Abstract

‘Honeycrisp’ is a widely grown and acclaimed apple cultivar that is commonly used in breeding programs. It also has a well-documented tendency to develop the physiological disorder, zonal leaf chlorosis (ZLC). This disorder causes reduced photosynthetic capacity and is thought to be due to a problem with phloem loading, although the underlying genetics of the disorder have not previously been discerned. In order to understand the breeding implications of the disorder, six families with ‘Honeycrisp’ as a parent and one family with ‘Honeycrisp’ as both a maternal and paternal grandparent were evaluated for ZLC incidence over two years. One major quantitative trait locus (QTL) for ZLC incidence was identified on linkage group (LG) 9. A haplotype in ‘Honeycrisp’ that originated from grandparent ‘Duchess of Oldenburg’ was associated with increased ZLC incidence in offspring in both years and all families evaluated. The LG9 QTL was 5 to 10 cM from MdMYB1, which is a major gene regulating fruit skin anthocyanin production. ‘Honeycrisp’ is heterozygous for red fruit skin overcolor at MdMYB1. The ‘Honeycrisp’ haplotype at the LG9 QTL associated with increased ZLC is in linkage phase with the allele at MdMYB1 associated with red color. Selection for the red allele from ‘Honeycrisp’ at MdMYB1 will result in most offspring also inheriting the haplotype at the LG9 QTL associated with high ZLC. The occurrence of two copies of this haplotype was sub-lethal in seedlings of a family where both parents inherited both the red overcolor allele at MdMYB1 and the haplotype at the LG9 QTL associated with high ZLC. This is the first study to have identified a genetic component of ZLC with clear breeding implications.

Highlights

  • It is possible that the specific diplotype of ‘Honeycrisp’ at the LG9 quantitative trait locus (QTL) for zonal leaf chlorosis (ZLC) is responsible for its high level of ZLC

  • Severe segregation distortion was observed at the LG9 QTL for ZLC in the ‘Honeycrisp’ x ‘Pitmaston Pineapple’ family, with no individuals having the LG9-H1 + LG9-H3 diplotype and very few individuals with the diplotype LG9-H2 + LG9-H6 present for evaluation in the field

  • The consistent location of the LG9 QTL for ZLC and the consistent effect of the LG9-H1 haplotype from ‘Honeycrisp’ in its offspring indicate this genomic region is a good target for further inquiries into the genetics and physiology of ZLC

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Summary

Methods

Plant materialTwo populations were evaluated in this study (S1 and S2 Tables). The first population consisted of 409 genotypes from five families that share ‘Honeycrisp’ as a common parent. The other parents in this population were ‘Jonafree’ [19], ‘Monark’ ( known as the University of Arkansas selection AA-44) [20], ‘Pitmaston Pineapple’ (USDA PI 279323) and the UMN apple selections MN1702 (‘Fireside’ x ‘Frostbite’) and MN1764 (parentage unknown). These families have been previously described in McKay et al [21] and were used to create the integrated genetic map described in Howard et al, 2017 [22] that is used in this study. The trees from these families were budded onto ‘Budagovsky 9’ rootstock in 2008 (buds that failed or were damaged were rebudded, primarily in 2009 and 2010)

Results
Discussion
Conclusion
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