热带作物学报

处理Treatment	处理组合Treatment combination
CK₁	H₂O
CK₂	200 mmol/L NaCl
A₁	50 µmol/L ABA+200 mmol/L NaCl
A₂	100 µmol/L ABA+200 mmol/L NaCl
A₃	200 µmol/L ABA+200 mmol/L NaCl
C₁	10 mmol/L CaCl₂+200 mmol/L NaCl
C₂	20 mmol/L CaCl₂+200 mmol/L NaCl
C₃	40 mmol/L CaCl₂+200 mmol/L NaCl
S₁	50 µmol/L SA+200 mmol/L NaCl
S₂	100 µmol/L SA+200 mmol/L NaCl
S₃	200 µmol/L SA+200 mmol/L NaCl

处理Treatment	处理组合Treatment combination
CK₁	H₂O
CK₂	200 mmol/L NaCl
A₁	50 µmol/L ABA+200 mmol/L NaCl
A₂	100 µmol/L ABA+200 mmol/L NaCl
A₃	200 µmol/L ABA+200 mmol/L NaCl
C₁	10 mmol/L CaCl₂+200 mmol/L NaCl
C₂	20 mmol/L CaCl₂+200 mmol/L NaCl
C₃	40 mmol/L CaCl₂+200 mmol/L NaCl
S₁	50 µmol/L SA+200 mmol/L NaCl
S₂	100 µmol/L SA+200 mmol/L NaCl
S₃	200 µmol/L SA+200 mmol/L NaCl

处理Treatment	株高增长量Plant height growth rate/cm	全株干质量Whole plant dry weight/g	根冠比Root shoot ratio
CK₁	1.30±0.25^e	3.77±0.69^e	0.300±0.11^f
CK₂	2.17±0.54^c	5.13±0.46^bc	0.500±0.19^a
A₁	2.34±0.29^c	4.54±0.50^d	0.351±0.09^e
A₂	1.80±0.42^d	5.48±0.73^b	0.397±0.18^c
A₃	1.45±0.37^e	5.09±0.42^c	0.458±0.05^b
C₁	2.27±0.3^c	4.43±0.28^d	0.482±0.09^ab
C₂	2.69±0.49^b	3.93±0.31^e	0.364±0.08^de
C₃	3.33±0.67^a	5.97±0.82^a	0.398±0.13^cd
S₁	2.21±0.45^c	4.57±0.45^d	0.455±0.12^b
S₂	2.35±0.28^c	5.93±0.39^a	0.344±0.06^e
S₃	3.51±0.34^a	5.23±0.40^bc	0.351±0.05^e

处理Treatment	株高增长量Plant height growth rate/cm	全株干质量Whole plant dry weight/g	根冠比Root shoot ratio
CK₁	1.30±0.25^e	3.77±0.69^e	0.300±0.11^f
CK₂	2.17±0.54^c	5.13±0.46^bc	0.500±0.19^a
A₁	2.34±0.29^c	4.54±0.50^d	0.351±0.09^e
A₂	1.80±0.42^d	5.48±0.73^b	0.397±0.18^c
A₃	1.45±0.37^e	5.09±0.42^c	0.458±0.05^b
C₁	2.27±0.3^c	4.43±0.28^d	0.482±0.09^ab
C₂	2.69±0.49^b	3.93±0.31^e	0.364±0.08^de
C₃	3.33±0.67^a	5.97±0.82^a	0.398±0.13^cd
S₁	2.21±0.45^c	4.57±0.45^d	0.455±0.12^b
S₂	2.35±0.28^c	5.93±0.39^a	0.344±0.06^e
S₃	3.51±0.34^a	5.23±0.40^bc	0.351±0.05^e

处理Treatment	可溶性蛋白含量Soluble protein content/(mg·g^-1)	可溶性糖含量Soluble sugar content/(mmol·g^-1)	质膜透性Membrane permeability/%	MDA含量MDA content/(μmol·g^-1)	H₂O₂含量H₂O₂ content/(μmol·g^-1)
CK₁	0.59±0.025^f	25.14±1.29^de	25.28±1.01^g	6.55±0.44^c	22.34±2.90^g
CK₂	0.81±0.021^c	27.47±1.50^bc	36.93±2.04^cd	7.28±0.33^ab	183.49±17.51^c
A₁	0.61±0.031^f	30.85±1.67^a	35.02±1.95^de	7.99±0.52^a	168.04±10.70^c
A₂	0.68±0.035^e	29.01±0.85^ab	34.42±1.73^de	6.79±0.30^bc	162.14±16.00^cd
A₃	0.74±0.040^d	23.91±1.52^ef	35.82±1.10^de	5.53±0.48^de	118.34±17.02^ef
C₁	0.76±0.073^cd	29.69±1.35^ab	33.66±1.30^e	7.79±0.57^a	285.22±14.01^a
C₂	0.91±0.045^b	28.90±1.63^ab	36.97±2.34^cd	7.51±0.89^a	255.04±15.28^b
C₃	0.93±0.040^b	26.70±1.78^cd	30.20±1.55^f	6.49±0.25^c	169.29±22.01^c
S₁	0.73±0.059^d	22.46±2.49^fg	38.86±1.25^bc	6.18±0.29^cd	164.34±20.54^c
S₂	0.93±0.025^b	21.43±3.07^gh	40.76±2.30^ab	5.57±0.27^de	132.47±15.89^de
S₃	1.01±0.030^a	19.59±0.59^h	42.38±1.65^a	5.34±0.67^e	100.86±14.65^e

处理Treatment	可溶性蛋白含量Soluble protein content/(mg·g^-1)	可溶性糖含量Soluble sugar content/(mmol·g^-1)	质膜透性Membrane permeability/%	MDA含量MDA content/(μmol·g^-1)	H₂O₂含量H₂O₂ content/(μmol·g^-1)
CK₁	0.59±0.025^f	25.14±1.29^de	25.28±1.01^g	6.55±0.44^c	22.34±2.90^g
CK₂	0.81±0.021^c	27.47±1.50^bc	36.93±2.04^cd	7.28±0.33^ab	183.49±17.51^c
A₁	0.61±0.031^f	30.85±1.67^a	35.02±1.95^de	7.99±0.52^a	168.04±10.70^c
A₂	0.68±0.035^e	29.01±0.85^ab	34.42±1.73^de	6.79±0.30^bc	162.14±16.00^cd
A₃	0.74±0.040^d	23.91±1.52^ef	35.82±1.10^de	5.53±0.48^de	118.34±17.02^ef
C₁	0.76±0.073^cd	29.69±1.35^ab	33.66±1.30^e	7.79±0.57^a	285.22±14.01^a
C₂	0.91±0.045^b	28.90±1.63^ab	36.97±2.34^cd	7.51±0.89^a	255.04±15.28^b
C₃	0.93±0.040^b	26.70±1.78^cd	30.20±1.55^f	6.49±0.25^c	169.29±22.01^c
S₁	0.73±0.059^d	22.46±2.49^fg	38.86±1.25^bc	6.18±0.29^cd	164.34±20.54^c
S₂	0.93±0.025^b	21.43±3.07^gh	40.76±2.30^ab	5.57±0.27^de	132.47±15.89^de
S₃	1.01±0.030^a	19.59±0.59^h	42.38±1.65^a	5.34±0.67^e	100.86±14.65^e

指标Index	株高Plant height growth rate	全株干质量Whole plant dry weight	根冠比Root shoot ratio	可溶性蛋白含量Soluble protein content	可溶性糖含量Soluble sugar content	质膜透性Membrane permeability	MDA含量MDA content	H₂O₂含量H₂O₂ content	SOD活性SOD activity	POD活性POD activity
株高	1.000	0.383^*	-0.109	0.796^**	-0.147	0.409^*	-0.090	0.272	0.247	0.560^**
全株干质量		1.000	0.088	0.269	0.009	0.545^**	0.072	0.319	0.509^**	0.120
根冠比			1.000	0.003	0.203	0.174	0.137	0.527^**	0.129	0.498^**
可溶性蛋白含量				1.000	-0.395^*	0.545^**	-0.399^*	0.211	0.100	0.501^**
可溶性糖含量					1.000	-0.371^*	0.848^*	0.551^**	0.391^*	0.282
质膜透性						1.000	-0.340	0.215	0.311	0.300
MDA含量							1.000	0.571^**	0.416^*	0.243
H₂O₂含量								1.000	0.533^**	0.736^**
SOD活性									1.000	0.352^*
POD活性										1.000

指标Index	株高Plant height growth rate	全株干质量Whole plant dry weight	根冠比Root shoot ratio	可溶性蛋白含量Soluble protein content	可溶性糖含量Soluble sugar content	质膜透性Membrane permeability	MDA含量MDA content	H₂O₂含量H₂O₂ content	SOD活性SOD activity	POD活性POD activity
株高	1.000	0.383^*	-0.109	0.796^**	-0.147	0.409^*	-0.090	0.272	0.247	0.560^**
全株干质量		1.000	0.088	0.269	0.009	0.545^**	0.072	0.319	0.509^**	0.120
根冠比			1.000	0.003	0.203	0.174	0.137	0.527^**	0.129	0.498^**
可溶性蛋白含量				1.000	-0.395^*	0.545^**	-0.399^*	0.211	0.100	0.501^**
可溶性糖含量					1.000	-0.371^*	0.848^*	0.551^**	0.391^*	0.282
质膜透性						1.000	-0.340	0.215	0.311	0.300
MDA含量							1.000	0.571^**	0.416^*	0.243
H₂O₂含量								1.000	0.533^**	0.736^**
SOD活性									1.000	0.352^*
POD活性										1.000

处理Treatment	隶属函数值Membership function value	总得分Total Points	位次Precedence
CK₁	0.000	0.618	1.000	0.000	0.493	1.000	0.543	1.000	0.000	0.000	4.036	10
CK₂	0.394	0.000	0.000	0.524	0.700	0.319	0.268	0.387	0.587	0.557	4.354	9
A₁	0.471	0.350	0.745	0.048	1.000	0.430	0.000	0.446	0.978	0.403	4.871	6
A₂	0.226	0.777	0.515	0.214	0.837	0.465	0.928	0.468	0.932	0.505	5.392	4
A₃	0.068	0.600	0.210	0.357	0.384	0.384	0.453	0.635	0.712	0.460	4.738	7
C₁	0.439	0.300	0.190	0.405	0.897	0.510	0.075	0.000	0.616	1.000	4.432	8
C₂	0.629	0.073	0.680	0.762	0.827	0.316	0.181	0.115	1.000	0.641	5.224	5
C₃	0.919	1.000	0.510	0.810	0.631	0.712	0.566	0.441	0.721	0.646	6.956	1
S₁	0.412	0.364	0.225	0.333	0.256	0.206	0.683	0.581	0.686	0.374	3.946	11
S₂	0.475	0.982	0.780	0.810	0.163	0.095	0.913	0.460	0.671	0.321	5.844	3
S₃	1.000	0.664	0.745	1.000	0.000	0.000	1.000	0.701	0.521	0.706	6.337	2

处理Treatment	隶属函数值Membership function value	总得分Total Points	位次Precedence
CK₁	0.000	0.618	1.000	0.000	0.493	1.000	0.543	1.000	0.000	0.000	4.036	10
CK₂	0.394	0.000	0.000	0.524	0.700	0.319	0.268	0.387	0.587	0.557	4.354	9
A₁	0.471	0.350	0.745	0.048	1.000	0.430	0.000	0.446	0.978	0.403	4.871	6
A₂	0.226	0.777	0.515	0.214	0.837	0.465	0.928	0.468	0.932	0.505	5.392	4
A₃	0.068	0.600	0.210	0.357	0.384	0.384	0.453	0.635	0.712	0.460	4.738	7
C₁	0.439	0.300	0.190	0.405	0.897	0.510	0.075	0.000	0.616	1.000	4.432	8
C₂	0.629	0.073	0.680	0.762	0.827	0.316	0.181	0.115	1.000	0.641	5.224	5
C₃	0.919	1.000	0.510	0.810	0.631	0.712	0.566	0.441	0.721	0.646	6.956	1
S₁	0.412	0.364	0.225	0.333	0.256	0.206	0.683	0.581	0.686	0.374	3.946	11
S₂	0.475	0.982	0.780	0.810	0.163	0.095	0.913	0.460	0.671	0.321	5.844	3
S₃	1.000	0.664	0.745	1.000	0.000	0.000	1.000	0.701	0.521	0.706	6.337	2

外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗生长及生理的影响

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刘艳菊 , 徐玉芬 , 于钊妍 , 吴坤林 , 曹秋林 , 龚涵 , 贾效成 ^*

热带作物学报 | 作物栽培与生理生化 2025,46(3): 695-704

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热带作物学报 |作物栽培与生理生化 2025 , 46 (3) : 695 -704

外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗生长及生理的影响

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刘艳菊, 徐玉芬, 于钊妍, 吴坤林, 曹秋林, 龚涵, 贾效成^*

作者信息

中国热带农业科学院椰子研究所/海南省热带油料作物生物学重点实验室，海南文昌　571339

通讯作者:

^* 贾效成（JIA Xiaocheng），E-mail：xcjia1@163.com。

Growth and Physiological Effects of Exogenous ABA, CaCl₂, SA under NaCl Stress on Camellia oleifera Seedlings

Yanju LIU, Yufen XU, Zhaoyan YU, Kunlin WU, Qiulin CAO, Han GONG, Xiaocheng JIA^*

Affiliations

Coconut Research Institute, Chinese Academy of Tropical Agricultural Sciences / Hainan Key Laboratory of Tropical Oil Crops Biology, Wenchang, Hainan 571339, China

出版时间: 2025-03-25 doi: 10.3969/j.issn.1000-2561.2025.03.017

文章导航

摘要

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探讨外源脱落酸（ABA）、氯化钙（CaCl₂）、水杨酸（SA）对NaCl胁迫下油茶幼苗生长和生理特性的影响，为提高油茶抗性提供理论依据。以一年生油茶幼苗为材料，NaCl胁迫下外施不同浓度ABA、CaCl₂、SA处理，研究其对油茶幼苗生长及相关生理指标的影响。结果表明：（1）单独NaCl胁迫下油茶幼苗株高增长量、全株干质量、根冠比、可溶性蛋白含量、可溶性糖含量、质膜透性、丙二醛（MDA）含量、H₂O₂含量、SOD和POD活性均显著提升；（2）NaCl胁迫下外施不同浓度ABA、CaCl₂、SA处理后可增加油茶幼苗株高增长量、全株干质量，提高幼苗可溶性蛋白含量、可溶性糖含量、SOD和POD活性，降低油茶幼苗根冠比、质膜透性、MDA含量；（3）利用隶属法进行综合评价结果发现，外施40 mmol/L CaCl₂最能减少NaCl胁迫对油茶幼苗的伤害，其次是200 µmol/L SA、100 µmol/L SA处理。

关键词

油茶 / NaCl胁迫 / ABA / CaCl₂ / SA / 生理

Abstract

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This study investigated the effects of exogenous abscisic acid (ABA), calcium chloride (CaCl₂), and salicylic acid (SA) on the growth and physiological characterisics of Camellia oleifera seedlings subjected to NaCl stress, aiming to provide theoretical basis for enhancing the resistance of tea oil plants. One-year-old C. oleifera seedlings were utilized, and the impact of varying concentrations of ABA, CaCl₂, and SA on growth parameters and related physiological indicators was assessed under NaCl stress. Under sole NaCl stress conditiongs, significant increase was observed in height growth, total dry weight, root-to-shoot ratio, soluble protein content, soluble sugar levels, membrane permeability, MDA, H₂O₂ content, as well as SOD and POD activities in C. oleifera seedlings. Furthermore, external application of different substances under NaCl stress enhanced height growth, total dry weight, soluble protein content, soluble sugar levels while reducing root-to-shoot ratio along with membrane permeability and MDA content in the seedlings. Comprehensive evaluation using membership function analysis indicated that treatment with 40 mmol/L CaCl₂ was most effective in mitigating NaCl-induced damage to C. oleifera seedlings; this was followed by treatments with 200 µmol/L SA and 100 µmol/L SA.

Key words

Camellia oleifera / NaCl stress / ABA / CaCl₂ / SA / physiology

引用本文

刘艳菊, 徐玉芬, 于钊妍, 吴坤林, 曹秋林, 龚涵, 贾效成. 外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗生长及生理的影响. 热带作物学报, 2025 , 46 (3) : 695 -704 . DOI: 10.3969/j.issn.1000-2561.2025.03.017

Yanju LIU, Yufen XU, Zhaoyan YU, Kunlin WU, Qiulin CAO, Han GONG, Xiaocheng JIA. Growth and Physiological Effects of Exogenous ABA, CaCl₂, SA under NaCl Stress on Camellia oleifera Seedlings[J]. Chinese Journal of Tropical Crops, 2025 , 46 (3) : 695 -704 . DOI: 10.3969/j.issn.1000-2561.2025.03.017

正文

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油茶（Camellia vietnamensis）为山茶科山茶属（Camellia L.）常绿小乔木，是我国南方主要的经济树种，与椰子、油棕、油橄榄称为世界四大木本食用油料树种^[1]。具有较高的经济价值和生态效益，是农林业产业结构调整、精准扶贫的重要经济作物之一^[2]。滨海盐碱地是我国重要的盐碱地土地类型，主要受海水影响形成，具有盐分高、养分低和危害大等特点。海南岛是我国第二大岛，海岸线总长约1800 km，拥有丰富的滨海资源，但也因台风等天气原因，造成海水倒灌形成盐碱地。同时，不合理施肥、工业污染和土壤资源过度开发等也会导致土壤盐碱化，阻碍植物正常生长，严重影响生态发展。研究表明，盐胁迫下利用外源植物生长调节剂可提高抗逆性，维持植物生长发育^[3-4]。脱落酸（ABA）能调节种子萌发、幼苗生长及对非生物胁迫的耐受性等生理过程^[5]。当植物叶片遭受盐胁迫时，叶片中的ABA含量增加。而当植物适应了盐胁迫后，ABA含量降低，叶片恢复正常的代谢水平和生长状态^[6]。ABA可关闭植物气孔，减少蒸腾速率，减少地上部盐离子含量，从而增强植物适应盐胁迫的能力^[7-8]。ABA能诱导植物选择性地吸收Ca²⁺，提高植物对Na⁺的耐受性^[9]。

水杨酸（salicyic aicd, SA）作为天然的信号分子，通过提高植物叶绿素含量^[10]和抗氧化酶活性^[11-12]、降低植物蒸腾速率^[13]、增加渗透物质含量^[14]等生理变化，进而提高植株耐盐性。外源SA可提高植物抗氧化酶活性（SOD、POD、CAT等），降低MDA含量，同时增加生物量，进而提高抗盐能力^[15-18]，能促进小麦幼苗生长和增强根系活力^[19]，增加脯氨酸含量^[20]，增加可溶性糖含量^[21]。Ca²⁺作为植物细胞内第二信使，对低温、干旱、盐等逆境胁迫中的信号转导有着重要作用。适宜浓度的Ca²⁺可通过钙调节系统来维持细胞质膜稳定性，进而提高植物抗盐性，浓度过高的Ca²⁺则会对植物产生毒害作用^[22-23]。有研究结果表明，外源Ca²⁺能缓解盐胁迫对植物生长有较好的效果，可能是因为太多的Na⁺、Cl^-抑制Ca²⁺、K⁺的吸收，致使离子缺失严重^[24]。Ca²⁺能改善苜蓿幼苗离子平衡，缓解盐胁迫造成的伤害^[25]。不同浓度CaCl₂可缓解盐胁迫下对小麦种子萌发的抑制，维持小麦幼苗叶片渗透压，增强小麦种子和幼苗的耐盐性^[26]。

目前，针对油茶逆境生理生化的研究大多集中在低温^[27]、高温^[28]、干旱^[29-30]等方面，开展盐胁迫对油茶的研究较少，仅有外源NO^[31]、芸苔素内酯、水杨酸、赤霉素和褪黑素^[32]对NaCl胁迫下油茶幼苗生理变化的研究，而外施不同浓度ABA、SA和CaCl₂对盐胁迫下油茶生理变化方面的研究未见报道，因此本研究以油茶嫁接苗为研究对象，开展NaCl胁迫下不同浓度ABA、SA和CaCl₂对油茶幼苗生长和生理指标的影响，为油茶抗盐栽培提供理论依据和技术支持。

1　材料与方法

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1.1　材料

试验在中国热带农业科学院椰子研究所油茶种苗圃进行，油茶品种为海南省品种审定委员会认定的热研1号。选取长势一致，健康良好，苗高40 cm左右的一年生油茶盆栽幼苗用于试验。

1.2　方法

1.2.1　试验设计

将200 mmol/L NaCl溶液浸没营养杯的1/3处，3 d更换一次NaCl溶液以维持NaCl处理浓度，同时每天上午9：00对油茶幼苗喷施40 mL不同浓度的ABA、CaCl₂、SA溶液（表1），每个处理10盆，3次重复。等体积清水处理的幼苗为对照1（CK₁），单独NaCl处理为对照2（CK₂）。处理7 d后，取心叶外围第4、5片叶进行相关指标的测定。

1.2.2　指标测定

（1）生长指标的测定。油茶幼苗土面到顶端新芽芽尖的距离为株高，喷施外源物质处理前和NaCl胁迫7 d后分别测量单株幼苗的株高，其差值为株高增加量；全株收获洗净，吸干表面水分后称重以测量鲜质量，于110 ℃烘箱杀青15 min，以80 ℃烘干至恒质量；茎叶干质量和根干质量的总和为全株干质量，根干质量与茎叶干质量的比值为根冠比。

（2）生理指标的测定。可溶性蛋白含量测定采用考马斯亮蓝G-250法，可溶性糖含量测定采用蒽酮比色法^[33]，H₂O₂含量测定采用硫酸钛法^[34]，MDA含量测定采用硫代巴比妥酸法^[35]，脯氨酸含量测定采用磺基水杨酸法^[36]，质膜透性测定采用电导法^[37]，SOD和POD活性应用南京建成试剂盒进行测定；吸光值由日本岛津UV-2600紫外分光光度计测定。

1.3　数据处理

试验数据用Excel 2010软件进行收集与整理，数据采用SPSS 25软件进行差异显著性分析。采用隶属函数值法对耐盐性进行综合评价，分别计算NaCl胁迫下外施不同物质油茶幼苗的株高、全株干质量、根冠比、可溶性蛋白含量、可溶性糖含量、MDA含量、质膜透性及SOD、POD活性等指标变化的隶属函数值。

分析指标与耐盐性呈正相关，则计算公式为：X=(X_ij-X_min)/(X_max-X_min)；分析指标与耐盐性呈负相关，则计算公式为：X=1-(X_ij-X_min)/(X_max-X_min)。式中：X_ij表示第i处理第j个指标的隶属函数值；X_max和X_min分别表示测定值中的最大值和最小值。分别计算各指标隶属函数值，再计算所有指标平均值，值越大，耐盐性越强，反之越弱。

2　结果与分析

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2.1　外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗生长指标的影响

由表2可知，单独NaCl胁迫下，幼苗株高增长量、全株干质量和根冠比均显著上升，与CK₁相比，分别增加66.92%、36.07%、66.67%。外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗株高增长量、全株干质量、根冠比产生不同的影响。ABA处理下，株高增长量随ABA浓度增加呈下降趋势，其中A₁处理的株高增长量与CK₂相比差异不显著，但比CK₁显著增加80%，A₂处理也比CK₁显著增加38.46%，但比CK₂显著减少17.05%，A₃处理与CK₁相比差异不显著，但比CK₂显著减少33.18%；株高增长量随CaCl₂浓度增加呈上升趋势，C₁处理与CK₂相比差异不显著，但比CK₁显著增加74.62%，C₂处理分别比CK₁、CK₂显著增加106.92%、23.96%，C₃处理分别比CK₁、CK₂、C₁、C₂、A₁显著增加156.15%、53.46%、46.70%、23.79%、42.71%；株高增长量随SA浓度增加呈上升趋势，但S₁、S₂均与CK₂差异不显著，但均比CK₁显著增加70%、80.77%，S₃处理的株高增长量达到最大，分别比CK₁、CK₂、S₂显著增加170%、61.75%、49.36%，但与C₃处理差异不显著。

全株干质量均随ABA浓度的增大呈先升高后下降趋势，A₁处理比CK₂显著下降11.50%，A₂、A₃处理与CK₂差异不显著；而在CaCl₂处理下，全株干质量呈先下降后上升趋势，C₁、C₂分别比CK₂显著下降13.65%、23.39%，而C₃则比CK₂显著增加16.37%。SA处理下，全株干质量在S₂处理下较大，分别比CK₂、S₁、S₃显著增加15.59%、29.76%、11.80%。

不同浓度ABA、CaCl₂、SA处理的根冠比与CK₂相比均显著下降，其中C₁处理与CK₂差异不显著，A₃、S₁分别比CK₂减少8.4%、9%，A₁、C₂、S₂、S₃下降幅度较大，与CK₂相比，分别减少29.8%、27.2%、31.2%、29.8%。

2.2　外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗生理指标的影响

NaCl胁迫下外源ABA、CaCl₂、SA对油茶幼苗7个生理指标的影响如表3所示。单独NaCl胁迫下，油茶幼苗可溶性蛋白含量、可溶性糖含量、质膜透性、丙二醛含量、脯氨酸含量、H₂O₂含量较CK₁均有显著提高；ABA处理下，幼苗可溶性蛋白含量随ABA浓度的增加呈升高趋势，A₁与CK₁差异不大，A₂、A₃分别比CK₁显著增加15.25%、25.42%，但比CK₂均显著下降；C₃处理的可溶性蛋白含量比CK₂显著增加14.81%，S₃处理的可溶性蛋白含量比CK₂显著增加24.69%，说明高浓度SA处理比ABA、CaCl₂更能提高油茶幼苗可溶性蛋白含量；随着ABA、CaCl₂、SA处理浓度的增加，可溶性糖含量均呈逐渐下降趋势，A₁、C₁处理的可溶性糖含量分别比CK₂增加12.30%、8.08%，而A₃处理比CK₂显著降低12.96%，C₃处理与CK₂差异不显著，不同浓度SA处理比CK₂均显著降低，分别减少18.24%、21.99%、28.69%。说明低浓度ABA、CaCl₂可提高油茶幼苗可溶性糖含量。

不同浓度ABA处理的质膜透性与CK₂差异不显著，而CaCl₂处理下，与CK₂相比，C₁、C₃处理显著减少8.85%、18.22%。SA处理下，S₂、S₃处理的质膜透性比CK₂分别显著增加10.37%、14.76%。说明高浓度CaCl₂处理可降低油茶幼苗质膜透性，而较高浓度SA处理则提高油茶幼苗质膜透性。

不同浓度ABA、CaCl₂、SA处理的幼苗MDA随外源物质浓度的增加均呈逐渐下降趋势，A₁、C₁与CK₂差异不显著，A₃、C₃、S₃分别比CK₂减少24.04%、10.85%、26.65%，不同浓度SA处理均能显著降低MDA含量，减少NaCl对油茶幼苗的氧化伤害。

ABA、CaCl₂、SA处理对NaCl胁迫下油茶幼苗H₂O₂含量产生不同的影响，A₁、A₂处理的H₂O₂含量与CK₂差异不显著，但A₃处理显著下降35.51%，而较低浓度CaCl₂处理则显著提高幼苗H₂O₂含量，C₁、C₂分别显著增加55.44%、38.99%，C₃处理与CK₂差异不显著。S₂、S₃处理显著降低，分别比CK₂减少27.81%、45.03%，而S₁处理与CK₂差异不显著。

2.3　外源ABA、CaCl₂、SA对NaCl胁迫下油茶幼苗SOD、POD活性的影响

NaCl胁迫下外源ABA、CaCl₂、SA对油茶幼苗SOD活性的影响如图1所示。单独NaCl胁迫下，油茶幼苗SOD活性较CK₁有显著提高，增加3.19倍；较低浓度ABA处理下，A₁、A₂的SOD活性显著提高，分别比CK₂增加20.30%、17.92%，而A₃处理与CK₂差异不显著；C₁、C₃处理与CK₂差异不显著，C₂处理的SOD活性增幅最高，比CK₂增加21.46%。不同浓度SA处理与CK₂差异均不显著，S₁、S₂处理仅比CK₂增加5.11%、4.37%，S₃处理比CK₂减少3.45%。说明较低浓度ABA和适当浓度CaCl₂能提高油茶幼苗SOD活性。

单独NaCl胁迫下，油茶幼苗POD活性较CK₁有显著提高，分别增加3.19倍；由图2所示，不同浓度ABA处理的油茶幼苗POD活性均较CK₂有所降低，其中A₁处理比CK₂显著减少20.97%，A₂、A₃处理与CK₂差异不显著；不同浓度CaCl₂处理油茶幼苗POD活性较CK₂呈显著提升，其中C₁处理的POD活性增幅最高，比CK₂增加60.61%，C₂、C₃处理分别增加11.51%、12.16%；S₁、S₂处理的POD活性显著下降，比CK₂减少24.93%、32.28%，而S₃处理的POD活性则比CK₂显著增加20.45%。说明CaCl₂和高浓度SA处理可提高油茶幼苗POD活性。

2.4　外源ABA、CaCl₂、SA对NaCl胁迫下各指标的相关性分析

由表4可知，NaCl胁迫下油茶幼苗株高与可溶性蛋白含量、POD活性呈极显著正相关（P<0.01），与全株干质量、质膜透性呈显著正相关（P<0.05）；全株干质量与质膜透性、SOD活性呈极显著正相关（P<0.01）；根冠比与H₂O₂含量、POD活性呈极显著正相关（P<0.01）；可溶性蛋白含量与可溶性糖含量、MDA含量呈显著负相关（P<0.05），与质膜透性、POD活性呈极显著正相关（P<0.01）；可溶性糖含量与质膜透性呈显著负相关（P<0.05），与MDA含量、SOD活性呈显著正相关（P<0.01）；MDA含量与H₂O₂含量呈极显著正相关（P<0.01），与SOD活性呈显著正相关（P<0.05）；过氧化氢含量与SOD活性、POD活性呈极显著正相关（P<0.01）；SOD活性与POD活性呈显著正相关（P<0.05）。

2.5　外源ABA、CaCl₂、SA处理下油茶幼苗耐盐胁迫的综合评价

通过NaCl胁迫下油茶幼苗的株高、全株干质量、根冠比、可溶性蛋白含量、可溶性糖含量、质膜透性、MDA含量、SOD活性、POD活性等10项抗盐相关生长和生理指标变化幅度的隶属函数值及总分排序（表5），总分越高表示外源物质处理越能提高油茶幼苗耐盐性，不同外源物质和浓度处理的幼苗耐盐性排序为C₃>S₃>S₂>A₂>C₂>A₁>A₃>C₁>CK₂>CK₁>S₁。说明外施40 mmol/L CaCl₂最能减少NaCl胁迫对油茶幼苗的伤害，其次是200 µmol/L SA、100 µmol/L SA处理。

3　讨论

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超过一定浓度的盐胁迫会显著抑制植物生长，表现为株高减少、生物量减少、根冠比增加等^[38-39]。本研究中，单独200 mmol/L NaCl胁迫下油茶幼苗株高增长量、全株干质量、根冠比、可溶性蛋白、可溶性糖含量、质膜透性、MDA含量、H₂O₂含量均显著升高，与作者之前研究结果相一致^[40]，说明油茶幼苗具有一定的耐盐性。

可溶性蛋白和可溶性糖可作为重要的渗透调节物质，能有效地缓解盐胁迫对玉米幼苗根系和地上部生长的抑制作用^[41]，本研究中，较高浓度的CaCl₂和SA处理均显著提高幼苗可溶性蛋白含量，较低浓度的ABA和CaCl₂处理显著提高幼苗可溶性糖含量，这与粟莉圆等^[42]对火力楠幼苗研究结果一致，可溶性糖含量的积累通过调节盐胁迫下植物细胞渗透电位及渗透保护作用，降低细胞渗透压，吸收更多的水分维持细胞膨压^[43]，缓解盐害，进而维持植株正常生长。

过度积累的Na⁺使细胞膜的选择透性丧失，质膜透性增加，电解质大量渗透^[44]。本研究发现，ABA处理对幼苗质膜透性影响不大，较高浓度的CaCl₂显著降低幼苗质膜透性，而较高浓度SA则可提高幼苗质膜透性，说明外施不同物质对油茶幼苗质膜透性产生不同影响，其机制原因还有待进一步实验验证。作为植物细胞膜膜质过氧化的产物MDA，能破坏细胞膜结构和功能^[45]，其含量越高说明植物受损伤越严重^[46]。本研究发现，SA处理和较高浓度的ABA、CaCl₂处理可显著降低幼苗MDA含量，可能是Ca²⁺减轻幼苗细胞膜质过氧化程度，减轻NaCl胁迫对油茶幼苗的氧化伤害，这与WANG等^[47]对黄瓜幼苗研究结果一致。H₂O₂含量能直接反映植物受逆境胁迫的程度^[48]。本研究发现，外施ABA和SA均显著降低油茶幼苗H₂O₂含量，其中不同浓度的SA处理降幅更为明显，200 µmol/L SA处理较CK₂显著减少45.03%。而较低浓度CaCl₂则显著提高幼苗H₂O₂含量，仅在40 mmol/L CaCl₂处理下与CK₂处理的差异不显著；

SOD和POD作为抗氧化酶类物质，能降低植物体内自由基、活性氧含量，避免膜质过氧化，进而提高植物的抗逆能力^[49]。本研究发现，较低浓度ABA和适当浓度CaCl₂能提高油茶幼苗SOD活性，CaCl₂和高浓度SA处理可提高油茶幼苗POD活性。这可能是Ca²⁺和钙调蛋白相结合，进而提高抗氧化酶活性，这与屈施旭等^[50]对火麻幼苗、PALMA等^[51]对菜豆、周希琴等^[52]对木麻黄幼苗的研究结果一致。

综上所述，外施适宜浓度的ABA、CaCl₂和SA对油茶幼苗NaCl胁迫均有缓解作用，经过综合评价发现外施40 mmol/L CaCl₂最能减少NaCl胁迫对油茶幼苗的伤害，其次是200 µmol/L SA和100 µmol/L SA处理。在海南，靠近海边种植油茶幼苗，可适当喷施40 mmol/L CaCl₂，提高幼苗耐盐性，促进其生长。

基金

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海南省重点研发计划专项(ZDYF2023XDNY054)
海南省自然科学基金青年基金项目(322QN414)

参考文献

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文献

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2025年第46卷第3期

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doi: 10.3969/j.issn.1000-2561.2025.03.017

接收时间：2024-09-30
首发时间：2026-06-25
出版时间：2025-03-25

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收稿日期：2024-09-30
录用日期：2024-10-14

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海南省重点研发计划专项(ZDYF2023XDNY054)

海南省自然科学基金青年基金项目(322QN414)

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中国热带农业科学院椰子研究所/海南省热带油料作物生物学重点实验室，海南文昌　571339

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^* 贾效成（JIA Xiaocheng），E-mail：xcjia1@163.com。

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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处理Treatment	处理组合Treatment combination
CK₁	H₂O
CK₂	200 mmol/L NaCl
A₁	50 µmol/L ABA+200 mmol/L NaCl
A₂	100 µmol/L ABA+200 mmol/L NaCl
A₃	200 µmol/L ABA+200 mmol/L NaCl
C₁	10 mmol/L CaCl₂+200 mmol/L NaCl
C₂	20 mmol/L CaCl₂+200 mmol/L NaCl
C₃	40 mmol/L CaCl₂+200 mmol/L NaCl
S₁	50 µmol/L SA+200 mmol/L NaCl
S₂	100 µmol/L SA+200 mmol/L NaCl
S₃	200 µmol/L SA+200 mmol/L NaCl

处理Treatment

处理组合Treatment combination

CK₁

H₂O

CK₂

200 mmol/L NaCl

A₁

50 µmol/L ABA+200 mmol/L NaCl

A₂

100 µmol/L ABA+200 mmol/L NaCl

A₃

200 µmol/L ABA+200 mmol/L NaCl

C₁

10 mmol/L CaCl₂+200 mmol/L NaCl

C₂

20 mmol/L CaCl₂+200 mmol/L NaCl

C₃

40 mmol/L CaCl₂+200 mmol/L NaCl

S₁

50 µmol/L SA+200 mmol/L NaCl

S₂

100 µmol/L SA+200 mmol/L NaCl

S₃

200 µmol/L SA+200 mmol/L NaCl

处理Treatment	株高增长量Plant height growth rate/cm	全株干质量Whole plant dry weight/g	根冠比Root shoot ratio
CK₁	1.30±0.25^e	3.77±0.69^e	0.300±0.11^f
CK₂	2.17±0.54^c	5.13±0.46^bc	0.500±0.19^a
A₁	2.34±0.29^c	4.54±0.50^d	0.351±0.09^e
A₂	1.80±0.42^d	5.48±0.73^b	0.397±0.18^c
A₃	1.45±0.37^e	5.09±0.42^c	0.458±0.05^b
C₁	2.27±0.3^c	4.43±0.28^d	0.482±0.09^ab
C₂	2.69±0.49^b	3.93±0.31^e	0.364±0.08^de
C₃	3.33±0.67^a	5.97±0.82^a	0.398±0.13^cd
S₁	2.21±0.45^c	4.57±0.45^d	0.455±0.12^b
S₂	2.35±0.28^c	5.93±0.39^a	0.344±0.06^e
S₃	3.51±0.34^a	5.23±0.40^bc	0.351±0.05^e

处理Treatment

株高增长量Plant height growth rate/cm

全株干质量Whole plant dry weight/g

根冠比Root shoot ratio

CK₁

1.30±0.25^e

3.77±0.69^e

0.300±0.11^f

CK₂

2.17±0.54^c

5.13±0.46^bc

0.500±0.19^a

A₁

2.34±0.29^c

4.54±0.50^d

0.351±0.09^e

A₂

1.80±0.42^d

5.48±0.73^b

0.397±0.18^c

A₃

1.45±0.37^e

5.09±0.42^c

0.458±0.05^b

C₁

2.27±0.3^c

4.43±0.28^d

0.482±0.09^ab

C₂

2.69±0.49^b

3.93±0.31^e

0.364±0.08^de

C₃

3.33±0.67^a

5.97±0.82^a

0.398±0.13^cd

S₁

2.21±0.45^c

4.57±0.45^d

0.455±0.12^b

S₂

2.35±0.28^c

5.93±0.39^a

0.344±0.06^e

S₃

3.51±0.34^a

5.23±0.40^bc

0.351±0.05^e

处理Treatment	可溶性蛋白含量Soluble protein content/(mg·g^-1)	可溶性糖含量Soluble sugar content/(mmol·g^-1)	质膜透性Membrane permeability/%	MDA含量MDA content/(μmol·g^-1)	H₂O₂含量H₂O₂ content/(μmol·g^-1)
CK₁	0.59±0.025^f	25.14±1.29^de	25.28±1.01^g	6.55±0.44^c	22.34±2.90^g
CK₂	0.81±0.021^c	27.47±1.50^bc	36.93±2.04^cd	7.28±0.33^ab	183.49±17.51^c
A₁	0.61±0.031^f	30.85±1.67^a	35.02±1.95^de	7.99±0.52^a	168.04±10.70^c
A₂	0.68±0.035^e	29.01±0.85^ab	34.42±1.73^de	6.79±0.30^bc	162.14±16.00^cd
A₃	0.74±0.040^d	23.91±1.52^ef	35.82±1.10^de	5.53±0.48^de	118.34±17.02^ef
C₁	0.76±0.073^cd	29.69±1.35^ab	33.66±1.30^e	7.79±0.57^a	285.22±14.01^a
C₂	0.91±0.045^b	28.90±1.63^ab	36.97±2.34^cd	7.51±0.89^a	255.04±15.28^b
C₃	0.93±0.040^b	26.70±1.78^cd	30.20±1.55^f	6.49±0.25^c	169.29±22.01^c
S₁	0.73±0.059^d	22.46±2.49^fg	38.86±1.25^bc	6.18±0.29^cd	164.34±20.54^c
S₂	0.93±0.025^b	21.43±3.07^gh	40.76±2.30^ab	5.57±0.27^de	132.47±15.89^de
S₃	1.01±0.030^a	19.59±0.59^h	42.38±1.65^a	5.34±0.67^e	100.86±14.65^e

处理Treatment

可溶性蛋白含量Soluble protein content/(mg·g^-1)

可溶性糖含量Soluble sugar content/(mmol·g^-1)

质膜透性Membrane permeability/%

MDA含量MDA content/(μmol·g^-1)

H₂O₂含量H₂O₂ content/(μmol·g^-1)

CK₁

0.59±0.025^f

25.14±1.29^de

25.28±1.01^g

6.55±0.44^c

22.34±2.90^g

CK₂

0.81±0.021^c

27.47±1.50^bc

36.93±2.04^cd

7.28±0.33^ab

183.49±17.51^c

A₁

0.61±0.031^f

30.85±1.67^a

35.02±1.95^de

7.99±0.52^a

168.04±10.70^c

A₂

0.68±0.035^e

29.01±0.85^ab

34.42±1.73^de

6.79±0.30^bc

162.14±16.00^cd

A₃

0.74±0.040^d

23.91±1.52^ef

35.82±1.10^de

5.53±0.48^de

118.34±17.02^ef

C₁

0.76±0.073^cd

29.69±1.35^ab

33.66±1.30^e

7.79±0.57^a

285.22±14.01^a

C₂

0.91±0.045^b

28.90±1.63^ab

36.97±2.34^cd

7.51±0.89^a

255.04±15.28^b

C₃

0.93±0.040^b

26.70±1.78^cd

30.20±1.55^f

6.49±0.25^c

169.29±22.01^c

S₁

0.73±0.059^d

22.46±2.49^fg

38.86±1.25^bc

6.18±0.29^cd

164.34±20.54^c

S₂

0.93±0.025^b

21.43±3.07^gh

40.76±2.30^ab

5.57±0.27^de

132.47±15.89^de

S₃

1.01±0.030^a

19.59±0.59^h

42.38±1.65^a

5.34±0.67^e

100.86±14.65^e

指标Index	株高Plant height growth rate	全株干质量Whole plant dry weight	根冠比Root shoot ratio	可溶性蛋白含量Soluble protein content	可溶性糖含量Soluble sugar content	质膜透性Membrane permeability	MDA含量MDA content	H₂O₂含量H₂O₂ content	SOD活性SOD activity	POD活性POD activity
株高	1.000	0.383^*	-0.109	0.796^**	-0.147	0.409^*	-0.090	0.272	0.247	0.560^**
全株干质量		1.000	0.088	0.269	0.009	0.545^**	0.072	0.319	0.509^**	0.120
根冠比			1.000	0.003	0.203	0.174	0.137	0.527^**	0.129	0.498^**
可溶性蛋白含量				1.000	-0.395^*	0.545^**	-0.399^*	0.211	0.100	0.501^**
可溶性糖含量					1.000	-0.371^*	0.848^*	0.551^**	0.391^*	0.282
质膜透性						1.000	-0.340	0.215	0.311	0.300
MDA含量							1.000	0.571^**	0.416^*	0.243
H₂O₂含量								1.000	0.533^**	0.736^**
SOD活性									1.000	0.352^*
POD活性										1.000

指标Index

株高Plant height growth rate

全株干质量Whole plant dry weight

根冠比Root shoot ratio

可溶性蛋白含量Soluble protein content

可溶性糖含量Soluble sugar content

质膜透性Membrane permeability

MDA含量MDA content

H₂O₂含量H₂O₂ content

SOD活性SOD activity

POD活性POD activity

株高

1.000

0.383^*

-0.109

0.796^**

-0.147

0.409^*

-0.090

0.272

0.247

0.560^**

全株干质量

1.000

0.088

0.269

0.009

0.545^**

0.072

0.319

0.509^**

0.120

根冠比

1.000

0.003

0.203

0.174

0.137

0.527^**

0.129

0.498^**

可溶性蛋白含量

1.000

-0.395^*

0.545^**

-0.399^*

0.211

0.100

0.501^**

可溶性糖含量

1.000

-0.371^*

0.848^*

0.551^**

0.391^*

0.282

质膜透性

1.000

-0.340

0.215

0.311

0.300

MDA含量

1.000

0.571^**

0.416^*

0.243

H₂O₂含量

1.000

0.533^**

0.736^**

SOD活性

1.000

0.352^*

POD活性

1.000

处理Treatment	隶属函数值Membership function value	总得分Total Points	位次Precedence
CK₁	0.000	0.618	1.000	0.000	0.493	1.000	0.543	1.000	0.000	0.000	4.036	10
CK₂	0.394	0.000	0.000	0.524	0.700	0.319	0.268	0.387	0.587	0.557	4.354	9
A₁	0.471	0.350	0.745	0.048	1.000	0.430	0.000	0.446	0.978	0.403	4.871	6
A₂	0.226	0.777	0.515	0.214	0.837	0.465	0.928	0.468	0.932	0.505	5.392	4
A₃	0.068	0.600	0.210	0.357	0.384	0.384	0.453	0.635	0.712	0.460	4.738	7
C₁	0.439	0.300	0.190	0.405	0.897	0.510	0.075	0.000	0.616	1.000	4.432	8
C₂	0.629	0.073	0.680	0.762	0.827	0.316	0.181	0.115	1.000	0.641	5.224	5
C₃	0.919	1.000	0.510	0.810	0.631	0.712	0.566	0.441	0.721	0.646	6.956	1
S₁	0.412	0.364	0.225	0.333	0.256	0.206	0.683	0.581	0.686	0.374	3.946	11
S₂	0.475	0.982	0.780	0.810	0.163	0.095	0.913	0.460	0.671	0.321	5.844	3
S₃	1.000	0.664	0.745	1.000	0.000	0.000	1.000	0.701	0.521	0.706	6.337	2

处理Treatment

隶属函数值Membership function value

总得分Total Points

位次Precedence

株高Plant height

全株干质量Wholeplant dry weight

根冠比Root shoot ratio

可溶性蛋白含量Soluble protein content

可溶性糖含量Soluble sugar content

质膜透性Membrane permeability

MDA含量MDA content

H₂O₂含量H₂O₂ content

SOD活性SOD activity

POD活性POD activity

CK₁

0.000

0.618

1.000

0.000

0.493

1.000

0.543

1.000

0.000

4.036

CK₂

0.394

0.000

0.524

0.700

0.319

0.268

0.387

0.587

0.557

4.354

A₁

0.471

0.350

0.745

0.048

1.000

0.430

0.000

0.446

0.978

0.403

4.871

A₂

0.226

0.777

0.515

0.214

0.837

0.465

0.928

0.468

0.932

0.505

5.392

A₃

0.068

0.600

0.210

0.357

0.384

0.453

0.635

0.712

0.460

4.738

C₁

0.439

0.300

0.190

0.405

0.897

0.510

0.075

0.000

0.616

1.000

4.432

C₂

0.629

0.073

0.680

0.762

0.827

0.316

0.181

0.115

1.000

0.641

5.224

C₃

0.919

1.000

0.510

0.810

0.631

0.712

0.566

0.441

0.721

0.646

6.956

S₁

0.412

0.364

0.225

0.333

0.256

0.206

0.683

0.581

0.686

0.374

3.946

S₂

0.475

0.982

0.780

0.810

0.163

0.095

0.913

0.460

0.671

0.321

5.844

S₃

1.000

0.664

0.745

1.000

0.000

1.000

0.701

0.521

0.706

6.337