黄淮冬麦区小麦品种植酸含量与植酸酶活性聚类分析

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李颖睿, 陈茹梅, 朱伟, 阎俊, 何中虎, 张勇. 黄淮冬麦区小麦品种植酸含量与植酸酶活性聚类分析. 作物学报, 2014, 40(2): 329-336[LI Ying-Rui, CHEN Ru-Mei, ZHU Wei, YAN Jun, HE Zhong-Hu, ZHANG Yong. Variability of Phytate Content and Phytase Activity among Wheat Cultivars from Yellow and Huai River Valleys. 作物学报, 2014, 40(2): 329-336] 复制到剪切板

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作物学报编辑部

黄淮冬麦区小麦品种植酸含量与植酸酶活性聚类分析

李颖睿¹, 陈茹梅², 朱伟³, 阎俊⁴, 何中虎^1,⁵, 张勇^1,^*

¹中国农业科学院作物科学研究所/国家小麦改良中心, 北京 100081

²中国农业科学院生物技术研究所, 北京 100081

³河南省商丘市农林科学院, 河南商丘476000

⁴中国农业科学院棉花研究所, 河南安阳 455000

⁵国际玉米小麦改良中心(CIMMYT)中国办事处, 北京 100081

^* 通讯作者(Corresponding author): 张勇, E-mail:zhangyong05@caas.cn, Tel: 010-82108745

收稿日期:2013-11-24

基金:本研究由中央级公益性科研院所基本科研业务费专项资金, Harvest-Plus挑战计划项目(HPC#8273), 国家高技术研究发展计划(863计划)项目(2012AA101105)和中国农业科学院创新工程项目资助。

摘要

植酸含量与植酸酶活性是影响铁、锌等微量元素生物有效性的关键因子。2009—2010和2010—2011年度, 在河南安阳种植212个黄淮麦区代表性小麦品种和高代品系, 分析其籽粒植酸含量和植酸酶活性。结果表明, 这2个指标变异范围较大, 植酸含量为2.18~13.37 g kg^-1, 平均5.72 g kg^-1; 植酸酶活性为10~1759 U kg^-1, 平均657 U kg^-1。品种及品种与年度互作效应显著影响植酸含量和植酸酶活性, 以品种效应较大。根据植酸含量与植酸酶活性将参试品种分别聚为5类, 类间植酸含量和植酸酶活性差异显著。石麦12、衡4568、洛麦21和济麦096141的植酸含量较低, 且植酸酶活性较高, 可作为进一步改良植酸含量和植酸酶活性的亲本。

关键词: 普通小麦; 植酸含量; 植酸酶活性; 营养品质

Variability of Phytate Content and Phytase Activity among Wheat Cultivars from Yellow and Huai River Valleys

LI Ying-Rui¹, CHEN Ru-Mei², ZHU Wei³, YAN Jun⁴, HE Zhong-Hu^1,⁵, ZHANG Yong^1,^*

¹ Institute of Crop Science / National Wheat Improvement Center, Chinese Academy of Agricultural Sciences (CAAS), Beijing 100081, China

² Biotechnology Research Institute, CAAS, Beijing 100081, China

³ Shangqiu Research Institute of Agriculture and Forestry Science, Shangqiu 476000, China

⁴ Cotton Research Institute, CAAS, Anyang 455004, China

⁵ CIMMYT-China Office, c/o CAAS, Beijing 100081, China

Abstract

Phytate content and phytase activity are key factors influencing bioavailability of iron and zinc. To understand the status of phytate content and phytase activity in wheats from the Yellow and Huai River Valleys Winter Wheat Region, 212 representative cultivars and advanced lines were sown in Anyang, Henan Province, China in 2009-2010 and 2010-2011 cropping seasons. Phytate content and phytase activity varied greatly among these cultivars, ranging from 2.18 to 13.37 g kg^-1 of phytate content and from 10 to 1759 U kg^-1 of phytase activity, with the mean values of 5.72 g kg^-1 and 657 U kg^-1, respectively. Both indices were significantly affected by genotype and genotype × season interaction, with genotype effect being predominant. All cultivars were classified into five groups based on the seasonal standardized values of phytate content and phytase activity, with significant difference among groups. Four cultivars, i.e., Shimai 12, Heng 4568, Luomai 21, and Jimai 096141 exhibited low phytate content and high phytase activity, and can be used in wheat breeding program aiming at improving iron and zinc nutritional quality.

Keyword: Bread wheat; Phytate content; Phytase activity; Nutritional quality

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微量营养元素缺乏造成的营养不良非常严重。全世界约20亿人患有不同程度的贫血, 其中约12%由缺铁所致, 而铁缺乏者的数量约为缺铁性贫血患病率的2.5倍^{[ 1, 2]}; 超过一半人口摄入的锌和维生素A等营养元素不足^{[ 2]}。发展中国家超过50%的妇女和儿童及约58%的孕妇存在缺铁性贫血^{[ 3]}, 每年约有800万儿童死于锌缺乏症^{[ 4]}。我国的缺铁性贫血发病率为20%左右, 贫困地区儿童和孕妇则高达45%和35%^{[ 1]}。

小麦是我国北方地区的主要粮食作物, 黄淮麦区是最重要的商品粮基地, 提高人体对小麦籽粒中矿物质元素的吸收利用, 对于解决我国人民由于矿物质元素含量摄入不足造成的健康问题具有重要意义^{[ 4, 5]}。人体对食物中铁锌等矿物质元素的吸收利用取决于其含量和生物有效性的高低^{[ 2, 3, 4]}, 植酸含量和植酸酶活性是影响微量矿物质元素生物有效性的主要因子^{[ 6]}。影响铁锌等生物有效性的限制性因子包括植酸、纤维素、丹宁和重金属等, 以植酸最为重要^{[ 2, 3, 4]}。植酸是磷在小麦籽粒中的主要贮存形式, 通过与铁、锌、钙等二价金属离子结合, 形成螯合态植酸盐^{[ 7]}, 降低人对铁、锌等的吸收利用^{[ 8]}, 植酸含量与锌含量呈显著正相关^{[ 9]}。食物中的植酸盐可以被植酸酶有效分解, 但人和猪等单胃动物的消化系统中植酸酶活性非常低^{[ 10]}, 导致铁、锌和磷等矿物质元素无法被有效吸收, 同时大量的有机磷还随排泄物排出, 引起土壤和水污染^{[ 11]}。植酸虽然具有抗癌^{[ 12]}、预防心脏病与糖尿病等代谢类疾病的功能^{[ 13]}, 并可提高种子活力, 降低籽粒中黄曲霉毒素的含量^{[ 4]}, 但对微量矿物质元素生物有效性的阻遏作用非常显著, 这对婴儿和孕妇等需要大量铁、锌等元素的人群来说非常重要^{[ 1, 3]}。植酸酶可有效分解植酸, 通过提高铁、锌和磷等元素的生物有效性, 大大改善人体因铁锌等元素缺乏所导致的营养不良问题^{[ 2]}。

植酸酶已作为添加剂广泛用于食品和饲料工业^{[ 14]}, 采用微生物方法进行工业化生产成本较高, 生产过程中大量消耗能源并严重污染环境^{[ 6]}。转植酸酶基因玉米已获生产应用安全证书并进行环境释放^{[ 15]}, 但转基因作物受大众认知和接受程度的严重影响, 目前还难以大规模应用^{[ 16]}。因此, 有必要对现有小麦品种的植酸酶活性进行筛选, 以充分利用籽粒本身所含的植酸酶。

国内外对小麦的铁、锌等元素含量已进行了较深入研究, 品种间存在显著差异, 并受环境显著影响^{[ 17]}。铁含量相关基因/主效QTL定位在2A、4A、5A、7A和4D等染色体上, 锌含量相关基因/主效QTL定位在1A、2D、3A、4A、4D、5A和7A等染色体上^{[ 18, 19]}。对印度和国际玉米小麦改良中心(CIMMYT)品种的植酸含量和植酸酶活性已进行了初步报道^{[ 20]}, Liu等^{[ 9]}和吴澎等^{[ 21]}曾对我国的地方品种和河南、山东、陕西、江苏和四川等少数地区的品种进行了植酸含量和植酸酶活性分析。但总体来说, 有关植酸含量与植酸酶活性分析的资料十分有限, 黄淮主产麦区主要品种的植酸含量与植酸酶活性尚不清楚。本研究选用该区212个主栽品种和高代品系, 连续两年度种植在中国农业科学院作物科学研究所安阳试验站, 检测其植酸含量和植酸酶活性, 为通过育种途径改良铁锌等营养品质提供理论依据。

1 材料与方法

1.1 品种及其田间种植

212份供试材料(表1)中, 来自河北80份、河南92份和山东34份, 均为当地主栽品种或苗头品系; 还包括已被主要育种单位用作优质亲本的中优9507和西农979, 面包、面条品质优良的澳大利亚品种Sunco和Sunstate, 7Dl.7Ag易位系品种Wheatear, 以及在国际上广泛用作条锈、叶锈和白粉病抗病亲本的RL6077 (Weebill*2/Brambling)。

2009—2010和2010—2011年度, 将所有品种种植于中国农业科学院作物科学研究所安阳试验站。田间采用拉丁方alpha格子设计^{[ 5, 22]}, 2次重复, 每隔15个品种加一个当地对照。双行播种, 行长1.5 m, 行间距0.20 m, 每行50粒, 按当地常规进行田间管理。

表1 212份材料代号及其名称和来源 Table 1 Code, name, and origin of the 212 cultivars and advanced lines used in this study

代号 Code	品种 Variety	来源 Origin	代号 Code	品种 Variety	来源 Origin	代号 Code	品种 Variety	来源 Origin	代号 Code	品种 Variety	来源 Origin
1	冀麦36	中国河北	54	科农20	中国河北	107	矮抗58	中国河南	160	曙麦05-2	中国河南
2	冀麦38	中国河北	55	科农2009	中国河北	108	丰德存麦1号	中国河南	161	许科316	中国河南
3	冀麦112	中国河北	56	科农2011	中国河北	109	丰德存麦6号	中国河南	162	许农5号	中国河南
4	冀麦5265	中国河北	57	冀师02-1	中国河北	110	丰德存麦10号	中国河南	163	许科99087	中国河南
5	冀矮1号	中国河北	58	师栾08-2	中国河北	111	百农64	中国河南	164	豫保2号	中国河南
6	冀麦325	中国河北	59	师栾08-4	中国河北	112	中育5号	中国河南	165	长海大穗	中国河南
7	冀麦518	中国河北	60	藁城9415	中国河北	113	中麦349	中国河南	166	保丰10-82	中国河南
8	冀麦585	中国河北	61	藁优5218	中国河北	114	中麦895	中国河南	167	内乡188	中国河南
9	冀麦602	中国河北	62	藁优5766	中国河北	115	中麦875	中国河南	168	豫麦34	中国河南
10	冀麦867	中国河北	63	U07-6308	中国河北	116	中麦871	中国河南	169	豫安208	中国河南
11	石4185	中国河北	64	振桥大穗	中国河北	117	中892	中国河南	170	偃4110	中国河南
12	石家庄8号	中国河北	65	新麦8号	中国河北	118	00-62154	中国河南	171	偃展9998	中国河南
13	石家庄10号	中国河北	66	新麦9号	中国河北	119	01-52289	中国河南	172	豫教0520	中国河南
14	石家庄11	中国河北	67	邢04-1135	中国河北	120	04中36	中国河南	173	济麦19	中国山东
15	石麦12	中国河北	68	邢05-1241	中国河北	121	04中70	中国河南	174	济麦20	中国山东
16	石06-6136	中国河北	69	邢麦9号	中国河北	122	06CA25	中国河南	175	济麦22	中国山东
17	石08-4741	中国河北	70	邢麦10号	中国河北	123	07CA255	中国河南	176	济麦6097	中国山东
18	石09-4276	中国河北	71	邢麦13	中国河北	124	07CA266	中国河南	177	济麦8186	中国山东
19	石5341	中国河北	72	邢台456	中国河北	125	08CA101	中国河南	178	济麦0836262	中国山东
20	石6136	中国河北	73	邯6172	中国河北	126	08CA062	中国河南	179	济麦0850187	中国山东
21	石83H-366	中国河北	74	邯07-6092	中国河北	127	08CA137	中国河南	180	济麦0860223	中国山东
22	石B05-7388	中国河北	75	邯09-41307	中国河北	128	08CA307	中国河南	181	济麦096141	中国山东
23	石B07-4056	中国河北	76	邯09-41344	中国河北	129	09CA034	中国河南	182	鲁原502	中国山东
24	石B07-4179	中国河北	77	邯5093	中国河北	130	09CA163	中国河南	183	烟农19	中国山东
25	石新633	中国河北	78	邯5672	中国河北	131	09CA170	中国河南	184	烟农24	中国山东
26	石新703	中国河北	79	邯5849	中国河北	132	09CA175	中国河南	185	烟99102	中国山东
27	石新733	中国河北	80	邯9983	中国河北	133	09CA86	中国河南	186	山农05-066	中国山东
28	石新811	中国河北	81	郑麦005	中国河南	134	10CA23	中国河南	187	山农055843	中国山东
29	石新828	中国河北	82	郑麦106	中国河南	135	10CA79	中国河南	188	山农2149	中国山东
30	衡观33	中国河北	83	郑7698	中国河南	136	10CA70	中国河南	189	PH6911	中国山东
31	衡观35	中国河北	84	郑麦98	中国河南	137	11CA105	中国河南	190	泰农2413	中国山东
32	衡05-4444	中国河北	85	郑丰4431	中国河南	138	11CA26	中国河南	191	泰农8681	中国山东
33	衡09-4061	中国河北	86	郑麦110	中国河南	139	11CA40	中国河南	192	泰农8968	中国山东
34	衡10-5197	中国河北	87	郑麦366	中国河南	140	9705-0-1-2-3	中国河南	193	泰农9862	中国山东
35	衡10S99-2	中国河北	88	郑育麦9987	中国河南	141	94462-0-30-5	中国河南	194	泰山064199	中国山东
36	衡4041	中国河北	89	花培5号	中国河南	142	濮麦9号	中国河南	195	泰山6195	中国山东
37	衡4422	中国河北	90	花培87-3	中国河南	143	丰优66	中国河南	196	良星619	中国山东
38	衡4568	中国河北	91	新麦18	中国河南	144	金育麦2号	中国河南	197	良星66	中国山东
39	衡5114	中国河北	92	新麦19	中国河南	145	浚K8	中国河南	198	良星99	中国山东
40	衡5218	中国河北	93	新麦0208	中国河南	146	兰考15	中国河南	199	临麦2号	中国山东
41	衡5229	中国河北	94	新麦21	中国河南	147	洛麦21	中国河南	200	潍麦8号	中国山东
42	衡5364	中国河北	95	新麦26	中国河南	148	洛麦22	中国河南	201	汶农14	中国山东
43	衡6421	中国河北	96	新选2039	中国河南	149	洛麦23	中国河南	202	YB66180	中国山东
44	衡辐418	中国河北	97	周麦16	中国河南	150	洛麦24	中国河南	203	莱州127	中国山东
45	衡辐9103	中国河北	98	周麦18	中国河南	151	洛新998	中国河南	204	枣9119	中国山东
46	河农6049	中国河北	99	周9823	中国河南	152	洛麦02133	中国河南	205	淄995015	中国山东
47	河农7069	中国河北	100	周98343	中国河南	153	洛麦05123	中国河南	206	淄麦12	中国山东
48	河农9311	中国河北	101	周麦22	中国河南	154	平安8号	中国河南	207	中优9507	中国北京
49	金丰7183	中国河北	102	周麦24	中国河南	155	平安10号	中国河南	208	西农979	中国陕西
50	金禾2417	中国河北	103	周麦25	中国河南	156	平安11	中国河南	209	Sunco	澳大利亚
51	金禾9123	中国河北	104	周麦26	中国河南	157	漯麦9908	中国河南	210	Sunstate	澳大利亚
52	科奥08-6	中国河北	105	周麦27	中国河南	158	漯麦6082	中国河南	211	RL6077	墨西哥
53	科农1006	中国河北	106	周麦28	中国河南	159	汝州0319	中国河南	212	Wheatear	墨西哥

表1 212份材料代号及其名称和来源 Table 1 Code, name, and origin of the 212 cultivars and advanced lines used in this study

1.2 性状测定

两年度收获前均无有效降雨, 未受穗发芽影响, 籽粒样品饱满度均较好。收获后各样品随机数200粒种子, 3次重复, 称重, 换算成千粒重。采用近红外分析仪(Foss 1241, Sweden)测定籽粒蛋白质含量(14%湿基)。在-20℃冷库中保存籽粒样品, 用高通量组织研磨机(SPEX GENO 2010 GRINDER, 美国)磨粉。按Chen等^{[ 15]}的方法分析植酸含量, 并适当改进。将30 mg全粉置于1.5 mL离心管中, 加入0.4 mol L^-1 HCl 1 mL和15%的TCA提取液, 室温下振荡3 h, 再以2000 × g离心10 min, 然后取50 μL上清液, 置于1.5 mL离心管中(内装36.3 mmol L^-1 NaOH 550 μL); 加入200 μL显色液(含0.03%氯化铁, 0.3%磺基水杨酸), 反应后取200 μL溶液, 用酶标仪(SPECTRA max PLUS³⁸⁴)在500 nm下读数, 测定植酸含量。参考Chen等^{[ 15]}的方法, 并略加调整, 分析植酸酶活性。100 mg全粉置1.5 mL离心管中, 加入0.25 mL抽提缓冲液(含50 mmol L^-1 NaAc、1 mmol L^-1 CaCl₂、0.5% BSA、0.075% Triton X-100, pH 5.15)。室温振荡1 h, 3000 × g离心15 min后取上清液, 按Wyss等^{[ 23]}的方法, 通过测定无机磷的增加量来确定植酸酶的活性。20 μL上清液50℃预热5 min, 加入6.25 mmol L^-1植酸钠溶液80 μL, 30 min后用15%TCA终止反应。20 μL反应后溶液中加80 μL去离子水和100 μL显色液(7.35% FeSO₄∙7H₂O、1%钼酸铵、0.6 mmol L^-1 H₂SO₄), 37℃反应10 min后,

700 nm下测定吸光值。空白对照除先加TCA再加植酸钠以外, 其余操作均相同。需要注意的是, 在小麦植酸酶提取液与植酸钠反应时, 需将最佳反应温度调整为50℃, 溶液pH值调整为5.15。

1.3 数据处理

采用合适的空间模型^{[ 17]}分析试验数据, 将两年度所得品种各性状最佳线性无偏预测值用于数据处理和分析^{[ 24]}。采用SAS (Statistical Analysis System) 8.0软件, 调用PROC MEANS、PROC MIXED、PROC CLUSTER和PROC CORR分别进行基本统计量、方差、聚类和相关等分析。其中调用PROC MIXED进行方差分析时, 将基因型类作为固定效应, 年度及年度相关互作、类内基因型和年度内重复作为随机效应。调用PROC CLUSTER时, 将植酸含量和植酸酶活性数据分别按年度进行标准化后, 以欧氏距离为标准, 按Ward类平方和法分别对品种进行聚类^{[ 25]}。

2 结果与分析

2.1 基本统计量分析

品种和年度间籽粒植酸含量和植酸酶活性均存在较大差异(表2)。植酸含量和植酸酶活性平均值分别为5.72 g kg^-1和657 U kg^-1, 变异范围分别为2.18~13.37 g kg^-1和10~1759 U kg^-1, 品种间差异远大于年度间差异。千粒重和蛋白质含量平均为46.0 g和12.9%, 变异范围分别为30.6~63.1 g和9.6%~ 16.7%。由此可见, 我国黄淮冬麦区品种间植酸含量与植酸酶活性变异范围广, 通过品种筛选降低籽粒植酸含量或提高植酸酶活性的潜力较大。

表2 212个小麦品种千粒重、蛋白质含量、植酸含量和植酸酶活性的变异 Table 2 Variations of thousand-kernel weight, protein content, phytate content, and phytase activity in 212 wheat cultivars

2.2 聚类分析

将所有品种的植酸含量和植酸酶活性数据分别按年度进行标准化, 在决定系数( R²)为90.0%和93.6%水平分别将品种聚为5类(表3和表4), 不同地区来源品种间植酸含量和植酸酶活性差异不显著(表略)。方差分析表明, 植酸含量的品种、年度及其互作效应均达0.001显著水平, 以品种效应较大, 其次为品种和年度互作效应; 品种效应中, 品种类别效应较大, 达0.001显著水平, 类内品种效应不显著。植酸酶活性的品种及品种和年度互作效应均达0.001显著水平, 以品种效应较大, 年度效应不显著; 品种效应中, 品种类别效应较大, 达0.001显著水平, 类内品种效应不显著(表5)。

5类植酸含量存在显著差异的品种中, 第3和第5极端类的品种数量较少(分别包括19个和15个品种), 绝大多数品种表现植酸含量中等。第3类品种的平均植酸含量最高, 达10.33 g kg^-1, 显著高于其他4类。第5类品种的植酸含量最低, 为2.78 g kg^-1; 其次是第4类, 平均为4.04 g kg^-1; 这两类品种的植酸含量变异范围都较小, 在低植酸含量育种中具有较高的利用价值。从品种来源看, 各类型在多个地区均有分布, 黄淮麦区广泛应用的重要亲本中, 只有周麦16属于第4类。墨西哥抗病品种RL6077也属于第4类(表3), 表现植酸含量较低。

5类植酸酶活性存在显著差异的品种类群中所包含的品种与基于植酸含量的聚类结果不尽相同。其中, 第4类品种的植酸酶活性最高, 达1146 U kg^-1; 其次是第5类, 平均植酸酶活性为930 U kg^-1; 这两类分别包括6个和27个品种, 数量较少, 分别来自多个地区, 类间植酸酶活性没有显著差异, 但都显著高于其他3类, 可作为重要的高植酸酶活性育种亲本。值得一提的是, 澳大利亚品种Sunstate和Sunco分别属于第4和第5类(表4), 表现植酸酶活性较高。

综合植酸含量和植酸酶活性的聚类分析结果, 发现低植酸含量和高植酸酶活性的共同品种有石麦12、衡4568、洛麦21和济麦096141, 分别来自河北、河南和山东省。这4个品种有可能成为重要的育种亲本, 在低植酸含量和高植酸酶活性小麦育种中发挥作用。

表3 两年度212份品种植酸含量聚类分析结果 Table 3 Cluster of 212 cultivars based on phytate content across two seasons

类别 Group	植酸含量 Phytate content (g kg^-1)	变幅 Range (g kg^-1)	品种来源 Origin of cultivar	品种代号 Code of cultivar
PC1 (90)	6.05±1.49 b	4.09-9.72	河北 Hebei (24)	49, 78, 58, 41, 12, 60, 77, 1, 29, 26, 68, 21, 69, 7, 40, 46, 32, 4, 13, 28, 11, 8, 52, 30
			河南 Henan (47)	137, 110, 140, 90, 124, 101, 84, 102, 118, 151, 132, 172, 158, 121, 113, 105, 122, 104, 165, 85, 131, 81, 159, 127, 144, 133, 125, 117, 96, 123, 89, 161, 129, 160, 88, 92, 93, 148, 112, 141, 91, 168, 150, 98, 162, 157, 149
			山东 Shandong (16)	199, 201, 202, 197, 196, 190, 175, 174, 205, 200, 188, 198, 187, 195, 203, 182
			其他 Others (3)	207, 208, 209
PC2 (62)	5.08±1.01 c	3.51-6.89	河北 Hebei (26)	45, 67, 10, 50, 24, 25, 37, 53, 47, 36, 14, 70, 63, 66, 39, 19, 3, 51, 22, 31, 27, 16, 5, 2, 72, 49
			河南 Henan (27)	134, 170, 106, 139, 164, 108, 145, 114, 142, 100, 107, 120, 95, 103, 126, 153, 143, 115, 152, 116, 167, 128, 154, 156, 163, 171, 119
			山东 Shandong (7)	206, 204, 176, 189, 173, 191, 184
			其他 Others (2)	210, 212
PC3 (15)	10.33±4.01 a	7.13-13.36	河北 Hebei (9)	65, 20, 42, 75, 73, 23, 64, 57, 48
			河南 Henan (4)	109, 99, 111, 87
			山东 Shandong (2)	186, 183
PC4 (26)	4.04±0.88 d	2.67-5.11	河北 Hebei (11)	56, 71, 35, 59, 76, 54, 62, 55, 74, 15, 79
			河南 Henan (9)	130, 169, 86, 97, 135, 138, 136, 83,147
			山东 Shandong (5)	177, 192, 181, 193, 194
			其他 Others (1)	211
PC5 (19)	2.78±0.35 e	2.25-3.24	河北 Hebei (11)	44, 18, 6, 38, 34, 33, 43, 61, 80, 17, 9
			河南 Henan (4)	94, 155, 166, 82
			山东 Shandong (4)	178, 179, 180, 185

括号中数字表示品种数。植酸含量为平均值±标准差, 数据后不同字母表示类间有显著差异( P<0.05)。各类中品种排序按植酸含量从高到低, 品种名称见表1。

The figure in parentheses is the number of cultivars. Phytate content is shown in mean ± standard deviation, and different letters afterwards indicate significant difference among groups at P< 0.05. In each group, cultivars were sorted by phytate content with value from high to low. Cultivar names are given in Table 1.

表3 两年度212份品种植酸含量聚类分析结果 Table 3 Cluster of 212 cultivars based on phytate content across two seasons

表4 两年度212份品种植酸酶活性聚类分析结果 Table 4 Cluster of 212 cultivars based on phytase activity across two seasons

类别 Group	植酸酶活性 Phytase activity (U kg^-1)	变幅 Range (U kg^-1)	品种来源 Origin of cultivar	品种代号 Code of cultivar
PA1 (56)	339±247 e	12-930	河北 Hebei (28)	21, 29, 52, 27, 20, 24, 75, 34, 79, 6, 68, 44, 14, 37, 3, 55, 32, 77, 78, 2, 26, 66, 7, 19, 70, 1, 41, 25
			河南 Henan (18)	108, 115, 123, 106, 113, 111, 92, 116, 86, 127, 128, 146, 156, 105, 99, 100, 130, 104
			山东Shandong (9)	198, 176, 187, 189, 191, 204, 185, 195, 193
			其他 Others (1)	208
PA2 (79)	757±135 c	415-978	河北 Hebei (30)	74, 71, 80, 33, 45, 10, 76, 51, 9, 56, 64, 11, 59, 4, 16, 58, 18, 13, 61, 62, 53, 35, 50, 43, 60, 49, 28, 17, 47, 57
			河南 Henan (36)	139, 122, 134, 160, 117, 83, 137, 82, 126, 169, 136, 110, 88, 155, 109, 135, 154, 132, 170, 153, 125, 166, 94, 89, 157, 159, 163, 112, 143, 142, 168, 96, 133, 98, 150, 158
			山东Shandong (10)	174, 177, 180, 179, 197, 188, 192, 205, 186, 200
			其他 Others (3)	212, 211, 207
PA3 (44)	590±160 d	235-772	河北 Hebei (14)	22, 5, 39, 54, 42, 46, 72, 23, 36, 8, 12, 31, 48, 30
			河南 Henan (21)	171, 118, 85, 120, 145, 81, 148, 97, 161, 172, 164, 138, 141, 150, 114, 95, 144, 107, 167, 162, 102
			山东Shandong (9)	182, 201, 190, 173, 178, 194, 183, 203, 175
PA4 (6)	1146±411 a	859-1736	河北 Hebei (2)	73, 40
			河南 Henan (2)	151, 124
			山东Shandong (1)	199
			其他 Others (1)	210
PA5 (27)	930±185 ab	644-1085	河北 Hebei (6)	65, 67, 69, 63, 15, 38
			河南 Henan (15)	103, 121, 165, 87, 131, 84, 119, 129, 140, 152, 147, 91, 90, 93, 101
			山东Shandong (5)	202, 196, 206, 184, 181
			其他 Others (1)	209

括号中数字表示品种数。植酸酶活性为平均值±标准差, 数据后不同字母表示类间有显著差异( P<0.05)。各类中品种排序按植酸酶活性从高到低, 品种名称见表1。

The figure in parentheses is the number of cultivars. Phytase activity is shown in mean ± SD, and different letters afterwards indicate significant difference among groups at P< 0.05. In each group, cultivars were sorted by phytase activity with value from high to low. Cultivar names are given in Table 1.

表4 两年度212份品种植酸酶活性聚类分析结果 Table 4 Cluster of 212 cultivars based on phytase activity across two seasons

表5 两年度212份品种植酸含量与植酸酶活性方差分析表 Table 5 Analysis of variance for phytate content and phytase activity of 212 cultivars across two seasons

3 讨论

对我国76份小麦地方品种和62份来自黄淮、长江中下游和西南麦区品种的分析表明, 小麦植酸含量为5.16~9.87 g kg^-1, 植酸酶活性为620~2192 U kg^{-1 [ 9]}。在我国137份微核心种质资源中, 白蚂蚱的植酸含量较低, 为9.59 g kg^-1, 辐w 070261的植酸含量则高达29.63 g kg^{-1 [ 21]}。400份印度及CIMMYT品种和人工合成种植酸含量与植酸酶活性的变异范围分别为11.7~19.3 g kg^-1与284~962 U kg^{-1 [ 20]}。本研究表明, 黄淮麦区212份代表性品种的植酸含量和植酸酶活性分别为2.18~13.37 g kg^-1和10~1759 U kg^-1, 与前人报道结果^{[ 9, 20, 26]}基本一致, 但本研究中植酸含量和植酸酶活性的变异范围更大, 可能与所选取材料的代表性和数量有关。目前还没有对植酸含量和植酸酶活性进行育种选择, 因而其变异范围较大。因此, 在改良小麦籽粒的微量矿物质营养品质时, 首先应对现有品种的目标性状进行筛选, 我们在对小麦品种铁锌等微量元素含量的分析时, 也有类似结论^{[ 5]}。本研究发现4个品种的植酸含量较低且植酸酶活性较高, 分别是石麦12、衡4568、洛麦21和济麦096141, 可作为进一步改良植酸含量和植酸酶活性的亲本。此外, 相关分析结果(资料未列出)表明, 籽粒中植酸含量和植酸酶活性呈显著正相关( r = 0.31, P< 0.001), 说明植酸含量与植酸酶活性间可能存在某种程度的依存关系, 要同时提高小麦籽粒中的植酸酶活性并降低其植酸含量存在一定困难。鉴于目前尚未对我国的大多数品种进行植酸含量和植酸酶活性分析, 建议在本研究的基础上, 进一步筛选我国现有品种特别是各地主栽品种的植酸含量和植酸酶活性, 对植酸含量低且植酸酶活性高的主栽品种加大推广力度, 同时将其用于育种, 为提高我国品种铁锌等微量矿物质元素的生物有效性奠定基础。

本研究发现籽粒植酸含量与植酸酶活性受品种、年度及品种和年度互作效应的显著影响; 植酸含量比植酸酶活性更易受年度效应的影响, 这些结果与前人报道一致^{[ 26, 27]}。Kim等^{[ 27]}选用3个品种在多个环境中种植, 结果表明植酸含量受降雨量等环境因素的显著影响, 而植酸酶活性的环境效应不显著。因此, 有必要进一步分析温度、降水量、日照时数等环境因子对植酸含量与植酸酶活性的影响。

4 结论

植酸含量和植酸酶活性变异范围均较大, 受品种、年度及其互作效应的显著影响, 其中品种效应较大。石麦12、衡4568、洛麦21和济麦096141的植酸含量低且植酸酶活性较高, 可望在小麦营养品质育种中发挥重要作用。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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22	Barreto H J, Edmeades G O, Chapman S C, Crossa J. The alpha lattice design in plant breeding and agronomy: generation and analysis. In: Edmeades G O, Banziger M, Mickelson H R, Peña-Valdivia C B, eds. Developing Drought- and Low N-Tolerant Maize. Proceedings of a Symposium. Mexico, D. F. : CIMMYT, 1997. pp544-551 [本文引用:1]
23	Wyss M, Pasamontes L, Friedlein A, Rémy R, Tessier M, Kronenberger A, Middendorf A, Lehmann M, Schnoebelen L, Röthlisberger U, Kusznir E, Wahl G, Müller F, Lahm H W, Vogel K, van Loon A P. Biophysical characterization of fungal phytases (myo-inositol hexakisphosphate phosphohydrolases): molecular size, glycosylation pattern, and engineering of proteolytic resistance. Appl Environ Microb, 1999, 65: 359-366 [本文引用:1] [JCR: 3.678]
24	张勇, 吴振录, 张爱民, van Ginkel M, 何中虎. CIMMYT小麦在中国春麦区的适应性分析. 中国农业科学, 2006, 39: 655-663 Zhang Y, Wu Z L, Zhang A M, van Ginkel M, He Z H. Adaptation of CIMMYT wheat germplasm in China’s spring wheat regions. Sci Agric Sin, 2006, 39: 655-663 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
25	Ward J H. Hierarchical grouping to optimize an objective function. J Am Stat Assoc, 1963, 58: 236-244 [本文引用:1] [JCR: 1.834]
26	Liu Z H, Wang H Y, Wang X E, Zhang G P, Chen P D, Liu D J. Phytase activity, phytate, iron, and zinc contents in wheat pearling fractions and their variation across production locations. J Cereal Sci, 2007, 45: 319-326 [本文引用:2] [JCR: 2.088]
27	Kim J C, Mullan B P, Selle P H, Pluske J R. Levels of total phosphorus, phytate phosphorus and phytase activity in three varieties of Western Australian wheats in response to growing region, growing season and storage. Aust J Agric Res, 2002, 53: 1361-1366 [本文引用:2] [JCR: 1.328]

2004

0.0

... 5倍^[1,2] ...

... 我国的缺铁性贫血发病率为20%左右, 贫困地区儿童和孕妇则高达45%和35%^[1] ...

... 植酸虽然具有抗癌^[12]、预防心脏病与糖尿病等代谢类疾病的功能^[13], 并可提高种子活力, 降低籽粒中黄曲霉毒素的含量^[4], 但对微量矿物质元素生物有效性的阻遏作用非常显著, 这对婴儿和孕妇等需要大量铁、锌等元素的人群来说非常重要^[1,3] ...

1999

2.474

0.0

... 5倍^[1,2] ...

... 超过一半人口摄入的锌和维生素A等营养元素不足^[2] ...

... 人体对食物中铁锌等矿物质元素的吸收利用取决于其含量和生物有效性的高低^[2,3,4], 植酸含量和植酸酶活性是影响微量矿物质元素生物有效性的主要因子^[6] ...

... 影响铁锌等生物有效性的限制性因子包括植酸、纤维素、丹宁和重金属等, 以植酸最为重要^[2,3,4] ...

... 植酸酶可有效分解植酸, 通过提高铁、锌和磷等元素的生物有效性, 大大改善人体因铁锌等元素缺乏所导致的营养不良问题^[2] ...

2000

2.106

0.0

... 发展中国家超过50%的妇女和儿童及约58%的孕妇存在缺铁性贫血^[3], 每年约有800万儿童死于锌缺乏症^[4] ...

... 影响铁锌等生物有效性的限制性因子包括植酸、纤维素、丹宁和重金属等, 以植酸最为重要^[2,3,4] ...

2007

2.088

0.0

J Cereal Sci. 2007, 46(3):293 - 307 DOI:10.1016/j.jcs.2007.06.005

Enhancing the mineral and vitamin content of wheat and maize through plant breeding

Ortiz-Monasterio

J I

,Palacios-Rojas

,Meng

,Pixley

,Trethowan

,Pena

R J

.Enhancing the mineral and vitamin content of wheat and maize through plant breeding.J Cereal Sci,2007,46:293-307

Abstract

More than half of the world's population suffers micronutrient undernourishment. The main sources of vitamins and minerals (iron, zinc, and vitamin A) for low-income rural and urban populations are staple foods of plant origin that often contain low levels or low bioavailability of these micronutrients. Biofortification aims to develop micronutrient-enhanced crop varieties through conventional plant breeding. HarvestPlus, the CGIAR's biofortification initiative, seeks to breed and disseminate crop varieties with enhanced micronutrient content that can improve the nutrition of the “hard to reach” (by fortification or supplementation programmes) rural and urban poor in targeted countries/regions. In attempting to enhance micronutrient levels in maize and wheat through conventional plant breeding, it is important to identify genetic resources with high levels of the targeted micronutrients, to consider the heritability of the targeted traits, to explore the availability of high throughput screening tools and to gain a better understanding of genotype by environment interactions. Biofortified maize and wheat varieties must have the trait combinations which encourage adoption such as high yield potential, disease resistance, and consumer acceptability. When defining breeding strategies and targeting micronutrient levels, researchers need to consider the desired micronutrient increases, food intake and retention and bioavailability as they relate to food processing, anti-nutritional factors and promoters. Finally,ex ante studies are required to quantify the burden of micronutrient deficiency and the potential of biofortification to achieve a significant improvement in human micronutrient status in the deficient target population in order to determine whether a biofortification program is cost-effective.

... 发展中国家超过50%的妇女和儿童及约58%的孕妇存在缺铁性贫血^[3], 每年约有800万儿童死于锌缺乏症^[4] ...

... 小麦是我国北方地区的主要粮食作物, 黄淮麦区是最重要的商品粮基地, 提高人体对小麦籽粒中矿物质元素的吸收利用, 对于解决我国人民由于矿物质元素含量摄入不足造成的健康问题具有重要意义^[4,5] ...

... 影响铁锌等生物有效性的限制性因子包括植酸、纤维素、丹宁和重金属等, 以植酸最为重要^[2,3,4] ...

2007

0.0

1.889

Sci Agric Sin. 2007, 40(9):1871 - 1876

Variation of major mineral elements content and their relationships in Chinese wheats

北方冬麦区小麦品种籽粒主要矿物质元素含量分布及其相关性分析

(张勇),(王德森),(张艳),(何中虎).(北方冬麦区小麦品种籽粒主要矿物质元素含量分布及其相关性分析).(中国农业科学),2007,40:1871-1876

张勇,王德森,张艳,何中虎

中国农业科学院作物科学研究所/国家小麦改良中心/国家农作物基因资源与基因改良重大科学工程

Mineral elements including micronutrients as Fe and Zinc are very essential for human life. Two hundred and forty cultivars and advanced lines, originated from Beijing, Heibei, Henan, Shandong, Shanxi, and Shaanxi Provinces, were collected and sown in the experiment station of Crop Science Research Institute, in Chinese Academy of Agriculture Sciences in 1997-1998 wheat season, to evaluate the contents of major mineral elements including micronutrients as Fe, Ze, Mn, Cu, and macronutrients as Ca, Mg, K, P, S. Large variation for most of the mineral elements contents except Mg and S, and big difference between cultivars for all the mineral elements contents were observed. Among the micronutrients, Fe performed the highest average content, with value being of 41.9 mg/Kg, while the lowest for Cu, with value being of 5.54 mg/Kg. Among the macronutrients, K had the highest average content, with value being of 4747 mg/Kg, while the lowest for Ca, with value being of 465 mg/Kg. There were large differences for the mineral elements contents among cultivars with different origin, with cultivars from Beijing and Shandong performed high contents for most of the micronutrients, cultivars from Hebei and Shandong had high contents for most of the macronutrients, while cultivars from Henan Province had the lowest content for most of the mineral elements including micro- and macronutrients. Highly significant positive correlations between the contents of Fe and Zn (r = 0.630), Mg and P (r = 0.571), Mg and S (r = 0.521), Zn and P (r = 0.502), and P and S (r = 0.537) were also observed, indicating possibility for elevating these mineral elements contents at the same time. There was obvious pedigree relationship among the cultivars and advanced lines with high content of mineral elements as Fe and Zn, suggesting that parents with high mineral elements content and grain yield should be used first of all in crossing with the main objective being to improve mineral elements contents in wheat breeding program. It also indicated that in the current situation leading cultivars should be screened for mineral elements contents in order to be used directly of indirectly in wheat breeding program economically and effectively.

【目的】小麦品种籽粒铁和锌等微量营养元素含量研究对人体健康具有重要意义。【方法】来自北京、河北、河南、山东、山西和陕西等6省区240个小麦品种和高代品系于1997～1998年度种植在中国农业科学院作物科学研究所农场试验田，收获后分析籽粒中包括铁、锌、锰、铜等微量元素和钙、镁、钾、磷、硫等常量元素在内的主要矿物质元素含量。【结果】品种间各微量和常量矿物质元素含量均存在明显差异。微量元素中，铁平均含量最高，为41.9 mg•kg-1；铜含量最低，为5.54 mg•kg-1。常量元素中，钾平均含量最高，达4 747 mg•kg-1；钙含量最低，为465 mg•kg-1。除镁和硫元素外，其它各元素含量变幅均较大。不同地区来源品种籽粒中的矿物质元素含量差别较大，北京和山东品种铁、锌、铜等微量元素含量较高，河北和山东品种钙、镁、磷等常量元素含量较高，河南品种大部分微量和常量元素含量都较低。籽粒中铁和锌（r = 0.630）、镁与磷（r = 0.571）、镁与硫（r = 0.521）、锌与磷（r = 0.502）、磷与硫（r = 0.537）等元素含量间相关系数均较高，达0.001显著水平，表明可以同时提高小麦籽粒中铁、锌和磷等矿物质元素的含量。【结论】铁和锌等元素含量高的品种（系）间存在明显的亲源关系，说明在育种中通过选择或选用矿物质元素含量高的亲本可以提高矿物质元素的含量。并指出现阶段应进一步从中国现有小麦品种特别是各地主栽品种中筛选矿物质元素特别是铁和锌含量高的基因型，以经济有效地直接或间接利用这些材料。

... 田间采用拉丁方alpha格子设计^[5,22], 2次重复, 每隔15个品种加一个当地对照 ...

... 因此, 在改良小麦籽粒的微量矿物质营养品质时, 首先应对现有品种的目标性状进行筛选, 我们在对小麦品种铁锌等微量元素含量的分析时, 也有类似结论^[5] ...

2001

3.689

0.0

Appl Microbiol Biot. 2001, 57(4):474 - 481 DOI:10.1007/s002530100795

Biotechnological development of effective phytases for mineral nutrition and environmental protection

Lei

X G

,Stahl

C H

.Biotechnological development of effective phytases for mineral nutrition and environmental protection.Appl Microbiol Biot,2001,57:474-481

1.Department of Animal Science, 252 Morrison Hall, Cornell University, Ithaca, New York 14853, USA

... 植酸酶已作为添加剂广泛用于食品和饲料工业^[14], 采用微生物方法进行工业化生产成本较高, 生产过程中大量消耗能源并严重污染环境^[6] ...

1966

3.386

0.0

... 植酸是磷在小麦籽粒中的主要贮存形式, 通过与铁、锌、钙等二价金属离子结合, 形成螯合态植酸盐^[7], 降低人对铁、锌等的吸收利用^[8], 植酸含量与锌含量呈显著正相关^[9] ...

1970

0.0

Ann. N. Y. Acad. Sci.. 1969, 165(2):801-14

Formation of phytic acid in cereal grains.

Asada K, Tanaka K, Kasai Z

PMID: 5259621

2006

2.088

0.0

... 对印度和国际玉米小麦改良中心(CIMMYT)品种的植酸含量和植酸酶活性已进行了初步报道^[20], Liu等^[9]和吴澎等^[21]曾对我国的地方品种和河南、山东、陕西、江苏和四川等少数地区的品种进行了植酸含量和植酸酶活性分析 ...

... 87 g kg^-1, 植酸酶活性为620~2192 U kg^{-1 [9]} ...

... 37 g kg^-1和10~1759 U kg^-1, 与前人报道结果^[9,20,26]基本一致, 但本研究中植酸含量和植酸酶活性的变异范围更大, 可能与所选取材料的代表性和数量有关 ...

1996

0.0

DTW. Dtsch. Tierarztl. Wochenschr.. 1996, 103(6):209-14

Mechanisms of intestinal phosphorus absorption and availability of dietary phosphorus in pigs.

Schröder B, Breves G, Rodehutscord M

Department of Veterinary-Physiology, Justus-Liebig-University, Giessen, Germany.

Phosphorus (P) is an essential component of many organic and inorganic compounds in vertebrates such as pigs. Therefore, adequate dietary P supply is important to meet daily requirements in order to maintain P homeostasis. Under normal circumstances regulation of P homeostasis occurrs by controlling the absorption rate of inorganic phosphate (Pi) in the upper small intestines and by renal Pi excretion. These processes are mainly mediated by parathyroid hormone (PTH) and calcitriol (1,25-dihydroxycholecalciferol, 1,25-(OH)2D3). If, for example, the Pi level in plasma decreases, renal calcitriol production is stimulated and higher amounts of the hormone are released into the circulation. Calcitriol increases Pi absorption from the intestinal tract by stimulation of a secondary active, sodium-coupled Pi-cotransport system in the upper small intestines. In addition, calcitriol is involved in the mobilization of bone and soft tissue P. Simultaneously, hypercalcemia develops, which can be induced by either increased intestinal Ca absorption and/or Ca mobilization from bone. Hypophosphatemia and hypercalcemia suppress PTH release from the parathyroid glands and thus minimize urinary Pi losses. The concerted action of increased/decreased circulating calcitriol/PTH on the intestinal tract, bone and kidneys normalizes Pi levels in plasma. With respect to adequate P supply in animal nutrition, it must be considered that utilization of dietary P not only depends on absorption capacity of the pig intestinal tract but also on differences in availability of dietary P between ingredients. In feedstuffs of plant origin most of the P is bound as phytate-P and can only be absorbed after enzymatic breakdown of phytic acid by phytases. Intrinsic phytase activity differs between plant materials such as wheat, wheat bran, barley and triticale with higher activities than found in maize and legume seeds subjected to thermal treatments. Supplementation of microbial phytase increased P digestibility more pronounced in those feedstuffs which showed very limited intrinsic phytase activity. At present, a digestibility of about 70% seems to be the upper level for digestibility of P from plant material. From the environmental point of view, an increased digestibility resulting from phytase supplementation offers the possibility to reduce the supplementation of phosphates and the concentration of total P in the diet. Therefore, the amount of P being excreted by the pig can be remarkably reduced. However, the first step for minimizing faecal P excretion should be to supply P in accordance with the animal's requirement.

PMID: 8766902

... 食物中的植酸盐可以被植酸酶有效分解, 但人和猪等单胃动物的消化系统中植酸酶活性非常低^[10], 导致铁、锌和磷等矿物质元素无法被有效吸收, 同时大量的有机磷还随排泄物排出, 引起土壤和水污染^[11] ...

2002

11.808

0.0

Trends Plant Sci.. 2002, 7(3):118-25

Engineering crop plants: getting a handle on phosphate.

Brinch-Pedersen H, Sørensen LD, Holm PB

Danish Institute of Agricultural Sciences, Dept of Plant Biology, Research Centre Flakkebjerg, DK-4200, Slagelse, Denmark. henrik.brinchpedersen@agrsci.dk

In plant seeds, most of the phosphate is in the form of phytic acid. Phytic acid is largely indigestible by monogastric animals and is the single most important factor hindering the uptake of a range of minerals. Engineering crop plants to produce a heterologous phytase improves phosphate bioavailability and reduces phytic acid excretion. This reduces the phosphate load on agricultural ecosystems and thereby alleviates eutrophication of the aquatic environment. Improved phosphate availability also reduces the need to add inorganic phosphate, a non-renewable resource. Iron and zinc uptake might be improved, which is significant for human nutrition in developing countries.

PMID: 11906835

1999

1.713

0.0

2002

0.0

2004

3.689

0.0

Appl Microbiol Biot. 2004, 63(4):362 - 372 DOI:10.1007/s00253-003-1345-0

Biochemical properties and substrate specificities of alkaline and histidine acid phytases

B.-C. Oh(1) W.-C. Choi(1) S. Park(2) Y.-o. Kim(3) T.-K. Oh(1)

1.Microbial Genomics Laboratory Korea Research Institute of Bioscience and Biotechnology P.O. Box?115 305-600 Yusong Taejon Korea
2.Comparative Pharmacokinetics and Microbial Pharmacology College of Veterinary Medicine Kyungpook National University 702-701 Taegu Korea
3.Biotechnology Research Center National Fisheries Research and Development Institute 619-902 Busan Korea

Phytases are a special class of phosphatase that catalyze the sequential hydrolysis of phytate to less-phosphorylated myo-inositol derivatives and inorganic phosphate. Phytases are added to animal feedstuff to reduce phosphate pollution in the environment, since monogastric animals such as pigs, poultry, and fish are unable to metabolize phytate. Based on biochemical properties and amino acid sequence alignment, phytases can be categorized into two major classes, the histidine acid phytases and the alkaline phytases. The histidine acid phosphatase class shows broad substrate specificity and hydrolyzes metal-free phytate at the acidic pH range and produces myo-inositol monophosphate as the final product. In contrast, the alkaline phytase class exhibits strict substrate specificity for the calcium–phytate complex and produces myo-inositol trisphosphate as the final product. This review describes recent findings that present novel viewpoints concerning the molecular basis of phytase classification.

... 植酸酶已作为添加剂广泛用于食品和饲料工业^[14], 采用微生物方法进行工业化生产成本较高, 生产过程中大量消耗能源并严重污染环境^[6] ...

2008

2.609

0.0

Transgenic Res. 2008, 17(4):633 - 643 DOI:10.1007/s11248-007-9138-3

Transgenic maize plants expressing a fungal phytase gene

Rumei Chen(1) Guangxing Xue(2) Ping Chen(1) Bin Yao(3) Wenzhu Yang(1) Qianli Ma(1) Yunliu Fan(1) Zuoyu Zhao(4) Mitchell C. Tarczynski(4) Jinrui Shi(4)

1.Chinese Academy of Agricultural Sciences Biotechnology Research Institute Beijing 100081 P.R. China
2.Chinese Academy of Agricultural Sciences Crop Sciences Institute Beijing 100081 P.R. China
3.Chinese Academy of Agricultural Sciences Feed Research Institute Beijing 100081 P.R. China
4.Pioneer Hi-Bred International, A DuPont Company 7300 NW 62nd Ave Johnston IA 50131 USA

Maize seeds are the major ingredient of commercial pig and poultry feed. Phosphorus in maize seeds exists predominately in the form of phytate. Phytate phosphorus is not available to monogastric animals and phosphate supplementation is required for optimal animal growth. Undigested phytate in animal manure is considered a major source of phosphorus pollution to the environment from agricultural production. Microbial phytase produced by fermentation as a feed additive is widely used to manage the nutritional and environmental problems caused by phytate, but the approach is associated with production costs for the enzyme and requirement of special cares in feed processing and diet formulation. An alternative approach would be to produce plant seeds that contain high phytase activities. We have over-expressedAspergillus nigerphyA2 gene in maize seeds using a construct driven by the maize embryo-specific globulin-1 promoter. Low-copy-number transgenic lines with simple integration patterns were identified. Western-blot analysis showed that the maize-expressed phytase protein was smaller than that expressed in yeast, apparently due to different glycosylation. Phytase activity in transgenic maize seeds reached approximately 2,200 units per kg seed, about a 50-fold increase compared to non-transgenic maize seeds. The phytase expression was stable across four generations. The transgenic seeds germinated normally. Our results show that the phytase expression lines can be used for development of new maize hybrids to improve phosphorus availability and reduce the impact of animal production on the environment.

... 转植酸酶基因玉米已获生产应用安全证书并进行环境释放^[15], 但转基因作物受大众认知和接受程度的严重影响, 目前还难以大规模应用^[16] ...

... 按Chen等^[15]的方法分析植酸含量, 并适当改进 ...

... 参考Chen等^[15]的方法, 并略加调整, 分析植酸酶活性 ...

2010

0.0

0.49

Biotechnol Bull. 2010, 0(7):1 - 6

(转基因作物的生态安全性问题及其对策).(生物技术通报),2010, (7):1-6

.Ecological risks and countermeasures of genetically modified crops

转基因作物的生态安全性问题及其对策

(王延锋),(郎志宏),(赵奎军),(黄大昉).(转基因作物的生态安全性问题及其对策).(生物技术通报),2010, (7):1-6

王延锋;郎志宏;赵奎军;黄大昉;

随着转基因作物商业化进程的加快,转基因作物的安全性引起了人们越来越多的关注,主要就转基因作物生态安全性问题以及产生的原因进行综述,并提出我国现阶段应采取的对策。

... 转植酸酶基因玉米已获生产应用安全证书并进行环境释放^[15], 但转基因作物受大众认知和接受程度的严重影响, 目前还难以大规模应用^[16] ...

2010

1.643

0.0

Euphytica. 2010, 174(3):303 - 313 DOI:10.1007/s10681-009-0082-6

Mineral element concentrations in grains of Chinese wheat cultivars

Yong Zhang(1) Qichao Song(2) Jun Yan(3) Jianwei Tang(1) Rongrong Zhao(2) Yueqiang Zhang(2) Zhonghu He(14) Chunqin Zou(2) Ivan Ortiz-Monasterio(5)

1.Institute of Crop Science, National Wheat Improvement Centre/The National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agriculture Sciences (CAAS), 12 Zhongguancun South Street, Beijing, 100081 China
2.Department of Plant Nutrition, China Agricultural University, 2 West Yuanmingyuan Road, Haidian District, Beijing, 100193 China
3.Cotton Research Institute, Chinese Academy of Agricultural Sciences (CAAS), Huanghedadao, Anyang, 455000 Henan Province China
4.CIMMYT-China Office, c/o CAAS, 12 Zhongguancun South Street, Beijing, 100081 China
5.CIMMYT, Apdo. Postal 6-641, 066000 Mexico, DF Mexico

Investigations on concentration of mineral elements including Fe and Zn in wheat grains are important for human health. Two hundreds and sixty-five cultivars and advanced lines were collected and sown at Anyang experimental station of the Institute of Crop Science of the Chinese Academy of Agriculture Sciences in season 2005–2006 to evaluate the genetic variation of major mineral element concentrations in wheat grain. Twenty-four selected cultivars were also planted at seven representative locations in seasons 2005–2006 and 2006–2007 to evaluate the effects of genotype, environment, and genotype by environment interaction on mineral element concentrations. The 265 genotypes displayed a large variation for all mineral elements investigated including Fe and Zn, ranging from 28.0 to 65.4 mg kg⁻¹ and 21.4 to 58.2 mg kg⁻¹ for Fe and Zn, with mean values of 39.2 and 32.3 mg kg⁻¹, respectively. Jimai 26, Henong 326, and Jingdong 8 displayed high Fe and Zn concentrations, and Jimai 26 and Henong 326 also displayed high concentrations of Cu, Mg, K, P, and protein content. Jingdong 8 is the most promising leading cultivar for increasing Fe and Zn concentrations. All mineral element concentrations including Fe and Zn were largely influenced by environment effects. Production of high Fe concentration can be best secured at Jiaozuo and Jinan, and high Zn concentration can be best secured at Jinan and Xuzhou, since samples from these locations in the two seasons are characterized by high Fe or Zn concentration, compared with the other locations. High and significant genotype by environment interaction effects on all mineral element concentrations were also observed, with ratios of genotype by environment to genotype variances all larger than 1.20. Grain Fe concentration was highly significant and positively correlated with that of Zn, indicating a high possibility to combine high Fe and Zn traits in wheat breeding. It also indicated strong positive correlations between concentrations of Fe, Zn, and protein content.

... 国内外对小麦的铁、锌等元素含量已进行了较深入研究, 品种间存在显著差异, 并受环境显著影响^[17] ...

... 3 数据处理采用合适的空间模型^[17]分析试验数据, 将两年度所得品种各性状最佳线性无偏预测值用于数据处理和分析^[24] ...

2009

1.995

0.0

J. Hered.. 2009, 100(6):771-6 DOI:10.1093/jhered/esp030

Mapping of quantitative trait Loci for grain iron and zinc concentration in diploid A genome wheat.

Tiwari VK, Rawat N, Chhuneja P, Neelam K, Aggarwal R, Randhawa GS, Dhaliwal HS, Keller B, Singh K

Department of Biotechnology, Indian Institute of Technology, Uttarakhand, India.

Micronutrients, especially iron (Fe) and zinc (Zn), are deficient in the diets of people in underdeveloped countries. Biofortification of food crops is the best approach for alleviating the micronutrient deficiencies. Identification of germplasm with high grain Fe and Zn and understanding the genetic basis of their accumulation are the prerequisites for manipulation of these micronutrients. Some wild relatives of wheat were found to have higher grain Fe and Zn concentrations compared with the cultivated bread wheat germplasm. One accession of Triticum boeoticum (pau5088) that had relatively higher grain Fe and Zn was crossed with Triticum monococcum (pau14087), and a recombinant inbred line (RIL) population generated from this cross was grown at 2 locations over 2 years. The grains of the RIL population were evaluated for Fe and Zn concentration using atomic absorption spectrophotometer. The grain Fe and Zn concentrations in the RIL population ranged from 17.8 to 69.7 and 19.9 to 64.2 mg/kg, respectively. A linkage map available for the population was used for mapping quantitative trait loci (QTL) for grain Fe and Zn accumulation. The QTL analysis led to identification of 2 QTL for grain Fe on chromosomes 2A and 7A and 1 QTL for grain Zn on chromosome 7A. The grain Fe QTL were mapped in marker interval Xwmc382-Xbarc124 and Xgwm473-Xbarc29, respectively, each explaining 12.6% and 11.7% of the total phenotypic variation and were designated as QFe.pau-2A and QFe.pau-7A. The QTL for grain Zn, which mapped in marker interval Xcfd31-Xcfa2049, was designated as QZn.pau-7A and explained 18.8% of the total phenotypic variation.

PMID: 19520762

... 铁含量相关基因/主效QTL定位在2A、4A、5A、7A和4D等染色体上, 锌含量相关基因/主效QTL定位在1A、2D、3A、4A、4D、5A和7A等染色体上^[18,19] ...

2008

2.638

0.0

Plant Soil. 2008, 306(1-2):95 - 104 DOI:10.1007/s11104-007-9483-2

Identification of quantitative trait locus of zinc and phosphorus density in wheat (Triticum aestivum L.) grain

Rongli Shi(1) Hongwei Li(2) Yiping Tong(2) Ruilian Jing(3) Fusuo Zhang(1) Chunqin Zou(1)

1.China Agricultural University Department of Plant Nutrition 2 West Yuanmingyuan Road, Haidian District Beijing 100094 China
2.Chinese Academy of Sciences The State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology Beijing 100101 China
3.Chinese Academy of Agricultural Sciences Institute of Crop Science 100081 Beijing China

Zinc (Zn) is an essential micronutrient for human beings. However, Zn malnutrition has become a major problem throughout the world. Wheat is the most important food crop in the world, therefore, developing Zn-enriched wheat varieties provides an effective approach to reduce Zn malnutrition in human beings. The aim of this study was to understand the genetic control of grain Zn density in wheat and hence, to provide genetic basis for breeding wheat with high grain Zn density using molecular approach. A doubled haploid (DH) population developed from a cross between winter wheat varieties Hanxuan10 and Lumai 14 was used to identify quantitative trait loci (QTLs) for Zn concentration and content in wheat grains. In addition, phosphorus (P) concentration and content in wheat grain were also investigated to examine possible interactions between these two nutrients. The wheat grains used in this study were harvested from the plants grown under normal condition in a field trial. We found the grain Zn concentrations of the DH population varied from 25.9 to 50.5 mg/kg and the Zn content varied from 0.90 to 2.21 μg/seed. The grain P concentrations of the DH population varied from 0.258 to 0.429 mg/kg, and the P contents varied from 0.083 to 0.186 mg/seed. A significant positive correlation was observed between Zn and P density in this experiment. The results showed that both grain Zn and P densities were controlled by polygenes. Four and seven QTLs for Zn concentration and Zn content were detected, respectively. All the four QTLs for Zn concentration co-located with the QTLs for Zn content, suggesting a possibility to improve both grain Zn concentration and content simultaneously. Four and six QTLs for P concentration and P content were detected, respectively. The two QTLs for grain Zn concentration on chromosomes 4A and 4D co-located with the QTLs for P concentration. The four QTLs for grain Zn content on chromosome 2D, 3A and 4A co-located with the QTLs for P contents, reflecting the positive correlations between the grain Zn and P density, and providing possibility of improving grain micro- and macronutrient density simultaneously in wheat. In order to improve human health, the effect of P–Zn relation in grain on the Zn bioavailability should also be considered in the future work.

... 铁含量相关基因/主效QTL定位在2A、4A、5A、7A和4D等染色体上, 锌含量相关基因/主效QTL定位在1A、2D、3A、4A、4D、5A和7A等染色体上^[18,19] ...

2010

2.088

0.0

J Cereal Sci. 2010, 52(3):486 - 490 DOI:10.1016/j.jcs.2010.08.003

Large variability exits in phytase levels among Indian wheat varieties and synthetic hexaploids

Ram

,Verma

,Sharma

.Large variability exits in phytase levels among Indian wheat varieties and synthetic hexaploids.J Cereal Sci,2010,52:486-490

Abstract

Four hundred wheat genotypes including released varieties in India, advanced lines and synthetic hexaploids were used in the present investigation to assess the variability in phytase and phytate levels. Three different extraction processes such as direct, syringe and spin column methods were used for assessing phytase levels. The direct assay method showed greater potential for use in assaying enzyme activity in large numbers of samples over other methods and gave reproducible results (R² = 0.87). The extraction and assay of the enzyme was faster and more cost effective compared to other methods. There were 3.4 fold variations in phytase levels among varieties and 5.9 fold among synthetic hexaploids. In contrast, lower variability (1.6 fold in varieties and 2.2 fold in synthetic hexaploids) was observed in phytate levels. Ten varieties grown in 4 locations in India were used to assess the genotypic and environmental effect on phytase and phytate levels. There were significant effects of both genotype and environment on phytate and phytase levels; it was more genetic with heritability of 0.98 and 0.82 for phytase and phytate, respectively. Higher variability in phytase levels and more genotypic effect on the enzyme activity demonstrated the potential of improving phytase levels through breeding.

... 3 g kg^-1与284~962 U kg^{-1 [20]} ...

2010

0.0

... 63 g kg^{-1 [21]} ...

1997

0.0

... 田间采用拉丁方alpha格子设计^[5,22], 2次重复, 每隔15个品种加一个当地对照 ...

1999

3.678

0.0

... g离心15 min后取上清液, 按Wyss等^[23]的方法, 通过测定无机磷的增加量来确定植酸酶的活性 ...

2006

0.0

1.889

Sci Agric Sin. 2006, 39(4):655 - 663

Adaptation of CIMMYT Wheat Germplasm in China's Spring Wheat Regions

CIMMYT小麦在中国春麦区的适应性分析

,,,,

张勇,吴振录,张爱民,Maarten van Ginkel,何中虎

中国农业科学院作物科学研究所

【Objective】Information on adaptation of CIMMYT wheat germplasm in China would greatly enhance wheat breeding efficiency in China's spring wheat regions. 【Methed】Twenty-five spring wheat cultivars including 10 from CIMMYT and 15 from China, were sown at 9 locations in China and 4 management environments in Obregon Station at CIMMYT in 2001 and 2002 seasons, and grain yield, yield components, and other agronomic traits were investigated. 【Result】CIMMYT cultivars were characterized by more spikes and grains per spike, medium thousand kernel weight, and high grain yield with broad adaptability. The photoperiod sensitive cultivars from Heilongjiang province performed tall plant height, late maturity, medium spike number, but low grains per spike, thousand-kernel weight, and grain yield. The other Chinese cultivars had medium plant height, early maturity, low spike number and medium grains per spike, but high thousand kernel weight, and medium grain yield. CIMMYT cultivars performed short plant height, earlier maturity, and a little lower grain yield when planted in China's spring wheat regions compared with that in CIMMYT. The photoperiod sensitive cultivars had taller plant height, later maturity, and much lower grain yield, while the other Chinese cultivars showed taller plant height, a little later maturity, and a little lower grain yield with almost the same thousand kernel weight when they were planted in CIMMYT. 【Conclusion】It is recommended to directly use CIMMYT wheat in production in Yunnan, Qinghai, and Xinjiang provinces. CIMMYT wheat can also be used in production in such provinces as Inner Mongolia, Gansu, and Ningxia provinces, and are more suitable as crossing parents in these areas; while only can be used as crossing parents in Heilongjiang province. In order to improve the shuttle breeding efficacy, and taking into account the genotype mean repeatability for major traits, Chinese scientists should focus on lines with large grain size when selecting materials in CIMMYT for China, choosing lines with short plant height, late maturity for Yunnan, and lines with a little tall plant height and early maturity for Inner Mongolia, Gansu, Ningxia and Xinjiang, lines with a little tall plant height and around the same maturity for Qinghai, while lines for high latitude environment with a little tall plant height and late maturity for Heilongjiang.

【目的】研究CIMMYT小麦在中国的适应性有助于提高春麦区的育种水平。【方法】10份CIMMYT代表性品种和15份中国春麦主栽品种于2001和2002年种植在中国春麦区的9个试点和CIMMYT的4种不同处理环境，分析产量、产量构成因子和农艺性状的变化趋势。【结果】CIMMYT品种穗数和穗粒数多，千粒重中等，具有广泛适应性，比中国品种具有更高的产量优势；黑龙江光敏感品种植株高、抽穗和成熟晚、穗数中等、穗粒数少、千粒重和产量低；中国其它品种株高中等、抽穗和成熟早、穗数少、穗粒数中等、千粒重高、产量中等。CIMMYT品种引种到中国后，株高降低，抽穗和成熟提早，并略减产；黑龙江光敏感品种在CIMMYT种植时株高增加，抽穗和成熟推迟，千粒重降低，并显著减产；中国其它品种在CIMMYT种植时株高增加、抽穗和成熟略推迟、千粒重变化较小，并略减产。【结论】CIMMYT品种可在云南、青海和新疆直接推广种植；内蒙古、甘肃和宁夏为其次适宜地区，可以直接推广应用，但主要用作杂交亲本；在黑龙江以作杂交亲本为宜。为提高引种效率，并考虑到性状的重复力大小，在CIMMYT为中国选种时应重点选择籽粒较大的材料。为云南所选材料可略矮、适当晚熟，内蒙古、甘肃、宁夏和新疆所选材料可略高、较早熟，青海所选材料可较高、熟期相当，黑龙江应主要选择高纬度材料、植株偏高且晚熟。

... 3 数据处理采用合适的空间模型^[17]分析试验数据, 将两年度所得品种各性状最佳线性无偏预测值用于数据处理和分析^[24] ...

1963

1.834

0.0

... 调用PROC CLUSTER时, 将植酸含量和植酸酶活性数据分别按年度进行标准化后, 以欧氏距离为标准, 按Ward类平方和法分别对品种进行聚类^[25] ...

2007

2.088

0.0

J Cereal Sci. 2007, 45(3):319 - 326 DOI:10.1016/j.jcs.2006.10.004

Phytase activity, phytate, iron, and zinc contents in wheat pearling fractions and their variation across production locations

Liu

Z H

,Wang

H Y

,Wang

X E

,Zhang

G P

,Chen

P D

,Liu

D J

.Phytase activity, phytate, iron, and zinc contents in wheat pearling fractions and their variation across production locations.J Cereal Sci,2007,45:319-326

Abstract

Pearling is an effective method for evaluating the distribution of chemical components in wheat grain. Twelve pearling fractions (P₁–P₁₂) of wheat grain were obtained using two rice polishers for 10 cultivars (six soft red wheats and four hard white wheats) grown at two locations with different environmental conditions in Jiangsu Province, China. The results show that the effects of cultivar, location, and pearling on wheat flour phytase activity, phytate, iron, and zinc contents were all significant, with pearling having the greatest effect. All the four components showed a diminishing trend as pearling progressed from the outer layers to the inner part of wheat grain. Generally, the P₂ fraction (the outer 4–8% layer of wheat grain) had the highest phytase activity and phytate and iron contents, whereas the P₁ fraction (the outer 0–4% layer) ranked the highest for zinc content. Growing location had a large influence on grain phytase, phytate, and iron, but the differences between locations decreased as pearling level increased.

... 植酸含量比植酸酶活性更易受年度效应的影响, 这些结果与前人报道一致^[26,27] ...

2002

1.328

0.0

... 植酸含量比植酸酶活性更易受年度效应的影响, 这些结果与前人报道一致^[26,27] ...

... Kim等^[27]选用3个品种在多个环境中种植, 结果表明植酸含量受降雨量等环境因素的显著影响, 而植酸酶活性的环境效应不显著 ...