Hybrid seed production: an applied usage of maternal inheritance

Plant breeders have recognized for many years that the progeny from a specific cross can out yield either of the two parents used in the cross. This is called heterosis, the phenomenon were the phenotypic value of the heterozygote is greater than either of the two parents. For plant breeding, the observation of heterosis for yield has lead to the development of inbred lines that exhibit a heterotic yield advantage. Corn was the first crop species in which heterosis was exploited. The original approach to develop hybrid corn seed required the manual detasseling of the female parent to prevent self -pollination. The field would be planted with the two lines and high school students would walk the field and detassel the female parent. As you can imagine this was a labor intensive proposition. Because it was realized that manual detasseling of corn plants would not be required if male sterile system could be developed, attemp ts were made to apply the two types of male sterility systems, genic and cyotplasmic male sterility. Genic male sterility is controlled by nuclear genes. The weakness of each of the genetic systems, though, was that a portion of the F2 were male fertile, a nd thus a portion of the seed that was developed was not hybrid. The ultimate solution to this problem was the use of cytoplasmic male sterility ( cms). As the name suggests, this type of cytoplasmic male sterility is controlled by a cytoplasmic factor and is maternally inherited . (Current molecular research suggests, but has not conclusively proven, that the sterility is a mitochondrial encoded function.) Thus all the males that contain the appropriate cytoplasm would be sterile. But this is only part of the solution for hybrid seed production. The seed company sells hybrid seed to the farmer, and the farmer expects this seed to be fertile. If the hybrid seed sold to the farmer was sterile, the seed company would have to provide a pollinator source to be p lanted along with with the hybrid seed to obtain the seed. The need for this step though is alleviated by the use of restorer of fertility ( Rf) genes . These dominant nuclear genes can override the cytoplasmic male sterility factors. Thus plants that have t he cms cytoplasm contain a dominant Rf allele will be male fertile. Taking the above discussion into consideration, the following is a procedure to produce hybrid corn seed without manual detasseling. A line that contains that contains a male sterile cytoplasm and is recessive for the restorer of fertility alleles (rfrf) is the female parent in a cross with a male that has male sterile cytoplasm and is heterozygous for the restorer of fertility alleles ( Rfrf). The F1 progeny from this cross will exhibit th e heterotic effects for yield. Furthermore, all the plants will be cytoplasmically male sterile because they contain the cms cytoplasm. The plants will also segregate 1 Rfrf :1 rfrf. Those that are recessive for the restorer factor will still produce seed because the other heterozygous Rfrf plants will produce

ample pollen to pollinate those plant Thus, the farmer will realize the advantages of hybrid seed, and the seed company will not have to use resoruces for manual detasseling. The following diagram de monstrates the process.

Although this system is functional, some difficulties have arisen with it practical application. In corn breeding, the cytoplasm that was used initially to provide the male sterility was the Texas or T cytoplasm. Nearly all of th e hybrid corn grown until 1970 contained this cytoplasm. During that year a fungal disease (Southern corn leaf blight) appeared that preferentially attacked plants with the T cytoplasm. This is an example of genetic vulnerability. This term refers to the genetic condition were all of the individuals in a region have the same genotype that makes them all vulnerable to a single disease or pathogen that could destroy the whole population. Thus, all of the hybrid corn was at great risk the following year. Fortunately, USDA scientists recognized the upcoming problem, and seed stocks were developed that contained other cytoplasms. These seed stocks had to be manually detasseled, and its hybrid performance was not as good as the previous material, but it did provide a source of seed that was resistant to the disease and saved the United States from losing the entire hybrid corn harvest that year. Since that time though sources of the T cytoplasm have been developed that are resistant to the disease, and hybrid see d production utilizing cytoplasmic male sterility has resumed.
Copyright 1997. Phillip McClean

The use of cytoplasmic male -sterility in maize seed production --Has, V, Has, I, Grecu, C The use of cytoplasmic male sterility (cms) in maize hybrid seed production is of economic importance and is also advantageous for genetic purity of seeds. Three types of male sterile cytoplasms in maize are used as cms maternal parents to produce hybrids: cms-C, cms-S, cms-T. The concern that in a few years all maize might again be in C or S cytoplasms gave rise to ideas as to how to prevent this narrowing of the cytoplasmic gene base. Thus, a new technique of producing hybrids was proposed, using multiplasm, respectively a blend of several kinds of male sterile cytoplasms. The aim of this investigation was: 1) to detect the presence of dominant Rf genes in more than 600 inbred lines by crossing with different types of cms: C, ES, M, T; 2) to compare some registered "TURDA" hybrids developed with normal and cms and/or Rf parental forms, in different environmental conditions, for three agronomic traits. Restoration reactions of 600 inbreds lines on the cms: C, ES, M and T were scored using Josephson s scale (Josephson et al., 1978). The observations were performed at the Agricultural Research Station -Turda, between 1995-2001. Nine registered "Turda" hybrids carrying both fertile and sterile cytoplasms were grown in two years at five locations. When using cms in maize breeding programs it is as necess ary as it is difficult to identify the inbred lines by their composition of Rf genes. Identifying restorers of cms-C and cms-ES becomes much more complicated, due both to the involvement of at least two-three complementary Rf4, Rf5, Rf6 genes, and to certain modifying factors, probably quantitative ones which, in some specific environmental conditions, act in the absence of the Rf gene, influencing the reactions of lines by the "late break" phenomenon. The percentage of non -restorer genotypes was 40% both t o cms-C and to cms-ES (Table 1). Table 1. The distribution of inbred lines according to their reaction in crosses to four cms types.

% inbred lines cms types No. cms tester lines partially fully cmsC cmsES 5 3 198 94 34 35 4 5 55 51 7 9 No. studied lines Non restorers Restorers Different reactions

cmsM cmsT

2 7

121 223

20 74

33 1

20 16

27 9

Total-studied lines 636 121 inbred lines have been tested with cms -M, only 20% of them being identified as Rf3/Rf3. The inbred lines which partially restore fertility or have a variable reaction according to the environmental conditions represent 27% of the inbred lines tested with cms-M. Because cms-T is only used in areas less favorable to the disease caused by Helminthosporium maydis T-race, research on the use of this cms type is limited. Table 2 presents the synthetic results of the comparison between the cytoplasmic (N or cms) effects on certain agronomic traits of registered hybrids developed at the Agricultural Research Station, Turda, Romania. Trial conditions (years, locations) have emphasized a series of significant differences between the two cytoplasms as far as grain yield is concerned. These differences are greatly determined by nuclear -cytoplasmic interaction or by hybrid x local conditions interaction. Table 2. Cytoplasmic male sterility effect for some traits in 9 registered "TURDA" hybrids.

Hybrid

Cytoplasms

Grain yield q/ha

Dry matter of grain %

Erect Synthetic plants relative at index harvest % % 5 85.4 79.3 93 79.0 81.8 103 81.6 6 100 90 100 100 100

1 Turda-SU 182

2 N cmsC (%)cms/N

3 98.6 95.0 96 101.1 97.3 96 85.1

4 76.6 77.1 101 77.7 77.9 100 75.3

TurdaMold 188

N cmsC (%)cms/N

Turda

Super cmsC (%)cms/N Saturn N cmsC (%)cms/N Turda 215 N cmsT (%)cms/N Turda-SU 210 N cmsC (%)cms/N Turda Favorite N cmsC (%)cms/N Turda 198 N cmsES (%)cms/N Turda 160 N cmsC (%)cms/N Trial mean N cms (%)cms/N 86.7 102 78.2 92.1 118** 97.5 86.2 880 84.9 89.2 105 93.3 104.5 112* 102.1 101.2 99 90.0 87.8 97 92.3 93.4 101 76.0 101 73.2 75.1 102** 76.0 74.4 9800 76.0 77.0 101 75.8 74.7 99 76.8 76.5 100 77.9 77.1 99 76.1 76.1 100 78.2 96 78.0 83.2 107 73.5 70.0 95 73.6 75.8 103 80.7 74.3 92 76.1 75.5 99 78.7 81.2 100 78.5 77.7 99 98 100 129 100 82 100 110 100 102 100 98 100 97 100 100 -

*** Significant at 5% and 1%, respectively *Si%= {col.3x4x5(cms)/col.3x4x5(N)}. 100 The nine hybrids carrying cms did not differ generally from their counterparts with fertile cytoplasm (N) for yield and for two other traits.

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