CN114651622A - 植物磺肽素在提高农作物果实风味和营养品质中的应用 - Google Patents
植物磺肽素在提高农作物果实风味和营养品质中的应用 Download PDFInfo
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Abstract
本发明公开了植物磺肽素在提高农作物果实风味和营养品质以及在制备提高农作物果实风味和营养品质的制剂中的应用。本发明制剂以植物磺肽素为主要有效成分，通过植物磺肽素被植物体内PSKR1受体识别激活下游信号通路，调控代谢，并辅以海藻糖的作用，可显著提高果实品质。本发明制剂可在采前生长成熟阶段或在采后对农作物果实进行处理，适用性好，且环境友好、可在环境和生物体内降解，使用后也无残留。采用本发明制剂提高果实品质简单易行，成本较低，可显著提高农作物果实中可溶性糖和挥发性芳香物质、类胡萝卜素等风味物质和营养物质的含量，进而可以显著提高农作物果实的风味品质和营养品质。
Description
技术领域
本发明涉及农业生物技术领域，尤其涉及植物磺肽素在提高农作物果实风味和营养品质中的应用。
背景技术
随着生活水平的提高，人们对蔬菜水果等天然健康食品的需求发生了从“量”到“质”的变化。天然食品的安全和品质逐渐成为大众关注的重点。番茄是我国重要的蔬菜作物，在我国的栽培面积和总产量均位居世界首位，因其外形端正美观，饱满红亮，且鲜美多汁，带有独特的果香，广受消费者欢迎。番茄果实品质包括外观品质、风味品质、营养品质和加工贮藏品质，其中风味品质和营养品质是近年来市场关注的重点。
番茄风味品质主要由果实中的可溶性糖、挥发性化合物等物质所决定。番茄果实中的可溶性糖主要为葡萄糖、果糖和蔗糖，可溶性糖含量高的番茄往往更受消费者喜爱；挥发性芳香物质在番茄果实中的含量很少，约为2mg/kg，但同样是影响番茄风味的重要因素，研究发现正乙醛等芳香物质是构成番茄香气的主要成分。
番茄富含维生素，矿物元素和类胡萝卜素，具有很高的营养价值。类胡萝卜素作为强抗氧化剂，可以有效清除人体的氧自由基，减轻生物体的氧胁迫，是番茄营养品质中的重要指标之一。番茄中的类胡萝卜素主要为番茄红素、β-胡萝卜素和叶黄素，其中番茄红素是目前自然界中发现的最强抗氧化剂之一，可以抑制人体与衰老有关的疾病，减少引发癌症和心血管疾病的机率；其次，食用含β-胡萝卜素等维生素A原类胡萝卜素是人类摄取维生素A的唯一途径，维生素A具有维持骨骼、上皮组织、视力和粘膜上皮正常分泌等多种生理功能；另外，叶黄素则可以有效保护人类视网膜中的黄斑，是帮助眼睛发育的关键营养元素。越来越多的研究表明，经常摄入叶黄素可以减缓或预防视力退化、近视、视网膜病变等症状。
现在市场中番茄品质下滑，“番茄味”消失，而果实营养及风味较好的番茄品种往往无法兼顾抗性，生产上管理复杂。因此，寻找一种高效、安全绿色的果实品质提高方法至关重要。
植物激素如生长素、赤霉素、脱落酸和乙烯在调节果实生长发育过程中发挥重要作用。植物磺肽素(PSK)是一种新型“肽类激素”，最初在芦笋叶肉细胞悬浮培养的培养基中鉴定分离。PSK由约80个氨基酸长的前体肽经酪氨酸残基的翻译后磺化和蛋白水解加工而产生(Yang等,“Oryza sativa PSK gene encodes a precursor of phytosulfokine-α,asulfated peptide growth factor found in plants”,Proceedings of the NationalAcademy of Sciences of the United States of America,1999,96:13560-13565)。剪切修饰后成熟的PSK为磺化的五肽([H-Tyr(SO3H)-Ile-Tyr(SO3H)-Thr-Gln-OH])，结构式如下所示。
PSK在细胞增殖和分化、分生组织形成、自交不亲和性、器官脱落、气孔运动、抗性反应等过程中均具有植物生长调节物质的属性(Matsubayashi等,“Peptide hormones inplants”,Annual Review of Plant Biology,2006,57:649-674)。Song等研究发现外源PSK处理可以提高枇杷果实抗冷性(Song等,“A cold-induced phytosulfokine peptide isrelated to the improvement of loquat fruit chilling tolerance”,Foodchemistry,2017,232:434-442)；公开号为CN 105900983A和CN107114369A的中国专利文献分别公开了植物磺肽素-α在提高植物黄化曲叶病毒病抗性和提高植物灰霉病抗性中的应用；但当前关于PSK调控果实生长发育及品质的研究较少。
发明内容
本发明通过实验发现，植物磺肽素(PSK)可以激活农作物果实生长发育相关转录因子的信号途径，进一步调控代谢，提高果实品质。
基于以上发现，本发明提供了植物磺肽素在提高农作物果实风味和营养品质中的应用。
PSK是由约80个氨基酸长的前体肽经酪氨酸残基的翻译后磺化和蛋白水解加工而产生的五肽；可通过被其受体PSKR1(Solyc01g008140)识别，从而激活果实生长发育相关转录因子SlC2H2(Solyc07g053570)、SlTCP(Solyc02g089020)、SlHB(Solyc03g120910)信号途径，进一步调控代谢，提高果实品质。
本发明还提供了植物磺肽素在制备提高农作物果实风味和营养品质的制剂中的应用。
所述农作物果实包括番茄。
本发明还提供了一种提高农作物果实风味和营养品质的制剂，所述制剂中包含植物磺肽素、海藻糖和表面活性剂，其中，植物磺肽素、海藻糖和表面活性剂的配比为6.7～10.2g：30～40g：20～30mL。
所述表面活性剂可采用有机硅、吐温60或Silwet-L77中的一种。
所述制剂在使用过程中，将植物磺肽素、海藻糖加水配制为含6.7～10.2g/L植物磺肽素和30～40g/L海藻糖的混合溶液，混合溶液先后加入水和表面活性剂得到稀释液；稀释液中，植物磺肽素的浓度为67.7～101.6mg/L，海藻糖的浓度为0.3～0.4g/L，表面活性剂的体积百分数为0.02％～0.03％。
所述制剂以水为分散介质，植物磺肽素为主要有效成分，其中，海藻糖的加入可以起到辅助作用，使果实成熟后保持较高的风味品质。
表面活性剂可使制剂稀释液在果实表面的延展性和渗透性能显著增强，有效减少制剂稀释液喷施后随风漂移的情况，提高制剂稀释液的抗雨水冲刷能力和药效，同时有利于浸果时有效成分的渗透，减少制剂稀释液的用量并延长有效期。
优选的，所述表面活性剂为有机硅，有机硅表面活性剂价格更为低廉，且在提高制剂稀释液的延展性、降低制剂稀释液表面张力上效果更为显著，使有效成分更易被植株吸收。
本发明还提供了一种提高农作物果实风味和营养品质的方法，将所述制剂配制的稀释液采前喷施于农作物果实上或对采后农作物果实进行浸果处理。该方法可在果实采前生长成熟阶段或在果实采后进行处理，适用性好。
制剂稀释液的使用浓度、次数，尤其是PSK的浓度可根据植物的苗龄、具体生长情况、农作物生长环境以及农作物果实的成熟程度来确定。
进一步优选的，利用所述制剂配制的稀释液对绿熟期番茄进行采前喷施或采后浸果处理。绿熟期的番茄果实已充分长成，果实表面绿色且不再膨大，物质积累过程已完成，可在植株上或贮藏中完成后熟过程。在绿熟期对番茄进行上述处理，易于实施且效果显著。
与现有技术相比，本发明的有益效果在于：
(1)本发明对于已知化合物PSK发掘了新的提高农作物果实品质的用途，开拓了一个新的应用领域。
(2)本发明制剂环境友好、可在环境和生物体内降解，使用后也无残留，对人畜和环境无毒害。
(3)本发明制剂以PSK为主要有效成分，通过被受体PSKR1(Solyc01g008140)识别激活果实生长发育相关转录因子SlC2H2(Solyc07g053570)、SlTCP(Solyc02g089020)、SlHB(Solyc03g120910)信号途径，并辅以海藻糖的作用，可显著增强果实品质，提高农作物生产经济效率。
(4)本发明方法可在果实采前生长成熟阶段或在果实采后进行处理，适用性好。
(5)采用本发明制剂提高果实品质简单易行，成本较低，可显著提高农作物果实中可溶性糖和挥发性芳香物质、类胡萝卜素等风味物质和营养物质的含量，进而可以显著提高农作物果实的风味和营养品质。
附图说明
图1为实施例1中制剂稀释液处理实验组和对照组中番茄可溶性糖含量的对照图，其中，A为葡萄糖，B为果糖，C为蔗糖，a，b代表5％水平上的差异显著。
图2为实施例1中制剂稀释液处理实验组和对照组中番茄挥发性物质正己醛含量的对照图，a，b代表5％水平上的差异显著。
图3为实施例1中制剂稀释液处理实验组和对照组中番茄类胡萝卜素含量的对照图，其中，A为番茄红素，B为β胡萝卜素，C为叶黄素，a，b代表5％水平上的差异显著。
图4为实施例1中制剂稀释液处理实验组和对照组处理7天后的番茄图片。
图5为实施例2中制剂稀释液处理实验组和对照组中番茄可溶性糖含量的对照图，其中，A为葡萄糖，B为果糖，C为蔗糖，a，b代表5％水平上的差异显著。
图6为实施例2中制剂稀释液处理实验组和对照组中番茄挥发性物质正己醛含量的对照图，a，b代表5％水平上的差异显著。
图7为实施例2中制剂稀释液处理实验组和对照组中番茄类胡萝卜素含量的对照图，其中，A为番茄红素，B为β胡萝卜素，C为叶黄素，a，b代表5％水平上的差异显著。
图8为实施例2中制剂稀释液处理实验组和对照组处理7天后的番茄图片。
图9为实施例3中pskr#1基因编辑突变体突变类型以及pskr#1基因编辑突变体与普通番茄的绿熟期果实生长发育相关转录因子相对表达量对比图，其中，A为pskr#1基因编辑突变体突变类型，B为SlC2H2，C为SlTCP，D为SlHB，a，b代表5％水平上的差异显著。
图10为实施例3中pskr#1基因编辑突变体与普通番茄花后49天可溶性糖含量的对照图，其中，A为葡萄糖，B为果糖，C为蔗糖，a，b代表5％水平上的差异显著。
图11为实施例3中pskr#1基因编辑突变体与普通番茄花后49天挥发性物质正己醛含量的对照图，a，b代表5％水平上的差异显著。
图12为实施例3中pskr#1基因编辑突变体与普通番茄花后49天番茄类胡萝卜素含量的对照图，其中，A为番茄红素，B为β胡萝卜素，C为叶黄素，a，b代表5％水平上的差异显著。
具体实施方式
下面结合附图与实施例，进一步阐明本发明。应理解，这些实施例仅用于说明本发明，而不用于限制本发明的范围。
实施例1制剂稀释液采前喷施处理绿熟期番茄
本实施例中，制剂的成分为：0.85g植物磺肽素(药品购自杭州德晟医药有限公司)，3.78g海藻糖(沪试)和2.5mL有机硅。
一、制剂稀释液的制备
将0.85g植物磺肽素、3.78g海藻糖缓慢加入100mL的水，搅拌直至充分溶解得到混合溶液；往100mL混合溶液中先后加入9.9L水和2.5mL有机硅混合均匀成稀释液后使用。
二、所述制剂稀释液的施用和果实品质指标的检测
将步骤(一)的制剂稀释液倒入喷壶，在傍晚均匀喷施于绿熟期果实(果实表面绿色且不再膨大)表面，直至果实表面湿润为止，持续喷施2天，以喷施纯水的番茄植株作为对照。将上述处理的所有植株置于温度25℃/21℃(昼/夜)，光周期为12h/12h(昼/夜)，光照强度为200μmol·m-2·s-1，相对湿度80％的环境条件中培养。
每升制剂稀释液大约能处理80棵番茄。
处理后7天分别取制剂稀释液处理组和对照组30个果实样品，10个为1组，测定其可溶性糖包括葡萄糖、果糖、蔗糖，挥发性芳香物质包括正己醛，以及类胡萝卜素包括番茄红素、β-胡萝卜素、叶黄素的含量。
可溶性糖(葡萄糖、果糖、蔗糖)测定方法如下：
称取0.1g果实冻样粉末，加入1.0mL双蒸水，涡旋，80℃恒温水浴30min后12000×g离心10min，收集上清液；再次向沉淀中加入1.0mL双蒸水，重复上述操作并将两次上清液混合。分别取2μL和100μL上清液，用80％色谱级乙腈定容至1mL，各取200μL溶液用于测定蔗糖以及葡萄糖、果糖的含量。参考Zhang等(Zhang等,“Postharvest responses of Chinesebayberry fruit”,Postharvest Biology and Technology,2005,37,241-251)通过高效液相色谱的方法测定三种糖的含量。葡萄糖、果糖、蔗糖标准品均购自Sigma-Aldrich公司。
挥发性物质正己醛测定方法如下：
称取5g果肉组织，液氮磨成粉末后转移至装有5mL饱和氯化钠溶液的15mL小瓶中。在密封小瓶之前，加入20μL 2-辛醇(0.8mg mL-1)作为内标并涡旋10秒。参考Li等方法(Li等，“Roles of RIN and ethylene in tomato fruit ripening and ripening-associated traits”,New Phytologist,2020,226(2):460-475)采用顶空固相微萃取结合气相质谱技术检测番茄果实挥发性物质正己醛含量的变化。正己醛标准品购自Sigma-Aldrich公司。
类胡萝卜素(番茄红素、β-胡萝卜素、叶黄素)测定方法如下：
称取番茄果实冻干粉0.1g于2mL离心管中，依次加入350μL甲醇、700μL氯仿和350μL超纯水，充分涡旋，10000rpm离心10min，收集氯仿相。向剩余液体中再次加入700μL氯仿，混匀并10000rpm离心10min，收集氯仿相并与之前的氯仿相合并。合并后的氯仿相用旋蒸仪30℃蒸干，加入350μL 6％KOH甲醇溶液，60℃水浴锅暗中放置30min，进行皂化反应，后加入700μL超纯水进行萃取，直至水相颜色为无色。萃取后的氯仿相用氮气吹干，溶于100μL色谱级乙酸乙酯中，10000rpm离心10min，取50μL上清液参考Zheng等方法(Zheng等,“Naturalvariation in CCD4 promoter underpins species-specific evolution of redcoloration in citrus peel”,Molecular Plant,2019,12(9),1294-1307)进行高效液相色谱分析。番茄红素、β-胡萝卜素和叶黄素标准品均购于北京索莱宝公司。
制剂稀释液处理组和对照组果实品质指标的检测结果如图1-4所示，上述制剂稀释液处理实验组可溶性糖包括葡萄糖(图1中的A)、果糖(图1中的B)、蔗糖的含量(图1中的C)，挥发性物质正己醛的含量(图2)，以及类胡萝卜素包括番茄红素(图3中的A)、β-胡萝卜素(图3中的B)、叶黄素(图3中的C)的含量较对照组均显著提高。如图4所示，制剂稀释液处理组的果实颜色更红。即制剂稀释液处理实验组的番茄果实风味和营养品质更好。
实施例2所述制剂对采后绿熟期番茄进行浸果处理
本实施例中，制剂的成分为：2.02g植物磺肽素(药品购自杭州德晟医药有限公司)，8g海藻糖(沪试)和6mL有机硅。
一、制剂稀释液的制备
将2.02g植物磺肽素、8g海藻糖缓慢加入200mL的水，搅拌直至充分溶解得到混合溶液；往200mL混合溶液中先后加入19.8L水和6mL有机硅混合均匀成稀释液后使用。
二、所述制剂的施用和果实品质指标的检测
将步骤(一)的2L制剂稀释液倒入长20cm宽10cm高10cm容器，挑选成熟度均一，无机械伤，无病虫害，大小相对一致的绿熟期番茄放入其中，盖好盖子，使果实整个浸入溶液中，静置3小时后取出，常温晾干。连续浸果处理2天，以同样方法清水处理的果实作为对照。上述所有果实处理后于温度25℃/21℃(昼/夜)，光周期为12h/12h(昼/夜)，光照强度为200μmol·m-2·s-1，相对湿度80％的环境条件中静置。
2L制剂稀释液处理的果实量可根据容器和果实大小决定，需保证每个果实都完全浸泡在制剂中。
处理9天后分别取30个制剂稀释液处理组和对照组果实样品，10个为一组，测定其可溶性糖包括葡萄糖、果糖、蔗糖，挥发性芳香物质包括正己醛，以及类胡萝卜素包括番茄红素、β-胡萝卜素、叶黄素的含量。测量方法同实施例1。
制剂稀释液处理组和对照组果实品质指标的检测结果如图5-8所示，上述制剂稀释液处理实验组可溶性糖包括葡萄糖(图5中的A)、果糖(图5中的B)、蔗糖的含量(图5中的C)，挥发性物质正己醛的含量(图6)，以及类胡萝卜素包括番茄红素(图7中的A)、β-胡萝卜素(图7中的B)、叶黄素(图7中的C)的含量较对照组均显著提高。如图8所示，制剂稀释液处理组的果实颜色更红。即制剂稀释液处理实验组的番茄果实风味和营养品质更好。
实施例3番茄pskr#1基因编辑突变体果实品质变化
(1)pskr#1基因编辑突变体与未经过基因编辑的普通番茄相比，突变体在sgRNA位置发生碱基缺失，分别取9个pskr#1基因编辑突变体(突变类型如图9中的A所示)和对照番茄绿熟期(花后43天)果实样品；3个为一组，利用Omega Bio-Tek公司的RNA提取试剂盒(货号R6827)和Invitrogen公司生产的Superscript II试剂盒进行RNA提取纯化及反转录，后利用美国Applied Biosystems公司生产的Step ONE Plus Real-Time PCR System仪器及该公司的SYBR RT-PCR Kit荧光染料试剂盒测定样品中果实生长发育相关转录因子SlC2H2、SlTCP、SlHB的相对表达量。
结果如图9所示，SlC2H2(图9中的B)、SlTCP(图9中的C)、SlHB(图9中的D)在普通番茄中转录水平的相对表达量均高于pskr#1基因编辑突变体。
(2)分别取9个pskr#1基因编辑突变体和对照番茄花后49天果实样品，3个为一组，测定其可溶性糖包括葡萄糖、果糖、蔗糖，挥发性芳香物质包括正己醇，以及类胡萝卜素包括番茄红素、β-胡萝卜素、叶黄素的含量。测量方法同实施例1。
pskr#1基因编辑突变体与未经过基因编辑的普通番茄品质指标的检测结果如图10-12所示，pskr#1基因编辑突变体果实中可溶性糖包括葡萄糖(图10中的A)、果糖(图10中的B)、蔗糖(图10中的C)，挥发性物质包括正己醛(图11)，以及类胡萝卜素包括番茄红素(图12中的A)、β-胡萝卜素(图12中的B)、叶黄素(图12中的C)的含量均显著下降。
以上所述的实施例对本发明的技术方案进行了详细说明，应理解的是以上所述的仅为本发明的具体实施例，并不用于限制本发明，凡在本发明的原则范围内所做的任何修改、补充或类似方式替代等，均应包含在本发明的保护范围之内。
序列表
<110> 浙江大学
<120> 植物磺肽素在提高农作物果实风味和营养品质中的应用
<141> 2022-02-21
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 19
<212> DNA
<213> Artificial Sequence
<400> 1
atgggttccg tggttagtt 19
<210> 2
<211> 20
<212> DNA
<213> Lycopersicon esculentum
<400> 2
atgggttccg tgggttagtt 20
Claims (8)
1.植物磺肽素在提高农作物果实风味和营养品质中的应用。
2.植物磺肽素在制备提高农作物果实风味和营养品质的制剂中的应用。
3.根据权利要求1或2所述应用，其特征在于，所述农作物果实包括番茄。
4.一种提高农作物果实风味和营养品质的制剂，其特征在于，所述制剂中包含植物磺肽素、海藻糖和表面活性剂，其中，植物磺肽素、海藻糖和表面活性剂的配比为6.7～10.2g：30～40g：20～30mL。
5.根据权利要求4所述制剂，其特征在于，所述表面活性剂为有机硅、吐温60或Silwet-L77中的一种。
6.根据权利要求4或5所述制剂，其特征在于，将植物磺肽素、海藻糖加水配制为含6.7～10.2g/L植物磺肽素和30～40g/L海藻糖的混合溶液，混合溶液先后加入水和表面活性剂得到稀释液；稀释液中，植物磺肽素的浓度为67.7～101.6mg/L，海藻糖的浓度为0.3～0.4g/L，表面活性剂的体积百分数为0.02％～0.03％。
7.一种提高农作物果实风味和营养品质的方法，其特征在于，将权利要求6所述制剂配制的稀释液采前喷施于农作物果实上或对采后农作物果实进行浸果处理。
8.根据权利要求7所述方法，其特征在于，利用所述制剂配制的稀释液对绿熟期番茄进行采前喷施或采后浸果处理。
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