Current IssueEditorial BoardOnline First ArticlesArchive
Current IssueEditorial BoardOnline First ArticlesArchive

Full Research Article

Effects of Drought Stress During Branching and Pod Formation Stages on Morpho-Physiology and Yield in Soybean

F
Feifei Huang1
H
Hongwei Yuan2,3
J
Jiwei Yang2,3
X
Xiaoliang Li1,*
1College of Resource and Environment, Anhui Science and Technology University, Fengyang, Anhui-233 100, China.
2Anhui and Huaihe River Institute of Hydraulic Research, Hefei, Anhui-230 088, China.
3Anhui Provincial Key Laboratory of Water Science and Intelligent Water Conservancy, Bengbu, Anhui-233 000, China.

  • Submitted04-10-2025|

  • Accepted23-02-2026|

  • First Online 05-03-2026|

  • doi 10.18805/LRF-908

ABSTRACT

Background: Recurrent drought events in the Huaibei Plain of China pose a major constraint to soybean production. The peak drought period in July and September coincides with two critical  stages of soybean, branching (B) and pod formation (PF), yet current research still focuses predominantly on the flowering-podding (FP) stage, leaving the physiological and yield response mechanisms to drought stress during these stages poorly understood. This study was designed to investigate the differential responses of soybean to varying intensities of drought stress during critical growth stages.

Methods: A pot experiment was conducted under drought conditions in the Huaibei Plain in 2024, with three limits of soil water content set at both the B and PF stages: light drought (LD, 55%), moderate drought (MD, 45%) and well-watered control (CK, 75%). Morphological, photosynthetic, yield and pod-related traits of soybean under varying drought levels were examined and Pearson correlation analysis was applied to assess their relationships with yield.

Result: The results indicated that, overall, drought stress during the B stage resulted in severe reductions in plant height (PH), root length (RL) and seed moisture content (MC) compared to the PF stage. In contrast, photosynthetic parameters (SPAD value, stomatal conductance (Gs), net photosynthetic rate (Pn), transpiration rate (Tr) and intercellular CO2 concentration (Ci), yield, pod-related traits (total seed number (TSN), total pod number (TPN), filled pod number (FPN), empty pod number (EPN) and pod wall weight (PWW), as well as seed oil content (OC), exhibited an opposite trend. Furthermore, the magnitude of change in all aforementioned traits (except for EPN) relative to the CK followed the pattern MD > LD at the same growth stage. Correlation analysis showed that soybean yield was significantly and positively associated with TSN, TPN, FPN and PWW (p<0.05), while exhibiting negative correlations with EPN, Ci and MC.

INTRODUCTION

Amid the current trend of increasingly extreme global climate patterns, drought events are increasingly affecting agricultural production regions worldwide, characterized by their high frequency, prolonged duration and broad spatial extent (Mangena, 2020; Christian et al., 2023; Quat et al., 2025). In particular, it has resulted in a marked increase in drought risk and the extent of affected areas across varying regions in China (Song et al., 2021), imposing substantial economic losses on grain production. As one of the most important oil crops in China, soybean [Glycine max (L.) Merr.] is highly sensitive to water deficit during its growth stages, with severe drought potentially leading to yield losses of up to 40% (Zhou et al., 2022). However, China’s domestic soybean production remains at approximately 20.65 million tons per year (2024 statistics), while import dependence remains as high as 80%, undoubtedly creating a critical bottleneck for national strategic food security (Jiang et al., 2025). Therefore, evaluating soybean adaptation under drought stress is critical to mitigating yield losses.
       
Drought stress substantially influences the growth and development of various soybean organs (Dietz et al., 2021). Notably, the responses of phenotypic traits to such stress can serve as valuable indicators for selecting drought-resistant soybean varieties (Kim et al., 2023). Studies have indicated that crop responses to drought follow a progressive sequence from physiological alterations to growth adjustments and ultimately yield reduction, with photosynthesis-a key physiological process-being among the first to be adversely affected (Chaves et al., 2009). Moreover, drought stress occurring at any growth stage of soybean can impair photosynthetic performance, primarily due to reduced chlorophyll content, which is widely recognized as a major cause of alterations in photosynthetic characteristics (Wijewardana et al., 2019; Elsalahy and Reckling, 2022). These physiological alterations can disrupt the synthesis and transport of assimilates in flowers and pods and also adversely affect morphological traits such as plant height and root architecture (da Silva et al., 2018; García-Rodríguez et al., 2024). Ultimately, this results in reduced yield and quality parameters such as protein and oil content in soybean (Gholamhoseini et al., 2018).
       
The Huaibei Plain in Anhui Province, China, a major soybean-producing region with an annual summer sowing area of 700,000-800,000 hectares, is frequently affected by droughts that significantly limit grain yield (Cui et al., 2021). Droughts frequently occur in July and September in the Huaihe River Basin of Anhui Province, which overlap with the branching and pod formation stages of soybean. In contrast to previous studies focusing on the flowering-podding stage, this study examines soybean responses to drought during these critical phases in the Huaibei Plain, to support breeding of drought-resistant varieties and development of targeted irrigation strategies.

MATERIALS AND METHODS

Experimental materials and crop management
 
The pot-based drought experiment on soybean was conducted from June to September 2024 at the Xinmaqiao Comprehensive Agricultural Water Experimental Station (33°09′N, 117°22′E), situated 25 km north of Bengbu City, Anhui Province, China. Meteorological records indicate that the experimental site is located within the warm temperate semi-humid monsoon climate region of the Huang-Huai-Hai Plain, where the long-term average annual precipitation and evaporation measure 917 mm and 916 mm, respectively. The pot experiment utilized a lime concretion black soil of heavy texture and low permeability, containing >95% silt and clay. Key soil properties are provided in Table 1.

Table 1: Main physicochemical properties of the topsoil used in the experiment.


       
Based on the long-term agronomic experience of the experimental station, the soybean growth period in this study was categorized into four stages: seedling (S, 19 June-14 July), branching (B, 15-29 July), flowering-podding (FP, 30 July-20 August) and pod formation (PF, 21 August-28 September). Pots measuring 33 cm in top diameter, 22 cm in bottom diameter and 28 cm in height were used for sowing soybean seeds of the cultivar ‘Zhonghuang 13’ at a density of 8-10 seeds per pot. Seven days after emergence, seedlings were thinned to three uniform plants per pot. All plants were uniformly managed-including fertilization, pest and weed control-throughout the growing season. All potted plants were placed on outdoor shelves and were equipped with a movable rain shelter to minimize interference from natural precipitation during the experiment.
 
Experimental design
 
The experiment included two factors: growth stages (B and PF) and drought intensity, the latter was implemented by maintaining specific limits of soil moisture content (SMC) in the pots. Based on the station’s long-term agronomic experience and established protocols (Cui et al., 2021), three SMC levels were applied: well-watered control (CK, 75%), light drought (LD, 55%) and moderate drought (MD, 45%). The experiment comprised one control group and four drought-treated groups, with three replicates each, resulting in a total of 15 pots. Table 2 for the complete experimental layout.

Table 2: Experimental design for drought stress at different growth stages of soybean.


 
Morphological and yield traits measurement
 
At harvest, plant height (PH) and root length (RL) were measured using a 1-m standard ruler. The total pod number (TPN), filled pod number (FPN), empty pod number (EPN) and total seed number (TSN) were enumerated and recorded. Soybean grain yield and pod wall weight (PWW) were determined using an electronic balance.
 
SPAD and photosynthetic traits measurement

Measurements of relative chlorophyll content (SPAD) and photosynthetic traits-including intercellular CO2 concentration (Ci), stomatal conductance (Gs), net photosynthetic rate (Pn) and transpiration rate (Tr)-were conducted from 9:00 to 12:00 on sunny days during the late B and late PF stages using a SPAD-502 Plus chlorophyll meter (Konica Minolta Inc., Japan) and a CIRAS-4 portable photosynthesis system (PP Systems International Inc., USA), respectively.
 
Quality traits measurement
 
Oil content (OC) and moisture content (MC) of soybean seeds were determined using an Infratec™ near-infrared grain analyzer (FOSS Analytical A/S, Denmark).
 
Data processing and analysis
 
Experimental data were organized using Microsoft Excel 2010, followed by one-way ANOVA performed with IBM Statistics SPSS 27.0 software. Visualization including column charts, vertical multi-panel line plots and correlation heatmaps was generated using Origin 2021 software.

RESULTS AND DISCUSSION

Effects of drought stress on PH and RL in soybean
 
As shown in Fig 1a, PH declined with increasing drought intensity at the same growth stage, though the reduction was less pronounced during the PF stage compared to the B stage. All treatment values were lower than those of the CK, but no significant differences were detected (p>0.05). Specifically, PH in drought-stressed groups at the B stage (T1, T2) decreased by 9.97% and 15.03%, respectively, compared to CK, whereas at the PF stage (T3, T4), the reductions were about 5.55% and 6.04%. Greater drought intensity resulted in a more pronounced reduction in PH; however, as the PF stage coincides with limited vegetative growth in soybean, plants were less affected during this stage than at the B stage (Wei et al., 2018), consistent with our findings. Then, a similar response pattern was observed for RL under drought stress (Fig 1b). Compared with CK, RL in the B stage groups (T1, T2) declined by roughly 9.21% and 11.93%, whereas at the PF stage (T3, T4), the decreases were approximately 4.40% and 11.72%. No significant differences were found among any treatment groups (p>0.05). These results demonstrated that increasing drought severity exerts a more pronounced inhibitory effect on soybean root growth (García-Rodríguez et al., 2024). Ndlovu et al., (2025) observed a greater effect of drought stress on root length in sorghum seedlings at the early growth stage than at later stages, which aligns with the findings of this study. Therefore, MD was more inhibitory to PH and RL in soybean than LD. Since these traits were largely developed by the PF stage, drought stress during this phase had a much smaller effect compared to the B stage.

Fig 1: Effects of different drought treatments on PH and RL.


 
Effects of drought stress on SPAD and photosynthetic traits in soybean
 
Drought stress impairs plant photosynthesis primarily by triggering stomatal closure, which subsequently leads to the degradation of chlorophyll and disruption of the electron transport chain in chloroplasts (El Amine et al., 2025). As shown in Table 3, SPAD and photosynthetic traits (Ci, Gs, Pn, Tr) were generally higher during the B than the PF stage across all treatments. These traits showed no significant differences during the B stage, but all except SPAD differed significantly (p<0.05) during the PF stage. Compared with CK, the SPAD values in T1 and T2 decreased by approximately 5.73% and 10.53%, respectively, during the B stage, while greater reductions of about 10.13% and 18.79% were observed in T3 and T4 at the PF stage. Previous studies have reported a marked decline in soybean SPAD values with progression of growth stages and intensification of drought stress (Elsalahy and Reckling, 2022), a trend consistent with the findings of the present study. In terms of photosynthetic traits, compared with CK, the treatment groups (T1, T2) at the B stage exhibited decreases in Gs, Pn and Tr of approximately (36.94%, 0.36%), (52.25%, 86.25%) and (-1.78%, 8.77%), respectively. In contrast, at the PF stage, the treatment groups (T3, T4) showed reductions in these parameters of about (21.90%, 73.20%), (52.43%, 91.94%) and (26.27%, 70.90%), respectively. Liu et al., (2019) reported that Gs, Pn and Tr in persimmon exhibited a decreasing trend with increasing intensity of PEG-6000-simulated drought stress, a response consistent with the observations in the present study. However, in both the B (T1, T2) and PF (T3, T4) stage treatment groups of soybean, the Ci increased rather than decreased with intensifying drought stress when compared to CK, with specific increases of (48.20%, 61.15%) and (11.47%, 29.07%), respectively. Fugate et al., (2018) observed a similar response pattern of the Ci to drought stress in sugar beet, as did Liu et al., (2024) in Quercus acutissima.

Table 3: Effects of different drought treatments on SPAD and photosynthetic traits.



Effects of drought stress on yield and pod traits in soybean
 
Soybean pods and seeds, serving as the primary water-storing organs, critically influence grain yield through their size and number per plant (Desclaux et al., 2000). As shown in Table 4, with the progression of growth stages and the intensification of drought stress, soybean yield, TSN, TPN, FPN and PWW displayed a decreasing trend (i.e., T1 > T2 > T3 > T4). Compared to CK, all five traits showed reductions, with percentage decreases from T1 to T4 as follows: yield (7.09%, 16.48%, 42.59%, 44.36%), TSN (20.38%, 23.27%, 39.23%, 45.58%), TPN (9.69%, 14.96%, 28.03%, 31.08%), FPN (12.81%, 15.31%, 30.24%, 34.34%) and PWW (8.18%, 23.26%, 37.70%, 45.69%). Furthermore, these traits exhibited significant differences (p<0.05) from CK under drought stress at the PF stage. In contrast to other traits, the EPN in soybean showed an inverse trend under drought stress, with T1-T4 increasing by 100.00%, 12.36%, 174.16% and 83.15% compared to CK, though only T3 exhibited a significant difference (p>0.05). Previous studies (Wei et al., 2018; Cui et al., 2019; Kisman et al., 2025) have shown that drought stress significantly reduces soybean yield and pod traits, with water deficit during the reproductive stage having a greater impact than during the vegetative stage, consistent with our findings. Increasing drought intensity further amplified these effects, with the PF stage being more affected than the B stage.

Table 4: Effects of different drought treatments on yield and pod traits.


 
Effects of drought stress on quality in soybean
 
Within the same growth stage, both OC and MC of soybean seeds decreased with increasing drought stress intensity (Fig 2). Compared with CK, the OC in soybean kernels increased by approximately 1.43% and 0.84% under the T1 and T2 treatments at the B stage, respectively, but decreased by 1.63% and 4.19% under T3 and T4 at the PF stage. Meanwhile, the MC increased by 1.71%, 0.85%, 3.08% and 0.09% under T1-T4 across both the B and PF stages, respectively. Haghaninia et al., (2025) reported that severe drought stress leads to a reduction in OC compared with optimal irrigation conditions. In the present study, drought stress applied at the B stage increased the OC of soybean seeds, whereas a reduction was observed under drought at the PF stage. Furthermore, Kouighat et al., (2025) observed an increase in seed MC in sesame under drought stress conditions, which aligns with our results. Therefore, drought stress at the B stage enhanced the OC of soybean seeds, while that applied at the PF stage led to a reduction. In contrast, seed MC increased under drought stress regardless of the growth stage.

Fig 2: Effects of different drought treatments on OC and MC.



Correlation analysis between soybean yield and agronomic traits
 
Pearson correlation analysis showed that yield was positively associated with most traits except MC, Ci and EPN (Fig 3). Strong significant correlations (r = 0.95-0.99, p<0.05) were observed with pod-related traits (TPN, FPN, TSN, PWW), while correlations with photosynthetic traits were positive but not significant, consistent with previous studies (Aminian et al., 2007; Dou et al., 2023; Basavarajeshwari et al., 2025). 

Fig 3: Correlation analysis among agronomic traits of soybean under drought stress.

CONCLUSION

Pot experiments under drought stress in the Huaibei Plain showed that drought at the B stage mainly affected PH and RL, while at the PF stage it had greater impacts on photosynthetic traits, yield and pod-related traits. OC increased at B but declined at PF, whereas MC rose consistently. Moderate drought caused more severe effects than light drought. Yield correlated strongly with TPN, FPN, TSN and PWW, highlighting the importance of optimizing irrigation during the PF stage.

ACKNOWLEDGEMENT

The present study was supported by the Sub-project of the National Key Research and Development Program of China (Grant No. 2018YFD0300901-2) and the Natural Science Foundation of Anhui Province (Grant Nos. 2208085US03, 2208085US15).
 
Disclaimers
 
The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.

CONFLICT OF INTEREST

The authors declare that there are no conflicts of interest regarding the publication of this article. No funding or sponsorship influenced the design of the study, data collection, analysis, decision to publish, or preparation of the manuscript.

REFERENCES


  1. Aminian, R., Khodambashi, M. and Yadegari, M. (2007). Study of seed yield correlation with different traits of common bean under stress condition. In: International Conference on Mathematical Biology (ICMB07). pp 255.

  2. Basavarajeshwari, H., Kuchanur, P.H., Zaidi, P.H., Vinayan, M.T., Patil, A., Patil, R.P., Nidagundi, J.M. and Arunkumar, B. (2025). Performance assessment of F2:3 testcrosses of maize (Zea mays L.) for physiological traits Vis-a-Vis grain yield under heat stress and drought conditions. Journal of Agronomy and Crop Science. 211(2). doi: 10.1111/jac.70027.

  3. Chaves, M.M., Flexas, J. and Pinheiro, C. (2009). Photosynthesis under drought and salt stress: Regulation mechanisms from whole plant to cell. Annals of Botany. 103(4): 551- 560. doi: 10.1093/aob/mcn125.

  4. Christian, J.I., Martin, E.R., Basara, J.B., Furtado, J.C., Otkin, J.A., Lowman, L.E.L., Hunt, E.D., Mishra, V. and Xiao, X.M. (2023). Global projections of flash drought show increased risk in a warming climate. Communications Earth and Environment. 4(1). doi: 10.1038/s43247-023- 00826-1.

  5. Cui, Y., Jiang, S.M., Jin, J.L., Ning, S.W. and Feng, P. (2019). Quantitative assessment of soybean drought loss sensitivity at different growth stages based on S-shaped damage curve. Agricultural Water Management. 213: -832. doi: 10.1016/j.agwat.2018.11.020.

  6. Cui, Y., Ning, S.W., Jin, J.L., Jiang, S.M., Zhou, Y.L. and Wu, C.G. (2021). Quantitative lasting effects of drought stress at a growth stage on soybean evapotranspiration and aboveground biomass. Water. 13(1). doi: 10.3390/ w13010018.

  7. da Silva, A.J., Magalhaes, J.R., Sales, C.R.G., Pires, R.C.D. and Machado, E.C. (2018). Source-sink relationships in two soybean cultivars with indeterminate growth under water deficit. Bragantia. 77(1): 23-35. doi: 10.1590/1678- 4499.2017010.

  8. Desclaux, D., Huynh, T.T. and Roumet, P. (2000). Identification of soybean plant characteristics that indicate the timing of drought stress. Crop Science. 40(3): 716-722. doi: 10.2135/cropsci2000.403716x.

  9. Dietz, K.J., Zörb, C. and Geilfus, C.M. (2021). Drought and crop yield. Plant Biology. 23(6): 881-893. doi: 10.1111/plb.13304.

  10. Dou, Z.Y., Feng, H.L., Zhang, H., Abdelghany, A.E., Zhang, F.C., Li, Z.J. and Fan, J.L. (2023). Silicon application mitigated the adverse effects of salt stress and deficit irrigation on drip-irrigated greenhouse tomato. Agricultural Water Management. 289. doi: 10.1016/j.agwat.2023.108526.

  11. El Amine, B., Mosseddaq, F., Houssa, A.A., Bouaziz, A., Moughli, L. and Oukarroum, A. (2025). Physiological and agronomic effects of regulated-deficit irrigation on soybean grown under arid climatic conditions. Crop Journal. 13(1): 281- 291. doi: 10.1016/j.cj.2024.10.011.

  12. Elsalahy, H.H. and Reckling, M. (2022). Soybean resilience to drought is supported by partial recovery of photosynthetic traits. Frontiers in Plant Science. 13. doi: 10.3389/ fpls.2022.971893.

  13. Fugate, K.K., Lafta, A.M., Eide, J.D., Li, G.L., Lulai, E.C., Olson, L.L., Deckard, E.L., Khan, M.F.R. and Finger, F.L. (2018). Methyl jasmonate alleviates drought stress in young sugar beet (Beta vulgaris L.) plants. Journal of Agronomy and Crop Science. 204(6): 566-576. doi: 10.1111/ jac.12286.

  14. García-Rodríguez, J.C., Manzo-Valencia, M.K., Olalde-Portugal, V. and Valdés-Rodríguez, S.E. (2024). Exploring drought responses in mexican soybeans: plant water status, shoot and root biomass and root system architecture. Food and Energy Security. 13(6). doi: 10.1002/fes3.70017.

  15. Gholamhoseini, M., Ebrahimian, E., Habibzadeh, F., Ataei, R. and Dezfulizadeh, M.S. (2018). Interactions of shading conditions and irrigation regimes on photosynthetic traits and seed yield of soybean (Glycine max L.). Legume Research. 41(2): 230-238. doi: 10.18805/lr-359.

  16. Haghaninia, M., Mashhouri, S.M., Najafifar, A., Soleimani, F. and Wu, Q.S. (2025). Combined effects of zinc oxide nanoparticles and arbuscular mycorrhizal fungi on soybean yield, oil quality and biochemical responses under drought stress. Future Foods. 11. doi: 10.1016/ j.fufo.2025.100594.

  17. Jiang, X.Y., Li, H.S., Dai, X., Li, J.D. and Liu, Y. (2025). An empirical analysis of global soybean supply potential and china’s diversified import strategies based on global agro- ecological zones and multi-objective nonlinear programming models. Agriculture-Basel. 15(5). doi: 10.3390/agriculture 15050529.

  18. Kim, J., Lee, C.W., Park, J.E., Mansoor, S., Chung, Y.S. and Kim, K. (2023). Drought stress restoration frequencies of phenotypic indicators in early vegetative stages of soybean (Glycine max L.). Sustainability. 15(6). doi: 10.3390/su15064852.

  19. Kisman, Hemon, A.F., Listiana, B.E., Dewi, S.M. and Mustikarini, E.D. (2025). Evaluation of drought tolerance and adaptation of large-seeded soybean genotypes under various drought stress levels. Indian Journal of Agricultural Research. 59(8): 1189-1200. doi: 10.18805/IJARe.AF-929.

  20. Kouighat, M., Moussaoui, F.E., Adiba, A., Hafid, A., Bouchyoua, A., El Fechtali, M. and Nabloussi, A. (2025). Evaluation and selection of sesame mutants (Sesamum indicum L.) for optimal nutritional profiles in seeds under field drought conditions. Ocl-Oilseeds and Fats Crops and Lipids. 32. doi: 10.1051/ocl/2025004.

  21. Liu, D., Guo, H.L., Yan, L.P., Gao, L., Zhai, S.S. and Xu, Y. (2024). Physiological, photosynthetic and stomatal ultrastructural responses of quercus acutissima seedlings to drought stress and rewatering. Forests. 15(1). doi: 10.3390/ f15010071.

  22. Liu, Z.C., Bao, D.E., Hu, H.L. and Miao, Y.C. (2019). Changes of photosynthetic characteristics in persimmon (Diospyros Spp.) rootstocks during drought stress simulated by PEG-6000. Bangladesh Journal of Botany. 48(3): 913- 918.

  23. Mangena, P. (2020). Effect of hormonal seed priming on germination, growth, yield and biomass allocation in soybean grown under induced drought stress. Indian Journal of Agricultural Research. 54(5): 592-598. doi: 10.18805/IJARe.A-441.

  24. Ndlovu, E., Maphosa, M. and van Staden, J. (2025). Morphological responses of sorghum seedlings to drought, heat and combined stresses. Experimental Agriculture. 61. doi: 10.1017/s0014479725100082.

  25. Quat, N.N., Dang, B.Q., Trang, N.T., Nhung, N.T. and Cuc, L.M. (2025). Evaluating drought tolerance in common beans using drought indices and molecular markers. Indian Journal of Agricultural Research. 59(8): 1177-1183. doi: 10.18805/IJARe.AF-940.

  26. Song, Y.L., Tian, J.F., Linderholm, H.W., Wang, C.Y., Ou, Z.R. and Chen, D.L. (2021). The contributions of climate change and production area expansion to drought risk for maize in China over the last four decades. International Journal of Climatology. 41: E2851-E2862. doi: 10.1002/joc.6885.

  27. Wei, Y.Q., Jin, J.L., Jiang, S.M., Ning, S.W. and Liu, L. (2018). Quantitative response of soybean development and yield to drought stress during different growth stages in the huaibei plain, China. Agronomy-Basel. 8(7). doi: 10.3390/ agronomy8070097.

  28. Wijewardana, C., Alsajri, F.A., Irby, J.T., Krutz, L.J., Golden, B., Henry, W.B., Gao, W. and Reddy, K.R. (2019). Physiological assessment of water deficit in soybean using midday leaf water potential and spectral features. Journal of Plant Interactions. 14(1): 533-543. doi: 10.1080/17429145. 2019.1662499.

  29. Zhou, Q., Li, Y.P., Wang, X.J., Yan, C., Ma, C.M., Liu, J. and Dong, S.K. (2022). Effects of different drought degrees on physiological characteristics and endogenous hormones of soybean. Plants-Basel. 11(17). doi: 10.3390/plants 11172282.
Published In
Legume Research
Article Metrics
Metrics Icon
0
Views
0
Citations
Reviewed By
Share Article
Download Article
In this Article
    Contact us
    Follow us

    Full Research Article

    Effects of Drought Stress During Branching and Pod Formation Stages on Morpho-Physiology and Yield in Soybean

    F
    Feifei Huang1
    H
    Hongwei Yuan2,3
    J
    Jiwei Yang2,3
    X
    Xiaoliang Li1,*
    1College of Resource and Environment, Anhui Science and Technology University, Fengyang, Anhui-233 100, China.
    2Anhui and Huaihe River Institute of Hydraulic Research, Hefei, Anhui-230 088, China.
    3Anhui Provincial Key Laboratory of Water Science and Intelligent Water Conservancy, Bengbu, Anhui-233 000, China.

    • Submitted04-10-2025|

    • Accepted23-02-2026|

    • First Online 05-03-2026|

    • doi 10.18805/LRF-908

    ABSTRACT

    Background: Recurrent drought events in the Huaibei Plain of China pose a major constraint to soybean production. The peak drought period in July and September coincides with two critical  stages of soybean, branching (B) and pod formation (PF), yet current research still focuses predominantly on the flowering-podding (FP) stage, leaving the physiological and yield response mechanisms to drought stress during these stages poorly understood. This study was designed to investigate the differential responses of soybean to varying intensities of drought stress during critical growth stages.

    Methods: A pot experiment was conducted under drought conditions in the Huaibei Plain in 2024, with three limits of soil water content set at both the B and PF stages: light drought (LD, 55%), moderate drought (MD, 45%) and well-watered control (CK, 75%). Morphological, photosynthetic, yield and pod-related traits of soybean under varying drought levels were examined and Pearson correlation analysis was applied to assess their relationships with yield.

    Result: The results indicated that, overall, drought stress during the B stage resulted in severe reductions in plant height (PH), root length (RL) and seed moisture content (MC) compared to the PF stage. In contrast, photosynthetic parameters (SPAD value, stomatal conductance (Gs), net photosynthetic rate (Pn), transpiration rate (Tr) and intercellular CO2 concentration (Ci), yield, pod-related traits (total seed number (TSN), total pod number (TPN), filled pod number (FPN), empty pod number (EPN) and pod wall weight (PWW), as well as seed oil content (OC), exhibited an opposite trend. Furthermore, the magnitude of change in all aforementioned traits (except for EPN) relative to the CK followed the pattern MD > LD at the same growth stage. Correlation analysis showed that soybean yield was significantly and positively associated with TSN, TPN, FPN and PWW (p<0.05), while exhibiting negative correlations with EPN, Ci and MC.

    INTRODUCTION

    Amid the current trend of increasingly extreme global climate patterns, drought events are increasingly affecting agricultural production regions worldwide, characterized by their high frequency, prolonged duration and broad spatial extent (Mangena, 2020; Christian et al., 2023; Quat et al., 2025). In particular, it has resulted in a marked increase in drought risk and the extent of affected areas across varying regions in China (Song et al., 2021), imposing substantial economic losses on grain production. As one of the most important oil crops in China, soybean [Glycine max (L.) Merr.] is highly sensitive to water deficit during its growth stages, with severe drought potentially leading to yield losses of up to 40% (Zhou et al., 2022). However, China’s domestic soybean production remains at approximately 20.65 million tons per year (2024 statistics), while import dependence remains as high as 80%, undoubtedly creating a critical bottleneck for national strategic food security (Jiang et al., 2025). Therefore, evaluating soybean adaptation under drought stress is critical to mitigating yield losses.
           
    Drought stress substantially influences the growth and development of various soybean organs (Dietz et al., 2021). Notably, the responses of phenotypic traits to such stress can serve as valuable indicators for selecting drought-resistant soybean varieties (Kim et al., 2023). Studies have indicated that crop responses to drought follow a progressive sequence from physiological alterations to growth adjustments and ultimately yield reduction, with photosynthesis-a key physiological process-being among the first to be adversely affected (Chaves et al., 2009). Moreover, drought stress occurring at any growth stage of soybean can impair photosynthetic performance, primarily due to reduced chlorophyll content, which is widely recognized as a major cause of alterations in photosynthetic characteristics (Wijewardana et al., 2019; Elsalahy and Reckling, 2022). These physiological alterations can disrupt the synthesis and transport of assimilates in flowers and pods and also adversely affect morphological traits such as plant height and root architecture (da Silva et al., 2018; García-Rodríguez et al., 2024). Ultimately, this results in reduced yield and quality parameters such as protein and oil content in soybean (Gholamhoseini et al., 2018).
           
    The Huaibei Plain in Anhui Province, China, a major soybean-producing region with an annual summer sowing area of 700,000-800,000 hectares, is frequently affected by droughts that significantly limit grain yield (Cui et al., 2021). Droughts frequently occur in July and September in the Huaihe River Basin of Anhui Province, which overlap with the branching and pod formation stages of soybean. In contrast to previous studies focusing on the flowering-podding stage, this study examines soybean responses to drought during these critical phases in the Huaibei Plain, to support breeding of drought-resistant varieties and development of targeted irrigation strategies.

    MATERIALS AND METHODS

    Experimental materials and crop management
     
    The pot-based drought experiment on soybean was conducted from June to September 2024 at the Xinmaqiao Comprehensive Agricultural Water Experimental Station (33°09′N, 117°22′E), situated 25 km north of Bengbu City, Anhui Province, China. Meteorological records indicate that the experimental site is located within the warm temperate semi-humid monsoon climate region of the Huang-Huai-Hai Plain, where the long-term average annual precipitation and evaporation measure 917 mm and 916 mm, respectively. The pot experiment utilized a lime concretion black soil of heavy texture and low permeability, containing >95% silt and clay. Key soil properties are provided in Table 1.

    Table 1: Main physicochemical properties of the topsoil used in the experiment.


           
    Based on the long-term agronomic experience of the experimental station, the soybean growth period in this study was categorized into four stages: seedling (S, 19 June-14 July), branching (B, 15-29 July), flowering-podding (FP, 30 July-20 August) and pod formation (PF, 21 August-28 September). Pots measuring 33 cm in top diameter, 22 cm in bottom diameter and 28 cm in height were used for sowing soybean seeds of the cultivar ‘Zhonghuang 13’ at a density of 8-10 seeds per pot. Seven days after emergence, seedlings were thinned to three uniform plants per pot. All plants were uniformly managed-including fertilization, pest and weed control-throughout the growing season. All potted plants were placed on outdoor shelves and were equipped with a movable rain shelter to minimize interference from natural precipitation during the experiment.
     
    Experimental design
     
    The experiment included two factors: growth stages (B and PF) and drought intensity, the latter was implemented by maintaining specific limits of soil moisture content (SMC) in the pots. Based on the station’s long-term agronomic experience and established protocols (Cui et al., 2021), three SMC levels were applied: well-watered control (CK, 75%), light drought (LD, 55%) and moderate drought (MD, 45%). The experiment comprised one control group and four drought-treated groups, with three replicates each, resulting in a total of 15 pots. Table 2 for the complete experimental layout.

    Table 2: Experimental design for drought stress at different growth stages of soybean.


     
    Morphological and yield traits measurement
     
    At harvest, plant height (PH) and root length (RL) were measured using a 1-m standard ruler. The total pod number (TPN), filled pod number (FPN), empty pod number (EPN) and total seed number (TSN) were enumerated and recorded. Soybean grain yield and pod wall weight (PWW) were determined using an electronic balance.
     
    SPAD and photosynthetic traits measurement

    Measurements of relative chlorophyll content (SPAD) and photosynthetic traits-including intercellular CO2 concentration (Ci), stomatal conductance (Gs), net photosynthetic rate (Pn) and transpiration rate (Tr)-were conducted from 9:00 to 12:00 on sunny days during the late B and late PF stages using a SPAD-502 Plus chlorophyll meter (Konica Minolta Inc., Japan) and a CIRAS-4 portable photosynthesis system (PP Systems International Inc., USA), respectively.
     
    Quality traits measurement
     
    Oil content (OC) and moisture content (MC) of soybean seeds were determined using an Infratec™ near-infrared grain analyzer (FOSS Analytical A/S, Denmark).
     
    Data processing and analysis
     
    Experimental data were organized using Microsoft Excel 2010, followed by one-way ANOVA performed with IBM Statistics SPSS 27.0 software. Visualization including column charts, vertical multi-panel line plots and correlation heatmaps was generated using Origin 2021 software.

    RESULTS AND DISCUSSION

    Effects of drought stress on PH and RL in soybean
     
    As shown in Fig 1a, PH declined with increasing drought intensity at the same growth stage, though the reduction was less pronounced during the PF stage compared to the B stage. All treatment values were lower than those of the CK, but no significant differences were detected (p>0.05). Specifically, PH in drought-stressed groups at the B stage (T1, T2) decreased by 9.97% and 15.03%, respectively, compared to CK, whereas at the PF stage (T3, T4), the reductions were about 5.55% and 6.04%. Greater drought intensity resulted in a more pronounced reduction in PH; however, as the PF stage coincides with limited vegetative growth in soybean, plants were less affected during this stage than at the B stage (Wei et al., 2018), consistent with our findings. Then, a similar response pattern was observed for RL under drought stress (Fig 1b). Compared with CK, RL in the B stage groups (T1, T2) declined by roughly 9.21% and 11.93%, whereas at the PF stage (T3, T4), the decreases were approximately 4.40% and 11.72%. No significant differences were found among any treatment groups (p>0.05). These results demonstrated that increasing drought severity exerts a more pronounced inhibitory effect on soybean root growth (García-Rodríguez et al., 2024). Ndlovu et al., (2025) observed a greater effect of drought stress on root length in sorghum seedlings at the early growth stage than at later stages, which aligns with the findings of this study. Therefore, MD was more inhibitory to PH and RL in soybean than LD. Since these traits were largely developed by the PF stage, drought stress during this phase had a much smaller effect compared to the B stage.

    Fig 1: Effects of different drought treatments on PH and RL.


     
    Effects of drought stress on SPAD and photosynthetic traits in soybean
     
    Drought stress impairs plant photosynthesis primarily by triggering stomatal closure, which subsequently leads to the degradation of chlorophyll and disruption of the electron transport chain in chloroplasts (El Amine et al., 2025). As shown in Table 3, SPAD and photosynthetic traits (Ci, Gs, Pn, Tr) were generally higher during the B than the PF stage across all treatments. These traits showed no significant differences during the B stage, but all except SPAD differed significantly (p<0.05) during the PF stage. Compared with CK, the SPAD values in T1 and T2 decreased by approximately 5.73% and 10.53%, respectively, during the B stage, while greater reductions of about 10.13% and 18.79% were observed in T3 and T4 at the PF stage. Previous studies have reported a marked decline in soybean SPAD values with progression of growth stages and intensification of drought stress (Elsalahy and Reckling, 2022), a trend consistent with the findings of the present study. In terms of photosynthetic traits, compared with CK, the treatment groups (T1, T2) at the B stage exhibited decreases in Gs, Pn and Tr of approximately (36.94%, 0.36%), (52.25%, 86.25%) and (-1.78%, 8.77%), respectively. In contrast, at the PF stage, the treatment groups (T3, T4) showed reductions in these parameters of about (21.90%, 73.20%), (52.43%, 91.94%) and (26.27%, 70.90%), respectively. Liu et al., (2019) reported that Gs, Pn and Tr in persimmon exhibited a decreasing trend with increasing intensity of PEG-6000-simulated drought stress, a response consistent with the observations in the present study. However, in both the B (T1, T2) and PF (T3, T4) stage treatment groups of soybean, the Ci increased rather than decreased with intensifying drought stress when compared to CK, with specific increases of (48.20%, 61.15%) and (11.47%, 29.07%), respectively. Fugate et al., (2018) observed a similar response pattern of the Ci to drought stress in sugar beet, as did Liu et al., (2024) in Quercus acutissima.

    Table 3: Effects of different drought treatments on SPAD and photosynthetic traits.



    Effects of drought stress on yield and pod traits in soybean
     
    Soybean pods and seeds, serving as the primary water-storing organs, critically influence grain yield through their size and number per plant (Desclaux et al., 2000). As shown in Table 4, with the progression of growth stages and the intensification of drought stress, soybean yield, TSN, TPN, FPN and PWW displayed a decreasing trend (i.e., T1 > T2 > T3 > T4). Compared to CK, all five traits showed reductions, with percentage decreases from T1 to T4 as follows: yield (7.09%, 16.48%, 42.59%, 44.36%), TSN (20.38%, 23.27%, 39.23%, 45.58%), TPN (9.69%, 14.96%, 28.03%, 31.08%), FPN (12.81%, 15.31%, 30.24%, 34.34%) and PWW (8.18%, 23.26%, 37.70%, 45.69%). Furthermore, these traits exhibited significant differences (p<0.05) from CK under drought stress at the PF stage. In contrast to other traits, the EPN in soybean showed an inverse trend under drought stress, with T1-T4 increasing by 100.00%, 12.36%, 174.16% and 83.15% compared to CK, though only T3 exhibited a significant difference (p>0.05). Previous studies (Wei et al., 2018; Cui et al., 2019; Kisman et al., 2025) have shown that drought stress significantly reduces soybean yield and pod traits, with water deficit during the reproductive stage having a greater impact than during the vegetative stage, consistent with our findings. Increasing drought intensity further amplified these effects, with the PF stage being more affected than the B stage.

    Table 4: Effects of different drought treatments on yield and pod traits.


     
    Effects of drought stress on quality in soybean
     
    Within the same growth stage, both OC and MC of soybean seeds decreased with increasing drought stress intensity (Fig 2). Compared with CK, the OC in soybean kernels increased by approximately 1.43% and 0.84% under the T1 and T2 treatments at the B stage, respectively, but decreased by 1.63% and 4.19% under T3 and T4 at the PF stage. Meanwhile, the MC increased by 1.71%, 0.85%, 3.08% and 0.09% under T1-T4 across both the B and PF stages, respectively. Haghaninia et al., (2025) reported that severe drought stress leads to a reduction in OC compared with optimal irrigation conditions. In the present study, drought stress applied at the B stage increased the OC of soybean seeds, whereas a reduction was observed under drought at the PF stage. Furthermore, Kouighat et al., (2025) observed an increase in seed MC in sesame under drought stress conditions, which aligns with our results. Therefore, drought stress at the B stage enhanced the OC of soybean seeds, while that applied at the PF stage led to a reduction. In contrast, seed MC increased under drought stress regardless of the growth stage.

    Fig 2: Effects of different drought treatments on OC and MC.



    Correlation analysis between soybean yield and agronomic traits
     
    Pearson correlation analysis showed that yield was positively associated with most traits except MC, Ci and EPN (Fig 3). Strong significant correlations (r = 0.95-0.99, p<0.05) were observed with pod-related traits (TPN, FPN, TSN, PWW), while correlations with photosynthetic traits were positive but not significant, consistent with previous studies (Aminian et al., 2007; Dou et al., 2023; Basavarajeshwari et al., 2025). 

    Fig 3: Correlation analysis among agronomic traits of soybean under drought stress.

    CONCLUSION

    Pot experiments under drought stress in the Huaibei Plain showed that drought at the B stage mainly affected PH and RL, while at the PF stage it had greater impacts on photosynthetic traits, yield and pod-related traits. OC increased at B but declined at PF, whereas MC rose consistently. Moderate drought caused more severe effects than light drought. Yield correlated strongly with TPN, FPN, TSN and PWW, highlighting the importance of optimizing irrigation during the PF stage.

    ACKNOWLEDGEMENT

    The present study was supported by the Sub-project of the National Key Research and Development Program of China (Grant No. 2018YFD0300901-2) and the Natural Science Foundation of Anhui Province (Grant Nos. 2208085US03, 2208085US15).
     
    Disclaimers
     
    The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.

    CONFLICT OF INTEREST

    The authors declare that there are no conflicts of interest regarding the publication of this article. No funding or sponsorship influenced the design of the study, data collection, analysis, decision to publish, or preparation of the manuscript.

    REFERENCES


    1. Aminian, R., Khodambashi, M. and Yadegari, M. (2007). Study of seed yield correlation with different traits of common bean under stress condition. In: International Conference on Mathematical Biology (ICMB07). pp 255.

    2. Basavarajeshwari, H., Kuchanur, P.H., Zaidi, P.H., Vinayan, M.T., Patil, A., Patil, R.P., Nidagundi, J.M. and Arunkumar, B. (2025). Performance assessment of F2:3 testcrosses of maize (Zea mays L.) for physiological traits Vis-a-Vis grain yield under heat stress and drought conditions. Journal of Agronomy and Crop Science. 211(2). doi: 10.1111/jac.70027.

    3. Chaves, M.M., Flexas, J. and Pinheiro, C. (2009). Photosynthesis under drought and salt stress: Regulation mechanisms from whole plant to cell. Annals of Botany. 103(4): 551- 560. doi: 10.1093/aob/mcn125.

    4. Christian, J.I., Martin, E.R., Basara, J.B., Furtado, J.C., Otkin, J.A., Lowman, L.E.L., Hunt, E.D., Mishra, V. and Xiao, X.M. (2023). Global projections of flash drought show increased risk in a warming climate. Communications Earth and Environment. 4(1). doi: 10.1038/s43247-023- 00826-1.

    5. Cui, Y., Jiang, S.M., Jin, J.L., Ning, S.W. and Feng, P. (2019). Quantitative assessment of soybean drought loss sensitivity at different growth stages based on S-shaped damage curve. Agricultural Water Management. 213: -832. doi: 10.1016/j.agwat.2018.11.020.

    6. Cui, Y., Ning, S.W., Jin, J.L., Jiang, S.M., Zhou, Y.L. and Wu, C.G. (2021). Quantitative lasting effects of drought stress at a growth stage on soybean evapotranspiration and aboveground biomass. Water. 13(1). doi: 10.3390/ w13010018.

    7. da Silva, A.J., Magalhaes, J.R., Sales, C.R.G., Pires, R.C.D. and Machado, E.C. (2018). Source-sink relationships in two soybean cultivars with indeterminate growth under water deficit. Bragantia. 77(1): 23-35. doi: 10.1590/1678- 4499.2017010.

    8. Desclaux, D., Huynh, T.T. and Roumet, P. (2000). Identification of soybean plant characteristics that indicate the timing of drought stress. Crop Science. 40(3): 716-722. doi: 10.2135/cropsci2000.403716x.

    9. Dietz, K.J., Zörb, C. and Geilfus, C.M. (2021). Drought and crop yield. Plant Biology. 23(6): 881-893. doi: 10.1111/plb.13304.

    10. Dou, Z.Y., Feng, H.L., Zhang, H., Abdelghany, A.E., Zhang, F.C., Li, Z.J. and Fan, J.L. (2023). Silicon application mitigated the adverse effects of salt stress and deficit irrigation on drip-irrigated greenhouse tomato. Agricultural Water Management. 289. doi: 10.1016/j.agwat.2023.108526.

    11. El Amine, B., Mosseddaq, F., Houssa, A.A., Bouaziz, A., Moughli, L. and Oukarroum, A. (2025). Physiological and agronomic effects of regulated-deficit irrigation on soybean grown under arid climatic conditions. Crop Journal. 13(1): 281- 291. doi: 10.1016/j.cj.2024.10.011.

    12. Elsalahy, H.H. and Reckling, M. (2022). Soybean resilience to drought is supported by partial recovery of photosynthetic traits. Frontiers in Plant Science. 13. doi: 10.3389/ fpls.2022.971893.

    13. Fugate, K.K., Lafta, A.M., Eide, J.D., Li, G.L., Lulai, E.C., Olson, L.L., Deckard, E.L., Khan, M.F.R. and Finger, F.L. (2018). Methyl jasmonate alleviates drought stress in young sugar beet (Beta vulgaris L.) plants. Journal of Agronomy and Crop Science. 204(6): 566-576. doi: 10.1111/ jac.12286.

    14. García-Rodríguez, J.C., Manzo-Valencia, M.K., Olalde-Portugal, V. and Valdés-Rodríguez, S.E. (2024). Exploring drought responses in mexican soybeans: plant water status, shoot and root biomass and root system architecture. Food and Energy Security. 13(6). doi: 10.1002/fes3.70017.

    15. Gholamhoseini, M., Ebrahimian, E., Habibzadeh, F., Ataei, R. and Dezfulizadeh, M.S. (2018). Interactions of shading conditions and irrigation regimes on photosynthetic traits and seed yield of soybean (Glycine max L.). Legume Research. 41(2): 230-238. doi: 10.18805/lr-359.

    16. Haghaninia, M., Mashhouri, S.M., Najafifar, A., Soleimani, F. and Wu, Q.S. (2025). Combined effects of zinc oxide nanoparticles and arbuscular mycorrhizal fungi on soybean yield, oil quality and biochemical responses under drought stress. Future Foods. 11. doi: 10.1016/ j.fufo.2025.100594.

    17. Jiang, X.Y., Li, H.S., Dai, X., Li, J.D. and Liu, Y. (2025). An empirical analysis of global soybean supply potential and china’s diversified import strategies based on global agro- ecological zones and multi-objective nonlinear programming models. Agriculture-Basel. 15(5). doi: 10.3390/agriculture 15050529.

    18. Kim, J., Lee, C.W., Park, J.E., Mansoor, S., Chung, Y.S. and Kim, K. (2023). Drought stress restoration frequencies of phenotypic indicators in early vegetative stages of soybean (Glycine max L.). Sustainability. 15(6). doi: 10.3390/su15064852.

    19. Kisman, Hemon, A.F., Listiana, B.E., Dewi, S.M. and Mustikarini, E.D. (2025). Evaluation of drought tolerance and adaptation of large-seeded soybean genotypes under various drought stress levels. Indian Journal of Agricultural Research. 59(8): 1189-1200. doi: 10.18805/IJARe.AF-929.

    20. Kouighat, M., Moussaoui, F.E., Adiba, A., Hafid, A., Bouchyoua, A., El Fechtali, M. and Nabloussi, A. (2025). Evaluation and selection of sesame mutants (Sesamum indicum L.) for optimal nutritional profiles in seeds under field drought conditions. Ocl-Oilseeds and Fats Crops and Lipids. 32. doi: 10.1051/ocl/2025004.

    21. Liu, D., Guo, H.L., Yan, L.P., Gao, L., Zhai, S.S. and Xu, Y. (2024). Physiological, photosynthetic and stomatal ultrastructural responses of quercus acutissima seedlings to drought stress and rewatering. Forests. 15(1). doi: 10.3390/ f15010071.

    22. Liu, Z.C., Bao, D.E., Hu, H.L. and Miao, Y.C. (2019). Changes of photosynthetic characteristics in persimmon (Diospyros Spp.) rootstocks during drought stress simulated by PEG-6000. Bangladesh Journal of Botany. 48(3): 913- 918.

    23. Mangena, P. (2020). Effect of hormonal seed priming on germination, growth, yield and biomass allocation in soybean grown under induced drought stress. Indian Journal of Agricultural Research. 54(5): 592-598. doi: 10.18805/IJARe.A-441.

    24. Ndlovu, E., Maphosa, M. and van Staden, J. (2025). Morphological responses of sorghum seedlings to drought, heat and combined stresses. Experimental Agriculture. 61. doi: 10.1017/s0014479725100082.

    25. Quat, N.N., Dang, B.Q., Trang, N.T., Nhung, N.T. and Cuc, L.M. (2025). Evaluating drought tolerance in common beans using drought indices and molecular markers. Indian Journal of Agricultural Research. 59(8): 1177-1183. doi: 10.18805/IJARe.AF-940.

    26. Song, Y.L., Tian, J.F., Linderholm, H.W., Wang, C.Y., Ou, Z.R. and Chen, D.L. (2021). The contributions of climate change and production area expansion to drought risk for maize in China over the last four decades. International Journal of Climatology. 41: E2851-E2862. doi: 10.1002/joc.6885.

    27. Wei, Y.Q., Jin, J.L., Jiang, S.M., Ning, S.W. and Liu, L. (2018). Quantitative response of soybean development and yield to drought stress during different growth stages in the huaibei plain, China. Agronomy-Basel. 8(7). doi: 10.3390/ agronomy8070097.

    28. Wijewardana, C., Alsajri, F.A., Irby, J.T., Krutz, L.J., Golden, B., Henry, W.B., Gao, W. and Reddy, K.R. (2019). Physiological assessment of water deficit in soybean using midday leaf water potential and spectral features. Journal of Plant Interactions. 14(1): 533-543. doi: 10.1080/17429145. 2019.1662499.

    29. Zhou, Q., Li, Y.P., Wang, X.J., Yan, C., Ma, C.M., Liu, J. and Dong, S.K. (2022). Effects of different drought degrees on physiological characteristics and endogenous hormones of soybean. Plants-Basel. 11(17). doi: 10.3390/plants 11172282.
    In this Article
      Contact us
      Follow us
      Published In
      Legume Research

      Editorial Board

      View all (0)