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Principles of Plant Genetics and Breeding

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192 CHAPTER 11<br />

Table 1 Estimated gain per cycle for grain yield, <strong>and</strong> ear <strong>and</strong> plant traits <strong>of</strong> SP <strong>and</strong> SA synthetic populations<br />

averaged for 4 years <strong>of</strong> testing.<br />

In our experiments, selection was carried out for ear size. Each season, 1,000–2,000 haploid plants from an improved synthetic<br />

population were planted in the field. Of these, 200–300 haploids were pollinated by diploid representatives <strong>of</strong> the same synthetic<br />

population. At harvest time about 20–30 haploids with the largest ears were selected for the next cycle <strong>of</strong> selection. Three cycles<br />

<strong>of</strong> haploid sib recurrent selection were completed for two synthetic populations, SP <strong>and</strong> SA. Initial <strong>and</strong> improved synthetics were<br />

evaluated in the field for 4 years. The performance results <strong>of</strong> synthetic SA are presented in the Figure 2. The data indicate that<br />

selection for ear size, utilizing haploids, can result in a significant increase <strong>of</strong> grain yield.<br />

An important method that effectively estimates the efficiency <strong>of</strong> a recurrent selection program is the determination <strong>of</strong> gain per<br />

cycle. This parameter is used for comparing the efficiency <strong>of</strong> different recurrent selection schemes. Normally gain per cycle for<br />

grain yield in a recurrent selection scheme in maize approximates 2–5% <strong>and</strong> seldom exceeds 7% (Gardner 1977; Weyhrich et al.<br />

1998). The results obtained by haploid sib recurrent selection are presented in Table 1. For synthetic population SA, the gain per<br />

cycle was 12.0%, <strong>and</strong> for synthetic SP it was 13.1%. Gain per cycle was distinctly higher than that which is observed when utilizing<br />

conventional recurrent selection methods. The conclusion drawn from the experiment is that the utilization <strong>of</strong> haploid plants<br />

for the selection <strong>of</strong> favorable genotypes greatly increases the efficiency <strong>of</strong> recurrent selection.<br />

Overall, numerous experiments have indicated that haploid generation can be successfully applied to several species on a<br />

large scale. The methods <strong>and</strong> citations given above provide only a few examples <strong>of</strong> this useful <strong>and</strong> efficient method. The utilization<br />

<strong>of</strong> haploids <strong>and</strong> doubled haploids can simplify the identification <strong>of</strong> genotypes that can provide a significant improvement in<br />

a variety <strong>of</strong> agronomic traits. In addition, haploids <strong>and</strong> doubled haploids can accelerate the generation <strong>of</strong> homozygous lines <strong>and</strong><br />

pure cultivars. Therefore, haploid-inducement technologies have a bright future in plant breeding.<br />

References<br />

Gain per cycle (%)<br />

SP SA<br />

Traits 1998 1999 2000 2001 Average 1998 1999 2000 2001 Average<br />

Grain yield 11.0 16.4 1.7 17.8 13.1 16.7 21.0 10.3 8.4 12.0<br />

Ear length 9.2 6.6 4.4 7.5 7.8 4.5 9.2 4.4 5.8 6.2<br />

Seeds per row 2.6 6.7 1.1 9.6 6.2 7.5 11.4 4.8 4.8 6.3<br />

Rows number 7.3 6.2 3.0 5.2 2.9 1.4 4.7 3.0 4.4 6.1<br />

Ear diameter 5.7 4.8 0.6 4.1 4.1 3.5 3.7 2.2 2.6 2.0<br />

<strong>Plant</strong> height 9.3 9.6 10.5 7.7 10.0 12.1 3.7 7.3 4.8 4.8<br />

Ear height 10.0 7.6 10.0 11.8 10.9 16.6 8.4 13.8 11.1 6.7<br />

Leaf length 8.5 6.2 3.8 9.0 7.8 3.3 2.8 1.5 3.2 2.6<br />

Chase, S.S. 1969. Monoploids <strong>and</strong> monoploid-derivatives <strong>of</strong> maize (Zea mays L.). Bot. Rev. 35:117–167.<br />

Coe, E.H. 1959. A line <strong>of</strong> maize with high haploid frequency. Am. Nat. 93:381–382.<br />

Eder, J., <strong>and</strong> S. Chalyk. 2002. In vivo haploid induction in maize. Theor. Appl. Genet. 104:703–708.<br />

Gardner, C.O. 1977. Population improvement in maize. In: Maize breeding <strong>and</strong> genetics (J. Janick, ed.), pp. 207–228. Wiley,<br />

New York.<br />

Kasha, K.J., <strong>and</strong> K.N. Kao. 1970. High frequency haploid production in barley (Hordeum vulgare L.). Nature 225:874–876.<br />

Kasha, K.J., E. Simion, R. Oro, Q.A. Yao, T.C. Hu, <strong>and</strong> A.R. Carlson. 2001. An improved in vitro technique for isolated microspore<br />

culture <strong>of</strong> barley. Euphytica 120:379–385.<br />

Kermicle, J.L. 1969. Androgenesis conditioned by a mutation in maize. Science 116:1422–1424.<br />

Kindiger, B. 1993. Registration <strong>of</strong> 10 genetic stocks <strong>of</strong> maize for the transfer <strong>of</strong> cytoplasmic male sterility. Crop Sci. 34:321–322.<br />

Kindiger, B., <strong>and</strong> S. Hamann. 1994. Generation <strong>of</strong> haploids in maize: A modification <strong>of</strong> the indeterminate gametophyte (ig) system.<br />

Crop Sci. 33:342–344.<br />

Thomas, W.T.B., B.P. Forster, <strong>and</strong> B. Gertsson. 2003. Doubled haploids in breeding. In: Doubled haploid production in crop<br />

plants (Maluszynski, M., K. Kasha, B.P. Forster, <strong>and</strong> I. Szarejko, eds), pp. 337–350. Kluwer Academic, Dordrecht.<br />

Weyhrich, R.A., K.R. Lamkey, <strong>and</strong> A.R. Hallauer. 1998. Responses to seven methods <strong>of</strong> recurrent selection in the BS11 maize<br />

population. Crop Sci. 38:308–321.

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