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Principles of Plant Genetics and Breeding

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246 CHAPTER 14<br />

sequence, <strong>and</strong> RIP coding sequence. <strong>Plant</strong> B consists<br />

<strong>of</strong> a promoter that is active during germination <strong>and</strong><br />

a CRE coding sequence. When the seed from a cross<br />

<strong>of</strong> A × B is planted, the floxing reaction occurs.<br />

Because the LEA promoter is inactive after late<br />

embryogenesis, the expression <strong>of</strong> the RIP is restricted<br />

to the seeds <strong>of</strong> the resulting mature plants from the<br />

hybridization.<br />

3 The use <strong>of</strong> an inducible promoter. This strategy is<br />

similar to using the hybridization process. However,<br />

in this case, the germination-specific promoter is<br />

replaced by a promoter that is controlled directly by<br />

an exogenous substance.<br />

Current status <strong>of</strong> GURT<br />

Since the first GURT patent was awarded jointly to<br />

USDA <strong>and</strong> Delta <strong>and</strong> Pine L<strong>and</strong>, various entities,<br />

including universities <strong>and</strong> private corporations, have<br />

pursued the development <strong>of</strong> a variety <strong>of</strong> technologies<br />

for seed sterilization. These include Syngenta (with at<br />

least eight GURT patents), Dupont, Monsanto, BASF,<br />

Iowa State University, <strong>and</strong> Cornell University. The TPS<br />

technology has so far not been commercially exploited.<br />

However, it appears various companies are working<br />

towards this objective. The Convention on Biological<br />

Diversity continues to discuss the issue. Like every technology,<br />

there are those who see the promise <strong>of</strong> TPS <strong>and</strong><br />

those who describe it in the most unflattering terms.<br />

Some <strong>of</strong> the stated potential advantages are:<br />

1 TPS would be an incentive for further research <strong>and</strong><br />

development <strong>of</strong> value-added cultivars.<br />

2 It could possibly reduce the unintended gene flow<br />

from transgenic cultivars to conventional cultivars.<br />

3 It could reduce the incidence <strong>of</strong> volunteer weeds.<br />

Distracters counter that:<br />

1 The only reason for developing <strong>and</strong> deploying the<br />

technology is to maximize the pr<strong>of</strong>its <strong>of</strong> seed<br />

companies.<br />

2 Poor farmers cannot afford the seed; further, they<br />

cannot save seed to plant if they wanted to.<br />

3 The protection provided lasts longer than any other<br />

similar protection system already in place.<br />

Molecular plant breeding<br />

Molecular breeding may be defined as the use <strong>of</strong><br />

molecular markers, in conjunction with linkage maps<br />

<strong>and</strong> genomics, to select plants with desirable traits on<br />

the basis <strong>of</strong> genetic assays. The potential <strong>of</strong> indirect<br />

selection in plant breeding was recognized in the 1920s,<br />

but indirect selection using markers was first proposed<br />

in 1961 by Thoday. The lack <strong>of</strong> suitable markers slowed<br />

the adoption <strong>of</strong> this concept. Molecular breeding<br />

gained new momentum in the 1980s <strong>and</strong> has since<br />

made rapid progress, with the evolution <strong>of</strong> DNA marker<br />

technologies.<br />

Molecular markers are used for several purposes in<br />

plant breeding.<br />

1 Gaining a better underst<strong>and</strong>ing <strong>of</strong> breeding materials<br />

<strong>and</strong> breeding system. The success <strong>of</strong> a breeding<br />

program depends to a large extent on the materials<br />

used to initiate it. Molecular markers can be used to<br />

characterize germplasm, develop linkage maps, <strong>and</strong><br />

identify heterotic patterns. An underst<strong>and</strong>ing <strong>of</strong> the<br />

breeding material will allow breeders to select the<br />

appropriate parents to use in crosses. Usually, breeders<br />

select genetically divergent parents for crossing.<br />

Molecular characterization will help to select parents<br />

that are complementary at the genetic level. Molecular<br />

markers can be especially useful in identifying markers<br />

that co-segregate with QTLs (quantitative trait loci)<br />

to facilitate the breeding <strong>of</strong> polygenic traits.<br />

2 Rapid introgression <strong>of</strong> simply inherited traits.<br />

Introgression <strong>of</strong> genes into another genetic background<br />

involves several rounds <strong>of</strong> tedious backcrosses.<br />

When the source <strong>of</strong> desirable genes is a<br />

wild species, issues <strong>of</strong> linkage drag becomes more<br />

important because the dragged genes are <strong>of</strong>ten undesirable,<br />

requiring additional backcrosses to accomplish<br />

breeding objectives. Using markers <strong>and</strong> QTL<br />

analysis, the genome regions <strong>of</strong> the wild genotype<br />

containing the genes encoding the desirable trait<br />

can be identified more precisely, thereby reducing<br />

the fragment that needs to be introgressed, <strong>and</strong> consequently<br />

reducing linkage drag.<br />

3 Early generation testing. Unlike phenotypic markers<br />

that <strong>of</strong>ten manifest in the adult stage, molecular<br />

markers can be assayed at an early stage in the development<br />

<strong>of</strong> the plant. <strong>Breeding</strong> for compositional<br />

traits such as high lysine <strong>and</strong> high tryptophan genes<br />

in maize can be advanced with early detection <strong>and</strong><br />

selection <strong>of</strong> desirable segregants.<br />

4 Unconventional problem-solving. The use <strong>of</strong><br />

molecular markers can bring about novel ways <strong>of</strong><br />

solving traditional problems, or solving problems<br />

traditional breeding could not h<strong>and</strong>le. When linkage<br />

drag is recessive <strong>and</strong> tightly linked, numerous rounds<br />

<strong>of</strong> backcrosses may never detect <strong>and</strong> remove it.<br />

Disease resistance is <strong>of</strong>ten a recessive trait. When the

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