梨果肉质地及香气研究进展

徐钰清,田路明*,曹玉芬*,董星光,张 莹,霍宏亮,齐 丹,徐家玉,刘 超

(中国农业科学院果树研究所·农业农村部园艺作物种质资源利用重点实验室,辽宁兴城 125100)

摘 要:梨因其独特的口感和风味而备受消费者的喜爱。果肉质地和香气是梨果实重要的经济性状,针对梨果肉质地和香气研究的相关报道较多,且取得了较大的研究进展。围绕梨果肉质地和香气研究进行了概述,重点对软肉型梨、脆肉型梨果肉质地和香气形成机制,以及对影响梨果肉质地和香气形成因素的研究成果进行了综述分析,展望了果肉质地与香气研究的方向,以期为梨果肉质地评价研究、果实香气研究和培育不同香型梨果新种质提供参考。

关键词:梨;果肉质地;香气

梨是最重要的温带果树之一,世界范围内栽培较广泛,基于地理分布梨一般被分为东方梨种群和西方梨种群,大多数西方梨栽培品种果肉具有质地软溶、多汁和香气浓郁的特点,而大多数东方梨栽培品种果肉质地清脆,汁液较多但香气较少[1]。原产我国的栽培梨品种有4 个种,分别是白梨、砂梨、秋子梨和新疆梨,秋子梨与东方梨其他品种不同,是一种典型的呼吸跃变型水果品种,但它与西洋梨栽培品种有相似之处,都需经后熟软化果实变软后才可食用,并且在成熟期间会产生大量的乙烯和浓郁的香气[2]。果实的品质通常由颜色、形状、质地和香气等因素决定。其中,果肉质地是衡量果实品质的一个重要指标,被用来直接表示果实的组织状态、口感和风味[3];香气是影响果实最终感官品质以及消费者满意度的重要因素,果实中的香气代表了许多挥发性化合物的复杂混合物,包括酯类、醇类、醛类、酮类、萜类、苯类、烃类、酸类等[4],而挥发性酯是梨果肉香气的主要贡献者,不同品种中不同浓度的挥发性酯决定了其感官属性和消费者偏好[5]。果肉质地和香气是决定梨果在水果市场竞争力的主要因素,直接影响梨果的经济效益,笔者在本文中阐述了梨果肉质地类型的分类和香气的研究概况,概述了软肉型梨和脆肉型梨果肉质地研究和香气形成的机制研究,综述了近些年影响梨果肉质地及香气形成的研究进展,探讨了梨果肉质地与香气形成的相关影响因子,展望了相关的研究方向,旨在为开展果肉质地研究、香气研究和培育富有香气梨种质研究提供参考。

1 梨果肉质地的研究

1.1 梨果肉质地的类型

梨果肉质地是一个综合性指标,能够直接反映果实的口感[6],主要包含硬度、弹性、凝聚性、回复性、咀嚼性、破裂力、破裂位移、黏附性等质地参数,这些参数在一定程度上反映了梨果肉的质地特性[7]。不同梨品种之间的果肉质地有很大差异,一般梨属种质资源按果肉质地可分为软肉型和脆肉型,其中大多数软肉型梨果较不耐贮藏,在刚采收时果肉粗硬、口味生淡或酸涩,食用品质差,但经后熟果实变软,肉质细腻多汁,香甜适口;而大多数脆肉型梨果实脆嫩多汁,贮藏时间比较长[3]。西洋梨和秋子梨品种为软肉型梨果,要经过后熟软化才能食用[8],如康佛伦斯(Conference)、阿巴特(Abbé Fetel)、南果梨、京白梨等,而白梨、砂梨和新疆梨栽培种为脆肉型梨果,不用经过后熟即可食用,食用品质较好[8],如砀山酥梨、鸭梨、丰水、黄金梨、库尔勒香梨等。此外,一些软肉型梨和脆肉型梨杂交的新品种的果肉质地表现出种间杂合的过渡类型,苏艳丽等[9]的研究发现,红宝石为白梨、砂梨和西洋梨的杂交后代,常温货架期较短,两周内会松软失去商品价值。另外,安萌萌等[10]在野生秋子梨上的研究发现,果实的硬度随着贮藏时间的延长而逐渐降低,在常温贮藏20 d后果肉软化,汁液丰富,鲜食品质最佳。

1.2 软肉型梨果肉质地机制研究

果胶是细胞壁的重要组成部分,梨果实的软化与贮藏和成熟过程中细胞壁果胶多糖的解聚和溶解以及细胞壁相关修饰酶的活性有关,可通过影响果胶多糖的解聚和溶解以及细胞壁相关修饰酶的活性来调节梨果实的软化进程[11-12]。Dong等[13]的研究发现,考密斯(Comice)和安久(d’Anjou)在贮藏5个月后,两个梨品种果肉均具有软肉多汁的质地且水溶性果胶(WSP)的含量和果胶甲酯酶(PME)的活性较高,但梨果肉硬度降低,使梨在成熟过程中形成软肉质地。梨果实中难溶性果胶在细胞壁相关修饰酶的作用下降解、水解成可溶性果胶,导致细胞间的黏合力下降,细胞结构受损和离散,从而使果实质地软化,硬度下降[8,13]。齐秀东等[8]在京白梨上的研究发现,在贮藏期间,β-半乳糖糖苷酶(β-GAL)和α-阿拉伯呋喃糖苷酶(α-ARF)活性迅速升高,WSP 和离子结合果胶(ISP)含量呈上升趋势,而共价结合果胶(CSP)含量呈下降趋势,WSP、CSP 和ISP 含量均与硬度呈显著或极显著相关,且各果胶组分由高分子质量降解为低分子质量,促进了果实质地的软化。Zdunek 等[14]通过对Xenia 和康佛伦斯这2 个西洋梨细胞壁中的多糖结构进行研究,发现梨果实质地与细胞壁中多糖的纳米结构特征有关,较少降解、较厚和较多的分支的果胶分子,细胞间的黏附性更强,使果实硬度更高。Wei等[15]的研究表明,京白梨果实中的碱溶性果胶(NSP)含量随贮藏时间的延长迅速下降,而WSP 含量显著增加;多聚半乳糖醛酸酶(PG)的活性在梨果实成熟过程中迅速增加,且与果肉硬度呈负相关。霍宏亮等[16]在秋子梨上的研究发现,果肉的硬度与WSP 和ISP 的含量均呈显著负相关,而与CSP 的含量呈极显著正相关,即当WSP 和ISP的含量均上升而CSP的含量下降时,果肉硬度下降。

软肉型果实成熟的特点是果实软化、着色和变甜,主要由糖、有机酸、木质素、黄酮类化合物的代谢来调控,这些代谢过程均表现为果实成熟过程中差异蛋白(DEP)的上调和下调[17]。蔗糖是许多水果甜味和风味的主要贡献者,蔗糖酶、蔗糖合酶(SUS)和蔗糖磷酸合成酶(SPS)是深度参与梨果实蔗糖代谢的3种关键酶[18]。齐秀东等[19]的研究表明,蔗糖代谢参与了京白梨果实的后熟软化,酸性转化酶(AI)活性和基因表达的上调主要发生在呼吸跃变后期,参与了京白梨果实后期的软化过程;SPS 和蔗糖合酶(SS)随果实软化具有较高的活性和基因表达,可通过调控可溶性糖分的组成和含量来参与果实软化的生理过程。

1.3 脆肉型梨果肉质地机制研究

由于果实质地属性不同,脆肉梨与软肉梨果实在软化过程中果胶降解途径差异显著,使得两个品种梨果实具有不同的后熟软化和贮藏特性。齐秀东等[8]在鸭梨的研究上发现,鸭梨果实在发育期间,梨果实中的CSP、ISP 和WSP 含量逐渐增加;在贮藏期,鸭梨果实中的CSP 含量较高且保持稳定,WSP含量增加缓慢;相关性分析表明,在果实发育期的WSP、CSP、ISP 含量均与果实硬度呈显著负相关。Wei 等[15]的研究表明,鸭梨果实中的NSP 含量几乎没有发生变化,随着贮藏时间的延长,WSP 含量逐渐增加,而CSP 含量稍有下降;而α-ARF 活性在贮藏过程中非常低且略有增加。

在脆肉梨果实中,与果肉质地相关蛋白在梨果实发育和成熟过程中的差异表达影响果实的品质、质地和口感[20]。苯丙烷途径中的木质素生物合成与梨果实中木质素的形成有关,对石细胞含量的增加起着主导作用[21],而石细胞的含量以及石细胞团的大小和密度决定着梨果实的口感和质地[22- 23]。Cheng等[24]研究了LAC基因家族在砀山酥梨果实木质素代谢中的作用,发现PbLAC1、PbLAC6、PbLAC29 和PbLAC36 在果实不同发育阶段的表达变化与石细胞的形成一致,说明这4 种基因在果实木质素合成和石细胞发育中发挥着重要作用,影响果实的硬度。

2 不同果肉质地类型的梨果实香气研究

2.1 梨果实香气研究概况

果实香气能客观反映不同水果的风味特点,是衡量果实品质的重要指标之一。目前,在梨果实中已经鉴定出300 多种挥发性物质,包括酯类、醇类、醛类、烯烃类、酮类、酸类等,果实中香气成分的种类、含量及香气成分种类的多少直接决定着果实的香味[25]。梨果实中的香气成分根据人的感官分类,可分为果香型、青香型和醛香型等[26]。白梨和砂梨果实中的香气成分主要是乙酸己酯和己醛,因此白梨和砂梨果实具有果香的同时还具有一定的青草香[27]。秦改花[28]的研究发现,红安久、五九香、南果梨和福安尖把具有浓郁的果香味;库尔勒香梨具有果香和花香;而丰水和砀山酥梨等具有较淡的青香味。此外,一些野生品种也具有其独特的香味,Prakash 等[29]的研究发现,印度野生类型Kainth(Pyrus pashia Buch.-Ham. ex D. Don)梨果中挥发性化合物以酯类和烷烃类物质为主,具有类似于柑橘类水果的果香味、甜味和玫瑰香味。

梨果实香气物质的合成代谢由多途径共同调控,其中起到决定作用的酶主要有脂氧合酶(LOX)、丙酮酸脱羧酶(PDC)、氢过氧化物裂解酶(HPL)、醇脱氢酶(ADH)和醇酰基转移酶(AAT)等,由于前体物质的不同,从而产生不同的香气物质[30]。目前,常通过固相微萃法(SPME)、顶空固相微萃取法(HSSPME)或同时蒸馏萃取法(SDE)等[31]与气相色谱-质谱联用技术(GC-MS)联用的方法对梨果实中的香气物质进行提取和鉴定。张文君等[32]联用SPME与GC-MS技术,对不同种间杂交梨果的香气成分进行分析鉴定。李国鹏等[33]应用HS-SPME 与GC-MS技术联用的方法分析了油红梨果实后熟的不同阶段香气成分的变化。

2.2 软肉型梨果实香气机制研究

脂肪酸代谢的LOX 途径是参与梨果实挥发性物质形成的最基本途径,可通过调控与酯类合成相关基因的表达来调节梨果实成熟过程中挥发性物质的生物合成。Luo 等[34]在南果梨上的研究发现,在采后成熟过程中,果实中PuADH3PuLOX3PuAAT 的表达水平与总酯、主要酯类物质的含量呈正相关,是调节南果梨采后成熟过程中酯类合成的关键基因。Li 等[2]研究表明,1-MCP 抑制了软儿梨果实中PuLOXsPuAAT的转录水平的升高以及果实在贮藏期间酯类浓度的增加,说明PuLOXs 和PuAAT对梨果实中酯类的形成至关重要。Li等[35]发现苹香梨果实中与酯生物合成相关的基因在果实成熟过程中表现出不同的转录模式,PuLOX1PuLOX8PuAAT 的转录水平在成熟期间第3 天达到峰值,同时丁酸乙酯、2-甲基丁酸乙酯、己酸乙酯和乙酸己酯的浓度均在增加,因此PuLOX1PuLOX8PuAAT 对酯的形成至关重要。Li等[36]分析了南果梨脂肪酸代谢相关基因的差异表达,发现在采收10 d后果实中PuFAD2.1PuLOX2-1PuLOX5PuADH1PuAAT的转录水平较高,且均与油酸、亚油酸(LA)和亚麻酸(LNA)的含量呈正相关,表明挥发性酯的形成与这些基因的表达以及芳香挥发物前体的积累有直接关系。脂肪酸去饱和酶(FAD)在油酸转化为LA 和LNA 方面发挥着关键作用,FAD 基因表达水平的提高可以显著增加LA 和LNA 的含量,增强果实对低温的抗性[37-38]。Zhou 等[37]的研究发现,Pu-FAD2PuFAD3 的表达与不饱和脂肪酸的含量呈显著正相关,因此这两个基因表达水平的显著上调可提高南果梨果实中酯类物质的含量。Shi 等[38]在南果梨上的研究发现,脂肪酸含量和脂肪酸代谢中关键基因的调控对冷藏果实香气酯的增加至关重要,fadD、fadE、fadJ 和atoB 的下调减缓了冷藏果实中棕榈酸和硬脂酸的降解。Wei 等[39]的研究发现,在南果梨采收前喷施钙肥,导致编码丙酮酸脱羧酶同工酶1(PDC1)、脂肪酶、LOX 相关基因的表达显著上调,使3 个编码FAD 的基因差异表达,说明PDCs和FAD可能在香气代谢产物和释放过程中起重要作用。

植物激素在香气的代谢过程中发挥作用,植物激素信号转导基因在香气形成过程中差异表达,影响着梨果实香气物质的合成。Luo 等[40]的研究表明,总酯含量的增加与乙烯生物合成直接相关的基因PuACO1PuACO2PuACS 的表达呈正相关,经甘氨酸甜菜碱(GB)处理的果实在货架期内乙烯合成基因的表达上调,说明GB 处理促进了乙烯的生物合成,进而促进了酯类物质的积累。Zhou等[41]研究了南果梨果实中乙烯信号转导相关基因表达的变化,结果表明,在货架期的第0天至第18天经间歇升温(IW)处理的果实中PuACS1和PuACO1的表达显著高于冷藏处理的果实;而PuACS1 和PuACO1 更高的翻译水平有利于促进乙烯生成,影响果实香气物质的形成。Shi 等[42]发现南果梨果实在冷藏后的货架期内,参与植物激素信号转导的基因DELLASnRK2PR-1 的表达在香气形成过程中上调,AUX1IAAARFGID1TFPYR/PYLNPR1 的表达在香气形成过程中下调,说明植物激素信号的变化影响了冷藏中果实正常的生理代谢。

转录因子通常在生物过程中的转录水平上调控基因表达,编码转录因子基因的表达在香气形成过程中存在显著差异。Wei等[39]在南果梨上的研究表明,分别编码MADS-box转录因子、AGL6、AGL9的3 个基因Pbr022012.1Pbr007181.1Pbr018801.2在香气散发过程中表达特异性上调,说明MADS-box转录因子、AGL9和AGL6可能调控香气释放。Luo等[43]的研究表明,过表达的PuMYB308-likePuMYB6-like促进了PuAAT的表达,提高了南果梨果实中丁酸乙酯和乙酸己酯的含量;此外,增强PuMYB308-likePuMYB6-like 表达的同时,抑制PuNAC68-like 的负调控,可以促进果实中酯类物质的形成和积累。Li等[36]的研究发现,与采收后0 d相比,南果梨果实中PuWRKY24 和PuAUX/-IAA29 的表达在采收后10 d时显著上调,而AP2/ERF家族的PuTINY 的表达显著下调,说明PuWRKY24、PuIAA29 和PuTINY 等转录因子对调控南果梨香气物质的形成起重要作用。Luo 等[44]的研究表明,经茉莉酸甲酯(MeJA)处理的南果梨在最佳食用期,PuMYB108-likePuWRKY61PuWRKY72 的表达显著上调;且与PuLOX3PuADH5PuADH9PuAAT的表达呈显著正相关,表明它们在挥发性酯类物质的积累中起重要作用。Yao等[5]的研究发现,低温调节(LTC)处理可以提高PuERS1PuEIN4PuEIN2PuERF 等乙烯信号转导基因的表达水平,促进果实在冷藏和货架期内芳香挥发性化合物,说明LTC 减少了冷藏后果实香气相关挥发性物质的损失。Luo等[40]的研究发现,南果梨中总酯含量与乙烯信号转导相关基因PuERF015PuERF114PuRAV1-likePu-ERF113-likePuERF2PuERF071-likePuERF109-like的变化呈正相关,经GB处理的果实在货架期内乙烯信号转导相关基因的表达上调,增强了乙烯信号转导,从而提高了果实中酯类物质含量。

2.3 脆肉型梨果实香气机制研究

LOX 基因家族参与果实中醛类和醇类化合物的合成,在梨果实香气挥发性物质的合成方面发挥着重要作用,影响了果实香气的形成。Li 等[45]在转录组水平分析了白梨果实发育过程中LOX 基因的表达,发现白梨果实的LOX 基因在发育前期(开花后15 d)表达水平较低,发育中期(开花后80 d 或110 d)表达水平达到高峰,近熟期(开花后145 d)表达水平降至0,随后在开花后167 d 略有升高,表明大多数LOX基因具有功能活性。

ADH对梨果中挥发性化合物的形成至关重要,ADH基因家族参与果实挥发性化合物的生物合成,且与果实成熟过程中醇类的形成密切相关。Zeng等[46]通过RNA-Seq 和qRT-PCR 分析发现砀山酥梨果实中的3 个ADH 基因(Pbr013912.1Pbr026289.1Pbr01252.1)在梨果实成熟期催化醛转化为醇,在果实发育早期的表达水平较高,在发育后期表达水平迅速下降,说明这3个ADH基因的表达水平与果实发育过程中芳香族化合物的含量密切相关。

BAHD 酰基转移酶超家族具有调节果实品质[47],催化合成酯类、萜烯类和酚类物质等作用。Liu 等[48]对7 种蔷薇科植物的BAHD 超家族进行全基因组比较分析,发现从7 个蔷薇科物种的基因组中获得的717个BAHD基因中有114个属于白梨系品种砀山酥梨,通过qRT-PCR 分析发现Pbr020016.1Pbr029551.1Pbr019034.1Pbr014028.1 是砀山酥梨果实发育过程中参与香气形成的重要候选基因。

3 影响梨果肉质地和香气的因素研究

3.1 品种

不同梨品种间的果肉质地存在一定的差异,王斐等[6]对2个脆肉梨品种华酥和圆黄研究发现,在同一成熟度圆黄梨果实各部分的脆度、果肉硬度、胶黏性、咀嚼性均显著高于华酥。王燕霞等[7]通过对14个品种梨果实的果肉质地进行测定发现,梨果肉质地各参数指标中果肉硬度、破裂力和脆度等对梨果肉质地的影响较大。苏艳丽等[9]的研究表明,东方梨间的杂交后代品种的果皮硬度、韧性和果肉硬度均在货架前期缓慢降低,无显著差异且货架期较长;而西洋梨与东方梨的杂交后代品种的果皮硬度、韧性和果肉硬度均在货架前期快速降低,有显著差异且货架期较短。张文等[49]对黄金、玉冠和丰水3 个砂梨品种在贮藏过程中的质地变化进行研究,发现这3 个品种梨果实的泰勒(MT)硬度、果肉硬度、果肉弹性率均随贮藏时间的延长而逐渐下降。高海生等[3]的研究发现,鸭梨在常温下贮藏60 d 后果实的硬度、破裂力和内聚性变化不大;而京白梨果实硬度和破裂力均随着贮藏时间的延长而迅速降低,在常温下贮藏18 d左右果实完全软化。Pham等[50]通过2次压缩试验对2个砂梨品种的果实质地进行对比分析,发现新高果实的硬度、脆度、咀嚼性、弹性模量和断裂应变均高于丰水,这2 个梨品种果实中部的硬度和脆度均略高于顶部和底部。

不同品种间梨果实中香气物质的种类、相对含量以及特征香气成分的种类也有较大差异,因此不同品种的梨果具有不同香味。田长平等[27]的研究表明,鸭梨、雪花梨、茌梨这3个白梨品种的香气总量、醛类物质相对含量、己醛和乙酸己酯含量均明显高于黄金梨、圆黄、晚秀这3个砂梨品种。Yi等[51]的研究发现,2-甲基丁酸乙酯和己酸乙酯是地方梨品种香面梨的主要香味物质,其香味浓郁,且与南果梨接近,可作为潜在亲本基因型进行白梨的香气改良育种。Taiti等[52]的研究表明,与丰水和鸭梨相比,威廉姆斯(Williams)果实中萜烯、倍半萜烯和丁醛的含量最高,所含的挥发性化合物的种类和含量最多。Chen 等[1]对12 个西洋梨成熟果实的香气浓度和香气成分进行了比较分析,发现好本号(Alexandrine Douillard)总香气浓度最高,而红巴梨(Bartlett-Max Red)最低;酯类是西洋梨果实中最重要的挥发性有机化合物,酯类的数量在博斯克(Beurre Bosc)中最多,而在茄梨(Clapp Favorite)中最少。Qin等[53]对33个秋子梨品种的香气成分进行了测定比较,发现热梨、锦香、红八里香、白八里香和伏五香等品种的酯类含量高,烃类含量低;秋香、福安尖把、龙香、官红宵、山梨24 和五香梨等品种的烃类含量高,酯类含量低;砂糖梨和满园香果实中醛类浓度较高。乌云塔娜等[54]研究发现,安乔梨和秋子梨品种的香气成分主要为烯烃类和酯类,但安乔梨中存在芳烃类物质,秋子梨品种中不含芳烃类物质。刘婉君等[4]研究了18个梨品种授粉对鸭梨果实香气成分的影响,发现与雪花梨授粉的果实相比,南水、库尔勒香梨、Crispel、绿宝石和翠玉授粉的果实中酯类物质的相对含量显著增加,说明不同的品种授粉可以显著改善鸭梨果实的香味。向思敏等[55]对不同倍性库尔勒香梨果实中香气成分和相关代谢物质进行比较分析,发现三倍体香梨果皮中挥发性物质的总含量高于普通二倍体香梨,但种类少于二倍体香梨。向思敏等[56]的研究表明,杂交后代新梨7 号果实中的挥发性物质的种类与父本早酥和母本库尔勒香梨相比有所减少;其主要挥发性物质与亲本相似,挥发性物质的总含量高于父本,但低于母本。刘园等[57]对库尔勒香梨挥发性物质进行分析发现,(E)-2-己烯醛、己醛、α-法呢烯、1-壬醇、乙酸己酯是其主要挥发性物质,且香梨果皮比果肉含有更丰富的挥发性物质。安萌萌等[58]的研究发现,野生秋子梨含有的香气种类显著多于小香水、京白、南果梨这3个栽培品种,野生秋子梨和栽培品种中均含有醇类、酯类、烃类、醛类、酮类、杂环类、醚类等化合物,但酸类和烃衍生物类化合物仅在野生秋子梨中存在。

3.2 成熟度

采收的成熟度对梨果肉质地会产生一定程度的影响,适期采收可使果实达到应有的外观和风味。王斐等[6]对不同成熟度华酥和圆黄进行果肉质地研究,发现在成熟前期这2 种脆肉梨果实不同部位的果肉脆度、硬度、胶黏性和咀嚼性均存在差异,萼部最大,中部次之,梗部最小,且随着成熟度增加,差异显著性逐渐减弱。Hendges等[59]的研究发现,康佛伦斯和卢卡斯(Alexander Lucas)在第一成熟期采收后果实未软化,果皮颜色未变黄;第二成熟期采收的果实具有更多的芳香挥发性物质;延迟采收的果实中乙烯产生量和呼吸频率显著增加,果肉硬度降低和果皮颜色变浅。齐秀东等[8]在京白梨上的研究发现,果实的硬度和破裂力在贮藏期间呈下降的趋势,而弹性、咀嚼性、黏附性和内聚性在果实完成后熟后显著增加。贾晓辉等[60]的研究表明,随着采收成熟度的增加,玉露香果实中可溶性固形物含量、可滴定酸含量和维生素C 含量不断增加,而果实硬度不断下降。阚超楠等[61]在翠冠上的研究发现,随着货架期的延长,果实的硬度、脆度和咀嚼性逐渐降低;较早采收能使果实保持较好的果实色泽和质地品质;延迟采收的果实成熟度较高,易发生果皮褐变和质地软化的现象,影响果实在货架期间的品质。

梨果实在不同成熟阶段香气成分的种类、含量和特征香味物质也会有所不同。廖凤玲等[62]对爱甘水梨不同发育期果实中的香气物质研究发现,随着果实的生长发育,酯类物质的含量逐渐增多,醛类物质的含量呈现出先增后降的变化趋势,醇类物质的含量呈现出“M”形的变化趋势。秦改花等[63]的研究表明,不同采收期南果梨果实中特征香气和果实香味有所不同,晚采收的果实中比早采收的果实中多了乙酸丁酯和2-己烯醛两种特征香气;晚采收的果实比早采收的果实果香味浓郁,而早采收的果实中具有青香味。Zlatic 等[64]分别在商熟期和商熟期后7 d 对巴梨(Bartlett)进行采收,发现在成熟过程中,果实中的乙醛含量增加,在商熟期后7 d采收的果实中的乙醛含量较高。张军等[65]对不同时期库尔勒香梨香气合成相关酶活性进行测定,发现LOX、ADH和β-葡萄糖苷酶(CB)的活性均呈先升高后降低的趋势;AAT 活性在果实发育前期变化不稳定,但随着果实的成熟逐渐升高,在9月达到最大值,随后下降至平缓。Chen 等[66]对库尔勒香梨果实中挥发性物质的产生进行研究,发现随着果实的成熟,醛类和醇类的浓度显著增加,在商熟期形成的高浓度醛和醇为香梨提供了特有的青香味。

3.3 采后处理

果实的采后处理技术可显著提高果实在贮藏期间和货架期间的品质,延长果实的货架期,以达到长期贮藏的目的。张雪等[67]的研究表明,经F-MAP保鲜处理的苹果梨果实中可溶性果胶含量呈上升趋势,但始终低于纸箱贮藏,说明F-MAP 保鲜贮藏效果好于纸箱贮藏,可有效延缓梨果实品质的下降。王聪等[68]的研究表明,花椒精油处理抑制了苹果梨贮藏后期果实胶黏性和黏着性的升高以及果实硬度和脆度的下降,维持了果实的质地品质。Zhou等[37]的研究表明,南果梨经间歇升温(IW)处理后,没有缩短果实的货架期,但在货架期结束时加快了果实的软化进程,果实硬度显著降低。Ferreira等[69]的研究表明,晒干处理显著地降低梨果实的硬度和脆度,提高梨果实的凝聚性、弹性和黏附性。梨果实采后经1-MCP 处理可抑制梨果实的软化以及贮藏过程中乙烯和其他挥发物的产生,显著延缓果实的衰老和劣变,维持果实的质地品质。王晓曼等[70]的研究表明,1-MCP延缓了黄金梨果实在常温贮藏下果肉脆度、硬度、凝聚性和咀嚼性的降低,能较好地延缓果肉质地的劣变,维持果肉的质地品质。Rizzolo等[71]的研究表明,1-MCP 显著降低了阿巴特果实在冷藏期间的乙烯产生量,延缓果实的果皮黄化和果实软化,抑制叶绿素含量的降低,但有时经1-MCP处理的果实在货架期仍保持较大的硬度。

由于采后处理方式的不同,梨果实中香气物质的种类和含量存在一定的差异。Zhou 等[72]研究了LTC 处理对南果梨在常温贮藏期间香气的影响,结果表明,经LTC 处理的果实具有更高水平的乙烯产生量和呼吸速率,AAT、LOX、ADH、1-氨基环丙基-1-羧酸合成酶(ACS)和1-氨基环丙基-1-羧酸氧化酶(ACO)的活性显著升高,果实中酯类含量以及LA和LNA 含量显著增加。Yao 等[5]也得出相同的研究结果,说明LTC 处理可有效缓解南果梨在长期冷藏过程中与香气相关酯类的损失,延长南果梨的货架期。Zhou等[41]在南果梨上的研究发现,IW处理的果实在冷藏后货架期内香气相关酯类和总酯的含量均显著增加,促进了果实中LA和LNA的积累,为酯类物质的合成提供更多的底物。Zhou等[37]研究发现,南果梨经IW 处理可延缓果实在冷藏90 d 后货架期内香气相关酯类的减少,PuLOX1PuADH3PuAAT1的表达水平显著上调,说明它们在南果梨香气相关酯的合成中发挥着重要作用。Chen 等[73]的研究表明,鲜梨与冻梨中的香气成分表现出显著的差异,冻梨表现出浓郁的甜味和香味,反复的冷冻处理使梨产生了独特的香气,有助于梨新产品的研发。Jiang等[74]发现冷冻干燥(FD)保留了鲜梨果实的特征香气,具有更高的挥发性化合物相对含量;热风干燥(HD)处理损失了部分与鲜梨相关的挥发性化合物,但癸醛、苯甲醛、壬酸、乙酮等挥发性化合物含量增加,使果实具有甜味和浓郁的香气。此外,由于梨果实贮藏前施用的调节物质不同,果实中的香气成分在贮藏期间和货架期存在一定差异。Li等[2]在软儿梨上的研究发现,果实在采后冷藏过程中,1-MCP抑制了酯类物质浓度的增加;在采后贮藏期间的各个阶段,1-MCP处理果实中挥发性物质的总浓度均低于未经处理的果实。张丽萍等[75]研究了外源乙烯对1-MCP 处理南果梨香气代谢的影响,发现2 mmol·L-1外源乙烯处理提高了LOX、HPL和AAT的活性,提高了冷藏后南果梨果实在货架期间的香气品质。Chen等[66]对库尔勒香梨的果肉进行体外LA和LNA的喂养试验,表明经过处理的果实中挥发性酯的浓度在培养12 h后显著升高,同时伴随着醛类和醇类浓度的降低。Liu 等[76]的研究发现,褪黑素(MT)处理降低了库尔勒香梨和阿巴特梨在贮藏过程中果实己醛和(E)-2-己烯醛的含量;促进了阿巴特果实中PbAAT 基因表达,增强AAT 的活性,增加了乙酸丙酯和乙酸己酯的含量。Luo等[43]在南果梨上的研究发现,水杨酸(SA)处理增加了冷藏的南果梨中酯类物质的种类和含量,促进了乙烯的合成,减少了冷藏南果梨的香气损失。Luo等[40]的研究发现,GB处理可促进南果梨果实中酯类物质的积累,显著增加果实中的酯类物质的种类和含量,有效缓解了南果梨在长期冷藏后香味的损失,提高了果实的香味品质。Luo 等[44]的研究表明,MeJA 处理增加冷藏后货架期南果梨酯类物质含量,提高LOX、ADH 和AAT 的活性,促进果实酯类物质的合成,提高冷藏南果梨品质。

3.4 贮藏条件

此外,贮藏方式和贮藏时间的不同也会对果实的质地产生不同的影响。Rizzolo等[71]的研究表明,与在-0.5 ℃下贮藏的果实相比,阿巴特在1 ℃贮藏下果实的硬度、脆度和破裂能较低,但乙烯的产生速率较高;动态气调保鲜贮藏(DCA)的果实色度、硬度和乙烯产生速率均高于气调贮藏(CA)和常温贮藏(NA),但弹性低于CA 和NA。Dong 等[13]的研究发现,与贮藏5个月相比,考密斯果实在贮藏8个月后成熟过程中乙烯产生量较低且呼吸速率较高;PG和果胶酸裂解酶(PL)活性显著升高,PME、β-GAL和α-ARF 活性降低;安久果实在贮藏8 个月后成熟过程中乙烯产生量和呼吸速率显著提高;PME和PL的活性升高,而PG、纤维素酶(CEL)、β-GAL 和α-ARF的活性降低。娄来峰等[77]对不同贮藏期库尔勒香梨的质地进行分析,发现果肉硬度、脆度、弹性、黏聚性、咀嚼性、回复性随贮藏时间增加逐渐下降,而附着性随贮藏时间增加逐渐上升。

3.5 栽培管理

果树在生长发育过程中,一些栽培管理技术,如套袋、肥料种类和用量等,都会对果实香气物质的种类及相对含量产生一定的影响。果实套袋是我国果品生产中常用的一项技术措施,不同类型的果袋对果实香气物质的组成和含量有不同的影响。魏树伟等[78]在秋洋梨上的研究发现,与不套袋相比,套袋降低了果实香气物质的总含量,且套单层袋果实香气物质的含量高于套双层袋。李芳芳等[79]的研究表明,套袋减少了库尔勒香梨果实香气物质的种类,尤其会使酯类物质的合成受到抑制,不利于香气物质的合成;但与单层白色纸袋相比,紫色塑料膜袋对果实香气物质的影响较小。

不同种类的肥料和用量也是造成梨果实中香气物质种类和含量差异的重要因素。童盼盼等[80]的研究表明,施用不同类型的有机肥和菌肥均使库尔勒香梨果实中增加了9-癸烯醇、2,6,10-三甲基-9-十一烯醛、壬癸烷三种香气物质,此外有机肥和菌肥均提高香梨果实香气物质和特征香气的含量,其中海藻酸冲施有机肥处理的香气物质种类及含量均高于其他施肥处理。Wei 等[81-82]在南果梨采收前15 d 喷施氯化钙发现,在采后成熟过程中,葡萄糖苷酶、ADH、PDC 和LOX 的活性有所提高,促进了果实乙醇和乙醛的合成,提高了酯类物质的含量;而且钙处理增加了亚油酸和亚麻酸的相对含量,为酯类的合成代谢提供了更多的前体,从而提高了果实挥发性香气物质的含量。此外,南果梨经钙处理后的果实果皮蜡质层脱落较多,果皮蜡质层中酯类、脂肪酸和醛类的相对含量和总酯含量显著高于未经处理的果实,促进了果实香气物质的释放,使果实香气更加浓郁[83]。童盼盼等[84]的研究发现,在库尔勒香梨生长发育季节喷施特定微量元素“优多锌”和“优多硼”可提高香梨果实特征香气含量,其中经“优多锌”处理后的香梨果实中香气含量最高。

4 结论与展望

通过对比梨果肉质地和香气的研究发现,梨果肉质地有软肉型、脆肉型和种间杂合的过渡型之分。跃变型梨品种随着果实成熟乙烯产生量和呼吸速率高,而非跃变型梨品种乙烯产生量和呼吸速率较低,秋子梨作为重要的亚洲梨品种,它经过后熟后果实才能软化、可食用,随着果实成熟,它在产生大量乙烯的同时产生了浓郁的香气[35]。软肉型梨和脆肉型梨果肉质地和香气因不同因素的影响造成不同的差异,其中梨果肉质地和果肉中的香气物质因品种、成熟度、采后处理方式的不同而存在一定的差异;与脆肉型梨相比,软肉型梨在贮藏期间的果肉硬度、破裂力等质地参数指标较低,但所含的香气物质的种类和相对含量较高,香气更为浓郁。梨果采后经1-MCP处理显著抑制了果实的软化,达到了长期贮藏的目的,但降低了果实中挥发性物质的种类和浓度,通过提高贮藏期间的温度或在贮藏前施用外源调节物质,如乙烯、水杨酸、甘氨酸甜菜碱等,可有效减少果实香味的损失,维持果实的品质。

目前,国内外关于梨果肉质地的研究主要集中在果实成熟软化过程中细胞壁结构和物质的变化,以及细胞壁相关修饰酶活性的变化;而关于梨果实香气的研究大多集中于酯类合成相关基因、植物激素信号转导基因和编码转录因子的基因在香气形成过程中的差异表达。研究表明,脆肉梨果实和软肉梨果实在果肉质地和香气成分上存在较大差异,软肉型梨和脆肉型梨的果肉质地和香气存在差异的机制还需更全面深入的比较研究,调控不同类型梨果肉质地变化和香气形成的关键基因以及挥发性物质释放的调控机制等还有待进一步解析。

果实的质地和香气是决定水果市场价值的重要因素,随着人们生活水平的提高,培育出货架期长、香味浓郁的优质果实成为种植者和消费者的目标。为提升梨果实的品质,满足人们对高质量梨果品的要求,要着眼于培育不同果肉类型且富有浓郁香气的梨品种。一方面要加强梨果肉质地与香气形成差异的机制研究,挖掘出与梨果肉质地变化和香气形成相关的关键基因,逐步探索研究果实香气释放的机制和调控机制,从而为产生香气和调控香气提供理论依据;另一方面,利用基因编辑技术在分子水平上进行香气形成精准调控研究,控制梨果实香气物质的形成、挥发性物质释放量和释放时间,为创制出不同香型的梨果实提供理论支撑,比如玫瑰香型梨果。不断加强梨果肉质地和香气形成的机制研究,结合利用最新的基因工程技术,进行原始创新利用,从而培育出不同特色、不同风味且具有更多香气成分的软肉型梨和脆肉型梨新种质,以期促进我国梨产业高质量健康发展。

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Research progress in flesh texture and aroma of pears

XU Yuqing,TIAN Luming*,CAO Yufen*,DONG Xingguang,ZHANG Ying,HUO Hongliang,QI Dan,XU Jiayu,LIU Chao
(Institute of Pomology, Chinese Academy of Agricultural Sciences/Key Laboratory of Horticultural Crop Germplasm Resources Utilization,Ministry of Agriculture and Rural Affairs,Xingcheng 125100,Liaoning,China)

Abstract: Pears are popular by consumers because of their unique taste and flavor. Flesh texture and aroma are important economic characteristics of pear fruits. There are many reports on the research of flesh texture and aroma of pears, and more research progress had been made. First of all, the results of research on flesh texture and aroma of pears are summarized in this paper.According to the flesh texture,pear germplasm resources can be divided into melting-flesh type,crisp-flesh type and interspecific hybrid type.Among them, Pyrus bretschneideri, P. pyrifolia and P. sinkangensis are planted and cultivated as crisp-flesh pear fruits, which are crisp and juicy, with good eating quality, while P. communis and P. ussuriensis were melting-flesh pear fruits, which could be eaten only after ripening and softening.The flesh texture of fruit is often directly used to reflect the taste of the fruit,and aroma is an important factor affecting the sensory quality of the fruit.The volatile substances in pear fruit mainly include esters, alcohols, alkenes, ketones and acids. The type and content of these volatile compounds determine the flavor of the fruit and consumer preferences.Secondly,the mechanism of flesh texture and aroma formation of melting-flesh pear and crisp-flesh pear are elaborated.The degradation pathway of pectin in the softening process of melting-flesh pear and crisp-flesh pear is significantly different, which leads to the different ripening and softening characteristics between two types of pear varieties. When the contents of water-soluble polyuronides(WSP)and ionic-soluble polyuronides(ISP)in melting-flesh pear fruits increase and the contents of CDTA-soluble polyuronides (CSP) decrease, the flesh hardness decreases;However,the content of WSP in crisp-flesh pear fruit increases slowly,the content of CSP is higher and decreases slightly with the extension of storage time,and the flesh hardness decreases slowly.In addition,sucrose metabolism is also involved in the ripening and softening of melting-flesh pears,which could be regulated by regulating the composition and content of soluble sugar.LAC gene family plays an important role in the lignin synthesis and stone cell development of Dangshansuli, which affects the fruit hardness.The aroma reflects the flavor characteristics of different pear varieties. It could affect the synthesis of aroma substances in melting-flesh pear fruits by regulating the expression of genes related to ester synthesis and the differential expression of plant hormone signal transduction genes and genes encoding transcription factors.In crisp-flesh pears,LOX gene family is involved in the synthesis of aldehydes and alcohols in the fruit.BAHD acyltransferase superfamily catalyzes the synthesis of esters,terpenes and other substances,The ADH gene family is closely related to the formation of alcohols during fruit ripening. Furthermore, the factors affecting the flesh texture and aroma formation are reviewed.The flesh texture and aroma substances in the fruit of pear are different due to different varieties,maturity and post-harvest treatment methods.There are significant differences in flesh hardness,brittleness, stickiness, chewiness and other texture parameters among different pear varieties’fruits.The hardness and breaking force of most soft flesh pears decrease rapidly with the extension of storage time, while the hardness and breaking force of most crisp flesh pears cannot change much with the extension of storage time. There are also great differences in the types and contents of aroma substances and the types of characteristic aroma substances among different pear varieties’fruits.Therefore,different pear varieties'fruits show different flavor type.The maturity of harvest also affects the flesh texture of pear fruit and the type and content of aroma substances in the fruit. Properly harvested pear fruits could show good fruit texture and quality, while delayed harvest fruits with higher maturity could lead to texture softening and short shelf life.But compared with the fruits harvested early,the fruits harvested late show more fragrant flavor,and the fruits harvested early have a green flavor.Postharvest 1-MCP treatment for pear fruits could significantly inhibit the deterioration and softening of the fruit, maintain the quality of the fruit during storage,but lead to the reduction of the type and concentration of volatile substances in the fruit. The loss of fruit flavor could be reduced and the quality of the fruit could be maintained by increasing the temperature during storage or applying ethylene,salicylic acid,glycine betaine and other exogenous regulators before storage.In addition,the flesh quality of pear fruits is affected by the different storage methods and storage time.And the types and relative content of aroma substances are affected by bagging,fertilization and other treatments during development of pear fruits.Finally, the study directions in flesh texture and aroma is suggested, in order to provide reference for the study of evaluation on flesh texture,fruit aroma and the cultivation of new pear germplasm with different aroma types.

Key words:Pear;Flesh texture;Aroma

中图分类号:S661.2

文献标志码:A

文章编号:1009-9980(2023)04-0757-14

DOI:10.13925/j.cnki.gsxb.20220487

收稿日期:2022-09-21

接受日期:2022-11-21

基金项目:国家现代农业产业技术体系(CARS-28-01);中国农业科学院科技创新工程(CAAS-ASTIP-RIP);国家科技资源共享服务平台项目(NHGRC2022-NH02-1)

作者简介:徐钰清,女,在读硕士研究生,研究方向为梨种质资源。Tel:15532996887,E-mail:xuyuqing0325@163.com

*通信作者Author for correspondence.Tel:0429-3598119,E-mail:tianluming@caas.cn;Tel:13942936219,E-mail:yfcaas@263.net