Insights into the role of cytokinin and gibberellic acid in improving waterlogging tolerance of mung bean

Insights into the role of cytokinin and gibberellic acid in improving waterlogging tolerance of mung bean | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Insights into the role of cytokinin and gibberellic acid in improving waterlogging tolerance of mung bean M Rafiqul Islam, Md. Mezanur Rahman, Munny Akter, Erin Zama, Sanjida Sultana Keya, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-690868/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Mung bean ( Vigna radiata ) is one of the most important pulse crops, well-known for its protein rich seeds, which growth and productivity are severely undermined by waterlogging. In this study, we aim to evaluate how two promising phytohormones, namely cytokinin (CK) and gibberellic acid (GA 3 ), can improve waterlogging tolerance in mung bean by investigating key morphological, physiological, biochemical and yield-related attributes. Our results showed that foliar application of CK and GA 3 under 5-days of waterlogged conditions improved mung bean growth and biomass, which was associated to increased levels of photosynthetic rate and pigments. Waterlogged-induced accumulation of reactive oxygen species, and the consequent elevated levels of malondialdehyde, were considerably reduced by CK and GA 3 treatments. Mung bean plants sprayed with either CK or GA 3 suffered less oxidative stress due to the enhancement of total phenolics and flavonoids levels. Improvement in the contents of proline and total soluble sugars indicating a better osmotic adjustment following CK and GA 3 treatments in waterlogged-exposed plants. Most fundamentally, CK or GA 3 -sprayed waterlogged-stressed mung bean plants demonstrated an increased tendency of the above-mentioned parameters after the 15-day recovery period as compared to water-sprayed waterlogged-exposed plants. Our results also revealed that CK and GA 3 treatments increased yield-associated features in waterlogged-stressed plant. Importantly, both phytohormones are efficient in improving mung bean resistance to waterlogging; however, CK was found to be more effective. Overall, our findings suggested that CK or GA 3 could be used for the management of waterlogging-induced damage in mung bean, and perhaps in other cash crops. Agronomy Antioxidants osmoprotectants phytohormone pulse crop reactive oxygen species yield Figures Figure 1 Figure 2 Figure 3 1. Introduction Waterlogging has become a major environmental problem in global agriculture, arises from heavy rainfall, impromptu irrigation, inadequate drainage systems, unleveled fields, and soil compaction (Kaur et al., 2020 ). Under waterlogged conditions, the mass of oxygen in the soil attenuates partially (hypoxic) or fully (anoxic) due to high microbial activity, and the resulting carbon dioxide accumulation in the root zone restricts root metabolism, aerobic respiration, ATP synthesis, and nutrient acquisition, resulting in a significant reduction in the growth, development and biomass of roots and shoots (Najeeb et al., 2015 ; Gill et al., 2018 ; Dossa et al., 2019 ). Moreover, waterlogging-induced excessive oxygen scarcity in water impairs root permeability and causes root injury, leading to a decrement of hydraulic conductivity and consequent stomatal closure, resulting in a significant reduction in net photosynthetic and transpiration rates (Else et al., 2001 ; Anee et al., 2019 ). While photosynthesis is disrupted in waterlogged plants, lethal levels of reactive oxygen species (ROS) such as superoxide (O 2 •− ), singlet oxygen ( 1 O 2 ), hydrogen peroxide (H 2 O 2 ) and hydroxyl radical ( • OH) are produced, which are harmful to cellular components and cause DNA and RNA damage, lipid peroxidation, protein oxidation, and enzyme inactivation (Yeung et al., 2018 ; Zhou et al., 2020 ). Plants can acclimatise the deletorious consequence of waterlogging-mediated soil oxygen dearth through adapting various morphological, physiological and biochemical mechanisms. Increased adventitious root development and faster stem elongation are some examples of morphological adaptations (Yamauchi et al., 2018 ), whereas reduced stomatal conductance and subsequent decrease in net photosynthetic rate are some short-term physiological adaptation process (Jacobsen et al., 2007 ; Bhusal et al., 2020 ). Additionally, to shield themselves from stress, waterlogged plants accumulate a number of osmoprotectants such as proline (Pro), soluble sugars, and sucrose (Tewari and Mishra, 2018 ). However, the most important adaptive mechanism is a well-balanced antioxidant defense system that involves both enzymatic (e.g., superoxide dismutase, SOD; catalase, CAT; ascorbate peroxidase, APX; glutathione S -transferase, GST; glutathione peroxidase, GPX) and non-enzymatic (e.g., total phenolics, total flavonoids, carotenoids) antioxidants that scavenge overaccumulation of ROS (Doupis et al., 2017 ; Fukao et al., 2019 ; Garcia et al., 2020 ). Mung bean ( Vigna radiata ) is the third most important pulse crop after grass pea ( Lathyrus sativus ) and lentil ( Lens culinaris ) and considered as a major source of pulse protein because of its high protein content in the seeds (20.97–31.32%) (AESA, 2017; Yi-Shen et al., 2018 ; Rahman et al., 2019 ). Additionally, mung bean has been well-documented for its detoxification bioactivities along with its use as a treating agent for various health conditions, including stimulation of human mental function, allaying of heat stroke and regulation of gastrointestinal upset (Yi-Shen et al., 2018 ; Hou et al., 2019 ). More fundamentally, mung bean gaining its popularity in recent years and extensively cultivated throughout Bangladesh as a cash crop in the extensive rice-based cropping system, owing to its low fertilizer and pesticide requirements, which provides farmers with good economic benefits and nutritional security (Rahman et al., 2019 ). According to the estimation in the years 2016-17, cultivation of mung bean covered approximately 0.3188 million hectares (Mha) out of 0.9976 Mha of total pulse crop areas and yielded 0.21 million metric tonnes of mung bean (DAE. Krishi Dairy, 2018). However, recently the emerging waterlogging problem in major mung bean growing areas like Barishal, Patuakhali, Mymensingh, Sylhet, Chittagong hill tract, Natore, Khulna, Brahmanbaria and Pabna region make the land untenable for growing mung bean ( http://www.ffwc.gov.bd/index.php/map/inundation-map/bangladesh-today ). Therefore, the country must devise a strategy for effectively alleviating the effects of waterlogging stress in order to improve mung bean production in flash flood-prone areas. In this context, the search of a simple and cost-effective technology propelled us to use phytohormone such as cytokinin (CK) and gibberellic acid (GA 3 ) for overcoming the deleterious effects of waterlogging on the growth and yield-related attributes of mung bean. In the present study, we first examined whether CK and GA 3 provides protection to mung bean from waterlogging stress, and to our expectations it really did. Next, we also examined the regulatory roles of CK and GA 3 in improving mung bean tolerance to waterlogging by investigating the morpho-physiological and biochemical mechanisms through assessments of the following key attributes: (i) plant growth performance and biomass production, (ii) gas-exchange features, (iii) photosynthetic pigment status, (iv) excessive waterlogging-mediated oxidative damage in terms of elevated ROS levels and lipid peroxidation, (v) different types of osmolyte accumulations, and (vi) improvement of non-enzymatic antioxidant defense. Additionally, we also record the yield-associated features of waterlogged-stressed and waterlogged-devoid mung bean plants. 2. Materials And Methods 2.1. Plant growth conditions and treatments imposition Mung bean ( Vigna radiata ) genotype, VC6173-B, was used in the present study owing to their properties of short duration (80 days to maturity), greater grain weight (44g/1000 grain) and high yield potential (1.5 tons ha − 1 ). Healthy seeds of VC6173-B were germinated following the procedures of Rahman et al. ( 2019 ). Afterward, six germinated seeds were sown in each plastic pot (height × diameter = 20 cm × 16 cm) having 8 kg of soil per pot. The soil used was prepared by intermixing of cow dung, sand and soil at the ratio of 1:1:3 (in a weight basis). Additionally, the soil of the pots was also fertilized with urea, triple super phosphate (TSP) and muriate of potash (MP) at the rate of 0.176, 0.400 and 0.347 g corresponding to 40-60-40 Kg N, P 2 O 5 and K 2 O per hectare (Islam et al., 2008 ). It is worth noting that one-third of urea and the whole amount of TSP, MP and gypsum were mixed with the soil as basal dose during final pot preparation. The remaining amount of urea was applied at 10 and 25 days after emergence. In order to protect the seedlings from insect-induced damage, Ripcord 10 EC (BASE Bangladesh Limited) at 1 mL L −1 water was sprayed at vegetative stage of plant growth. The average maximum and minimum temperature of the experimental area were lies in the range of 32.6 to 22.8°C, with 85.6% of relative humidity. Notably, the number of seedlings was thinned to two in each pot after eighth day of germination and continues to grow in normal condition up to the imposition of treatments. Prior to waterlogging exposure for the next five days, fifteen-day-old mung bean seedlings at vegetative V1 stage (when first trifoliate were fully developed) were grouped into four sets as follows: water-sprayed waterlogged-stress-free plants (Control) water-sprayed waterlogged-stressed plants (WL) cytokinin (CK)-sprayed (50 mg L −1 ) waterlogged-stressed plants (CK + WL) gibberellic acid (GA 3 )-sprayed (50 mg L −1 ) waterlogged-stressed plants (GA 3 + WL) It is worth noting that foliar spray with either CK or GA 3 or water (20 mL to each pot) was done for two times in a day (9.00 AM to 10.00 AM, and 3.00 PM to 4.00 PM). Tween-20 surfactant (0.2%, v/v) was used to ensure maximum adherence of CK, GA 3 and water to the leaves. Notably, the level of water for creating waterlogging stress was maintained at 2.5 cm above the soil surface. Following the stress period, one set of seedlings (Set I) were immediately harvested, and another set (Set II) was harvested 15-days after recovery, to record the morphological, physiological and biochemical parameters. Importantly, after stress exposure, one set of seedlings (Set III) was allowed to grow until harvest with normal irrigation to determine yield-related attributes, including pod length, total number of pods per plant, thousand seed weight, and seed yield per plant. Each set of experiments were repeated four times under the same experimental conditions. Importantly, for determining various morphological, physiological and biochemical parameters, the first trifoliate leaves from the bottom of the plants were collected. 2.2. Determination of growth-related parameters The growth performance of mung bean plants was assessed by measuring the shoot height, and fresh weight (FW) of both shoots and roots. Additionally, stem girth of the mung bean plants was determined using a slide caliper. 2.3. Measurement of gas exchange features Gas exchange features, including photosynthesis rate ( Pn ), stomatal conductance to water (H 2 O) ( g s ) and transpiration rate ( E ) of mung bean plants were measured using a LI-6400XT portable photosynthesis system (LI-COR Biosciences, Lincoln, Nebraska, USA) from 11.00 AM to 2.00 PM under a full sun-light condition. The parameters Pn , g s and E were used for assessing the instantaneous water-use efficiency (WUEins; ratio Pn / E ) and intrinsic water-use efficiency (WUEint; ratio Pn / g s ). 2.4. Quantification of photosynthetic pigments, and total phenolic and flavonoid contents in mung bean leaves The freshly harvested mung bean leaves were used to determine the contents of chlorophylls (Chls) [Chl a , Chl b and Chl ( a + b )] by using the visible spectrophotometer (Model: T60 UV, PG Instruments Limited, Leicestershire, UK) according to the protocols described by Lichtenthaler and Wellburn ( 1983 ). The contents of total phenolic and total flavonoid were quantified following the methods of Ainsworth and Gillespie ( 2007 ), and Zhishen et al. ( 1999 ), respectively. 2.5. Quantification of H 2 O 2 , MDA, proline and total soluble sugars Freshly harvested leaf tissues were used for the quantification of hydrogen peroxide (H 2 O 2 ), malondialdehyde (MDA) (a lipid peroxidation product), proline (Pro) and total soluble sugars levels following the comprehensive protocols described by Yu et al. ( 2003 ), Health and Packer (1968), Bates et al. ( 1973 ) and Somogyi ( 1952 ), respectively. 2.6. Statistical analysis The obtained data were subjected to a one-way analysis of variance (ANOVA) using Statistix software (version 10). Different alphabetical letters were used to denote the significant variations among different treatments at the P < 0.05 level following a least significant difference (LSD) test. All the numerical data in figures and tables are presented as means ± standard errors (SEs) of four independent replications. 3. Results 3.1. CK and GA 3 supplementation improved morphological features of mung bean plants under waterlogging stress and recovery period In comparison with ‘Control’ plants, ‘WL’ plants displayed significant decrease in shoot height, shoot FW, root FW and stem girth width by 33.46, 37.04, 41.50 and 37.07%, respectively; however, following 15-days of recovery, these parameters were reduced by 35.31, 44.05, 35.88 and 32.18%, respectively (Table 1). On the other hand, ‘CK+WL’ and ‘GA 3 +WL’ plants showed noteworthy improvement in shoot height by 35.46 and 25.54%, shoot FW by 37.93 and 26.87%, root FW by 36.05 and 25.58%, and stem girth width by 36.06 and 30.61%, respectively, when compared with ‘WL’ plants (Table 1). Following the recovery period, a substantial enhancement in shoot height (34.70 and 23.90%), shoot FW (37.93 and 26.87%), root FW (36.05 and 25.58%) and stem girth width (36.06 and 30.61%, respectively) was observed in the ‘CK+WL’ and ‘GA 3 +WL’ plants, in comparison with ‘WL’ plants (Table 1). It was also observed that ‘CK+WL’ and ‘GA 3 +WL’ plants exhibited notable decrease in shoot height by 9.86 and 16.46%, shoot FW by 13.17 and 20.13%, root FW by 20.41 and 26.53% and stem girth width by 14.40 and 17.83%, respectively, relative to that of ‘Control’ plants (Table 1). Similarly, in contrast to ‘Control’ plants, significant reductions in shoot height (by 12.86 and 19.85%), shoot FW (24.21 and 29.94%), root FW (20.58% for GA 3 +WL) and stem girth width (8.99 and 15.86%, respectively) were observed following recovery period in the ‘CK+WL’ and ‘GA 3 +WL’ plants (Table 1). 3.2. CK and GA 3 supplementation improved gas exchange features of mung bean plants under waterlogging stress and recovery period Mung bean plants subjected to waterlogging stress and followed by stress recovery period showed a decrease in Pn (by 44.12 and 41.97%), gs (90.28 and 53.25%) and E (73.92 and 35.56%, respectively) when compared with ‘Control’ plants (Fig. 1A-C). Nonetheless, the levels of WUEint and WUEins in ‘WL’ plants increased by 514.96 and 117.90%, respectively relative to ‘Control’ plants, whereas the level of WUEint and WUEins in ‘Control’ and ‘WL’ plants was comparable after recovery period (Fig. 1D,E). On the other hand, ‘CK+WL’ and ‘GA 3 +WL’ plants exhibited decreased levels of gs (by 40.03 and 27.36%) and E (32.87 and 22.55%), and increased levels of Pn (by 36.80 and 26.65%), WUEint (119.44 and 63.82%) and WUEins (100.06 and 61.36%, respectively) when compared with that of ‘WL’ plants (Fig. 1A-E). Similarly, following the recovery period, decreased levels of gs (by 35.60 and 27.61%) and E (34.14 and 21.37%), and increased levels of Pn (36.80 and 26.65%), WUEint (114.55 and 72.68%) and WUEins (107.42 and 59.09%, respectively) were recorded in ‘CK+WL’ and ‘GA 3 +WL’ plants, as compared with that of ‘WL’ plants (Fig. 1A-E). In relation to the ‘Control’, a sharp reduction in the levels of Pn (by 23.56 and 29.23%), gs (94.17 and 92.94%) and E (82.49 and 79.80%) was noticed in ‘CK+WL’ and ‘GA 3 +WL’ plants, respectively. Similarly, the levels of Pn , gs and E were noticeably reduced by 20.85, 69.89 and 57.56% in ‘CK+WL’, and by 27.28, 66.15 and 49.33% in ‘GA 3 +WL’ plants followed by the recovery period over the corresponding values of ‘Control’ plants (Fig. 1A-C). Additionally, in relation to the ‘Control’, the level of WUEint and WUEins was substantially enhanced by 1249.50 and 335.94% in ‘CK+WL’ and by 907.44 and 251.60% in ‘GA 3 +WL’ plants followed by exposure to waterlogging stress. Whereas after recovery period, augmented level of WUEint (by 165.23 and 113.47%) and WUEins (87.07 and 43.47%) was observed in ‘CK+WL’ and ‘GA 3 +WL’ plants, respectively (Fig. 1D,E). 3.3. CK and GA 3 supplementation improved photosynthetic pigments status of mung bean plants under waterlogging stress and recovery period In relation to ‘Control’ plants, a sharp decline in the contents of Chl a (by 50.10 and 43.29%), Chl b (65.44 and 62.99%) and Chl ( a + b ) (54.35 and 48.96%) were observed in the leaves of ‘WL’ plants following stress and recovery periods, respectively (Fig. 2A-C). In contrast, CK and GA 3 supplementation protected the photosynthetic pigments from waterlogged-induced deleterious effects by enhancing the contents of Chl a (by 74.01 and 60.52%), Chl b (142.38 and 124.40%) and Chl ( a + b ) (88.37 and 73.94%, respectively) in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants, respectively (Fig. 2A-C). Furthermore, following the recovery period, the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants also displayed a significant rise in the content of Chl a (by 59.75 and 49.54%), Chl b (134.31 and 120.36%) and Chl ( a + b ) (75.32 and 64.33%, respectively) in comparison with ‘WL’ plants (Fig. 2A-C). On the other hand, CK and GA 3 supplementation resulted in a decrement of Chl a (by 13.16 and 19.90%), Chl b (16.23 and 22.44%) and Chl ( a + b ) (14.01 and 20.60%) in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants, respectively, when compared with ‘Control’ plants (Fig. 2A-C). Likewise, following the recovery period, a noteworthy decrement in the contents of Chl a (by 9.40 and 15.19 %), Chl b (13.28 and 18.44%) and Chl ( a + b ) (10.51 and 16.13%) were observed in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants, respectively, in comparison with ‘Control’ plants (Fig. 2A-C). 3.4. CK and GA 3 supplementation suppressed the levels of oxidative stress markers of mung bean plants under waterlogging stress and recovery period Imposition of waterlogging stress and following recovery period substantially increased the contents of H 2 O 2 (by 113.40 and 62.08%) and MDA (178.67 and 100.18%, respectively) in the leaves of ‘WL’ plants, relative to that of ‘Control’ plants (Fig. 3A,B). Fascinatingly, CK and GA 3 treatment resulted in reductions of H 2 O 2 (by 57.31 and 45.40%) and MDA (29.01 and 19.04%, respectively) contents in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants, in comparison with ‘WL’ plants (Fig. 3A,B). Similarly, compared with the ‘WL’ plants, substantial decreases in the contents of H 2 O 2 (by 36.61 and 26.44 %) and MDA (33.79 and 20.70%, respectively) were noticed in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants followed by the recovery period (Fig. 3A,B). On the other hand, ‘CK+WL’ and ‘GA 3 +WL’ plants exhibited noteworthy improvements in the levels of H 2 O 2 (by 38.52 and 59.69%) and MDA (71.34 and 102.46%, respectively), when contrasted with that of ‘Control’ plants (Fig. 3A,B). After the recovery period, the contents of H 2 O 2 (by 2.75 and 19.22%) and MDA (32.53 and 58.74%, respectively) in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants were substantially higher compared with the respective value of ‘Control’ plants (Fig. 3A,B). 3.5. CK and GA 3 supplementation improved the levels of Proline (Pro), total soluble sugars, total phenolics and flavonoids of mung bean plants under waterlogging stress and recovery period Exposure of mung bean plants to waterlogging stress and followed by a recovery period exhibited a rise in the contents of Pro (63.01 and 42.17%), total soluble sugars (33.20 and 20.29%), total phenolics (57.65 and 23.16%) and total flavonoids (30.98 and 15.89%, respectively) in the leaves of ‘WL’ plants, when compared with that of ‘Control’ plants (Fig. 3C-F). In comparison with ‘WL’ plants, ‘CK+WL’ and ‘GA 3 +WL’ plants displayed enhanced levels of Pro (by 33.75 and 21.01%), total soluble sugars (32.24 and 21.79%), total phenolics (36.48 and 24.53%) and total flavonoids (37.34 and 28.19%, respectively) (Fig. 3C-F). Similarly, in relation to the ‘WL’ plants, the levels of Pro, total soluble sugars, total phenolics and flavonoids were noticeably enhanced in ‘CK+WL’ leaves by 34.61, 35.02, 34.59 and 35.04%, respectively, and in ‘GA 3 +WL’ leaves by 23.45, 27.21, 23.45 and 25.08%, respectively, followed by a recovery period (Fig. 3C-F). Significant improvement in the contents of Pro (by 118.02 and 97.25%), total soluble sugars (76.15 and 62.23%), total phenolics (115.16 and 96.32%) and total flavonoids (79.90 and 67.91%, respectively) were recorded in the leaves of ‘CK+WL’ and ‘GA 3 +WL’ plants, when contrasted with the corresponding ‘Control’ values (Fig. 3C-F). Likewise, in contrasted with the corresponding values obtained from the ‘Control’, the contents of Pro, total soluble sugars, total phenolics and total flavonoids increased by 91.38, 62.41, 65.76 and 56.51%, respectively, in ‘CK+WL’ plants, and by 75.50, 53.02, 52.04 and 44.96%, respectively, in ‘GA 3 +WL’ plants, followed by the stress recovery period (Fig. 3C-F). 3.6. CK and GA 3 supplementation improved yield-related features of mung bean plants Compared to the ‘Control’, ‘WL’ plants displayed notable decrements in pod length (by 13.18%), total number of pods per plant (36.36%), thousand seed weight (13.48%) and seed yield per plant (42.23%) (Table 2). Intriguingly, significant improvement in pod length (by 9.29 and 7.77%), total number of pods per plant (23.81 and 19.05%), and seed yield per plant (25.52 and 20.76%, respectively) were observed in ‘CK+WL’ and ‘GA 3 +WL’ plants when compared with that of ‘WL’ plants (Table 2). On the other hand, ‘CK+WL’ and ‘GA 3 +WL’ plants displayed noticeable reduction in total number of pods per plant (21.21 and 24.24%) and seed yield per plant (27.49 and 30.24%, respectively), in relation to the ‘Control’ plants (Table 2). Nonetheless, no distinct difference was observed in pod length between ‘Control’ and ‘CK+WL’ plants, while 6.43% reduction in pod length was recorded in ‘GA 3 +WL’ plants versus ‘Control’ plants (Table 2). 4. Discussion Waterlogging in agricultural field is generally triggered by climate change-induced intensive and/or extensive rainfall over a period of time, which has detrimental consequences on a number of crop plants, including mung bean (Donat et al., 2016 ; Amin et al., 2015 ). The current study investigated the effective mitigation measure of waterlogged-induced damage to mung bean plants by exploring the potential role of CK and GA 3 . In the present study, waterlogging-induced deleterious effects was evident as a reduction of shoot height, shoot and root FW, and stem girth width; however, intriguingly the foliar application of CK and GA 3 alleviated those waterlogged-mediated detrimental effects (Table 1 ). More fundamentally, the positive effects of CK and GA 3 also remain persistent after 15-days of recovery (Table 1 ). Our results also clearly indicated that CK was more effective than GA 3 in ameliorating waterlogged-induced growth retardation (Table 1 ). It is well reported that both CK and GA 3 played a pivotal role in protecting plants from a variety of stresses (Ahanger et al., 2018 ; Wang et al., 2019 ; Rady et al., 2021 ; Saleem et al., 2020 ); nonetheless, their underlying mechanisms in alleviation of waterlogged-induced damages are yet to be investigated. Thus, we used various physiological and biochemical assay to investigate on how CK and GA 3 could improve the growth response of mung bean plants in response to waterlogging stress. The waterlogged-induced growth inhibition and biomass reduction in mung bean plants might be a consequence of impeded photosynthesis (Table 1 ; Fig. 1 A). It is envisaged that at the beginning of waterlogging stress, plant roots rapidly transmit a xylem-borne signals to the leaves in the form of hormones, most notably abscisic acid (ABA) to slow down the process of transpiration through stomatal closure, and thus attenuating carbon dioxide availability in leaves (Jackson et al., 2003 ; Najeeb et al., 2015 ). Coupled with decreased transpiration rate because of enhanced stomatal closure, the waterlogged-mediated anaerobic respiration triggered a drop in root hydraulic conductivity and an upsurge in root cell death, resulting in inadequate ATP for active delivery of water and nutrients to the shoots, thereby diminishing net photosynthetic rate (Steffens et al., 2005 ; Kaur et al., 2019 ). Henceforth, the supply of photoassimilates from leaves to different plant parts is also reduced, which ultimately leading to poor plant growth and biomass production (Kogawara et al., 2006 ). Apart from these, waterlogged-induced demolition of photosynthetic pigments, including Chl a and b , also responsible for reduced net photosynthetic rate (Ren et al., 2016 ; Zhang et al., 2020 ), as also observed in the present study (Fig. 1 A and 3 A-C). On the other hand, mung bean plants treated with CK and GA 3 significantly improved the net photosynthetic, as well as the contents of Chl a and Chl b under waterlogged conditions, implying that CK and GA 3 played a pivotal role in improving photosynthetic process during waterlogging stress (Fig. 1 A,C and 3 A-C). Improvements of photosynthesis as well as photosynthetic pigments were also observed after recovery period in response to CK and GA 3 supplementation (Fig. 1 A,C and 3 A-C). Importantly, our results clearly showed that CK supplementation plausibly played more decisive role than GA 3 supplementation in repairing the growth and photosynthetic performance of mung bean under waterlogging stress. Similar to our findings, protection of photosynthetic pigments and consequent improvement in plant growth performance were also reported in drought-stressed pomegranate ( Punica granatum ) and salt-stressed tomato ( Solanum lycopersicum ) plant in response to CK application, while GA 3 provided positive results in salt-stressed tomato, drought-stressed faba bean ( Vicia faba ) and copper-exposed jute ( Corchorus capsularis ) plants (Bompadre et al., 2015 ; Ahanger et al., 2018 ; Siddiqui et al., 2020 ; Saleem et al., 2020 ; Rady et al., 2021 ). We also observed that, in comparison with ‘WL’ plants, CK or GA 3 -treated plants demonstrated remarkably improved WUE (Fig. 2 D,E), indicating that CK and GA 3 might enabled mung bean plants to produce more biomass under the conditions of limited water uptake. Surprisingly, after recovery period, we have seen similar positive effects of CK and GA 3 in boosting WUE (Fig. 2 D,E). Importantly, our findings revealed that the spraying of mung bean plants with CK was more efficient than GA 3 in preventing waterlogged-induced deterioration of water use efficiency (Fig. 2 D,E). It is well-known that improving WUE in abiotic-stressed plants without deteriorating growth and yield is scrutinized as a paramount goal of current plant breeding programs (Yang et al., 2019). Our findings support this phenomenon and corroborate with the previous findings where external application of CK to cold-stressed coffee ( Coffea arabica ) and GA 3 to drought-stressed faba bean seedlings improved WUE (Acidri et al., 2020; Rady et al., 2021 ). Water stagnation because of waterlogging conditions causes poor gas exchange between soils and plant roots, resulting in hypoxia or anoxia in plant tissues (Matin and Jalali, 2017 ). The scarcity of oxygen induces leakage of electrons from mitochondria, which together with impaired photosynthesis, triggers a burst of excessive ROS production, causing oxidative damage to waterlogged-stressed seedlings (Xu et al., 2013), and was clearly manifested in our study with increasing H 2 O 2 and MDA in ‘WL’ plants (Fig. 3 A,B). Intriguingly, external application of either CK or GA 3 to waterlogged-stressed mung bean plants resulted in a dramatic reduction in ROS accumulation (Fig. 3 A,B), implying that CK and GA 3 plays an important role in reducing oxidative burden induced by ROS and providing protection to plasma membrane integrity. In line with our study, previous reports also demonstrated that the contents of H 2 O 2 and MDA were noticeably diminished in salt-stressed tomato and okra ( Abelmoschus esculentus ) by CK application, and in drought-stressed wheat ( Triticum aestivum ) and boron-stressed tomato by GA 3 application (Ahanger et al., 2018 ; Wang et al., 2019 ; Moumita et al., 2019; Javed et al, 2021 ). Our results also demonstrated that, compared with ‘WL’, the leaves of ‘CK + WL’ and ‘GA 3 + WL’ plants accumulated higher levels of non-enzymatic antioxidants, namely total phenolics and total flavonoids (Fig. 3 E,F). More fundamentally, following the recovery period, there was a similar pattern of improvement in total phenolics and total flavonoids levels in response to CK or GA 3 application (Fig. 3 A,B,E,F). Phenolics and flavonoids are two well-known secondary metabolites that play a pivotal role in scavenging free radicals and preventing lipid peroxidation, thus maintaining membrane fluidity and shielding cell membrane damage from oxidative damage under different abiotic stresses, including waterlogging (Ahmad et al., 2010 ; Elkelish et al., 2020 ; Patel et al., 2020 ). These results implied that either CK or GA 3 -induced alleviation of ROS burden was most likely due to the heightened level of non-enzymatic antioxidants (Fig. 3 A,E,F). It is worth noting that the application of CK to mung bean plants displayed greater role in alleviating the contents of H 2 O 2 and MDA by enhancing the levels of total phenolics and flavonoids when compared with the plants treated with GA 3 (Fig. 3 A,B,E,F). Nonetheless, in consistent with our findings, Ahanger et al. ( 2018 ) reported that CK application to salt-stressed tomato plants increased the level of total phenolics and flavonoids, while Rady et al. ( 2021 ) revealed that GA 3 supplementation to drought-stressed faba bean plants increased phenolics accumulation. To overcome waterlogged-induced osmotic stress, plants also accumulate a number of compatible solutes, including Pro and total soluble sugars, which play vital roles in maintaining water balance, retrieving photosynthetic functions, stabilizing cellular components, ROS scavenging, improving cellular signalling and secondary metabolite biosynthesis (Sami et al., 2016 ; Barickman et al., 2019; Chávez-Arias et al., 2019 ; Xiao et al., 2020 ). Our data on compatible solutes revealed that the levels of Pro and total soluble sugar were dramatically enhanced in ‘WL’ plants (Fig. 3 C,D). Importantly, exogenous application of CK and GA 3 further enhanced the contents of Pro and total soluble sugars in the leaves of ‘CK + WL’ and ‘GA 3 + WL’ plants, and this trend was found surprisingly similar following the recovery period (Fig. 3 C,D). The results of our study clearly indicated that CK-treated waterlogged-stressed mung bean plants were able to accumulate more Pro and soluble sugars than GA 3 -treated plant (Fig. 3 C,D). Our findings coincided with the results of Sarafraz-ardakani et al. (2014), who also observed that CK supplementation increased the amount of Pro and soluble sugars in drought-stressed wheat plants. Apart from the physiological and biochemical assay, we also focused on the yield-related attributes of mung bean plants to confirm the beneficiary role of the CK and GA 3 application. Waterlogged-stressed mung bean plants showed a decrease in yield-associated features such as pod length, total number of pods per plant, thousand seed weight, and seed yield per plant (Table 2 ), which might be correlated with poor plant growth because of impaired photosynthesis and enhanced accumulation of ROS (Fig. 2 A-C and 3 A). In contrast, foliar application of either CK or GA 3 to waterlogged-exposed mung bean plants resulted in a dramatic increase in pod length, total number of pods per plant, thousand seed weight, and seed yield per plant (Table 2 ). These results suggested that CK or GA 3 supplementation improved photosynthetic efficiency of mung bean to sustain higher levels of total soluble sugars (Fig. 3 F), as well as heightened the levels of Pro and non-enzymatic antioxidants to resist the phytotoxic effects of waterlogging through the reductions of ROS-mediated oxidative stress (Fig. 3 A, C-E), resulting in improvement of yield performance (Table 2 ). Importantly, CK-treated waterlogged-stressed mung bean plants provided greater yield-associated features compared with GA 3 -treated plants. Yield improvement in wheat plants under heat and drought stress was also reported upon supplementation of CK and GA 3 , respectively (Gupta et al., 2012 ; Yang et al., 2016 ). 5. Conclusion In summary, our results concluded that treating plants with either CK or GA 3 positively regulates defense responses in waterlogged-stressed mung bean plants through regulating several physiological and biochemical mechanisms. In comparison with water-sprayed stressed plants, CK and GA 3 -sprayed plants demonstrated improvement in plant height, stem girth width and plant biomass, which might be interlinked with delayed of photosynthetic pigment destruction, improvement of photosynthetic rate and WUE. Under waterlogged conditions, application of CK and GA 3 successfully decreased the levels of ROS accumulation and consequent MDA level; implying that CK and GA 3 played a decisive role in controlling ROS-induced cellular damage. These observations were closely interlinked to heightened levels of non-enzymatic antioxidants, including total phenolics and total flavonoids. Elevated levels of proline and total soluble sugars in CK and GA 3 -sprayed plants also contributed to the maintenance of better water status and osmotic adjustment under waterlogging. Intriguingly, the positive regulatory roles of CK and GA 3 in the improvement of above-mentioned parameters were also persist after 15-days of recovery period. All of these physiological and biochemical mechanisms in CK and GA 3 -sprayed plants contribute to improve yield-associated features, including pod length, total number of pods per plant, thousand seed weight, and seed yield per plant, in relation of water-sprayed plants. Importantly, plants treated with CK performed better compared with GA 3 -treated plants. Taken together, our findings suggests that either CK or GA 3 could be a potential chemical for mitigating waterlogged-induced detrimental effects on crop performance to ensure sustainable agriculture in flash flood-prone areas. However, together with field trials, comprehensive studies are recommended to decipher the underlying molecular mechanisms that are regulated by either CK or GA 3 under waterlogged conditions. Declarations Author Contributions MRI: conceived, designed and supervised the experiment, wrote and edited the manuscript, and gave final approval; MMR: analyzed and interpreted the data, wrote and revised the manuscript; MA and EJ: performed the experiment, prepared the materials, collected and analyzed the data; SSK: helped in statistical analysis, interpretation of the data and results, and writing the manuscript; MH: monitoring the experiment and reviewed the manuscript. Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgements The work was supported by a grant from the Bangladesh Bureau of Educational Information and Statistics (BANBEIS), Ministry of Education, Government of the People's Republic of Bangladesh under the Higher Educational Research Program. References Ahanger, M.A., Alyemeni, M.N., Wijaya, L., Alamri, S.A., Alam, P., Ashraf, M., Ahmad, P., 2018. Potential of exogenously sourced kinetin in protecting Solanum lycopersicum from NaCl-induced oxidative stress through up-regulation of the antioxidant system, ascorbate-glutathione cycle and glyoxalase system. PLOS One 13, e0202175. Ahmad, P., Jaleel, C.A., Salem, M.A., Nabi, G., Sharma, S., 2010. Roles of enzymatic and non-enzymatic antioxidants in plants during abiotic stress. Crit. Rev. Biotechnol. 30, 161–175. 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The effects of exogenous cytokinin and gibberellic acid on shoot height, shoot dry weight, root dry weight and stem girth width of mung bean plants after 5-days of stress and 15-days of recovery periods Treatments Shoot height (cm) Shoot FW (g) Root FW (g) Stem girth width (mm) After stress After recovery After stress After recovery After stress After recovery After stress After recovery Control 16.96 ± 0.65 a 33.21 ± 1.58 a 2.34 ± 0.09 a 9.68 ± 0.44 a 0.37 ± 0.02 a 1.90 ± 0.07 a 4.82 ± 0.31 a 7.04 ± 0.04 a WL 11.29 ± 0.42 c 21.49 ± 0.90 d 1.47 ± 0.09 c 5.42 ± 0.17 c 0.22 ± 0.01 c 1.22 ± 0.10 c 3.03 ± 0.14 c 4.77 ± 0.27 c CK+WL 15.29 ± 0.46 b 28.94 ± 0.58 b 2.03 ± 0.06 b 7.34 ± 0.26 b 0.29 ± 0.01 b 1.65 ± 0.06 ab 4.12 ± 0.08 b 6.41 ± 0.20 b GA 3 +WL 14.17 ± 0.33 b 26.62 ± 0.48 c 1.87 ± 0.03 b 6.78 ± 0.30 b 0.27 ± 0.01 b 1.51 ± 0.12 bc 3.96 ± 0.12 b 5.92 ± 0.18 b Values are means ± standard errors ( n = 4). Different alphabetical letters as superscripted within the same column indicate significant differences among various treatments according to a least significant difference test (LSD) at P < 0.05. WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L - 1 ) waterlogged-stressed plants; GA 3 +WL, gibberellic acid (GA)-sprayed (50 mg L - 1 ) waterlogged-stressed plants; FW, fresh weight. Table 2. The effects of exogenous cytokinin and gibberellic acid on yield-contributing parameters of stressed and stress-free mung bean plants Treatments Pod length (cm) Total number of pods per plant Thousand seed weight (g) Seed yield (g per plant) Control 8.46 ± 0.19 a 33.00 ± 1.29 a 51.10 ± 0.86 a 16.47 ± 0.59 a WL 7.35 ± 0.19 c 21.00 ± 0.41 c 44.21 ± 1.07 b 9.51 ± 0.30 c CK+WL 8.03 ± 0.10 ab 26.00 ± 1.47 b 47.12 ± 2.31 ab 11.94 ± 0.64 b GA 3 +WL 7.92 ± 0.09 b 25.00 ± 0.41 b 46.23 ± 1.88 ab 11.49 ± 0.33 b Values are means ± standard errors ( n = 4). Different alphabetical letters as superscripted within the same column indicate significant differences among various treatments according to a least significant difference test (LSD) ( P < 0.05). WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L - 1 ) waterlogged-stressed plants; GA 3 +WL, gibberellic acid (GA)-sprayed (50 mg L - 1 ) waterlogged-stressed plants. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Mezanur Rahman","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2ElEQVRIiWNgGAWjYHACNhAhZwDnHwCK8BChxZh0LYkbiNaiO/vws0c3au6kb+c/nfbhZxuDHN+NBLYHb/BoMTuXZm6cc+xZ7s4ZuZtn9rYxGEveSGA3nINPyxkGM+kctsO5G27wbmbgbQO6EGiLND6HmZ1h/yad8+9wusH5s5sZ/7Yx1BOhhcdMOrftcILBgdzNzEBbEgyI0FImndt32BDkF2aZcxKGM888bCfgF/Zt0jnfDsub8wMd9qbMRp7vePIxvCGGDiSAmLGBBA2jYBSMglEwCrABACQATw45h10lAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0001-8822-9683","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Md.","middleName":"Mezanur","lastName":"Rahman","suffix":""},{"id":37675940,"identity":"389b0a49-72ee-4232-9db6-6229a6c82eb0","order_by":2,"name":"Munny Akter","email":"","orcid":"","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Munny","middleName":"","lastName":"Akter","suffix":""},{"id":37675941,"identity":"d6c477d0-3ae8-407b-8108-2ed03f5c1fde","order_by":3,"name":"Erin Zama","email":"","orcid":"","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Erin","middleName":"","lastName":"Zama","suffix":""},{"id":37675942,"identity":"e2b60b08-c1c5-478d-9d4e-986f18c78ea0","order_by":4,"name":"Sanjida Sultana Keya","email":"","orcid":"","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Sanjida","middleName":"Sultana","lastName":"Keya","suffix":""},{"id":37675943,"identity":"09d97a56-4bc4-47db-85ac-c54409044536","order_by":5,"name":"Mehfuz Hasan","email":"","orcid":"","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Mehfuz","middleName":"","lastName":"Hasan","suffix":""}],"badges":[],"createdAt":"2021-07-06 22:08:59","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":true,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false,"coiExplicitlySet":false},"doi":"10.21203/rs.3.rs-690868/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-690868/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":11301475,"identity":"8f9d1241-37b9-4881-bf4e-bd049f0391e0","added_by":"auto","created_at":"2021-07-09 14:38:28","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":220066,"visible":true,"origin":"","legend":"Effects of exogenous cytokinin and gibberellic acid on (A) photosynthetic rate (Pn), (B) stomatal conductance to H2O (gs), (C) transpiration rate (E) (D) intrinsic water-use efficiency (WUEint) and (E) instantaneous water-use efficiency (WUEins) in the leaves of mung bean plants after 5-days of stress and 15-days of recovery period. Data represent means of four independent replicates (n = 4). Vertical bars indicate standard errors. Different letters represent significant differences at P \u003c 0.05 (least significant difference test). WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L1) waterlogged-stressed plants; GA3+WL, gibberellic acid (GA)-sprayed (50 mg L1) waterlogged-stressed plants.","description":"","filename":"Onlinefloatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-690868/v1/16f52b2d2bd75b691761747f.png"},{"id":11301476,"identity":"591d56de-4ed3-4d64-a568-e439ecee5273","added_by":"auto","created_at":"2021-07-09 14:38:28","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":130841,"visible":true,"origin":"","legend":"Effects of exogenous cytokinin and gibberellic acid on (A) Chlorophyll (Chl) a, (B) Chl b and (C) Chl (a+b) in the leaves of mung bean plants after 5-days of stress and 15-days of recovery period. Data represent means of four independent replicates (n = 4). Vertical bars indicate standard errors. Different letters represent significant differences at P \u003c 0.05 (least significant difference test). WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L1) waterlogged-stressed plants; GA3+WL, gibberellic acid (GA)-sprayed (50 mg L1) waterlogged-stressed plants; FW, fresh weight.","description":"","filename":"Onlinefloatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-690868/v1/8065815ef18aa61cd1324bdf.png"},{"id":11301477,"identity":"f88ae514-242f-48af-9dae-e01840978d78","added_by":"auto","created_at":"2021-07-09 14:38:28","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":250696,"visible":true,"origin":"","legend":"Effects of exogenous cytokinin and gibberellic acid on contents of (A) malondialdehyde (MDA), (B) hydrogen peroxide (H2O2), (C) proline, (D) total soluble sugars, (E) total phenolics and (F) total flavonoids in the leaves of mung bean plants after 5-days of stress and 15-days of recovery period. Data represent means of four independent replicates (n = 4). Vertical bars indicate standard errors. Different letters represent significant differences at P \u003c 0.05 (least significant difference test). WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L1) waterlogged-stressed plants; GA3+WL, gibberellic acid (GA)-sprayed (50 mg L1) waterlogged-stressed plants; FW, fresh weight; GAE, gallic acid equivalent; QE, quercetin equivalent.","description":"","filename":"Onlinefloatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-690868/v1/bd7e886c4c71436d0b4418cb.png"},{"id":13703060,"identity":"536d55c7-895c-4b80-9954-cf65eb600cfa","added_by":"auto","created_at":"2021-09-17 13:39:06","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1077339,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-690868/v1/05126201-37b6-415d-9b87-9e2e321b90a0.pdf"}],"financialInterests":"","formattedTitle":"\u003cp\u003eInsights into the role of cytokinin and gibberellic acid in improving waterlogging tolerance of mung bean\u003c/p\u003e","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eWaterlogging has become a major environmental problem in global agriculture, arises from heavy rainfall, impromptu irrigation, inadequate drainage systems, unleveled fields, and soil compaction (Kaur et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Under waterlogged conditions, the mass of oxygen in the soil attenuates partially (hypoxic) or fully (anoxic) due to high microbial activity, and the resulting carbon dioxide accumulation in the root zone restricts root metabolism, aerobic respiration, ATP synthesis, and nutrient acquisition, resulting in a significant reduction in the growth, development and biomass of roots and shoots (Najeeb et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Gill et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Dossa et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Moreover, waterlogging-induced excessive oxygen scarcity in water impairs root permeability and causes root injury, leading to a decrement of hydraulic conductivity and consequent stomatal closure, resulting in a significant reduction in net photosynthetic and transpiration rates (Else et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Anee et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). While photosynthesis is disrupted in waterlogged plants, lethal levels of reactive oxygen species (ROS) such as superoxide (O\u003csub\u003e2\u003c/sub\u003e\u003csup\u003e\u0026bull;\u0026minus;\u003c/sup\u003e), singlet oxygen (\u003csup\u003e1\u003c/sup\u003eO\u003csub\u003e2\u003c/sub\u003e), hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e) and hydroxyl radical (\u003csup\u003e\u0026bull;\u003c/sup\u003eOH) are produced, which are harmful to cellular components and cause DNA and RNA damage, lipid peroxidation, protein oxidation, and enzyme inactivation (Yeung et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Zhou et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePlants can acclimatise the deletorious consequence of waterlogging-mediated soil oxygen dearth through adapting various morphological, physiological and biochemical mechanisms. Increased adventitious root development and faster stem elongation are some examples of morphological adaptations (Yamauchi et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), whereas reduced stomatal conductance and subsequent decrease in net photosynthetic rate are some short-term physiological adaptation process (Jacobsen et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Bhusal et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Additionally, to shield themselves from stress, waterlogged plants accumulate a number of osmoprotectants such as proline (Pro), soluble sugars, and sucrose (Tewari and Mishra, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, the most important adaptive mechanism is a well-balanced antioxidant defense system that involves both enzymatic (e.g., superoxide dismutase, SOD; catalase, CAT; ascorbate peroxidase, APX; glutathione \u003cem\u003eS\u003c/em\u003e-transferase, GST; glutathione peroxidase, GPX) and non-enzymatic (e.g., total phenolics, total flavonoids, carotenoids) antioxidants that scavenge overaccumulation of ROS (Doupis et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Fukao et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Garcia et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMung bean (\u003cem\u003eVigna radiata\u003c/em\u003e) is the third most important pulse crop after grass pea (\u003cem\u003eLathyrus sativus\u003c/em\u003e) and lentil (\u003cem\u003eLens culinaris\u003c/em\u003e) and considered as a major source of pulse protein because of its high protein content in the seeds (20.97\u0026ndash;31.32%) (AESA, 2017; Yi-Shen et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Rahman et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Additionally, mung bean has been well-documented for its detoxification bioactivities along with its use as a treating agent for various health conditions, including stimulation of human mental function, allaying of heat stroke and regulation of gastrointestinal upset (Yi-Shen et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hou et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). More fundamentally, mung bean gaining its popularity in recent years and extensively cultivated throughout Bangladesh as a cash crop in the extensive rice-based cropping system, owing to its low fertilizer and pesticide requirements, which provides farmers with good economic benefits and nutritional security (Rahman et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). According to the estimation in the years 2016-17, cultivation of mung bean covered approximately 0.3188\u0026nbsp;million hectares (Mha) out of 0.9976 Mha of total pulse crop areas and yielded 0.21\u0026nbsp;million metric tonnes of mung bean (DAE. Krishi Dairy, 2018). However, recently the emerging waterlogging problem in major mung bean growing areas like Barishal, Patuakhali, Mymensingh, Sylhet, Chittagong hill tract, Natore, Khulna, Brahmanbaria and Pabna region make the land untenable for growing mung bean (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.ffwc.gov.bd/index.php/map/inundation-map/bangladesh-today\u003c/span\u003e\u003c/span\u003e). Therefore, the country must devise a strategy for effectively alleviating the effects of waterlogging stress in order to improve mung bean production in flash flood-prone areas. In this context, the search of a simple and cost-effective technology propelled us to use phytohormone such as cytokinin (CK) and gibberellic acid (GA\u003csub\u003e3\u003c/sub\u003e) for overcoming the deleterious effects of waterlogging on the growth and yield-related attributes of mung bean.\u003c/p\u003e \u003cp\u003eIn the present study, we first examined whether CK and GA\u003csub\u003e3\u003c/sub\u003e provides protection to mung bean from waterlogging stress, and to our expectations it really did. Next, we also examined the regulatory roles of CK and GA\u003csub\u003e3\u003c/sub\u003e in improving mung bean tolerance to waterlogging by investigating the morpho-physiological and biochemical mechanisms through assessments of the following key attributes: (i) plant growth performance and biomass production, (ii) gas-exchange features, (iii) photosynthetic pigment status, (iv) excessive waterlogging-mediated oxidative damage in terms of elevated ROS levels and lipid peroxidation, (v) different types of osmolyte accumulations, and (vi) improvement of non-enzymatic antioxidant defense. Additionally, we also record the yield-associated features of waterlogged-stressed and waterlogged-devoid mung bean plants.\u003c/p\u003e"},{"header":"2. Materials And Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Plant growth conditions and treatments imposition\u003c/h2\u003e \u003cp\u003eMung bean (\u003cem\u003eVigna radiata\u003c/em\u003e) genotype, VC6173-B, was used in the present study owing to their properties of short duration (80 days to maturity), greater grain weight (44g/1000 grain) and high yield potential (1.5 tons ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Healthy seeds of VC6173-B were germinated following the procedures of Rahman et al. (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Afterward, six germinated seeds were sown in each plastic pot (height \u0026times; diameter\u0026thinsp;=\u0026thinsp;20 cm \u0026times; 16 cm) having 8 kg of soil per pot. The soil used was prepared by intermixing of cow dung, sand and soil at the ratio of 1:1:3 (in a weight basis). Additionally, the soil of the pots was also fertilized with urea, triple super phosphate (TSP) and muriate of potash (MP) at the rate of 0.176, 0.400 and 0.347 g corresponding to 40-60-40 Kg N, P\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e5\u003c/sub\u003e and K\u003csub\u003e2\u003c/sub\u003eO per hectare (Islam et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). It is worth noting that one-third of urea and the whole amount of TSP, MP and gypsum were mixed with the soil as basal dose during final pot preparation. The remaining amount of urea was applied at 10 and 25 days after emergence. In order to protect the seedlings from insect-induced damage, Ripcord 10 EC (BASE Bangladesh Limited) at 1 mL L\u003csup\u003e\u0026minus;1\u003c/sup\u003e water was sprayed at vegetative stage of plant growth. The average maximum and minimum temperature of the experimental area were lies in the range of 32.6 to 22.8\u0026deg;C, with 85.6% of relative humidity. Notably, the number of seedlings was thinned to two in each pot after eighth day of germination and continues to grow in normal condition up to the imposition of treatments.\u003c/p\u003e \u003cp\u003ePrior to waterlogging exposure for the next five days, fifteen-day-old mung bean seedlings at vegetative V1 stage (when first trifoliate were fully developed) were grouped into four sets as follows:\u003c/p\u003e \u003cp\u003e \u003col\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003ewater-sprayed waterlogged-stress-free plants (Control)\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003ewater-sprayed waterlogged-stressed plants (WL)\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003ecytokinin (CK)-sprayed (50 mg L\u003csup\u003e\u0026minus;1\u003c/sup\u003e) waterlogged-stressed plants (CK\u0026thinsp;+\u0026thinsp;WL)\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003cspan\u003e \u003cli\u003e \u003cp\u003egibberellic acid (GA\u003csub\u003e3\u003c/sub\u003e)-sprayed (50 mg L\u003csup\u003e\u0026minus;1\u003c/sup\u003e) waterlogged-stressed plants (GA\u003csub\u003e3\u003c/sub\u003e\u0026thinsp;+\u0026thinsp;WL)\u003c/p\u003e \u003c/li\u003e \u003c/span\u003e \u003c/ol\u003e \u003c/p\u003e \u003cp\u003eIt is worth noting that foliar spray with either CK or GA\u003csub\u003e3\u003c/sub\u003e or water (20 mL to each pot) was done for two times in a day (9.00 AM to 10.00 AM, and 3.00 PM to 4.00 PM). Tween-20 surfactant (0.2%, v/v) was used to ensure maximum adherence of CK, GA\u003csub\u003e3\u003c/sub\u003e and water to the leaves. Notably, the level of water for creating waterlogging stress was maintained at 2.5 cm above the soil surface. Following the stress period, one set of seedlings (Set I) were immediately harvested, and another set (Set II) was harvested 15-days after recovery, to record the morphological, physiological and biochemical parameters. Importantly, after stress exposure, one set of seedlings (Set III) was allowed to grow until harvest with normal irrigation to determine yield-related attributes, including pod length, total number of pods per plant, thousand seed weight, and seed yield per plant. Each set of experiments were repeated four times under the same experimental conditions. Importantly, for determining various morphological, physiological and biochemical parameters, the first trifoliate leaves from the bottom of the plants were collected.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Determination of growth-related parameters\u003c/h2\u003e \u003cp\u003eThe growth performance of mung bean plants was assessed by measuring the shoot height, and fresh weight (FW) of both shoots and roots. Additionally, stem girth of the mung bean plants was determined using a slide caliper.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Measurement of gas exchange features\u003c/h2\u003e \u003cp\u003eGas exchange features, including photosynthesis rate (\u003cem\u003ePn\u003c/em\u003e), stomatal conductance to water (H\u003csub\u003e2\u003c/sub\u003eO) (\u003cem\u003eg\u003c/em\u003e\u003csub\u003e\u003cem\u003es\u003c/em\u003e\u003c/sub\u003e) and transpiration rate (\u003cem\u003eE\u003c/em\u003e) of mung bean plants were measured using a LI-6400XT portable photosynthesis system (LI-COR Biosciences, Lincoln, Nebraska, USA) from 11.00 AM to 2.00 PM under a full sun-light condition. The parameters \u003cem\u003ePn\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003e\u003cem\u003es\u003c/em\u003e\u003c/sub\u003e and \u003cem\u003eE\u003c/em\u003e were used for assessing the instantaneous water-use efficiency (WUEins; ratio \u003cem\u003ePn\u003c/em\u003e/\u003cem\u003eE\u003c/em\u003e) and intrinsic water-use efficiency (WUEint; ratio \u003cem\u003ePn\u003c/em\u003e/\u003cem\u003eg\u003c/em\u003e\u003csub\u003e\u003cem\u003es\u003c/em\u003e\u003c/sub\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Quantification of photosynthetic pigments, and total phenolic and flavonoid contents in mung bean leaves\u003c/h2\u003e \u003cp\u003eThe freshly harvested mung bean leaves were used to determine the contents of chlorophylls (Chls) [Chl \u003cem\u003ea\u003c/em\u003e, Chl \u003cem\u003eb\u003c/em\u003e and Chl (\u003cem\u003ea\u003c/em\u003e\u0026thinsp;+\u0026thinsp;\u003cem\u003eb\u003c/em\u003e)] by using the visible spectrophotometer (Model: T60 UV, PG Instruments Limited, Leicestershire, UK) according to the protocols described by Lichtenthaler and Wellburn (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). The contents of total phenolic and total flavonoid were quantified following the methods of Ainsworth and Gillespie (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), and Zhishen et al. (\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), respectively.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Quantification of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e, MDA, proline and total soluble sugars\u003c/h2\u003e \u003cp\u003eFreshly harvested leaf tissues were used for the quantification of hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e), malondialdehyde (MDA) (a lipid peroxidation product), proline (Pro) and total soluble sugars levels following the comprehensive protocols described by Yu et al. (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), Health and Packer (1968), Bates et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1973\u003c/span\u003e) and Somogyi (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1952\u003c/span\u003e), respectively.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.6. Statistical analysis\u003c/h2\u003e \u003cp\u003eThe obtained data were subjected to a one-way analysis of variance (ANOVA) using Statistix software (version 10). Different alphabetical letters were used to denote the significant variations among different treatments at the \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 level following a least significant difference (LSD) test. All the numerical data in figures and tables are presented as means\u0026thinsp;\u0026plusmn;\u0026thinsp;standard errors (SEs) of four independent replications.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cp\u003e\u003cem\u003e3.1. CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation improved morphological features of mung bean plants under waterlogging stress and recovery period\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn comparison with \u0026lsquo;Control\u0026rsquo; plants, \u0026lsquo;WL\u0026rsquo; plants displayed significant decrease in shoot height, shoot FW, root FW and stem girth width by 33.46, 37.04, 41.50 and 37.07%, respectively; however, following 15-days of recovery, these parameters were reduced by 35.31, 44.05, 35.88 and 32.18%, respectively (Table 1). On the other hand, \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants showed noteworthy improvement in shoot height by 35.46 and 25.54%, shoot FW by 37.93 and 26.87%, root FW by 36.05 and 25.58%, and stem girth width by 36.06 and 30.61%, respectively, when compared with \u0026lsquo;WL\u0026rsquo; plants (Table 1). Following the recovery period, a substantial enhancement in shoot height (34.70 and 23.90%), shoot FW (37.93 and 26.87%), root FW (36.05 and 25.58%) and stem girth width (36.06 and 30.61%, respectively) was observed in the \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, in comparison with \u0026lsquo;WL\u0026rsquo; plants (Table 1). It was also observed that \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants exhibited notable decrease in shoot height by 9.86 and 16.46%, shoot FW by 13.17 and 20.13%, root FW by 20.41 and 26.53% and stem girth width by 14.40 and 17.83%, respectively, relative to that of \u0026lsquo;Control\u0026rsquo; plants (Table 1). Similarly, in contrast to \u0026lsquo;Control\u0026rsquo; plants, significant reductions in shoot height (by 12.86 and 19.85%), shoot FW (24.21 and 29.94%), root FW (20.58% for GA\u003csub\u003e3\u003c/sub\u003e+WL) and stem girth width (8.99 and 15.86%, respectively) were observed following recovery period in the \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants (Table 1).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.2. CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation improved gas exchange features of mung bean plants under waterlogging stress and recovery period\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eMung bean plants subjected to waterlogging stress and followed by stress recovery period showed a decrease in \u003cem\u003ePn\u003c/em\u003e (by 44.12 and 41.97%),\u003cem\u003e\u0026nbsp;gs\u003c/em\u003e (90.28 and 53.25%) and \u003cem\u003eE\u003c/em\u003e (73.92 and 35.56%, respectively) when compared with \u0026lsquo;Control\u0026rsquo; plants (Fig. 1A-C). Nonetheless, the levels of WUEint and WUEins in \u0026lsquo;WL\u0026rsquo; plants increased by 514.96 and 117.90%, respectively relative to \u0026lsquo;Control\u0026rsquo; plants, whereas the level of WUEint and WUEins in \u0026lsquo;Control\u0026rsquo; and \u0026lsquo;WL\u0026rsquo; plants was comparable after recovery period (Fig. 1D,E). On the other hand, \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants exhibited decreased levels of \u003cem\u003egs\u003c/em\u003e (by 40.03 and 27.36%) and \u003cem\u003eE\u003c/em\u003e (32.87 and 22.55%), and increased levels of \u003cem\u003ePn\u003c/em\u003e (by 36.80 and 26.65%), WUEint (119.44 and 63.82%) and WUEins (100.06 and 61.36%, respectively) when compared with that of \u0026lsquo;WL\u0026rsquo; plants (Fig. 1A-E). Similarly, following the recovery period, decreased levels of \u003cem\u003egs\u003c/em\u003e (by 35.60 and 27.61%) and \u003cem\u003eE\u003c/em\u003e (34.14 and 21.37%), and increased levels of \u003cem\u003ePn\u003c/em\u003e (36.80 and 26.65%), WUEint (114.55 and 72.68%) and WUEins (107.42 and 59.09%, respectively) were recorded in \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, as compared with that of \u0026lsquo;WL\u0026rsquo; plants (Fig. 1A-E). In relation to the \u0026lsquo;Control\u0026rsquo;, a sharp reduction in the levels of \u003cem\u003ePn\u003c/em\u003e (by 23.56 and 29.23%), \u003cem\u003egs\u003c/em\u003e (94.17 and 92.94%) and \u003cem\u003eE\u003c/em\u003e (82.49 and 79.80%) was noticed in \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, respectively. Similarly, the levels of \u003cem\u003ePn\u003c/em\u003e, \u003cem\u003egs\u003c/em\u003e and \u003cem\u003eE\u003c/em\u003e were noticeably reduced by 20.85, 69.89 and 57.56% in \u0026lsquo;CK+WL\u0026rsquo;, and by 27.28, 66.15 and 49.33% in \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants followed by the recovery period over the corresponding values of \u0026lsquo;Control\u0026rsquo; plants (Fig. 1A-C). Additionally, in relation to the \u0026lsquo;Control\u0026rsquo;, the level of WUEint and WUEins was substantially enhanced by 1249.50 and 335.94% in \u0026lsquo;CK+WL\u0026rsquo; and by 907.44 and 251.60% in \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants followed by exposure to waterlogging stress. Whereas after recovery period, augmented level of WUEint (by 165.23 and 113.47%) and WUEins (87.07 and 43.47%) was observed in \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, respectively (Fig. 1D,E).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.3. CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation improved photosynthetic pigments status of mung bean plants under waterlogging stress and recovery period\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn relation to \u0026lsquo;Control\u0026rsquo; plants, a sharp decline in the contents of Chl \u003cem\u003ea\u003c/em\u003e (by 50.10 and 43.29%), Chl \u003cem\u003eb\u003c/em\u003e (65.44 and 62.99%) and Chl (\u003cem\u003ea\u003c/em\u003e+\u003cem\u003eb\u003c/em\u003e) (54.35 and 48.96%) were observed in the leaves of \u0026lsquo;WL\u0026rsquo; plants following stress and recovery periods, respectively (Fig. 2A-C). In contrast, CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation protected the photosynthetic pigments from waterlogged-induced deleterious effects by enhancing the contents of Chl \u003cem\u003ea\u003c/em\u003e (by 74.01 and 60.52%), Chl \u003cem\u003eb\u003c/em\u003e (142.38 and 124.40%) and Chl (\u003cem\u003ea\u003c/em\u003e+\u003cem\u003eb\u003c/em\u003e) (88.37 and 73.94%, respectively) in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, respectively (Fig. 2A-C). Furthermore, following the recovery period, the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants also displayed a significant rise in the content of Chl \u003cem\u003ea\u003c/em\u003e (by 59.75 and 49.54%), Chl \u003cem\u003eb\u003c/em\u003e (134.31 and 120.36%) and Chl (\u003cem\u003ea\u003c/em\u003e+\u003cem\u003eb\u003c/em\u003e) (75.32 and 64.33%, respectively) in comparison with \u0026lsquo;WL\u0026rsquo; plants (Fig. 2A-C). On the other hand, CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation resulted in a decrement of Chl \u003cem\u003ea\u003c/em\u003e (by 13.16 and 19.90%), Chl \u003cem\u003eb\u003c/em\u003e (16.23 and 22.44%) and Chl (\u003cem\u003ea\u003c/em\u003e+\u003cem\u003eb\u003c/em\u003e) (14.01 and 20.60%) in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, respectively, when compared with \u0026lsquo;Control\u0026rsquo; plants (Fig. 2A-C). Likewise, following the recovery period, a noteworthy decrement in the contents of Chl \u003cem\u003ea\u003c/em\u003e (by 9.40 and 15.19 %), Chl \u003cem\u003eb\u003c/em\u003e (13.28 and 18.44%) and Chl (\u003cem\u003ea\u003c/em\u003e+\u003cem\u003eb\u003c/em\u003e) (10.51 and 16.13%) were observed in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, respectively, in comparison with \u0026lsquo;Control\u0026rsquo; plants (Fig. 2A-C).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.4. CK and GA\u003csub\u003e3\u0026nbsp;\u003c/sub\u003esupplementation suppressed the levels of oxidative stress markers of mung bean plants under waterlogging stress and recovery period\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eImposition of waterlogging stress and following recovery period substantially increased the contents of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (by 113.40 and 62.08%) and MDA (178.67 and 100.18%, respectively) in the leaves of \u0026lsquo;WL\u0026rsquo; plants, relative to that of \u0026lsquo;Control\u0026rsquo; plants (Fig. 3A,B). Fascinatingly, CK and GA\u003csub\u003e3\u003c/sub\u003e treatment resulted in reductions of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (by 57.31 and 45.40%) and MDA (29.01 and 19.04%, respectively) contents in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, in comparison with \u0026lsquo;WL\u0026rsquo; plants (Fig. 3A,B). Similarly, compared with the \u0026lsquo;WL\u0026rsquo; plants, substantial decreases in the contents of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (by 36.61 and 26.44 %) and MDA (33.79 and 20.70%, respectively) were noticed in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants followed by the recovery period (Fig. 3A,B). On the other hand, \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants exhibited noteworthy improvements in the levels of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (by 38.52 and 59.69%) and MDA (71.34 and 102.46%, respectively), when contrasted with that of \u0026lsquo;Control\u0026rsquo; plants (Fig. 3A,B). After the recovery period, the contents of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (by 2.75 and 19.22%) and MDA (32.53 and 58.74%, respectively) in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants were substantially higher compared with the respective value of \u0026lsquo;Control\u0026rsquo; plants (Fig. 3A,B).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.5. CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation improved the levels of Proline (Pro), total soluble sugars, total phenolics and flavonoids of mung bean plants under waterlogging stress and recovery period\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eExposure of mung bean plants to waterlogging stress and followed by a recovery period exhibited a rise in the contents of Pro (63.01 and 42.17%), total soluble sugars (33.20 and 20.29%), total phenolics (57.65 and 23.16%) and total flavonoids (30.98 and 15.89%, respectively) in the leaves of \u0026lsquo;WL\u0026rsquo; plants, when compared with that of \u0026lsquo;Control\u0026rsquo; plants (Fig. 3C-F). In comparison with \u0026lsquo;WL\u0026rsquo; plants, \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants displayed enhanced levels of Pro (by 33.75 and 21.01%), total soluble sugars (32.24 and 21.79%), total phenolics (36.48 and 24.53%) and total flavonoids (37.34 and 28.19%, respectively) (Fig. 3C-F). Similarly, in relation to the \u0026lsquo;WL\u0026rsquo; plants, the levels of Pro, total soluble sugars, total phenolics and flavonoids were noticeably enhanced in \u0026lsquo;CK+WL\u0026rsquo; leaves by 34.61, 35.02, 34.59 and 35.04%, respectively, and in \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; leaves by 23.45, 27.21, 23.45 and 25.08%, respectively, followed by a recovery period (Fig. 3C-F). Significant improvement in the contents of Pro (by 118.02 and 97.25%), total soluble sugars (76.15 and 62.23%), total phenolics (115.16 and 96.32%) and total flavonoids (79.90 and 67.91%, respectively) were recorded in the leaves of \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, when contrasted with the corresponding \u0026lsquo;Control\u0026rsquo; values (Fig. 3C-F). Likewise, in contrasted with the corresponding values obtained from the \u0026lsquo;Control\u0026rsquo;, the contents of Pro, total soluble sugars, total phenolics and total flavonoids increased by 91.38, 62.41, 65.76 and 56.51%, respectively, in \u0026lsquo;CK+WL\u0026rsquo; plants, and by 75.50, 53.02, 52.04 and 44.96%, respectively, in \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants, followed by the stress recovery period (Fig. 3C-F).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e3.6. CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation improved yield-related features of mung bean plants\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eCompared to the \u0026lsquo;Control\u0026rsquo;, \u0026lsquo;WL\u0026rsquo; plants displayed notable decrements in pod length (by 13.18%), total number of pods per plant (36.36%), thousand seed weight (13.48%) and seed yield per plant (42.23%) (Table 2). Intriguingly, significant improvement in pod length (by 9.29 and 7.77%), total number of pods per plant (23.81 and 19.05%), and seed yield per plant (25.52 and 20.76%, respectively) were observed in \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants when compared with that of \u0026lsquo;WL\u0026rsquo; plants (Table 2). On the other hand, \u0026lsquo;CK+WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants displayed noticeable reduction in total number of pods per plant (21.21 and 24.24%) and seed yield per plant (27.49 and 30.24%, respectively), in relation to the \u0026lsquo;Control\u0026rsquo; plants (Table 2). Nonetheless, no distinct difference was observed in pod length between \u0026lsquo;Control\u0026rsquo; and \u0026lsquo;CK+WL\u0026rsquo; plants, while 6.43% reduction in pod length was recorded in \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e+WL\u0026rsquo; plants versus \u0026lsquo;Control\u0026rsquo; plants (Table 2).\u0026nbsp;\u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eWaterlogging in agricultural field is generally triggered by climate change-induced intensive and/or extensive rainfall over a period of time, which has detrimental consequences on a number of crop plants, including mung bean (Donat et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Amin et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The current study investigated the effective mitigation measure of waterlogged-induced damage to mung bean plants by exploring the potential role of CK and GA\u003csub\u003e3\u003c/sub\u003e.\u003c/p\u003e \u003cp\u003eIn the present study, waterlogging-induced deleterious effects was evident as a reduction of shoot height, shoot and root FW, and stem girth width; however, intriguingly the foliar application of CK and GA\u003csub\u003e3\u003c/sub\u003e alleviated those waterlogged-mediated detrimental effects (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). More fundamentally, the positive effects of CK and GA\u003csub\u003e3\u003c/sub\u003e also remain persistent after 15-days of recovery (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Our results also clearly indicated that CK was more effective than GA\u003csub\u003e3\u003c/sub\u003e in ameliorating waterlogged-induced growth retardation (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). It is well reported that both CK and GA\u003csub\u003e3\u003c/sub\u003e played a pivotal role in protecting plants from a variety of stresses (Ahanger et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Rady et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Saleem et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2020\u003c/span\u003e); nonetheless, their underlying mechanisms in alleviation of waterlogged-induced damages are yet to be investigated. Thus, we used various physiological and biochemical assay to investigate on how CK and GA\u003csub\u003e3\u003c/sub\u003e could improve the growth response of mung bean plants in response to waterlogging stress.\u003c/p\u003e \u003cp\u003eThe waterlogged-induced growth inhibition and biomass reduction in mung bean plants might be a consequence of impeded photosynthesis (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). It is envisaged that at the beginning of waterlogging stress, plant roots rapidly transmit a xylem-borne signals to the leaves in the form of hormones, most notably abscisic acid (ABA) to slow down the process of transpiration through stomatal closure, and thus attenuating carbon dioxide availability in leaves (Jackson et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Najeeb et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Coupled with decreased transpiration rate because of enhanced stomatal closure, the waterlogged-mediated anaerobic respiration triggered a drop in root hydraulic conductivity and an upsurge in root cell death, resulting in inadequate ATP for active delivery of water and nutrients to the shoots, thereby diminishing net photosynthetic rate (Steffens et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Kaur et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Henceforth, the supply of photoassimilates from leaves to different plant parts is also reduced, which ultimately leading to poor plant growth and biomass production (Kogawara et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Apart from these, waterlogged-induced demolition of photosynthetic pigments, including Chl \u003cem\u003ea\u003c/em\u003e and \u003cem\u003eb\u003c/em\u003e, also responsible for reduced net photosynthetic rate (Ren et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), as also observed in the present study (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003eA and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA-C). On the other hand, mung bean plants treated with CK and GA\u003csub\u003e3\u003c/sub\u003e significantly improved the net photosynthetic, as well as the contents of Chl \u003cem\u003ea\u003c/em\u003e and Chl \u003cem\u003eb\u003c/em\u003e under waterlogged conditions, implying that CK and GA\u003csub\u003e3\u003c/sub\u003e played a pivotal role in improving photosynthetic process during waterlogging stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003eA,C and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA-C). Improvements of photosynthesis as well as photosynthetic pigments were also observed after recovery period in response to CK and GA\u003csub\u003e3\u003c/sub\u003e supplementation (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003eA,C and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA-C). Importantly, our results clearly showed that CK supplementation plausibly played more decisive role than GA\u003csub\u003e3\u003c/sub\u003e supplementation in repairing the growth and photosynthetic performance of mung bean under waterlogging stress. Similar to our findings, protection of photosynthetic pigments and consequent improvement in plant growth performance were also reported in drought-stressed pomegranate (\u003cem\u003ePunica granatum\u003c/em\u003e) and salt-stressed tomato (\u003cem\u003eSolanum lycopersicum\u003c/em\u003e) plant in response to CK application, while GA\u003csub\u003e3\u003c/sub\u003e provided positive results in salt-stressed tomato, drought-stressed faba bean (\u003cem\u003eVicia faba\u003c/em\u003e) and copper-exposed jute (\u003cem\u003eCorchorus capsularis\u003c/em\u003e) plants (Bompadre et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ahanger et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Siddiqui et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saleem et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Rady et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWe also observed that, in comparison with \u0026lsquo;WL\u0026rsquo; plants, CK or GA\u003csub\u003e3\u003c/sub\u003e-treated plants demonstrated remarkably improved WUE (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003eD,E), indicating that CK and GA\u003csub\u003e3\u003c/sub\u003e might enabled mung bean plants to produce more biomass under the conditions of limited water uptake. Surprisingly, after recovery period, we have seen similar positive effects of CK and GA\u003csub\u003e3\u003c/sub\u003e in boosting WUE (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003eD,E). Importantly, our findings revealed that the spraying of mung bean plants with CK was more efficient than GA\u003csub\u003e3\u003c/sub\u003e in preventing waterlogged-induced deterioration of water use efficiency (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003eD,E). It is well-known that improving WUE in abiotic-stressed plants without deteriorating growth and yield is scrutinized as a paramount goal of current plant breeding programs (Yang et al., 2019). Our findings support this phenomenon and corroborate with the previous findings where external application of CK to cold-stressed coffee (\u003cem\u003eCoffea arabica\u003c/em\u003e) and GA\u003csub\u003e3\u003c/sub\u003e to drought-stressed faba bean seedlings improved WUE (Acidri et al., 2020; Rady et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWater stagnation because of waterlogging conditions causes poor gas exchange between soils and plant roots, resulting in hypoxia or anoxia in plant tissues (Matin and Jalali, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The scarcity of oxygen induces leakage of electrons from mitochondria, which together with impaired photosynthesis, triggers a burst of excessive ROS production, causing oxidative damage to waterlogged-stressed seedlings (Xu et al., 2013), and was clearly manifested in our study with increasing H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e and MDA in \u0026lsquo;WL\u0026rsquo; plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA,B). Intriguingly, external application of either CK or GA\u003csub\u003e3\u003c/sub\u003e to waterlogged-stressed mung bean plants resulted in a dramatic reduction in ROS accumulation (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA,B), implying that CK and GA\u003csub\u003e3\u003c/sub\u003e plays an important role in reducing oxidative burden induced by ROS and providing protection to plasma membrane integrity. In line with our study, previous reports also demonstrated that the contents of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e and MDA were noticeably diminished in salt-stressed tomato and okra (\u003cem\u003eAbelmoschus esculentus\u003c/em\u003e) by CK application, and in drought-stressed wheat (\u003cem\u003eTriticum aestivum\u003c/em\u003e) and boron-stressed tomato by GA\u003csub\u003e3\u003c/sub\u003e application (Ahanger et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Moumita et al., 2019; Javed et al, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur results also demonstrated that, compared with \u0026lsquo;WL\u0026rsquo;, the leaves of \u0026lsquo;CK\u0026thinsp;+\u0026thinsp;WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e\u0026thinsp;+\u0026thinsp;WL\u0026rsquo; plants accumulated higher levels of non-enzymatic antioxidants, namely total phenolics and total flavonoids (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eE,F). More fundamentally, following the recovery period, there was a similar pattern of improvement in total phenolics and total flavonoids levels in response to CK or GA\u003csub\u003e3\u003c/sub\u003e application (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA,B,E,F). Phenolics and flavonoids are two well-known secondary metabolites that play a pivotal role in scavenging free radicals and preventing lipid peroxidation, thus maintaining membrane fluidity and shielding cell membrane damage from oxidative damage under different abiotic stresses, including waterlogging (Ahmad et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Elkelish et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Patel et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). These results implied that either CK or GA\u003csub\u003e3\u003c/sub\u003e-induced alleviation of ROS burden was most likely due to the heightened level of non-enzymatic antioxidants (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA,E,F). It is worth noting that the application of CK to mung bean plants displayed greater role in alleviating the contents of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e and MDA by enhancing the levels of total phenolics and flavonoids when compared with the plants treated with GA\u003csub\u003e3\u003c/sub\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA,B,E,F). Nonetheless, in consistent with our findings, Ahanger et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) reported that CK application to salt-stressed tomato plants increased the level of total phenolics and flavonoids, while Rady et al. (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) revealed that GA\u003csub\u003e3\u003c/sub\u003e supplementation to drought-stressed faba bean plants increased phenolics accumulation.\u003c/p\u003e \u003cp\u003eTo overcome waterlogged-induced osmotic stress, plants also accumulate a number of compatible solutes, including Pro and total soluble sugars, which play vital roles in maintaining water balance, retrieving photosynthetic functions, stabilizing cellular components, ROS scavenging, improving cellular signalling and secondary metabolite biosynthesis (Sami et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Barickman et al., 2019; Ch\u0026aacute;vez-Arias et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Xiao et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Our data on compatible solutes revealed that the levels of Pro and total soluble sugar were dramatically enhanced in \u0026lsquo;WL\u0026rsquo; plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eC,D). Importantly, exogenous application of CK and GA\u003csub\u003e3\u003c/sub\u003e further enhanced the contents of Pro and total soluble sugars in the leaves of \u0026lsquo;CK\u0026thinsp;+\u0026thinsp;WL\u0026rsquo; and \u0026lsquo;GA\u003csub\u003e3\u003c/sub\u003e\u0026thinsp;+\u0026thinsp;WL\u0026rsquo; plants, and this trend was found surprisingly similar following the recovery period (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eC,D). The results of our study clearly indicated that CK-treated waterlogged-stressed mung bean plants were able to accumulate more Pro and soluble sugars than GA\u003csub\u003e3\u003c/sub\u003e-treated plant (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eC,D). Our findings coincided with the results of Sarafraz-ardakani et al. (2014), who also observed that CK supplementation increased the amount of Pro and soluble sugars in drought-stressed wheat plants.\u003c/p\u003e \u003cp\u003eApart from the physiological and biochemical assay, we also focused on the yield-related attributes of mung bean plants to confirm the beneficiary role of the CK and GA\u003csub\u003e3\u003c/sub\u003e application. Waterlogged-stressed mung bean plants showed a decrease in yield-associated features such as pod length, total number of pods per plant, thousand seed weight, and seed yield per plant (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), which might be correlated with poor plant growth because of impaired photosynthesis and enhanced accumulation of ROS (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003eA-C and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA). In contrast, foliar application of either CK or GA\u003csub\u003e3\u003c/sub\u003e to waterlogged-exposed mung bean plants resulted in a dramatic increase in pod length, total number of pods per plant, thousand seed weight, and seed yield per plant (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These results suggested that CK or GA\u003csub\u003e3\u003c/sub\u003e supplementation improved photosynthetic efficiency of mung bean to sustain higher levels of total soluble sugars (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eF), as well as heightened the levels of Pro and non-enzymatic antioxidants to resist the phytotoxic effects of waterlogging through the reductions of ROS-mediated oxidative stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003eA, C-E), resulting in improvement of yield performance (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Importantly, CK-treated waterlogged-stressed mung bean plants provided greater yield-associated features compared with GA\u003csub\u003e3\u003c/sub\u003e-treated plants. Yield improvement in wheat plants under heat and drought stress was also reported upon supplementation of CK and GA\u003csub\u003e3\u003c/sub\u003e, respectively (Gupta et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eIn summary, our results concluded that treating plants with either CK or GA\u003csub\u003e3\u003c/sub\u003e positively regulates defense responses in waterlogged-stressed mung bean plants through regulating several physiological and biochemical mechanisms. In comparison with water-sprayed stressed plants, CK and GA\u003csub\u003e3\u003c/sub\u003e-sprayed plants demonstrated improvement in plant height, stem girth width and plant biomass, which might be interlinked with delayed of photosynthetic pigment destruction, improvement of photosynthetic rate and WUE. Under waterlogged conditions, application of CK and GA\u003csub\u003e3\u003c/sub\u003e successfully decreased the levels of ROS accumulation and consequent MDA level; implying that CK and GA\u003csub\u003e3\u003c/sub\u003e played a decisive role in controlling ROS-induced cellular damage. These observations were closely interlinked to heightened levels of non-enzymatic antioxidants, including total phenolics and total flavonoids. Elevated levels of proline and total soluble sugars in CK and GA\u003csub\u003e3\u003c/sub\u003e-sprayed plants also contributed to the maintenance of better water status and osmotic adjustment under waterlogging. Intriguingly, the positive regulatory roles of CK and GA\u003csub\u003e3\u003c/sub\u003e in the improvement of above-mentioned parameters were also persist after 15-days of recovery period. All of these physiological and biochemical mechanisms in CK and GA\u003csub\u003e3\u003c/sub\u003e-sprayed plants contribute to improve yield-associated features, including pod length, total number of pods per plant, thousand seed weight, and seed yield per plant, in relation of water-sprayed plants. Importantly, plants treated with CK performed better compared with GA\u003csub\u003e3\u003c/sub\u003e-treated plants. Taken together, our findings suggests that either CK or GA\u003csub\u003e3\u003c/sub\u003e could be a potential chemical for mitigating waterlogged-induced detrimental effects on crop performance to ensure sustainable agriculture in flash flood-prone areas. However, together with field trials, comprehensive studies are recommended to decipher the underlying molecular mechanisms that are regulated by either CK or GA\u003csub\u003e3\u003c/sub\u003e under waterlogged conditions.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMRI:\u003c/strong\u003e conceived, designed and supervised the experiment, wrote and edited the manuscript, and gave final approval;\u0026nbsp;\u003cstrong\u003eMMR:\u003c/strong\u003e analyzed and interpreted the data, wrote and revised the manuscript; \u003cstrong\u003eMA and EJ:\u003c/strong\u003e performed the experiment, prepared the materials,\u0026nbsp;collected and analyzed the data;\u003cstrong\u003e\u0026nbsp;SSK:\u003c/strong\u003e helped in\u0026nbsp;statistical analysis, interpretation of the data and results, and writing the manuscript;\u003cstrong\u003e\u0026nbsp;MH:\u003c/strong\u003e monitoring the experiment and reviewed the manuscript.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Competing Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe work was supported by a grant from the Bangladesh Bureau of Educational Information and Statistics (BANBEIS), Ministry of Education, Government of the People\u0026apos;s Republic of Bangladesh under the Higher Educational Research Program.\u003c/p\u003e"},{"header":"References","content":"\u003cp\u003eAhanger, M.A., Alyemeni, M.N., Wijaya, L., Alamri, S.A., Alam, P., Ashraf, M., Ahmad, P., 2018. 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Plant. 148, 228\u0026ndash;236.\u003c/p\u003e"},{"header":"tables","content":"\u003cp style='margin-top:0in;margin-right:0in;margin-bottom:8.0pt;margin-left:0in;line-height:150%;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"color:black;\"\u003eTable 1.\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003cspan style=\"color:black;\"\u003eThe effects of exogenous cytokinin and gibberellic acid on shoot height, shoot dry weight, root dry weight and stem girth width of mung bean plants after 5-days of stress and 15-days of recovery periods\u003c/span\u003e\u003c/p\u003e\n\u003ctable style=\"width:487.7pt;border-collapse:collapse;border:none;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 63.25pt;border: 1pt solid windowtext;padding: 0in 5.4pt;height: 11.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eTreatments\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 106.15pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 11.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eShoot height (cm)\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 106.1pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 11.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eShoot FW (g)\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 106.1pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 11.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eRoot FW (g)\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 106.1pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 11.4pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eStem girth width (mm)\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 55.75pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter stress\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter recovery\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter stress\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter recovery\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter stress\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter recovery\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter stress\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 34.3pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eAfter recovery\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 63.25pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eControl\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.75pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e16.96\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.65\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e33.21\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; \u0026nbsp; \u0026nbsp;1.58\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e2.34\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.09\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e9.68\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.44\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e0.37\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.02\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n 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style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e4.82\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.31\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e7.04\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.04\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n 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none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e1.47\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.09\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e5.42\u0026nbsp;\u003c/span\u003e\u003cspan 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style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e1.22\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.10\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e3.03\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.14\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n 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style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eCK+WL\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.75pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e15.29\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.46\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e28.94\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.58\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e2.03\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.06\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e7.34\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.26\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e0.29\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.01\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e1.65\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.06\u003csup\u003eab\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e4.12\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.08\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e6.41\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.20\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 63.25pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003eGA\u003csub\u003e3\u003c/sub\u003e+WL\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n 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style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e26.62\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.48\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e1.87\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.03\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e6.78\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.30\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e0.27\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.01\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e1.51\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.12\u003csup\u003ebc\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 55.7pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e3.96\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.12\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 50.35pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 10.85pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e5.92\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.18\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:150%;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"color: rgb(0, 0, 0);\"\u003eValues are means \u0026plusmn; standard errors (\u003cem\u003en\u003c/em\u003e = 4). \u003c/span\u003eDifferent alphabetical letters as superscripted within the same column indicate significant differences among various treatments according to a least significant difference test (LSD) at \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05.\u0026nbsp;WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L\u003csup\u003e\u003cspan style=\"font-family:Symbol;color:black;\"\u003e-\u003c/span\u003e\u003cspan style=\"color:black;\"\u003e1\u003c/span\u003e\u003c/sup\u003e) waterlogged-stressed plants; GA\u003csub\u003e3\u003c/sub\u003e+WL, gibberellic acid (GA)-sprayed (50 mg L\u003csup\u003e\u003cspan style=\"font-family:Symbol;color:black;\"\u003e-\u003c/span\u003e\u003cspan style=\"color:black;\"\u003e1\u003c/span\u003e\u003c/sup\u003e) waterlogged-stressed plants; FW, fresh weight.\u003c/p\u003e\n\u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:150%;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"color:black;\"\u003e\u0026nbsp;\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp style='margin-top:0in;margin-right:0in;margin-bottom:8.0pt;margin-left:0in;line-height:150%;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003eTable 2.\u003c/strong\u003e The effects of exogenous cytokinin and gibberellic acid on yield-contributing parameters of stressed and stress-free mung bean plants\u0026nbsp;\u003c/p\u003e\n\u003ctable style=\"border: none;width:478.6pt;border-collapse:collapse;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89.75pt;border: 1pt solid windowtext;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003eTreatments\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 94.5pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003ePod length (cm)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 99pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003eTotal number of pods per plant\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108.9pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003eThousand seed weight (g)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 86.45pt;border-color: windowtext windowtext windowtext currentcolor;border-style: solid solid solid none;border-width: 1pt 1pt 1pt medium;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003eSeed yield (g per plant)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89.75pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;\"\u003eControl\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 94.5pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e8.46 \u0026plusmn; 0.19\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 99pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e33.00 \u0026plusmn; 1.29\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108.9pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e51.10\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 0.86\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 86.45pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13.65pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e16.47 \u0026plusmn; 0.59\u003csup\u003ea\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89.75pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;\"\u003eWL\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 94.5pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e7.35 \u0026plusmn; 0.19\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 99pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e21.00 \u0026plusmn; 0.41\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108.9pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e44.21\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 1.07\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 86.45pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e9.51 \u0026plusmn; 0.30\u003csup\u003ec\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89.75pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;\"\u003eCK+WL\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 94.5pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e8.03 \u0026plusmn; 0.10\u003csup\u003eab\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 99pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e26.00 \u0026plusmn; 1.47\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108.9pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e47.12\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 2.31\u003csup\u003eab\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 86.45pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e11.94 \u0026plusmn; 0.64\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89.75pt;border-color: currentcolor windowtext windowtext;border-style: none solid solid;border-width: medium 1pt 1pt;border-image: none 100% / 1 / 0 stretch;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cstrong\u003e\u003cspan style=\"font-size:15px;\"\u003eGA\u003csub\u003e3\u003c/sub\u003e+WL\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 94.5pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e7.92 \u0026plusmn; 0.09\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 99pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e25.00 \u0026plusmn; 0.41\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 108.9pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;color:black;\"\u003e46.23\u0026nbsp;\u003c/span\u003e\u003cspan style=\"font-size:15px;\"\u003e\u0026plusmn; 1.88\u003csup\u003eab\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 86.45pt;border-color: currentcolor windowtext windowtext currentcolor;border-style: none solid solid none;border-width: medium 1pt 1pt medium;padding: 0in 5.4pt;height: 13pt;vertical-align: top;\"\u003e\n \u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:normal;font-size:16px;font-family:\"Times New Roman\",serif;'\u003e\u003cspan style=\"font-size:15px;\"\u003e11.49 \u0026plusmn; 0.33\u003csup\u003eb\u003c/sup\u003e\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp style='margin-top:0in;margin-right:0in;margin-bottom:0in;margin-left:0in;line-height:150%;font-size:16px;font-family:\"Times New Roman\",serif;'\u003eValues are means \u0026plusmn; standard errors (\u003cem\u003en\u003c/em\u003e = 4). Different alphabetical letters as superscripted within the same column indicate significant differences among various treatments according to a least significant difference test (LSD) (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05).\u0026nbsp;WL, water-sprayed waterlogged-stressed plants; CK+WL, cytokinin (CK)-sprayed (50 mg L\u003csup\u003e\u003cspan style=\"font-family:Symbol;color:black;\"\u003e-\u003c/span\u003e\u003cspan style=\"color:black;\"\u003e1\u003c/span\u003e\u003c/sup\u003e) waterlogged-stressed plants; GA\u003csub\u003e3\u003c/sub\u003e+WL, gibberellic acid (GA)-sprayed (50 mg L\u003csup\u003e\u003cspan style=\"font-family:Symbol;color:black;\"\u003e-\u003c/span\u003e\u003cspan style=\"color:black;\"\u003e1\u003c/span\u003e\u003c/sup\u003e) waterlogged-stressed plants.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"Bangabandhu Sheikh Mujibur Rahman Agricultural University","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Antioxidants, osmoprotectants, phytohormone, pulse crop, reactive oxygen species, yield","lastPublishedDoi":"10.21203/rs.3.rs-690868/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-690868/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMung bean (\u003cem\u003eVigna radiata\u003c/em\u003e) is one of the most important pulse crops, well-known for its protein rich seeds, which growth and productivity are severely undermined by waterlogging. In this study, we aim to evaluate how two promising phytohormones, namely cytokinin (CK) and gibberellic acid (GA\u003csub\u003e3\u003c/sub\u003e), can improve waterlogging tolerance in mung bean by investigating key morphological, physiological, biochemical and yield-related attributes. Our results showed that foliar application of CK and GA\u003csub\u003e3\u003c/sub\u003e under 5-days of waterlogged conditions improved mung bean growth and biomass, which was associated to increased levels of photosynthetic rate and pigments. Waterlogged-induced accumulation of reactive oxygen species, and the consequent elevated levels of malondialdehyde, were considerably reduced by CK and GA\u003csub\u003e3\u003c/sub\u003e treatments. Mung bean plants sprayed with either CK or GA\u003csub\u003e3\u003c/sub\u003e suffered less oxidative stress due to the enhancement of total phenolics and flavonoids levels. Improvement in the contents of proline and total soluble sugars indicating a better osmotic adjustment following CK and GA\u003csub\u003e3\u003c/sub\u003e treatments in waterlogged-exposed plants. Most fundamentally, CK or GA\u003csub\u003e3\u003c/sub\u003e-sprayed waterlogged-stressed mung bean plants demonstrated an increased tendency of the above-mentioned parameters after the 15-day recovery period as compared to water-sprayed waterlogged-exposed plants. Our results also revealed that CK and GA\u003csub\u003e3\u003c/sub\u003e treatments increased yield-associated features in waterlogged-stressed plant. Importantly, both phytohormones are efficient in improving mung bean resistance to waterlogging; however, CK was found to be more effective. Overall, our findings suggested that CK or GA\u003csub\u003e3\u003c/sub\u003e could be used for the management of waterlogging-induced damage in mung bean, and perhaps in other cash crops.\u003c/p\u003e","manuscriptTitle":"Insights into the role of cytokinin and gibberellic acid in improving waterlogging tolerance of mung bean","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2021-07-09 14:38:27","doi":"10.21203/rs.3.rs-690868/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"f9d3497b-d282-4c6f-bb9a-0127e2d4eedd","owner":[],"postedDate":"July 9th, 2021","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":5523747,"name":"Agronomy"}],"tags":[],"updatedAt":"2021-07-09T14:38:27+00:00","versionOfRecord":[],"versionCreatedAt":"2021-07-09 14:38:27","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-690868","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-690868","identity":"rs-690868","version":["v1"]},"buildId":"WrCJVZZCHTDjtuVLN7oU0","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}