褐藻寡糖与虾青素对盐胁迫下生菜生长的缓解效果及机制

doi:10.15933/j.cnki.1004-3268.2026.01.011

摘要/Abstract

摘要： 为了研究不同添加量褐藻寡糖和虾青素对盐胁迫下生菜生长、根系形态、养分吸收及抗逆性的影响，为逆境下生菜种植提供科学依据与实践经验，以生菜品种大速生菜为供试材料进行水培试验，用NaCl将营养液调节为含盐120 mmol/L溶液模拟盐胁迫环境（S），分别设置2种添加生物刺激素处理：褐藻寡糖添加量0、0.005%、0.010%、0.015%、0.020%、0.025%，即S-B0、S-B1、S-B2、S-B3、S-B4、S-B5；虾青素添加量0、0.05%、0.10%、0.15%、0.20%、0.25%，即S-A0、S-A1、S-A2、S-A3、S-A4、S-A5。同时以无盐营养液为对照（CK），分析生菜植株生物量、长势、养分吸收量、根系形态、叶片渗透调节物质以及抗氧化关键酶活性等指标。结果表明，与S-B0处理相比，添加褐藻寡糖可促进根系生物量累积，改善叶片叶绿素相对含量并增加叶片数，S-B1—S-B5 处理植株氮、磷、钾吸收量分别提高47.32%~89.09%（p<0.05）、4.32%~20.38%和2.29%~24.79%，总根长、根表面积及根体积分别增加58.83%~139.04%、53.57%~92.33%和36.07%~72.50%（p<0.05）。根系分级评价显示，添加褐藻寡糖主要促进Ⅰ~Ⅲ级根系生长，同时根平均直径与根冠比分别提高11.54%~34.62%和18.60%~32.56%。与S-A0处理相比，添加虾青素可促进根系生物量累积，改善叶片SPAD值并增加叶片数，S-A1—S-A5处理植株氮、磷、钾吸收量分别提高32.11%~145.95%、10.07%~66.91%和25.42%~64.51%，总根长、根表面积及根体积分别增加26.57%~128.97%、26.59%~129.01%和26.67%~193.33%。根系分级评价显示，添加虾青素主要促进Ⅰ~Ⅱ级根系生长，同时根平均直径提高10.00%~33.33%，各处理根冠比差异不显著。盐胁迫下添加褐藻寡糖和虾青素均可显著增加叶片脯氨酸（Pro）含量以调节渗透平衡，同时提高超氧化物歧化酶（SOD）、过氧化物酶（POD）与过氧化氢酶（CAT）活性并降低丙二醛（MDA）的累积，减少细胞氧化损伤。冗余分析表明，添加褐藻寡糖主要通过提高SOD活性缓解盐胁迫对生菜生长的损伤，而虾青素驱动生菜抗逆生长的主要因子为SOD活性和Pro含量。依据隶属函数法对生菜各指标进行评价，各褐藻寡糖处理中S-B3处理得分最高，虾青素处理中S-A2处理得分最高。将褐藻寡糖或虾青素应用于水培蔬菜生产中均可促进生菜根系生长与养分吸收，有效改善叶片细胞渗透平衡，同时提高抗氧化保护酶活性以缓解盐胁迫对叶菜造成的负面影响；增强生菜抗盐性的褐藻寡糖与虾青素的最适添加量分别为0.015%和0.10%。

关键词: 生菜, 盐胁迫, 褐藻寡糖, 虾青素, 生物量, 根系形态, 抗氧化

Abstract: This study investigated the effects of different addition levels of alginate oligosaccharides and astaxanthin separately on the growth，root morphology，nutrient uptake，and stress resistance of lettuce under salt stress to provide scientific basis and practical experience for lettuce cultivation under adverse conditions.A hydroponic experiment was conducted using Dasu lettuce. We adjusted the nutrient solution to a salt concentration of 120 mmol/L using NaCl solid to simulate the salt stress environment（S）.Two types of treatments were set up：alginate oligosaccharides were added at concentrations of 0，0.005%，0.010%，0.015%，0.020% and 0.025%，corresponding to S‐B0，S‐B1，S‐B2，S‐B3，S‐B4 and S‐B5，respectively；astaxanthin was added at concentrations of 0，0.05%，0.10%，0.15%，0.20% and 0.25%，corresponding to S‐A0，S‐A1，S‐A2，S‐A3，S‐A4 and S‐A5，respectively. Meanwhile，the nutrient solution without NaCl added was used as the control treatment（CK），with three replicates pertreatment.We analyzed various indicators of lettuce plants，including biomass，growth status，nutrient uptake，root morphology，leaf osmotic regulators and the activity of key antioxidant enzymes.Our experimental results indicated that alginate oligosaccharides promoted the accumulation of root biomass，and the increase of leaf SPAD value and the number of leaves compared to S‐B0 treatment.The nitrogen，phosphorus and potassium uptake in S‐B1 to S‐B5 treatments increased by 47.32% to 89.09%（p<0.05），4.32% to 20.38% and 2.29% to 24.79%，respectively.The total root length，root surface area and root volume increased by 58. 83% to 139.04%，53.57% to 92.33% and 36.07% to 72.50%（p<0.05），respectively.Root classification evaluation showed that the addition of alginate oligosaccharides primarily promoted the growth of roots at levels Ⅰ to Ⅲ.At the same time，the average root diameter and root‐to‐shoot ratio increased by 11.54% to 34.62% and 18.60% to 32.56%，respectively.Compared to S‐A0 treatment，the astaxanthin promoted the accumulation of root biomass，leaf SPAD values and the number of leaves.The nitrogen，phosphorus，and potassium uptake in S‐A1 to S‐A5 treatments increased by 32.11% to 145.95%，10.07% to 66.91% and 25.42% to 64.51%，respectively.The total root length，root surface area and root volume increased by 26.57% to 128.97%，26.59% to 129.01% and 26.67% to 193.33%，respectively.Root classification evaluation showed that the addition of astaxanthin primarily promoted the growth of roots at levels Ⅰ to Ⅱ. At the same time，the average root diameter increased by 10.00% to 33.33%，and there were no significant differences in the root‐to‐shoot ratio among the treatments.Under salt stress，the addition of alginate oligosaccharides and astaxanthin significantly increased the proline（Pro）content in the leaves，helping to regulate osmotic balance. At the same time，both treatments enhanced the activities of superoxide dismutase（SOD），peroxidase（POD）and catalase（CAT）as well as reduced the accumulation of malondialdehyde（MDA），thereby decreasing cellular oxidative damage.Redundancy analysis indicated that the addition of alginate oligosaccharides mainly enhanced the activity of SOD，which helped alleviate the damage caused by salt stress to lettuce growth. On the other hand，astaxanthin promoted lettuce’s stress tolerance primarily through the combined effects of SOD and Pro，driving its adaptive growth under stress conditions. According to the membership function method，which was used to evaluate various indicators of lettuce，the highest score in alginate oligosaccharide treatments was observed in S‐B3 treatment，while in the astaxanthin treatments，the highest score was found in S‐A2 treatment.The application of alginate oligosaccharides or astaxanthin in hydroponic vegetable production can both promote the growth of lettuce roots and enhance nutrient absorption.These treatments effectively improve the osmotic balance of leaf cells，while also increase the activity of antioxidant protective enzymes to mitigate the negative effects of salt stress on leafy vegetables.The optimal addition rates for improving the salt tolerance of lettuce are 0.015% for alginate oligosaccharides and 0. 10% for astaxanthin.

Key words: Lettuce, Salt stress, Alginate oligosaccharides, Astaxanthin, Biomass, Root morphology, Antioxidation

中图分类号:

S636.2

李喜凤, 胥子航, 吴鑫雨, 苑征征, 陈玉凤, 郭景丽. 褐藻寡糖与虾青素对盐胁迫下生菜生长的缓解效果及机制[J]. 河南农业科学, 2026, 55(1): 118-131.

LI Xifeng, XU Zihang, WU Xinyu, YUAN Zhengzheng, CHEN Yufeng, GUO Jingli. Alleviation Effects and Mechanisms of Alginate Oligosaccharides and Astaxanthin on Lettuce Growth under Salt Stress[J]. Journal of Henan Agricultural Sciences, 2026, 55(1): 118-131.

图/表 12

参考文献

［1］ZHAO W，ZHOU Q，TIAN Z Z，et al. Apply biochar to ameliorate soda saline‐alkali land，improve soil function and increase corn nutrient availability in the Songnen Plain［J］. Science of the Total Environment，2020，722：137428.
［2］唐茜，王佳丽，李文强. 胞外多糖对盐胁迫下荆芥种子萌发、幼苗生长及生理特性的影响［J］. 河南农业科学，2024，53（10）：96‐105.
TANG Q，WANG J L，LI W Q. Effects of extracellular polysaccharides on seed germination，seedling growth and physiological characteristics of Schizonepeta tenuifolia under salt stress［J］.Journal of Henan Agricultural Sciences，2024，53（10）：96‐105.
［3］刘梅，郑青松，刘兆普，等. 盐胁迫下氮素形态对油菜和水稻幼苗离子运输和分布的影响［J］.植物营养与肥料学报，2015，21（1）：181‐189.
LIU M，ZHENG Q S，LIU Z P，et al. Effects of nitrogen forms on transport and accumulation of ions in canola（B.napus L.）and rice（Oryza sativa L.）under saline stress［J］. Journal of Plant Nutrition and Fertilizer，2015，21（1）：181‐189.

［4］陈选阳，张招娟，郑佳伟，等.水培对叶菜型甘薯茎尖营养品质与硝酸盐含量的影响［J］. 中国农业科学，2013，46（17）：3736‐3742.

CHEN X Y，ZHANG Z J，ZHENG J W，et al. Effects of hydroponics on nutrient components and nitrate contents in tips of leaf vegetable sweet potato［J］.Scientia Agricultura Sinica，2013，46（17）：3736‐3742.

［5］FARRUGGIA D，TORTORICI N，IACUZZI N，et al.Biostimulants improve plant performance of rosemary growth in agricultural organic system［J］. Agronomy，2024，14（1）：158.

［6］李炳言，宋大利，王秀斌，等.不同生物刺激素对玉米生长及土壤微生物群落结构的影响［J］.植物营养与肥料学报，2023，29（11）：2172‐2180.

LI B Y，SONG D L，WANG X B，et al.Effects of biostimulants on maize growth and soil microbial community structure［J］.Journal of Plant Nutrition and Fertilizers，2023，29（11）：2172‐2180.

［7］白炬，刘晓林，李申，等.污泥热碱液对干旱胁迫下小青菜生长的缓解机制［J］.干旱区研究，2024，41（1）：80‐91.

BAI J，LIU X L，LI S，et al. Mechanism of sludge alkaline thermal hydrolysis liquid on the growth of Brassica chinensis under drought stress［J］.Arid Zone Research，2024，41（1）：80‐91.

［8］吴娟，施国新，夏海威，等. 外源钙对汞胁迫下菹草（Potamogeton crispus L.）叶片抗氧化系统及脯氨酸代谢的调节效应［J］.生态学杂志，2014，33（2）：380‐387.

WU J，SHI G X，XIA H W，et al.Effects of exogenous calcium on antioxidant system and proline metabolism of Potamogeton crispus L.leaves under mercury stress ［J］.Chinese Journal of Ecology，2014，33（2）：380‐387.

［9］刘琛，丁能飞，傅庆林，等. 盐胁迫对3种蔬菜幼苗抗氧化酶活性的影响［J］.安徽农业科学，2010，38（1）：115‐116.

LIU C，DING N F，FU Q L，et al. Effect of the salt stress on the anti‐oxidative enzyme in seedlings of three vegetable species［J］. Journal of Anhui Agricultural Sciences，2010，38（1）：115‐116.

［10］王语，张渝鹏，朱冠亚，等. 局部供氮对干旱胁迫下玉米苗期生长发育和水氮利用的影响［J］.中国农业科学，2024，57（5）：919‐934.

WANG Y，ZHANG Y P，ZHU G Y，et al. Effects of localized nitrogen supply on plant growth and water and nitrogen use efficiencies of maize seedling under drought stress［J］. Scientia Agricultura Sinica，2024，57（5）：919‐934.

［11］李雪婷，任昊，王洪章，等.盐胁迫对不同耐盐型玉米品种叶片光合性能和干物质积累与分配的影响［J］.作物学报，2025，51（4）：1091‐1101.

LI X T，REN H，WANG H Z，et al.Effects of salt stress on photosynthetic performance and dry matter accumulation and distribution in leaves of different salt‐tolerant maize varieties［J］. Acta Agronomica Sinica，2025，51（4）：1091‐1101.

［12］杨娅坤，赵艳，李亮锴，等. 盐胁迫对水稻幼苗生长发育及生理特性的影响［J］.分子植物育种，2024，22（23）：7803‐7810.

YANG Y K，ZHAO Y，LI L K，et al. Effects of salt stress on the growth，development and physiological characteristics of rice seedlings［J］.Molecular Plant Breeding，2024，22（23）：7803‐7810.

［13］ZHANG M M，SONG D P，PU X，et al. Effect of different straw returning measures on resource use efficiency and spring maize yield under a plastic film mulch system［J］. European Journal of Agronomy，2022，134：126461.

［14］于晟玥，牛银星，王泽平，等.黄腐酸添加量对低氮胁迫下小麦生长和根系形态的影响［J］.植物营养与肥料学报，2023，29（2）：323‐333.

YU S Y，NIU Y X，WANG Z P，et al. Effects of fulvic acid addition rate on wheat growth and root morphology under low nitrogen stress［J］.Journal of Plant Nutrition and Fertilizers，2023，29（2）：323‐333.

［15］王学江，李峰，迟艳，等. 褐藻寡糖对蕹菜生长量及生长速率的影响［J］. 分子植物育种，2022，20（14）：4851‐4857.

WANG X J，LI F，CHI Y，et al. The effect of brown algae oligosaccharides on the growth and growth rate of water spinach［J］. Molecular Plant Breeding，2022，20（14）：4851‐4857.

［16］刘航.褐藻胶寡糖提高小麦抗旱能力的初步研究［J］. 天然产物研究与开发，2015，27（9）：1520‐1525.

LIU H. Alginate oligosaccharides enhanced wheat resistant ability to drought stress［J］. Natural Product Research and Development，2015，27（9）：1520‐1525.

［17］MA L J，LI X M，BU N，et al. An alginate‐derived oligosaccharide enhanced wheat tolerance to cadmium stress［J］. Plant Growth Regulation，2010，62（1）：71‐76.

［18］苗丽青，马旭辉，李素贞，等. 虾青素的生物合成与产业化应用［J］. 中国农业科技导报，2023，25（3）：21‐29.

MIAO L Q，MA X H，LI S Z，et al. Biosynthesis and industrial application of astaxanthin［J］. Journal of Agricultural Science and Technology，2023，25（3）：21‐29.

［19］WANG Y Y，CHEN L P，ZHAO J R，et al. Astaxanthin esters as functional food：A review of their nutrition，phytochemical structure，biological features，and food industry prospects［J］. Journal of Agricultural and Food Chemistry，2024，72（24）：13467‐13475.

［20］杨梦煊，王宝杰，刘梅，等. 盐度胁迫下凡纳滨对虾体内虾青素在着色与抗氧化的资源权衡［J］.海洋科学，2023，47（10）：32‐42.

YANG M X，WANG B J，LIU M，et al. Resource tradeoff between coloring and antioxidation of astaxanthin in Litopenaeus vannamei under salinity stress［J］. Marine Sciences，2023，47（10）：32‐42.

［21］高俊凤. 植物生理学实验指导［M］. 北京：高等教育出版社，2006.

GAO J F. Experimental guidance of plant physiology ［M］. Beijing：Higher Education Press，2006.

［22］金江，黄依妮，刘鸿宇，等. 减施氮肥及玉米间作绿肥对混合饲草产量及品质的影响［J］. 中国土壤与肥料，2024（9）：61‐69.
JIN J，HUANG Y N，LIU H Y，et al. Effects of reduced nitrogen fertilizer and green manure intercropping of maize on yield and quality of mixed forage［J］. Soil and Fertilizer Sciences in China，2024（9）：61‐69.

［23］陈晨，杨凯波，王奎萍. 不同胁迫对植物生长发育的影响及植物抗胁迫机制综述［J］. 江苏农业科学，2024，52（19）：15‐24.
CHEN C，YANG K B，WANG K P. Effects of different stresses on plant growth and mechanisms of plant resistance to stresses： A review ［J］. Jiangsu Agricultural Sciences，2024，52（19）：15‐24.

［24］朱芝宜，李培根，陆晓彤，等. 假单胞菌与褐藻寡糖协同提高梨果实产量及品质［J］. 植物营养与肥料学报，2024，30（10）：1964‐1973.
ZHU Z Y，LI P G，LU X T，et al. Synergistic effects of Pseudomonas CD1 and brown algal oligosaccharides on pear fruit yield and quality［J］. Journal of Plant Nutrition and Fertilizers，2024，30（10）：1964‐1973.

［25］ZHANG J X，MENG Q X，YANG Z P，et al. Humic acid promotes the growth of switchgrass under salt stress by improving photosynthetic function［J］.Agronomy，2024，14（5）：1079.

［26］AZAB E S，ALSHALLASH K S，ALQAHTANI M M，et al. Physiological，anatomical，and agronomic responses of Cucurbita pepo to exogenously sprayed potassium silicate at different concentrations under varying water regimes［J］. Agronomy，2022，12（9）：2155.

［27］LYNCH J. Root architecture and plant productivity［J］.Plant Physiology，1995，109（1）：7‐13.

［28］许猛，袁亮，李伟，等. 复合氨基酸肥料增效剂对NaCl 胁迫下小白菜种子萌发和苗期生长的影响［J］.植物营养与肥料学报，2018，24（4）：992‐1000.
XU M，YUAN L，LI W，et al. Effects of a fertilizer synergist containing compound amino acids on seed germination and seedling growth of pakchoi under NaCl stress［J］.Journal of Plant Nutrition and Fertilizers，2018，24（4）：992‐1000.

［29］LI Z M，DUAN S P，OUYANG X，et al. Coupled soil moisture management and alginate oligosaccharide strategies enhance citrus orchard production，water and potassium use efficiency by improving the rhizosphere soil environment［J］. Agricultural Water Management，2024，297：108828.

［30］杨凤军，李天来，臧忠婧，等. 外源钙施用时期对缓解盐胁迫番茄幼苗伤害的作用［J］. 中国农业科学，2010，43（6）：1181‐1188.
YANG F J，LI T L，ZANG Z J，et al. Effects of timing of exogenous calcium application on the alleviation of salt stress in the tomato seedlings［J］.Scientia Agricultura Sinica，2010，43（6）：1181‐1188.
［31］YANG L，QIAO X，LIU J，et al. Preparation，characterization and antioxidant activity of astaxanthin esters with different molecular structures［J］. Journal of the Science of Food and Agriculture，2021，101（6）：2576‐2583.

［32］GOTO S，KOGURE K，ABE K，et al. Efficient radical trapping at the surface and inside the phospholipid membrane is responsible for highly potent antiperoxidative activity of the carotenoid astaxanthin［J］. Biochimica et Biophysica Acta，2001，1512（2）：251‐258.
［33］王恒，曹佳茵，侯剑，等. 喷施褐藻寡糖对茶树抗旱性的影响［J］. 山东农业科学，2023，55（12）：63‐70.

WANG H，CAO J Y，HOU J，et al. Effect of spraying algal oligosaccharide on drought resistance of tea plant［J］. Shandong Agricultural Sciences，2023，55（12）：63‐70.
［34］石锦，李仁，王晋芳，等. 柳枝稷质膜型水通道蛋白基因PvPIP1的克隆和序列分析［J］. 草地学报，2014，22（4）：840‐846.
SHI J，LI R，WANG J F，et al. Cloning and sequence analysis of the plasma membrane aquaporins gene PvPIP1 in switchgrass［J］. Acta Agrestia Sinica，2014，22（4）：840‐846.

[1]	苟瑞丽, 舍杨梦斐, 方晶莹, 田浩天, 马国林, 田蕾, 罗成科. 不同耐盐碱类型水稻根系对盐碱胁迫的生理及分子响应差异[J]. 河南农业科学, 2025, 54(9): 34-42.
[2]	王琳, 师莹莹, 王光安. 鼓槌石斛GATA 基因家族的鉴定及其在盐胁迫下的功能分析[J]. 河南农业科学, 2025, 54(9): 72-83.
[3]	库朝锋, 王献伟, 吕玲燕, 张家庆, 刘洋, 宋伟宜, 张俊霞. 艾草粉对育肥猪生长性能、养分表观消化率、抗氧化功能及粪便微生物数量的影响[J]. 河南农业科学, 2025, 54(9): 141-148.
[4]	范小垒, 孟岭超, 李丹丹, 周强. 新型转光膜覆盖对温室切花月季生长及气孔特征的影响[J]. 河南农业科学, 2025, 54(7): 126-134.
[5]	田野, 杨培华, 阳婷倩, 刘小岑, 刘奕清, 王娇, 胡海骏, 蒋昕晨, 张万顺, 朱永兴. 外源纳米硅对盐胁迫下辣椒种子萌发的影响[J]. 河南农业科学, 2025, 54(6): 110-120.
[6]	邹泉, 邢伟明, 王若玎, 周文俊. 根施丛枝菌根真菌和独脚金内酯对盐碱胁迫下多年生黑麦草叶绿素荧光及抗氧化系统的影响[J]. 河南农业科学, 2025, 54(6): 72-83.
[7]	陈春, 种春斌, 路雷. 丹参WOX 家族成员鉴定及SmWOX8基因在盐胁迫下的功能分析[J]. 河南农业科学, 2025, 54(6): 43-54.
[8]	龙威, 王靓, 金羽琨, 刘佳遥, 魏尊苗, 程艳, 牟忠生. 苗期油莎豆在盐胁迫下的生理特性及转录组学分析[J]. 河南农业科学, 2025, 54(6): 11-20.
[9]	杨雯, 张静, 何卫凯, 周旭. 不同光照强度对4 种罗勒属植物叶绿素荧光和光合生理的影响[J]. 河南农业科学, 2025, 54(5): 113-123.
[10]	彭波, 马梦梅, 赵苹. 基于转录组的芍药WOX基因家族鉴定及PlWOX5在盐胁迫下的功能分析[J]. 河南农业科学, 2025, 54(4): 57-65.
[11]	梁亚男, 邬梦晞, 肖雍琴, 曾鲸津, 段益莉. 植物根际促生菌和褪黑素对大花金鸡菊抗氧化特性和砷吸收的影响[J]. 河南农业科学, 2025, 54(3): 87-98.
[12]	邓聪, 马璐, 汪青松, 付健, 王玉凤, 杨克军. 芽孢杆菌对盐碱胁迫下玉米种子萌发及生理生化特性的影响[J]. 河南农业科学, 2025, 54(3): 20-30.
[13]	彭超军, 华夏, 王松峰, 高崇, 董海滨, 胡琳. 干旱胁迫对不同小麦品种灌浆期旗叶光合及生理特性的影响[J]. 河南农业科学, 2025, 54(2): 40-47.
[14]	韩锐锋, 任荣魁, 董向阳, 胡泽彬, 郭方君, 方甜甜, 马文豪, 张书红. 不同类型肥料施用深度和施用量对冬小麦生长发育的影响[J]. 河南农业科学, 2025, 54(2): 30-39.
[15]	张朝冉, 彭漪, 杨凯丽, 袁宁鸽, 黎远金, 刘小娟, 吴慧丽. 基于光合有效吸收的水稻穗生物量遥感估测[J]. 河南农业科学, 2024, 53(6): 154-161.