, Volume 250, Issue 5, pp 1491–1504 | Cite as

Two types of mutations in the HEUKCHEEM gene functioning in cucumber spine color development can be used as signatures for cucumber domestication

  • Chunying Zhang
  • Khin Thanda Win
  • Young-Cheon Kim
  • Sanghyeob LeeEmail author
Original Article


Main conclusion

The HEUKCHEEM gene plays an important role in spine color formation. A white spine occurs due to two mutations in HEUKCHEEM and is closely related to the regional distribution of these mutants.


Mapping analysis revealed that the HEUKCHEEM gene is co-segregated with the B locus in the regulation of black spine color development in cucumber fruit. HEUKCHEEM induced the expression of the genes involved in the anthocyanin biosynthetic pathway, leading to the accumulation of anthocyanins in black spines. The transiently over-expressed HEUKCHEEM in cucumber and tobacco plants enhanced the expression of anthocyanin biosynthesis-related genes, leading to anthocyanin accumulation. However, two mutations—insertion of the 6994 bp mutator-like transposable element (MULE) sequence into the second intron and one single-nucleotide polymorphism (SNP) of C to T in the second exon of HEUKCHEEM—were identified in white spines, leading to no accumulation of anthocyanin biosynthesis-related gene transcripts and anthocyanins. Furthermore, association analysis using 104 cucumber accessions with different geographical origins revealed that the types of mutations in HEUKCHEEM are strongly linked to geographical origins. The MULE insertion is found extensively in cucumbers with white spines in East Asia and Australia. However, cucumbers with white spines in other areas could be significantly influenced by a single SNP mutation. Our results provide fundamental information on spine color development in cucumber fruits and spine color-based cucumber breeding programs.


Anthocyanin B locus Cucumber (Cucumis sativusHEUKCHEEM R2R3-MYB Spine color 



Anthocyanin synthase


(Derived) Cleaved amplified polymorphic sequences


Chalcone isomerase


Chalcone synthase


Dihydroflavonol 4-reductase




Insertions and deletions




Mutator-like transposable element


Single-nucleotide polymorphism


UDP-glucose flavonoid 3-O-glucosyltransferase


WD-repeat protein



This work was supported by grants from the Bio-industry Technology Development Program (Grant nos. 117043-3, 316087-4) of iPET (Korea Institute of Planning and Evaluation for Technology in Food, Agriculture and Forestry) and the Next-Generation BioGreen 21 Program (Plant Molecular Breeding Center) (Grant no. PJ01329601) of the Rural Development Administration, Republic of Korea.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

425_2019_3244_MOESM1_ESM.pdf (951 kb)
Supplementary material 1 (PDF 951 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Plant Genomics Laboratory, Department of Bio-resource Engineering, College of Life SciencesSejong UniversitySeoulRepublic of Korea
  2. 2.Plant Engineering Research InstituteSejong UniversitySeoulRepublic of Korea

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