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Article|18 Jul 2023|OPEN
Basic leucine zipper gene VvbZIP61 is expressed at a quantitative trait locus for high monoterpene content in grape berries
Yuyu Zhang1,2 ,† , Cuixia Liu3 ,† , Xianju Liu1,2 ,† , Zemin Wang4 , Yi Wang1 , Gan-yuan Zhong5 , Shaohua Li1 , and Zhanwu Dai1,2 , Zhenchang Liang1,2 , , Peige Fan,1,2 ,
1Beijing Key Laboratory of Grape Science and Enology, and CAS Key Laboratory of Plant Resources, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
2University of Chinese Academy of Sciences, Beijing 100049, China
3Centre for Special Economic Plant Studies, Guangxi Institute of Botany, Guangxi Zhuang Autonomous Region and Chinese Academy of Sciences, Guilin 541006, Guangxi, China
4College of Life Science and Technology, Gansu Agricultural University, Lanzhou 730070, China
5Grape Genetics Research Unit, USDA-ARS, Geneva 14456, USA
*Corresponding author. E-mail:,,
Yuyu Zhang and Cuixia Liu,Xianju Liu contributed equally to the study.

Horticulture Research 10,
Article number: uhad151 (2023)
Views: 79

Received: 16 Feb 2023
Revised: 26 Jul 2023
Published online: 18 Jul 2023


The widely appreciated muscat flavor of grapes and wine is mainly attributable to the monoterpenes that accumulate in ripe grape berries. To identify quantitative trait loci (QTL) for grape berry monoterpene content, an F1 mapping population was constructed by a cross between two grapevine genotypes, one with neutral aroma berries (cv. ‘Beifeng’) and the other with a pronounced muscat aroma (elite Vitis vinifera line ‘3–34’). A high-density genetic linkage map spanning 1563.7 cM was constructed using 3332 SNP markers that were assigned to 19 linkage groups. Monoterpenes were extracted from the berry of the F1 progeny, then identified and quantified by gas chromatography–mass spectrometry. Twelve stable QTLs associated with the amounts of 11 monoterpenes in berries were thus identified. In parallel, the levels of RNA in berries from 34 diverse cultivars were estimated by RNA sequencing and compared to the monoterpene content of the berries. The expression of five genes mapping to stable QTLs correlated well with the monoterpene content of berries. These genes, including the basic leucine zipper VvbZIP61 gene on chromosome 12, are therefore considered as potentially being involved in monoterpene metabolism. Overexpression of VvbZIP61 in Vitis amurensis callus through Agrobacterium-mediated transformation significantly increased the accumulation of several monoterpenes in the callus, including nerol, linalool, geranial, geraniol, β-myrcene, and D-limonene. It is hypothesized that VvbZIP61 expression acts to increase muscat flavor in grapes. These results advance our understanding of the genetic control of monoterpene biosynthesis in grapes and provide important information for the marker-assisted selection of aroma compounds in grape breeding.