Did backcrossing contribute to the origin of hybrid edible bananas?

Background

Bananas and plantains (Musa spp.) provide a staple food for many millions of people living in the humid tropics. The cultivated varieties (cultivars) are seedless parthenocarpic clones of which the origin remains unclear. Many are believed to be diploid and polyploid hybrids involving the A genome diploid M. acuminata and the B genome M. balbisiana, with the hybrid genomes consisting of a simple combination of the parental ones. Thus the genomic constitution of the diploids has been classified as AB, and that of the triploids as AAB or ABB. However, the morphology of many accessions is biased towards either the A or B phenotype and does not conform to predictions based on these genomic formulae.

Scope

On the basis of published cytotypes (mitochondrial and chloroplast genomes), we speculate here that the hybrid banana genomes are unbalanced with respect to the parental ones, and/or that inter-genome translocation chromosomes are relatively common. We hypothesize that the evolution under domestication of cultivated banana hybrids is more likely to have passed through an intermediate hybrid, which was then involved in a variety of backcrossing events. We present experimental data supporting our hypothesis and we propose a set of experimental approaches to test it, thereby indicating other possibilities for explaining some of the unbalanced genome expressions. Progress in this area would not only throw more light on the origin of one of the most important crops, but provide data of general relevance for the evolution under domestication of many other important clonal crops. At the same time, a complex origin of the cultivated banana hybrids would imply a reconsideration of current breeding strategies.     

太古核不育小麦显性不育基因的染色体组测定

以太谷核不育小麦为母本与5个四倍体小麦种——硬粒小麦、波斯小麦、波兰小麦、埃及小麦、东方小麦为父本,进行杂交和回交。杂种F_1育性分离比例是:不育株与可育株为1:1,染色体构型为14Ⅱ+7Ⅰ。不育株的回交后代是随着回交次数的增加,分离出的不育株数逐次减少,可育株数逐次增多,极显著偏离1:1;BC_3不育株绝大多数染色体构型为14 Ⅱ+1 Ⅰ,带有一个单价体,BC_3可育株染色体构型为14Ⅱ。杂种F_1的可育株,其回交后代全是可育株,没有分离出1株不育株。以太谷核不育小麦为母本与3个六倍体小麦种—印度矮生小麦、瓦维洛夫小麦、密穗小麦为父本,进行杂交和回交,其后代育性分离比始终为1:1。测定结果表明:太谷核不育小麦的显性不育基因是在D组染色体的某个染色体上。因而这个基因不能直接导入不含有D组染色体的小麦种中去,如硬粒小麦。     

Breeding for ”low-gossypol seed and high-gossypol plants” in upland cotton. Analysis of tri-species hybrids and backcross progenies using AFLPs and mapped RFLPs

This work aims at breeding upland cotton [Gossypium hirsutum L., 2(AD)₁ genome] with a reduced level of gossypol in the seeds for optimal food and feed uses, and a high gossypol level in the remaining organs for resistance to pests. Two tri-species Gossypium hybrids, (G. thurberi–G. sturtianum–G. hirsutum and G. hirsutum–G. raimondii–G. sturtianum) including G. sturtianum (2C₁) as a donor, G. thurberi (2D₁) and G. raimondii (2D₅) as a bridge species, were created. Recurrent selection initiated with these tri-species hybrids produced backcross (BC) progenies expressing the ”low-gossypol seed and high-gossypol plant” trait at different levels. We used AFLP markers to assess the genetic similarity among the germplasm and RFLP probes to tag the introgression of specific chomosome segments from the parental species. Five pairs of AFLP primers generated 477 fragments, among which 417 (87.4%) were polymorphic. The genetic similarity between the upland cotton and the wild species ranged from 29.5 to 43.2%, while similarity reached 80% between upland cotton and BC3 plants. Introgression of species-specific AFLPs was evident from all the parental species and confirmed the hybrid origin of the analyzed progenies. Southern-blot analysis based on 49 RFLP probes allowed us to trace the introgression of parental DNA segments in the tri-species hybrids and in three generations of backcross. Introgression was evident from 11, 8 and 7 linkage groups of G. sturtianum, G. raimondii and G. thurberi respectively. The types of introgression revealed by RFLP probes are discussed, and breeding schemes to enhance recombination are proposed. The ability to trace DNA segments of known chromosomal locations from the donor G. sturtianum through segregating generations is a starting point to map the ”low-gossypol seed and high-gossypol plant” traits. Received: 5 January 1999 / Accepted: 17 June 1999     

Recurrent selection alternating with haploid steps — a rapid breeding procedure for combining agronomic traits in inbreeders

Summary Doubled haploid lines are increasingly used in practical breeding programs. Depending on the nature of the starting material as well as on the breeding aims, one or more haploid steps are recommended, and F₁ hybrids or selfed progenies in later generations serve as starting material for the haploidization. It is demonstrated that one haploid step followed by selection in the greenhouse and in the field during the first androgenetic (A₁) and two subsequent selfed generations (A₂; A₃) is the most efficient procedure, if characters from related varieties are to be combined. For breeding programs that include wild types or unrelated genotypes, recurrent selection alternating with several haploid steps is the most efficient. A prerequisite for successful application of this combination of repeated haploidization and selection steps alternating with backcrosses is an effective selection system. The different approaches are demonstrated on a winter barley breeding program, with the central aim of combining the qualitatively inherited character resistance to barley yellow mosaic virus with quantitatively inherited complexes, particularly yield.     

Expectations of means and genetic variances in backcross populations

Summary The genetic variance among F₂-derived lines of backcrosses (BC_gF₂-derived lines) depends on the backcross generation (g), the number of F₁ plants crossed and selfed in generations 1 through g, and the number of BC_gF₂-derived lines evaluated. Additive genetic variance decreases linearly with backcrossing when one BCF₁ plant per generation is crossed and selfed. The relationship is curvilinear if more than one BCF₁ plant is used; as the number of BCF₁ plants increases, additive genetic variance among BC₁F₂-derived lines approaches that among BC₀F₂-derived lines. The effect of population size on genetic variance is due both to fixation of alleles in previous generations and to sampling of genotypes in the population being evaluated. Dominance and repulsion linkage can cause small increases in genetic variance from BC₀ to BC₁.Joint contribution of USDA-ARS and Journal Paper No. J-11095 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa. Project No. 2471     

Sex linkage of CpGV resistance in a heterogeneous field strain of the codling moth Cydia pomonella (L.)

The occurrence of codling moth populations in European apple orchards that were not controlled by Cydia pomonella granulovirus (CpGV) is the first reported case of field resistance against a baculovirus control agent. A monogenic dominant sex-linked mode of inheritance was previously demonstrated in single-pair crosses between a homogeneous resistant (CpRR1) and a susceptible (CpS) laboratory strain of codling moth. However, resistant field populations (CpR) are more heterogeneous in their levels of resistance, and the possibility that they could harbor different resistance genes to CpRR1 had not been directly addressed. Here we report single pair crossing experiments using a resistant codling moth strain collected from an apple orchard in the southwest of Germany. Single-pair crosses within the field strain revealed a genetic basis to the heterogeneity of CpR concerning CpGV resistance. Hybrid crosses to a susceptible laboratory strain and backcrosses of the F1 generation to the resistant CpR strain confirmed that the homogeneous CpRR1 and the heterogeneous field strain CpR share the same mode of inheritance. Thus the variable levels of CpGV resistance in field populations is likely due to frequency differences of the same resistance-conferring gene, rather than different genes, which will facilitate future efforts to monitor and manage resistance.