Review

Abstract
References
1

Al-Aswad RMA, Al-Azzawi QKZ (2021) Control of downy mildew disease on cucumber caused by the fungus Pseudoperonospora cubensis by using environmentally friendly materials. Euphrates J Agric Sci 13:98-110. https://doi.org/10.1038/s41598-024-81643-0

10.1038/s41598-024-81643-040188117PMC11972287
2

Ali A, Elrys AS, Liu L, Zhao J, Cheng Z, Cai Z (2023) Deciphering the synergies of reductive soil disinfestation combined with biochar and antagonistic microbial inoculation in cucumber Fusarium wilt suppression through rhizosphere microbiota structure. Microb Ecol 85:980-997. https://doi.org/10.1007/s00248-022-02097-3

10.1007/s00248-022-02097-3
3

Atallah OO, Osman A, Ali MA, Sitohy M (2021) Soybean β‐conglycinin and catfish cutaneous mucous p22 glycoproteins deteriorate sporangial cell walls of Pseudoperonospora cubensis and suppress cucumber downy mildew. Pest Manag Sci 77:3313-3324. https://doi.org/10.1002/ps.6375

10.1002/ps.6375
4

Ávalos-Sánchez E, Moreno-Teruel MÁ, Molina-Aiz FD, López-Martínez A, Peña-Fernández A, Baptista F, Valera-Martínez DL (2022) Influence of the diffusivity and transmittance of a plastic greenhouse cover on the development of fungal diseases in a cucumber crop. Agronomy 12:2743. https://doi.org/10.3390/agronomy12112743

10.3390/agronomy12112743
5

Bai ZL, Yuan XJ, Cai R, Liu LZ, He HL, Zhou HF, Pan JS (2008) QTL mapping of resistance gene to downy mildew in cucumber. Prog Nat Sci 18:706-710. https://doi.org/10.3321/j.issn:1002-008X.2008.06.015

10.3321/j.issn:1002-008X.2008.06.015
6

Barnea D, Yermiyahu U, Rav-David D, Elad Y (2022) Effect of mineral nutrition and salt spray on cucumber downy mildew (Pseudoperonospora cubensis). Plants 11:1007. https://doi.org/10.3390/plants11081007

10.3390/plants1108100735448736PMC9024561
7

Barnes WC, Epps WM (1954) An unreported type of resistance to cucumber downy mildew. Plant Dis Rep 38:620-620

8

Berg JA (2019) Cucumber mildew resistance: identification of cucumber genes involved in susceptibility and resistance to powdery and downy mildew. Wageningen UR, https://doi.org/10.18174/497042

10.18174/497042
9

Bi YM, Zhang Y, Signorelli T, Zhao R, Zhu T, Rothstein S (2005) Genetic analysis of Arabidopsis GATA transcription factor gene family reveals a nitrate‐inducible member important for chlorophyll synthesis and glucose sensitivity. Plant J 44:680-692. https://doi.org/10.1111/j.1365-313X.2005.02568.x

10.1111/j.1365-313X.2005.02568.x
10

Caldwell D, Chan E, De Vries J, Joobeur T, King J (2011) Methods and compositions for identifying downy mildew resistant cucumber plants: U.S. Patent WO2011050296. 28.04.2011

11

Call AD (2010) Studies on resistance to downy mildew in cucumber (Cucumis sativus L.) caused by Pseudoperonospora cubensis. NC State. https://api.semanticscholar.org/CorpusID:87386007

12

Call AD, Criswell AD, Wehner TC, Klosinska U, Kozik EU (2012) Screening cucumber for resistance to downy mildew caused by Pseudoperonospora cubensis (Berk. and Curt.) Rostov. Crop Science, 52:577-592. https://doi.org/10.2135/cropsci2011.06.0296

10.2135/cropsci2011.06.0296
13

Call AD, Wehner TC (2010.) Search for higher resistance to the new race of downy mildew in cucumber

14

Cavatorta J, Moriarty G, Henning M, Glos M, Kreitinger M, Munger HM, Jahn M (2007) ‘Marketmore 97’: a monoecious slicing cucumber inbred with multiple disease and insect resistances. HortScience 42:707-709. https://doi.org/10.21273/HORTSCI.42.3.707

10.21273/HORTSCI.42.3.707
15

Cohen Y (1971) Field and growth chamber approach to epidemiology of Pseudoperonospora cubensis on cucumbers. Phytopathology 61:736. https://doi.org/10.1094/Phyto-61-736

10.1094/Phyto-61-736
16

Cohen Y, Eyal H (1977) Growth and differentiation of sporangia and sporangiophores of Pseudoperonospora cubensis on cucumber cotyledons under various combinations of light and temperature. Physiol Plant Pathol 10:93-103. https://doi.org/10.1016/0048-4059(77)90013-3

10.1016/0048-4059(77)90013-3
17

Colucci SJ, Wehner TC, Holmes GJ (2006) The downy mildew epidemic of 2004 and 2005 in the eastern United States. pp. 403–410. https://api.semanticscholar.org/CorpusID:127582147

18

Cortés AJ, López-Hernández F, Blair MW (2022) Genome–environment associations, an innovative tool for studying heritable evolutionary adaptation in orphan crops and wild relatives. Front Genet 13:910386. https://doi.org/10.3389/fgene.2022.910386.

10.3389/fgene.2022.91038635991553PMC9389289
19

Elsharkawy M, Kamel S, El-Khateeb N (2014) Biological control of powdery and downy mildews of cucumber under greenhouse conditions. Egypt J Biol Pest Co 24:407-414

20

Evangelisti E, Rey T, Schornack S (2014) Cross-interference of plant development and plant–microbe interactions. Curr Opin Plant Biol 20:118-126. https://doi.org/10.1016/j.pbi.2014.05.014

10.1016/j.pbi.2014.05.014
21

Fan YT, Chung KR, Huang JW (2019) Fungichromin production by Streptomyces padanus PMS-702 for controlling cucumber downy mildew[J/OL]. Plant Pathology J 35:341-350. https://doi.org/10.5423/PPJ.OA.03.2019.0057

10.5423/PPJ.OA.03.2019.005731481857PMC6706012
22

Fanourakis NE, Simon PW (1987) Analysis of genetic linkage in the cucumber. J Hered 78:238-242. https://doi.org/10.1093/oxfordjournals.jhered.a110374

10.1093/oxfordjournals.jhered.a110374
23

Guo SM, Niu ZF (2012) The effect of temperature and humidity on the survival of cucumber downy mildew sporangia. North Hortic 23:142-144

24

Hammer RS, Cohen Y (2025) Non-Sikkim cucumber accessions resistant to downy mildew (Pseudoperonospora cubensis). Seeds, 4:8. https://doi.org/10.3390/seeds4010008

10.3390/seeds4010008
25

Holdsworth W, Summers CF, Glos M, Smart CD, Mazourek M (2014) Development of downy mildew-resistant cucumbers for late-season production in the northeastern United States. HortScience 49:10-17. https://doi.org/10.21273/HORTSCI.49.1.10

10.21273/HORTSCI.49.1.10
26

Holmes G, Thomas C (2009) The history and re-emergence of cucurbit downy mildew. Phytopathology 99:S171-S171

27

Holmes G, Wehner T, Thornton A (2006) An old enemy re-emerges. American Vegetable Grower 54:14-15

28

Holmes GJ, Main CE, Zeev ZT (2004) Cucurbit downy mildew: a unique pathosystem for disease forecasting. pp 69-80. https://doi.org/10.1007/978-1-4020-2658-4_3

10.1007/978-1-4020-2658-4_3
29

Horejsi T, Staub JE, Thomas C (2000) Linkage of random amplified polymorphic DNA markers to downy mildew resistance in cucumber (Cucumis sativus L.). Euphytica 115:105–113. https://doi.org/10.1023/A:1003942228323

10.1023/A:1003942228323
30

Innark P, Panyanitikoon H, Khanobdee C, Samipak S, Jantasuriyarat C (2020) QTL identification for downy mildew resistance in cucumber using genetic linkage map based on SSR markers. J Genet 99:81. https://doi.org/10.1007/s12041-020-01242-6

10.1007/s12041-020-01242-6
31

Jenkins JM (1942) Downy mildew resistance in cucumbers. J Hered 33:35-38. https://doi.org/10.1093/oxfordjournals.jhered.a105122

10.1093/oxfordjournals.jhered.a105122
32

Kanetis L, Holmes GJ, Ojiambo PS (2010) Survival of Pseudoperonospora cubensis sporangia exposed to solar radiation. Plant Pathol 59:313-323. https://doi.org/10.1111/j.1365-3059.2009.02211.x

10.1111/j.1365-3059.2009.02211.x
33

Kelly JD, Vallejo V (2006) QTL analysis of multigenic disease resistance in plant breeding, In: Tuzun, S., Bent, E. (eds) Multigenic and Induced Systemic Resistance in Plants. Springer, Boston, MA, US, pp 21-48. https://doi.org/10.1007/0-387-23266-4_3

10.1007/0-387-23266-4_3
34

Khudhair AA, Aljarah NS (2023) The role of the relative humidity on the development of downy mildew infection of cucumber in the greenhouse in Baghdad. Acta Fytotech Zootech IOP Conference Series: Earth and Environmental Science 1252:012019. https://doi.org/10.1088/1755-1315/1252/1/012019

10.1088/1755-1315/1252/1/012019
35

Lebeda A, Urban J (2004) Distribution, harmfulness and pathogenic variability of cucurbit downy mildew in the Czech Republic. 7:170-173

36

Lebeda A, Urban J (2007) Temporal changes in pathogenicity and fungicide resistance in Pseudoperonospora cubensis populations. Acta Hortic 731:327-336. https://doi.org/10.17660/ActaHortic.2007.731.44

10.17660/ActaHortic.2007.731.44
37

Li DG, Xie Z (2024) Occurrence and integrated control measures of cucumber downy mildew. Northw Hortic 3:35-37

38

Li JA (2025) Prevention and Control Techniques For Cucumber Downy Mildew In Early Spring Greenhouses. Henan Agric 19:72. https://doi.org/10.15904/j.cnki.hnny.2025.19.038

10.15904/j.cnki.hnny.2025.19.038
39

Li L, He H, Zou Z, Li Y (2018) QTL analysis for downy mildew resistance in cucumber inbred line PI 197088. Plant Dis 102:1240-1245. https://doi.org/10.1094/PDIS-04-17-0491-RE

10.1094/PDIS-04-17-0491-RE
40

Li X (2024) Research progress on molecular marker-assisted selection breeding in crops. Ningxia J Agric For Sci Technol 65:35–43

41

Li Y, Ge XZ (2022) Construction of cucumber downy mildew early warning system based on PCA and multiple regression algorithm. J Anhui Agric Sci 50:232-234

42

Li YQ, Liu HQ, Wang Y, Yang LJ, Gao W (2024) The effect of temperature on the growth of Pseudoperonospora cubensis spores and its infection process on cucumber. Proceedings of the 2024 Annual Meeting of the Chinese Society for Plant Pathology, Changchun, Jilin, China, 6, August

43

Liu D, Xin M, Zhou X, Wang C, Zhang Y, Qin Z (2017) Expression and functional analysis of the transcription factor-encoding gene CsERF004 in cucumber during Pseudoperonospora cubensis and Corynespora cassiicola infection. BMC Plant Biol 17:96. https://doi.org/10.1186/s12870-017-1049-8

10.1186/s12870-017-1049-828583084PMC5460474
44

Liu X, Lu H, Liu P, Miao H, Bai Y, Gu X, Zhang S (2020) Identification of novel loci and candidate genes for cucumber downy mildew resistance using GWAS. Plants 9:1659. https://doi.org/10.3390/plants9121659

10.3390/plants912165933260843PMC7768435
45

Luan Q, Chen C, Liu M, Li Q, Wang L, Ren Z (2019) CsWRKY50 mediates defense responses to Pseudoperonospora cubensis infection in Cucumis sativus. Plant Sci 279:59-69. https://doi.org/10.1016/j.plantsci.2018.11.002

10.1016/j.plantsci.2018.11.002
46

Palti J, Cohen Y (1980) Downy mildew of Cucurbits (Pseudoperonospora Cubensis): the Fungus and its hosts, distribution, epidemiology and control. Phytoparasitica 8:109-147. https://doi.org/10.1007/BF02994506

10.1007/BF02994506
47

Pang X, Zhou X, Wan H, Chen J (2013) QTL mapping of downy mildew resistance in an introgression line derived from interspecific hybridization between cucumber and Cucumis hystrix. J Phytopathol 161:536-543. https://doi.org/10.1111/jph.12103

10.1111/jph.12103
48

Pershin AF, Medvedeva NI, Nedvedev AV (1988) Quantitative approach to studying the genetics of disease resistance. IV. Interaction of the genetic systems for resistance to powdery and downy mildews in cucumber. Geneticka, Ussr, 24:484-493

49

Qiao C, Han M, Gao W, Li K, Zhu X, Zhang L (2023) Detection of cucumber downy mildew spores based on Faster-NAM-YOLO. Trans Chin Soc Agric Mach 54:288-299. https://doi.org/10.1016/j.inpa.2024.05.002

10.1016/j.inpa.2024.05.002
50

Qiao C, Li K, Zhu X, Jing J, Gao W, Zhang L (2024) Detection of cucumber downy mildew spores based on improved YOLOv5s. Inform Process Agr 2214-3173. https://doi.org/10.1016/j.inpa.2024.05.002

10.1016/j.inpa.2024.05.002
51

Savory EA, Granke LL, Quesada-Ocampo LM, Varbanova M, Hausbeck MK, Day B (2011) The cucurbit downy mildew pathogen Pseudoperonospora cubensis. Mol Plant Pathol 12:217-226. https://doi.org/10.1111/j.1364-3703.2010.00670.x

10.1111/j.1364-3703.2010.00670.x21355994PMC6640371
52

Schuster C, Schmitt A (2018) Efficacy of a bacterial preparation of Aneurinibacillus migulanus against downy mildew of cucumber (Pseudoperonospora cubensis). Eur J Plant Pathol 151:439-450. https://doi.org/10.1007/s10658-017-1385-4

10.1007/s10658-017-1385-4
53

Shimizu S, Kanazawa K, Kato A (1963) Studies on the breeding of cucumber for resistance to downy mildew: Part 2. Difference of resistance to downy mildew among cucumber varieties and the utility of cucumber variety resistance to downy mildew. Bull Hort Res Sta Jpn 2:80-81

54

Sun Z, Yu S, Hu Y, Wen Y (2022) Biological control of the cucumber downy mildew pathogen Pseudoperonospora cubensis. Horticulturae 8:410. https://doi.org/10.3390/horticulturae8050410

10.3390/horticulturae8050410
55

Szczech M, Nawrocka J, Felczyński K, Małolepsza U, Sobolewski J, Kowalska B, Maciorowski R, Jas K, Kancelista A (2017) Trichoderma atroviride TRS25 isolate reduces dowFny mildew and induces systemic defence responses in cucumber in field conditions. Sci Hortic 224:17-26. https://doi.org/10.1016/j.scienta.2017.05.035

10.1016/j.scienta.2017.05.035
56

Tan J, Wang Y, Dymerski R, Wu Z, Weng Y (2022) Sigma factor binding protein 1 (CsSIB1) is a putative candidate of the major-effect QTL dm5.3 for downy mildew resistance in cucumber (Cucumis sativus). Theor Appl Genet 135:4197-4215. https://doi.org/10.1007/s00122-022-04212-x

10.1007/s00122-022-04212-x
57

Van Vliet GJA, Meijsing WD (1977) Relation in the inheritance of resistance to Pseudoperonospora cubensis ROST and Sphaerotheca fuliginea POLL. in cucumber (Cucumis sativus L.). Euphytica 26:793-796. https://doi.org/10.1007/BF00021708

10.1007/BF00021708
58

Vandenlangenberg KM (2015) Studies on downy mildew resistance in cucumber (Cucumis sativus L.). Raleigh: NC State

59

Wang B, Xue Z, Lan J, Sun M, Sun Q, Huang Z, Zhang C, Liu X (2023) Activity of the new OSBP inhibitor Y18501 against Pseudoperonospora cubensis and its application for the control of cucumber downy mildew. Pestic Biochem Phys 194:105415. https://doi.org/10.1016/j.pestbp.2023.105415

10.1016/j.pestbp.2023.105415
60

Wang Y, Fu SR, Wang SW, Wang SY, Li YL, Hu B, Cao JJ, Zhang XL, Zhang AH (2024a) Control measures for tomato gray mold and cucumber downy mildew in facility cultivation. Agric Eng Technol 44:96-98. https://doi.org/10.16815/j.cnki.11-5436/s.2024.21.039

10.16815/j.cnki.11-5436/s.2024.21.039
61

Wang Y, Jin GT (2025) Symptoms and control measures of cucumber downy mildew. Mod Rural Sci Technol 4:68-69

62

Wang Y, Tan J, Wu Z, VandenLangenberg K, Wehner TC, Wen C, Zheng X, Owens K, Thornton A, et al. (2018a) STAYGREEN, STAY HEALTHY: a loss-of-susceptibility mutation in the STAYGREEN gene provides durable, broad-spectrum disease resistances for over 50 years of US cucumber production. New Phytol 221:415-430. https://doi.org/10.1111/nph.15353

10.1111/nph.15353
63

Wang Y, VandenLangenberg K, Wehner TC, Kraan PAG, Suelmann J, Zheng X, Owens K, Weng Y (2016) QTL mapping for downy mildew resistance in cucumber inbred line WI7120 (PI 330628). Theor Appl Genet 129:943-953. https://doi.org/10.1007/s00122-016-2719-x

10.1007/s00122-016-2719-x
64

Wang Y, VandenLangenberg K, Wen C, Wehner TC, Weng Y (2018b) QTL mapping of downy and powdery mildew resistances in PI 197088 cucumber with genotyping-by-sequencing in RIL population. Theor Appl Genet 131:597-611. https://doi.org/10.1007/s00122-017-3022-1

10.1007/s00122-017-3022-1
65

Wang Z, Yin M, Han J, Wang X, Chang J, Ren Z, Wang L (2024b) Pan-Genome-Wide Identification and Transcriptome-Wide Analysis of ZIP Genes in Cucumber. Agriculture 14:133. https://doi.org/10.3390/agriculture14010133

10.3390/agriculture14010133
66

Wen DM, Chen MX, Zhao L, Ji T, Li M, Yang XT (2019) Use of thermal imaging and Fourier transform infrared spectroscopy for the pre-symptomatic detection of cucumber downy mildew. Eur J Plant Pathol 155:405-416. https://doi.org/10.1007/s10658-019-01775-2

10.1007/s10658-019-01775-2
67

Weng Y (2008) 2008 public sector cucumber research priority survey. Cucurbit Genetics Cooperative Report, 31-32:1-4.

68

Win KT, Vegas J, Zhang C, Song K, Lee S (2017) QTL mapping for downy mildew resistance in cucumber via bulked segregant analysis using next-generation sequencing and conventional methods. Theor Appl Genet 130:199-211. https://doi.org/10.1007/s00122-016-2806-z

10.1007/s00122-016-2806-z
69

Wu RF (2024) Identification and control of cucumber downy mildew. J Changjiang Veg 9:58-59.

70

Xu GL (2024) Study on the transmission pattern and early warning technology of cucumber downy mildew. JSU, Jiangsu, China.

71

Xu SC, Wang HB, Feng JJ, Xiang HF, Wu MD, Wang ZM, Wei DY, Zhang HC, Tang QL (2022) Research progress on cucumber downy mildew and host resistance mechanisms. J Biotechnol 38:1724-1737. https://doi.org/10.13345/j.cjb.210513

10.13345/j.cjb.210513
72

Yang F, Wang H, Zhi C, Chen B, Zheng Y, Qiao L, Gao J, Pan Y, Cheng Z (2021) Garlic volatile diallyl disulfide induced cucumber resistance to downy mildew. Int J Mol Sci 22:12328. https://doi.org/10.3390/ijms222212328

10.3390/ijms22221232834830208PMC8625977
73

Yoshioka Y, Sakata Y, Sugiyama M, Fukino N (2014) Identification of quantitative trait loci for downy mildew resistance in cucumber (Cucumis sativus L.). Euphytica 198:265-276. https://doi.org/10.1007/s10681-014-1102-8

10.1007/s10681-014-1102-8
74

Zhang K, Wang X, Zhu W, Qin X, Xu J, Cheng C, Lou Q, Li J, Chen J (2018) Complete resistance to powdery mildew and partial resistance to downy mildew in a Cucumis hystrix introgression line of cucumber were controlled by a co-localized locus. Theor Appl Genet 131:2229-2243. https://doi.org/10.1007/s00122-018-3150-2

10.1007/s00122-018-3150-2
75

Zhang SP, Liu MM, Miao H, Zhang SQ, Yang YH, Xie BY, Wehner TC, Gu XF (2013) Chromosomal mapping and QTL analysis of resistance to downy mildew in Cucumis sativus. Plant Dis 97:245-251. https://doi.org/10.1094/PDIS-11-11-0941-RE

10.1094/PDIS-11-11-0941-RE
76

Zhang X (2022) Research on early detection method of cucumber downy mildew in greenhouse based on hyperspectral imaging. NWAFU, Shaanxi, China

77

Zhang Y (2020) The relationship between environmental temperature and humidity and the infection and incubation stages of cucumber downy mildew (Pseudoperonospora cubensis). SYAU, Shenyang, China

78

Zheng X, Yang J, Lou T, Zhang J, Yu W, Wen C (2019) Transcriptome profile analysis reveals that CsTCP14 induces susceptibility to foliage diseases in cucumber. Int J Mol Sci 20:2582. https://doi.org/10.3390/ijms20102582

10.3390/ijms2010258231130701PMC6567058
Information
  • Publisher :KOREAN SOCIETY FOR HORTICULTURAL SCIENCE
  • Publisher(Ko) :한국원예학회
  • Journal Title :Horticultural Science and Technology
  • Journal Title(Ko) :원예과학기술지
  • Received Date : 2025-12-05
  • Revised Date : 2026-01-20
  • Accepted Date : 2026-04-06