采前调节剂处理对采后果实保鲜效应的研究进展

孙闽子1,2,3,张 珅1*,吴光斌1,陈发河1,倪 辉1,3,陈兴麟4,5*,许 旻1,2,林河通4,5

1集美大学海洋食品与生物工程学院,福建厦门 361021;2福建省海洋生物增养殖与高植化利用重点实验室,福建厦门 361000;3福建省食品微生物与酶工程重点实验室,福建厦门 361021;4亚热带特色农产品采后生物学福建省高校重点实验室,福州 350002;5福建农林大学食品科学学院,福州 350002)

摘 要采前因素是果实采后品质和贮藏保鲜的重要影响因素,近年来愈发受到重视。国内外科研工作者围绕果实采前处理技术开展了大量研究工作。笔者对相关研究进展进行综述,概述了果实采前处理技术特点和处理剂种类,探讨了采前处理对果实采后品质、贮藏生理和耐贮性、病害控制等方面的影响及作用机制,同时对果实采前处理技术的发展方向进行展望,以期为果实采前保鲜处理技术的研究、应用和发展提供参考。

关键词:果实;采前处理;贮藏品质;采后生理;耐贮性;诱导抗性

采后果实作为独立的有机体,其生理代谢变化既是商品性状变化的内在诱因,也是采前生命活动的延续。由于采收时生长发育状况决定机体代谢的物质和能量基础,因此采前因素是果蔬采后品质的重要先决条件。几十年来,果蔬采后商品化处理、包装、贮运和销售等环节通过各种物理、化学和生物方法进行保鲜,取得了丰厚的研究成果,其中气调技术、1-甲基环丙烯(1-methylcyclopropene,1-MCP)处理和壳聚糖涂膜等技术已在生产中广泛应用,大幅减少了采后损失[1]。同时,采前定期施用化学或生物制剂由于可以改善果蔬采收品质并增强采后抗性,因此一直是抑制采后品质败坏的重要措施。国内外大量研究发现,许多具有调节植物生长作用的物质不仅可以用于采后施用以延长果蔬货架期,还可以用于采前处理以诱导增强果蔬采后品质和耐贮性,这与常用的农用植物营养剂以及化学杀菌和除虫剂类似,因而得到越来越多的关注[2]。笔者就近年来国内外有关采前处理提高果实采后品质和贮藏适应性的研究进行综述,为采前处理技术在提高采后果实品质和延长保鲜期方面的应用和相关研究工作提供参考。

1 果实采前调节剂处理及其保鲜作用

果实的采前处理技术是指在果实盛花期、幼果期、果实膨大期及采摘前期使用处理剂对树体及果体进行3~5 次喷施,采前处理的时间和次数依据果实种类和品种而定,通常选择在晴天上午进行。采前保鲜处理方法主要有化学法、生物法以及复合法,其中化学法是目前研究较多的采前处理手段,如使用二氧化氯、赤霉素(gibberellin,GA3)、胺酰脂(diethyl aminoethyl hexanate,DA-6)、二乙基二硫代氨基甲酸钠(diethy dithiocarbamate,DDTC)以及茉莉酸甲酯(methyl jasmonate,MeJA)等采前喷布蓝莓果实、番茄叶片和樱桃树体等,可延缓果实采后的成熟衰老、提高贮藏期抗性和品质并延长货架期[3-4]。常见的采前调节处理剂主要分为植物生长调节剂类、壳聚糖类、钙盐类以及其他化学处理剂,因其性质和所引起的植物生理效应不同,对采后果实的保鲜作用及机制也各有特点。

1.1 植物生长调节剂类

植物生长调节剂是指与植物激素具有相似生物学效应并可调节植物生长发育的天然激素或人工合成物质,其施用受果实种类、品种、生长期以及外界环境因素的影响。植物生长调节剂具有用量小、作用面广、速度快、残留少等优点,低浓度即可达到促进或抑制果实生长成熟、增强抗逆性、影响风味物质和色素积累等效果,从而实现果实外部性状与内部生理过程的双调控。常用于增强果实采后保鲜效果研究的植物生长调节剂主要有GA3、DA-6、水杨酸(salicylic acid,SA)和茉莉酸等。研究发现,采前GA3喷施处理能够调节杧果体内叶绿素酶、苯丙氨酸解氨酶(phenylalanine ammonia-lyase,PAL)与查尔酮异构酶等多种酶活性,抑制果实采后色泽的转变,延缓果实衰老,提升贮藏品质[2]。杜云霞[5]研究发现,采前DA-6喷施处理龙眼可通过调节果实体内活性氧代谢、细胞膜脂肪酸代谢和能量代谢等多种生理过程控制龙眼果肉自溶,维持采后品质。SA是公认的植物抗病诱导剂,用于采前处理能够诱导植物病程相关蛋白(pathogenesis ralated proteins,PRs)的积累及PR1PR2PR3基因的表达,提高番茄果实的抗病性[6]。植物生长调节剂的前体物质或供体也常用于果实采前处理研究,如采前MeJA 处理可改善梨果实的色泽和风味,诱导抗逆性,抑制果实发病[7]

1.2 壳聚糖类

壳聚糖是从虾、蟹壳中提取的甲壳素经脱乙酰反应得到的碱性天然多糖,其中聚合度在2~20之间的低聚壳聚糖为壳寡糖。壳聚糖化学性质稳定且无毒,因其良好的成膜性、机械性和保湿性在果实采后保鲜处理中得到了广泛应用[8]。同时,壳聚糖具有广谱抗菌性和诱导抗性等多种生物活性,也常被用于果实的采前处理研究。采前对火龙果喷施壳聚糖溶液,可一定程度维持果实细胞膜的完整性,延缓果皮变薄和果实失重率上升,维持火龙果贮藏品质,延长其保鲜期[9]。Bhaskara等[10]报道,采前对草莓喷施壳聚糖溶液可显著降低采后草莓果实灰霉病的发生率,且高浓度壳聚糖的防腐保鲜效果较好。

1.3 钙盐类

钙是植物细胞分裂和果实生长发育过程中的重要营养元素,可作为第二信使在植物体内发挥信号传导作用,参与调控乙烯的合成,也可通过维持植物细胞的蛋白质合成能力,保护细胞膜的完整性,抑制成熟衰老相关酶的活性,进而延缓果实后熟和衰老[11]。Goutam 等[12]使用0.5%、1.0%和1.5%(w)的Ca(NO32溶液对番石榴进行采前喷施,发现质量分数为1.0%的Ca(NO32溶液处理能够较好维持番石榴果实可溶性固形物及可滴定酸含量,并降低腐烂率,保持采后贮藏品质。

1.4 其他化学处理剂

由于采前处理通常在自然环境下进行,具有周期长、间隔久、果实残留量少以及安全性高的特点,所以许多化学处理剂被用于果实采前处理技术的研究探索。1-MCP 作为近20 年来被广泛关注的乙烯抑制剂,能不同程度地抑制呼吸跃变型及非跃变型果实乙烯的产生和释放,延缓果实成熟衰老进程,已被广泛应用于果实保鲜[13-15],在采前处理的应用潜力也非常植得探索[16]。Li等[17]发现采前使用1-MCP对梨进行喷施处理,可有效抑制采后梨果实乙烯的产生,推迟呼吸高峰的出现,延缓梨果实的采后成熟。DDTC 是一种褐变抑制剂,林福兴[18]研究发现采前使用DDTC喷施龙眼可有效延缓采后龙眼果实果皮褐变,并延长其贮藏保鲜期。此外,复合类处理剂近年来也逐渐受到关注,常见的复合处理剂有植物生长调节剂复合试剂和壳聚糖复合试剂等[19-20]

2 采前处理对果实采后品质的影响

2.1 对果实色泽的影响

果皮色泽可直观反映果实成熟度和新鲜度,是果品外观性状的首要考量因素之一。叶绿素、花青素和类胡萝卜素是决定果实采后色泽的主要色素,在果实生长、成熟和衰老阶段进行不同程度的合成、积累与降解。果实色素合成机制是果实品质形成与调控领域的研究热点,近些年的研究表明,采前处理可调控色素合成基因,改变色素合成相关酶活性,从而影响采后果实色泽的形成。Shafiq等[21]使用浓度为5 mmol·L-1的MeJA 对粉红女士苹果进行采前处理,促进了苹果果皮中花青素和花色苷等黄酮类物质的积累,加快果实表皮转红,进而改善果实色泽。曾凤[22]研究发现,采前喷施4.0 g·L-1的GA3溶液能够降低贵妃杧果果实采后八氢番茄红素合成酶、查尔酮合成酶及查尔酮异构酶基因的表达水平,降低果皮查尔酮异构酶活性,从而减缓花青素和类胡萝卜素积累,推迟采后成熟期的果皮转色。采前处理对采后果实色素分解方向的影响作用也得到越来越多的关注。Saracoglu 等[23]研究发现,使用MeJA 对甜樱桃进行采前喷施,可通过延缓果实花色苷含量降低从而减缓果实颜色的变化,其中以浓度10 mmol·L-1 MeJA 为佳。刘豆豆等[24]报道,采前用0.05%(w)的壳寡糖溶液对杏树进行喷施,可抑制杏果实叶绿素酶和脱镁叶绿素酶活性,显著延缓叶绿素的降解,维持果实表皮颜色。

2.2 对果实质地的影响

果实的硬度是判断成熟度和耐贮运性的重要品质因素,综合反映果实的质地特性。研究表明,在果实采后流通和贮藏过程中,果实硬度变化的内因主要与果实后熟或衰老过程中果胶、纤维素和半纤维素等细胞壁多糖物质组分和含量的改变,以及细胞壁水解酶的作用密切相关[25]。采前处理剂调控果实采后硬度的研究也多基于抑制细胞壁多糖降解和调控细胞壁多糖降解酶活性展开。李红震等[26]在采前8、12 d 分别对泰山早霞苹果喷施含量为2%(w)的1-MCP 溶液,能够较好地保持苹果的果肉硬度,并对果实绵化起到一定防治效果。Sharma[27]研究发现,采前对亚热带桃果实使用1%(w)Ca(NO32和2%CaCl2 进行叶面喷施,均可改善果实的硬度,且Ca(NO32 的作用效果要优于CaCl2。Siddiqui等[28]在采前每隔7 d对苹果喷施1.2%(w)CaCl2,发现CaCl2可通过提高果实细胞壁的机械强度提高果实的平均硬度,但对小果实的硬度影响不显著。邓佳[19]使用GA3(25 mg·L-1)和SA(200 mg·L-1)复合溶液对葡萄柚进行采前喷施处理,可通过抑制多聚半乳糖醛酸酶、纤维素酶(cellulase,Cx)、果胶甲酯酶、α-L-阿拉伯呋喃糖苷酶和β-半乳糖苷酶基因的表达及酶的活性,减缓采后葡萄柚果实中果胶及半纤维素的降解、减缓细胞壁有效物质释放、维持果实硬度。

2.3 对果实风味的影响

果实的风味由果实组织中的糖、有机酸及挥发性芳香物质等共同决定。糖类不仅是影响果实感官质量的重要因素,还是果实采后主要代谢底物,与成熟衰老生理过程密切相关。因此,采前处理对果实中糖等有机物积累的影响作用既有助于风味品质形成,又可调节采后生理品质。Erogul等[29]发现,采前分别使用1、2 mmol·L-1 SA喷施Cresthaven桃,均可延缓果实中可溶性固形物及可滴定酸含量的下降,较好地维持采后桃果实的风味。Lal 等[30]于收获前使用0.5%、1.0%和1.5%(w)的CaCl2溶液对杏叶面进行喷施,显著提高了果实采收时的可溶性糖含量,使其糖酸比显著高于对照,果实风味得到提升。黄仁华等[31]报道,采前喷施SA溶液能够显著提高贮藏期内红肉脐橙果实的葡萄糖和果糖含量,以低浓度(0.25 mmol·L-1)处理效果较优,并且不同浓度处理均能抑制柠檬酸和苹果酸的消耗,这可能与SA处理可以延缓果实呼吸代谢以及推迟物质消耗和果实衰老有关。尚琪[32]使用1.0 mmol·L-1乙酰水杨酸在玛瑙厚皮甜瓜幼果期、膨大期、网纹形成期和采前2 d进行喷施处理,发现该处理增加了果实采后醛类、醇类、酯类物质以及总挥发性香气物质含量,并且在贮藏末期显著提高了2-甲基丁酸乙酯、壬烯醛、β-紫罗酮等特征香气物质的释放量。

综上,不同采前处理剂在适宜浓度和处理条件下有助于采后果实的色泽、质地、风味品质的形成与维持,从而有助于延长保鲜期。同时,采前处理对果实采后品质的影响作用与其对生理性状的影响密切相关。

3 采前处理对采后果实贮藏生理及耐贮性的调控作用

3.1 影响采后果实呼吸强度和乙烯释放

呼吸作用是采后果实物质代谢和能量代谢的中枢,直接关乎商品品质变化和货架期。果实自身产生的乙烯通常会促进呼吸作用,加速营养物质的消耗,加快失鲜和生理衰老进程。对果实进行适宜的采前处理,可通过调节果实生长和采收时的生理状态,影响采后呼吸作用和乙烯产生。张鹏等[33]将1-MCP 应用于阳光玫瑰葡萄的采前处理,发现采前1 d 喷施1µL·L-1的1-MCP 溶液可降低采后葡萄果实的乙烯释放量和呼吸强度。Mcartney等[34]采前使用1-MCP 处理Law Rome 苹果,抑制了果实采后30 d 内的乙烯释放浓度和呼吸速率,认为这可能与1-MCP 影响了果实乙烯与受体蛋白的结合有关。Reddy 等[35]报道,使用0.2 g·L-1的SA 对杧果进行采前处理,可有效抑制果实呼吸速率和乙烯合成速率,延迟杧果采后成熟衰老。Tezotto-Uliana等[36]的研究发现,从树莓果实生长转红期开始,每7 d 喷施1 次壳聚糖溶液,持续21 d,可在果实周围形成屏障,控制果实与环境中氧气的交换,进而影响1-氨基环丙烷-1-羧酸(ACC)氧化酶和ACC合酶的作用,有效减缓采后果实乙烯的生成和呼吸速率的升高,并降低果实腐烂率,且壳聚糖浓度越高,乙烯产量越低。Kumarihami等[37]的研究发现采前猕猴桃果实经100、500 mg·L-1壳聚糖溶液浸泡处理后,采后乙烯合成相关基因AdACS2AdACO2的表达受抑制,减缓了乙烯的合成与释放,且在贮藏后期抑制作用尤为明显。因此,适当采前处理可通过抑制果实呼吸和乙烯释放速率,延缓果实衰老进程,提高采后果实贮藏品质,但是其控制作用机制仍待进一步研究解析。

3.2 抑制采后果实失水失重

随着果实采后呼吸作用和蒸腾作用的进行,营养物质和水分不断散失,不可避免地造成果实失鲜和失重,加剧代谢紊乱,导致果实耐贮性和外观性状变差。目前,许多研究工作着眼于将常见的采后商品化处理保鲜剂用于采前措施以控制果实失水和失鲜。杨雪梅等[38]的研究发现,采前对泰山红石榴喷施1.25%(w)壳聚糖溶液,可减弱皮孔蒸腾作用,降低细胞膜渗透率,维持果皮内含水量,从而保持石榴表皮新鲜程度,提高贮藏期内的外观品质。Ehtesham 等[39]报道,采前发育阶段对葡萄喷施2%和3%(w)的壳聚糖溶液,可减少采后贮藏期间葡萄果实质量损失,而且3%(w)的壳聚糖溶液喷施叶面联合采后33%(w)芦荟凝胶覆膜处理,能够更好地保持果实的整体质量指数、硬度、花青素含量及抗氧化能力,缓解葡萄果实贮藏期老化。Ahmad 等[40]对Lane Late和Valencia Late柑橘树喷施不同浓度SA溶液,发现8、9 mmol·L-1的SA 溶液处理能够显著延缓果实质量下降,提高果皮紧实程度,降低采后损失。在冬枣果实白熟期向叶和果喷施含钙量为0.27%(w)的氯化钙或醋酸钙溶液,能够降低果实采后失水率,且醋酸钙处理对果实采后水分保持效果更佳[41]。因此,将壳聚糖等保鲜和保水剂用于果实采前处理,可能具有较好地延缓果实采后失重效果,但其作用机制与采后涂膜处理不尽相同,目前鲜有较为深入的研究报道。

3.3 控制采后果实代谢失调

果实采后在自身衰老过程中或在不适宜环境因素下,物质和能量消耗、组织细胞结构破坏及胁迫反应,引发生理代谢紊乱,其中活性氧对生物高分子和细胞膜系统的破坏,以及膜脂过氧化等作用被认为是导致植物果实细胞结构和功能失调、衰老和品质败坏的重要因素[42]。果实组织细胞抗氧化和能量水平的调控一直是采前和采后处理控制果实衰老、品质劣变的关注热点。Vicente 等[43]报道,使用浓度分别为0.1、0.5、1.0 mmol·L-1的MeJA 溶液喷施柠檬,可诱导贮藏期间果实中POD、PAL 和抗坏血酸过氧化物酶(Ascorbate Peroxidase,APX)等酶活性的提高,保持果实较高的抗氧化活性,减缓活性氧的产生和积累,进而推迟果实衰老。酚类和黄酮类物质是果实中重要的抗氧化和抗胁迫成分,对维持采后贮藏品质有显著作用[44]。He 等[45]的研究发现,采前使用50 mg·L-1壳寡糖溶液在幼苗期、开花前、盛花期和果实着色期分别喷施草莓植株,可有效增加果实中总酚和类黄酮含量,提高果实体外抗氧化能力,增强贮藏适应性。此外,能量作为物质代谢的必要条件,维持细胞系统的修复以及结构和功能完整性,可延缓膜脂过氧化作用,减轻生理代谢紊乱造成的伤害。林钟铨[44]的研究发现,采前使用0.5 g·L-1 SA处理龙眼,可通过抑制龙眼果皮中脂氧合酶和磷脂酶D的活性,减少膜脂过氧化,保持果皮细胞膜结构和功能的完整性,进而延缓代谢失调,提高耐贮性;而喷施含量(w)为10 mg·kg-1的DA-6可使采后龙眼果皮保持较高的ATP 和ADP 含量,延缓能荷水平下降,维持组织细胞区室化功能,有效延缓果皮褐变的发生[46]。王斌等[47]采用0.5 mmol·L-1 硝普钠对厚皮甜瓜植株进行多次喷施,显著提高了果实贮藏期间琥珀酸脱氢酶和苹果酸脱氢酶活性,延缓ATP 含量下降速率,提高了ADP 含量,维持了甜瓜果实较高的能荷植,从而延长贮藏期。

3.4 调控采后果实低温适应性

随着现代冷链技术的发展,低温流通系统已成为国内外新鲜果实贮运的普遍手段。对于冷敏型水果,不适宜的低温贮运流通环境极易造成果实冷害,出现表皮凹陷变色、水渍状斑点、不能正常后熟、芳香气味减退以及组织软化溃烂等症状,成为限制长期贮藏和远距离运销的重要因素,因此控制采后冷害、增强冷藏适应性一直是果实保鲜技术研究的热点[48]。冷害的发生与低温下果实正常生理代谢失调、活性氧积累导致膜脂过氧化作用以及膜脂发生相变造成细胞膜组织损伤密切相关。采前使用不同诱导剂对果实进行喷施处理,可在一定程度上增强采后果实的抗冷性,减少冷害现象的发生。许多研究表明,采前喷施包括MeJA、SA和GA3在内的植物生长调节剂可诱导石榴、菠萝和李等果实提高抗冷性进而减少采后果实的冷害[49]。活性氧清除系统可调节果实内部活性氧的产生与积累,对维持果实细胞膜结构的完整性具有重要作用,诱导果实维持较高的活性氧清除活力,是采前处理诱导果实采后抗冷性的主要机制之一。Lu等[50]的研究发现,采前喷施2.0 mmol·L-1 SA 溶液可显著抑制菠萝冷藏期间PPO 和PAL 活性,缓解冷藏期间菠萝果实的内部褐变,提高果实抗性,减缓冷害发生。司敏等[51]在红宝石李果实发育期连续喷施3次质量浓度为100 mg·L-1的GA3溶液,发现贮藏25 d 时果实超氧阴离子产生速率和过氧化氢含量较对照组分别低80.36%和46.44%,超氧化物歧化酶(superoxide dismutase,SOD)和过氧化氢酶(catalase,CAT)活性分别较对照组高57.98%和35.69%,因此认为GA3可通过提高抗氧化酶活性,降低果实中自由基水平,以维持李果实细胞膜的稳定,提高低温胁迫抗性,减缓冷害症状。

4 采前调节剂处理对采后果实病害的控制作用

果实采后流通过程中微生物病害的发生受采前田间带病、采后机械损伤和贮藏环境不适宜等诸多因素影响。多数果实的采后病害和田间病害由同一种病原菌侵染引起,即潜伏性侵染是诱发果实采后病害的主要方式[44]。因此,单靠采后措施防治果实病害的效果往往欠佳,采前喷施适宜诱导剂以控制采后病害的方法也广受研究人员关注。研究发现,喷施SA[6]、MeJA[52]、DA-6[5]、壳聚糖[53]等处理剂可通过抑制病原菌或诱导果实抗病性增强等方式有效减少果实采后贮藏期间的病害发生。

4.1 对病原菌的抑制作用

SA、GA3、MeJA 和壳聚糖等物质具有杀灭果皮表面潜伏菌、抑制病菌孢子的萌发以及降低病斑扩展的作用,有助于抑制病害[54]。弓德强等[55]使用SA溶液对红芒6 号杧果进行采前处理,并于采后接种炭疽菌,发现SA处理能有效降低接种果实的发病率及病斑扩展速度,同时显著降低果实贮藏期间自然发病指数,有效抑制杧果果实采后病害发生。Yao等[54]指出,采前对甜樱桃喷施2 mmol·L-1的SA溶液可对果实褐腐病菌菌丝生长和菌落扩展有明显抑制作用。采前处理与采后物理处理方法结合也是颇具应用潜力的病害控制措施。在苹果梨的幼果期、膨大期和成熟期分别使用50 mg·L-1的GA3溶液进行处理,结合采后热处理,可使果实采后黑皮病的发病指数降低69.2%[56]。也有研究表明,采前处理虽可抑制果实主要腐败菌的菌斑形成,但对潜伏性较强的病原菌作用有限[57]

4.2 诱导果实抗病性

采前处理剂诱导抗病的作用主要体现在提高或维持果实体内抗性物质含量与抗病蛋白活性方面,从而提升果实自身防御系统,并且其诱导效应与浓度相关,低浓度喷施时通常能促进果实产生抗病性,而较高浓度可能会使果实抗病性下降,加速果实发病[44]。王斌等[58]使用1.0 mmol·L-1的乙酰水杨酸在哈密瓜果实坐果期、膨大期、网纹形成期及后熟期(采前2 d)进行喷施处理,在采后接种粉红单端孢,发现乙酰水杨酸处理能显著提高哈密瓜果实PAL、4-香豆酰-辅酶A连接酶和肉桂酸羟化酶活性,提高总酚和类黄酮含量,诱导果实增强抗病性,显著降低和减少了哈密瓜果实贮藏期间的发病率和病斑面积。Cao 等[59]的研究发现,使用2.0 mmol·L-1的SA在开花后30、60、90、110 d 对枣进行喷施,可诱导果实体内β-1,3-葡聚糖酶、POD及SOD等抗病蛋白活性增强,从而诱导果实对病原菌的抗性增强,且该抗性可在采后持续存在。Felipe 等[60]报道,用250µmoL·L-1的MeJA溶液采前处理能够诱导采后草莓果实几丁质酶、β-1,3-葡聚糖酶和多半乳糖醛酸酶的活性上升,并提高相关防御基因FcBG2-1FcBG2-3FcPGIP1FcPGIP2FcCHI2-2FcCHI3-1 的表达水平,激活了采后草莓果实的防御系统,从而提高了抗病性。

5 展 望

果实采收前适时喷施适当的处理剂,能够有效提高果实采后品质、减缓生理代谢失调、提高贮藏适应性和抗性并延缓果实衰老败坏,并且具有无毒、无污染和长效性等优点,因此在果实采后保鲜方面的作用广受研究人员关注,得到了许多具有前景的应用性研究成果,但在生产上还需进一步推广。在应用方面,采前处理剂的使用除根据处理剂种类确定最佳处理时间、次数及方式外,还需注意不同处理剂和处理方式间的协同作用。因此,应针对果实种类、生长特性和环境因素确定处理剂的种类、剂量以及配套措施,开发实用新型采前处理剂和复合处理技术体系。有关采前处理延长果实保鲜期的研究目前仍多集中于贮藏品质和采后生理层面,而采前物质和能量积累对耐贮性的影响以及深层次生物学机制,尤其是采前生长阶段到采后流通过程中的持续性生物学效应也植得更多关注。随着植物生理学和采后生物学的发展,以及科研人员对影响果实采后品质和安全的采前因素的愈发重视,采前处理在果实质量和保鲜领域的研究应用仍具有广阔的前景。

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Advances in the effects of preharvest treatments on fresh-keeping of postharvest fruits

SUN Minzi1, 2, 3,ZHANG Shen1*,WU Guangbin1,CHEN Fahe1,NI Hui1, 3,CHEN Xinglin4, 5*,XU Min1, 2,LIN Hetong4,5
(1College of Ocean Food and Biological Engineering, Jimei University, Xiamen 361021, Fujian, China;2Key Laboratory of Cultivation and High-value Utilization of Marine Organisms in Fujian Province,Xiamen 361000,Fujian,China;3Fujian Provincial Key Laboratory of Food Microbiology and Enzyme Engineering, Xiamen 361021, Fujian, China;4Key Laboratory of Postharvest Biology of Subtropical Special Agricultural Products,Fujian Province University,Fuzhou 350002,Fujian,China;5 College of Food Science,Fujian Agriculture and Forestry University,Fuzhou 350002,Fujian,China)

Abstract:The life activities of the postharvest fruits determine the commercial quality of the fruits,and the preharvest treatments are important for the life activities, quality and shelf-life of the postharvest fruits.Researchers have paid more attentions on the importance of preharvest application of plant regulators for enhancing the quality and storability of postharvest fruits for several decades. Thus, lots of work focusing on the postharvest effectiveness of preharvest treatments have been carried out constantly.This paper reviews the domestic and abroad efforts on developing preharvest treating strategy and investigating their effects on fresh-keeping and physiological activities. Generally, the preharvest treatment refers to the handlings of spraying regulator solutions three to five times on the leaves or fruits at full-bloom stage, young fruit stage, rapid growth stage and/or fruit ripening stage. The treating agents are usually naturally acquired chemicals eco-friendly and easy to absorb.The plant growth regulator is the largest category of the treating agents, which mainly includes gibberellin, diethyl aminoethyl hexanoate,salicylic acid,jasmonic acid and their precursors.Moreover,chitosan and calcium salts are widely and frequently used agents.Other chemical regulators like 1-methylcyclopropene and sodium diethyldithiocarbamate are also employed in the studies.Preharvest spraying with these agents is capable of enhancing and maintaining fruit quality at harvest and during postharvest storage:the accumulation of pigments like anthocyanin could be elevated before harvest, and the postharvest de-greening or coloring might be postponed via regulating ripeness and senescence;the fruit firmness is closely associated with cell wall polysaccharides like pectin,cellulose,and hemi-cellulose,which could be retained after preharvest handlings via suppressing the degrading enzymes like polygalacturonic acid,pectin methylesterase,cellulase and corresponding genes;the flavor characteristic largely depends on the carbohydrates,organic acid and volatiles,and their accumulation can be improved by preharvest treatments to retard postharvest flavor decay.These effects of preharvest treatment on quality retention are reflections of its postharvest physiological impacts, which mainly include the inhibitions on the respiration, ethylene synthesis and action, water loss, and metabolic disorder, and the improvements in the refrigeration adaptability.As the key metabolism of postharvest fruit,respiration signifies the intensity of ripeness and senescence progress in relation with the storability and shelf-life, while the endogenous ethylene is well acknowledged for dominating the respiration and accelerating fruit aging. Application of preharvest regulator could inhibit the respiration and ethylene production via influencing the physiological status during growth and at harvest, or blocking the gas exchange between fruits and environment, and further suppressing ethylene synthesis.Postharvest fruit water loss due to respiration and transpiration is an important accelerator of metabolic dysfunction.Some agents used in postharvest water retention handling like chitosan are also employed in preharvest treatment and exhibited sound effects, but its function would not be the same as postharvest coating and would deserve more investigation. Another key factor of physiological dysfunction is the loss of cellular structural and functional integrity,which is closely associated with the attack of reactive oxygen species(ROS)and energy deficit.Preharvest spraying with regulators might induce higher oxidation resistance to delay the production and accumulation of ROS via improving levels of antioxidants and ROS-scavenging enzymes.It has been reported that the energy status during storage could be maintained after preharvest treatment, protecting the cellular structure and retarding the metabolic disorder. The cold chain has become a common measurement for postharvest handling, distribution, and marketing, while chilling injury is hard to avoid during these processes for cold sensitive fruits. Many researches found that preharvest treatment with plant growth regulators like gibberellin, salicylic acid, and jasmonic acid has notable effect of inducing the cold resistance of the cold sensitive fruit like peach,plum and pineapple.The main inducing mechanism has been reported to be related to higher ROS-scavenging system protecting the compartmentalization in cell and reducing the membrane lipid peroxidation. On the other hand, preharvest handlings may also prolong fruit storage period via inhibiting postharvest diseases.Since most fruit pathogenic invasions are latent infection,preharvest prevention is a critical measure. Most preharvest regulators such as salicylic acid, jasmonic acid, gibberellin, and chitosan are natural antimicrobial agents capable of preventing spoilage organisms.More importantly,they are also disease resistance inducers that has long-term enhancement on the resistant proteins such as phenylalnine ammonialyase, β-1,3-glucanase and peroxidase, and increases the disease resistant compounds like phenolics.Therefore,preharvest application of regulators is a highly feasible and effective handling to improve fruit commercial quality, storage adaptability and resistance to diseases, and its mechanism of action is an integrated function of physiological impacts that still demands lots of constant investigation.This paper would provide an important reference for the research,extension,and development of fresh-keeping oriented preharvest handling technology.

Key words: Fruit; Preharvest treatment; Storage quality; Postharvest physiology; Storability; Induced resistance

中图分类号:S66

文献标志码:A

文章编号:1009-9980(2022)06-1111-10

DOI:10.13925/j.cnki.gsxb.20210417

收稿日期:2021-09-03

接受日期:2022-03-02

基金项目:福建省自然科学基金项目(2020J01675);福建省教育厅中青年教师教育科研项目(JAT190338);福建省海洋生物增养殖与高植化利用重点实验室开放基金(2020fjscq02);福建省食品微生物与酶工程重点实验室开放基金(Z820260)

作者简介:孙闽子,女,在读硕士研究生,研究方向为农产品加工及贮藏工程。Tel:18710959665,E-mail:mz529@163.com

*通信作者Author for correspondence.Tel:18850561192,E-mail:szhang8811@jmu.edu.cn;Tel:13860450177,E-mail:chenxinglin@tio.org.cn