“功能性小麦新品种及其研发利用”特约专栏文章之六
随着科技的发展和人民生活水平的提高,人民的饮食也从“吃得饱”转变为“吃得好”。高端农作物市场需求增大,特殊功能性农作物新品种亟需丰富[1]。2014年,世界粮农组织(FAO)与世界卫生组织(WHO)联合举办的第二届国际营养大会上提出了“营养导向性农业(NSA)”的概念。2017年,我国发布《主要农作物品种审定标准》,首次正式提出了“高产稳产品种、绿色优质品种和特殊类型品种”三类品种的要求和标准。
特殊颜色作物作为一类重要的特殊类型品种,具有观赏性和自然的功能性和特殊营养,愈来愈受到人们的青睐,已成为科学研究热点和高端作物产业市场的重要增长点[2]。特殊颜色谷物由于在果皮、种皮或糊粉层与正常谷物相比较含有色素的种类和数量不同,从而呈现不同颜色[3]。例如紫薯、紫米、红米、红麦、紫麦、蓝麦、紫玉米、黑玉米等。
色素是一类能使有机体呈现不同颜色的物质,因特异化学结构吸收特定波长光而显色[4],植物中的色素主要包括脂溶性色素和水溶性色素,脂溶性色素主要包括叶绿素和类胡萝卜素,水溶性色素主要是一些类黄酮类化合物[5-6]。叶绿素主要存在于叶片中,而特殊颜色谷物主要是其果皮、种皮或糊粉层中含有花青素、黄酮和类胡萝卜素等色素,从而呈现不同的颜色,花青素由于pH值和修饰基团不同会呈现紫、蓝、黑色等[7],麦黄酮会呈现黄色[8],在蔬菜和水果中类胡萝卜素常呈黄色、橙色和红色等[9]。
特殊颜色谷物中常见色素包括花青素、麦黄酮和类胡萝卜素等三种色素,其中花青素和麦黄酮是类黄酮类化合物,属于酚类化合物。类胡萝卜素是含40个碳的类异戊二烯聚合物,属于萜类化合物。
类黄酮是植物次生代谢的产物,属于酚类化合物,广泛分布于各种植物中,在植物中已经发现了超过9 000种类黄酮化合物,而且这一数目还在不断增加[10],类黄酮是由两个芳香环通过三碳桥连接起来的15碳化合物,其结构是由两部分组成的,分别来自两个生物合成途径,一个芳香环和三碳桥是来自苯丙氨酸的莽草酸途径,另一个芳香环来自丙二醛途径。类黄酮是由p-香豆酰CoA和丙二酸单酰CoA在查尔酮合酶催化下缩合合成的。根据三碳桥的氧化程度,类黄酮包括花青素、黄酮、黄酮醇和异黄酮等。
花青素(Anthocyanidin)是自然界中广泛存在于植物中的天然水溶性色素,属于类黄酮类化合物[11-12],花青素经过糖基化、酰基化和甲基化修饰,从而产生不同类型的花青素。到目前为止,已从各种植物中鉴定出七百多个花青素,其中大约有 23种花青素[13],常见广泛存在于植物中的花青素主要有6类,包括矢车菊色素(Cyanidin)、飞燕草色素(Delphinidin)、天竺葵色素(Pelargonidin)、芍药色素(Peonidin)、矮牵牛色素(Petunidin)和锦葵色素(Malvidin)。目前最常见的糖基是葡萄糖,其次是鼠李糖、半乳糖、木糖和阿拉伯糖。此外,这些糖类还可以和芳香族或脂肪族酸进行酰化作用。
花青素在某些谷物以及叶类和根系蔬菜中含量丰富,例如有色土豆、茄子、卷心菜、豆类和红洋葱[14]。新育成的11S91白菜中,BrMYB2基因大量表达显著提高花青素含量,从而使白菜心呈现紫色[15]。与蔬菜相比,水果中的花青素更为复杂,在水果中,没有酰基和仅有一个或两个单糖的花青素比例为74%,而蔬菜中超过77%的花青素具有一个或多个酰基,通常为芳香族酰基[16]。花青素不仅为植物提供鲜艳的色彩,吸引动物传粉,还具有强大的自由基清除能力[17]。在彩色土豆、黑胡萝卜、红萝卜或红卷心菜等某些蔬菜中所存在的花青素,通常具有更高的热稳定性和光稳定性,因此它们更适合用作着色剂[18]。研究发现吡喃花青素,对pH的变化具有更大的稳定性,并显示出从黄色到亮蓝色的多种颜色[19]。
花青素具有重要营养价值[20]。Kazemi等用从石榴皮中提取的花青素加入面条中,强化了面条,显著提高了面条的抗氧化能力[21]。将富含各种水平花青素的黑米提取物,加入面包中,发现面包消化率受花青素含量的影响[22]。花青素有利于人类健康,具有较好的抗氧化活性,能提高抗肿瘤和抗突变的能力[23],有利于改善糖尿病和心脏病的不利影响[24-25]。有研究发现,14种花青素对氧自由基具有明显的清除能力[26]。Tsuda等发现,矢车菊素-3-葡萄糖可降低小鼠体内血清蛋白和脂质体的过氧化作用[27]。Sarma等发现色苷-DNA复合体有助于防御DNA被氧化损伤[28]。
麦黄酮是黄酮类化合物,广泛存在于禾本科植物中[29],最初是从一种抗锈病的小麦中分离出来的[30]。黄酮类化合物多为结晶状固体,少数为非晶形粉末。其颜色以黄色为主,这主要由分子中交叉共轭体系及助色团的数目、取代的位置所决定。黄酮醇类化合物具有高氧化还原电位,能够充当还原剂、氧供体和单线态氧猝灭剂,因此是重要的抗氧化剂[31]。黄酮类化合物的作用包括保护植物免受各类胁迫在细胞水平上所造成的氧化损伤[32]。
麦黄酮和花青素早期的生物合成途径相同,后期柚皮素在黄酮酶的催化下形成芹菜素,之后在类黄酮 3′5′羟化酶和 O-甲基转移酶的催化下形成麦黄酮(图 1)。麦黄酮能显著提高植物抗病虫害的能力[35]。研究表明,麦黄酮能够有效抑制氧化偶氮甲烷和葡聚糖硫酸钠诱导的小鼠结肠癌变[36],麦黄酮具有较强的抗克隆能力,能抑制癌细胞生长[37],这是由于其具有降低环加氧酶活性的能力。此外麦黄酮能抑制人乳腺癌细胞中的P-糖蛋白活性,从而延迟自发性乳腺肿瘤的发生并抑制氧化应激诱导的细胞凋亡[38]。
图1 花青素和麦黄酮合成途径[33-34]
Fig.1 Synthesis pathway of anthocyanin and tricin[33-34]
注:PAL:苯丙氨酸裂解酶;C4H:肉桂酸-4-羟基化酶;4CL:4-香豆酸-辅酶 A连接酶;ACC:氨基环丙烷羧酸;CHI:查尔酮异构酶;F3H:黄烷酮3-羟化酶;ANS:花青素合成酶;FNS:黄酮酶;F3′5′H:类黄酮3′5′羟化酶;OMT:O-甲基转移酶。
Note:PAL:phenylalanine ammonia-lyase; C4H:cinnamic-4-hydroxylase; 4CL:4-coumaric acid CoA ligase; ACC:aminocyclopropane carboxylic acid; CHI:chalcone isomerase; F3H:Flavanone 3-hydroxylase; ANS:anthocyanin synthase; FNS:flavonoid enzyme; F3′5′H:flavonoid 3′,5′-hydroxylase; OMT:O-methyltransferase.
四萜化合物是含有八个异戊二烯单位的化合物,其合成的前体物质是异戊二烯基焦磷酸(IPP),该前体物质来自两条合成途径,分别是甲羟戊酸途径和磷酸甘油醛-丙酮酸途径[39-40]。根据分子组成不同,类胡萝卜素可以分为两类(图2)。一类是胡萝卜素、另一类是叶黄素。胡萝卜素只含有碳氢两种元素,有 α-胡萝卜素、β-胡萝卜素和番茄红素等。叶黄素是胡萝卜素的含氧衍生物,有叶黄素、玉米黄质等。类胡萝卜素基本化学结构中含有共轭双键,可以吸收可见光,并且因共轭双键的数量不同从而具有浅黄色、深橙色等颜色[43]。目前已知的类胡萝卜素大约有800多种[44],在生物体内发挥着重要作用。
图2 类胡萝卜素合成途径[41-42]
Fig.2 Carotenoid synthesis pathway[41-42]
注:GGPS:牻牛儿基牻牛儿基焦磷酸合成酶;PSY:八氢番茄红素合成酶;PDS:八氢番茄红素脱氢酶;ZDS:ζ-胡萝卜素脱氢酶;LYCB:番茄红素β-环化酶;LCYE:番茄红素ε-环化酶。
Note:GGPS:geranylgeranyl pyrophosphate synthase; PSY:octahydro lycopene synthase; PDS:octahydro lycopene dehydrogenase;ZDS:ζ-carotene dehydrogenase; LYCB:Lycopene β-cyclase; LCYE:lycopene ε-cyclase.
植物细胞质中,异戊二烯基焦磷酸形成牻牛儿基牻牛儿基焦磷酸(GGPP),然后在八氢番茄红素合成酶(PSY)、八氢番茄红素脱氢酶(PDS)、ζ-胡萝卜素脱氢酶(ZDS)的催化下下形成番茄红素[45-46]。番茄红素是类胡萝卜素合成途径的关键分支点,可分别合成α-胡萝卜素和β-胡萝卜素。这两条合成路径均是由番茄红素 ε-环化酶(LCYE)和番茄红素β-环化酶(LCYB)的催化的区别在于其作用的底物不同[47]。
在蔬菜和水果中类胡萝卜素常呈黄色、橙色和红色等,有助于光收集和光保护、充当信号分子、作为引诱剂吸引昆虫传粉等[48]。类胡萝卜素含有交替的双键,可以吸收多余的能量,从而减轻氧化胁迫[49]。类胡萝卜素还可以预防视网膜退化、晒伤[50],预防肝癌[51],并增强免疫系统,此外还具有抗衰老和抗炎特性,并通过影响转录因子参与细胞内信号传导级联反应[52]。但人和动物无法合成类胡萝卜素,因此只能从食物中获取[53]。
市场上常见的有紫薯、紫米、红米、紫玉米等,已经育成的特色颜色谷物品种,包括紫薯徐紫薯5号[54]、紫米滇香紫1号[55]、红米南两优红3号[56]、紫玉米闽紫糯1号[57]等。
紫薯的食用块根中合成了大量花青素,使薯肉和薯皮呈现紫红色。紫薯中的的花青素是由矢车菊色素或芍药色素经过糖基化再经过酰基化形成的[58],故紫薯中的花青素主要有两种,一种是矢车菊素-槐糖昔-葡糖苷(Cy类),另一种是芍药素-槐糖苷-葡糖苷(Pn类)。紫薯所含的花青素等色素有助于人体抗氧化,提高机体免疫力,并且从紫薯中提取的天然食用色素,具有安全、无毒、无异味等优势,有利于人体健康。李泓烨研究发现紫薯花青素不仅可以降低谷草转氨酶(AST)、谷丙转氨酶(ALT)活力,消除细胞肿胀,维持细胞结构,还可以参与介导肝癌细胞SNU-387的凋亡过程,通过抑制肝癌细胞中的超氧化物歧化酶(SOD)的活力来提高活性氧含量,从而促进癌细胞的凋亡[59]。Cho研究发现紫薯中提取的花青素可以有效抑制 Fe2+和抗坏血酸引起的小鼠大脑的膜脂过氧化,提高小鼠认知[60]。目前培育的紫薯品种有川紫薯2号、宁紫薯1号、湘紫薯174号、济紫薯18号、京紫薯6号等。
紫米因果皮内积累色素而呈现紫色,其积累的色素主要为花青素[61-63]。傅翠珍等研究发现黑米的色素主要组成成分是矢车菊-3-葡萄糖苷[64]。Reddy等通过分析紫米中色素成分认为紫米种皮中的花青素主要是矢车菊素,其次是芍药素[65]。孙田垒研究发现,墨江紫米中主要的花青素是矢车菊素-3-葡萄糖苷,其次是芍药素-3-葡萄糖苷、矢车菊素-3-丙二酰-葡萄糖苷和天竺葵素-3-丙二酰-葡萄糖苷[66]。紫米的色素是一种纯天然的色素,稳定性较高,在食品加工领域可以作为色素使用,具有多种功效[67-68]。目前育成的品种有紫香糯861、紫米132、墨紫1号等。
紫玉米籽粒中因含有花青素,故呈现紫色,其基本结构主要由花青素在3位酰化一个葡萄糖基及丙二酰等形成。与紫薯和紫米相似,Hiromisu研究表明,紫玉米花青素主要是矢车菊色素衍生物,其次是芍药色素衍生物,还有少量是天竺葵色素衍生物[69]。Tsuda研究发现从紫玉米中提取的花青素可以有效降低脂肪酸和甘油三脂的合成酶的mRNA表达,预防肥胖[70]。目前育成的紫玉米品种有紫玉194、河“琢紫1号”、967号紫玉米等。
有些谷物还有红色品种,例如红玉米、红稻米等。红色品种的色素主要是原花青素,与紫色品种中的花青素不同,原花青素也被称为缩合单宁,是一种由黄烷-3-醇组成的低聚物,能有效清除人体内部自由基,并具有降血脂、改善贫血、保护肝脏、增强人体免疫力等功能[71-72]。李清华等研究发现,红米中的黄酮含量显著高于白米[73]。目前常见的品种有红 401、红 239、舟山红米等。
特殊用途功能性小麦品种是全世界小麦育种的新趋势。通过研究小麦中黄酮类物质生物合成的机理,为进一步培育富含色素的小麦品种奠定了坚实的基础[74]。孙兰珍教授自1998年进行红、黑粒小麦选育研究,通过多年努力,于2006年育出了17个种植品质较好的红、黑粒小麦品系[75],山西省农科院育成了黑色小麦“河东乌麦526”[76],中国科学院西北高原生物研究所育成了紫黑色春小麦“高原115”[77],推广面积较少[78]。
近日山东农业大学农学院田纪春教授最新育成色素类功能性小麦新品种“山农蓝麦1号”和“山农 101”(山农黄酮麦 1号),通过审定,进入推广种植和加工应用阶段。
“山农蓝麦1号”,该品种籽粒因糊粉层含有大量蓝色花青色素而呈深蓝色。经过3年的区域和生产试验,2020年通过山东省农作物品种审定委员会审定。该小麦不仅富含花青素,还富含多种其它色素和VE、VB等多种维生素,具有清除人体内自由基,维持血管正常渗透压,改善心肌营养和抑制癌细胞发生等生理功能。该品种半冬性,株型较紧凑,叶片深绿,旗叶上举,抗倒伏,长芒、白壳、蓝粒,籽粒半角质,越冬抗寒性好。在2017—2020年特殊用途小麦区域试验中,平均亩产488.48公斤;2019—2020年生产试验中,平均亩产502.17公斤,两年试验皆比对照品种增产2%左右。
“山农101”是国内审定的第一个高黄酮小麦新品种。经山东省农科院农业质量标准与检测技术研究所测定,“山农101”麦黄酮含量1.013 mg/kg,是普通小麦平均含量的 3.5~5倍。黄酮类物质是小麦籽粒中重要的生理活性物质,具有扩张血管、抗菌消炎和增强人体免疫力等多重功效,对高血脂、心血管疾病、动脉硬化和肿瘤等疾病有一定疗效。该品种田间表现良好,株型半紧凑,旗叶上举,较抗倒伏,穗长方形,顶芒、白壳、白粒,籽粒硬质,越冬抗寒性好。在2017—2019年特殊用途区域试验中,两年平均亩产508.52公斤,比2个对照品种分别增产5.27%和4.13%;在2018—2019年生产试验,平均亩产537.16公斤,比对照品种分别增产4.28%和4.20%。
这两个专用型小麦既含有丰富的营养,包括花青素、黄酮、多种人体必需氨基酸和维生素,有益于人体健康,又具有较高的产量。在生产上若能将色素类功能小麦与相关食品企业联合,既能提高小麦价格,提高农民收入,又能提高企业食品营养,增加食品附加值。
2017 年国务院办公厅印发《国民营养计划(2017—2030年)》,提出“提升营业型农产品的占比,推进传统食品的升级换代,创立营业型农产品推广体系,丰富营养健康产品供给,提高国民营养健康水平”。功能性食品将是大农业到大健康的基础和桥梁,通过健康饮食提升免疫力,是未来高端食品的重要发展方向。近年来,我国先后出台《国务院关于实施健康中国行动的意见》《健康中国行动组织实施和考核方案》和《健康中国行动(2019—2030年)》等政策,明确指出“鼓励研发生产符合健康需求的产品,增加健康产品供给”,营养导向型农业和功能农业迎来了前所未有的发展机遇。小麦生产在产量提高和加工品质改善之后,提升营养和功能性品质是今后育种研究的重点。目前,对色素类功能性小麦的研究仍处于起始阶段,如何更有效的培育色素类功能性谷物、检测功能性品种的代谢组差异及保证功能性育种的稳定性、丰富性和可靠性,都是未来的重点发展方向。
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Research Progress on Special Colored Grains and Creation of New Pigment Functional Wheat Varieties