柠檬酸浸种引发对低磷胁迫下番茄幼苗生长及生理特性的影响

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靳琇,
陈浩婷,
石玉,
白龙强,
侯雷平,
张毅^,
山西农业大学园艺学院/山西省设施蔬菜提质增效协同创新中心太谷 030801
基金项目: 山西省重点研发计划重点项目201903D211011
山西省重点研发计划重点项目201703D211001

详细信息

作者简介:靳琇, 研究方向为设施蔬菜栽培生理。E-mail: xiaojinyuboer@163.com

通讯作者:张毅, 研究方向为设施园艺。E-mail: harmony1228@163.com

中图分类号:S641.2

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出版历程

收稿日期:2020-11-29
录用日期:2021-03-13
刊出日期:2021-07-01

Effect of citric acid seed priming on the growth and physiological characteristics of tomato seedlings under low phosphorus stress

JIN Xiu,
CHEN Haoting,
SHI Yu,
BAI Longqiang,
HOU Leiping,
ZHANG Yi^,
College of Horticulture, Shanxi Agricultural University/Collaborative Innovation Center for Improving Quality and Efficiency of Facilities Vegetables, Taigu 030801, China
Funds: the Shanxi Province Key R&D Plan201903D211011
the Shanxi Province Key R&D Plan201703D211001

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Corresponding author:ZHANG Yi, E-mail: harmony1228@163.com

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摘要
摘要:本试验以‘中杂9号’番茄为试材，采用砂培法，设置对照（0.66 mmol·L^-1）、轻度缺磷（0.44 mmol·L^-1）、重度缺磷（0.22 mmol·L^-1）和无磷（0 mmol·L^-1）4个磷浓度，研究了柠檬酸（7.5 mmol·L^-1）浸种对低磷胁迫下番茄种子萌发、幼苗生长及生理特性的影响，旨在阐明柠檬酸浸种缓解番茄低磷胁迫的生理机制。结果表明：与蒸馏水浸种相比，不同浓度柠檬酸浸种引发显著提高了番茄发芽率、发芽势、发芽指数，最佳浓度为7.5 mmol·L^-1。与对照相比，低磷胁迫下番茄幼苗生长受到严重抑制，丙二醛（MDA）、活性氧（$ $\text{O}_2^{\bar \cdot }$$、H₂O₂）含量显著升高，超氧化物歧化酶（SOD）、过氧化物酶（POD）和过氧化氢酶（CAT）活性显著减低；可溶性蛋白、可溶性糖及甜菜碱含量显著升高，渗透势降低；而柠檬酸浸种引发明显促进了番茄幼苗的生长，增加了生物量的积累；提高了番茄中抗氧化酶活性，降低了MDA和活性氧的积累，减轻了番茄幼苗的氧化胁迫；显著提高了有机酸、可溶性糖等的含量，缓解了渗透势的下降；显著提高了番茄幼苗中的磷含量，缓解了植株的低磷胁迫症状。综上所述，柠檬酸浸种引发通过缓解低磷胁迫对番茄造成的氧化损伤，提高磷素吸收，维持正常生长代谢，从而增强番茄幼苗对低磷胁迫的抵抗能力。
关键词:番茄/
低磷胁迫/
种子引发/
柠檬酸/
抗氧化系统/
渗透调节
Abstract:To explore the effects of seed priming with citric acid on tomato seed germination, seedling growth and physiological characteristics under low phosphorus stress, 'Zhongza 9' was cultured in sand at four phosphorus concentrations: control (0.66 mmol·L^-1), mild phosphorus deficiency (0.44 mmol·L^-1), severe phosphorus deficiency (0.22 mmol·L^-1), and non-phosphorus (0 mmol·L^-1). The tomato seeds were soaked in citric acid (7.5 mmol·L^-1), and the seed germination, seedlings growth and physiological characteristics were examined. This experiment aimed to clarify the physiological mechanisms of citric acid seed priming in alleviating low phosphorus stress in tomatoes. The results showed that, compared with seed soaking in distilled water, the different concentrations of citric acid seed soaking significantly increased the germination rate, germination potential, and germination index; the optimal concentration was 7.5 mmol·L^-1. Compared with the control, tomato seedling growth under low phosphorus stress was severely inhibited; the malondialdehyde (MDA) and reactive oxygen species ($ $\text{O}_2^{\bar \cdot }$$, H₂O₂) contents significantly increased; and the superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) activities significantly decreased. The soluble protein, soluble sugar, and betaine contents significantly increased, and the osmotic potential was low. The seed priming with citric acid significantly promoted tomato seedling growth, increased biomass accumulation, increased the activities of antioxidant enzymes, reduced the accumulation of MDA and reactive oxygen species, and reduced the oxidative stress in tomato seedlings. It also significantly improved the organic acid and soluble sugar contents, alleviated osmotic potential decline, increased the phosphorus content, and lessened the symptoms of low phosphorus stress. In summary, priming tomato seeds with citric acid alleviated the oxidative damage caused by low phosphorus, improved the absorption efficiency of phosphorus, maintained normal growth and metabolism, and enhanced the resistance of tomato seedlings to low phosphorus stress.
Key words:Tomato/
Low phosphorus stress/
Seed priming/
Citric acid/
Antioxidant system/
Osmotic regulation

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图1不同浓度柠檬酸浸种引发对番茄种子萌发指标的影响
CK: 蒸馏水浸种。不同小写字母表示不同处理间差异显著(P < 0.05)。
Figure1.Effects of seed priming with different concentrations of citric acid on seeds germination indexes of tomato
CK: soaking seeds in distilled water. Different lowercase letters indicate significant differences among treatments (P < 0.05).

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图2柠檬酸浸种引发对低磷胁迫下番茄幼苗根系活力的影响
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。不同小写字母表示处理间差异显著(P < 0.05)。
Figure2.Effect of seeds priming with citric acid on root activity of tomato seedlings under low phosphorus stress
CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters indicate significantly difference (P < 0.05).

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图3柠檬酸浸种引发对低磷胁迫下番茄幼苗丙二醛(MDA)、超氧化阴离子($\text{O}_2^{\bar \cdot }$)和过氧化氢(H₂O₂)含量的影响
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。不同小写字母表示处理间差异显著(P < 0.05)。
Figure3.Effect of seeds priming with citric acid on malondialdehyde (MDA), superoxide ion (${\rm{O}}_{\rm{2}}^{\bar \cdot }$), and H₂O₂ contents of tomato seedlings under low phosphorus stress
CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters indicate significantly difference (P < 0.05).

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图4柠檬酸浸种引发对低磷胁迫下番茄幼苗渗透势及渗透调节物质含量的影响
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。不同小写字母表示处理间差异显著(P < 0.05)。
Figure4.Effect of seeds priming with citric acid on osmotic potential and osmoregulation substances contents of tomato seedlings under low phosphorus stress
CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters indicate significantly difference (P < 0.05).

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图5柠檬酸浸种引发对低磷胁迫下番茄幼苗磷含量的影响
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。不同小写字母表示处理间差异显著(P < 0.05)。
Figure5.Effect of seeds priming with citric acid on phosphorus content of tomato seedlings under low phosphorus stress
CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters indicate significantly difference (P < 0.05).

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表1柠檬酸浸种引发对低磷胁迫下番茄幼苗形态指标的影响
Table1.Effects of seeds priming with citric acid on morphological indexes of tomato seedlings under low phosphorus stress

处理 Treatment	单株鲜质量 Fresh weight per plant (g)		单株干质量 Dry weight per plant (g)		株高 Plant height (cm)	茎粗 Stem diameter (mm)	总根长 Total root length (cm)	总根系表面积 Total root surface area (cm²)
处理 Treatment	地上部Shoots	地下部Roots	地上部Shoots	地下部Roots	株高 Plant height (cm)	茎粗 Stem diameter (mm)	总根长 Total root length (cm)	总根系表面积 Total root surface area (cm²)
CK	3.38±0.28b	0.38±0.03ab	0.22±0.02b	0.02±0.01bc	22.56±0.23b	3.59±0.21b	243.06±20.50b	73.70±10.13b
CK+A	4.58±0.29a	0.42±0.04ab	0.25±0.02a	0.02±0.01f	26.61±1.25a	4.00±0.08a	319.89±14.36a	120.72±9.32a
P0	1.57±0.02e	0.22±0.02ab	0.13±0.01de	0.02±0.01a	13.84±0.48e	2.60±0.08g	138.06±4.87e	17.98±2.59ef
P0+A	1.73±0.03e	0.34±0.04b	0.15±0.01d	0.01±0.01ef	14.34±0.57de	2.71±0.06ef	159.78±9.31d	24.57±0.28de
P1	2.61±0.18d	0.22±0.03ab	0.11±0.01e	0.02±0.01a	15.02±0.34d	2.73±0.01ef	197.96±6.51c	28.46±1.03de
P1+A	3.09±0.05c	0.28±0.00a	0.18±0.02c	0.03±0.01b	20.51±0.91c	3.07±0.09c	221.88±18.92b	38.55±6.00c
P2	2.91±0.08c	0.24±0.02b	0.12±0.02e	0.01±0.01de	19.60±0.65c	2.84±0.08d	65.61±3.42f	8.28±1.28f
P2+A	3.49±0.08b	0.34±0.03ab	0.2±0.01b	0.02±0.01cd	22.68±0.48b	3.45±0.18b	120.34±7.36e	16.49±2.32ef
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。同列数据后标不同小写字母表示各处理间差异显著(P < 0.05)。CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters in each column indicate significantly difference (P < 0.05).

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表2柠檬酸浸种引发对低磷胁迫下番茄幼苗叶绿素的影响
Table2.Effect of seeds priming with citric acid on chlorophyll contents in tomato seedlings under low phosphorus stress

处理 Treatment	叶绿素a Chl a (mg·g^–1)	叶绿素b Chl b (mg·g^–1)	类胡萝卜素 Carotenoid (mg·g^–1)	总叶绿素 Chl a+b (mg·g^–1)	叶绿素a/b Chl a/b
CK	16.27±0.08c	4.58±0.22b	1.43±0.17ab	20.85±0.21c	3.56±0.18bcd
CK+A	18.17±0.15a	5.19±0.11a	1.53±0.06ab	23.37±0.26a	3.50±0.05cd
P0	7.75±0.05h	1.75±0.06f	0.33±0.04d	9.50±0.03h	4.42±0.19a
P0+A	9.01±0.04g	1.96±0.01e	0.97±0.02c	10.97±0.03g	4.60±0.03a
P1	12.98±0.03f	3.61±0.05d	1.13±0.02c	16.58±0.03f	3.60±0.05bc
P1+A	14.38±0.1e	4.09±0.07c	1.46±0.18ab	18.47±0.14e	3.52±0.06bcd
P2	15.52±0.04d	4.62±0.2b	1.39±0.04b	20.14±0.2d	3.36±0.15d
P2+A	17.31±0.05b	4.66±0.04b	1.58±0.02ab	21.96±0.04b	3.72±0.04b
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。同列数据后不同小写字母表示各处理间差异显著(P < 0.05)。CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters in each column indicate significantly difference (P < 0.05).

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表3柠檬酸浸种引发对低磷胁迫下番茄幼苗抗氧化酶活性的影响
Table3.Effect of seed priming with citric acid on antioxidant enzymes activities of tomato seedlings under low phosphorus stress?U·g^–1·min^–1

处理 Treatment	叶片Leaf			根系Root
处理 Treatment	超氧化物歧化酶 SOD	过氧化氢酶 CAT	过氧化物酶 POD	超氧化物歧化酶 SOD	过氧化氢酶 CAT	过氧化物酶 POD
CK	10.80±0.32e	35.87±1.47f	367.70±5.23f	15.49±0.92e	13.91±0.86f	1929.98±38.16h
CK+A	21.09±10.29b	78.32±3.82a	984.32±27.03a	25.55±0.91a	55.73±0.77a	5674.12±34.27a
P0	23.24±30.29a	67.05±0.89b	562.67±2.34c	24.81±0.17ab	44.24±0.62b	5034.89±23.48b
P0+A	13.96±30.02d	40.86±0.92e	423.28±3.86e	21.57±0.07c	14.91±0.54f	2048.31±16.16g
P1	16.69±60.02c	50.89±0.04d	501.72±2.47d	24.03±0.27b	27.61±0.93d	3972.91±3.59d
P1+A	13.96±30.44d	41.64±1.49e	560.94±8.84c	19.60±0.56d	28.53±0.09d	3765.95±131.21e
P2	14.64±0.89d	41.45±0.12e	429.30±2.41e	21.39±0.13c	21.36±0.71e	3282.32±47.58f
P2+A	13.45±30.76d	54.01±1.86c	640.21±1.35b	20.48±0.71cd	33.99±1.33c	4584.88±104.88c
CK: 对照(P浓度为0.66 mmol?L^–1); P0: 无磷(P浓度为0 mmol?L^–1)处理; P1: 重度缺磷(P浓度为0.22 mmol?L^–1)处理; P2: 轻度缺磷(P浓度为0.44 mmol?L^–1)处理; A: 7.5 mmol?L^–1柠檬酸浸种。同列数据后标不同小写字母表示各处理间差异显著(P < 0.05)。CK: control (with P concentration of 0.66 mmol?L^–1); P0: non-phosphorus (with P concentration of 0 mmol·L^–1); P1: severe phosphorus deficiency (with P concentration of 0.22 mmol·L^–1); P2: mild low phosphorus deficiency (with P concentration of 0.44 mmol·L^–1); A: soaking seed in 7.5 mmol?L^–1 citric acid. Different lowercase letters in each column indicate significantly difference (P < 0.05).

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