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The plants of all the six genotypes showed smaller size and less leaf areas under waterlogging stress as compared with the controls (Supplemental Fig. S1). The plant height of all the six tomato genotypes significantly decreased ranging from 40.4% (No. 10, 'Jinzhu') to 54.8% (No. 16, 'Ruifen 882') under waterlogging stress than the respective controls (Fig. 1a). Similarly, waterlogging stress induced significant reduction in the stem diameter of all the six genotypes ranging from 21.5% (No. 9, 'Hezuo 908') to 61.4% (No. 16, 'Ruifen 882') (Fig. 1b). The number of leaves in plants except for genotype No. 3 was significantly lower under waterlogging stress than the respective controls (Fig. 1c). More importantly, the number of inflorescences of all the six tomato genotypes significantly decreased 64.2%, 57.1%, 41.8%, 60.1%, 46.6% and 60.1%, respectively, under waterlogging stress as compared with the respective controls (Fig. 1d).
Figure 1.
(a) Plant height, (b) stem diameter, (c) number of leaves, and (d) number of inflorescences of six tomato genotypes at anthesis stage under CK and WL. The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). The percentages (blue square) referred to significantly decreased percentages of the parameters in each genotype under waterlogging treatment as compared with the respective controls (p < 0.05). The percentages without color indicated no significant difference (p < 0.05).
Chlorophyll content
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Leaf chlorophyll a content of genotype No. 1, No. 3 and No. 10 significantly decreased 66.7%, 35.2% and 62.5%, respectively, and leaf chlorophyll b content of genotype No. 1 and No. 16 significantly decreased 31.9% and 20.0%, under waterlogging stress as compared with the respective controls (Fig. 2a, b). Waterlogging stress significantly decreased leaf total chlorophyll content of No. 1 (51.1%), No. 3 (21.0%) and No. 16 (44.5%) (Fig. 2c).
Figure 2.
(a) Chlorophyll a content, (b) chlorophyll b content, (c) total chlorophyll content of six tomato genotypes at anthesis stage under CK and WL. The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). The percentages (blue square) referred to significantly decreased percentages of the parameters in each genotype under waterlogging treatment as compared with the respective controls (p < 0.05). The percentages without color indicated no significant difference (p < 0.05).
Leaf gas exchange
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The PN of genotype No. 1 significantly decreased 17.5% under waterlogging stress as compared with the respective controls (Fig. 3a). Waterlogging stress induced significant drops in the Ci of No. 7 (42.6%), No. 9 (24.5%) and No. 16 (18.9%) (Fig. 3b). The Gs of No. 1 and No. 3 significantly decreased 40.6% and 20.7% respectively, and the Tr of all the six genotypes except for No. 7 significantly decreased, under waterlogging stress as compared with the respective controls (Fig. 3c, d).
Figure 3.
(a) Net photosynthetic rate (PN), (b) intercellular CO2 concentration (Ci), (c) stomatal conductance (Gs), and (d) transpiration rate (Tr) of six tomato genotypes at anthesis stage under CK and WL. The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). The percentages (blue square) referred to significantly decreased percentages of the parameters in each genotype under waterlogging treatment as compared with the respective controls (p < 0.05). The percentages without color indicated no significant difference (p < 0.05).
Activities of antioxidant enzymes
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The SOD activity of No.1, No. 3 and No. 16 significantly decreased 10.6%, 6.5% and 25.9%, respectively under waterlogging stress than the respective controls (Fig. 4a). As compared with the respective controls, the CAT activity of No. 9, and No. 16 significantly increased 74.9% and 159.9%, but that of No. 10 significantly decreased 67.3% under waterlogging stress (Fig. 4b). Waterlogging stress induced significant increases in the POD activity of No. 1 and No. 9, but significant drops in the POD activity of No. 3, No. 7 and No. 10 (Fig. 4c). By comparison, the APX activity of No. 1 and No. 7 significantly increased, but that of No. 10 and No. 16 significantly decreased under waterlogging stress as compared with the respective controls (Fig. 4d).
Figure 4.
Activities of (a) superoxide dismutase (SOD), (b) catalase (CAT), (c) peroxidase (POD), and (d) ascorbate peroxidase (APX) of six tomato genotypes at anthesis stage under CK and WL The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). The percentages (blue/orange square) referred to significantly decreased/increased percentages of the parameters in each genotype under waterlogging treatment as compared with the respective controls (p < 0.05). The percentages without color indicated no significant difference (p < 0.05).
Levels of leaf membrane lipid peroxidation under waterlogging stress
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Waterlogging stress induced significant increases in the MDA content and H2O2 content of all the six genotypes (Fig. 5a, b). The proline content of No. 1 and No. 9 significantly decreased, while that of the rest four genotypes significantly increased under waterlogging stress as compared with the respective controls (Fig. 5c). Waterlogging stress significantly increased the O2·− production rate of No. 1, No. 3 and No. 16, but decreased that of No. 10 (Fig. 5d).
Figure 5.
(a) Malondialdehyde (MDA) content, (b) hydrogen peroxide (H2O2) content, (c) proline (Pro) content, and (d) superoxide anion (O2•─) production rate of six tomato genotypes at anthesis stage under CK and WL. The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). The percentages (blue/orange square) referred to significantly decreased/increased percentages of the parameters in each genotype under waterlogging treatment as compared with the respective controls (p < 0.05). The percentages without color indicated no significant difference (p < 0.05).
Changes in tomato yield under waterlogging stress
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Moreover, the flower number per plant of all the genotypes except for No. 3 significantly decreased under waterlogging stress than the respective controls (Table 1). Waterlogging stress significantly decreased single fruit weight and fruit weight per plant of genotype No. 1, 7, 10 and 16, where the genotype No. 3, 9 failed to harvest tomato fruits (Table 1). The fruit number per plant of genotype No. 1 and No. 16 significantly decreased under waterlogging stress as compared with the respective controls (Table 1).
Table 1. Effects of waterlogging on yield of six tomato genotypes at the anthesis stage.
NO. Tomato genotype Treatment Flower number
per plantSingle fruit
weight (g)Fruit number per plant Fruit weight per plant (Kg) 1 MIX-002 CK 48 ± 5.0* 20.7 ± 1.22* 20 ± 1.0* 0.41 ± 0.02* WL 5 ± 1.5 9.0 ± 0.91 5 ± 3.0 0.04 ± 0.03 2 LA4440 CK 15 ± 2.8 69.7 ± 4.35 7 ± 0.3 0.46 ± 0.02 WL 4 ± 0.3 ─ ─ ─ 3 Fenbeibei CK 59 ± 10.4* 20.5 ± 0.28* 198 ± 4.0 4.06 ± 0.08* WL 21 ± 5.2 6.6 ± 0.06 70 ± 48.3 0.46 ± 0.32 4 Hezuo 908 CK 18 ± 2.3* 171.9 ± 14.52 16 ± 4.7 2.75 ± 0.81 WL 6 ± 1.2 ─ ─ ─ 5 Jinzhu CK 90 ± 14.4* 8.7 ± 0.36* 54 ± 7.2 0.47 ± 0.06* WL 24 ± 6.5 3.6 ± 0.40 26 ± 7.6 0.09 ± 0.03 6 Ruifen 882 CK 16 ± 2.2* 142.3 ± 13.71* 50 ± 2.9* 7.16 ± 0.41* WL 5 ± 2.9 16.0 ± 3.16 5 ± 0.6 0.05 ± 0.03 Note: The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). '*' indicated the significant difference of the same genotype between CK and WL (p < 0.05). Fruit characteristic of six tomato genotypes
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The transverse and longitudinal diameter of fruits harvested from genotype No. 1, No. 7, No. 10 and No. 16 significantly decreased under waterlogging stress as compared with the respective controls (Fig. 6a, b). The fruit hardness of genotype No. 16 was lower, while the fruit soluble solid content of genotype No. 10 was higher under waterlogging stress than control (Fig. 6c, d). The content of soluble protein, soluble sugar, and VC of genotype No. 7, 10 and 16 was significantly higher under waterlogging stress than control, except the content of soluble sugar of genotype No. 16 (Fig. 6e−g).
Figure 6.
(a) Transverse diameter, (b) longitudinal diameter, (c) hardness, (d) soluble solid content, (e) soluble protein content, (f) soluble sugar content, and (g) vitamin C (VC) content of the fruits from six tomato genotypes at anthesis stage under CK and WL. The CK and WL indicated control and waterlogging treatment, respectively. The values indicated mean ± SEM (n = 3). '*' indicated the significant difference of the parameters in each genotype under waterlogging treatment as compared with the respective controls; no marker indicated no significant difference (p < 0.05). Plants of genotype No. 2 and No. 4 under waterlogging treatment exhibited less than three replicates due to less than three fruits, resulting in blank bars.
Correlation between the physiological traits during tomato flowering period under waterlogging stress
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We found that the leaf MDA and H2O2 content showed extremely significantly negative correlation with yield per plant (correlation coefficient = −0.53** and −0.55**, respectively) (Fig. 7). Meanwhile, the yield per plant was extremely significantly positive correlated with transverse (0.64**) and longitudinal diameter (0.60**) (Fig. 7). The PN, POD and CAT activities, proline content, O2·− production rate and fruit soluble solid content and protein content showed weaker correlation with the rest indicators (Fig. 7), which was removed during the PCA. Six principal components (F1−F6) were extracted with 87.254% accumulative contribution percentage based on the PCA (Supplemental Table S1). The evaluation formula for overall traits of tomato plants was F = 0.313 × F1 + 0.284 × F2 + 0.166 × F3 + 0.109 × F4 + 0.069 × F5 + 0.060 × F6 (Supplemental Tables S1, S2). Based on the formula, the F value of each tomato genotype was calculated, where Jinzhu and Ruifen 882 showed the highest and lowest F value (0.814 and −0.384, respectively) (Supplemental Table S3).
Figure 7.
Correlation analysis of all the measured parameters. PN: Net photosynthetic rate, Ci: intercellular CO2 concentration, Gs: stomatal conductance, Tr: transpiration rate, SOD: superoxide dismutase, CAT: catalase, POD: peroxidase, APX: ascorbate peroxidase, MDA: Malondialdehyde, Pro: proline, VC: vitamin C.
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All data generated during this study are included within the article or its supplementary files.
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About this article
Cite this article
Yin J, Niu L, Li Y, Song X, Ottosen CO, et al. 2023. The effects of waterlogging stress on plant morphology, leaf physiology and fruit yield in six tomato genotypes at anthesis stage. Vegetable Research 3:31 doi: 10.48130/VR-2023-0031
The effects of waterlogging stress on plant morphology, leaf physiology and fruit yield in six tomato genotypes at anthesis stage
- Received: 30 July 2023
- Accepted: 21 September 2023
- Published online: 05 December 2023
Abstract: Waterlogging stress caused by concentrated and heavy rainfall has become an increasingly popular abiotic stress, especially for tomato production. The aim is to clarify the leaf physiological and biochemical responses as well as flower and fruit set of tomato plants at reproductive stage under waterlogging stress. Six tomato genotypes including 'MIX-002', 'LA4440', 'Fenbeibei', 'Hezuo 908', 'Jinzhu' and 'Ruifen 882' at anthesis stage were treated under control and waterlogging stress. We found that plant height, stem diameter, inflorescences number, single fruit weight and fruit weight per plant of all the six tomato genotypes significantly decreased under waterlogging stress as compared with the control. Chlorophyll a content of 'MIX-002', 'LA4440' and 'Jinzhu' and total chlorophyll content of 'MIX-002', 'LA4440' and 'Ruifen 882' were significantly lower under waterlogging conditions than the control. Waterlogging stress induced low net photosynthetic rate of 'MIX-002' and decreased transpiration rate of six genotypes except 'Fenbeibei'. Antioxidant enzymes activity and proline content responded to waterlogging stress by up/down regulation, resulting in high malondialdehyde and H2O2 content in leaves of six genotypes. Overall, waterlogging stress caused low gas diffusion, negatively affected pigment content, decreased transpiration rate, and induced antioxidant system disorder in tomato plants at anthesis stage. These adverse effects contributed to few buds and flowers and finally resulted in yield loss of tomato plants under waterlogging conditions. The significant correlation between malondialdehyde and H2O2 content and tomato yield indicated that the malondialdehyde and H2O2 content in leaves could be applied to predict the tomato yield under waterlogging condition.
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Key words:
- Tomato /
- Anthesis period /
- Waterlogging stress /
- Leaf physiology /
- Fruit yield and quality