Studies Referencing Molecular Hydrogen for use in Plants

Premise: The basic biological effects of hydrogen on animal cells are similar and translational on plant cells. See Review Articles.

Reported biological effects in plants

· Involved in phytohormone signaling (Zeng, Zhang, and Sun 2013; Liu, Li, and Liu 2016)

· Involved in stress response (Zeng, Zhang, and Sun 2013; Liu et al 2017)

· Involved in sugar accumulation and metabolism (Zhao et al 2021)

· Involved in antioxidant signaling and defense (Cao et al 2017; Su et al 2014; Ren et al 2017; Torres, Ballesteros, and Fernandez 1986)

· A 100% hydrogen atmosphere results in reversible metabolic arrest (Zerveas et al 2021) (i.e. to extend shelf life by using hydrogen in packaging)

Reported beneficial effects in plants

· Alleviates metal toxicity due to Cadmium (Wu et al 2015; Cui et al 2020; Dai et al 2017; Su et al 2019; Wang et al 2019; Wu et al 2019; Wu et al 2020; Zhang et al 2017; Wu et al 2021; Cui et al 2013; Wu et al 2015; Zhao et al 2021), Aluminum (Chen et al 2014; Zhao et al 2017; Xu et al 2017), Mercury (Cui et al 2014),

· Alleviates salt stress (Xu et al 2013; Zeng, Zhang, and Sun 2013; Su et al 2021; Qi et al 2020; Xie et al 2012; Zhang et al 2020; Fu et al 2020; Su et al 2018; Jin et al 2016; Felix et al 2018; Zhao et al 2021)

· Alleviates drought stress (Zeng, Zhang, and Sun 2013; Jin et al 2015; Xie et al 2014; Chen et al 2017; Zhao et al 2021)

· Alleviates pesticide toxicity (Zhang et al 2022; Jin et al 2013)

· Alleviates ultraviolet (UV) damage (Su et al 2014; Xie et al 2015; Zhang et al 2018; Zhang et al 2019; Zhang et al 2018)

· Alleviates chilling injury/cold tress (Xu et al 2017; Fengjiao et al 2020; Liu et al 2017)

· Alleviates heat stress (Chen et al 2017)

· Alleviates high-light induced injury (Zhang et al 2015)

· Improves rooting (Zhao et al 2021; Cao et al 2017; Li et al 2020; Lin et al 2014; Wang et al 2021; Wu et al 2020; Zhu et al 2016; Zhu and Liao 2017; Huang et al 2020; Zhu et al 2016; Liu et al 2021)

· Increased microbial metabolism/plant growth in H-treated soils (Liu et al 2010; Zhang, He, and Dong 2009; Irvine, Smith, and Dong 2002; Maimaiti et al 2006; Dong and Layzell 2001; Piche-Choquette and Constant 2019; Dong et al 2003)

Reported commercial effects in produce

· Prolonging vase life of cut flowers (Nguyen and Lim 2022; Li et al 2021; Huo et al 2018; Li et al 2020; Ren et al 2017; Su et al 2019; Wang et al 2020; Li et al 2021; Fang et al 2021)

· Prolonging shelf life in mushrooms (Chen et al 2017), chives (Jiang et al 2021), kiwifruit (Hu et al 2014; Hu et al 2018; Zhao et al 2021), daylily buds (Hu, Li, and Shen 2021), rice (Cai et al 2022), litchi (Yun et al 2021)

· Decreases nitrite (Zhang et al 2019) and nitrate (Wei et al 2021) in produce

· Increasing resistance of produce to diseases and pathogens (Hui et al 2017)

· Improved grain quality in rice (Cheng et al 2021)

· Increasing the quantity of medicinal compounds in plants (Zheng and Yu 2022)

______________________________________________________________________________________

Review Articles

Hancock JT. Editorial for Special Issue: "Production and Role of Molecular Hydrogen in Plants". Plants (Basel). 2022 Aug 5;11(15):2047. doi: 10.3390/plants11152047. PMID: 35956525; PMCID: PMC9370376.

Hancock JT, LeBaron TW, May J, Thomas A, Russell G. Molecular Hydrogen: Is This a Viable New Treatment for Plants in the UK? Plants (Basel). 2021 Oct 22;10(11):2270. doi: 10.3390/plants10112270. PMID: 34834633; PMCID: PMC8618766.

Hancock JT, Russell G. Downstream Signalling from Molecular Hydrogen. Plants (Basel). 2021 Feb 14;10(2):367. doi: 10.3390/plants10020367. PMID: 33672953; PMCID: PMC7918658.

Li C, Gong T, Bian B, Liao W. Roles of hydrogen gas in plants: a review. Funct Plant Biol. 2018 Jul;45(8):783-792. doi: 10.1071/FP17301. PMID: 32291062.

Li L, Lou W, Kong L, Shen W. Hydrogen Commonly Applicable from Medicine to Agriculture: From Molecular Mechanisms to the Field. Curr Pharm Des. 2021;27(5):747-759. doi: 10.2174/1381612826666201207220051. PMID: 33290194.

Li, L., Zeng, Y., Cheng, X., & Shen, W. (2021). The applications of molecular hydrogen in horticulture. Horticulturae, 7(11), 513.

Nguyen TK, Lim JH. Is It a Challenge to Use Molecular Hydrogen for Extending Flower Vase Life? Plants (Basel). 2022 May 10;11(10):1277. doi: 10.3390/plants11101277. PMID: 35631701; PMCID: PMC9146928.

Renwick GM, Giumarro C, Siegel SM. Hydrogen Metabolism in Higher Plants. Plant Physiol. 1964 May;39(3):303-6. doi: 10.1104/pp.39.3.303. PMID: 16655917; PMCID: PMC550076.

Russell G, Zulfiqar F, Hancock JT. Hydrogenases and the Role of Molecular Hydrogen in Plants. Plants (Basel). 2020 Sep 2;9(9):1136. doi: 10.3390/plants9091136. PMID: 32887396; PMCID: PMC7569912.

Wang YQ, Liu YH, Wang S, Du HM, Shen WB. Hydrogen agronomy: research progress and prospects. J Zhejiang Univ Sci B. 2020 Nov.;21(11):841-855. doi: 10.1631/jzus.B2000386. PMID: 33150769; PMCID: PMC7691689.

Zeng, J., Ye, Z. & Sun, X. Progress in the study of biological effects of hydrogen on higher plants and its promising application in agriculture. Med Gas Res 4, 15 (2014). https://doi.org/10.1186/2045-9912-4-15

Zulfiqar F, Russell G, Hancock JT. Molecular hydrogen in agriculture. Planta. 2021 Aug 21;254(3):56. doi: 10.1007/s00425-021-03706-0. PMID: 34420086.

Primary Articles

Cai C, Zhao Z, Zhang Y, Li M, Li L, Cheng P, Shen W. Molecular Hydrogen Improves Rice Storage Quality via Alleviating Lipid Deterioration and Maintaining Nutritional Values. Plants (Basel). 2022 Sep 30;11(19):2588. doi: 10.3390/plants11192588. PMID: 36235453; PMCID: PMC9571184.

Cao Z, Duan X, Yao P, Cui W, Cheng D, Zhang J, Jin Q, Chen J, Dai T, Shen W. Hydrogen Gas Is Involved in Auxin-Induced Lateral Root Formation by Modulating Nitric Oxide Synthesis. Int J Mol Sci. 2017 Oct 3;18(10):2084. doi: 10.3390/ijms18102084. PMID: 28972563; PMCID: PMC5666766.

Chen H, Zhang J, Hao H, Feng Z, Chen M, Wang H, Ye M. Hydrogen-rich water increases postharvest quality by enhancing antioxidant capacity in Hypsizygus marmoreus. AMB Express. 2017 Dec 20;7(1):221. doi: 10.1186/s13568-017-0496-9. PMID: 29264772; PMCID: PMC5738332.

Chen M, Cui W, Zhu K, Xie Y, Zhang C, Shen W. Hydrogen-rich water alleviates aluminum-induced inhibition of root elongation in alfalfa via decreasing nitric oxide production. J Hazard Mater. 2014 Feb 28;267:40-7. doi: 10.1016/j.jhazmat.2013.12.029. Epub 2013 Dec 27. PMID: 24413050.

Chen, Q., Zhao, X., Lei, D. et al. Hydrogen-rich water pretreatment alters photosynthetic gas exchange, chlorophyll fluorescence, and antioxidant activities in heat-stressed cucumber leaves. Plant Growth Regul 83, 69–82 (2017). https://doi.org/10.1007/s10725-017-0284-1

Chen Y, Wang M, Hu L, Liao W, Dawuda MM, Li C. Carbon Monoxide Is Involved in Hydrogen Gas-Induced Adventitious Root Development in Cucumber under Simulated Drought Stress. Front Plant Sci. 2017 Feb 7;8:128. doi: 10.3389/fpls.2017.00128. PMID: 28223992; PMCID: PMC5293791.

Cheng P, Wang J, Zhao Z, Kong L, Lou W, Zhang T, Jing D, Yu J, Shu Z, Huang L, Zhu W, Yang Q, Shen W. Molecular Hydrogen Increases Quantitative and Qualitative Traits of Rice Grain in Field Trials. Plants (Basel). 2021 Oct 28;10(11):2331. doi: 10.3390/plants10112331. PMID: 34834694; PMCID: PMC8624507.

Cui W, Gao C, Fang P, Lin G, Shen W. Alleviation of cadmium toxicity in Medicago sativa by hydrogen-rich water. J Hazard Mater. 2013 Sep 15;260:715-24. doi: 10.1016/j.jhazmat.2013.06.032. Epub 2013 Jun 21. PMID: 23846121.

Cui W, Fang P, Zhu K, Mao Y, Gao C, Xie Y, Wang J, Shen W. Hydrogen-rich water confers plant tolerance to mercury toxicity in alfalfa seedlings. Ecotoxicol Environ Saf. 2014 Jul;105:103-11. doi: 10.1016/j.ecoenv.2014.04.009. Epub 2014 May 7. PMID: 24793520.

Cui W, Yao P, Pan J, Dai C, Cao H, Chen Z, Zhang S, Xu S, Shen W. Transcriptome analysis reveals insight into molecular hydrogen-induced cadmium tolerance in alfalfa: the prominent role of sulfur and (homo)glutathione metabolism. BMC Plant Biol. 2020 Feb 4;20(1):58. doi: 10.1186/s12870-020-2272-2. PMID: 32019510; PMCID: PMC7001311.

Dai C, Cui W, Pan J, Xie Y, Wang J, Shen W. Proteomic analysis provides insights into the molecular bases of hydrogen gas-induced cadmium resistance in Medicago sativa. J Proteomics. 2017 Jan 30;152:109-120. doi: 10.1016/j.jprot.2016.10.013. Epub 2016 Oct 28. PMID: 27989938.

Dong, Z., Layzell, D. H2 oxidation, O2 uptake and CO2 fixation in hydrogen treated soils. Plant and Soil 229, 1–12 (2001). https://doi.org/10.1023/A:1004810017490

Dong, Z., Wu, L., Kettlewell, B., Caldwell, C. D., & Layzell, D. B. (2003). Hydrogen fertilization of soils–is this a benefit of legumes in rotation?. Plant, Cell & Environment, 26(11), 1875-1879.

Hua Fang, Chunlei Wang, Shuya Wang, Weibiao Liao, Hydrogen gas increases the vase life of cut rose ‘Movie star’ by regulating bacterial community in the stem ends, Postharvest Biology and Technology, Volume 181, 2021, 111685, ISSN 0925-5214, https://doi.org/10.1016/j.postharvbio.2021.111685.

Felix, K., Su, J., Lu, R. et al. Hydrogen-induced tolerance against osmotic stress in alfalfa seedlings involves ABA signaling. Plant Soil 445, 409–423 (2019). https://doi.org/10.1007/s11104-019-04328-y

Fengjiao, L., Xiaowei, Z., Fude, L., Jiang, Z., Huangai, B., & Xizhen, A.A. (2020). Effect of Exogenous Hydrogen on Photosynthetic Carbon Assimilation and Nitrogen Metabolism of Cucumber Seedlings Under Low Temperature.

Fu X, Ma L, Gui R, et al. Hydrogen rich water (HRW) induces plant growth and physiological attributes in fragrant rice varieties under salt stress. Research Square; 2020. DOI: 10.21203/rs.3.rs-21074/v1.

Hu H, Li P, Wang Y, Gu R. Hydrogen-rich water delays postharvest ripening and senescence of kiwifruit. Food Chem. 2014 Aug 1;156:100-9. doi: 10.1016/j.foodchem.2014.01.067. Epub 2014 Jan 30. PMID: 24629944.

Huali Hu, Pengxia Li, Wenbiao Shen, Preharvest application of hydrogen-rich water not only affects daylily bud yield but also contributes to the alleviation of bud browning, Scientia Horticulturae, Volume 287, 2021, 110267, ISSN 0304-4238, https://doi.org/10.1016/j.scienta.2021.110267.

Huali Hu, Suping Zhao, Pengxia Li, Wenbiao Shen, Hydrogen gas prolongs the shelf life of kiwifruit by decreasing ethylene biosynthesis, Postharvest Biology and Technology, Volume 135, 2018, Pages 123-130, ISSN 0925-5214, https://doi.org/10.1016/j.postharvbio.2017.09.008.

Huang D, Bian B, Zhang M, Wang C, Li C, Liao W. The role and proteomic analysis of ethylene in hydrogen gas-induced adventitious rooting development in cucumber (Cucumis sativus L.) explants. PeerJ. 2020 Apr 7;8:e8896. doi: 10.7717/peerj.8896. PMID: 32292654; PMCID: PMC7147439.

Lu, H., Wu, B., Wang, Y., Liu, N., Meng, F., Hu, Z., ... & Zhao, H. (2017). Effects of hydrogen-rich water treatment on defense responses of postharvest tomato fruit to Botrytis cinerea. Journal of Henan Agricultural Sciences, 46(2), 64-68.

Huo J, Huang D, Zhang J, Fang H, Wang B, Wang C, Ma Z, Liao W. Comparative Proteomic Analysis during the Involvement of Nitric Oxide in Hydrogen Gas-Improved Postharvest Freshness in Cut Lilies. Int J Mol Sci. 2018 Dec 9;19(12):3955. doi: 10.3390/ijms19123955. PMID: 30544843; PMCID: PMC6320913.

Irvine, P., Smith, M., & Dong, Z. (2002, August). Hydrogen fertilizer: bacteria or fungi?. In XXVI International Horticultural Congress: Issues and Advances in Transplant Production and Stand Establishment Research 631 (pp. 239-242).

Jiang K, Kuang Y, Feng L, Liu Y, Wang S, Du H, Shen W. Molecular Hydrogen Maintains the Storage Quality of Chinese Chive through Improving Antioxidant Capacity. Plants (Basel). 2021 May 29;10(6):1095. doi: 10.3390/plants10061095. PMID: 34072565; PMCID: PMC8227461.

Jin, Q., Cui, W., Dai, C. et al. Involvement of hydrogen peroxide and heme oxygenase-1 in hydrogen gas-induced osmotic stress tolerance in alfalfa. Plant Growth Regul 80, 215–223 (2016). https://doi.org/10.1007/s10725-016-0159-x

Jin, Q., Zhu, K., Cui, W. et al. Hydrogen-Modulated Stomatal Sensitivity to Abscisic Acid and Drought Tolerance Via the Regulation of Apoplastic pH in Medicago sativa . J Plant Growth Regul 35, 565–573 (2016). https://doi.org/10.1007/s00344-015-9561-2

Jin Q, Zhu K, Cui W, Xie Y, Han B, Shen W. Hydrogen gas acts as a novel bioactive molecule in enhancing plant tolerance to paraquat-induced oxidative stress via the modulation of heme oxygenase-1 signalling system. Plant Cell Environ. 2013 May;36(5):956-69. doi: 10.1111/pce.12029. Epub 2012 Nov 19. PMID: 23094798.

Li C, Huang D, Wang C, Wang N, Yao Y, Li W, Liao W. NO is involved in H2-induced adventitious rooting in cucumber by regulating the expression and interaction of plasma membrane H+-ATPase and 14-3-3. Planta. 2020 Jun 29;252(1):9. doi: 10.1007/s00425-020-03416-z. PMID: 32602044.

Li L, Liu Y, Wang S, Zou J, Ding W, Shen W. Magnesium Hydride-Mediated Sustainable Hydrogen Supply Prolongs the Vase Life of Cut Carnation Flowers via Hydrogen Sulfide. Front Plant Sci. 2020 Dec 9;11:595376. doi: 10.3389/fpls.2020.595376. PMID: 33362825; PMCID: PMC7755932.

Ying Li, Longna Li, Shu Wang, Yuhao Liu, Jianxin Zou, Wenjiang Ding, Hongmei Du, Wenbiao Shen, Magnesium hydride acts as a convenient hydrogen supply to prolong the vase life of cut roses by modulating nitric oxide synthesis, Postharvest Biology and Technology, Volume 177, 2021, 111526, ISSN 0925-5214, https://doi.org/10.1016/j.postharvbio.2021.111526.

Li L, Yin Q, Zhang T, Cheng P, Xu S, Shen W. Hydrogen Nanobubble Water Delays Petal Senescence and Prolongs the Vase Life of Cut Carnation (Dianthus caryophyllus L.) Flowers. Plants (Basel). 2021 Aug 12;10(8):1662. doi: 10.3390/plants10081662. PMID: 34451707; PMCID: PMC8401707.

Lin Y, Zhang W, Qi F, Cui W, Xie Y, Shen W. Hydrogen-rich water regulates cucumber adventitious root development in a heme oxygenase-1/carbon monoxide-dependent manner. J Plant Physiol. 2014 Jan 15;171(2):1-8. doi: 10.1016/j.jplph.2013.08.009. Epub 2013 Nov 14. PMID: 24331413.

Liu, F., Cai, B., Sun, S., Bi, H., & Ai, X. (2017). Effect of hydrogen-rich water soaked cucumber seeds on cold tolerance and its physiological mechanism in cucumber seedlings. Scientia Agricultura Sinica, 50(5), 881-889.

Liu, F., Li, J. & Liu, Y. Molecular hydrogen can take part in phytohormone signal pathways in wild rice. Biol Plant 60, 311–319 (2016). https://doi.org/10.1007/s10535-016-0591-9

Liu F, Lou W, Wang J, Li Q, Shen W. Glutathione produced by γ-glutamyl cysteine synthetase acts downstream of hydrogen to positively influence lateral root branching. Plant Physiol Biochem. 2021 Oct;167:68-76. doi: 10.1016/j.plaphy.2021.07.034. Epub 2021 Jul 28. PMID: 34333372.

Liu, Fang & Jiang, Wanxiang & Han, Weijuan & li, Junsheng & Liu, Yongbo. (2017). Effects of Hydrogen-Rich Water on Fitness Parameters of Rice Plants. Agronomy Journal. 109. 10.2134/agronj2017.02.0109.

Liu, H., Wang, W., Cao, G., & Tang, M. (2010). Effect of hydrogen on microbial population and enzyme activity in Robinia pseudoacacia rhizosphere soil. Chin. J. Appl. Environ. Biol, 16, 515-518.

Maimaiti, J., Zhang, Y., Yang, J., Cen, Y. P., Layzell, D. B., Peoples, M., & Dong, Z. (2007). Isolation and characterization of hydrogen‐oxidizing bacteria induced following exposure of soil to hydrogen gas and their impact on plant growth. Environmental Microbiology, 9(2), 435-444.

Piché-Choquette S, Constant P. Molecular Hydrogen, a Neglected Key Driver of Soil Biogeochemical Processes. Appl Environ Microbiol. 2019 Mar 6;85(6):e02418-18. doi: 10.1128/AEM.02418-18. PMID: 30658976; PMCID: PMC6414374.

Qi Wu, Nana Su, Lana Shabala, Liping Huang, Min Yu, Sergey Shabala, Understanding the mechanistic basis of ameliorating effects of hydrogen rich water on salinity tolerance in barley, Environmental and Experimental Botany, Volume 177, 2020, 104136, ISSN 0098-8472, https://doi.org/10.1016/j.envexpbot.2020.104136.

Ren A, Liu R, Miao ZG, Zhang X, Cao PF, Chen TX, Li CY, Shi L, Jiang AL, Zhao MW. Hydrogen-rich water regulates effects of ROS balance on morphology, growth and secondary metabolism via glutathione peroxidase in Ganoderma lucidum. Environ Microbiol. 2017 Feb;19(2):566-583. doi: 10.1111/1462-2920.13498. Epub 2016 Sep 9. PMID: 27554678.

Ren, PJ., Jin, X., Liao, WB. et al. Effect of hydrogen-rich water on vase life and quality in cut lily and rose flowers. Hortic. Environ. Biotechnol. 58, 576–584 (2017). https://doi.org/10.1007/s13580-017-0043-2

Su J, Yang X, Shao Y, Chen Z, Shen W. Molecular hydrogen-induced salinity tolerance requires melatonin signalling in Arabidopsis thaliana. Plant Cell Environ. 2021 Feb;44(2):476-490. doi: 10.1111/pce.13926. Epub 2020 Nov 5. PMID: 33103784.

Jiuchang Su, Yang Nie, Gan Zhao, Dan Cheng, Ren Wang, Jun Chen, Shihai Zhang, Wenbiao Shen, Endogenous hydrogen gas delays petal senescence and extends the vase life of lisianthus cut flowers, Postharvest Biology and Technology, Volume 147, 2019, Pages 148-155, ISSN 0925-5214, https://doi.org/10.1016/j.postharvbio.2018.09.018.

Jiuchang Su, Yihua Zhang, Yang Nie, Dan Cheng, Ren Wang, Huali Hu, Jun Chen, Jiaofei Zhang, Yuanwei Du, Wenbiao Shen, Hydrogen-induced osmotic tolerance is associated with nitric oxide-mediated proline accumulation and reestablishment of redox balance in alfalfa seedlings, Environmental and Experimental Botany, Volume 147, 2018, Pages 249-260, ISSN 0098-8472, https://doi.org/10.1016/j.envexpbot.2017.12.022.

Su N, Wu Q, Chen H, Huang Y, Zhu Z, Chen Y, Cui J. Hydrogen gas alleviates toxic effects of cadmium in Brassica campestris seedlings through up-regulation of the antioxidant capacities: Possible involvement of nitric oxide. Environ Pollut. 2019 Aug;251:45-55. doi: 10.1016/j.envpol.2019.03.094. Epub 2019 Mar 25. PMID: 31071632.

Su N, Wu Q, Liu Y, Cai J, Shen W, Xia K, Cui J. Hydrogen-rich water reestablishes ROS homeostasis but exerts differential effects on anthocyanin synthesis in two varieties of radish sprouts under UV-A irradiation. J Agric Food Chem. 2014 Jul 9;62(27):6454-62. doi: 10.1021/jf5019593. Epub 2014 Jun 30. PMID: 24955879.

Torres V, Ballesteros A, Fernández VM. Expression of hydrogenase activity in barley (Hordeum vulgare L.) after anaerobic stress. Arch Biochem Biophys. 1986 Feb 15;245(1):174-8. doi: 10.1016/0003-9861(86)90202-x. PMID: 3511850.

Wang B, Bian B, Wang C, Li C, Fang H, Zhang J, Huang D, Huo J, Liao W. Hydrogen gas promotes the adventitious rooting in cucumber under cadmium stress. PLoS One. 2019 Feb 20;14(2):e0212639. doi: 10.1371/journal.pone.0212639. PMID: 30785953; PMCID: PMC6382157.

Wang C, Fang H, Gong T, Zhang J, Niu L, Huang D, Huo J, Liao W. Hydrogen gas alleviates postharvest senescence of cut rose 'Movie star' by antagonizing ethylene. Plant Mol Biol. 2020 Feb;102(3):271-285. doi: 10.1007/s11103-019-00946-3. Epub 2019 Dec 14. PMID: 31838617.

Yueqiao Wang, Peixun Lv, Lingshuai Kong, Wenbiao Shen, Qianjun He, Nanomaterial-mediated sustainable hydrogen supply induces lateral root formation via nitrate reductase-dependent nitric oxide, Chemical Engineering Journal, Volume 405, 2021, 126905, ISSN 1385-8947, https://doi.org/10.1016/j.cej.2020.126905.

Wei, X., Chen, J., Chen, H. et al. Hydrogen-rich water ameliorates the toxicity induced by Ca(NO3)2 excess through enhancing antioxidant capacities and re-establishing nitrate homeostasis in Brassica campestris spp. chinensis L. seedlings. Acta Physiol Plant 43, 50 (2021). https://doi.org/10.1007/s11738-021-03220-6

Wu Q, Huang L, Su N, Shabala L, Wang H, Huang X, Wen R, Yu M, Cui J, Shabala S. Calcium-Dependent Hydrogen Peroxide Mediates Hydrogen-Rich Water-Reduced Cadmium Uptake in Plant Roots. Plant Physiol. 2020 Jul;183(3):1331-1344. doi: 10.1104/pp.20.00377. Epub 2020 May 4. PMID: 32366640; PMCID: PMC7333692.

Wu Q, Su N, Cai J, Shen Z, Cui J. Hydrogen-rich water enhances cadmium tolerance in Chinese cabbage by reducing cadmium uptake and increasing antioxidant capacities. J Plant Physiol. 2015 Mar 1;175:174-82. doi: 10.1016/j.jplph.2014.09.017. Epub 2014 Nov 18. PMID: 25543863.

Wu, Q., Su, N., Chen, Q., Shen, W., Shen, Z., Xia, Y., & Cui, J. (2015). Cadmium-induced hydrogen accumulation is involved in cadmium tolerance in Brassica campestris by reestablishment of reduced glutathione homeostasis. PLoS One, 10(10), e0139956.

Wu Q, Su N, Huang X, Ling X, Yu M, Cui J, Shabala S. Hydrogen-rich water promotes elongation of hypocotyls and roots in plants through mediating the level of endogenous gibberellin and auxin. Funct Plant Biol. 2020 Aug;47(9):771-778. doi: 10.1071/FP19107. PMID: 32522330.

Wu X, Su N, Yue X, Fang B, Zou J, Chen Y, Shen Z, Cui J. IRT1 and ZIP2 were involved in exogenous hydrogen-rich water-reduced cadmium accumulation in Brassica chinensis and Arabidopsis thaliana. J Hazard Mater. 2021 Apr 5;407:124599. doi: 10.1016/j.jhazmat.2020.124599. Epub 2020 Dec 13. PMID: 33360184.

Wu X, Zhu ZB, Chen JH, Huang YF, Liu ZL, Zou JW, Chen YH, Su NN, Cui J. Transcriptome analysis revealed pivotal transporters involved in the reduction of cadmium accumulation in pak choi (Brassica chinensis L.) by exogenous hydrogen-rich water. Chemosphere. 2019 Feb;216:684-697. doi: 10.1016/j.chemosphere.2018.10.152. Epub 2018 Oct 25. PMID: 30391890.

Xie Y, Mao Y, Zhang W, Lai D, Wang Q, Shen W. Reactive Oxygen Species-Dependent Nitric Oxide Production Contributes to Hydrogen-Promoted Stomatal Closure in Arabidopsis. Plant Physiol. 2014 Jun;165(2):759-773. doi: 10.1104/pp.114.237925. Epub 2014 Apr 14. PMID: 24733882; PMCID: PMC4044830.

Xie Y, Mao Y, Lai D, Zhang W, Shen W. H(2) enhances arabidopsis salt tolerance by manipulating ZAT10/12-mediated antioxidant defence and controlling sodium exclusion. PLoS One. 2012;7(11):e49800. doi: 10.1371/journal.pone.0049800. Epub 2012 Nov 21. PMID: 23185443; PMCID: PMC3504229.

Xie Y, Zhang W, Duan X, Dai C, Zhang Y, Cui W, Wang R, Shen W. Hydrogen-rich water-alleviated ultraviolet-B-triggered oxidative damage is partially associated with the manipulation of the metabolism of (iso)flavonoids and antioxidant defence in Medicago sativa. Funct Plant Biol. 2015 Dec;42(12):1141-1157. doi: 10.1071/FP15204. PMID: 32480752.

Xu D, Cao H, Fang W, Pan J, Chen J, Zhang J, Shen W. Linking hydrogen-enhanced rice aluminum tolerance with the reestablishment of GA/ABA balance and miRNA-modulated gene expression: A case study on germination. Ecotoxicol Environ Saf. 2017 Nov;145:303-312. doi: 10.1016/j.ecoenv.2017.07.055. Epub 2017 Jul 27. PMID: 28756251.

Xu, S., Jiang, Y., Cui, W. et al. Hydrogen enhances adaptation of rice seedlings to cold stress via the reestablishment of redox homeostasis mediated by miRNA expression. Plant Soil 414, 53–67 (2017). https://doi.org/10.1007/s11104-016-3106-8

Xu, S., Zhu, S., Jiang, Y. et al. Hydrogen-rich water alleviates salt stress in rice during seed germination. Plant Soil 370, 47–57 (2013). https://doi.org/10.1007/s11104-013-1614-3

Yun Z, Gao H, Chen X, Chen Z, Zhang Z, Li T, Qu H, Jiang Y. Effects of hydrogen water treatment on antioxidant system of litchi fruit during the pericarp browning. Food Chem. 2021 Jan 30;336:127618. doi: 10.1016/j.foodchem.2020.127618. Epub 2020 Jul 25. PMID: 32771896.

Zeng J, Yu H. Integrated Metabolomic and Transcriptomic Analyses to Understand the Effects of Hydrogen Water on the Roots of Ficus hirta Vahl. Plants (Basel). 2022 Feb 24;11(5):602. doi: 10.3390/plants11050602. PMID: 35270073; PMCID: PMC8912395.

Zeng J, Zhang M, Sun X. Molecular hydrogen is involved in phytohormone signaling and stress responses in plants. PLoS One. 2013 Aug 12;8(8):e71038. doi: 10.1371/journal.pone.0071038. PMID: 23951075; PMCID: PMC3741361.

Zerveas S, Kydonakis E, Mente MS, Daskalakis V, Kotzabasis K. Hydrogen gas as a central on-off functional switch of reversible metabolic arrest - New perspectives for biotechnological applications. J Biotechnol. 2021 Jul 20;335:9-18. doi: 10.1016/j.jbiotec.2021.06.005. Epub 2021 Jun 4. PMID: 34090950.

Zhang J, Hao H, Chen M, Wang H, Feng Z, Chen H. Hydrogen-rich water alleviates the toxicities of different stresses to mycelial growth in Hypsizygus marmoreus. AMB Express. 2017 Dec;7(1):107. doi: 10.1186/s13568-017-0406-1. Epub 2017 May 30. PMID: 28565883; PMCID: PMC5449350.

Zhang Y, Cheng P, Wang Y, Li Y, Su J, Chen Z, Yu X, Shen W. Genetic elucidation of hydrogen signaling in plant osmotic tolerance and stomatal closure via hydrogen sulfide. Free Radic Biol Med. 2020 Dec;161:1-14. doi: 10.1016/j.freeradbiomed.2020.09.021. Epub 2020 Sep 25. PMID: 32987125.

Zhang T, Wang Y, Zhao Z, Xu S, Shen W. Degradation of Carbendazim by Molecular Hydrogen on Leaf Models. Plants (Basel). 2022 Feb 25;11(5):621. doi: 10.3390/plants11050621. PMID: 35270091; PMCID: PMC8912477.

Zhang X, Wei J, Huang Y, Shen W, Chen X, Lu C, Su N, Cui J. Increased Cytosolic Calcium Contributes to Hydrogen-Rich Water-Promoted Anthocyanin Biosynthesis Under UV-A Irradiation in Radish Sprouts Hypocotyls. Front Plant Sci. 2018 Jul 16;9:1020. doi: 10.3389/fpls.2018.01020. PMID: 30061912; PMCID: PMC6055044.

Xiaoyan Zhang, Junyu Wei, Jiyuan Tian, Na Li, Li Jia, Wenbiao Shen, Jin Cui, Enhanced anthocyanin accumulation of immature radish microgreens by hydrogen-rich water under short wavelength light, Scientia Horticulturae, Volume 247, 2019, Pages 75-85, ISSN 0304-4238, https://doi.org/10.1016/j.scienta.2018.11.060.

Zhang, X., Zhao, X., Wang, Z. et al. Protective effects of hydrogen-rich water on the photosynthetic apparatus of maize seedlings (Zea mays L.) as a result of an increase in antioxidant enzyme activities under high light stress. Plant Growth Regul 77, 43–56 (2015). https://doi.org/10.1007/s10725-015-0033-2

Zhang, X., Su, N., Jia, L. et al. Transcriptome analysis of radish sprouts hypocotyls reveals the regulatory role of hydrogen-rich water in anthocyanin biosynthesis under UV-A. BMC Plant Biol 18, 227 (2018). https://doi.org/10.1186/s12870-018-1449-4

Zhang, Y., He, X. & Dong, Z. Effect of hydrogen on soil bacterial community structure in two soils as determined by terminal restriction fragment length polymorphism. Plant Soil 320, 295–305 (2009). https://doi.org/10.1007/s11104-009-9894-3

Zhang, Yihua & Zhao, Gan & Pengfei, Cheng & Yan, Xinyu & Li, Ying & Cheng, Dan & Wang, Ren & Chen, Jun & Shen, Wenbiao. (2019). Nitrite accumulation during storage of tomato fruit as prevented by hydrogen gas. International Journal of Food Properties. 22. 1425-1438. 10.1080/10942912.2019.1651737.

Zhao G, Cheng P, Zhang T, Abdalmegeed D, Xu S, Shen W. Hydrogen-rich water prepared by ammonia borane can enhance rapeseed (Brassica napus L.) seedlings tolerance against salinity, drought or cadmium. Ecotoxicol Environ Saf. 2021 Aug 13;224:112640. doi: 10.1016/j.ecoenv.2021.112640. Epub ahead of print. PMID: 34392154.

Zhao X, Chen Q, Wang Y, Shen Z, Shen W, Xu X. Hydrogen-rich water induces aluminum tolerance in maize seedlings by enhancing antioxidant capacities and nutrient homeostasis. Ecotoxicol Environ Saf. 2017 Oct;144:369-379. doi: 10.1016/j.ecoenv.2017.06.045. Epub 2017 Jun 22. PMID: 28647604.

Zhao, X., Meng, X., Li, W. et al. Effect of hydrogen-rich water and slightly acidic electrolyzed water treatments on storage and preservation of fresh-cut kiwifruit. Food Measure 15, 5203–5210 (2021). https://doi.org/10.1007/s11694-021-01000-x

Zhao Z, Li C, Liu H, Yang J, Huang P, Liao W. The Involvement of Glucose in Hydrogen Gas-Medicated Adventitious Rooting in Cucumber. Plants (Basel). 2021 Sep 17;10(9):1937. doi: 10.3390/plants10091937. PMID: 34579469; PMCID: PMC8469787.

Zhu, Y., Liao, W. The metabolic constituent and rooting-related enzymes responses of marigold explants to hydrogen gas during adventitious root development. Theor. Exp. Plant Physiol. 29, 77–85 (2017). https://doi.org/10.1007/s40626-017-0085-y

Zhu Y, Liao W, Niu L, Wang M, Ma Z. Nitric oxide is involved in hydrogen gas-induced cell cycle activation during adventitious root formation in cucumber. BMC Plant Biol. 2016 Jun 28;16(1):146. doi: 10.1186/s12870-016-0834-0. PMID: 27352869; PMCID: PMC4924243.

Zhu Y, Liao W, Wang M, Niu L, Xu Q, Jin X. Nitric oxide is required for hydrogen gas-induced adventitious root formation in cucumber. J Plant Physiol. 2016 May 20;195:50-8. doi: 10.1016/j.jplph.2016.02.018. Epub 2016 Mar 15. PMID: 27010347.