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2026 Vol.39, Issue 5 Preview Page

Research Article

1 October 2026. pp. 338-354
Abstract
References
1

Albrecht, T., B. Büttner, S.B. Carey, R. Seidenberger, A. Lutz, A. Harkess, and S. Gresset. 2024. Independent validation of molecular markers for sex determination on diverse sex chromosomes in hops (Humulus lupulus L.). Brewing Sci. 77:172-183.

2

Almaguer, C., C. Schönberger, M. Gastl, E.K. Arendt, and T. Becker. 2014. Humulus lupulus: A story that begs to be told. A Review. J. Inst. Brewing 120:289-314.

10.1002/jib.160
3

APG IV. 2016. An update of the Angiosperm phylogeny group classification for the orders and families of flowering plants. Bot. J. Linnean Soc. 181:1-20.

10.1111/boj.12385
4

BarthHaas. 2005. BarthHaas Report 2004/2005: The 2004 hop harvest and market. BarthHaas GmbH & Co. KG. p. 24.

5

BarthHaas. 2025. BarthHaas Report 2024/2025: The 2024 hop harvest and market. BarthHaas GmbH & Co. KG. p. 40.

6

Bassil, N.V., B. Gilmore, J.M. Oliphant, K.E. Hummer, and J.A. Henning. 2008. Genic SSRs for European and North American hop (Humulus lupulus L.). Gen. Resour. Crop Evol. 55:119-128.

10.1007/s10722-007-9303-9
7

Beatson, R.A., K.A. Ansell, and T. Graham. 2003. Breeding, development, and characteristics of the hop (Humulus lupulus) cultivar ‘Nelson Sauvin’. New Zealand J. Crop & Horti. Sci. 31:303-309.

10.1080/01140671.2003.9514265
8

Becker, L., I. Boyer, and B. Heldreth. 2024. Safety assessment of hops as used in cosmetics. Inter. J. Toxi. 43 (Supplement 1).

10.1177/10915818231221796
9

Behre, K.E. 1999. The history of beer additives in Europe: A review. Vegetation History and Archaeobotany 8:35-48.

10.1007/BF02042841
10

Bocquet, L., S. Sahpaz, L.L. Hilbert, C. Rambaud, and C. Riviere. 2018. Humulus lupulus L., a very popular beer ingredient and medicinal plant: overview of its phytochemistry, its bioactivity, and its biotechnology. Phytochem Rev. 17:1047-1090.

10.1007/s11101-018-9584-y
11

Brownie, S. and S. Salter. 2010. Treating primary insomnia: The efficacy of valerian and hops. Aust. Family Phys. 39:433-437.

12

Čerenak, A., Z. Satovic, and B. Javornik. 2006. Genetic mapping of hop (Humulus lupulus L.) applied to the detection of QTLs for alpha-acid content. Genome 49:485-494.

10.1139/g06-007
13

Christenhusz, M.J.M., L. Percival-Alwyn, K. Hajdu, J. Connell, and I.J. Leitch. 2025. The genome sequence of the common hop, Humulus lupulus L. Wellcome Open Res. 10:210.

10.12688/wellcomeopenres.24025.140528995PMC12171724
14

Clare, S., P. Schmuker, and K. Altendorf. 2026. Association mapping for hop cone chemistry and morphology identifies natural beneficial allele stacks. The Plant Genome. 19:e70238.

10.1002/tpg2.7023842051043PMC13125966
15

Clark, S.M., V. Vaitheeswaran, S.J. Ambrose, R.W. Purves, and J.E. Page. 2013. Transcriptome analysis of bitter acid biosynthesis and precursor pathways in hop (Humulus lupulus). BMC Plant Bio. 13:12.

10.1186/1471-2229-13-1223347725PMC3564914
16

Clarke, R.C. and M.D. Merlin. 2013. Cannabis: evolution and ethnobotany. Univ. of California Press. p. 452.

17

Dabbous-Wach, D., L. Majidi, J.V. Lorenzetti, J. Paolini, and J. Costa. 2025. Quantification of α-acids, β-acids, and phenolic compounds in Corsican hops using LC-MS/MS and metabolomic approach through molecular networks. Phytochem. Analysis 36:438-457.

10.1002/pca.348139609247PMC11876718
18

Danilova, T.V. and J.H. Henning. 2005. Sex molecular markers reveal differences of Y-chromosome molecular structure among hops of European and American origin. Acta Horticulturae 668:85-90.

10.17660/ActaHortic.2005.668.11
19

Darby, P. 2005. The history of hop breeding and development. Brewery History 121:94-112.

20

Divashuk, M.G., O.S. Alexandrov, P.Y. Kroupin, and G.I. Karlov. 2011. Molecular cytogenetic mapping of Humulus lupulus sex chromosomes. Cytogen. Genome Res. 134:213- 219.

10.1159/000328831
21

Doosan Nongsan. 1995. Hop cultivation history in Korea. Donga Press. Korea. p. 326 (in Korean).

22

Duarte, L.M., T.L. Amorim, R.M. Grazul, M. Augustom, and L. de Oliveira. 2020. Differentiation of aromatic, bittering and dual-purpose commercial hops from their terpenic profiles: An approach involving batch extraction, GC-MS and multivariate analysis. Food Res. Inter. 140:109768.

10.1016/j.foodres.2020.109768
23

Dzienisik, M., M. Marzec, and I. Nowak. 2026. Antioxidant constituents of Hops (Humulus lupulus L.) as functional raw materials for cosmetic applications. Materials 19:821.

10.3390/ma1904082141753540PMC12941662
24

Eyres, G. and J.-P. Dufour. 2009. Hop essential oil: Analysis, chemical composition and odor characteristics. In Beer in Health and Disease Prevention, Academic Press. pp. 239-254.

10.1016/B978-0-12-373891-2.00022-5
25

Gerhäuser, C. 2005. Beer constituents as potential cancer chemopreventive agents. Eur. J. Cancer. 41:1941-1954.

10.1016/j.ejca.2005.04.012
26

Ha, T.H., J.H. Lee, S.Y. Yi, and S.-Y. Kang. 2024a. The effect of plant growth regulators on generation rate during tissue culture of hop (Humulus lupulus L.). Korean J. Plant Res. 37:431-438 (in Korean).

27

Ha, T.H., J.I. Lyu, S.Y. Yi, and S.-Y. Kang. 2024b. Developing DNA markers based on male-specific chromosomal regions for selecting male plants in hop (Humulus lupulus). Plant Breed. Biotech. 12:10-16.

10.9787/PBB.2024.12.10
28

Ha, T.H., J.I. Lyu, J.H. Lee, J. Ryu, S.H. Park, and S.-Y. Kang. 2023. Studies on growth characteristics and propagation method of introduced hop (Humulus lupulus L.) cultivars. Korean J. Plant Res. 36:181-190 (in Korean).

10.7732/KJPR.2023.36.2.181
29

Han, N.Y., S.I. Park, and M. Kim. 2025. Antibacterial activity of Humulus lupulus supercritical fluid extract as a cosmeceutical ingredient of skin microbiome balance. J. Convert. Cul. Tech. 11:1073-1086.

30

Henning, J.A., J. Coggins, S. Hill, D. Hendrix, and S. Townsend. 2019. Genome-wide association study on ten traits of economic importance in hop (Humulus lupulus L.). Acta Horticulturae 1236:113-120.

10.17660/ActaHortic.2019.1236.13
31

Hieronymus, S. 2012. For the love of hops: The practical guide to aroma, bitterness and the culture of hops. Brewers Publications, Boulder, Colorado. USA. p. 326.

32

Hill, S.T., R. Sudarsanam, J. Henning, and D. Hendrix. 2017. HopBase: A unified resource for Humulus genomics. Database 2017:1-10.

10.1093/database/bax00928415075PMC5467566
33

Kale, S.M., N. Kamal, H. Gundlach, O. Gericke, A. Almeida, G. Haberer, L. de Bang, T. Lux, N. Pitra, N. Price, F. Krsticevic, O. Kemp, A. Himmelbach, M.-T. Rabanus-Wallace, K. Nielsen, S. Padmarasu, L. Horáková, V. Bačovský, N. Bjarnholt, I. Kruse-Andersen, B. Skadhauge, N. Micic, B.L. Møller, C. Janfelt, P.D. Matthews, K.F.X. Mayer, N. Stein, M. Mascher, and I. Braumann. 2026. Extensive variation between chromosomes of North American and European hop. Nature Com. 17:4110.

10.1038/s41467-026-72379-842204144PMC13216280
34

Karabín, M., T. Hudcová, L. Jelínek, and P. Dostálek. 2016. Biologically active compounds from hops and prospects for their use. Com. Rev. Food Sci. Food Safety 15:542-567.

10.1111/1541-4337.12201
35

Kew Science. 2026. Humulus lupulus L., Humulus scandens (Lour.) Merr. Plants of the World Online. https://powo.science.kew.org

36

Kiofentzoglou, D., E.M. Andronidou, P.I. Kontou, P.G. Bagos, and G.G. Braliou. 2025. Antimicrobial activity of chemical hop (Humulus lupulus) compounds: A systematic review and meta-Analysis. Applied Science 15:7806.

10.3390/app15147806
37

Koetter, U., E. Schrader, R. Käufeler, and A. Brattström. 2007. A randomized, double blind, placebo-controlled, prospective clinical study to demonstrate clinical efficacy of a fixed valerian hops extract combination (Ze 91019) in patients suffering from non-organic sleep disorder. Phytotherapy Res. 21:847-851.

10.1002/ptr.2167
38

Kolenc, Z., T. Langerholc, G. Hostnik, M. Ocvirk, S. Štumpf, A.M. Pintaric, I.J. Košir, A. Cerenak, A. Garmut, and U. Bren. 2023. Antimicrobial properties of different hop (Humulus lupulus) genotypes, Plants 12:120.

10.3390/plants1201012036616249PMC9824274
39

Korpelainen, H. and M. Pietiläinen. 2021. Hop (Humulus lupulus): Traditional and present use, and future potential. Eco. Bot. 75:302-322.

10.1007/s12231-021-09528-1
40

KSVS. 2026. Online of Korea Seed & Variety Service. https://www.seed.go.kr/seed/252/subview.do.

41

Lee, H., S.H. Chung, D.J. Kwon, M.J. Nam, J.W. Choi, H.J. Suh, H.S. Choi, and S.H. Han. 2025. Sleep-enhancing effect of Hongcheon-hop (Humulus lupulus L.) extract containing xanthohumol and humulone through GABAA receptor. J. Ethnopharmacology 338:119019.

10.1016/j.jep.2024.119019
42

Ling, L.-Z. and S.H. Zhang. 2019. The complete chloroplast genome of Humulus yunnanensis and phylogenetic analysis of the genus Humulus. Mitochondrial DNA PART B. 4:2681-2682.

10.1080/23802359.2019.164424333365681PMC7706996
43

Lyu, J.I., J. Ryu, K.S. Seo, K.Y. Kang, S.H. Park, T.H. Ha, J.W. Ahn, and S.-Y. Kang. 2022. Comparative study on phenolic compounds and antioxidant activities of hop (Humulus lupulus L.) strobile extracts. Plants 11:135.

10.3390/plants1101013535009138PMC8747208
44

Mafakheri, M., M. Kordrostami, and P. Matthews. 2020. Evaluating genetic diversity and structure of a wild hop (Humulus lupulus L.) germplasm using morphological and molecular characteristics. Euphytica 216:58.

10.1007/s10681-020-02592-z
45

Marceddu, R.M., A. Carrubba, and M. Sarno. 2022. Resilience of hop (Humulus lupulus L.) to salinity, heat and drought stresses: A mini-review. Fron. Plant Sci. 13:1064922.

10.3389/fpls.2022.106492236531342PMC9749550
46

Matthews, P.D. and A. Lutz. 2017. Hop (Humulus lupulus L.). In J. Janick (Ed.), Plant Breeding Rev. 41:89-149.

47

Matthews, P.D., M.C. Coles, and N.J. Pitra. 2013. Next generation sequencing for a plant of great tradition: Application of NGS to SNP detection and validation in hops (Humulus lupulus L.). Brewing Sci. 66:185-191.

48

Mishra, A.K., T. Kocábek, V.S. Nath, P. Awasthi, A. Shrestha, U.K. Killi, J. Jakse, J. Patzak, K. Krofta, and J. Matoušek. 2020. Dissection of dynamic transcriptome landscape of leaf, bract, and lupulin gland in hop (Humulus lupulus L.). Inter. J. Mol. Sci. 21:233.

10.3390/ijms2101023331905722PMC6981390
49

Morin, C.M., U. Koetter, C. Bastien, J.C. Ware, and V. Wooten. 2005. Valerian-hops combination and diphenhydramine for treating insomnia: A randomized placebo-controlled clinical trial. Sleep 28:1465-1471.

10.1093/sleep/28.11.1465
50

Mozny, M., R. Tolasz, J. Nekovar, T. Sparks, M. Trnka, and Z. Zalud. 2009. The impact of climate change on the yield and quality of Saaz hops in the Czech Republic. Agri. Forest Meteoro. 149:913-919.

10.1016/j.agrformet.2009.02.006
51

Murakami, A., P. Darby, B. Javornik, M.S.S. Pais, E. Seigner, A. Lutz, and P. Svoboda. 2006a. Molecular phylogeny of wild Hops, Humulus lupulus L. Heredity 97:66-74.

10.1038/sj.hdy.6800839
52

Murakami, A., P. Darby, B. Javornik, M.S.S. Pais, E. Seigner, A. Lutz, and P. Svoboda. 2006b. Microsatellite DNA analysis of wild hops, Humulus lupulus L. Gen. Resour. Crop Evol. 53:1553-1562.

10.1007/s10722-005-7765-1
53

Natsume, S., H. Takagi, A. Shiraishi, J. Murata, H. Toyonaga, J. Patzak, M. Takagi, H. Yaegashi, A. Uemura, C. Mitsuoka, K. Yoshida, K. Krofta, H. Satake, R. Terauchi, and E. Ono. 2015. The draft genome of hop (Humulus lupulus), an essence for brewing. Plant Cell Physiol. 56:428-441.

10.1093/pcp/pcu169
54

Neve, R.A. 1991. Hops. Chapman and Hall. p. 266.

10.1007/978-94-011-3106-3
55

Oledzka, E. 2024. Xanthohumol—A miracle molecule with biological activities: A review of biodegradable polymeric carriers and naturally derived compounds for its delivery. Inter. J. Mol. Sci. 25:3398.

10.3390/ijms2506339838542371PMC10970401
56

Padgitt-Cobb, L.K., S.B. Kingan, J. Wells, E. Justin J., B. Kronmiller, D. Moore, G. Concepcion, P. Peluso, D. Rank, P. Jaiswal, J. Henning, and D.A. Hendrix. 2021. A draft phased assembly of the diploid Cascade hop (Humulus lupulus) genome. Plant Genome 14:e20072.

10.1002/tpg2.2007233605092PMC12807344
57

Padgitt-Cobb, L.K., N.J. Pitra, P.D. Matthews, J.A. Henning, and D.A. Hendrix. 2023. An improved assembly of the “Cascade” hop (Humulus lupulus L.) genome uncovers signatures of molecular evolution and refines time of divergence estimates for the Cannabaceae family. Horti. Res. 10:uhac281.

10.1093/hr/uhac28136818366PMC9930403
58

Paguet, A.S., A. Siah, G. Lefe`vre, S. Sahpaz, and C. Rivie`re. 2022. Agronomic, genetic and chemical tools for hop cultivation and breeding. Phytochem Rev. 21:667-708.

10.1007/s11101-022-09813-4
59

Patzak, J. 2001. Comparison of RAPD, STS, ISSR and AFLP molecular methods used for assessment of genetic diversity in hop (Humulus lupulus L.). Euphytica 121:9-18.

10.1023/A:1012099123877
60

Polley, A., M.W. Ganal, and E. Seigner. 1997. Identification of sex in hop (Humulus lupulus) using molecular markers. Genome 40:357-361.

10.1139/g97-048
61

Pusiak, R.J.P., D.P. Overy, R. Liu, and C.S. Harris. 2026. Fruit of the bine: exploring the chemodiversity of cultivated hops (Humulus lupulus L.). Botany 104:1-15.

10.1139/cjb-2025-0054
62

RDA Library Databases. 2026. Online Databases of Agricultural Science Library, RDA. https://lib.rda.go.kr/eng/ main.do.

63

Řeřicha, J., M. Kohútek, V. Vandírková, K. Krofta, F. Kumhála, and J. Kumhálová. 2025. Assessment of UAV imageries for estimating growth vitality, yield and quality of hop (Humulus lupulus L.) crops. Remote Sensing 17:970.

10.3390/rs17060970
64

Ruggeri, R., P. Loussert, F. Rossini, L.K. Rutto, S.R. Roberto, and S. Agehara. 2024. Development of hop cultivation in new growing areas: The state of the art and the way forward. Eur. J. Agro. 161:127335.

10.1016/j.eja.2024.127335
65

Salanta, L.C., A.C. Farcas, A. Borsa, and C.R. Pop. 2023. Current strategies for the management of valuable compounds from hops waste for a circular economy. Food Chem. X. 19:100876.

10.1016/j.fochx.2023.10087637780312PMC10534220
66

Santos, B., J. Garcia, I. Gouvinhas, M.J. Alves, O.R. Pereira, M.C. Seijo, and M.J. Sousa. 2026. Humulus lupulus L. and its potential application in dermocosmetics. Pharma. Res.-Nat. Prod. 11:100779.

10.1016/j.prenap.2026.100779
67

Schicktanz, N., C. Gerhards, T. Schlitt, A. Amanda, E. Müggler, D. de Quervain, A. Papassotiropoulos, G. Boonen, J. Drewe, and V. Butterwecket. 2025. Effects of a valerian-hops extract combination (Ze 91019) on sleep duration and daytime cognitive and psychological parameters in occasional insomnia: A randomized controlled feasibility trial. Brain & Behavior 15:e70600.

10.1002/brb3.7060040462685PMC12134488
68

Silva, G.V.A., G.D. Arend, A.A.F. Zielinski, M. Di Luccio, and A. Ambrosi. 2023. Xanthohumol properties and strategies for extraction from hops and brewery residues: A review. Food Chem. 404:134629.

10.1016/j.foodchem.2022.134629
69

Small, E. 1978. A numerical and nomenclatural analysis of morphogeographic taxa in Humulus. Syst. Bot. 3:37-76.

10.2307/2418532
70

Stevens, J.F. and J.E. Page. 2004. Xanthohumol and related prenylflavonoids from hops and beer: to your good health!. Phytochemstry 65:1317-1330.

10.1016/j.phytochem.2004.04.025
71

Stevens, J.F., C. Miranda, B. Frei, and D.R. Buhler. 1998. Inhibition of peroxynitrite-mediated LDL oxidation by prenylated flavonoids: The α,β-unsaturated keto functionality of 2′-hydroxychalcones as a novel antioxidant pharmacophore. Chem. Res. Toxicol. 16:1277-1286.

10.1021/tx020100d
72

Takoi, K. 2019. “Flavor hops” varieties and various flavor compounds contributing to their “varietal aromas”: A review. Technical Quarterly 56:113-123.

73

Tembrock, L.R., J.M. McAleer, and T.M. Gilligan. 2016. A revision of native North American Humulus (Cannabaceae). J. Bot. Res. Inst. Texas 10:11-30.

74

Thomas, G.G., M. Galaverni, I. Marchioni, and M. Rodolfi. 2026. Hop botany and physiology. In K. Carbone (ed). All about Hops: The crop, its cultivation and its uses in brewing and beyond. Springer. pp. 27-58.

10.1007/978-3-031-96606-4_2
75

Valente, J.V., R. Palmeira-de-Oliveira, L. Guiomar, C.V. Vaz, J. Rolo, C. Gaspar, A.S. Oliveira, D. Caramelo, L. Breitenfeld, J.C. Goncalves, F. Delgado, J. Martinez-de-Oliveira, and A. Martinez-de-Oliveira. 2024. Humulus lupulus aqueous extract and hydrolate as a potential ingredient for cosmetics: Chemical characterization and in vitro antimicrobial, cytotoxicity, antioxidant and anti-inflammatory assessment. Fitoterapia 175:105861.

10.1016/j.fitote.2024.105861
76

Zanoli, P. and M. Zavatti. 2008. Pharmacognostic and pharmacological profile of Humulus lupulus L. J. Ethnophamacology 116:383-39.

10.1016/j.jep.2008.01.011
77

Zugravu, C.-A., R.-E. Bohiltea, T. Salmen, E. Pogurschi, and M.-R. Otelea. 2022. Antioxidants in hops: Bioavailability, health effects and perspectives for new products. Antioxidants 11:241.

10.3390/antiox1102024135204124PMC8868281
Information
  • Publisher :The Plant Resources Society of Korea
  • Publisher(Ko) :한국자원식물학회
  • Journal Title :Korean Journal of Plant Resources
  • Journal Title(Ko) :한국자원식물학회지
  • Volume : 39
  • No :5
  • Pages :338-354
  • Received Date : 2026-07-29
  • Revised Date : 2026-08-20
  • Accepted Date : 2026-09-01