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Principles of Plant Genetics and Breeding

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F. J. Betrán<br />

Texas A&M University, College Station, TX 77843, USA<br />

Figure 1 Heterosis in maize: A × B hybrid ears in the middle<br />

<strong>and</strong> corresponding parental inbreds A <strong>and</strong> B in both sides.<br />

Figure 2 Relative grain yields <strong>of</strong> an open-pollinated population<br />

<strong>and</strong> a single-cross hybrid developed from selected inbreds from<br />

the same population.<br />

BREEDING CORN 491<br />

Industry highlights<br />

Hybrid breeding in maize<br />

Introduction<br />

Maize is a cross-pollinated species that shows high<br />

heterosis (i.e., superior performance <strong>of</strong> crosses relative<br />

to their parents) for grain yield (Figure 1). This high<br />

expression <strong>of</strong> heterosis is exploited in maize hybrids<br />

<strong>and</strong> constitutes the foundation <strong>of</strong> the maize seed<br />

industry. Maize hybrids were first developed in the<br />

United States in the mid-1930s <strong>and</strong> by the early 1960s<br />

practically all the maize area in the US was planted to<br />

hybrids (Duvick & Cassman 1999). Improved productivity<br />

<strong>and</strong> selection gain with the use <strong>of</strong> hybrids has<br />

stimulated increased investment in hybrid development,<br />

resulting in impressive genetic progress (Figure<br />

2). Shull (1909) outlined the pure-line method in<br />

maize breeding suggesting the use <strong>of</strong> self-fertilization<br />

to develop homozygous lines that would be <strong>of</strong> use<br />

in hybrid production. This combination <strong>of</strong> inbreeding<br />

<strong>and</strong> hybridization constitutes the basis <strong>of</strong> maize<br />

improvement. The general process to develop maize<br />

hybrids starts with the creation <strong>of</strong> a source segregating<br />

breeding population that it is used to develop inbred<br />

lines through inbreeding <strong>and</strong> selection (Figure 3).<br />

Selected inbreds are then evaluated in hybrid combinations<br />

across locations to select superior hybrids<br />

<strong>and</strong> to estimate their combining abilities. The following<br />

is a brief description <strong>of</strong> the main components <strong>of</strong><br />

this process from the starting breeding population to<br />

the commercial hybrids.<br />

Source breeding populations<br />

Different type <strong>of</strong> segregating populations can be used<br />

as the source in line development: open-pollinated<br />

cultivars (OPC), synthetic cultivars, single crosses,<br />

backcrosses, double crosses, related line crosses, <strong>and</strong><br />

exotic germplasm. Overall, major emphasis goes to<br />

the use <strong>of</strong> breeding populations created by hybridization<br />

<strong>of</strong> complementary inbreds <strong>and</strong> the selection <strong>of</strong><br />

progenies possessing the desirable traits from both parents<br />

(Hallauer 1990). Selection within F 2 <strong>and</strong> backcross<br />

populations using pedigree breeding is the most<br />

important breeding method to develop maize inbreds.<br />

<strong>Breeding</strong> programs that emphasize pedigree selection<br />

within populations developed from elite inbred lines<br />

are therefore cyclical creating second-, third-, fourth-, etc. generation recycled improved inbreds (Figure 3). The incorporation<br />

<strong>and</strong> introgression <strong>of</strong> exotic germplasm brings new desirable alleles <strong>and</strong> genetic diversity to this recycling <strong>of</strong> elite lines (Goodman<br />

et al. 2000). A backcross or multiple backcross to the best parental inbred is used commonly to increase the probability <strong>of</strong> maintaining<br />

favorable combinations <strong>of</strong> alleles (Troyer 2001). Maize breeders use multiple trait, multistage, <strong>and</strong> multiple environment<br />

selection methods (Betrán et al. 2003). Multiple environment <strong>and</strong> multiyear inbred general combining ability values with

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