Differences of waterlogging tolerance between two grain legume species at germination and reproductive stages

Differences of waterlogging tolerance between two grain legume species at germination and reproductive stages | 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 Differences of waterlogging tolerance between two grain legume species at germination and reproductive stages Md Shahin Uz Zaman, Md Mahamudul Hasan, Lutfun Nahar Luna, Md Shahin Iqbal, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4569640/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 Background and aims Summer pulse crops- mungbean and blackgram are increasingly exposed to waterlogging (WL). This study analyzed the effects of WL at germination and reproductive stages, and to identify the suitable cultivars for different cropping systems. Methods The study evaluated WL tolerance of three mungbean cultivars -BARI Mung-6, BARI Mung-8, Binamoog-8; and a blackgram cultivar- BARI Mash-3 at the germination and reproductive stages. The treatment levels at germination were drained control, 3, 5 and 7 d WL, while at reproductive stage were drained control, 3, 6 and 9 d WL. Results At germination, % emergence was significantly reduced as WL duration increased. After 7 d WL, BARI Mash-3 displayed 38% emergence, while mungbean cultivars had 14–18% emergence. At reproductive-stage, WL reduced plant height, tap root length, shoot and root dry mass compared to drained control. At recovery compared to WL of 9 d, the chlorophyll content increased by 15% in BARI Mash-3, while it decreased in mungbean cultivars. Shoot relative growth rate (RGR) of BARI Mash-3 was positive, while mungbean cultivars showed negative. Similarly, BARI Mash-3 had the higher root RGR than mungbean cultivars. BARI Mash-3 also produced the higher number of adventitious roots (27) than mungbean cultivars (10–19). The seed yield was reduced at 40% in BARI Mash-3 while 52–60% for mungbean cultivars. Conclusions Blackgram cultivar is WL tolerant at both stages compared to mungbean cultivars. These new findings will allow to select suitable crops for different cropping systems based on the perceived risk of WL. Mungean blackgram waterlogging recovery nodulation fibrous roots yield Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction Mungbean ( Vigna radiata L.) and blackgram ( Vigna mungo L.) are two of the most important legume crops in Asia, including Bangladesh, due to their multipurpose use as food, fodder and green manure. The seed of the crops are a good source of proteins and other necessary micronutrients that play an important role in ensuring nutritional security in developing country. Additionally, they have the ability to fix atmospheric nitrogen (58–109 kg ha − 1 ) into soil through a symbiotic relationship with Rhizobium bacteria (Ali 1992 ); which helps in the growth of succeeding crops. More than 7.3 million hectares of land are devoted to mungbean cultivation worldwide, producing around 5.3 million tons annually. India, Bangladesh and Myanmar collectively contribute approximately to 30% of global output (Nair and Schreinemachers 2020 ). In 2021, Bangladesh produced 41,189 million tons (MT) of mungbean and 37,344 MT Blackgram (Bangladesh Bureau of Statistics 2021 ). Mungbean and blackgram are two crops that account for approximately 22.8% of the total area under cultivation for pulses in Bangladesh. These crops are typically grown on marginal lands where other crops are not suitable. Mungbean is usually planted during pre-monsoon season (Late January to March) on poorly drained soil after the harvest of aman rice ( Oryza sativa ). This exposes mungbean to waterlogging (WL) which can be detrimental to its growth and development. On the other hand, blackgram is grown during post-monsoon after the flood water has receded. However, the high soil moisture and subsequent unseasonal rainfall can also expose blackgram to WL at germination and reproductive stages. Moreover, the sudden rainfall as a consequence of climate change causes severe WL stress at different growth phases of mungbean and blackgram (Amin et al. 2014 ). Excessive moisture or water in the soil pores, referred to as waterlogged soil, reduces exchange of gases between root tissues and the atmosphere (van Veen et al. 2014 ). This can quickly lead to a depletion of soil oxygen, resulting in hypoxia and anoxia within a few hours due to continued respiration by plant roots, soil microflora, and fauna (Gambrell and Patrick 1978 ). The lack of oxygen in waterlogged soil can limit nutrient uptake and transport by roots that affects growth and development in wheat ( Triticum aestivum L.) (Malik et al. 2002 ), oats ( Avena sativa L.) (Arduini et al. 2019 ), maize (Ren et al. 2021 ) and cowpea ( Vigna unguiculata L.) (Olorunwa et al. 2023 ). Moreover, soil nitrogen concentration decreases due to rapid volatilization and denitrification processes (Rasaei et al. 2012 ). Furthermore, persistent WL can cause certain ions, such as Mn2 + , Fe2 + accumulation to a harmful level for plants (McKee and McKevlin 1993 ). Mungbean and blackgram are particularly susceptible to soil WL, especially during the early stages of development (Bansal et al. 2019 ; Douglas et al. 2020 ). WL affects plants negatively due to oxygen deficiency, that inhibits root respiration (Jackson and Drew 1984 ). As a result, it causes a decrease in shoot and root growth and nutrient uptake by plants (Malik et al. 2002 ) leading to significant yield losses (Tian et al. 2021 ). Legumes are particularly more sensitive to WL and produce lower seed yield when exposed to WL. For instance, under WL at the reproductive stage yield was decreased by 60% in pea ( Pisum sativum L.) and lupin ( Lupinus albus L.) (Pampana et al. 2016 ), by 35% in chickpea ( Cicer arietinum L.) (Cowie et al. 1996 ), by 48% in cowpea ( Vigna unguiculata L.) (Minchin et al. 1978 ), and by 41–58% in soybean (Oosterhuis et al. 1990 ). Among other crops, significant yield losses (39–44%) have been reported in wheat even with relatively WL-tolerant cultivars (Collaku and Harrison 2002 ). The impact of WL on crop growth and yield varies depending on the crop species and genotype and growth stage (Setter and Waters 2003 ). Significant variation has observed at the germination stage, and waterlogged tolerant varieties have been identified in some dryland crop species such as pigeon pea (Sultana et al. 2013 ), soybean (Hou et al. 1995 ) and maize (Zaidi et al. 2012 ). Similarly, variation is reported at the developmental stages of seedling, vegetative, flowering and pod growth, but the level of tolerance varies depends on the specific growth stage. For instance, the pre-flowering stage of growth in pea is more susceptible to WL than any of the vegetative, flowering and pod-filling stages (Cannell and Gales 1979); whereas in mungbean (Ahmed et al. 2002 ) and chickpea (Cowie et al. 1996 ) the reproductive stage is the most sensitive growth stage. WL tolerance of mungbean and blackgram cultivars have been studied under the varying duration of WL stress at germination and seedling stages (Kyu et al. 2021 ). This study analyzed the effects of WL at different stages of crops on germination, growth and yield components and the physiological response of two economically and nutritionally important legume crops (mungbean and blackgram); and to identify the superior summer pulse crop cultivars for the farmers of the waterlogged affected areas. Materials and methods Plant materials Four most popular cultivars of two summer pulse crops in Bangladesh- Mungbean (BARI Mung-6, BARI Mung-8, Binamoog-8) and Blackgram (BARI Mash-3) were used in this study. All the cultivars are widely grown throughout the country ( Table 1 ). Experimental Conditions The experiments were carried out in a research field at PRC, BARI, Ishwardi, Pabna (24°9′N; 89° 4′E; 19 m a.s.l.) from March 2022 to June 2022. The average temperature ranged from 34 ± 7◦C (day) to 25 ± 4 (night) in both growth stages. The minimum and maximum temperatures ranged 18 to 42 °C respectively in the field. The growing media comprised soil and sand in 2:1 ratio. The soil was collected from PRC research field (30°78′ S, 118°31′ E), with soil pH of 7.8 (1: 5 soil and water suspension), similar to Malik et al. (2016). The soil was sun-dried for a week and sieved to 2 mm diameter. The water content (w/w) at field capacity (i.e. pot capacity when fully drained) was 18%. Experiment A: Waterlogging at Germination Stage The experiment was laid out in a split-plot design with four replications. The main plots were waterlogging (WL) treatments with four levels: drained control, 3 d, 5 d and 7 d WL. The cultivars were in sub-plots. The experimental unit consisted of plastic pots with free draining and sealed base. Free-draining pots contained 100 g gravel at the bottom, followed by 1.0 kg of mixed soil and sand. Each free-draining pots of 0.8 L (90 mm × 90 mm × 180 mm) with drainage holes (∼10 mm in diameter) at the bottom was placed in sealed 60 L plastic tanks. Ten platinum electrodes: five for drained and five for WL pots were placed at a depth of 100 mm to measure soil redox potential. For the waterlogged treatment, tap water was added to sealed pots so that free-draining pots could be waterlogged from the bottom to maintain a water table at soil surface. The pots were waterlogged for 4 d prior to sowing to ensure hypoxia at sowing. Water was added to sealed base pots daily as required to maintain the water table. For drained control treatments, there was no water in the sealed base pots, but the soil moisture in free-draining pots was maintained at ∼80% of field capacity. Seeds were treated with Provax 200FF (Carboxin 17.5% + Thiram 17.5% FF) @ 3 g kg -1 seed to control seed-borne and seedling root pathogens. The healthy seeds of all the cultivars with similar size, color and shape were selected for the experiment. Fifteen seeds of each crop were sown on 16 March, 2022 in a free-draining pot by dibbling at 5 mm soil depth as described by Zaman et al. (2019). After each duration of WL treatment pots were allowed to drain. Soil redox potential was measured daily using platinum electrodes (Pt) and an Ag–AgCl reference electrode with a handheld Digital Multimeter (Fluke 114, Everett, Washington, USA). Redox measurements were corrected according to the method developed by Patrick et al. (1996). Seeds with an epicotyl >5mm were recorded as emergence. Emergence was recorded every alternate day for each seed in both the waterlogged and recovery periods. The experiment was terminated at 22 d after sowing, when there was no sign of further emergence. Experiment B: Waterlogging at Reproductive Stage The experiment comprised of four treatments: drained control and 3 d, 6 d and 9 d WL). The design was factorial with crop (4 levels) x WL treatment (4) in a completely randomized design with four replications. The experimental unit was a plastic pot—free draining and sealed base. Each free draining pot contained 300 g gravel at the bottom followed by 5 kg mixed soil and sand. Each free draining pot of 8 L (145 mm×145 mm×220 mm) with 15 mm drainage holes at the bottom was placed in sealed based 10 L (310 mm × 620 mm × 455 mm) plastic pot. Ten platinum electrodes: five each for drained and for WL pots were placed at 100 mm depth to measure soil redox potential. For the waterlogged treatment, tap water was added to sealed pots so that free-draining pots could be waterlogged from the bottom to maintain a water table at soil surface. Water was added to sealed base pots daily as required to maintain the water table. For drained control treatments, there was no water in the sealed base pots, but the soil moisture in free-draining pots was maintained at ∼80% of field capacity. Five seeds were sown in each pot by dibbling at 5 mm soil depth on 17 March 2022. The plants were thinned to two at seven days after sowing. The WL treatments were started 5-7 d before flowering at 30 DAS. The 9 days WL treatment was imposed on 16 April 2022 at 30 DAS, the 6 days WL treatment was 3 days later, and so on for 3 days WL treatment. This approach allowed the same stress release time for all treatments (Zaman et al. 2024; Wang et al. 2017) and replicated a common recovery and drying-out period, thereby enabling to measures to morphological and physiological parameters during recovery. All the Waterlogged pots were drained on 25 April 2022 at 40 DAS. Four pots of each treatment were harvested at the time of draining to measure the plant growth; and the other four pots of each treatment were harvested 28 days after draining, during the recovery. The experiment was terminated at 68 DAS. The redox potential of the soil was measured daily in the WL and drained pots. The chlorophyll content was measured using a handheld Minolta SPAD 502 (Konica-Minolta, Japan) on the first trifoliate leaf of each plant at two points in time: at the end of WL at 40 DAS and 14 days after draining in recovery. The measurements were taken on the same leaf at both WL and recovery period. At harvest, plant height was measured from the collar (point on the stem where roots start to grow) to the leaf base of the youngest fully expanded leaf of the plant. After gently washing the soil from the roots, the maximum taproot length was measured. The number of emerged adventitious roots longer than 5mm were recorded, and their length recorded at the end of WL treatment. Although the soil was not inoculated with rhizobium, root nodulation was scored at 40 DAS by counting the nodules on the main taproot and lateral roots and using a 0–8 scoring scale according to Yates et al. (2016), where 0=no nodules, 0.5=white ineffective nodules, 1 = rare effective, 2 = scarce, 3 = moderate, 4 = adequate, 5 = ample, 6 = abundant, 7 = very abundant, and 8 = extremely abundant. Finally, the plants were divided into shoots and roots and dried in paper bags in at 60°C oven for 3 days to record shoot and root dry mass. Relative growth rate was measured at the time of draining to 28 days after recovery. The data for each pot (2 plants in each pot) were pooled and the mean was used as one replicate. Pods per plant and seed yield were recorded at recovery. Statistical analysis Data recorded for different parameters were compiled and tabulated in proper form for statistical analysis which was carried out in Minitab, LLC. (2021) statistical software package and R Ver 4.2.1 (R Core Team, 2020). Various statistical tests such as, mean performance, two-way ANOVA (Analysis of Variance were executed for different morpho-physiological traits to explore treatment, genotype and treatment x genotype interaction. For all analyses, the means were separated based on their significance levels at 0.05 probability using the Tukey test (Tukey 1949). The relative growth rate (RGR) for shoot and root was calculated according to Hunt (1990), RGR = {ln ( W 2) −ln ( W 1)} ( t 2− t 1), where W 1 and W 2 are shoot/root dry mass at times t 1 and t 2 , respectively. Results Waterlogging Tolerance at Germination Stage In drained control soil the redox potential was 559 ± 11 mV throughout the experimental period. By contrast, the redox potential in waterlogged pots at the time of seed sowing to draining period was 337 ± 12 mV and this increased on draining the pots to 588 ± 15 by 22 d. The analysis of variance showed that WL duration, cultivars and their interaction was highly significant (P < 0.001) for emergence (Supplemental Table 1). All the crop cultivars showed close to 100% emergence in drained soil. However, in waterlogged soil crop exhibited contrasting responses in emergence (Fig. 1) . Seedling emergence was reduced significantly by increased WL duration (Fig. 2) . Three days of WL reduced seedling emergence to 18% for BARI Mash-3, 24% for BARI Mung-6, 30% for Binamoog-8 and 34% for BARI Mung-8 by the end of the experiment. With the increase of WL duration to 7 d, emergence percentage was further reduced for BARI Mash-3 to 61%, for BARI Mung-6 to 83%, for Binamoog-8 to 81% and for BARI Mung-8 to 86%. Waterlogging Tolerance at Reproductive Stage Soil redox potential At the start of the WL treatment, soil redox potential in the drained control pots was 466 ± 12 mV which was almost similar throughout the experimental period. During WL, soil redox potential rapidly decreased to 237 ± 15 mV at two days after waterlogging that remained almost similar at the end of treatment duration. Upon recovery, the soil redox potential increased to 452 ± 9 mV within 7 days - almost similar to drained control soil. Effect on shoot growth There was significant variation among the waterlogging (WL) treatments, cultivars and their interaction for the traits in both WL and recovery phases (Table 2). During WL phase (after draining of 40 DAS), all the cultivars showed the highest shoot length (i.e. plant height) in the drained control as compared WL treatments (Fig. 3A) . With the lowest duration of WL (3 d), the shoot length was reduced by 3, 3.2, 10 and 3.4% in BARI Mung-6, BARI Mung-8, Binamoog-8 and BARI Mash-3, respectively, compared to drained control. When subjected to the longer duration of WL stress (9 d), the shoot length of BARI Mung-6 was decreased by 16.4%, BARI Mung-8 by 20.6% in, Binamoog-8 by 14.4% in and BARI Mash-3 by 8.2% as compared to respective drained control. During recovery phase (68 DAS), drained control showed the highest shoot length compared to waterlogged treatments. The highest waterlogged treatments at 9 d, BARI Mung-6 had the least effect on shoot length (i.e. decreased by 15%) followed by BARI Mash-3 (i.e. decreased by 16%) as compared to drained control (Fig. 3A) . Shoot dry mass varied depending on cultivars and treatments. During WL phase at 40 DAS, the drained control had higher shoot dry mass for all the cultivars compared to WL treatments. BARI Mung-6 had the highest dry mass (4.42 g plant -1 ) followed by BARI Mash-3 (3.95 g plant -1 ), while Binamoog-8 (3.79 g plant -1 ) had the lowest. However, at the highest level of WL (9 d), shoot dry mass was reduced by 28% in BARI Mash-3, 31% in BARI Mung-6, 34% in BARI Mung-8, and 36% in Binamoog-8 relative to the drained controls ( Fig. 3B ). During the recovery phase (68 DAS), the drained controls showed higher shoot dry mass in comparison to the WL treatments. After recovering from 9 d WL, BARI Mash-3 had the lowest reduction of shoot dry mass (36%) indicating the potential ability to recover from the damage of WL stress during the subsequent period of drainage. In contrast, the highest shoot dry mass reduction (53%) was in Binamoog-8 ( Fig. 3B ). The relative growth rate (RGR) of shoots was compared between two periods- at the time of draining of WL pots (40 DAS) and 28 days after draining (68 DAS). The RGR of shoots in drained control was positively increased for all the cultivars across the treatments, with the highest increase in BARI Mash-3. However, in waterlogged treatment, RGR of shoot was positive in BARI Mash-3 while it was negative for BARI Mung-6, BARI Mung-8 and Binamoog-8 across the WL treatments ( Fig. 3C ). At the highest level of 9 d WL, shoot relative growth rate of BARI Mash-3 was 111 mg -1 plant -1 d -1 in compared to 190 mg -1 plant -1 d -1 in drained control. In contrast, BARI Mung-6, BARI Mung-8 and Binamoog-8 showed a negatively decreased RGR of -5, -8 and -9 mg -1 plant -1 d -1 than drained control of 17, 16 and 27 mg -1 plant -1 d -1 respectively. Effect on root growth Waterlogging (WL) had a negative impact on the tap root growth for all the cultivars during WL and could not recover by end of the experimental period. Moreover, some loss of or decay of tap root was observed during recovery phase as evident by the shorter root length by the end of the experiment. During WL phase (40 DAS), all the cultivars in WL treatment exhibited the higher level of root length in compared to drained control (Figure 4A) . After 3 d of WL treatment, there was no remarkable reduction of root length in BARI Mung-6 compared to drained control. However, BARI Mash-3, BARI Mung-8 and Binamoog-8 experienced a reduction of 7.2%, 8.3% and 12.2%, respectively, compared to drained control. As the WL duration increased to 9 d, the reduction of root length was more significant, with BARI Mash-3 a 17%, BARI Mung-8 a 17.2%, Binamoog-8 a 20% and BARI Mung-6 a 22.5% reduction compared to drained control. During recovery phase (68 DAS), all cultivars experienced secession of the tap root growth as evidenced by the reduced length in waterlogged treatments compared to drained control (Figure 4A) . After 3 d of WL treatment, BARI Mash-3 had the smallest reduction of root length, with only a 18.6% decrease compared to drained control. However, BARI Mung-8, BARI Mung-6 and Binamoog-8 experienced a reduction of 34%, 36%, 37%, respectively, compared to drained control. As the WL duration increased to 9 d, the reduction of root length was more significant, with BARI Mung-8 a 50%, BARI Mung-6 a 55%, Binamoog-8 a 59% and BARI Mash-3 a 64% reduction compared to drained control. The effect of WL was highly significant (P < 0.001) on root dry mass for all cultivars ( Table 2 ). During the WL phase at 40 DAS, root dry mass was reduced by 27% in BARI Mash-3, 33% in BARI Mung-8, 43% in BARI Mung-6 and 47% in Binamoog-8 at 3 d WL treatment, relative to the drained controls; the values in the 9 d WL treatment were 47% in BARI Mash-3, 63% in BARI Mung-8, 65% in Binamoog-8 and 74% in BARI Mung-6 (Figure 4B) . During recovery phase (68 DAS) of 3 d WL, BARI Mash-3 exhibited the lowest reduction (23%) of root dry mass followed by BARI Mung-8 at 61% with its drained control (Figure 4B) . With the increased WL duration at 9 d the reduction of root dry mass was 46% in BARI Mash-3, 50% in BARI Mung-8, 64% in BARI Mung-6 and 70% in Binamoog-8. The relative growth rate (RGR) of roots for all the varieties in waterlogged treatments was decreased in compared to drained control ( Fig. 4C ). During 9 d WL, BARI Mash-3 exhibited the highest root RGR of 27 mg -1 plant -1 d -1 while in 7 mg -1 plant -1 d -1 for BARI Mung-6, 7.5 mg -1 plant -1 d -1 for BARI Mung-8 and -5 mg -1 plant -1 d -1 for Binamoog-8 compared to 55, 10, 7 and 19 mg -1 plant -1 d -1 respectively in drained control. Adventitious root formation Adventitious roots were not observed in any of the drained control plants. However, all the cultivars developed adventitious roots near the shoot-root junction (hypocotyl region) in WL treatments and the length ranged from 0.45 to 0.80 mm after 3-9 days of WL. ( Fig. 5A ). The number of adventitious roots increased with the increase of WL duration and varied within cultivars. After 3 day of WL, BARI Mash-3 exhibited the highest number (3.6) of adventitious roots followed by BARI Mung-8 (2.6). During 9 days of WL, the number of adventitious roots was increased to 27 in BARI Mash-3 followed by Binamoog-8 (19), while the lowest in BARI Mung-6 (10). Nodule formation In drained control, all the cultivars showed higher number of nodulation score compared to waterlogged soil, and the number and size of the nodule increased with crop growth ( Fig. 5B ). Binamoog-8 exhibited the higher score of nodulations (7.6), but the biggest size of nodules was observed in BARI Mash-3 in compared to other crop in drained soil. In waterlogged soil, the number of nodules decreased with the increasing duration of WL. The shortest WL period of 3 d did not affect nodule formation in all the cultivars compared to its drained control. With the increase of WL duration to 9 d, nodule score was decreased by 42% in BARI Mash-3, 57% in Binamoog-8, 72% in BARI Mung-6 and 86% in BARI Mung-8. In comparison of crops, the nodule formation in BARI Mash-3 were less affected than other mungbean cultivars. All the crop produced nodules on lateral and adventitious roots near the soil surface during WL, but the plants in drained control had nodules in lower the soil surface. The color of the nodules was brown in both drained control and WL indicating that they were functional. Effect on chlorophyll content The chlorophyll content (SPAD value) of the first trifoliate leaves were recorded at WL (at the time of draining/40 DAS) and 14 days after recovery phase (54 DAS) to understand the effect of WL on photosynthesis. In the drained control the chlorophyll content in both WL and recovery phase was higher than waterlogged plants for all the cultivars, but chlorophyll content at recovery was reduced compared to WL due to ageing the plants (Fig. 6) . For instance, the highest SPAD value (55) of BARI Mung-8 in drained control during WL phase was decreased to 47 at recovery phase. Among the waterlogged treatments, the SPAD value in recovery phase was increased in BARI Mash-3, while decreased in all the mungbean cultivars across the treatments than WL phase. At the highest level of WL treatments at 9 d, the SPAD value was increased 15% in BARI Mash-3 at recovery phase in compared to WL phase. In contrast, the SPAD value decreased 11% in BARI Mung-6, 21% in Binamoog-8 and 32% in BARI Mung-8 at 9 d WL during the recovery phase. Effect on pod formation and seed yield Exposer to waterlogging during reproductive stage had a negative impact on both pod number plant -1 and seed yield plant -1 (Fig. 7) . However, it did not affect pod length and seeds pod -1 ( (Fig. 7A&B) . In the drained control, BARI Mash-3 produced the highest number of pods plant -1 (57) followed by BARI Mung-8 (46). When exposed to the shortest duration of (3 d) WL treatment, the number of pods plant -1 was reduced by 19% in Binamoog-8, 43% in BARI Mash-3, 50 % in BARI Mung-6 and 49% in BARI Mung-8 compared to drained controls. During 9 d WL, the reduction of number of pods plant -1 was the highest in BARI Mung-8 (74%) while in other cultivars it ranged from 50-59% compared to drained controls (Fig. 7C) . The waterlogging drastically affected the yield of all cultivars (Fig. 7D) . The longer the waterlogging duration during the reproductive stage, the greater the yield losses. In the drained control, seed yield plant -1 ranged from 5.8 g in BARI Mash-3 to 9.0 g in BARI Mung-8. The shortest duration of 3 d WL reduced seed yield plant -1 by 34% in Binamoog-8, 41% in BARI Mash-3, 52% in BARI Mung-8 and 54 % in BARI Mung-6 compared to drained controls. During 9 d WL treatment, the reduction of seed yield plant -1 was 59% in BARI Mung-8, 58% in BARI Mung-6, 52% in Binamoog-8 and 40% in BARI Mash-3 compared to drained controls. Discussion This study investigated the tolerance of two summer grain legumes to waterlogging at the germination and reproductive stages, highlighting the importance of implementing appropriate measures to prevent waterlogging in fields for optimal crop growth and yield. a. Variation in waterlogging tolerance at germination The findings of the study indicate that exposing legumes cultivars to waterlogging conditions for a short period can have a significant impact on its growth in the long term and ultimately affects emergence and yield. During the early stages of germination in control treatment, it was observed that all cultivars of both species exhibited more than 90% emergence within four days of sowing. This initial emergence was consistent across all cultivars studied, indicating that the germination process is not significantly affected by genetic differences between cultivars. However, when the plants were subjected to waterlogging treatments, emergence was delayed and reduced, indicating that waterlogging has a significant impact on the germination process. In the waterlogged treatment, emergence only began after nine days of sowing and was completed by the 16th day. This delay in emergence indicates that waterlogging has a significant impact on the germination process and can lead to slower growth and development of plants. The delay in emergence is likely due to the fact that waterlogging creates an anaerobic environment in the soil, which leads to reduced oxygen availability for the developing seeds (Zaman et al. 2018 in peas). The insufficient oxygen levels prevent the respiration which is required for germination (Powell and Matthews 1978 , wheat and barley; Hou and Thseng 1992 , Soybean; Xiong et al. 2020 , maize). This lack of respiration inhibits the growth and development of the seeds, leading to slower emergence and reduced plant growth (Wiraguna et al. 2017 , lentil; Kyu et al. 2021 , mungbean). Such information is vital for farmers and researchers alike, as it can help them make informed decisions about crop management strategies that can improve plant growth under challenging environmental conditions. The longer the WL duration, the greater the reduction in seedling emergence. In the present study, with the longest WL duration of 7 days, emergence was reduced by 70%. However, there was a variation of emergence among the cultivars. The dark testa BARI Mash-3 exhibited the higher percent of emergence than shiny green and yellow mungbean cultivars which is consistent with previous studied where the dark testa genotypes of peas (Zaman et al. 2019 ), soybean (Hou and Thseng 1991 ), rapeseed (Zhang et al. 2008 ) and wheat (Ueno and Takahashi 1997 ) were found to be tolerant to WL compared to less pigmented seeds of respective crops. The WL tolerance of dark seeded genotypes is presumably due to the presence of higher levels of phenolic compounds compared to the sensitive yellow seeded lines (Zhang et al. 2008 ). The phenolic compounds are responsible for preventing testa leakage in the tolerant genotypes. Conversely, the testa is leaked and whitish seed solutes are observed on the seed surface of susceptible genotypes in absence of phenolic compounds (Zaman et al. 2019 ). In this study the black testa BARI Mash-3 was tolerant as in other crops but the shiny green testa mungbean cultivars showed susceptibility. b. Effect of waterlogging on growth After emerging, plant root systems undergo development, allowing plants to use energy through root respiration for optimal growth and development in the presence of oxygen in soil pores. In the waterlogged condition, the respiration is limited due to absence of oxygen that reduces root length in mungbean (Kumar et al. 2013 ), soybean (Miao et al. 2012 ), wheat (Herzog et al. 2016 ), maize (Ren et al. 2016 ) and in current study (Fig. 4 a). However, the WL tolerant genotypes produced adventitious roots (ARs) as the primary roots are damaged (Qi et al. 2020 ). The development of ARs has been observed in many species- sunflower ( Helianthus annuus L.) (Jackson 1955 ), rice (Lorbiecke and Sauter 1999 ), wheat (Yamauchi et al. 2014 ), maize (Mano et al. 2005 ), Rumex spp. (Visser et. Al. 1996 ), and tomato ( Solanum lycopersicum ; Vidoz et al. 2010 ). The newly developed ARs emerge from the hypocotyl, mesocotyl, or base of the stem and contain more aerenchyma than primary roots (Visser and Voesenek 2004). Consequently, ARs enhance gas exchange and water and nutrient uptake in plants, which contribute greatly to plant tolerance of poorly aerated soils (Steffens and Rasmussen 2016 ). The number of adventitious roots is increased with the increase of WL duration in mungbean and blackgram (Kyu et al. 2021 ; Kumar et al. 2013 ), cucumber (Xu et al. 2018 ), wheat (Koramutla et al. 2022 ) and yard long bean (Nawata et al. 1991 ). In this study, with the longest WL of 9 days, the highest number of adventitious roots was found in BARI Mash-3 while the lowest in all the mungbean cultivars during WL. However, there was no such root development in the drained controls. In a drained control soil, legume crops acquire nitrogen mostly through symbiotic nitrogen fixation in rhizobia nodules. Under WL conditions, as soil condition shifts from hypoxia to anoxia, there is reduced O 2 for the rizhobium . Thus, with prolonged WL, the number of rhizobia decrease due to unavailable of soil oxygen. In the present study, all the crops showed the higher number of nodules in drained control than WL treatments. During WL, the variation of nodulation was observed among the crops. After 9 days WL of this study, nodulation had decreased by 42% in BARI Mash-3, 57% in Binamoog-8, 72% in BARI Mung-6 and 86% BARI Mung-8. This suggests that the nodulation might have an influence for survival ability of crops during WL stress. Previous research indicated that in legumes, tissues of stem, root and nodule are connected by a well- developed network of aerenchyma that allow oxygen diffusion to nodules (Roberts et al. 2010 ). In soybean, the aerenchyma connects to the outer cortex of nodules, presumably, supporting their functioning for roots in WL soils (Shimamura et al. 2003 ; Thomas et al. 2005 ). Hence, the crop with more nodules have a beter ability to thrive in waterlogged soil. Nodules located below 5–10 cm of WL soil might have become ineffective as their color changed from pink to white (Roberts et al. 2010 ). In the current study, pink nodules continued to develop at the at the base of the soil with adventitious roots regardless of the duration of WL, suggesting that they were able to function during the recovery phase. In soybean, the aerenchyma attaches to the outer cortex of nodules, possibly allowing them to serve as roots in WL. soils (Shimamura et al. 2003 ; Thomas et al. 2005 ). The damage of the root systems under WL affected the shoots growth, through a reduced number of leaves and a smaller leaf area. Limitation of nutrient uptake following a longer duration of WL might be the cause of the reduction in shoot that has been reported in previous studies (Bacanamwo and Purcell 1999 - soybean; Malik et al. 2002 - wheat; Board 2008 - soybean; Malik et al. 2015 - pea, lentil, and grass pea; Najeeb et al. 2015 - cotton). Moreover, WL causes leaching of nutrient e.g., N, P and K, causing nutrient shortage symptoms thus resulting in shoot length reduction (Rhine et al. 2010 ). In this study, all cultivars showed significant variation in plant height and reduced shoot length under WL. With the increasing duration of WL shoot length also decreases. Highest shoot length reduction was observed in BARI Mung-6 and BARI Mash-3 following the WL recovery. The symptom of affecting shoot growth is visualized with leaf chlorosis. In the present study, chlorophyll content (SPAD value) decreased under WL – compared to drained control - for all the crop, indicating the degradation of chlorophyll in functional leaves and a reduction in photosynthetic capacity (Barickman et al. 2019 ). During the recovery period, SPAD chlorophyll increased in BARI Mash-3 for all the treatments; but was further decreased in mungbean cultivars compared to WL period. Chlorophyll content (greenness of leaves) is an important parameter to screen pulses for WL tolerance. Similarly, reduction in chlorophyll content under WL has been reported in blackgram (Bansal et al. 2022 ), mungbean (Kyu et al. 2021 ; Kumar et al. 2013 ), sesame (Mensah et al. 2006 ), wheat (Collaku and Harrison 2002 ) and maize (Prasad et al. 2004 ). Under WL, yellowing of the plant may be caused by a reduction in leaf nitrogen (Bacanamwo and Purcell 1999 ), nodulation and N fixation, and the production of toxic substances such as nitrites and sulfides that are carried upward in large quantities from the soil through the roots and into the leaf (Ezin et al. 2010 ). In addition, WL reduces nitrogen in the soil by accelerated volatilization and denitrification (Rasaei et al. 2012 ). c. Effect on waterlogging on yield The shoot and root dry matter significantly decreased under WL stress by damaging the existing root systems of mungbean and blackgram cultivars which is consistent to other studies in mungbean (Ahmed et al. 2002 ; Kumar et al. 2013 ); chickpea, fababean (Munir et al. 2019 ); soybean (Miao et al. 2012 ; Beutler et al. 2014 ; Kim et al. 2019 ) and wheat (Malik et al. 2002 ). Relative Growth Rate (RGR) of shoot was higher in BARI Mash-3 compared to drained control, however, it was decreased to all the mungbean cultivars for all the WL treatments indicating contrasting response of crop, tolerant- BARI Mash-3 and sensitive- mungbean cultivars. The response was clearest during the highest level of WL at 9 days. Similarly, the RGR of roots was higher in BARI Mash-3 in compared to drained control across the treatments. At 9 days WL, BARI Mash-3 exhibited 3-fold higher RGR than drained control. In contrast, RGR of roots in mungbean cultivars was negative similar to the WL response seen in rapeseed waterlogged for 14 days (Brisson et al. 2002 ; Ploschuk et al. 2018 ). Finally, WL directly affects on pod formations and ultimate yield. The of number of pods plant − 1 was gradually declined with increasing WL duration. As plants most energy distributes to the formation of adventitious root and dry matter to survive through the WL stress thus least amount of energy, sugar, metabolites were left for the yield and yield related characters such as pod setting, pod filling and seed formation (Kyu et al. 2021 ; Kumar et al. 2013 ). This decreases the yield 40–60% in compared to drained control for the cultivars with the lowest (40%) in BARI Mash-3. Previous studies also showed the reduction of yield in different crops such as soybean (Miao et al. 2012 ; Beutler et al. 2014 ), maize (Tian et al. 2019 ), wheat and barley (De San Celedonio et al. 2014), under WL. This indicates that, BARI Mash-3 have the higher ability to grow WL than those of mungbean cultivars. Conclusion Waterlogging at different growth stage is one of the major constraints to pulse crop in rice-based cropping. This study has demonstrated the variation of waterlogging tolerance between two summer grain legumes. BARI Mash-3 showed the higher ability to grow in waterlogging at both germination and reproductive stages in compared to the mungbean cultivars. Seed yield was decreased to all the cultivars of both species, but the lowest reduction (40%) was found in BARI Mash-3, while the highest (59%) in BARI Mung-8. It indicated that blackgram is more suited to waterlogging condition. This differences of waterlogging response with stages and genotypes will help to select crops to alleviate the waterlogging stress. Declarations Acknowledgements This work has been part of the project Incorporating salt-tolerant wheat and pulses into smallholder farming systems in southern Bangladesh (CIM/2014/076) funded by Australian Centre for International Agriculture Research (ACIAR) and Krishi Gobeshona Foundation (KGF), Bangladesh in collaboration with the Pulses Research Centre (PRC), Bangladesh Agricultural Research Institute, Bangladesh. The authors are thankful to the authorities of ACIAR, KGF and PRC for all sorts of supports and facilities. We are also grateful to the staff at PRC, Ishwardi, Pabna, Bangladesh for all their assistance during the field trials. Funding : There is no funds, grants, or other support received during the preparation of this manuscript. Competing Interests: We have no relevant financial or non-financial interests. Author Contributions: All authors contributed to the study conception and design. Material preparation, data collection and analysis were performed by Md Shahin Uz Zaman and Md Mahamudul Hasan. The first draft of the manuscript was written by Md Shahin Uz Zaman and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript. Data Availability: The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request. References Ahmed S, Nawata E, Sakuratani T (2002) Effects of waterlogging at vegetative and reproductive growth stages on photosynthesis, leaf water potential and yield in mungbean. Plant Prod Sci 5:117–123. https://doi.org/10.1626/pps.5.117 Ali M (1992) Weeds are a great threat to kharif pulses. Indian farming 42:27–29 Amin M, Amiruzzaman M, Ahmed A, Ali M (2014) Combining ability study in waterlogged tolerant maize ( Zea mays L.). Bangladesh J Agric Res 39:283–291. https://doi.org/10.3329/bjar.v39i2.20430 Arduini I, Baldanzi M, Pampana S (2019) Reduced growth and nitrogen uptake during waterlogging at tillering permanently affect yield components in late sown oats. Front Plant Sci 10:1087. https://doi.org/10.3389/fpls.2019.01087 Bacanamwo M, Purcell LC (1999) Soybean dry matter and N accumulation responses to flooding stress, N sources and hypoxia. Journal of Experimental Botany 50: 689-696. Bangladesh Bureau of Statistics (2021) Yearbook of agricultural statistics. Ministry of planning, government of the people’s republic Bangladesh, Dhaka Bansal R, Dikshit H, Singh A, et al (2022) Effect of waterlogging on physiological traits and yield in blackgram ( Vigna mungo L.) in field condition. Indian J Biochem Biophys 59:580–585. https://doi.org/10.56042/ijbb.v59i5.63013 Bansal R, Sharma S, Tripathi K, Kumar A (2019) Waterlogging tolerance in blackgram [ Vigna mungo (L.) Hepper] is associated with chlorophyll content and membrane integrity. Indian J Biochem Biophys 56: Barickman TC, Simpson CR, Sams CE (2019) Waterlogging causes early modification in the physiological performance, carotenoids, chlorophylls, proline, and soluble sugars of cucumber plants. Plants 8:160. https://doi.org/10.3390/plants8060160 Beutler AN, Giacomeli R, Alberto CM, et al (2014) Soil hydric excess and soybean yield and development in Brazil. Aust J Crop Sci 8:1461–1466 Board JE (2008) Waterlogging effects on plant nutrient concentrations in soybean. Journal of Plant Nutrition 31:828–838. https://doi.org/10.1080/01904160802043122 Brisson N, Rebiere B, Zimmer D, Renault P (2002) Response of the root system of a winter wheat crop to waterlogging. Plant and soil 243: 43-55. Cannell RQ, Gales K, Snaydon RW, Suhail BA (1979) Effects of short‐term waterlogging on the growth and yield of peas ( Pisum sativum ). Annals of Applied Biology 93:327–335. https://doi.org/10.1111/j.1744-7348.1979.tb06549.x Collaku A, Harrison SA (2002) Losses in wheat due to waterlogging. Crop Science 42: 444-450. Cowie AL, Jessop RS, MacLeod DA (1996) Effects of waterlogging on chickpeas II. Possible causes of decreased tolerance of waterlogging at flowering. Plant Soil 183:105–115. https://doi.org/10.1007/BF02185570 Douglas C, Pratap A, Rao BH, et al (2020) Breeding progress and future challenges: Abiotic stresses. In: Nair RM, Schafleitner R, Lee S-H (eds) The Mungbean Genome. Springer International Publishing, Cham, pp 81–96. https://doi.org/10.1007/978-3-030-20008-4_6 Ezin V, Pena RDL, Ahanchede A (2010) Flooding tolerance of tomato genotypes during vegetative and reproductive stages. Braz J Plant Physiol 22:131–142. https://doi.org/10.1590/S1677-04202010000200007 Gambrell RP, Patrick WH Jr (1978) Chemical and microbiological properties of anaerobic soils and sediments. In: Hook DD, Crawford RM (eds) Plant life in anaerobic environments. Ann Arbor Science, Ann Arbor, pp 375–423 Herzog M, Striker GG, Colmer TD, Pedersen O (2016) Mechanisms of waterlogging tolerance in wheat – a review of root and shoot physiology. Plant Cell & Environment 39:1068–1086. https://doi.org/10.1111/pce.12676 Hou FF, Thseng FS (1991) Studies on the flooding tolerance of soybean seed: varietal differences. Euphytica 57:169–173. https://doi.org/10.1007/BF00023075 Hou F-F, Thseng F-S (1992) Studies on the screening technique for pre-germination flooding tolerance in soybean. Jpn J Crop Sci 61:447–453. https://doi.org/10.1626/jcs.61.447 Hou F-F, Thseng F-S, Wu S-T, Takeda K (1995) Varietal differences and diallel analysis of pre-germination flooding tolerance in soybean seed. Bull Res Inst Bioresour Okayama Univ 3:35–41 Hunt R (1990) Relative growth rates. In: Hunt R (ed) Basic growth analysis: Plant growth analysis for beginners. Springer Netherlands, Dordrecht, pp 25–34 Jackson MB, Drew MC (1984) Effects of flooding on growth and metabolism of herbaceous plants. In: Flooding and Plant Growth. Elsevier, pp 47–128 Jackson WT (1955) The role of adventitious roots in recovery of shoots following flooding of the original root systems. American J of Botany 42:816–819. https://doi.org/10.1002/j.1537-2197.1955.tb10428.x Kim KH, Cho MJ, Kim J-M, et al (2019) Growth response and developing simple test method for waterlogging stress tolerance in soybean. J Crop Sci Biotechnol 22:371–378. https://doi.org/10.1007/s12892-019-0271-0 Koramutla MK, Tuan PA, Ayele BT (2022) Salicylic acid enhances adventitious root and aerenchyma formation in wheat under waterlogged conditions. IJMS 23:1243. https://doi.org/10.3390/ijms23031243 Kumar P, Pal M, Joshi R, Sairam RK (2013) Yield, growth and physiological responses of mungbean[ Vigna radiata (L.) Wilczek] genotypes to waterlogging at vegetative stage. Physiol Mol Biol Plants 19:209–220. https://doi.org/10.1007/s12298-012-0153-3 Kyu KL, Malik AI, Colmer TD, et al (2021) Response of mungbean (cvs. Celera II-AU and Jade-AU) and blackgram (cv. Onyx-AU) to transient waterlogging. Front Plant Sci 12:709102. https://doi.org/10.3389/fpls.2021.709102 Lorbiecke R, Sauter M (1999) Adventitious root growth and cell-cycle induction in deepwater rice1. Plant Physiology 119:21–30. https://doi.org/10.1104/pp.119.1.21 Malik AI, Ailewe TI, Erskine W (2015) Tolerance of three grain legume species to transient waterlogging. AoB PLANTS 7. https://doi.org/10.1093/aobpla/plv040 Malik AI, Ali MO, Zaman MS, et al (2016) Relay sowing of lentil ( Lens culinaris subsp. culinaris) to intensify rice-based cropping. J Agric Sci 154:850–857. https://doi.org/10.1017/S0021859614001324 Malik AI, Colmer TD, Lambers H, et al (2002) Short‐term waterlogging has long‐term effects on the growth and physiology of wheat. New Phytologist 153: 225-236. Mano Y, Muraki M, Fujimori M, et al (2005) Identification of QTL controlling adventitious root formation during flooding conditions in teosinte (Zea mays ssp. huehuetenangensis) seedlings. Euphytica 142:33–42. https://doi.org/10.1007/s10681-005-0449-2 McKee WH, McKevlin MR (1993) Geochemical processes and nutrient uptake by plants in hydric soils. Enviro Toxic and Chemistry 12:2197–2207. https://doi.org/10.1002/etc.5620121204 Mensah JK, Obadoni BO, Eruotor PG, Onome-Irieguna F (2006) Simulated flooding and drought effects on germination, growth, and yield parameters of sesame ( Sesamum indicum L.). African Journal of biotechnology 5:1249-1253. Miao S, Shi H, Jin J, et al (2012) Effects of short-term drought and flooding on soybean nodulation and yield at key nodulation stage under pot culture. J Food Agric Env 10:819–824 Minchin FR, Summerfield RJ, Eaglesham ARJ, Stewart KA (1978) Effects of short-term waterlogging on growth and yield of cowpea ( Vigna unguiculata ). J Agric Sci 90:355–366. https://doi.org/10.1017/S0021859600055465 Munir R, Konnerup D, Khan HA, et al (2019) Sensitivity of chickpea and faba bean to root‐zone hypoxia, elevated ethylene, and carbon dioxide. Plant Cell & Environment 42:85–97. https://doi.org/10.1111/pce.13173 Nair R, Schreinemachers P (2020) Global status and economic importance of mungbean. In: Nair RM, Schafleitner R, Lee S-H (eds) The mungbean genome. Springer International Publishing, Cham, pp 1–8. https://doi.org/10.1007/978-3-030-20008-4_1 Najeeb U, Bange MP, Tan DKY, Atwell BJ (2015) Consequences of waterlogging in cotton and opportunities for mitigation of yield losses. AoB PLANTS 7:plv080. https://doi.org/10.1093/aobpla/plv080 Nawata E, Yoshinaga S, Shigenaga S (1991) Effects of waterlogging duration on the growth and yield of yard long bean ( Vigna sinensis var. sesquipedalis). Sci Hortic 48:185–191. https://doi.org/10.1016/0304-4238(91)90126-J Olorunwa OJ, Adhikari B, Brazel S, et al (2023) Waterlogging stress reduces cowpea ( Vigna unguiculata L.) genotypes growth, seed yield, and quality at different growth stages: Implications for developing tolerant cultivars under field conditions. Agricultural Water Management 284:108336. https://doi.org/10.1016/j.agwat.2023.108336 Oosterhuis DM, Scott HD, Hampton RE, Wullschleger SD (1990) Physiological responses of two soybean [ Glycine max (L.) Merr] cultivars to short-term flooding. Environmental and Experimental Botany 30:85–92. https://doi.org/10.1016/0098-8472(90)90012-S Pampana S, Masoni A, Arduini I (2016) Response of cool-season grain legumes to waterlogging at flowering. Can J Plant Sci 597–603. https://doi.org/10.1139/cjps-2015-0268 Patrick WH, Gambrell RP, Faulkner SP (1996) Redox measurements of soils. In: Sparks DL, Page AL, Helmke PA, et al. (eds) SSSA Book Series. Soil Science Society of America, American Society of Agronomy, Madison, WI, USA, pp 1255–1273 Ploschuk RA, Miralles DJ, Colmer TD, et al (2018) Waterlogging of winter crops at early and late stages: impacts on leaf physiology, growth and yield. Front Plant Sci 9:1863. https://doi.org/10.3389/fpls.2018.01863 Powell AA, Matthews S (1978) The damaging effect of water on dry pea embryos during imbibition. J Exp Bot 29:1215–1229. https://doi.org/10.1093/jxb/29.5.1215 Prasad S, Ram P, Uma S (2004) Effect of waterlogging duration on chlorophyll content, nitrate reductase activity, soluble sugars and grain yield of maize. Ann Plant Physiol 18:1–5 Qi X, Li Q, Shen J, et al (2020) Sugar enhances waterlogging‐induced adventitious root formation in cucumber by promoting auxin transport and signalling. Plant Cell & Environment 43:1545–1557. https://doi.org/10.1111/pce.13738 Rasaei A, Ghobadi M-E, Jalali-Honarmand S, et al (2012) Waterlogging and its effects on nitrogen of soil and plant. Ann. Biol. Res 3: 119-124. Ren B, Hu J, Liu P, et al (2021) Responses of nitrogen efficiency and antioxidant system of summer maize to waterlogging stress under different tillage. PeerJ 9:e11834. https://doi.org/10.7717/peerj.11834 Ren B, Zhang J, Dong S, et al (2016) Effects of waterlogging on leaf mesophyll cell ultrastructure and photosynthetic characteristics of summer maize. PLOS ONE 11:e0161424. https://doi.org/10.1371/journal.pone.0161424 Rhine MD, Stevens G, Shannon G, et al (2010) Yield and nutritional responses to waterlogging of soybean cultivars. Irrig Sci 28:135–142. https://doi.org/10.1007/s00271-009-0168-x Roberts DM, Choi WG, Hwang JH (2010) Strategies for adaptation to waterlogging and hypoxia in nitrogen fixing nodules of legumes. In: Mancuso S, Shabala S (eds) Waterlogging signalling and tolerance in plants. Springer Berlin Heidelberg, Berlin, Heidelberg, pp 37–59 Setter TL, Waters I (2003) Review of prospects for germplasm improvement for waterlogging tolerance in wheat, barley and oats. Plant and Soil 253:1–34. https://doi.org/10.1023/A:1024573305997 Shiferaw W, Shelton H, So H (1992) Tolerance of some subtropical pasture legumes to waterlogging. Tropical Grasslands 26:187–187 Shimamura S, Mochizuki T, Nada Y, Fukuyama M (2003) Formation and function of secondary aerenchyma in hypocotyl, roots and nodules of soybean ( Glycine max ) under flooded conditions. Plant and Soil 251: 351-359. Steffens B, Rasmussen A (2016) The physiology of adventitious roots. Plant Physiology 170:603–617. https://doi.org/10.1104/pp.15.01360 Sultana R, Vales MI, Saxena KB, et al (2013) Waterlogging tolerance in pigeonpea ( Cajanus cajan (L.) Millsp.): genotypic variability and identification of tolerant genotypes. J Agric Sci 151:659–671. https://doi.org/10.1017/S0021859612000755 Thomas AL, Guerreiro SMC, Sodek L (2005) Aerenchyma formation and recovery from hypoxia of the flooded root system of nodulated soybean. Annals of Botany 96:1191–1198. https://doi.org/10.1093/aob/mci272 Tian L, Bi W, Liu X, et al (2019) Effects of waterlogging stress on the physiological response and grain-filling characteristics of spring maize ( Zea mays L.) under field conditions. Acta Physiol Plant 41:63. https://doi.org/10.1007/s11738-019-2859-0 Tian L, Zhang Y, Chen P, et al (2021) How does the waterlogging regime affect crop yield? A global meta-analysis. Front Plant Sci 12:634898. https://doi.org/10.3389/fpls.2021.634898 Tukey JW (1949) Comparing individual means in the analysis of variance. Biometrics 5:99. https://doi.org/10.2307/3001913 Ueno K, Takahashi H (1997) Varietal variation and physiological basis for inhibition of wheat seed germination after excessive water treatment. Euphytica 94:169–173. https://doi.org/10.1023/A:1002976732395 Van Veen H, Akman M, Jamar DCL, et al (2014) Group VII Ethylene Response Factor diversification and regulation in four species from flood‐prone environments. Plant Cell & Environment 37:2421–2432. https://doi.org/10.1111/pce.12302 Vidoz ML, Loreti E, Mensuali A, et al (2010) Hormonal interplay during adventitious root formation in flooded tomato plants. The Plant Journal 63:551–562. https://doi.org/10.1111/j.1365-313X.2010.04262.x Visser EJW, Voesenek LACJ (2005) Acclimation to soil flooding–sensing and signal-transduction. Plant Soil 274:197–214. https://doi.org/10.1007/s11104-004-1650-0 Visser W, Cohen JD, Barendse CWM (1996) An ethylene-mediated increase in sensitivity to auxin induces adventitious root formation in flooded Rumex pahstris Sm. Plant Physiology 112: 1687-1692. Wang X, Deng Z, Zhang W, et al (2017) Effect of waterlogging duration at different growth stages on the growth, yield and quality of cotton. PLoS ONE 12:e0169029. https://doi.org/10.1371/journal.pone.0169029 Wiraguna E, Malik AI, Erskine W (2017) Waterlogging tolerance in lentil ( Lens culinaris Medik. subsp. culinaris) germplasm associated with geographic origin. Genet Resour Crop Evol 64:579–586. https://doi.org/10.1007/s10722-016-0385-0 Xiong R, Sang L, Liu R, et al (2020) Effects of waterlogging on maize seedling growth during seed germination. IOP Conf Ser: Earth Environ Sci 598:012075. https://doi.org/10.1088/1755-1315/598/1/012075 Xu X, Ji J, Xu Q, et al (2018) The major‐effect quantitative trait locus Cs ARN 6.1 encodes an AAA ATP ase domain‐containing protein that is associated with waterlogging stress tolerance by promoting adventitious root formation. The Plant Journal 93:917–930. https://doi.org/10.1111/tpj.13819 Yamauchi T, Abe F, Kawaguchi K, et al (2014) Adventitious roots of wheat seedlings that emerge in oxygen-deficient conditions have increased root diameters with highly developed lysigenous aerenchyma. Plant Signaling & Behavior 9:e28506. https://doi.org/10.4161/psb.28506 Yates RJ, Howieson JG, Hungria M, et al (2016) Authentication of rhizobia and assessment of the legume symbiosis in controlled plant growth systems. In: Howieson JG, Dilworth MJ (eds) Working with rhizobia. Australian Centre for International Agricultural Research, Canberra, pp 73–108 Zaidi PH, Rashid Z, Vinayan M, Babu TA (2012) Pre-germination anaerobic stress tolerance in tropical maize ( Zea mays L.). Aust J Crop Sci 6:1703–1711 Zaman MS, Malik AI, Kaur P, Erskine W (2018) Waterlogging tolerance of pea at germination. J Agronomy Crop Science 204:155–164. https://doi.org/10.1111/jac.12230 Zaman MSU, Malik AI, Kaur P, et al (2019) Waterlogging tolerance at germination in field pea: variability, genetic control, and indirect selection. Front Plant Sci 10:953. https://doi.org/10.3389/fpls.2019.00953 Zaman MSU, Malik AI, Luna LN, et al (2024) Differences of waterlogging tolerance in winter pulse crop between emergence and vegetative stages. J Agronomy Crop Science 210:e12704. https://doi.org/10.1111/jac.12704 Zhang XK, Chen J, Chen L, et al (2008) Imbibition behavior and flooding tolerance of rapeseed seed ( Brassica napus L.) with different testa color. Genet Resour Crop Evol 55:1175–1184. https://doi.org/10.1007/s10722-008-9318-x Tables Table 1 Days to flower, days to maturity and testa colour of four cultivars of two respective pulse crops SL Crop species Cultivar Days to flower Days to maturity Testa color 1 Mungbean ( Vigna radiata ) BARI Mung-6 35 70 Shiny green 2 Mungbean ( Vigna radiata ) BARI Mung-8 36 68 Yellow 3 Mungbean ( Vigna radiata ) Binamoog-8 36 71 Shiny green 4 Blackgram ( Vigna mungo ) BARI Mash-3 37 72 Black Table 2 Degrees of freedom (df), F -values, and probabilities of two-way ANOVA at the reproductive stage during WL and recovery phases. Character Source of variation WL duration Crop species (C) WL X C df 3 3 9 Waterlogging Phase SPAD chlorophyll content F -value 27.79 18.76 4.38 Probability <0.001 <0.001 <0.001 Plant height (cm) F -value 33.22 71.22 0.46 Probability <0.001 <0.001 0.89 Shoot dry mass (g) F -value 17.80 6.26 1.12 Probability <0.001 <0.001 0.37 Tap root depth (cm) F -value 17.07 1.48 1.46 Probability <0.001 0.23 0.20 Root dry mass (g plant −1 ) F -value 54.71 9.75 1.39 Probability <0.001 <0.001 0.23 Adventitious roots F -value 59.65 7.48 4.71 Probability <0.001 <0.001 <0.001 Recovery Phase Plant height (cm) F -value 6.23 37.21 0.93 Probability <0.01 <0.001 0.50 Shoot dry mass (g) F -value 3.70 54.08 0.69 Probability <0.05 <0.001 0.70 Tap root depth (cm) F -value 72.01 11.48 4.21 Probability <0.001 <0.001 <0.001 Root dry mass (g plant −1 ) F -value 37.95 99.88 1.58 Probability <0.001 <0.001 0.16 Nodulation score F -value 76.95 32.42 6.53 Probability <0.001 <0.001 <0.001 Supplementary Files SupplementaryInformation.docx 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. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4569640","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":314203516,"identity":"994f7794-73b1-4a2f-a517-b3f85cc18f49","order_by":0,"name":"Md Shahin Uz Zaman","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA3ElEQVRIiWNgGAWjYDACCQY2hgQIw/ABkOLhI0WLsQFICxtRWqAMMwkQTVCL/OzmZw8e7riTzz+7eVvl1xw7GTYG5oePbuDRYnDnmLlB4plnljPuHCu7LbstGegwNmPjHHxaJBLMJBLbDhsYSOSY3ZbcxgzUwsMmjU+L/Iz0b3AtxZLb6glrYbiRg7CF8eO2w4S1GNzIKQNqeWYgcedYsTTjtuM8bMwE/AJ02DbJn213DIAhtvHjz23V9vzszQ8f43UYBBwAk8w8YJKwcoQWxh/EqR4Fo2AUjIIRBgAcWUa5lsuJMAAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-1040-5963","institution":"Bangladesh Agricultural Research Institute","correspondingAuthor":true,"prefix":"","firstName":"Md","middleName":"Shahin Uz","lastName":"Zaman","suffix":""},{"id":314203517,"identity":"ac433710-1478-45c8-9828-9bc138202203","order_by":1,"name":"Md Mahamudul Hasan","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Md","middleName":"Mahamudul","lastName":"Hasan","suffix":""},{"id":314203518,"identity":"f015320c-2cdc-4a14-a769-cb35c52cc17d","order_by":2,"name":"Lutfun Nahar Luna","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Lutfun","middleName":"Nahar","lastName":"Luna","suffix":""},{"id":314203519,"identity":"e09b1de0-a7f0-48de-b5b3-eb2fff32784c","order_by":3,"name":"Md Shahin Iqbal","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Md","middleName":"Shahin","lastName":"Iqbal","suffix":""},{"id":314203520,"identity":"704104c9-1153-42b7-a5d0-8b3fff6edee0","order_by":4,"name":"Al Imran Malik","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Al","middleName":"Imran","lastName":"Malik","suffix":""},{"id":314203521,"identity":"ebf7e864-1b51-49cd-b4b9-6225ba878466","order_by":5,"name":"Md. Amir Hossain","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Md.","middleName":"Amir","lastName":"Hossain","suffix":""}],"badges":[],"createdAt":"2024-06-12 10:51:35","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4569640/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4569640/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":59379995,"identity":"fea96b21-9ecf-4c99-8de1-f121256d0c9a","added_by":"auto","created_at":"2024-07-01 05:23:16","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":649616,"visible":true,"origin":"","legend":"\u003cp\u003eThe effect of waterlogging (WL) on percent of seedling emergence for four cultivars of two important summer pulse crops, Mungbean and Blackgram, after different duration (Drained control, 3, 5 and 7 d) of WL. Emergence started from 2 d after sowing and was completed by 16 d. Emergence was recorded alternate days for each seed in both the waterlogged and recovery periods. Seeds with an epicotyl longer than 5 mm were considered as germinated. Means within cultivars at a given treatment followed by the same letters are not significantly different at \u003cem\u003eP \u0026lt; \u003c/em\u003e0\u003cem\u003e.\u003c/em\u003e01 on Turkey’s test. Bars are mean (\u003cem\u003en\u003c/em\u003e = 4) ± SE.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/1fc1e068be7a71cec49ac3b7.png"},{"id":59380073,"identity":"de527e07-203c-4a40-aca2-ccb27b81071d","added_by":"auto","created_at":"2024-07-01 05:23:46","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":351427,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage of cumulative seedling emergence in drained control (A), 3 d (B), 5 d (C) and 7 d (D) waterlogging. Arrows indicate the start of draining. Emergence started from 2 d after sowing in drained control and 4 d after in treatment; and was completed by 18 d. Emergence was recorded alternate days for each seed in both the waterlogged and recovery periods.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/4c288adf5b90f947ac19f7c5.png"},{"id":59379964,"identity":"15718fd0-617e-48d4-ae34-b62aac62fc53","added_by":"auto","created_at":"2024-07-01 05:22:48","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":181260,"visible":true,"origin":"","legend":"\u003cp\u003eThe effect of different durations of waterlogging (WL) treatments (Drained control, 3 d, 6 d and 9 d WL) on shoot length (A), shoot dry mass (B) and shoot relative growth rate (C) of four cultivars of two summer pulse crops at WL and recovery (RE) phases. The 9-d WL treatment was imposed on 30 DAS, the 6-d WL treatment was 3 d later, and so on for 3-d WL treatment. All the treatments were harvested at 40 DAS at the end of WL treatment and 68 DAS for 28 days RE. Bars are means (\u003cem\u003en \u003c/em\u003e= 4) ± SE and least significant differences (LSD) at \u003cem\u003eP \u003c/em\u003e= 0.05 for crop.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/570693bc6a3ee8680a6cf6c3.png"},{"id":59380061,"identity":"76ae34bc-a8ba-4cab-9790-53d286eaeaa9","added_by":"auto","created_at":"2024-07-01 05:23:26","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":317780,"visible":true,"origin":"","legend":"\u003cp\u003eThe effect of different durations of waterlogging (WL) treatments (Drained control, 3 d, 6 d and 9 d WL) on tap root length (A), root dry mass (B) and root relative growth rate (C) of four cultivars of two summer pulse crops at WL and recovery (RE) phases. The 9-d WL treatment was imposed on 30 DAS, the 6-d WL treatment was 3 d later, and so on for 3-d WL treatment. All the treatments were harvested at 40 DAS at the end of WL treatment and 68 DAS for 28 days RE. Bars are means (\u003cem\u003en \u003c/em\u003e= 4) ± SE and least significant differences (LSD) at \u003cem\u003eP \u003c/em\u003e= 0.05 for crop.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/cc8650f305aae2e7bc8c5ed9.png"},{"id":59380063,"identity":"f11327c5-3899-4c9b-83a6-b267a1642384","added_by":"auto","created_at":"2024-07-01 05:23:38","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":184311,"visible":true,"origin":"","legend":"\u003cp\u003eThe effect of waterlogging (WL) on the development of adventitious roots (A) and nodule formation (B) for four cultivars of two summer pulse crops after different duration (Drained control, 3, 6 and 9 d) of WL. The 9-d WLtreatment was imposed on 30 DAS, 6-d on 33 DAS and 3-d on 36 DAS to ensure the same end time for all the experimental treatments. All the pots were drained on 40 DAS. The number of adventitious roots and nodule number was recorded at the draining of the pots. Bars are means (\u003cem\u003en \u003c/em\u003e= 4) ± SE and least significant differences (LSD) at \u003cem\u003eP \u003c/em\u003e= 0.05 for crop.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/e3dd55eadf34c1ca9edda434.png"},{"id":59380064,"identity":"6e4809d9-12d2-43b2-a139-3f736672f3c6","added_by":"auto","created_at":"2024-07-01 05:23:39","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":239383,"visible":true,"origin":"","legend":"\u003cp\u003eChlorophyll content (SPAD value) for four cultivars- BARI Mung-6, BARI Mung-8, Binamoog-8 and BARI Mash-3 of two summer pulse crops after different duration (Drained control, 3, 6 and 9 d) of WLand 14 days after draining. WL was imposed at different days after sowing (DAS). The 9-d WLtreatment was imposed on 30 DAS, 6-d on 33 DAS and 3-d on 36 DAS to ensure the same end time for all the experimental treatments. All the pots were drained on 40 DAS. Bars are means ± SE (\u003cem\u003en \u003c/em\u003e= 4).\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/80ee5a70cd99fc9e216ab68e.png"},{"id":59379996,"identity":"3cd415da-7460-42ae-8e3e-8f30e4b349ab","added_by":"auto","created_at":"2024-07-01 05:23:16","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":334028,"visible":true,"origin":"","legend":"\u003cp\u003eThe effect of different durations of waterlogging (WL) treatments (Drained control, 3 d, 6 d and 9 d WL) on pod length (A), Number of seeds per pod (B), pods per plant (C) and seed yield per plant (C) of four summer pulse crops cultivars. The 9-d WL treatment was imposed on 30 DAS, 6-d on 33 DAS and 3-d on 36 DAS to ensure the same end time for all the experimental treatments. All the pots were drained on 40 DAS. Bars are means (\u003cem\u003en \u003c/em\u003e= 4) ± SE and least significant differences (LSD) at \u003cem\u003eP \u003c/em\u003e= 0.05 for crop.\u003c/p\u003e","description":"","filename":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/ad1da760ae44b44d3c9bc0b0.png"},{"id":60612257,"identity":"427cb626-6648-4f59-aa08-e86c32555256","added_by":"auto","created_at":"2024-07-18 19:06:14","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3338992,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/211ee88a-95df-43cf-8c80-679f364a3702.pdf"},{"id":59379959,"identity":"7f1833ba-fd8c-4dd7-96a4-8d537310291b","added_by":"auto","created_at":"2024-07-01 05:22:45","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":15471,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryInformation.docx","url":"https://assets-eu.researchsquare.com/files/rs-4569640/v1/8ae615b016e47a535d235c0b.docx"}],"financialInterests":"","formattedTitle":"Differences of waterlogging tolerance between two grain legume species at germination and reproductive stages","fulltext":[{"header":"Introduction","content":"\u003cp\u003eMungbean (\u003cem\u003eVigna radiata\u003c/em\u003e L.) and blackgram (\u003cem\u003eVigna mungo\u003c/em\u003e L.) are two of the most important legume crops in Asia, including Bangladesh, due to their multipurpose use as food, fodder and green manure. The seed of the crops are a good source of proteins and other necessary micronutrients that play an important role in ensuring nutritional security in developing country. Additionally, they have the ability to fix atmospheric nitrogen (58\u0026ndash;109 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) into soil through a symbiotic relationship with Rhizobium bacteria (Ali \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1992\u003c/span\u003e); which helps in the growth of succeeding crops. More than 7.3\u0026nbsp;million hectares of land are devoted to mungbean cultivation worldwide, producing around 5.3\u0026nbsp;million tons annually. India, Bangladesh and Myanmar collectively contribute approximately to 30% of global output (Nair and Schreinemachers \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In 2021, Bangladesh produced 41,189\u0026nbsp;million tons (MT) of mungbean and 37,344 MT Blackgram (Bangladesh Bureau of Statistics \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Mungbean and blackgram are two crops that account for approximately 22.8% of the total area under cultivation for pulses in Bangladesh. These crops are typically grown on marginal lands where other crops are not suitable. Mungbean is usually planted during pre-monsoon season (Late January to March) on poorly drained soil after the harvest of aman rice (\u003cem\u003eOryza sativa\u003c/em\u003e). This exposes mungbean to waterlogging (WL) which can be detrimental to its growth and development. On the other hand, blackgram is grown during post-monsoon after the flood water has receded. However, the high soil moisture and subsequent unseasonal rainfall can also expose blackgram to WL at germination and reproductive stages. Moreover, the sudden rainfall as a consequence of climate change causes severe WL stress at different growth phases of mungbean and blackgram (Amin et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eExcessive moisture or water in the soil pores, referred to as waterlogged soil, reduces exchange of gases between root tissues and the atmosphere (van Veen et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This can quickly lead to a depletion of soil oxygen, resulting in hypoxia and anoxia within a few hours due to continued respiration by plant roots, soil microflora, and fauna (Gambrell and Patrick \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1978\u003c/span\u003e). The lack of oxygen in waterlogged soil can limit nutrient uptake and transport by roots that affects growth and development in wheat (\u003cem\u003eTriticum aestivum\u003c/em\u003e L.) (Malik et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), oats (\u003cem\u003eAvena sativa\u003c/em\u003e L.) (Arduini et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), maize (Ren et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and cowpea (\u003cem\u003eVigna unguiculata\u003c/em\u003e L.) (Olorunwa et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Moreover, soil nitrogen concentration decreases due to rapid volatilization and denitrification processes (Rasaei et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Furthermore, persistent WL can cause certain ions, such as Mn2\u003csup\u003e+\u003c/sup\u003e, Fe2\u003csup\u003e+\u003c/sup\u003e accumulation to a harmful level for plants (McKee and McKevlin \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1993\u003c/span\u003e). Mungbean and blackgram are particularly susceptible to soil WL, especially during the early stages of development (Bansal et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Douglas et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWL affects plants negatively due to oxygen deficiency, that inhibits root respiration (Jackson and Drew \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). As a result, it causes a decrease in shoot and root growth and nutrient uptake by plants (Malik et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) leading to significant yield losses (Tian et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Legumes are particularly more sensitive to WL and produce lower seed yield when exposed to WL. For instance, under WL at the reproductive stage yield was decreased by 60% in pea (\u003cem\u003ePisum sativum\u003c/em\u003e L.) and lupin (\u003cem\u003eLupinus albus\u003c/em\u003e L.) (Pampana et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), by 35% in chickpea (\u003cem\u003eCicer arietinum\u003c/em\u003e L.) (Cowie et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1996\u003c/span\u003e), by 48% in cowpea (\u003cem\u003eVigna unguiculata\u003c/em\u003e L.) (Minchin et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1978\u003c/span\u003e), and by 41\u0026ndash;58% in soybean (Oosterhuis et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). Among other crops, significant yield losses (39\u0026ndash;44%) have been reported in wheat even with relatively WL-tolerant cultivars (Collaku and Harrison \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe impact of WL on crop growth and yield varies depending on the crop species and genotype and growth stage (Setter and Waters \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Significant variation has observed at the germination stage, and waterlogged tolerant varieties have been identified in some dryland crop species such as pigeon pea (Sultana et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), soybean (Hou et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and maize (Zaidi et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Similarly, variation is reported at the developmental stages of seedling, vegetative, flowering and pod growth, but the level of tolerance varies depends on the specific growth stage. For instance, the pre-flowering stage of growth in pea is more susceptible to WL than any of the vegetative, flowering and pod-filling stages (Cannell and Gales 1979); whereas in mungbean (Ahmed et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) and chickpea (Cowie et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) the reproductive stage is the most sensitive growth stage. WL tolerance of mungbean and blackgram cultivars have been studied under the varying duration of WL stress at germination and seedling stages (Kyu et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This study analyzed the effects of WL at different stages of crops on germination, growth and yield components and the physiological response of two economically and nutritionally important legume crops (mungbean and blackgram); and to identify the superior summer pulse crop cultivars for the farmers of the waterlogged affected areas.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003ePlant materials\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFour most popular cultivars of two summer pulse crops in Bangladesh- Mungbean (BARI Mung-6, BARI Mung-8, Binamoog-8) and Blackgram (BARI Mash-3) were used in this study. All the cultivars are widely grown throughout the country (\u003cstrong\u003eTable 1\u003c/strong\u003e).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eExperimental Conditions\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe experiments were carried out in a research field at PRC, BARI, Ishwardi, Pabna (24\u0026deg;9\u0026prime;N; 89\u0026deg; 4\u0026prime;E; 19 m a.s.l.) from March 2022 to June 2022. The average temperature ranged from 34 \u0026plusmn; 7◦C (day) to 25 \u0026plusmn; 4 (night) in both growth stages. The minimum and maximum temperatures ranged 18 to 42 \u0026deg;C respectively in the field. The growing media comprised soil and sand in 2:1 ratio. The soil was collected from PRC research field (30\u0026deg;78\u0026prime; S, 118\u0026deg;31\u0026prime; E), with soil pH of 7.8 (1: 5 soil and water suspension), similar to Malik et al. (2016). The soil was sun-dried for a week and sieved to 2 mm diameter. The water content (w/w) at field capacity (i.e. pot capacity when fully drained) was 18%.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eExperiment A: Waterlogging at Germination Stage\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe experiment was laid out in a split-plot design with four replications. The main plots were waterlogging (WL) treatments with four levels: drained control, 3 d, 5 d and 7 d WL. The cultivars were in sub-plots. The experimental unit consisted of plastic pots with free draining and sealed base. \u0026nbsp;Free-draining pots contained 100 g gravel at the bottom, followed by 1.0 kg of mixed soil and sand. Each free-draining pots of 0.8 L (90 mm \u0026times; 90 mm \u0026times; 180 mm) with drainage holes (\u0026sim;10 mm in diameter) at the bottom was placed in sealed 60 L plastic tanks. Ten platinum electrodes: five for drained and five for WL pots were placed at a depth of 100 mm to measure soil redox potential. For the waterlogged treatment, tap water was added to sealed pots so that free-draining pots could be waterlogged from the bottom to maintain a water table at soil surface. The pots were waterlogged for 4 d prior to sowing to ensure hypoxia at sowing. Water was added to sealed base pots daily as required to maintain the water table. For drained control treatments, there was no water in the sealed base pots, but the soil moisture in free-draining pots was maintained at \u0026sim;80% of field capacity. Seeds were treated with Provax 200FF (Carboxin 17.5% + Thiram 17.5% FF) @ 3 g kg\u003csup\u003e-1\u003c/sup\u003e seed to control seed-borne and seedling root pathogens. The healthy seeds of all the cultivars with similar size, color and shape were selected for the experiment. Fifteen seeds of each crop were sown on 16 March, 2022 in a free-draining pot by dibbling at 5 mm soil depth as described by \u0026nbsp; Zaman et al. (2019). After each duration of WL treatment pots were allowed to drain.\u003c/p\u003e\n\u003cp\u003eSoil redox potential was measured daily using platinum electrodes (Pt) and an Ag\u0026ndash;AgCl reference electrode with a handheld Digital Multimeter (Fluke 114, Everett, Washington, USA). Redox measurements were corrected according to the method developed by Patrick et al. (1996). Seeds with an epicotyl \u0026gt;5mm were recorded as emergence. Emergence was recorded every alternate day for each seed in both the waterlogged and recovery periods. The experiment was terminated at 22 d after sowing, when there was no sign of further emergence.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eExperiment B: Waterlogging at Reproductive Stage\u003c/p\u003e\n\u003cp\u003eThe experiment comprised of four treatments: drained control and 3 d, 6 d and 9 d WL). The design was factorial with crop (4 levels) x WL treatment (4) in a completely randomized design with four replications. The experimental unit was a plastic pot\u0026mdash;free draining and sealed base. Each free draining pot contained 300 g gravel at the bottom followed by 5 kg mixed soil and sand. Each free draining pot of 8 L (145 mm\u0026times;145 mm\u0026times;220 mm) with 15 mm drainage holes at the bottom was placed in sealed based 10 L (310 mm \u0026times; 620 mm \u0026times; 455 mm) plastic pot. Ten platinum electrodes: five each for drained and for WL pots were placed at 100 mm depth to measure soil redox potential. For the waterlogged treatment, tap water was added to sealed pots so that free-draining pots could be waterlogged from the bottom to maintain a water table at soil surface. Water was added to sealed base pots daily as required to maintain the water table. For drained control treatments, there was no water in the sealed base pots, but the soil moisture in free-draining pots was maintained at \u0026sim;80% of field capacity.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFive seeds were sown in each pot by dibbling at 5 mm soil depth on 17 March 2022. The plants were thinned to two at seven days after sowing. The WL treatments were started 5-7 d before flowering at 30 DAS. The 9 days WL treatment was imposed on 16 April 2022 at 30 DAS, the 6 days WL treatment was 3 days later, and so on for 3 days WL treatment. This approach allowed the same stress release time for all treatments (Zaman et al. 2024; Wang et al. 2017) and replicated a common recovery and drying-out period, thereby enabling to measures to morphological and physiological parameters during recovery. All the Waterlogged pots were drained on 25 April 2022 at 40 DAS. Four pots of each treatment were harvested at the time of draining to measure the plant growth; and the other four pots of each treatment were harvested 28 days after draining, during the recovery. The experiment was terminated at 68 DAS.\u003c/p\u003e\n\u003cp\u003eThe redox potential of the soil was measured daily in the WL and drained pots. The chlorophyll content was measured using a handheld Minolta SPAD 502 (Konica-Minolta, Japan) on the first trifoliate leaf of each plant at two points in time: at the end of WL at 40 DAS and 14 days after draining in recovery. The measurements were taken on the same leaf at both WL and recovery period. At harvest, plant height was measured from the collar (point on the stem where roots start to grow) to the leaf base of the youngest fully expanded leaf of the plant. After gently washing the soil from the roots, the maximum taproot length was measured. The number of emerged adventitious roots longer than 5mm were recorded, and their length recorded at the end of WL treatment. Although the soil was not inoculated with rhizobium, root nodulation was scored at 40 DAS by counting the nodules on the main taproot and lateral roots and using a 0\u0026ndash;8 scoring scale according to Yates et al. (2016), where 0=no nodules, 0.5=white ineffective nodules, 1 = rare effective, 2 = scarce, 3 = moderate, 4 = adequate, 5 = ample, 6 = abundant, 7 = very abundant, and 8 = extremely abundant. Finally, the plants were divided into shoots and roots and dried in paper bags in at 60\u0026deg;C oven for 3 days to record shoot and root dry mass. Relative growth rate was measured at the time of draining to 28 days after recovery. The data for each pot (2 plants in each pot) were pooled and the mean was used as one replicate. Pods per plant and seed yield were recorded at recovery.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eStatistical analysis\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eData recorded for different parameters were compiled and tabulated in proper form for statistical analysis which was carried out in Minitab, LLC. (2021) statistical software package and R Ver 4.2.1 (R Core Team, 2020). Various statistical tests such as, mean performance, two-way ANOVA (Analysis of Variance were executed for different morpho-physiological traits to explore treatment, genotype and treatment x genotype interaction. For all analyses, the means were separated based on their significance levels at 0.05 probability using the Tukey test (Tukey 1949). The relative growth rate (RGR) for shoot and root was calculated according to Hunt (1990), \u003cem\u003eRGR\u003c/em\u003e= {ln (\u003cem\u003eW\u003c/em\u003e2) \u0026minus;ln (\u003cem\u003eW\u003c/em\u003e1)} (\u003cem\u003et\u003c/em\u003e2\u0026minus;\u003cem\u003et\u003c/em\u003e1),\u0026nbsp;where W\u003csub\u003e1\u003c/sub\u003e and W\u003csub\u003e2\u003c/sub\u003e are shoot/root dry mass at times t\u003csub\u003e1\u003c/sub\u003e and t\u003csub\u003e2\u003c/sub\u003e, respectively.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eWaterlogging Tolerance at Germination Stage\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn drained control soil the redox potential was 559 \u0026plusmn; 11 mV throughout the experimental period. By contrast, the redox potential in waterlogged pots at the time of seed sowing to draining period was 337 \u0026plusmn; 12 mV and this increased on draining the pots to 588 \u0026plusmn; 15 by 22 d.\u003c/p\u003e\n\u003cp\u003eThe analysis of variance showed that WL duration, cultivars and their interaction was highly significant (P \u0026lt; 0.001) for emergence (Supplemental Table 1). All the crop cultivars showed close to 100% emergence in drained soil. However, in waterlogged soil crop exhibited contrasting responses in emergence \u003cstrong\u003e(Fig. 1)\u003c/strong\u003e. Seedling emergence was reduced significantly by increased WL duration \u003cstrong\u003e(Fig. 2)\u003c/strong\u003e. Three days of WL reduced seedling emergence to 18% for BARI Mash-3, 24% for BARI Mung-6, 30% for Binamoog-8 and 34% for BARI Mung-8 by the end of the experiment. With the increase of WL duration to 7 d, emergence percentage was further reduced for BARI Mash-3 to 61%, for BARI Mung-6 to 83%, for Binamoog-8 to 81% and for BARI Mung-8 to 86%.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWaterlogging Tolerance at Reproductive Stage\u003c/p\u003e\n\u003cp\u003eSoil redox potential\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAt the start of the WL treatment, soil redox potential in the drained control pots was 466 \u0026plusmn; 12 mV which was almost similar throughout the experimental period. During WL, soil redox potential rapidly decreased to 237 \u0026plusmn; 15 mV at two days after waterlogging that remained almost similar at the end of treatment duration. Upon recovery, the soil redox potential increased to 452 \u0026plusmn; 9 mV within 7 days - almost similar to drained control soil.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eEffect on shoot growth\u003c/p\u003e\n\u003cp\u003eThere was significant variation among the waterlogging (WL) treatments, cultivars and their interaction for the traits in both WL and recovery phases (Table 2). During WL phase (after draining of 40 DAS), all the cultivars showed the highest shoot length (i.e. plant height) in the drained control as compared WL treatments \u003cstrong\u003e(Fig. 3A)\u003c/strong\u003e. With the lowest duration of WL (3 d), the shoot length was reduced by 3, 3.2, 10 and 3.4% in BARI Mung-6, BARI Mung-8, Binamoog-8 and BARI Mash-3, respectively, compared to drained control. When subjected to the longer duration of WL stress (9 d), the shoot length of BARI Mung-6 was decreased by 16.4%, BARI Mung-8 by 20.6% in, Binamoog-8 by 14.4% in and BARI Mash-3 by 8.2% as compared to respective drained control. During recovery phase (68 DAS), drained control showed the highest shoot length compared to waterlogged treatments. The highest waterlogged treatments at 9 d, BARI Mung-6 had the least effect on shoot length (i.e. decreased by 15%) followed by BARI Mash-3 (i.e. decreased by 16%) as compared to drained control \u003cstrong\u003e(Fig. 3A)\u003c/strong\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eShoot dry mass varied depending on cultivars and treatments. During WL phase at 40 DAS, the drained control had higher shoot dry mass for all the cultivars compared to WL treatments. BARI Mung-6 had the highest dry mass (4.42 g plant\u003csup\u003e-1\u003c/sup\u003e) followed by BARI Mash-3 (3.95 g plant\u003csup\u003e-1\u003c/sup\u003e), while Binamoog-8 (3.79 g plant\u003csup\u003e-1\u003c/sup\u003e) had the lowest. However, at the highest level of WL (9 d), shoot dry mass was reduced by 28% in BARI Mash-3, 31% in BARI Mung-6, 34% in BARI Mung-8, and 36% in Binamoog-8\u0026nbsp;relative to the drained controls\u0026nbsp;(\u003cstrong\u003eFig. 3B\u003c/strong\u003e).\u0026nbsp;During the recovery phase (68 DAS), the drained controls showed higher shoot dry mass in comparison to the WL treatments. After recovering from 9 d WL, BARI Mash-3 had the lowest reduction of shoot dry mass (36%) indicating the potential ability to recover from the damage of\u0026nbsp;WL\u0026nbsp;stress during the subsequent period of drainage. In contrast, the highest shoot dry mass reduction (53%) was in\u0026nbsp;Binamoog-8\u0026nbsp;(\u003cstrong\u003eFig. 3B\u003c/strong\u003e).\u003c/p\u003e\n\u003cp\u003eThe relative growth rate (RGR) of shoots was compared between two periods- at the time of draining of WL pots (40 DAS) and 28 days after draining (68 DAS). The RGR of shoots in drained control was positively increased for all the cultivars across the treatments, with the highest increase in BARI Mash-3. However, in waterlogged treatment, RGR of shoot was positive in BARI Mash-3 while it was negative for BARI Mung-6, BARI Mung-8 and Binamoog-8 across the WL treatments (\u003cstrong\u003eFig. 3C\u003c/strong\u003e). At the highest level of 9 d WL, shoot relative growth rate of BARI Mash-3 was 111 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e in compared to 190 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e in drained control. In contrast, BARI Mung-6, BARI Mung-8 and Binamoog-8 showed a negatively decreased RGR of -5, -8 and -9 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e than drained control of 17, 16 and 27 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e respectively.\u003c/p\u003e\n\u003cp\u003eEffect on root growth\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWaterlogging (WL) had a negative impact on the tap root growth for all the cultivars during WL and could not recover by end of the experimental period. Moreover, some loss of or decay of tap root was observed during recovery phase as evident by the shorter root length by the end of the experiment. \u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e During WL phase (40 DAS), all the cultivars in WL treatment exhibited the higher level of root length in compared to drained control \u003cstrong\u003e(Figure 4A)\u003c/strong\u003e. After 3 d of WL treatment, there was no remarkable reduction of root length in BARI Mung-6 compared to drained control. However, BARI Mash-3, BARI Mung-8 and Binamoog-8 experienced a reduction of 7.2%, 8.3% and 12.2%, respectively, compared to drained control. \u0026nbsp;As the WL duration increased to 9 d, the reduction of root length was more significant, with BARI Mash-3 a 17%, BARI Mung-8 a 17.2%, Binamoog-8 a 20% and BARI Mung-6 a 22.5% reduction compared to drained control. During recovery phase (68 DAS), all cultivars experienced secession of the tap root growth as evidenced by the reduced length in waterlogged treatments compared to drained control \u003cstrong\u003e(Figure 4A)\u003c/strong\u003e. After 3 d of WL treatment, BARI Mash-3 had the smallest reduction of root length, with only a 18.6% decrease compared to drained control. However, BARI Mung-8, BARI Mung-6 and Binamoog-8 experienced a reduction of 34%, 36%, 37%, respectively, compared to drained control. \u0026nbsp;As the WL duration increased to 9 d, the reduction of root length was more significant, with BARI Mung-8 a 50%, BARI Mung-6 a 55%, Binamoog-8 a 59% and BARI Mash-3 a 64% reduction compared to drained control.\u003c/p\u003e\n\u003cp\u003eThe effect of WL was highly significant (P \u0026lt; 0.001) on root dry mass for all cultivars (\u003cstrong\u003eTable 2\u003c/strong\u003e). During the WL phase at 40 DAS, root dry mass was reduced by 27% in BARI Mash-3, 33% in BARI Mung-8, 43% in BARI Mung-6 and 47% in Binamoog-8 at 3 d WL treatment, relative to the drained controls; the values in the 9 d WL treatment were 47% in BARI Mash-3, 63% in BARI Mung-8, 65% in Binamoog-8 and 74% in BARI Mung-6 \u003cstrong\u003e(Figure 4B)\u003c/strong\u003e. During recovery phase (68 DAS) of 3 d WL, BARI Mash-3 exhibited the lowest reduction (23%) of root dry mass followed by BARI Mung-8 at 61% with its drained control \u003cstrong\u003e(Figure 4B)\u003c/strong\u003e. With the increased WL duration at 9 d the reduction of root dry mass was 46% in BARI Mash-3, 50% in BARI Mung-8, 64% in BARI Mung-6 and 70% in Binamoog-8.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe relative growth rate (RGR) of roots for all the varieties in waterlogged treatments was decreased in compared to drained control (\u003cstrong\u003eFig. 4C\u003c/strong\u003e). During 9 d WL,\u0026nbsp;BARI Mash-3 exhibited the highest root RGR of 27\u0026nbsp;mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e while in 7 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e for BARI Mung-6, 7.5 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e for BARI Mung-8 and -5 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e for Binamoog-8 compared to 55, 10, 7 and 19 mg\u003csup\u003e-1\u003c/sup\u003eplant\u003csup\u003e-1\u003c/sup\u003ed\u003csup\u003e-1\u003c/sup\u003e respectively in drained control.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAdventitious root formation\u003c/p\u003e\n\u003cp\u003eAdventitious roots were not observed in any of the drained control plants. However, all the\u0026nbsp;cultivars\u0026nbsp;developed adventitious roots near the shoot-root junction (hypocotyl region) in WL treatments and the length ranged from 0.45 to 0.80 mm after 3-9 days of\u0026nbsp;WL. (\u003cstrong\u003eFig. 5A\u003c/strong\u003e). The number of adventitious roots increased with the increase of WL duration and varied within cultivars. After 3 day of WL, BARI Mash-3 exhibited the highest number (3.6) of adventitious roots followed by BARI Mung-8 (2.6). During 9 days of WL, the number of adventitious roots was increased to 27 in BARI Mash-3 followed by Binamoog-8 (19), while the lowest in BARI Mung-6 (10).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eNodule formation\u003c/p\u003e\n\u003cp\u003eIn drained control, all the cultivars showed higher number of nodulation score compared to waterlogged soil, and the number and size of the nodule increased with crop growth\u0026nbsp;(\u003cstrong\u003eFig. 5B\u003c/strong\u003e). \u0026nbsp;Binamoog-8 exhibited the higher score of nodulations (7.6), but the biggest size of nodules was observed in BARI Mash-3 in compared to other crop in drained soil. In waterlogged soil, the number of nodules decreased with the increasing duration of WL. The shortest WL period of 3 d did not affect nodule formation in all the cultivars compared to its drained control. With the increase of WL duration to 9 d, nodule score was decreased by 42% in BARI Mash-3, 57% in Binamoog-8, 72% in BARI Mung-6 and 86% in BARI Mung-8. In comparison of crops, the nodule formation in BARI Mash-3 were less affected than other mungbean cultivars. All the crop produced nodules on lateral and adventitious roots near the soil surface during WL, but the plants in drained control had nodules in lower the soil surface. The color of the nodules was brown in both drained control and WL indicating that they were functional.\u003c/p\u003e\n\u003cp\u003eEffect on chlorophyll content\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe chlorophyll content (SPAD value) of the first trifoliate leaves were recorded at WL (at the time of draining/40 DAS) and 14 days after recovery phase (54 DAS) to understand the effect of WL on photosynthesis. In the drained control the chlorophyll content in both WL and recovery phase was higher than waterlogged plants for all the cultivars, but chlorophyll content at recovery was reduced compared to WL due to ageing the plants \u003cstrong\u003e(Fig. 6)\u003c/strong\u003e. For instance, the highest SPAD value (55) of BARI Mung-8 in drained control during\u0026nbsp;WL\u0026nbsp;phase was decreased to 47 at recovery phase. Among the waterlogged treatments, the SPAD value in recovery phase was increased in BARI Mash-3, while decreased in all the mungbean cultivars across the treatments than\u0026nbsp;WL\u0026nbsp;phase. At the highest level of WL treatments at 9 d, the SPAD value was increased 15% in BARI Mash-3 at recovery phase in compared to WL phase. In contrast, the SPAD value decreased 11% in BARI Mung-6, 21% in Binamoog-8 and 32% in BARI Mung-8 at 9 d WL during the recovery phase.\u003c/p\u003e\n\u003cp\u003eEffect on pod formation and seed yield\u003c/p\u003e\n\u003cp\u003eExposer to waterlogging during reproductive stage had a negative impact on both pod number plant\u003csup\u003e-1\u003c/sup\u003e and seed yield plant\u003csup\u003e-1\u003c/sup\u003e \u003cstrong\u003e(Fig. 7)\u003c/strong\u003e. However, it did not affect pod length and seeds pod\u003csup\u003e-1\u003c/sup\u003e (\u003cstrong\u003e(Fig. 7A\u0026amp;B)\u003c/strong\u003e. In the drained control, BARI Mash-3 produced the highest number of pods plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003e(57) followed by BARI Mung-8 (46). When exposed to the shortest duration of (3 d) WL treatment, the number of pods plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003ewas reduced by 19% in Binamoog-8, 43% in BARI Mash-3, 50 % in BARI Mung-6 and 49% in BARI Mung-8 compared to drained controls. During 9 d WL, the reduction of number of pods plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003ewas the highest in BARI Mung-8 (74%) while in other cultivars it ranged from 50-59% compared to drained controls \u003cstrong\u003e(Fig. 7C)\u003c/strong\u003e.\u003c/p\u003e\n\u003cp\u003eThe waterlogging drastically affected the yield of all cultivars \u003cstrong\u003e(Fig. 7D)\u003c/strong\u003e. The longer the waterlogging duration during the reproductive stage, the greater the yield losses.\u0026nbsp;In the drained control, seed yield plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003eranged from 5.8 g in BARI Mash-3 to 9.0 g in BARI Mung-8. The shortest duration of 3 d WL reduced seed yield plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003eby 34% in Binamoog-8, 41% in BARI Mash-3, 52% in BARI Mung-8 and 54 % in BARI Mung-6 compared to drained controls. During 9 d WL treatment, the reduction of seed yield plant\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003ewas 59% in BARI Mung-8, 58% in BARI Mung-6, 52% in Binamoog-8 and 40% in BARI Mash-3 compared to drained controls.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThis study investigated the tolerance of two summer grain legumes to waterlogging at the germination and reproductive stages, highlighting the importance of implementing appropriate measures to prevent waterlogging in fields for optimal crop growth and yield.\u003c/p\u003e \u003cp\u003ea. Variation in waterlogging tolerance at germination\u003c/p\u003e \u003cp\u003eThe findings of the study indicate that exposing legumes cultivars to waterlogging conditions for a short period can have a significant impact on its growth in the long term and ultimately affects emergence and yield. During the early stages of germination in control treatment, it was observed that all cultivars of both species exhibited more than 90% emergence within four days of sowing. This initial emergence was consistent across all cultivars studied, indicating that the germination process is not significantly affected by genetic differences between cultivars. However, when the plants were subjected to waterlogging treatments, emergence was delayed and reduced, indicating that waterlogging has a significant impact on the germination process. In the waterlogged treatment, emergence only began after nine days of sowing and was completed by the 16th day. This delay in emergence indicates that waterlogging has a significant impact on the germination process and can lead to slower growth and development of plants. The delay in emergence is likely due to the fact that waterlogging creates an anaerobic environment in the soil, which leads to reduced oxygen availability for the developing seeds (Zaman et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2018\u003c/span\u003e in peas). The insufficient oxygen levels prevent the respiration which is required for germination (Powell and Matthews \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1978\u003c/span\u003e, wheat and barley; Hou and Thseng \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1992\u003c/span\u003e, Soybean; Xiong et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, maize). This lack of respiration inhibits the growth and development of the seeds, leading to slower emergence and reduced plant growth (Wiraguna et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, lentil; Kyu et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, mungbean). Such information is vital for farmers and researchers alike, as it can help them make informed decisions about crop management strategies that can improve plant growth under challenging environmental conditions.\u003c/p\u003e \u003cp\u003eThe longer the WL duration, the greater the reduction in seedling emergence. In the present study, with the longest WL duration of 7 days, emergence was reduced by 70%. However, there was a variation of emergence among the cultivars. The dark testa BARI Mash-3 exhibited the higher percent of emergence than shiny green and yellow mungbean cultivars which is consistent with previous studied where the dark testa genotypes of peas (Zaman et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), soybean (Hou and Thseng \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1991\u003c/span\u003e), rapeseed (Zhang et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and wheat (Ueno and Takahashi \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) were found to be tolerant to WL compared to less pigmented seeds of respective crops. The WL tolerance of dark seeded genotypes is presumably due to the presence of higher levels of phenolic compounds compared to the sensitive yellow seeded lines (Zhang et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The phenolic compounds are responsible for preventing testa leakage in the tolerant genotypes. Conversely, the testa is leaked and whitish seed solutes are observed on the seed surface of susceptible genotypes in absence of phenolic compounds (Zaman et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In this study the black testa BARI Mash-3 was tolerant as in other crops but the shiny green testa mungbean cultivars showed susceptibility.\u003c/p\u003e \u003cp\u003eb. Effect of waterlogging on growth\u003c/p\u003e \u003cp\u003eAfter emerging, plant root systems undergo development, allowing plants to use energy through root respiration for optimal growth and development in the presence of oxygen in soil pores. In the waterlogged condition, the respiration is limited due to absence of oxygen that reduces root length in mungbean (Kumar et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), soybean (Miao et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), wheat (Herzog et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), maize (Ren et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and in current study (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003ea). However, the WL tolerant genotypes produced adventitious roots (ARs) as the primary roots are damaged (Qi et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The development of ARs has been observed in many species- sunflower (\u003cem\u003eHelianthus annuus\u003c/em\u003e L.) (Jackson \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1955\u003c/span\u003e), rice (Lorbiecke and Sauter \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), wheat (Yamauchi et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), maize (Mano et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), Rumex spp. (Visser et. Al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e1996\u003c/span\u003e), and tomato (\u003cem\u003eSolanum lycopersicum\u003c/em\u003e; Vidoz et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The newly developed ARs emerge from the hypocotyl, mesocotyl, or base of the stem and contain more aerenchyma than primary roots (Visser and Voesenek 2004). Consequently, ARs enhance gas exchange and water and nutrient uptake in plants, which contribute greatly to plant tolerance of poorly aerated soils (Steffens and Rasmussen \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The number of adventitious roots is increased with the increase of WL duration in mungbean and blackgram (Kyu et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kumar et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), cucumber (Xu et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), wheat (Koramutla et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and yard long bean (Nawata et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). In this study, with the longest WL of 9 days, the highest number of adventitious roots was found in BARI Mash-3 while the lowest in all the mungbean cultivars during WL. However, there was no such root development in the drained controls.\u003c/p\u003e \u003cp\u003eIn a drained control soil, legume crops acquire nitrogen mostly through symbiotic nitrogen fixation in rhizobia nodules. Under WL conditions, as soil condition shifts from hypoxia to anoxia, there is reduced O\u003csub\u003e2\u003c/sub\u003e for the \u003cem\u003erizhobium\u003c/em\u003e. Thus, with prolonged WL, the number of rhizobia decrease due to unavailable of soil oxygen. In the present study, all the crops showed the higher number of nodules in drained control than WL treatments. During WL, the variation of nodulation was observed among the crops. After 9 days WL of this study, nodulation had decreased by 42% in BARI Mash-3, 57% in Binamoog-8, 72% in BARI Mung-6 and 86% BARI Mung-8. This suggests that the nodulation might have an influence for survival ability of crops during WL stress. Previous research indicated that in legumes, tissues of stem, root and nodule are connected by a well- developed network of aerenchyma that allow oxygen diffusion to nodules (Roberts et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In soybean, the aerenchyma connects to the outer cortex of nodules, presumably, supporting their functioning for roots in WL soils (Shimamura et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Thomas et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Hence, the crop with more nodules have a beter ability to thrive in waterlogged soil. Nodules located below 5\u0026ndash;10 cm of WL soil might have become ineffective as their color changed from pink to white (Roberts et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In the current study, pink nodules continued to develop at the at the base of the soil with adventitious roots regardless of the duration of WL, suggesting that they were able to function during the recovery phase. In soybean, the aerenchyma attaches to the outer cortex of nodules, possibly allowing them to serve as roots in WL. soils (Shimamura et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Thomas et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe damage of the root systems under WL affected the shoots growth, through a reduced number of leaves and a smaller leaf area. Limitation of nutrient uptake following a longer duration of WL might be the cause of the reduction in shoot that has been reported in previous studies (Bacanamwo and Purcell \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1999\u003c/span\u003e- soybean; Malik et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e- wheat; Board \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2008\u003c/span\u003e- soybean; Malik et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2015\u003c/span\u003e- pea, lentil, and grass pea; Najeeb et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2015\u003c/span\u003e- cotton). Moreover, WL causes leaching of nutrient e.g., N, P and K, causing nutrient shortage symptoms thus resulting in shoot length reduction (Rhine et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In this study, all cultivars showed significant variation in plant height and reduced shoot length under WL. With the increasing duration of WL shoot length also decreases. Highest shoot length reduction was observed in BARI Mung-6 and BARI Mash-3 following the WL recovery.\u003c/p\u003e \u003cp\u003eThe symptom of affecting shoot growth is visualized with leaf chlorosis. In the present study, chlorophyll content (SPAD value) decreased under WL \u0026ndash; compared to drained control - for all the crop, indicating the degradation of chlorophyll in functional leaves and a reduction in photosynthetic capacity (Barickman et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). During the recovery period, SPAD chlorophyll increased in BARI Mash-3 for all the treatments; but was further decreased in mungbean cultivars compared to WL period. Chlorophyll content (greenness of leaves) is an important parameter to screen pulses for WL tolerance. Similarly, reduction in chlorophyll content under WL has been reported in blackgram (Bansal et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), mungbean (Kyu et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kumar et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), sesame (Mensah et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), wheat (Collaku and Harrison \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) and maize (Prasad et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Under WL, yellowing of the plant may be caused by a reduction in leaf nitrogen (Bacanamwo and Purcell \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), nodulation and N fixation, and the production of toxic substances such as nitrites and sulfides that are carried upward in large quantities from the soil through the roots and into the leaf (Ezin et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In addition, WL reduces nitrogen in the soil by accelerated volatilization and denitrification (Rasaei et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ec. Effect on waterlogging on yield\u003c/p\u003e \u003cp\u003eThe shoot and root dry matter significantly decreased under WL stress by damaging the existing root systems of mungbean and blackgram cultivars which is consistent to other studies in mungbean (Ahmed et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Kumar et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e); chickpea, fababean (Munir et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e); soybean (Miao et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Beutler et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Kim et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and wheat (Malik et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Relative Growth Rate (RGR) of shoot was higher in BARI Mash-3 compared to drained control, however, it was decreased to all the mungbean cultivars for all the WL treatments indicating contrasting response of crop, tolerant- BARI Mash-3 and sensitive- mungbean cultivars. The response was clearest during the highest level of WL at 9 days. Similarly, the RGR of roots was higher in BARI Mash-3 in compared to drained control across the treatments. At 9 days WL, BARI Mash-3 exhibited 3-fold higher RGR than drained control. In contrast, RGR of roots in mungbean cultivars was negative similar to the WL response seen in rapeseed waterlogged for 14 days (Brisson et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ploschuk et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Finally, WL directly affects on pod formations and ultimate yield. The of number of pods plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e was gradually declined with increasing WL duration. As plants most energy distributes to the formation of adventitious root and dry matter to survive through the WL stress thus least amount of energy, sugar, metabolites were left for the yield and yield related characters such as pod setting, pod filling and seed formation (Kyu et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kumar et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This decreases the yield 40\u0026ndash;60% in compared to drained control for the cultivars with the lowest (40%) in BARI Mash-3. Previous studies also showed the reduction of yield in different crops such as soybean (Miao et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Beutler et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), maize (Tian et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), wheat and barley (De San Celedonio et al. 2014), under WL. This indicates that, BARI Mash-3 have the higher ability to grow WL than those of mungbean cultivars.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eWaterlogging at different growth stage is one of the major constraints to pulse crop in rice-based cropping. This study has demonstrated the variation of waterlogging tolerance between two summer grain legumes. BARI Mash-3 showed the higher ability to grow in waterlogging at both germination and reproductive stages in compared to the mungbean cultivars. Seed yield was decreased to all the cultivars of both species, but the lowest reduction (40%) was found in BARI Mash-3, while the highest (59%) in BARI Mung-8. It indicated that blackgram is more suited to waterlogging condition. This differences of waterlogging response with stages and genotypes will help to select crops to alleviate the waterlogging stress.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements \u0026nbsp; \u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work has been part of the project Incorporating salt-tolerant wheat and pulses into smallholder farming systems in southern Bangladesh (CIM/2014/076) funded by Australian Centre for International Agriculture Research (ACIAR) and Krishi Gobeshona Foundation (KGF), Bangladesh in collaboration with the Pulses Research Centre (PRC), Bangladesh Agricultural Research Institute, Bangladesh. The authors are thankful to the authorities of ACIAR, KGF and PRC for all sorts of supports and facilities. We are also grateful to the staff at PRC, Ishwardi, Pabna, Bangladesh for all their assistance during the field trials.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e: \u003cem\u003eThere is no funds, grants, or other support received during the preparation of this manuscript.\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interests:\u003c/strong\u003e \u003cem\u003eWe have no relevant financial or non-financial interests.\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions:\u0026nbsp;\u003c/strong\u003e\u003cem\u003eAll authors contributed to the study conception and design. Material preparation, data collection and analysis were performed by Md Shahin Uz Zaman and\u0026nbsp;\u003c/em\u003eMd Mahamudul Hasan.\u003cem\u003e\u0026nbsp;The first draft of the manuscript was written by Md Shahin Uz Zaman and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.\u003c/em\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability:\u0026nbsp;\u003c/strong\u003e\u003cem\u003eThe datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.\u003c/em\u003e\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAhmed S, Nawata E, Sakuratani T (2002) Effects of waterlogging at vegetative and reproductive growth stages on photosynthesis, leaf water potential and yield in mungbean. Plant Prod Sci 5:117\u0026ndash;123. https://doi.org/10.1626/pps.5.117\u003c/li\u003e\n\u003cli\u003eAli M (1992) Weeds are a great threat to kharif pulses. Indian farming 42:27\u0026ndash;29\u003c/li\u003e\n\u003cli\u003eAmin M, Amiruzzaman M, Ahmed A, Ali M (2014) Combining ability study in waterlogged tolerant maize (\u003cem\u003eZea mays\u003c/em\u003e L.). Bangladesh J Agric Res 39:283\u0026ndash;291. https://doi.org/10.3329/bjar.v39i2.20430\u003c/li\u003e\n\u003cli\u003eArduini I, Baldanzi M, Pampana S (2019) Reduced growth and nitrogen uptake during waterlogging at tillering permanently affect yield components in late sown oats. Front Plant Sci 10:1087. https://doi.org/10.3389/fpls.2019.01087\u003c/li\u003e\n\u003cli\u003eBacanamwo M, Purcell LC (1999) Soybean dry matter and N accumulation responses to flooding stress, N sources and hypoxia. Journal of Experimental Botany 50: 689-696.\u003c/li\u003e\n\u003cli\u003eBangladesh Bureau of Statistics (2021) Yearbook of agricultural statistics. Ministry of planning, government of the people\u0026rsquo;s republic Bangladesh, Dhaka\u003c/li\u003e\n\u003cli\u003eBansal R, Dikshit H, Singh A, et al (2022) Effect of waterlogging on physiological traits and yield in blackgram (\u003cem\u003eVigna mungo\u003c/em\u003e L.) in field condition. Indian J Biochem Biophys 59:580\u0026ndash;585. https://doi.org/10.56042/ijbb.v59i5.63013\u003c/li\u003e\n\u003cli\u003eBansal R, Sharma S, Tripathi K, Kumar A (2019) Waterlogging tolerance in blackgram [\u003cem\u003eVigna mungo \u003c/em\u003e(L.) Hepper] is associated with chlorophyll content and membrane integrity. Indian J Biochem Biophys 56:\u003c/li\u003e\n\u003cli\u003eBarickman TC, Simpson CR, Sams CE (2019) Waterlogging causes early modification in the physiological performance, carotenoids, chlorophylls, proline, and soluble sugars of cucumber plants. Plants 8:160. https://doi.org/10.3390/plants8060160\u003c/li\u003e\n\u003cli\u003eBeutler AN, Giacomeli R, Alberto CM, et al (2014) Soil hydric excess and soybean yield and development in Brazil. Aust J Crop Sci 8:1461\u0026ndash;1466\u003c/li\u003e\n\u003cli\u003eBoard JE (2008) Waterlogging effects on plant nutrient concentrations in soybean. Journal of Plant Nutrition 31:828\u0026ndash;838. https://doi.org/10.1080/01904160802043122\u003c/li\u003e\n\u003cli\u003eBrisson N, Rebiere B, Zimmer D, Renault P (2002) Response of the root system of a winter wheat crop to waterlogging. Plant and soil 243: 43-55.\u003c/li\u003e\n\u003cli\u003eCannell RQ, Gales K, Snaydon RW, Suhail BA (1979) Effects of short‐term waterlogging on the growth and yield of peas (\u003cem\u003ePisum sativum\u003c/em\u003e). Annals of Applied Biology 93:327\u0026ndash;335. https://doi.org/10.1111/j.1744-7348.1979.tb06549.x\u003c/li\u003e\n\u003cli\u003eCollaku A, Harrison SA (2002) Losses in wheat due to waterlogging. Crop Science 42: 444-450.\u003c/li\u003e\n\u003cli\u003eCowie AL, Jessop RS, MacLeod DA (1996) Effects of waterlogging on chickpeas II. Possible causes of decreased tolerance of waterlogging at flowering. Plant Soil 183:105\u0026ndash;115. https://doi.org/10.1007/BF02185570\u003c/li\u003e\n\u003cli\u003eDouglas C, Pratap A, Rao BH, et al (2020) Breeding progress and future challenges: Abiotic stresses. In: Nair RM, Schafleitner R, Lee S-H (eds) The Mungbean Genome. Springer International Publishing, Cham, pp 81\u0026ndash;96. https://doi.org/10.1007/978-3-030-20008-4_6\u003c/li\u003e\n\u003cli\u003eEzin V, Pena RDL, Ahanchede A (2010) Flooding tolerance of tomato genotypes during vegetative and reproductive stages. Braz J Plant Physiol 22:131\u0026ndash;142. https://doi.org/10.1590/S1677-04202010000200007\u003c/li\u003e\n\u003cli\u003eGambrell RP, Patrick WH Jr (1978) Chemical and microbiological properties of anaerobic soils and sediments. In: Hook DD, Crawford RM (eds) Plant life in anaerobic environments. Ann Arbor Science, Ann Arbor, pp 375\u0026ndash;423\u003c/li\u003e\n\u003cli\u003eHerzog M, Striker GG, Colmer TD, Pedersen O (2016) Mechanisms of waterlogging tolerance in wheat \u0026ndash; a review of root and shoot physiology. Plant Cell \u0026amp; Environment 39:1068\u0026ndash;1086. https://doi.org/10.1111/pce.12676\u003c/li\u003e\n\u003cli\u003eHou FF, Thseng FS (1991) Studies on the flooding tolerance of soybean seed: varietal differences. Euphytica 57:169\u0026ndash;173. https://doi.org/10.1007/BF00023075\u003c/li\u003e\n\u003cli\u003eHou F-F, Thseng F-S (1992) Studies on the screening technique for pre-germination flooding tolerance in soybean. Jpn J Crop Sci 61:447\u0026ndash;453. https://doi.org/10.1626/jcs.61.447\u003c/li\u003e\n\u003cli\u003eHou F-F, Thseng F-S, Wu S-T, Takeda K (1995) Varietal differences and diallel analysis of pre-germination flooding tolerance in soybean seed. Bull Res Inst Bioresour Okayama Univ 3:35\u0026ndash;41\u003c/li\u003e\n\u003cli\u003eHunt R (1990) Relative growth rates. In: Hunt R (ed) Basic growth analysis: Plant growth analysis for beginners. Springer Netherlands, Dordrecht, pp 25\u0026ndash;34\u003c/li\u003e\n\u003cli\u003eJackson MB, Drew MC (1984) Effects of flooding on growth and metabolism of herbaceous plants. In: Flooding and Plant Growth. Elsevier, pp 47\u0026ndash;128\u003c/li\u003e\n\u003cli\u003eJackson WT (1955) The role of adventitious roots in recovery of shoots following flooding of the original root systems. American J of Botany 42:816\u0026ndash;819. https://doi.org/10.1002/j.1537-2197.1955.tb10428.x\u003c/li\u003e\n\u003cli\u003eKim KH, Cho MJ, Kim J-M, et al (2019) Growth response and developing simple test method for waterlogging stress tolerance in soybean. J Crop Sci Biotechnol 22:371\u0026ndash;378. https://doi.org/10.1007/s12892-019-0271-0\u003c/li\u003e\n\u003cli\u003eKoramutla MK, Tuan PA, Ayele BT (2022) Salicylic acid enhances adventitious root and aerenchyma formation in wheat under waterlogged conditions. IJMS 23:1243. https://doi.org/10.3390/ijms23031243\u003c/li\u003e\n\u003cli\u003eKumar P, Pal M, Joshi R, Sairam RK (2013) Yield, growth and physiological responses of mungbean[\u003cem\u003eVigna radiata\u003c/em\u003e (L.) Wilczek] genotypes to waterlogging at vegetative stage. Physiol Mol Biol Plants 19:209\u0026ndash;220. https://doi.org/10.1007/s12298-012-0153-3\u003c/li\u003e\n\u003cli\u003eKyu KL, Malik AI, Colmer TD, et al (2021) Response of mungbean (cvs. Celera II-AU and Jade-AU) and blackgram (cv. Onyx-AU) to transient waterlogging. Front Plant Sci 12:709102. https://doi.org/10.3389/fpls.2021.709102\u003c/li\u003e\n\u003cli\u003eLorbiecke R, Sauter M (1999) Adventitious root growth and cell-cycle induction in deepwater rice1. Plant Physiology 119:21\u0026ndash;30. https://doi.org/10.1104/pp.119.1.21\u003c/li\u003e\n\u003cli\u003eMalik AI, Ailewe TI, Erskine W (2015) Tolerance of three grain legume species to transient waterlogging. AoB PLANTS 7. https://doi.org/10.1093/aobpla/plv040\u003c/li\u003e\n\u003cli\u003eMalik AI, Ali MO, Zaman MS, et al (2016) Relay sowing of lentil (\u003cem\u003eLens culinaris\u003c/em\u003e subsp. culinaris) to intensify rice-based cropping. J Agric Sci 154:850\u0026ndash;857. https://doi.org/10.1017/S0021859614001324\u003c/li\u003e\n\u003cli\u003eMalik AI, Colmer TD, Lambers H, et al (2002) Short‐term waterlogging has long‐term effects on the growth and physiology of wheat. New Phytologist 153: 225-236.\u003c/li\u003e\n\u003cli\u003eMano Y, Muraki M, Fujimori M, et al (2005) Identification of QTL controlling adventitious root formation during flooding conditions in teosinte (Zea mays ssp. huehuetenangensis) seedlings. Euphytica 142:33\u0026ndash;42. https://doi.org/10.1007/s10681-005-0449-2\u003c/li\u003e\n\u003cli\u003eMcKee WH, McKevlin MR (1993) Geochemical processes and nutrient uptake by plants in hydric soils. Enviro Toxic and Chemistry 12:2197\u0026ndash;2207. https://doi.org/10.1002/etc.5620121204\u003c/li\u003e\n\u003cli\u003eMensah JK, Obadoni BO, Eruotor PG, Onome-Irieguna F (2006) Simulated flooding and drought effects on germination, growth, and yield parameters of sesame (\u003cem\u003eSesamum indicum\u003c/em\u003e L.). African Journal of biotechnology 5:1249-1253. \u003c/li\u003e\n\u003cli\u003eMiao S, Shi H, Jin J, et al (2012) Effects of short-term drought and flooding on soybean nodulation and yield at key nodulation stage under pot culture. J Food Agric Env 10:819\u0026ndash;824\u003c/li\u003e\n\u003cli\u003eMinchin FR, Summerfield RJ, Eaglesham ARJ, Stewart KA (1978) Effects of short-term waterlogging on growth and yield of cowpea (\u003cem\u003eVigna unguiculata\u003c/em\u003e). J Agric Sci 90:355\u0026ndash;366. https://doi.org/10.1017/S0021859600055465\u003c/li\u003e\n\u003cli\u003eMunir R, Konnerup D, Khan HA, et al (2019) Sensitivity of chickpea and faba bean to root‐zone hypoxia, elevated ethylene, and carbon dioxide. Plant Cell \u0026amp; Environment 42:85\u0026ndash;97. https://doi.org/10.1111/pce.13173\u003c/li\u003e\n\u003cli\u003eNair R, Schreinemachers P (2020) Global status and economic importance of mungbean. In: Nair RM, Schafleitner R, Lee S-H (eds) The mungbean genome. Springer International Publishing, Cham, pp 1\u0026ndash;8. https://doi.org/10.1007/978-3-030-20008-4_1\u003c/li\u003e\n\u003cli\u003eNajeeb U, Bange MP, Tan DKY, Atwell BJ (2015) Consequences of waterlogging in cotton and opportunities for mitigation of yield losses. AoB PLANTS 7:plv080. https://doi.org/10.1093/aobpla/plv080\u003c/li\u003e\n\u003cli\u003eNawata E, Yoshinaga S, Shigenaga S (1991) Effects of waterlogging duration on the growth and yield of yard long bean (\u003cem\u003eVigna sinensis\u003c/em\u003e var. sesquipedalis). Sci Hortic 48:185\u0026ndash;191. https://doi.org/10.1016/0304-4238(91)90126-J\u003c/li\u003e\n\u003cli\u003eOlorunwa OJ, Adhikari B, Brazel S, et al (2023) Waterlogging stress reduces cowpea (\u003cem\u003eVigna unguiculata\u003c/em\u003e L.) genotypes growth, seed yield, and quality at different growth stages: Implications for developing tolerant cultivars under field conditions. Agricultural Water Management 284:108336. https://doi.org/10.1016/j.agwat.2023.108336\u003c/li\u003e\n\u003cli\u003eOosterhuis DM, Scott HD, Hampton RE, Wullschleger SD (1990) Physiological responses of two soybean [\u003cem\u003eGlycine max\u003c/em\u003e (L.) Merr] cultivars to short-term flooding. Environmental and Experimental Botany 30:85\u0026ndash;92. https://doi.org/10.1016/0098-8472(90)90012-S\u003c/li\u003e\n\u003cli\u003ePampana S, Masoni A, Arduini I (2016) Response of cool-season grain legumes to waterlogging at flowering. Can J Plant Sci 597\u0026ndash;603. https://doi.org/10.1139/cjps-2015-0268\u003c/li\u003e\n\u003cli\u003ePatrick WH, Gambrell RP, Faulkner SP (1996) Redox measurements of soils. In: Sparks DL, Page AL, Helmke PA, et al. (eds) SSSA Book Series. Soil Science Society of America, American Society of Agronomy, Madison, WI, USA, pp 1255\u0026ndash;1273\u003c/li\u003e\n\u003cli\u003ePloschuk RA, Miralles DJ, Colmer TD, et al (2018) Waterlogging of winter crops at early and late stages: impacts on leaf physiology, growth and yield. Front Plant Sci 9:1863. https://doi.org/10.3389/fpls.2018.01863\u003c/li\u003e\n\u003cli\u003ePowell AA, Matthews S (1978) The damaging effect of water on dry pea embryos during imbibition. J Exp Bot 29:1215\u0026ndash;1229. https://doi.org/10.1093/jxb/29.5.1215\u003c/li\u003e\n\u003cli\u003ePrasad S, Ram P, Uma S (2004) Effect of waterlogging duration on chlorophyll content, nitrate reductase activity, soluble sugars and grain yield of maize. Ann Plant Physiol 18:1\u0026ndash;5 \u003c/li\u003e\n\u003cli\u003eQi X, Li Q, Shen J, et al (2020) Sugar enhances waterlogging‐induced adventitious root formation in cucumber by promoting auxin transport and signalling. Plant Cell \u0026amp; Environment 43:1545\u0026ndash;1557. https://doi.org/10.1111/pce.13738\u003c/li\u003e\n\u003cli\u003eRasaei A, Ghobadi M-E, Jalali-Honarmand S, et al (2012) Waterlogging and its effects on nitrogen of soil and plant. Ann. Biol. Res 3: 119-124.\u003c/li\u003e\n\u003cli\u003eRen B, Hu J, Liu P, et al (2021) Responses of nitrogen efficiency and antioxidant system of summer maize to waterlogging stress under different tillage. PeerJ 9:e11834. https://doi.org/10.7717/peerj.11834\u003c/li\u003e\n\u003cli\u003eRen B, Zhang J, Dong S, et al (2016) Effects of waterlogging on leaf mesophyll cell ultrastructure and photosynthetic characteristics of summer maize. PLOS ONE 11:e0161424. https://doi.org/10.1371/journal.pone.0161424\u003c/li\u003e\n\u003cli\u003eRhine MD, Stevens G, Shannon G, et al (2010) Yield and nutritional responses to waterlogging of soybean cultivars. Irrig Sci 28:135\u0026ndash;142. https://doi.org/10.1007/s00271-009-0168-x\u003c/li\u003e\n\u003cli\u003eRoberts DM, Choi WG, Hwang JH (2010) Strategies for adaptation to waterlogging and hypoxia in nitrogen fixing nodules of legumes. In: Mancuso S, Shabala S (eds) Waterlogging signalling and tolerance in plants. Springer Berlin Heidelberg, Berlin, Heidelberg, pp 37\u0026ndash;59\u003c/li\u003e\n\u003cli\u003eSetter TL, Waters I (2003) Review of prospects for germplasm improvement for waterlogging tolerance in wheat, barley and oats. Plant and Soil 253:1\u0026ndash;34. https://doi.org/10.1023/A:1024573305997\u003c/li\u003e\n\u003cli\u003eShiferaw W, Shelton H, So H (1992) Tolerance of some subtropical pasture legumes to waterlogging. Tropical Grasslands 26:187\u0026ndash;187\u003c/li\u003e\n\u003cli\u003eShimamura S, Mochizuki T, Nada Y, Fukuyama M (2003) Formation and function of secondary aerenchyma in hypocotyl, roots and nodules of soybean (\u003cem\u003eGlycine max\u003c/em\u003e) under flooded conditions. Plant and Soil 251: 351-359.\u003c/li\u003e\n\u003cli\u003eSteffens B, Rasmussen A (2016) The physiology of adventitious roots. Plant Physiology 170:603\u0026ndash;617. https://doi.org/10.1104/pp.15.01360\u003c/li\u003e\n\u003cli\u003eSultana R, Vales MI, Saxena KB, et al (2013) Waterlogging tolerance in pigeonpea (\u003cem\u003eCajanus cajan\u003c/em\u003e (L.) Millsp.): genotypic variability and identification of tolerant genotypes. J Agric Sci 151:659\u0026ndash;671. https://doi.org/10.1017/S0021859612000755\u003c/li\u003e\n\u003cli\u003eThomas AL, Guerreiro SMC, Sodek L (2005) Aerenchyma formation and recovery from hypoxia of the flooded root system of nodulated soybean. Annals of Botany 96:1191\u0026ndash;1198. https://doi.org/10.1093/aob/mci272\u003c/li\u003e\n\u003cli\u003eTian L, Bi W, Liu X, et al (2019) Effects of waterlogging stress on the physiological response and grain-filling characteristics of spring maize (\u003cem\u003eZea mays\u003c/em\u003e L.) under field conditions. Acta Physiol Plant 41:63. https://doi.org/10.1007/s11738-019-2859-0\u003c/li\u003e\n\u003cli\u003eTian L, Zhang Y, Chen P, et al (2021) How does the waterlogging regime affect crop yield? A global meta-analysis. Front Plant Sci 12:634898. https://doi.org/10.3389/fpls.2021.634898\u003c/li\u003e\n\u003cli\u003eTukey JW (1949) Comparing individual means in the analysis of variance. Biometrics 5:99. https://doi.org/10.2307/3001913\u003c/li\u003e\n\u003cli\u003eUeno K, Takahashi H (1997) Varietal variation and physiological basis for inhibition of wheat seed germination after excessive water treatment. Euphytica 94:169\u0026ndash;173. https://doi.org/10.1023/A:1002976732395\u003c/li\u003e\n\u003cli\u003eVan Veen H, Akman M, Jamar DCL, et al (2014) Group VII Ethylene Response Factor diversification and regulation in four species from flood‐prone environments. Plant Cell \u0026amp; Environment 37:2421\u0026ndash;2432. https://doi.org/10.1111/pce.12302\u003c/li\u003e\n\u003cli\u003eVidoz ML, Loreti E, Mensuali A, et al (2010) Hormonal interplay during adventitious root formation in flooded tomato plants. The Plant Journal 63:551\u0026ndash;562. https://doi.org/10.1111/j.1365-313X.2010.04262.x\u003c/li\u003e\n\u003cli\u003eVisser EJW, Voesenek LACJ (2005) Acclimation to soil flooding\u0026ndash;sensing and signal-transduction. Plant Soil 274:197\u0026ndash;214. https://doi.org/10.1007/s11104-004-1650-0\u003c/li\u003e\n\u003cli\u003eVisser W, Cohen JD, Barendse CWM (1996) An ethylene-mediated increase in sensitivity to auxin induces adventitious root formation in flooded Rumex pahstris Sm. Plant Physiology 112: 1687-1692.\u003c/li\u003e\n\u003cli\u003eWang X, Deng Z, Zhang W, et al (2017) Effect of waterlogging duration at different growth stages on the growth, yield and quality of cotton. PLoS ONE 12:e0169029. https://doi.org/10.1371/journal.pone.0169029\u003c/li\u003e\n\u003cli\u003eWiraguna E, Malik AI, Erskine W (2017) Waterlogging tolerance in lentil (\u003cem\u003eLens culinaris\u003c/em\u003e Medik. subsp. culinaris) germplasm associated with geographic origin. Genet Resour Crop Evol 64:579\u0026ndash;586. https://doi.org/10.1007/s10722-016-0385-0\u003c/li\u003e\n\u003cli\u003eXiong R, Sang L, Liu R, et al (2020) Effects of waterlogging on maize seedling growth during seed germination. IOP Conf Ser: Earth Environ Sci 598:012075. https://doi.org/10.1088/1755-1315/598/1/012075\u003c/li\u003e\n\u003cli\u003eXu X, Ji J, Xu Q, et al (2018) The major‐effect quantitative trait locus \u003cem\u003e Cs ARN 6.1 \u003c/em\u003e encodes an AAA ATP ase domain‐containing protein that is associated with waterlogging stress tolerance by promoting adventitious root formation. The Plant Journal 93:917\u0026ndash;930. https://doi.org/10.1111/tpj.13819\u003c/li\u003e\n\u003cli\u003eYamauchi T, Abe F, Kawaguchi K, et al (2014) Adventitious roots of wheat seedlings that emerge in oxygen-deficient conditions have increased root diameters with highly developed lysigenous aerenchyma. Plant Signaling \u0026amp; Behavior 9:e28506. https://doi.org/10.4161/psb.28506\u003c/li\u003e\n\u003cli\u003eYates RJ, Howieson JG, Hungria M, et al (2016) Authentication of rhizobia and assessment of the legume symbiosis in controlled plant growth systems. In: Howieson JG, Dilworth MJ (eds) Working with rhizobia. Australian Centre for International Agricultural Research, Canberra, pp 73\u0026ndash;108\u003c/li\u003e\n\u003cli\u003eZaidi PH, Rashid Z, Vinayan M, Babu TA (2012) Pre-germination anaerobic stress tolerance in tropical maize (\u003cem\u003eZea mays\u003c/em\u003e L.). Aust J Crop Sci 6:1703\u0026ndash;1711\u003c/li\u003e\n\u003cli\u003eZaman MS, Malik AI, Kaur P, Erskine W (2018) Waterlogging tolerance of pea at germination. J Agronomy Crop Science 204:155\u0026ndash;164. https://doi.org/10.1111/jac.12230\u003c/li\u003e\n\u003cli\u003eZaman MSU, Malik AI, Kaur P, et al (2019) Waterlogging tolerance at germination in field pea: variability, genetic control, and indirect selection. Front Plant Sci 10:953. https://doi.org/10.3389/fpls.2019.00953\u003c/li\u003e\n\u003cli\u003eZaman MSU, Malik AI, Luna LN, et al (2024) Differences of waterlogging tolerance in winter pulse crop between emergence and vegetative stages. J Agronomy Crop Science 210:e12704. https://doi.org/10.1111/jac.12704\u003c/li\u003e\n\u003cli\u003eZhang XK, Chen J, Chen L, et al (2008) Imbibition behavior and flooding tolerance of rapeseed seed (\u003cem\u003eBrassica napus\u003c/em\u003e L.) with different testa color. Genet Resour Crop Evol 55:1175\u0026ndash;1184. https://doi.org/10.1007/s10722-008-9318-x\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1\u0026nbsp;\u003c/strong\u003eDays to flower, days to maturity and testa colour of four cultivars of two respective pulse crops\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"624\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.769230769230769%\" valign=\"top\"\u003e\n \u003cp\u003eSL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"35.57692307692308%\" valign=\"top\"\u003e\n \u003cp\u003eCrop species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"19.23076923076923%\" valign=\"top\"\u003e\n \u003cp\u003eCultivar\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.576923076923077%\" valign=\"top\"\u003e\n \u003cp\u003eDays to flower\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.538461538461538%\" valign=\"top\"\u003e\n \u003cp\u003eDays to maturity\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.307692307692307%\" valign=\"top\"\u003e\n \u003cp\u003eTesta color\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.769230769230769%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"35.57692307692308%\" valign=\"top\"\u003e\n \u003cp\u003eMungbean (\u003cem\u003eVigna radiata\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"19.23076923076923%\" valign=\"top\"\u003e\n \u003cp\u003eBARI Mung-6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.576923076923077%\" valign=\"top\"\u003e\n \u003cp\u003e35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.538461538461538%\" valign=\"top\"\u003e\n \u003cp\u003e70\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.307692307692307%\" valign=\"top\"\u003e\n \u003cp\u003eShiny green\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.769230769230769%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"35.57692307692308%\" valign=\"top\"\u003e\n \u003cp\u003eMungbean (\u003cem\u003eVigna radiata\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"19.23076923076923%\" valign=\"top\"\u003e\n \u003cp\u003eBARI Mung-8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.576923076923077%\" valign=\"top\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.538461538461538%\" valign=\"top\"\u003e\n \u003cp\u003e68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.307692307692307%\" valign=\"top\"\u003e\n \u003cp\u003eYellow\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.769230769230769%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"35.57692307692308%\" valign=\"top\"\u003e\n \u003cp\u003eMungbean (\u003cem\u003eVigna radiata\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"19.23076923076923%\" valign=\"top\"\u003e\n \u003cp\u003eBinamoog-8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.576923076923077%\" valign=\"top\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.538461538461538%\" valign=\"top\"\u003e\n \u003cp\u003e71\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.307692307692307%\" valign=\"top\"\u003e\n \u003cp\u003eShiny green\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"5.769230769230769%\" valign=\"top\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"35.57692307692308%\" valign=\"top\"\u003e\n \u003cp\u003eBlackgram (\u003cem\u003eVigna mungo\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"19.23076923076923%\" valign=\"top\"\u003e\n \u003cp\u003eBARI Mash-3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.576923076923077%\" valign=\"top\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.538461538461538%\" valign=\"top\"\u003e\n \u003cp\u003e72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.307692307692307%\" valign=\"top\"\u003e\n \u003cp\u003eBlack\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e Degrees of freedom (df), \u003cem\u003eF\u003c/em\u003e-values, and probabilities of two-way ANOVA at the reproductive stage during WL and recovery phases.\u0026nbsp;\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"628\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eCharacter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eSource of variation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003eWL duration\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003eCrop species (C)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003eWL X C\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003edf\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" colspan=\"5\" valign=\"top\"\u003e\n \u003cp\u003eWaterlogging Phase\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eSPAD chlorophyll content\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e27.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e18.76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e4.38\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003ePlant height (cm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e33.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e71.22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.89\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eShoot dry mass (g)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e17.80\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e6.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e1.12\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.37\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eTap root depth (cm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e17.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e1.48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e1.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e0.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.20\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eRoot dry mass (g plant\u003csup\u003e\u0026minus;1\u003c/sup\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e54.71\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e9.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e1.39\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.23\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eAdventitious roots\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e59.65\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e7.48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e4.71\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" colspan=\"5\" valign=\"top\"\u003e\n \u003cp\u003eRecovery Phase\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003ePlant height (cm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e6.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e37.21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.93\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.50\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eShoot dry mass (g)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e3.70\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e54.08\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.69\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.70\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eTap root depth (cm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e72.01 \u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e11.48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e4.21\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eRoot dry mass (g plant\u003csup\u003e\u0026minus;1\u003c/sup\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e37.95\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e99.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e1.58\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e0.16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003eNodulation score\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eF\u003c/em\u003e-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e76.95\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e32.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e6.53\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"34.07643312101911%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"21.97452229299363%\" valign=\"top\"\u003e\n \u003cp\u003eProbability\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.764331210191083%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.197452229299362%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.987261146496815%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","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":"Mungean, blackgram, waterlogging, recovery, nodulation, fibrous roots, yield","lastPublishedDoi":"10.21203/rs.3.rs-4569640/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4569640/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground and aims\u003c/h2\u003e \u003cp\u003eSummer pulse crops- mungbean and blackgram are increasingly exposed to waterlogging (WL). This study analyzed the effects of WL at germination and reproductive stages, and to identify the suitable cultivars for different cropping systems.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eThe study evaluated WL tolerance of three mungbean cultivars -BARI Mung-6, BARI Mung-8, Binamoog-8; and a blackgram cultivar- BARI Mash-3 at the germination and reproductive stages. The treatment levels at germination were drained control, 3, 5 and 7 d WL, while at reproductive stage were drained control, 3, 6 and 9 d WL.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eAt germination, % emergence was significantly reduced as WL duration increased. After 7 d WL, BARI Mash-3 displayed 38% emergence, while mungbean cultivars had 14\u0026ndash;18% emergence. At reproductive-stage, WL reduced plant height, tap root length, shoot and root dry mass compared to drained control. At recovery compared to WL of 9 d, the chlorophyll content increased by 15% in BARI Mash-3, while it decreased in mungbean cultivars. Shoot relative growth rate (RGR) of BARI Mash-3 was positive, while mungbean cultivars showed negative. Similarly, BARI Mash-3 had the higher root RGR than mungbean cultivars. BARI Mash-3 also produced the higher number of adventitious roots (27) than mungbean cultivars (10\u0026ndash;19). The seed yield was reduced at 40% in BARI Mash-3 while 52\u0026ndash;60% for mungbean cultivars.\u003c/p\u003e\u003ch2\u003eConclusions\u003c/h2\u003e \u003cp\u003eBlackgram cultivar is WL tolerant at both stages compared to mungbean cultivars. These new findings will allow to select suitable crops for different cropping systems based on the perceived risk of WL.\u003c/p\u003e","manuscriptTitle":"Differences of waterlogging tolerance between two grain legume species at germination and reproductive stages","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-01 05:14:39","doi":"10.21203/rs.3.rs-4569640/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":"0cb4b3f6-4c67-436a-bdf9-f2e325ada59f","owner":[],"postedDate":"July 1st, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-07-18T18:58:06+00:00","versionOfRecord":[],"versionCreatedAt":"2024-07-01 05:14:39","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4569640","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4569640","identity":"rs-4569640","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}