Research Progress on Synergistic Regulation of Endogenous Gamma-aminobutyric Acid Enrichment in Plants by Environmental Stress and Germination
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摘要: γ-氨基丁酸（γ-aminobutyric acid，GABA）作为一种新型功能因子，具有调节血压、缓解抑郁、治疗癫痫、延缓衰老和治疗糖尿病等功能，广泛分布在植物体内。研究证明植物发芽是可以提高内源性GABA含量的有效方式，结合不同的环境胁迫方式可以进一步促进发芽植物中GABA累积。本文主要阐述了植物富集GABA的代谢途径和影响因素，以及环境胁迫与发芽协同对植物内源性GABA富集的协同效应，以期为富含GABA植物基食品的开发与应用提供参考。Abstract: As a novel functional component, γ-aminobutyric acid (GABA) with many kinds of functions of regulating blood pressure, alleviating depression, treating epilepsy, delaying aging and treating diabetes, is widely distributed in a large variety of plants. It is reported that plant germination was recognized as an effective way to raise endogenous GABA levels, which in combination with several environmental stress modes could further promote GABA enrichment in germinating plants. This paper mainly discusses the metabolic pathways and influence factors of GABA enrichment in plants, as well as the synergistic effects of environmental stress and germination on endogenous GABA enrichment in plants, providing references for the development and application of GABA-rich plant-based foods.
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γ-氨基丁酸（γ-aminobutyric acid，GABA）化学式为C4H9NO2，结构式见图1，是一种自由态非蛋白质氨基酸[1]，属于次级代谢产物，也是细胞内一种信号分子。1883年GABA被首次合成，并于1949年在马铃薯中首次发现[2]，随后研究表明，GABA主要存在于植物的胚芽中，具有调节碳氮间平衡、抵御逆境胁迫和维持pH平衡等功能。1950年，研究发现GABA还存在于哺乳动物的神经系统中，同时后续研究表明在哺乳动物的神经细胞中GABA是重要的抑制性神经递质[3]，具有缓解焦虑、加快伤口愈合、调节内分泌和防止动脉硬化等功效[4−5]。
虽然GABA具有良好的生理功能，但人体内GABA含量较低，需要从外界食物中获得以满足机体需求[6]。2009年我国将GABA认定为新食品原料[7]，并允许在饮料、可可制品和巧克力等食品中添加（除婴儿食品外），添加量不能超过500 mg/d。因此，通过日常膳食结构的优化来补充GABA有助于人体健康，而如何提高食物中GABA含量以开发富含GABA的功能食品也逐步受到广泛关注。目前，我国关于GABA食品大多以糙米、大豆、玉米和燕麦等粮食作物为原料，通过GABA富集方式研制功能性食品和饮料。发芽是指植物种子吸水涨破后，籽粒膨胀、软化，呼吸和代谢作用增强，根芽生长的过程[8]，在发芽过程中植物籽粒中GABA等营养物质快速富集，同时，当受到外界环境刺激，如低温、低氧、超声等非生物胁迫时，植物会通过一系列生理代谢调节适应胁迫环境，该过程也会促进GABA等营养物质富集[9]。
本文将从GABA的制备、代谢途径、影响因素以及环境胁迫对植物发芽富集内源性GABA的协同效应进行分析总结，旨在为富含GABA的功能性食品开发与研究提供理论依据。
1. GABA的制备
目前，GABA的制备方法主要有化学合成法、植物富集法和微生物发酵（表1）。
表 1 GABA的制备Table 1. Preparation of GABA1.1 化学合成法
化学合成法主要有四种方法，分别为γ-氯丁氰法、α-吡咯烷酮开环法、丁酸和氨水法以及γ-丁内酯和亚硫酰氯法。其中，γ-氯丁氰具有成本低廉的特点，但由于工艺条件过于复杂，且易存在化学物质残留被限制推广使用。α-吡咯烷酮开环法具有温和、安全的特点，但所得产物并非天然产物，故而不能用于食品添加。γ-丁内酯和亚硫酰氯法虽然具有产率高的优势，但生产成本过高且有化学残留[13]，目前该方法主要应用在化工和医药领域，暂未在食品领域应用。
1.2 微生物发酵法
微生物发酵法早期以大肠杆菌为主要菌种，但因大肠杆菌在食品开发上仍存在安全问题，现在大多选择使用安全、有益的酵母菌和乳酸菌等菌种进行GABA的富集。夏亚男等[14]对高产GABA的菌株进行筛选，发现存在3株高产GABA菌株SMN10-3、SMN12-7、SMN15-6。邢宏博等[15]采用红曲霉固态发酵的方式对生产GABA的发酵工艺条件进行优化研究发现，在加水量30 mL、接种量25%、培养温度25 ℃、发酵培养8 d条件下，GABA含量为655 mg/100 g，为优化前的1.90倍。另外，有研究者对发酵培养基pH、组成成分[16]等培养条件进行优化，说明微生物发酵法富集GABA受多种条件影响。
1.3 植物富集法
植物在受到外界环境刺激时会引起GABA的富集。郭芳[17]对燕麦进行发芽处理，发现在25 ℃浸泡8 h、发芽16 h时，GABA含量为253.55 mg/100 g，为原料的12.34倍。童晓萌等[18]研究发现在20 ℃浸泡8 h、发芽98 h时，苦荞籽粒的GABA含量为286 mg/100 g，为原料的1.17倍。王淑芳等[19]发现发芽大豆在培养液pH5.0、发芽温度30 ℃、低氧胁迫48 h条件下，GABA含量可达到197 mg/100 g，为原料的1.56倍。研究发现不同胁迫方式对植物富集GABA效果有较大差异，本文将着重对不同胁迫方式对植物发芽过程中内源性GABA富集的影响效果进行探究。
2. 植物富集GABA的代谢途径及其影响因素
GABA在植物体内的代谢途径主要包括GABA支路和多胺降解途径。其中GABA支路为主要代谢途径，对GABA支路的贡献率可达70%左右，而多胺降解途径在植物体内的贡献率为30%左右[6]。在GABA支路中，植物GAD活性主要表现在pH依赖性和Ca2+依赖性两种调节水平[20]，通过调节GAD活性来影响GABA的富集。另外还可通过添加GABA支路和多胺降解途径相关底物物质来提高GABA的富集效果。
2.1 GABA的代谢途径
2.1.1 GABA支路
在植物体内，GABA支路（图2）[21]是指L-Glu在谷氨酸脱羧酶（Glutamate decarboxylase，GAD）催化下生成的GABA从细胞质转到线粒体后，在GABA转氨酶（GABA transaminase，GABA-T）催化下生成琥珀酸半醛（Succinic acid，SSA），再经琥珀酸半醛脱氢酶（Succinic semial dehyde dehydrogenase，SSADH）的催化作用转化为琥珀酸进入三羧酸循环的代谢过程[22−24]。其中，L-Glu能够参与碳氮平衡的协调，连接GABA支路和TCA循环，为GABA合成提供碳骨架和α-氨基，L-Glu的合成主要依赖谷氨酸合成酶（Glutamate synthetase，GOGAT）和谷氨酸脱氢酶（Glutamate dehydrogenase，GDH）的催化作用，其在GABA代谢途径中起重要作用[25]。GABA支路对细胞质内pH的调节、氧化应激保护、信号传导、氮代谢和渗透调节等生理反应均有影响[26]。
2.1.2 多胺降解途径
多胺（Polyamines，PAs）是一种多聚阳离子，主要分布在植物的细胞壁以及液泡内，在植物中多以游离、不溶性束缚和可溶性结合三种形态存在。多胺降解途径合成GABA的限速酶为由相同亚基构成的二聚体二胺氧化酶（Diamine oxidase，DAO）和单体酶多胺氧化酶（Polyamine oxidase，PAO）。如图2所示[21]，精胺（Spermine，Spm）在PAO催化作用下生成1,3-二氨基丙烷和1-（3-氨丙基）-2-吡哆啉，亚精胺（Spermidine，Spd）在PAO催化下生成4-氨基丁醛和1,3-二氨基丙烷，腐胺（Putyescine，Put）在DAO催化下生成4-氨基丁醛，三种多胺物质生成的4-氨基丁醛后经氨基醛脱氢酶（Aminoaldehyde dehydrogenase，AMADH）催化产生GABA。
2.2 植物内源性GABA富集的影响因素
2.2.1 环境pH
植物体内GAD活性最适pH在5.5~6.0左右[27]，植物在受到机械损伤、低氧胁迫和酸处理等情况时，胞质内H+浓度增加，胞质发生酸化，GAD被激活，GABA含量提高。同时，GABA在合成过程中会消耗一定量的H+，引起胞内pH的增加（图3）[28]。李楠等[29]研究发现玉米胚中GAD的最适pH为5.7，在pH4.5~7.5时GAD酶活可维持在80%以上。魏彤[30]发现绿豆GAD1和GAD2在强酸和中性环境中稳定性较差，在弱酸环境中稳定性较高。张晖等[31]发现在pH为8.0的微碱性环境下，米胚GAD蛋白的微环境构象和酶活无明显变化，但在pH为3.0的酸性环境中，酶蛋白构象发生改变并导致GAD失活，说明在适宜的pH环境有利于GAD活性表达。
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图 3 H+和Ca2+浓度对GABA富集的影响Figure 3. Effects of H+ and Ca2+ concentrations on GABA enrichment2.2.2 Ca2+浓度
植物GAD是一种钙调素协调蛋白（Calmodulin，CaM），可以与Ca2+结合形成Ca2+/CaM复合体[32]。GAD氨基酸C末端的22~25个氨基酸残基组成的空间结构起到保证植物GAD与Ca2+/CaM结合的作用，能够激发GAD活性并提高GABA含量（图3）[28]，但Ca2+和CaM独立存在时并不能激发GAD活性，必须是二者的复合物才能激发GAD活性[33]。植物在受到如低温、高温、机械损伤等外界刺激时，会引起细胞质内Ca2+浓度的增加，进而提高GABA含量。程建军等[34]研究发现发芽小米在CaCl2浓度3.5 mmol/L、浸泡温度35 ℃、浸泡时间13 h、发芽时间48 h、发芽温度31 ℃条件下，GABA含量为251.46 mg/100 g，为原料的2.90倍。
2.2.3 底物水平提高
在植物富集GABA的代谢途径中，GABA支路虽仍占据主要地位，除胁迫方式外，目前大多采用添加GABA支路相关底物来促进GABA的累积（表2），但多胺降解途径对GABA的富集仍有一定的贡献率。Spd为植物体内多胺降解途径的前体物质之一，在PAO催化下可以生成氨基丁醛进而生成GABA[35]。DAO对二胺类物质具有底物专一性，对Put有催化作用，当添加外源Put时，底物增加可以促进DAO的催化作用，提高GABA含量[36]。何根生等[37]研究发现0.5 mmol/L的Put可以提高发芽豇豆胚芽和子叶PAO活性，但当Put浓度过高时会抑制PAO活性。在GABA支路中，L-Glu是GAD唯一底物，外源添加L-Glu可以促进GABA的累积[38]。GAD以磷酸吡哆醛（Pyridoxal-5-phosphate，PLP）为辅酶专一性催化L-Glu脱羧，导致GABA累积，同时维生素B6因其与PLP具有相似的结构，是PLP的前体物质，故而亦可作为底物物质外源添加来富集GABA[39]。
表 2 底物对植物发芽GABA含量影响Table 2. Effects of different substrates on GABA enrichment during plant germination3. 环境胁迫对植物发芽富集GABA的影响及其作用机制
常见的胁迫方式包括超声胁迫、低温胁迫、低氧胁迫、盐胁迫、微酸性处理水以及高静水压技术等。不同环境胁迫方式对GABA富集效果不同，其富集机理也有所不同。
3.1 超声胁迫
超声是一种频率高于20 kHz的机械声波[47]，可以通过改变植物种子内部GAD酶的构象变化来改变酶的活性，从而加速酶的催化速率，同时能够增加细胞膜的通透性，提高胞内Ca2+和H+浓度，激活GAD活性，提高GABA含量[48−49]。张祎等[50]发现糙米在30 kHz频率下超声15 min，发芽16 h时，GABA含量为85.36 mg/100 g，为原料的2.7倍。Ding等[51]对燕麦进行超声协同发芽处理72 h后，GABA含量为原料的32.7倍，为仅发芽处理燕麦的1.12倍，进一步代谢产物分析表明，超声波胁迫可能通过影响细胞质Ca2+水平和GAD结构来提高GAD活性。Ding等[52]在红米萌发72 h后对其进行超声处理，通过代谢组学分析发现GABA含量显著提高。单迪[53]研究发现在超声功率225 W、超声时间28 min、超声温度38 ℃条件下，发芽粟米GABA含量为389 mg/100 g，为仅发芽粟米的1.83倍。超声是采用水为介质进行的非接触式胁迫处理，具有耗时短、耗能低、产热少、可操作性强和无需生物或化学试剂等优点，但目前使用该胁迫方式进行GABA富集的相关研究较少[54]。
3.2 低温胁迫
植物种子萌发过程中，低温胁迫是一种重要的胁迫方式[55]。低温胁迫分为冻害和冷害处理，冻害指0 ℃下产生的损害，会使植物的细胞结构破坏，回温后受害部位无法恢复，冷害的温度则在0 ℃以上，一般不会严重破坏植物细胞结构，只对其生长发育产生影响，回温后可通过救治使其恢复生命活动[56]。不同程度的低温胁迫会破坏植物细胞结构，增加H+浓度，激发GAD活性，同时抑制GABA-T的活性，达到富集GABA的目的。低温处理通过破坏细胞膜完整性提高胞质内Ca2+浓度[57]，形成Ca2+/CaM复合体，激发GAD活性，诱导GABA的累积。Yang等[58]研究发现在缺氧处理后，将发芽大豆在−18 ℃冷冻12 h后，置于25 ℃解冻6 h可使GABA发生累积，GABA含量为非冻融处理的7.21倍。孙威等[59]研究发现在冷冻胁迫2 h、浸泡6 h、培养24 h时，发芽小麦中GABA含量为139.83 mg/100 g，比未发芽和发芽的小麦高出93.5%和29.3%。尹永琪等[60]将经低氧处理的发芽玉米进行低温处理，发现在−18 ℃冷冻8 h后解冻4 h时GABA含量最高，推测解冻期是GABA累积的主要阶段，可能是因为冷冻使胞内的冰晶结构对细胞膜造成破坏，解冻时冰晶消失细胞液恢复成流动状态使Ca2+和H+进入胞质。Yu等[61]对冷胁迫协同发芽处理黑米的工艺条件进行优化，发现在0 ℃处理1 h、萌发72 h时，GABA含量为195.64 mg/100 g，比发芽黑米GABA含量高51.54%，该过程中Ca2+浓度增加导致GAD活性提高，而AMADH活性降低，使GABA得到累积。在低温条件下，发芽植物体内GABA大量累积，但不同植物对低温的耐受性存在差异，同时低温处理的温度、时间等因素均会对GABA含量产生较大影响，故而对于不同的植物原料的低温处理条件需重新进行探究。
3.3 低氧胁迫
植物在低氧胁迫环境下，体内电子传递链受到抑制，糖类物质易经过糖酵解途径产生丙酮酸，进而分解生成乙醇和乳酸，使胞质内pH升高，GAD在酸性环境中被激活，催化L-Glu进行脱羧反应生成GABA[62−63]。另外，低氧胁迫能够抑制植物种子有氧呼吸作用，使GABA-T活性被抑制，促进GABA的累积[64]。丁俊胄等[65]采用不同气体对发芽糙米进行厌氧胁迫，发现糙米在发芽66 h后持续通入6 h CO2时，GABA含量为965.44 mg/g，为原料的1.91倍，同时发现CO2的GABA富集效果优于N2，可能是因为CO2溶于水引起pH的降低或CO2参与发芽过程中的碳代谢活动。为了提高GABA含量，目前许多研究利用低氧胁迫协同其他胁迫的方式富集GABA。周新勇等[66]通过对发芽大麦进行低氧联合酸胁迫发现在柠檬酸缓冲液pH4.0、通氧量为4.5 L/min、发芽111 h时，GABA含量可达到0.335 mg/g，与原料比提高了33.6倍。朱云辉等[67]在低氧胁迫时添加10 mmol/L NaCl进行盐胁迫，GABA含量为仅低氧胁迫的1.1倍。综上所述，低氧胁迫能够使植物GABA富集，具有成本低，操作简单等特点，但对不同的植物原料GABA富集效果有较大差异。
3.4 盐胁迫
盐胁迫会导致植物的生理性干旱，此时细胞需要从细胞膜外吸收大量离子，其中Ca2+与CaM结合，激发GAD活性，同时植物体内的PAO、DAO和AMADH活性也会随之增加，说明盐胁迫条件下植物体内两条代谢途径均有参与[68]。Al-Quraan等[69]利用qRT-PCR技术研究发现，盐胁迫处理小麦萌发后GABA累积和GAD表达均显著提高，表明在盐和渗透胁迫下GAD均被激活。陈春旭等[70]将糙米置于NaCl培养液中进行发芽处理，发芽3 d后GABA含量为121.714 mg/100 g、GAD活性为5.7845 U/g，分别为未胁迫处理的1.12和1.24倍，同时对蛋白组成分析发现，盐胁迫处理后大分子蛋白组分会降解成小分子组分，而醇溶蛋白在该胁迫过程中未被利用。由于盐胁迫对植物生长有抑制作用，所以在进行盐胁迫处理时，大多选择加入外源添加物来缓解这一现象。郭元新等[8]发现，单纯低氧胁迫与低氧联合盐胁迫相比，GAD和DAO活性均提高，说明该种联合胁迫方式对两条代谢途径均有提高作用。盐胁迫是GABA富集的有效方式之一，但研究表明同一植物的不同器官感受盐胁迫的强度不同，GABA含量存在差异[71]。
3.5 微酸性电解水
微酸性电解水（SAEW）是在电解装置中电解稀盐或稀盐酸溶液生成的pH5.0~6.5的水溶液[72]，有效氯成分主要是次氯酸（HClO），作为杀菌剂在食品中广泛应用。微酸性电解水的瞬时杀菌效果良好，可迅速夺取细菌电子点位，短时间内快速杀死细菌[73]。研究发现微酸性电解水对植物种子发芽具有诱导作用，可以促进植物体内营养物质的富集[74]。Hao等[75]将荞麦放入pH5.83，有效氯20.3 mg/L的微酸性电解水中浸泡后进行发芽，发现在发芽6 d时GABA含量达到最大值143.20 mg/100 g，为原料的14.3倍，同时GAD活性显著提高。华建业等[76]对发芽小米进行微酸性电解水处理，发现在有效氯浓度24 mg/L、浸泡10.5 h、温度29 ℃、发芽时间40.5 h时，GABA含量为109.72 mg/100 g，为原料的1.7倍。Li等[77]研究发现在微酸性电解质水中加入15 mg/mL或30 mg/mL有效氯处理发芽粟谷，可促进GABA累积，较原料增加21%。微酸性电解水作为一种新型非热杀菌技术，具有绿色经济、无毒无害和光谱抑菌性等优点，目前主要集中在食品杀菌、保鲜、营养物质富集等方面。
3.6 高静水压技术
高静水压技术（High hydrostatic pressure，HHP）是指在一个密闭容器中以水为压力传递介质对其中物料进行均衡压力的施加[78]的技术。HPP会破坏植物体内细胞结构，加速植物细胞内物质运输速度，可以通过酶促反应诱导L-Glu转化为GABA。HHP可以破坏该分子物质结构，引起大分子物质改性，但对小分子物质影响变小。在研究过程中可以通过改变温度、压力和时间等相关工艺参数来影响植物体内化学反应及酶反应速率。Kim等[79]研究发现将发芽与HHP处理结合可以显著促进生理代谢物质的生物合成，加快酶促反应速率，提高GABA和总阿拉伯木聚糖等功能性化合物含量，发芽2 d的糙米在50 MPa压强环境下继续发芽24 h后，GABA含量为111.4 mg/100 g，为原料的1.76倍。高静水压技术作为一种非热物理改性技术，对食品本身的营养成分损害较小，能够最大程度保持食品本身的色、香、味和营养成分，具有纯天然、绿色、无公害的优点，但目前高静水压技术的相关研究仍比较欠缺[80]。
3.7 其他胁迫方式
除常见胁迫方式外，紫外光照[81]、等离子体处理[82]、高压灭菌[82]、等方式也可起到富集GABA的目的。紫外线波长能通过破坏生物体内DNA物质导致细菌死亡从而起到杀菌作用。范军等[83]研究发现将发芽24 h糙米置于距离紫外光线25 cm处，每隔6 h进行3 min紫外照射，GABA含量可达到55.7 mg/100 g，与未进行紫外处理的相比增加了1.11倍。冷等离子体（Cold Plasma，CP）是指在各类激发能激发作用下，能够提高气态分子、原子的动能，从低能态激发到高能态产生的等离子体，是一新型的非加热杀菌技术[84]。Chen等[85]研究发现对发芽荞麦进行冷等离子体胁迫处理时，GABA含量为222 mg/100 g，为原料的2.64倍。脉冲强光处理后植物体内的GAD活性增加，可通过三羧酸循环来提高GABA含量[86]。张良晨等[87]研究发现发芽糙米在照射距离9.0 cm、光照强度450 J、照射次数395次时GABA含量可达到170.10 mg/100 g，为原料的3.7倍。
4. 结论和展望
GABA作为一种具有缓解焦虑、促进睡眠、降低血糖等多种生理功能的新型功能因子，在我国食品科学基础研究及应用研究领域逐渐引起关注。现阶段对于GABA的研究大多集中在GABA生理功能机制研究以及食物原料富集GABA的多元化途径探究，多以评估盐胁迫、低氧胁迫、超声胁迫等常见的胁迫方式对GABA富集的影响，具有一定的局限性，缺乏对新型胁迫方式的探索研究以及不同胁迫方式作用机制的深度挖掘，同时，不同食物原料GABA富集效果的差异化机制还有待探讨。此外，市售产品所用食品原料单一，大多以发芽糙米或大豆为主，从口感风味、营养组成、健康功效等方面存在局限性，故而在保证食品本身营养价值和安全品质的前提下，开发更多高含量GABA且营养均衡的健康产品是实现GABA产业发展的前提。
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图 3 H+和Ca2+浓度对GABA富集的影响
Figure 3. Effects of H+ and Ca2+ concentrations on GABA enrichment
表 2 底物对植物发芽GABA含量影响
Table 2 Effects of different substrates on GABA enrichment during plant germination
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