褪黑素在果蔬采后保鮮的應(yīng)用研究進(jìn)展

瞿光凡，巴良杰，王瑞，馬超，吉寧，曹森

褪黑素在果蔬采后保鮮的應(yīng)用研究進(jìn)展

瞿光凡1,2，巴良杰1,2，王瑞1,2，馬超1,2，吉寧1,2，曹森1,2

（1.貴陽學(xué)院 食品與制藥工程學(xué)院，貴陽 550005；2.貴州省果品加工工程技術(shù)研究中心，貴陽 550005）

通過介紹褪黑素對采后果蔬生理、營養(yǎng)品質(zhì)、能量代謝、風(fēng)味變化、冷害調(diào)控和抗病性的影響，為褪黑素在果蔬采后保鮮的應(yīng)用提供理論借鑒。綜述國內(nèi)外褪黑素在不同品種果蔬采后上的保鮮效果。褪黑素可以調(diào)控果蔬的成熟與衰老，提高果蔬的抗冷性和抗病性，維持果蔬較高的營養(yǎng)物質(zhì)，保持果蔬更好的生理品質(zhì)。褪黑素可顯著延長采后果蔬的貯藏期，在未來采后果蔬保鮮領(lǐng)域具有廣闊的發(fā)展空間。

褪黑素；果蔬；保鮮效果；研究進(jìn)展

褪黑素（Melatonin, MT）是一種內(nèi)源性、具有多種調(diào)節(jié)功能的生物活性分子，可以參與果蔬的生長、分化、成熟、衰老和防御等多種生理活動[1]。褪黑素也是一種天然、高效、安全性高、無污染的新型生物保鮮劑[2]，其已廣泛應(yīng)用于果蔬的采后保鮮，可有效提高果實(shí)的抗冷性[3]，延緩果蔬的成熟衰老進(jìn)程[4-5]，保持鮮切果實(shí)較高的營養(yǎng)品質(zhì)[6]，提高果蔬的抗病性[7-8]。

近年來研究者發(fā)現(xiàn)，褪黑素對果蔬采后的保鮮作用主要通過降低果蔬的呼吸速率、降低乙烯生成量[9-10]、延緩果實(shí)成熟來達(dá)到；也有研究者報(bào)道，褪黑素作為一種信號分子，具有較強(qiáng)的抗氧化能力，不僅可以誘導(dǎo)內(nèi)源褪黑素積累，從而調(diào)控果蔬的生理、生化和分子機(jī)制，還能通過中和自由基、增強(qiáng)酶或非酶抗氧化活性、防止細(xì)胞內(nèi)抗氧化酶失活、抑制活性氧自由基的產(chǎn)生等模式來調(diào)控果實(shí)的成熟衰老[11-12]。文中主要綜述國內(nèi)外研究者報(bào)道的關(guān)于褪黑素對果蔬采后成熟衰老、品質(zhì)劣變、病害防控的調(diào)控作用等方面，以期為褪黑素在果蔬采后貯藏保鮮上的進(jìn)一步研究提供理論指導(dǎo)。

1 褪黑素對采后果蔬成熟衰老的調(diào)控

1.1 褪黑素對呼吸及乙烯的影響

果蔬在呼吸過程中伴隨著內(nèi)源乙烯的增加，內(nèi)源乙烯的積累加快了果蔬的成熟軟化進(jìn)程[13]。近年來，MT在改善采后果蔬品質(zhì)、延緩果蔬成熟衰老中發(fā)揮著重要作用。Hu等[14]研究發(fā)現(xiàn)，外源褪黑素處理能顯著抑制香蕉果實(shí)中乙烯合成相關(guān)基因ACS、ACO表達(dá)和淀粉的降解，推遲了香蕉果實(shí)呼吸高峰的出現(xiàn)，有效延遲了香蕉果實(shí)成熟的進(jìn)程。Tan等[5]研究發(fā)現(xiàn)，采用褪黑素噴灑處理的大白菜，其對照組的呼吸速率和乙烯釋放量顯著高于處理組，并且處理組的葉綠素含量顯著大于對照組，從而有效延緩了大白菜的衰老黃化進(jìn)程。研究者還發(fā)現(xiàn)，采用褪黑素處理可以顯著提升梨果實(shí)內(nèi)一氧化氮合酶的活性與TDC、T5H基因的表達(dá)量，從而導(dǎo)致梨果的乙烯生物合成量和呼吸速率下降[15]。MT還可通過調(diào)節(jié)蘋果貯藏期間的呼吸速率和乙烯的生成，降低果實(shí)細(xì)胞中丙二醛的積累和果實(shí)水分的流失，顯著延長蘋果的貯藏期[16]。此外，外源褪黑素處理還可降低桃果[3]、芒果[17]、白菜[5]的呼吸強(qiáng)度及內(nèi)源乙烯的生成量，使采后果蔬的呼吸處于一種低而正常的狀態(tài)，有利于延長果蔬貯藏期。

1.2 褪黑素對能量代謝的影響

采后貯藏的果蔬仍然可以正常進(jìn)行呼吸和生理機(jī)能代謝活動，而果蔬的生理機(jī)能代謝活動需ATP提供能量[18]，低能量狀態(tài)會增強(qiáng)果蔬的呼吸速率，消耗大量的有機(jī)物，加劇活性氧的積累，并加速膜脂過氧化，導(dǎo)致果蔬生理機(jī)能下降[19]，而高能量狀態(tài)可維持果蔬正常的生理機(jī)能代謝活動，延遲采后果蔬的后熟衰老進(jìn)程。胡苗[19]研究發(fā)現(xiàn)，外源褪黑素均能抑制冷藏期間水蜜桃SDH、CCO、H+-ATP酶和Ca2+-ATP酶的活力，提高桃果的能荷水平。Wang等[20]研究結(jié)果顯示，0.4 mmol/L褪黑素處理可抑制荔枝果皮SDH、CCO、H＋-ATP酶和Ca2＋-ATP酶的活性，維持荔枝果中較高的脂肪酸、ATP和ADP等含量，使果皮組織細(xì)胞處于高能量狀態(tài)，從而延緩荔枝果實(shí)的品質(zhì)劣變進(jìn)程。此外，褪黑素處理還能激活三磷酸腺苷酶和煙酰胺腺嘌呤二核苷酸激酶活性，維持大白菜葉片較高的能量狀態(tài)，延緩大白菜葉片黃化進(jìn)程[5]。由此可見，能量狀態(tài)與果蔬的生理活動密切相關(guān)，調(diào)控采后果蔬的能量狀態(tài)對延長果蔬的貯藏期尤其重要。

2 褪黑素對采后果蔬品質(zhì)劣變的調(diào)控

2.1 褪黑素對生理和營養(yǎng)的影響

果蔬在采摘后會快速衰老，表現(xiàn)為脫水、表皮皺縮、果肉軟化及腐爛等[13]，導(dǎo)致果蔬的品質(zhì)下降。Liu等[21]研究發(fā)現(xiàn)，采用0.1 mol/L外源褪黑素浸泡處理草莓果實(shí)，有效降低了草莓的腐爛率和質(zhì)量損失率，有效減緩了果實(shí)中水分的流失，提高了草莓果實(shí)的亮度。據(jù)Gao等[3]報(bào)道可知，外源褪黑素浸泡處理可誘導(dǎo)提高桃果實(shí)在貯藏期間的抗氧化酶活性，從而維持ROS代謝平衡，促進(jìn)了桃果實(shí)酚類物質(zhì)的積累。Rastegar等[22]研究了褪黑素浸泡處理采后芒果，發(fā)現(xiàn)處理后可顯著抑制芒果中抗壞血酸、總酚、總黃酮和可滴定酸等含量的下降，穩(wěn)定果實(shí)膜脂結(jié)構(gòu)、 增強(qiáng)膜脂功能，提高了芒果貯藏期的營養(yǎng)品質(zhì)。Xia等[23]的研究結(jié)果表明，褪黑素噴灑處理可上調(diào)葡萄漿果MYBA1和MYBA2基因表達(dá)，促進(jìn)果實(shí)中花青素的積累，增加可溶性糖的含量，并提高礦物質(zhì)營養(yǎng)素N、K、Cu、Fe、Zn的含量。研究還發(fā)現(xiàn)，經(jīng)褪黑素浸泡過的黃瓜，其處理組的可滴定酸、抗壞血酸、可溶性蛋白和葉綠素等含量顯著高于對照組，有效減少了黃瓜貯藏期間營養(yǎng)物質(zhì)的流失[24]。此外，褪黑素處理還可誘導(dǎo)雙孢菇帽酚類物質(zhì)和抗壞血酸含量的增加，改善雙孢蘑菇帽的營養(yǎng)品質(zhì)[25]。

2.2 褪黑素對風(fēng)味變化的影響

果蔬中的有機(jī)酸、糖類、酯類化合物、揮發(fā)性物質(zhì)等是形成果蔬風(fēng)味的重要物質(zhì)[26]。果蔬的風(fēng)味與果蔬的品質(zhì)密切相關(guān)，當(dāng)果蔬的品質(zhì)發(fā)生劣變，果蔬的風(fēng)味會受到一定的影響，并散發(fā)出令人不愉快的氣味。杜天浩等[27]研究發(fā)現(xiàn)，MT浸泡處理提高了在鹽脅迫的逆境環(huán)境下番茄果實(shí)的有機(jī)酸、糖酸比和揮發(fā)性芳香物質(zhì)等含量，增加了番茄果實(shí)的香氣成分。王紀(jì)忠等[28]發(fā)現(xiàn)，MT有效提高了梨果實(shí)貨架期的可溶固形物含量，增加了果實(shí)中酯類、醛類和酮類物質(zhì)，保持了梨果特有的風(fēng)味。朱賽賽等[26]探究不同濃度MT對采后蜂糖李果實(shí)保鮮效果的影響，結(jié)果表明，適宜的MT含量會將果實(shí)中抗壞血酸、可滴定酸和可溶性糖等含量維持在較高的水平，有利于果實(shí)揮發(fā)性成分的積累，有效保持了蜂糖李的鮮味。另外，劉建龍[29]研究結(jié)果表明，褪黑素處理可上調(diào)蔗糖磷酸合酶和蔗糖合酶基因表達(dá)，降低酸性轉(zhuǎn)化酶的基因表達(dá)，從而誘導(dǎo)增強(qiáng)蔗糖磷酸合酶和蔗糖合酶的活性，維持較低酸性轉(zhuǎn)化酶活性，有助于減緩梨果實(shí)中糖的降解進(jìn)程。Xia等[23]報(bào)道，褪黑激素會誘導(dǎo)增強(qiáng)蔗糖磷酸合酶的活性，增加葡萄果實(shí)的可溶性糖含量，從而促進(jìn)果實(shí)內(nèi)源MT的積累和激活花青素合成的通路。此外，Tan等[5]研究發(fā)現(xiàn)，外源MT處理可調(diào)節(jié)大白菜中葡萄糖-6-磷酸脫氫酶和6-磷酸葡萄糖酸脫氫酶活性的上升，使白菜葉片的總糖含量保持在較高的水平。由此可知，外源褪黑素處理通過誘導(dǎo)提高果蔬中有機(jī)酸、糖酸比、醛類和酮類等物質(zhì)的含量，從而維持果蔬的風(fēng)味。

2.3 褪黑素對冷害調(diào)控的影響

低溫能降低果蔬貯藏環(huán)境中氧氣與二氧化碳的比例，抑制果蔬中有機(jī)物的降解和病原菌的生長，限制采后果蔬中酶的活力，從而延長果蔬采后的貯藏期[9]。由于低溫貯藏容易導(dǎo)致果蔬發(fā)生冷害，果蔬低溫冷害的發(fā)生通常表現(xiàn)為冰點(diǎn)以上溫度對果蔬的細(xì)胞、組織和器官造成傷害，使果蔬的細(xì)胞、組織和器官的機(jī)能下降，引起果蔬代謝紊亂，對致病菌的抵抗能力下 降[30]。發(fā)生冷害的果蔬表面較黏滑，躍變型果實(shí)不能正常后熟，果蔬表皮出現(xiàn)皺縮凹陷、斑紋，果肉出現(xiàn)點(diǎn)蝕，且花萼變黑、組織水分流失，導(dǎo)致果實(shí)的硬度上升，從而出現(xiàn)僵果、木質(zhì)化加重、局部果肉部分浸水、果蔬組織褐變、果蔬風(fēng)味喪失、產(chǎn)生異味、腐爛等現(xiàn)象[30-31]。此外，有些果蔬在貨架時(shí)才表現(xiàn)出冷害癥狀，導(dǎo)致果蔬快速腐爛變質(zhì)，失去食用價(jià)值。

2.3.1 調(diào)節(jié)ROS代謝平衡

果蔬細(xì)胞中ROS自由基積累會引起果蔬膜完整性和功能的喪失，膜脂抗氧化能力下降，以及電解質(zhì)外滲，從而導(dǎo)致果蔬冷害的發(fā)生[32]。Shang等[33]研究結(jié)果顯示，采用0.05 mmol/L褪黑素處理會顯著提高藍(lán)莓中抗壞血酸過氧化物酶和谷胱甘肽S-轉(zhuǎn)移酶的活性，維持較高ROS自由基清除能力，有效提高藍(lán)莓果實(shí)的抗冷能力。Gao等[34]還發(fā)現(xiàn)，褪黑素處理促進(jìn)了草菇檸檬酸循環(huán)、谷胱甘肽代謝和脯氨酸代謝，激活了脂氧合酶、谷胱甘肽還原酶和SOD活性，以此減輕了鎘對草菇的損傷，有利于草菇對冷脅迫的抵御。另外，褪黑素還可誘導(dǎo)提高蘋果中抗壞血酸過氧化物酶、脂氧合酶和谷胱甘肽硫轉(zhuǎn)移酶等的活性，增強(qiáng)蘋果的抗冷性[35]。此外，MT會觸發(fā)番茄果實(shí)中ROS自由基清除系統(tǒng)，以及抑制細(xì)胞凋亡的蛋白質(zhì)積累，從而增強(qiáng)番茄果實(shí)的抗冷應(yīng)激能力[36]。

2.3.2 調(diào)節(jié)膜脂及酚類代謝

采后果蔬的冷害和衰老與細(xì)胞膜系統(tǒng)的完整性和功能緊密相關(guān)，膜結(jié)構(gòu)和功能的損傷導(dǎo)致果實(shí)細(xì)胞中膜脂降解，以及酚類物質(zhì)被酶氧化轉(zhuǎn)化為醌類物質(zhì)，從而引起果蔬褐變，且食用時(shí)感覺淡而無味[37-38]。研究發(fā)現(xiàn)，采用褪黑素處理荔枝果實(shí)，其磷脂酶D、脂肪酶和脂氧合酶的活性顯著上升，同時(shí)減緩了磷脂酰膽堿（PC）水解成磷脂酸（PA），使荔枝果實(shí)中油酸、亞油酸和亞麻酸的含量維持在較高水平，減輕了荔枝冷害及褐變的發(fā)生[20]。Gao等[3]報(bào)道了采用褪黑素浸泡桃果，維持了桃果較高的不飽和脂肪酸含量和較低的飽和脂肪酸含量，誘導(dǎo)G6PDH、SKDH和PAL活性增加，且促進(jìn)了總酚物質(zhì)和內(nèi)源性水楊酸的積累，減輕了桃果冷害的發(fā)生。Aghdam等[39]研究結(jié)果顯示，褪黑素處理能夠提高石榴果實(shí)在冷藏過程中APX、GR、G6PDH活性，促進(jìn)酚類、花青素、AA和谷胱甘肽的積累，增強(qiáng)石榴果實(shí)的抗冷能力。此外，褪黑素處理還可增加青椒脯氨酸、亞油酸與亞麻酸的比例[40]，激活番茄果實(shí)GABA分流通路活性[41]，從而減輕了果蔬的冷害癥狀。

2.3.3 激活精氨酸代謝通路

果蔬中聚胺、脯氨酸、γ-氨基丁酸和一氧化氮等物質(zhì)能夠在果蔬受冷激脅迫下作出響應(yīng)，而精氨酸不僅通過NOS反應(yīng)生成一氧化氮，還可通過精氨酸脫羧酶和精氨酸酶合成聚胺和鳥氨酸，鳥氨酸進(jìn)一步生成腐胺[42-43]。此外，精氨酸又可用于鳥氨酸脫羧酶或脯氨酸氨基轉(zhuǎn)移酶合成聚氨酸[42]?？梢姡彼岬拇x受到多種酶的調(diào)控，當(dāng)果蔬受到冷激脅迫或外界環(huán)境刺激時(shí)，酶活性的變化會調(diào)控精氨酸的代謝途徑和方向。Bajwa等[44]研究發(fā)現(xiàn)，褪黑激素通過上調(diào)ZAT10和ZAT12基因的表達(dá)，并激活CBFs基因表達(dá)，促進(jìn)擬南芥內(nèi)源性聚胺的積累，提高擬南芥的抗冷性。研究還發(fā)現(xiàn)，褪黑激處理上調(diào)了ZAT12基因的表達(dá)，提升了內(nèi)源性聚胺的積累和ROS清除系統(tǒng)的活性，增強(qiáng)了黃瓜幼苗的耐寒性[45]。此外，據(jù)Aghdam等[42]報(bào)道可知，褪黑素誘導(dǎo)了番茄果實(shí)ADC、ODC、P5CS、OAT和NOS基因表達(dá)上調(diào)，激活了CBF1信號觸發(fā)精氨酸的代謝通路，從而促進(jìn)了內(nèi)源性聚氨酸、脯氨酸和一氧化氮的積累，增強(qiáng)了番茄果實(shí)的耐寒性。

2.3.4 誘導(dǎo)采后果蔬基因表達(dá)

近年來，關(guān)于MT在果蔬采后保鮮的研究主要集于果蔬的生理層面，也有許多研究者從分子的角度解釋了MT在一些果蔬上的作用機(jī)制。果蔬的生理機(jī)能不僅受到外部環(huán)境的干擾，還受到內(nèi)部基因和轉(zhuǎn)錄因子的調(diào)控。據(jù)報(bào)道，MT通過抑制BrABF1、BrABF4和BrABI5表達(dá)，阻止了脫落酸的生成和葉綠素的降解，提高了大白菜葉片對逆境脅迫的抗性[46]。Shang等[33]研究表明，MT誘導(dǎo)了抗氧化基因VcAPX、VcGST和VcPAL的表達(dá)量上調(diào)，使藍(lán)莓的膜脂過氧化降低，延緩了藍(lán)莓果實(shí)冷害的發(fā)生。MT還能誘導(dǎo)上調(diào)基因MeCAT1、MeGPX、MePX3和MeGST的表達(dá)量，從而延緩木薯根的衰老[47]。另外，MT調(diào)節(jié)基因SND2、KNAT7、MYB20和MYB85表達(dá)量的下調(diào)，減輕了竹筍的木質(zhì)化[48]，觸發(fā)了TDC、T5S、SNAT和ASMT基因的表達(dá)，從而提高了內(nèi)源MT的積累，并調(diào)控了苯丙烷通路，增加了酚類、花青素及谷胱甘肽等含量，增強(qiáng)了石榴果實(shí)的抗冷能力[39]。此外，褪黑素處理誘導(dǎo)了miR528表達(dá)，降低了MaPPO1、MaPPO2和MaPPO3潛在靶基因表達(dá)，促進(jìn)了香蕉果實(shí)中磷脂和不飽和脂肪酸的積累，減輕了果皮褐變，提高了果實(shí)的抗冷脅迫能力[49]。Kong等[40]研究發(fā)現(xiàn)，采用100 μmol/L褪黑素處理上調(diào)節(jié)了CaSOD、CaPOD、CaCAT和CAAPX基因的轉(zhuǎn)錄水平，觸發(fā)了青椒中的抗氧化保護(hù)系統(tǒng)，從而抵抗了氧化損傷，增強(qiáng)了青椒對冷脅迫的抵抗。

3 采后果蔬病害的防控

果蔬采后在貯運(yùn)和貯藏過程中容易遭受損傷，引起果蔬代謝紊亂，導(dǎo)致果蔬的免疫力下降，從而促進(jìn)病原菌的生長繁殖。由病原菌引起果蔬采后品質(zhì)劣變是近年來仍需解決的一個(gè)難題。生吉萍等[50]采用100 mmol/L褪黑素采前噴施處理誘導(dǎo)番茄果實(shí)中幾丁質(zhì)酶、β-1,3-葡聚糖酶等抗病相關(guān)酶活力的上升，有效增強(qiáng)了貯藏期間番茄果實(shí)的抗病能力。孫子荀等[51]發(fā)現(xiàn)，外源褪黑素能有效抑制黑斑病致病菌鏈格孢菌的菌絲生長，增強(qiáng)采后草莓對黑斑病入侵的抵御。外源褪黑素也能誘導(dǎo)果實(shí)內(nèi)防御酶活性的上升，抑制蘋果灰霉病病斑的擴(kuò)大[52]，并且還可激活苯丙烷代謝的通路，促進(jìn)果實(shí)IAA的積累，增強(qiáng)荔枝對霜霉病的抵抗能力[53]。此外，褪黑素誘導(dǎo)提高PbOPR3基因表達(dá)來調(diào)控茉莉酸的通路[54]，激活SA信號通路，并誘導(dǎo)SA受體蛋白NPR1的核定位[55]，調(diào)控果蔬細(xì)胞H2O2的積累，調(diào)節(jié)細(xì)胞內(nèi)活性氧自由基平衡[56]，上調(diào)NO合成基因表達(dá)，促進(jìn)果實(shí)細(xì)胞內(nèi)NO的積累[42,56]，并且激活MaHSP90s信號分子，增加吲哚乙酸的積累[57]，從而提高采后果蔬的抗病能力。目前關(guān)于MT調(diào)控增強(qiáng)采后果蔬的抑菌機(jī)制的研究報(bào)道仍較少，還需進(jìn)一步探究MT誘導(dǎo)果蔬抗病的機(jī)制。

4 結(jié)語

褪黑素不僅可作為一種信號分子，參與調(diào)節(jié)果蔬采后生理代謝，保持果蔬較好的營養(yǎng)品質(zhì)，提高采后果蔬總糖含量和能量水平，改善果蔬風(fēng)味，提高果蔬的抗病性，還可作為一種抗氧化物質(zhì)誘導(dǎo)增加果蔬的采后抗氧化能力，調(diào)節(jié)ROS代謝平衡，增強(qiáng)果蔬對外界生物或非生物脅迫，提高果蔬對冷激脅迫的應(yīng)答，延遲果蔬采后成熟衰老。目前，褪黑素對采后果蔬的冷害機(jī)制以及冷害影響果蔬的生理生化的研究已比較深入，但褪黑素對采后果蔬的風(fēng)味變化和抗病性的研究報(bào)道還較少，且多數(shù)集中于果蔬的生理層面，未來可探究褪黑素對采后果蔬風(fēng)味及抗病性的影響，為褪黑素在果蔬保鮮方面的應(yīng)用提供更多理論支撐。

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Application of Melatonin in Postharvest Preservation of Fruits and Vegetables

QU Guang-fan1,2, BA Liang-jie1,2, WANG Rui1,2, MA Chao1,2, JI Ning1,2, CAO Sen1,2

(1.School of Food and Pharmaceutical Engineering, Guiyang University, Guiyang 550005, China; 2.Guizhou Engineering Research Center for Fruit Processing, Guiyang 550005, China)

The work aims to introduce the effects of melatonin on physiological and nutritional quality, energy metabolism, flavor change, chilling injury regulation and disease resistance of postharvest fruits and vegetables, so as to provide theoretical reference for the application of melatonin in postharvest preservation of fruits and vegetables. The effects of melatonin on postharvest preservation of different fruits and vegetables at home and abroad were reviewed. Melatonin could regulate the ripening and senescence of fruits and vegetables, improve the cold resistance and disease resistance, and maintain higher nutrients and better physiological quality. Melatonin can significantly prolong the postharvest storage period of fruits and vegetables, and has broad development space in the field of postharvest preservation of fruits and vegetables in the future.

melatonin; fruits and vegetables; preservation effect; research progress

TS255.3

A

1001-3563(2022)07-0045-07

10.19554/j.cnki.1001-3563.2022.07.006

2021-07-25

貴州省科技計(jì)劃（黔科合支撐[2020]1Y137號）；貴州省科技計(jì)劃（黔科合基礎(chǔ)-ZK[2021]一般173）；貴陽市科技計(jì)劃（筑科合同[2020]37-2號）

瞿光凡（1997—），男，貴陽學(xué)院碩士生，主攻農(nóng)產(chǎn)品貯藏與保鮮。

曹森（1988—），男，碩士，貴陽學(xué)院教授，主要研究方向?yàn)檗r(nóng)產(chǎn)品貯藏與保鮮。

責(zé)任編輯：彭颋