Effects of downy mildew fungi spore infection on cucumber plants under nitrogen stress
Abstract
Investigating the interplay between biotic stress from downy mildew and abiotic stress from nitrogen deficiency is crucial for improving crop management measures and enhancing cucumber production. A greenhouse experiment was conducted using two pathogen treatments—non-infected (B1) and infected with Pseudoperonospora cubensis (B0)—across three nitrogen levels: deficiency (N1: 50%), optimal (N2: 100%), and excess (N3: 150%). The resulting six treatment combinations (B1N1, B1N2/Control, B1N3, B0N1, B0N2, and B0N3) revealed that downy mildew infection and nitrogen application rates significantly influenced key physiological and biochemical parameters (p<0.05). These included sucrose, soluble sugar, hydrogen peroxide (H2O2), catalase (CAT), superoxide dismutase (SOD), polyphenol oxidase (PPO), and malondialdehyde (MDA), as well as the fresh and dry weights of the leaves, stems, and roots. Among all groups, the combination of infection and nitrogen deficiency (B0N1) had the most significant impact on biomass accumulation and hormone metabolism. Compared to the B1N2 control, B0N1 led to substantial reductions in sucrose (52.83%), soluble sugar (68.67%), leaf fresh weight (56.67%), leaf dry weight (55.51%), stem fresh weight (52.82%), stem dry weight (57.28%), root fresh weight (32.46%), and root dry weight (54.07%). This study clarifies the interactive physiological responses of cucumbers to combined biotic and abiotic stress. It is of great significance for facilitating the control of downy mildew and the improvement of cucumber yield in sustainable agriculture.
Keywords: Cucumis sativusL, biotic stress; abiotic stress, physiological metabolism, substance accumulation
DOI: 10.25165/j.ijabe.20261902.10264
Citation: Wang Y F, Shi Q, Du X X, Mao H P, Taha M F. Effects of downy mildew fungi spore infection on cucumber plants under nitrogen stress. Int J Agric & Biol Eng, 2026; 19(2): 58–64.
References
[1] Yan H F, Acquah S J, Zhang J Y, Wang G. Q, Zhang C, Darko R O. Overview of modelling techniques for greenhouse microclimate environment and evapotranspiration. Int J Agric & Biol Eng, 2021; 14: 1–8.
[2] Yan H F, Deng S S, Zhang C, Wang G Q, Zhao S, Li M, et al. Determination of energy partition of a cucumber grown Venlo-type greenhouse in southeast China. Agric. Water Manag., 2023; 276: 108047.
[3] Abubaker B A, Yan H F, Li H, Wu Y Y, Elshaikh N A, Hussei G, et al. Enhancement of depleted loam soil as well as cucumber productivity utilizing biochar under water stress. Commun. Soil Sci. Plant Anal, 2019; 50(1): 49–64.
[4] Zhu W D, Sun J, Wang S M, Shen J F, Yang K F, Zhou X. Identifying field crop diseases using transformer-embedded convolutional neural network. Agriculture-basel, 2022; 12(8): 1083.
[5] Yan H F, Ma J M, Zhang J Y, Wang G Q, Zhang C, Akhlaq M, et al. Effects of film mulching on the physiological and morphological parameters and yield of cucumber under insufficient drip irrigation. Irrigation and Drainage, 2022; 71(4): 897–911.
[6] Wang Y F, Shi Q, Lin J L, Lu X T, Ye B, Lv H X, et al. Hormone metabolism and substance accumulation in cucumber plants: Downy mildew infection and potassium stress. Agriculture-basel, 2025; 15(9): 994.
[7] Shi Q, You L, Wang Y F, Du X X, Chen T H. Effects of downy mildew infection and potassium on growth and physiological traits of greenhouse cucumber. Agronomy-basel, 2025; 15(5): 1017.
[8] Mustapha A T, Zhou C S. Novel assisted/unassisted ultrasound treatment: Effect on respiration rate, ethylene production, enzymes activity, volatile composition, and odor of cherry tomato. LWT-Food Science and Technology, 2021; 149: 111779.
[9] Li X, Dong J L, Duan J J, Shen W Z, Duan Z Q. Nitrogen slow-release behavior of oxamide granules in two different types of paddy soils. Pedosphere, 2022; 32(6): 856–865.
[10] Rasool G, Guo X P, Wang Z C, Ali M U, Chen S, Zhang S X, et al. Coupling fertigation and buried straw layer improves fertilizer use efficiency, fruit yield, and quality of greenhouse tomato. Agricultural Water Management, 2020; 239: 106239.
[11] Rasool G, Guo X P, Wang Z C, Chen S, Hamoud Y A, Javed Q. Response of fertigation under buried straw layer on growth, yield, and water-fertilizer productivity of Chinese cabbage under greenhouse conditions. Commun. Soil Sci. Plant Anal, 2019; 50(8): 1030–1043.
[12] Yan Z N, Cao X X, Bing L X, Song J X, Qi Y, Han Q Y, et al. Responses of growth, enzyme activity, and flower bud differentiation of pepper seedlings to nitrogen concentration at different growth stages. Agronomy-basel, 2024; 14(10): 2270.
[13] El-Sharkawy M, Li J, AL-Huqail A A, Hamed M A, Du D L, EL-Khamisy R R. Slow-released fertilizers optimization and experimental impacts on soil fertility and wheat- maize cropping system. Scientia Agricola, 2024; 81: e20230234.
[14] Rasool G, Guo X P, Wang Z C, Chen S, Ullah I, Ali M, et al. Effect of fertigation levels on water consumption, soil total nitrogen, and growth parameters of brassica chinensis under straw burial. Communications in Soil Science and Plant Analysis, 2021; 52(1): 32–34.
[15] Hou P F, Yuan W S, Li G H, Petropoulos E, Xue L X, Feng Y F, et al. Deep fertilization with controlled-release fertilizer for higher cereal yield and N utilization in paddies: The optimal fertilization depth. Agronomy Journal, 2021; 113(6): 5027–5039.
[16] GB/T 17980.26-2000. Pesticide--Guidelines for the field efficacy trials(I)--Fungicides against downy mildew of cucumber. Beijing, China, 2000.
[17] Shao C H, Qiu C F, Qian Y F, Liu G R. Nitrate deficiency decreased photosynthesis and oxidation-reduction processes, but increased cellular transport, lignin biosynthesis and flavonoid metabolism revealed by RNA-Seq in Oryza sativa leaves. PLoS One, 2020; 15(7): e0235975.
[18] Huang W T, Xie Y Z, Chen X F, Zhang J, Chen H H, Ye X, et al. Growth, mineral nutrients, photosynthesis and related physiological parameters of citrus in response to nitrogen deficiency. Agronomy-basel, 2021; 11(9): 1859.
[19] Gao K, Chen F J, Yuan L X, Zhang F S, Mi G H. A comprehensive analysis of root morphological changes and nitrogen allocation in maize in response to low nitrogen stress. Plant Cell Environ, 2015; 38(4): 740–750.
[20] Chen G D, Wang L, Fabrice M R, Tian Y A, Qi K, Chen Q, et al. Physiological and nutritional responses of pear seedlings to nitrate concentrations. Front. Plant Sci, 2018; 9: 1679.
[21] Sung J, Lee S, Lee Y, Ha S, Song B, Kim T, et al. Metabolomic profiling from leaves and roots of tomato (Solanum lycopersicum L.) plants grown under nitrogen, phosphorus or potassium-deficient condition. Plant Sci, 2015; 241: 55–64.
[22] Chen Z P, Li H P, Yang T Y, Chen T T, Dong C X, Gu Q, et al. Transcriptome analysis provides insights into the molecular bases in response to different nitrogen forms-induced oxidative stress in tea plant roots (Camellia sinensis). Functional Plant Biology, 2020; 47(12): 1073–1082.
[23] Matic M, Vukovic R, Vrandecic K, Camagajevac I S, Cosic J, Vukovic A, et al. Oxidative status and antioxidative response to fusarium attack and different nitrogen levels in winter wheat varieties. Plants-basel, 2021; 10(4): 611.
[24] Zhu B, Xu Q W, Zou Y G, Ma S M, Zhang X D, Xie X Y, et al. Effect of potassium deficiency on growth, antioxidants, ionome and metabolism in rapeseed under drought stress. Plant Growth Regulation, 2020; 90(3): 455–466.
[25] Boussadia O, Stepp.K, Zgallai H, El Hadj S B, Braham M, Lemeur R, et al. Effects of nitrogen deficiency on leaf photosynthesis, carbohydrate status and biomass production in two olive cultivars ‘Meski’ and ‘Koroneiki’. Scientia Horticulturae, 2010; 123(3): 336–342.
[26] Sun T T, Zhang J K, Zhang Q, Li X L, Li M J, Yang Y Z, et al. Integrative physiological, transcriptome, and metabolome analysis reveals the effects of nitrogen sufficiency and deficiency conditions in apple leaves and roots. Environmental and Experimental Botany, 2021; 192: 104633.
Downloads
Published
How to Cite
Issue
Section
License
Copyright (c) 2026 International Journal of Agricultural and Biological Engineering

This work is licensed under a Creative Commons Attribution 4.0 International License.
IJABE is an international peer reviewed, open access journal, adopting Creative Commons Copyright Notices as follows.
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are permitted and encouraged to post their work online (e.g., in institutional repositories or on their website) prior to and during the submission process, as it can lead to productive exchanges, as well as earlier and greater citation of published work (See The Effect of Open Access).