromosomes are irregularly arranged. Fertilization<br />

<strong>of</strong> an ovule with 21 + 7 chromosomes by pollen<br />

with the same genomic constitution will contain the full<br />

complement <strong>of</strong> chromosomes for wheat <strong>and</strong> rye (56<br />

chromosomes). This product is the synthetic alloploid<br />

called triticale. Hexaploid triticale (AABBRR, 2n = 6x<br />

= 42) is superior agronomically to octoploid triticale

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