Embryological，Physiological and Biochemical Mechanisms Underlying GA3-Induced Seedlessness in Grapes

doi:10.16420/j.issn.0513-353x.2025-0039

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (1): 65-81.doi: 10.16420/j.issn.0513-353x.2025-0039

• Genetic & Breeding·Germplasm Resources·Molecular Biology • Previous Articles Next Articles

Embryological，Physiological and Biochemical Mechanisms Underlying GA₃-Induced Seedlessness in Grapes

BAO Min¹, TIAN Yaping¹, LI Fan¹, WEI Xixi¹, LIU Miaomiao¹, HASSAN Jawad¹, LI Zhi¹^,², ZHAO Yijie¹^,², GAO Min¹^,², WANG Xiping¹^,²^,^*()

¹ State Key Laboratory of Crop Stress Resistance and High-Efficiency Production， Key Laboratory of Biology and Genetic Improvement of Horticultural Crops（Northwest Region），Ministry of Agriculture and Rural Affairs，College of Horticulture，Northwest A & F University，Yangling， Shaanxi 712100， China
² Xinjiang Research Institute of Agriculture in Arid Areas， Urumqi 830091， China

Received:2025-03-20 Revised:2025-11-19 Online:2026-01-25 Published:2026-01-26
Contact: WANG Xiping

Abstract

Abstract:

To clarify the underlying mechanisms of gibberellin（GA₃）-induced seedlessness in grape，Vitis labrusca‘Hutai 8’and V. vinifera‘Red Globe’were subjected to three treatments：GA₃，PAC（the GA₃ inhibitor），and GA₃ + PAC. By integrating fruit and seed morphological observations，ovule histological analyses，antioxidant enzyme activity measurements，and determinations of polyamine and endogenous hormone contents，the embryological，physiological，and biochemical mechanisms of GA₃- induced grape seedlessness were elucidated. GA₃ treatment significantly increased both the seedlessness rate and fruit set in the two cultivars. The fruit shape index was also markedly higher than that of the control and other treatments，whereas PAC reduced it below the control level. GA₃ induced embryo sac and endosperm nucleus abortion，with abortion occurring earlier in‘Hutai 8’（13 d）than in‘Red Globe’（15 d）. Moreover，GA₃ promoted the accumulation of hydrogen peroxide（H₂O₂）and malondialdehyde （MDA），while decreasing the activities of superoxide dismutase（SOD）and peroxidase（POD）in both cultivars，indicating that GA₃ induces embryo sac abortion by regulating oxidative stress levels. GA₃ treatment differentially affected putrescine（Put），spermidine（Spd），and spermine（Spm），resulting in a decreased（Spd + Spm）/Put ratio relative to the control；this reduction may contribute to the metabolic regulation of seedlessness formation. GA₃ treatment promoted the synthesis of endogenous GA₃，indole-3-acetic acid（IAA），and abscisic acid（ABA）in both cultivars，although IAA in‘Red Globe’showed an initial decline followed by an increase. At the early stage，the（IAA + GA₃）/ABA ratio was higher than that of the control，and as exogenous GA₃ declined through natural metabolism，this ratio decreased accordingly. Overall，this study reveals that GA₃ induces grape seedlessness by coordinately regulation of embryo abortion，oxidative stress，polyamines，and hormone balance.

Key words: grape, gibberellin, seedlessness, ovule, antioxidant activity, endogenous hormone

BAO Min, TIAN Yaping, LI Fan, WEI Xixi, LIU Miaomiao, HASSAN Jawad, LI Zhi, ZHAO Yijie, GAO Min, WANG Xiping. Embryological，Physiological and Biochemical Mechanisms Underlying GA₃-Induced Seedlessness in Grapes[J]. Acta Horticulturae Sinica, 2026, 53(1): 65-81.

Figures/Tables 10

References 38

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处理 Treatment	坐果率/% Fruit set rate		无核率/% Seedlessness rate
处理 Treatment	户太8号 Hutai 8	红地球 Red Globe	户太8号 Hutai 8	红地球 Red Globe
H₂O（对照Control）	52.7 ± 0.34 c	26.3 ± 0.21 c	4.2 ± 2.13 d	2.2 ± 1.18 c
GA₃	64.2 ± 1.05 a	34.5 ± 1.26 a	79.8 ± 1.25 a	61.3 ± 1.50 a
PAC	48.5 ± 1.84 d	25.1 ± 0.57 c	12.5 ± 4.01 c	4.2 ± 2.31 c
GA₃ + PAC	59.3 ± 0.95 b	30.4 ± 1.69 b	28.5 ± 1.51 b	23.5 ± 1.08 b

处理 Treatment	坐果率/% Fruit set rate		无核率/% Seedlessness rate
处理 Treatment	户太8号 Hutai 8	红地球 Red Globe	户太8号 Hutai 8	红地球 Red Globe
H₂O（对照Control）	52.7 ± 0.34 c	26.3 ± 0.21 c	4.2 ± 2.13 d	2.2 ± 1.18 c
GA₃	64.2 ± 1.05 a	34.5 ± 1.26 a	79.8 ± 1.25 a	61.3 ± 1.50 a
PAC	48.5 ± 1.84 d	25.1 ± 0.57 c	12.5 ± 4.01 c	4.2 ± 2.31 c
GA₃ + PAC	59.3 ± 0.95 b	30.4 ± 1.69 b	28.5 ± 1.51 b	23.5 ± 1.08 b

品种	处理	果形指数Fruit shape index
Cultivar	Treatment	6 d	12 d	18 d	24 d	30 d	36 d	42 d	48 d	54 d
户太8号 Hutai 8	H₂O	1.50 ± 0.03 c	1.35 ± 0.03 c	1.35 ± 0.03 c	1.31 ± 0.03 c	1.23 ± 0.03 c	1.28 ± 0.03 c	1.25 ± 0.03 c	1.23 ± 0.03 b	1.15 ± 0.03 b
	GA₃	1.77 ± 0.03 a	1.63 ± 0.03 a	1.72 ± 0.03 a	1.56 ± 0.03 a	1.50 ± 0.02 a	1.51 ± 0.02 a	1.45 ± 0.03 a	1.38 ± 0.03 a	1.28 ± 0.02 a
	PAC	1.31 ± 0.03 d	1.14 ± 0.02 d	1.33 ± 0.02 d	1.17 ± 0.02 d	1.22 ± 0.02 d	1.18 ± 0.02 d	1.09 ± 0.02 d	1.12 ± 0.02 c	1.06 ± 0.02 c
	GA₃ + PAC	1.66 ± 0.02 b	1.59 ± 0.02 b	1.48 ± 0.02 b	1.35 ± 0.02 b	1.33 ± 0.03 b	1.35 ± 0.02 b	1.27 ± 0.02 b	1.24 ± 0.02 b	1.16 ± 0.02 b
红地球 Red Globe	H₂O	1.80 ± 0.02 b	1.63 ± 0.02 b	1.55 ± 0.02 b	1.44 ± 0.02 b	1.50 ± 0.04 b	1.29 ± 0.03 b	1.38 ± 0.03 b	1.36 ± 0.03 b	1.16 ± 0.03 b
	GA₃	2.00 ± 0.03 a	1.81 ± 0.02 a	1.90 ± 0.02 a	1.51 ± 0.02 a	1.70 ± 0.02 a	1.53 ± 0.02 a	1.60 ± 0.02 a	1.52 ± 0.03 a	1.39 ± 0.03 a
	PAC	1.53 ± 0.02 c	1.24 ± 0.02 d	1.28 ± 0.02 d	1.29 ± 0.02 d	1.25 ± 0.02 d	1.25 ± 0.02 d	1.19 ± 0.02 d	1.19 ± 0.02 c	1.14 ± 0.02 c
	GA₃ + PAC	1.50 ± 0.01 c	1.42 ± 0.02 c	1.51 ± 0.02 c	1.36 ± 0.03 c	1.45 ± 0.02 c	1.28 ± 0.02 c	1.35 ± 0.02 c	1.36 ± 0.03 b	1.16 ± 0.03 b

品种	处理	果形指数Fruit shape index
Cultivar	Treatment	6 d	12 d	18 d	24 d	30 d	36 d	42 d	48 d	54 d
户太8号 Hutai 8	H₂O	1.50 ± 0.03 c	1.35 ± 0.03 c	1.35 ± 0.03 c	1.31 ± 0.03 c	1.23 ± 0.03 c	1.28 ± 0.03 c	1.25 ± 0.03 c	1.23 ± 0.03 b	1.15 ± 0.03 b
	GA₃	1.77 ± 0.03 a	1.63 ± 0.03 a	1.72 ± 0.03 a	1.56 ± 0.03 a	1.50 ± 0.02 a	1.51 ± 0.02 a	1.45 ± 0.03 a	1.38 ± 0.03 a	1.28 ± 0.02 a
	PAC	1.31 ± 0.03 d	1.14 ± 0.02 d	1.33 ± 0.02 d	1.17 ± 0.02 d	1.22 ± 0.02 d	1.18 ± 0.02 d	1.09 ± 0.02 d	1.12 ± 0.02 c	1.06 ± 0.02 c
	GA₃ + PAC	1.66 ± 0.02 b	1.59 ± 0.02 b	1.48 ± 0.02 b	1.35 ± 0.02 b	1.33 ± 0.03 b	1.35 ± 0.02 b	1.27 ± 0.02 b	1.24 ± 0.02 b	1.16 ± 0.02 b
红地球 Red Globe	H₂O	1.80 ± 0.02 b	1.63 ± 0.02 b	1.55 ± 0.02 b	1.44 ± 0.02 b	1.50 ± 0.04 b	1.29 ± 0.03 b	1.38 ± 0.03 b	1.36 ± 0.03 b	1.16 ± 0.03 b
	GA₃	2.00 ± 0.03 a	1.81 ± 0.02 a	1.90 ± 0.02 a	1.51 ± 0.02 a	1.70 ± 0.02 a	1.53 ± 0.02 a	1.60 ± 0.02 a	1.52 ± 0.03 a	1.39 ± 0.03 a
	PAC	1.53 ± 0.02 c	1.24 ± 0.02 d	1.28 ± 0.02 d	1.29 ± 0.02 d	1.25 ± 0.02 d	1.25 ± 0.02 d	1.19 ± 0.02 d	1.19 ± 0.02 c	1.14 ± 0.02 c
	GA₃ + PAC	1.50 ± 0.01 c	1.42 ± 0.02 c	1.51 ± 0.02 c	1.36 ± 0.03 c	1.45 ± 0.02 c	1.28 ± 0.02 c	1.35 ± 0.02 c	1.36 ± 0.03 b	1.16 ± 0.03 b

时间 Time	处理 Treatment	H₂O₂/ （nmol · g^-1 FW）	MDA/ （nmol · g^-1 FW）	POD/ （U · g^-1 FW）	SOD/ （U · g^-1 FW）
0.5 h	H₂O（对照Control）	226.75 ± 5.83 a	23.84 ± 0.43 a	9.22 ± 1.34 b	628.52 ± 5.26 c
	GA₃	182.25 ± 2.39 b	24.04 ± 0.56 a	6.53 ± 0.35 c	617.68 ± 17.30 c
	PAC	168.82 ± 7.22 c	24.79 ± 0.62 a	23.50 ± 0.34 a	744.93 ± 7.34 a
	GA₃ + PAC	228.34 ± 4.71 a	23.62 ± 0.21 a	7.12 ± 0.39 c	700.17 ± 14.94 b
1 h	H₂O（对照Control）	216.15 ± 9.39 a	25.40 ± 0.85 b	12.32 ± 0.46 b	734.61 ± 17.73 a
	GA₃	201.34 ± 5.66 b	25.35 ± 0.95 b	5.56 ± 1.38 d	728.57 ± 3.10 a
	PAC	185.21 ± 6.01 c	25.23 ± 0.60 b	15.99 ± 2.05 a	749.10 ± 13.20 a
	GA₃ + PAC	214.23 ± 5.30 a	27.32 ± 0.28 a	9.35 ± 1.10 c	729.81 ± 16.88 a
3 h	H₂O（对照Control）	254.06 ± 4.52 a	20.96 ± 0.33 c	12.62 ± 0.52 c	729.64 ± 8.51 a
	GA₃	216.15 ± 7.90 c	24.53 ± 1.13 b	5.88 ± 0.96 d	689.53 ± 2.53 b
	PAC	191.16 ± 2.92 d	28.13 ± 0.54 a	19.89 ± 0.32 a	726.69 ± 20.69 a
	GA₃ + PAC	237.07 ± 9.06 b	25.66 ± 1.61 b	14.97 ± 0.20 b	720.20 ± 6.64 a
12 h	H₂O（对照Control）	174.82 ± 8.59 b	17.67 ± 0.33 c	43.95 ± 1.61 a	687.19 ± 1.18 b
	GA₃	193.32 ± 6.18 a	18.60 ± 1.19 c	29.89 ± 0.06 c	646.47 ± 18.21 c
	PAC	165.63 ± 4.97 b	28.57 ± 0.41 a	38.26 ± 0.80 b	687.05 ± 5.19 b
	GA₃ + PAC	199.69 ± 6.31 a	24.59 ± 0.12 b	17.42 ± 1.34 d	714.06 ± 10.10 a
24 h	H₂O（对照Control）	195.58 ± 7.42 c	21.97 ± 0.81 c	23.37 ± 0.68 a	693.64 ± 5.26 b
	GA₃	215.22 ± 8.66 b	25.95 ± 0.61 b	20.94 ± 0.60 b	641.57 ± 5.19 d
	PAC	192.95 ± 6.55 c	27.82 ± 1.44 a	23.27 ± 1.43 a	662.32 ± 13.28 c
	GA₃ + PAC	233.46 ± 9.70 a	28.95 ± 0.51 a	11.59 ± 0.39 c	716.22 ± 6.26 a
48 h	H₂O（对照Control）	168.41 ± 5.76 b	30.58 ± 0.40 b	22.32 ± 0.25 a	723.72 ± 21.90 a
	GA₃	199.15 ± 9.88 a	36.03 ± 1.50 a	18.73 ± 1.34 b	699.14 ± 13.78 a
	PAC	181.00 ± 6.93 b	31.79 ± 0.64 b	22.27 ± 0.70 a	708.84 ± 13.79 a
	GA₃ + PAC	206.88 ± 4.06 a	32.72 ± 1.54 b	7.04 ± 0.71 c	663.89 ± 4.72 b
72 h	H₂O（对照Control）	156.29 ± 10.71 c	29.05 ± 0.84 b	47.32 ± 3.95 a	753.38 ± 24.68 a
	GA₃	172.97 ± 2.62 b	35.66 ± 0.34 a	22.31 ± 0.05 c	733.01 ± 25.01 a
	PAC	165.56 ± 6.08 bc	35.14 ± 1.43 a	27.50 ± 0.72 b	750.87 ± 1.65 a
	GA₃ + PAC	190.39 ± 5.56 a	24.23 ± 0.58 c	30.15 ± 1.96 b	678.52 ± 7.86 b
6 d	H₂O（对照Control）	135.26 ± 2.68 c	31.54 ± 0.36 c	13.12 ± 1.30 a	755.99 ± 13.74 a
	GA₃	159.58 ± 2.23 b	35.83 ± 0.64 a	10.33 ± 0.69 b	754.42 ± 14.35 a
	PAC	138.01 ± 5.28 c	33.81 ± 1.57 b	11.33 ± 1.67 ab	761.18 ± 15.14 a
	GA₃ + PAC	172.09 ± 3.37 a	32.70 ± 0.76 bc	12.90 ± 0.39 a	707.45 ± 3.50 b
9 d	H₂O（对照Control）	70.72 ± 3.12 c	27.62 ± 0.98 b	8.10 ± 0.52 a	713.35 ± 16.58 c
	GA₃	117.87 ± 5.13 b	29.10 ± 0.33 a	6.26 ± 1.52 b	795.29 ± 4.27 a
	PAC	113.98 ± 10.56 b	25.90 ± 0.85 c	5.80 ± 0.80 ab	798.08 ± 18.98 a
	GA₃ + PAC	137.05 ± 3.00 a	29.02 ± 0.61 a	10.95 ± 0.15 ab	742.92 ± 18.17 b
12 d	H₂O（对照Control）	123.21 ± 6.06 c	30.59 ± 0.93 b	2.15±0.21 a	635.36 ± 11.38 b
	GA₃	195.70 ± 4.33 a	37.40 ± 1.32 a	1.24±0.60 a	724.02 ± 5.32 a
	PAC	103.38 ± 5.81 d	32.74 ± 1.32 b	1.21±1.30 a	724.83 ± 21.98 a
	GA₃ + PAC	143.77 ± 8.74 b	30.64 ± 1.08 b	1.06±1.63 a	647.53 ± 17.60 b
18 d	H₂O（对照Control）	72.37 ± 4.21 a	27.31 ± 0.69 a	1.61 ± 0.56 a	551.75 ± 5.27 a
	GA₃	74.41 ± 4.79 a	14.77 ± 0.98 c	1.34 ± 1.80 a	358.40 ± 12.72 d
	PAC	39.52 ± 9.07 b	19.66 ± 1.18 b	1.31 ± 0.40 a	482.41 ± 12.47 b
	GA₃ + PAC	60.77 ± 2.78 c	18.34 ± 1.04 b	1.26 ± 1.16 a	420.49 ± 6.47 c
24 d	H₂O（对照Control）	38.88 ± 1.39 c	13.75 ± 0.17 c	1.30 ± 1.13 a	264.45 ± 7.57 ab
	GA₃	76.53 ± 2.54 a	16.83 ± 0.15 b	1.47 ± 1.37 a	220.18 ± 19.07 ab
	PAC	55.68 ± 2.42 b	17.87 ± 0.42 a	1.45 ± 0.36 a	332.47 ± 4.99 a
	GA₃ + PAC	43.11 ± 3.27 c	17.70 ± 0.12 a	1.32 ± 0.57 a	288.79 ± 10.88 b

Embryological，Physiological and Biochemical Mechanisms Underlying GA₃-Induced Seedlessness in Grapes

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时间 Time	处理 Treatment	H₂O₂/ （nmol · g^-1 FW）	MDA/ （nmol · g^-1 FW）	POD/ （U · g^-1 FW）	SOD/ （U · g^-1 FW）
0.5 h	H₂O（对照Control）	313.02 ± 2.68 d	36.50 ± 0.41 b	68.25 ± 1.09 a	616.19 ± 7.53 c
	GA₃	339.32 ± 10.98 c	31.91 ± 1.74 c	29.47 ± 0.49 c	660.50 ± 12.63 b
	PAC	409.72 ± 8.90 a	43.15 ± 0.61 a	52.33 ± 0.40 b	675.10 ± 12.22 ab
	GA₃ + PAC	372.62 ± 7.47 b	46.02 ± 3.08 a	54.12 ± 2.34 b	682.78 ± 12.22 a
1 h	H₂O（对照Control）	338.23 ± 10.76 c	45.14 ± 0.86 c	83.33 ± 3.91 a	675.91 ± 8.55 b
	GA₃	368.50 ± 7.50 a	51.63 ± 2.31 b	31.90 ± 0.67 c	676.23 ± 3.47 b
	PAC	353.28 ± 6.43 b	48.85 ± 0.99 b	50.91 ± 1.11 b	687.64 ± 2.26 ab
	GA₃ + PAC	328.48 ± 3.03 c	59.38 ± 1.89 a	51.00 ± 0.86 b	691.71 ± 9.39 a
3 h	H₂O（对照Control）	404.28 ± 2.98 b	49.11 ± 1.09 b	83.03 ± 1.00 a	686.57 ± 20.78 bc
	GA₃	461.41 ± 10.84 a	49.21 ± 1.33 b	32.30 ± 1.96 d	665.05 ± 9.69 c
	PAC	415.62 ± 8.42 b	50.23 ± 1.92 ab	71.09 ± 0.78 b	728.48 ± 1.87 a
	GA₃ + PAC	378.79 ± 8.60 c	52.42 ± 0.21 a	54.20 ± 1.19 c	712.38 ± 17.42 ab
12 h	H₂O（对照Control）	352.47 ± 11.77 b	50.83 ± 0.73 b	74.38 ± 2.06 a	711.91 ± 14.55 ab
	GA₃	354.85 ± 8.59 b	56.41 ± 0.84 a	45.74 ± 0.16 d	687.45 ± 7.87 b
	PAC	409.86 ± 13.90 a	54.68 ± 1.59 a	58.99 ± 2.42 b	723.26 ± 23.19 a
	GA₃ + PAC	353.80 ± 7.11 b	56.89 ± 1.08 a	52.55 ± 1.68 c	726.49 ± 21.67 a
24 h	H₂O（对照Control）	393.84 ± 7.02 ab	56.62 ± 0.80 b	83.80 ± 2.12 a	675.90 ± 3.77 b
	GA₃	383.56 ± 12.28 b	58.85 ± 1.38 ab	57.96 ± 1.25 d	669.41 ± 4.46 b
	PAC	401.47 ± 11.19 ab	58.50 ± 1.63 ab	79.26 ± 0.82 b	689.06 ± 2.46 a
	GA₃ + PAC	410.50 ± 3.63 a	59.32 ± 0.57 a	63.36 ± 0.76 c	685.35 ± 7.66 a
48 h	H₂O（对照Control）	347.63 ± 5.79 b	59.38 ± 0.67 b	131.40 ± 5.03 a	670.42 ± 18.60 b
	GA₃	349.06 ± 8.01 b	60.93 ± 0.75 b	56.03 ± 1.15 c	648.88 ± 23.38 b
	PAC	364.53 ± 7.40 ab	63.83 ± 2.19 a	66.74 ± 0.78 b	723.70 ± 21.54 a
	GA₃ + PAC	372.38 ± 14.19 a	65.43 ± 1.37 a	60.54 ± 2.36 c	687.07 ± 13.30 ab
72 h	H₂O（对照Control）	347.00 ± 4.97 b	57.76 ± 1.52 c	88.58 ± 0.58 b	688.49 ± 13.76 b
	GA₃	394.44 ± 11.51 a	61.60 ± 1.21 b	101.40 ± 0.21 a	686.48 ± 4.09 b
	PAC	352.87 ± 3.94 b	67.78 ± 0.99 a	101.36 ± 1.17 a	710.73 ± 13.68 b
	GA₃ + PAC	336.75 ± 7.54 b	57.98 ± 0.73 c	102.42 ± 0.87 a	737.26 ± 14.77 a
6 d	H₂O（对照Control）	237.73 ± 3.95 d	48.44 ± 0.28 c	29.76 ± 1.67 c	616.29 ± 17.26 bc
	GA₃	354.22 ± 11.69 a	51.12 ± 0.90 bc	55.93 ± 3.76 a	596.69 ± 23.54 c
	PAC	325.25 ± 6.41 b	65.80 ± 1.82 a	39.29 ± 0.76 b	656.47 ± 17.48 a
	GA₃ + PAC	270.36 ± 9.43 c	53.49 ± 2.61 b	42.55 ± 0.40 b	631.78 ± 4.85 ab
9 d	H₂O（对照Control）	220.54 ± 1.46 c	44.59 ± 1.62 c	27.02 ± 0.90 b	495.65 ± 15.69 c
	GA₃	311.73 ± 8.94 a	66.89 ± 3.48 a	35.14 ± 0.63 a	363.88 ± 5.80 d
	PAC	242.86 ± 4.26 b	52.95 ± 3.16 b	29.12 ± 0.19 b	663.84 ± 20.10 a
	GA₃ + PAC	205.66 ± 8.11 d	56.69 ± 1.40 b	34.88 ± 2.50 a	546.31 ± 14.46 b
12 d	H₂O（对照Control）	170.84 ± 4.89 c	28.23 ± 0.71 a	16.84 ± 5.22 b	414.14 ± 20.40 b
	GA₃	184.52 ± 7.88 ab	30.72 ± 0.06 a	22.94 ± 1.00 a	342.97 ± 14.46 c
	PAC	196.36 ± 7.09 a	18.50 ± 3.30 b	25.09 ± 0.67 a	475.88 ± 6.95 a
	GA₃ + PAC	180.39 ± 4.60 bc	29.77 ± 0.31 a	22.11 ± 0.15 a	430.56 ± 7.75 b
18 d	H₂O（对照Control）	81.07 ± 1.23 c	13.63 ± 2.71 a	22.03 ± 2.32 c	272.19 ± 6.21 c
	GA₃	80.56 ± 2.07 c	12.36 ± 1.37 a	32.10 ± 2.50 a	171.76 ± 5.46 d
	PAC	110.94 ± 0.49 a	11.78 ± 0.13 a	25.98 ± 0.55 b	377.40 ± 34.33 a
	GA₃ + PAC	101.64 ± 2.42 b	13.50 ± 2.16 a	28.11 ± 0.42 b	319.42 ± 1.11 b
24 d	H₂O（对照Control）	96.50 ± 1.42 a	12.89 ± 1.34 a	9.04 ± 1.30 d	294.72 ± 8.86 b
	GA₃	62.15 ± 4.65 b	10.62 ± 1.71 a	38.99 ± 1.12 a	241.63 ± 28.00 c
	PAC	65.93 ± 2.28 b	10.58 ± 1.40 a	20.31 ± 0.97 b	352.07 ± 16.62 a
	GA₃ + PAC	61.15 ± 3.82 b	9.78 ± 1.52 a	15.14 ± 0.88 c	349.27 ± 16.01 a

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