Theoretical and Applied Genetics

, Volume 131, Issue 7, pp 1469–1480 | Cite as

Identification of a novel SPLIT-HULL (SPH) gene associated with hull splitting in rice (Oryza sativa L.)

  • Gileung Lee
  • Kang-Ie Lee
  • Yunjoo Lee
  • Backki Kim
  • Dongryung Lee
  • Jeonghwan Seo
  • Su Jang
  • Joong Hyoun Chin
  • Hee-Jong Koh
Original Article


Key message

The split-hull phenotype caused by reduced lemma width and low lignin content is under control of SPH encoding a type-2 13-lipoxygenase and contributes to high dehulling efficiency.


Rice hulls consist of two bract-like structures, the lemma and palea. The hull is an important organ that helps to protect seeds from environmental stress, determines seed shape, and ensures grain filling. Achieving optimal hull size and morphology is beneficial for seed development. We characterized the split-hull (sph) mutant in rice, which exhibits hull splitting in the interlocking part between lemma and palea and/or the folded part of the lemma during the grain filling stage. Morphological and chemical analysis revealed that reduction in the width of the lemma and lignin content of the hull in the sph mutant might be the cause of hull splitting. Genetic analysis indicated that the mutant phenotype was controlled by a single recessive gene, sph (Os04g0447100), which encodes a type-2 13-lipoxygenase. SPH knockout and knockdown transgenic plants displayed the same split-hull phenotype as in the mutant. The sph mutant showed significantly higher linoleic and linolenic acid (substrates of lipoxygenase) contents in spikelets compared to the wild type. It is probably due to the genetic defect of SPH and subsequent decrease in lipoxygenase activity. In dehulling experiment, the sph mutant showed high dehulling efficiency even by a weak tearing force in a dehulling machine. Collectively, the results provide a basis for understanding of the functional role of lipoxygenase in structure and maintenance of hulls, and would facilitate breeding of easy-dehulling rice.







Poly unsaturated fatty acid


Linoleic acid


Linolenic acid


Days after flowering


Bulked segregant analysis


Open reading frame


Chloroplast transit peptide


Marginal region of lemma



We thank Dr. Hong Yeol Kim and Mi Kyeong Kang for their assistance in cultivating rice at the experimental Farm of Seoul National University. We would also like to thank Prof. Gynheung An of Kyunghee University for providing seed of T-DNA line. This study was supported by a grant from the Next-Generation BioGreen 21 Program (No. PJ013165) of the Rural Development Administration, Korea.

Compliance with ethical standards

Conflict of interest

On behalf of all authors, the corresponding author states that there is no conflict of interest.

Supplementary material

122_2018_3091_MOESM1_ESM.docx (10 mb)
Supplementary material 1 (DOCX 10243 kb)
122_2018_3091_MOESM2_ESM.xlsx (13 kb)
Supplementary material 2 (XLSX 13 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Department of Plant Science and Research Institute for Agriculture and Life Sciences, and Plant Genomics and Breeding InstituteSeoul National UniversitySeoulSouth Korea
  2. 2.Graduate School of Integrated BioindustrySejong UniversitySeoulSouth Korea

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