水稻锌吸收、转运、累积机理研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2020-1484

摘要/Abstract

摘要：

锌是水稻正常生长发育所必需的微量营养元素之一,是生长素、叶绿素合成等多种酶促反应的辅助因子。水稻缺锌时,植株矮小,叶片生长受到抑制且失绿变白,抽穗晚而少,严重影响产量;锌过量又容易对水稻产生毒害作用。因此研究水稻锌元素吸收、转运和累积机制,通过现代分子生物技术对水稻籽粒锌富集进行生物强化具有重要意义。基于近年来国内外研究报道,简要概述了锌在水稻体内的生理功能,水稻不同营养器官对锌的吸收途径以及影响锌元素吸收积累的因素,并对近年来水稻中鉴定出的锌转运蛋白及其家族成员的种类、结构以及功能进行了归纳总结,旨为进一步了解水稻体内的锌转运系统及其调控提供参考。

关键词: 水稻, 锌, 吸收转运, 锌转运蛋白, 生物强化

Abstract:

Zinc is one of the micronutrient elements necessary for the normal growth and development in rice,and is the auxiliary factor in biosynthesis of auxin,chlorophyll and other chemicals. Zinc deficiency can lead to drawf,slow growth of new leaves,chlorosis and white midrib in leaves,late and less heading,which seriously affects the yield. In addition,excessive zinc may cause toxic effects on rice. Therefore,it is of great significance to study the mechanisms of zinc absorption,transportation and accumulation in rice and to biofortificate the enrichment of zinc in rice grain by modern biotechnology. Based on the recent reports in domestic and abroad,this paper briefly reviewed the physiological function of zinc in rice,zinc absorption pathways in different tissues of rice,and factors affecting the absorption and accumulation of zinc. Further this paper sorted and summarized the types,structures and functions of zinc transporters,and their family members identified in rice in recent years,aiming to provide references for further understanding the zinc transport system and its regulatory mechanism in rice.

Key words: rice, zinc, absorption and transport, zinc transporter, biofortification

薛欣月, 于雪然, 刘晓刚, 马嘉欣, 田蕾, 李培富. 水稻锌吸收、转运、累积机理研究进展[J]. 生物技术通报, 2022, 38(4): 29-43.

XUE Xin-yue, YU Xue-ran, LIU Xiao-gang, MA Jia-xin, TIAN Lei, LI Pei-fu. Research Progress in Absorption,Transportation and Accumulation Mechanism of Zinc in Rice[J]. Biotechnology Bulletin, 2022, 38(4): 29-43.

图/表 9

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酶Enzyme	所在部位Location	可能功能Possible function	分类Classification	参考文献Reference
碳酸酐酶	叶绿体外膜	参与光合作用	含锌酶	[5-6]
蛋白酶	茎叶	水解蛋白质肽链	含锌酶	[7]
谷氨酸脱氢酶	线粒体	参与植物碳氮代谢	含锌酶	[8]
醛缩酶	细胞质	参与光合作用碳代谢	含锌酶	[7]
铜锌超氧化物歧化酶	叶绿体	提高植物抗逆性	含锌酶	[9-10]
RNA聚合酶	细胞核	参与蛋白质合成	需锌酶	[7]
过氧化氢酶	叶绿体	参与光呼吸、共生性氮固定	需锌酶	[7]
核酮糖-1,5-二磷酸羧化酶	叶绿体	参与光合作用碳同化、光呼吸	需锌酶	[10-11]
1,6-二磷酸果糖酶	叶绿体	光合产物转运	需锌酶	[7]
过氧化物酶	过氧化物酶体	促进光合作用、呼吸作用	需锌酶	[12]

酶Enzyme	所在部位Location	可能功能Possible function	分类Classification	参考文献Reference
碳酸酐酶	叶绿体外膜	参与光合作用	含锌酶	[5-6]
蛋白酶	茎叶	水解蛋白质肽链	含锌酶	[7]
谷氨酸脱氢酶	线粒体	参与植物碳氮代谢	含锌酶	[8]
醛缩酶	细胞质	参与光合作用碳代谢	含锌酶	[7]
铜锌超氧化物歧化酶	叶绿体	提高植物抗逆性	含锌酶	[9-10]
RNA聚合酶	细胞核	参与蛋白质合成	需锌酶	[7]
过氧化氢酶	叶绿体	参与光呼吸、共生性氮固定	需锌酶	[7]
核酮糖-1,5-二磷酸羧化酶	叶绿体	参与光合作用碳同化、光呼吸	需锌酶	[10-11]
1,6-二磷酸果糖酶	叶绿体	光合产物转运	需锌酶	[7]
过氧化物酶	过氧化物酶体	促进光合作用、呼吸作用	需锌酶	[12]

染色体 Chromosome	群体 Group	标记 Marker	LOD	贡献率/% Contribution rate/%	参考文献 Reference
Chr1	Asominori/IR24RILs	XNpb93-C3029C	6.00	21.90	[53]
Chr1	热研/华占F₁₂RILs	-	7.29	-	[54]
Chr1	龙锦1号/香软米1578F₂	RM446-RM9	1.61	4.39	[55]
Chr2	红香1号/松98-131F_2：3家系	RM450-RM327	7.24	35.40	[56]
Chr2	Nipponbare/MeridionalisW1627BC₂F₆	RM573	6.90	15.20	[57]
Chr2	Nipponbare/MeridionalisW1627BC₂F₆	RM6	6.30	17.60	[57]
Chr3	协青早B/中恢9308RILs	RM6283-RM7370	2.72	5.60	[58]
Chr3	奉新红米/明恢100F₂	RM186-RM168	1.87	12.75	[59]
Chr4	香黑糯195/中花紫香糯F₂	RM5414-RM6659	1.92	4.56	[60]
Chr4	川香29B/LemontRILs	RM3308-RM3471	4.57	13.82	[61]
Chr5	红香1号/松98-131F_2：3家系	RM206-RM430	9.81	24.80	[56]
Chr5	郑山97/明慧RILs	R3166-RG360	4.27	12.34	[62]
Chr6	奉新红米/明恢100F₂	RM435-RM469	1.58	4.97	[59]
Chr6	珍汕97B/密阳46RILs	RM204-RM225	4.18	6.50	[63]
Chr6	十田和/丽粳2号BC₅F₆	RM4608-RM6119	2.22	5.00	[64]
Chr7	宁粳35号/杨和白皮稻F₂	RM6018-RM5508	-	5.00-8.00	[65]
Chr8	龙锦1号/香软米1578F₂	RM152-RM3572	1.83	4.00	[55]
Chr8	香黑糯195/中花紫香糯F₂	RM447-RM7356	5.88	13.80	[60]
Chr8	香黑糯195/中花紫香糯F₂	RM7356-RM1109	5.60	14.37	[60]
Chr9	宁大62/宁粳40号F₂	RM285-RM219	-	11.00	[66]
Chr9	红香1号/松98-131F_2：3家系	RM410-RM566	10.85	24.07	[56]
Chr10	Asominori/IR24RILs	C751B-C148	2.20	7.60	[53]
Chr11	奉新红米/明恢100F₂	RM212-RM144	1.52	7.74	[59]
Chr11	郑山97/明慧RILs	C794-RG118	5.65	18.61	[62]
Chr12	龙锦1号/香软米1578F₂	RM1036-RM1986	1.55	5.21	[55]
Chr12	Madhukar/SwarnaRILs	RM260-RM7102	2.90	34.00	[67]
Chr12	红香1号/松98-131F_2：3家系	RM1300-RM1227	4.50	46.80	[56]

染色体 Chromosome	群体 Group	标记 Marker	LOD	贡献率/% Contribution rate/%	参考文献 Reference
Chr1	Asominori/IR24RILs	XNpb93-C3029C	6.00	21.90	[53]
Chr1	热研/华占F₁₂RILs	-	7.29	-	[54]
Chr1	龙锦1号/香软米1578F₂	RM446-RM9	1.61	4.39	[55]
Chr2	红香1号/松98-131F_2：3家系	RM450-RM327	7.24	35.40	[56]
Chr2	Nipponbare/MeridionalisW1627BC₂F₆	RM573	6.90	15.20	[57]
Chr2	Nipponbare/MeridionalisW1627BC₂F₆	RM6	6.30	17.60	[57]
Chr3	协青早B/中恢9308RILs	RM6283-RM7370	2.72	5.60	[58]
Chr3	奉新红米/明恢100F₂	RM186-RM168	1.87	12.75	[59]
Chr4	香黑糯195/中花紫香糯F₂	RM5414-RM6659	1.92	4.56	[60]
Chr4	川香29B/LemontRILs	RM3308-RM3471	4.57	13.82	[61]
Chr5	红香1号/松98-131F_2：3家系	RM206-RM430	9.81	24.80	[56]
Chr5	郑山97/明慧RILs	R3166-RG360	4.27	12.34	[62]
Chr6	奉新红米/明恢100F₂	RM435-RM469	1.58	4.97	[59]
Chr6	珍汕97B/密阳46RILs	RM204-RM225	4.18	6.50	[63]
Chr6	十田和/丽粳2号BC₅F₆	RM4608-RM6119	2.22	5.00	[64]
Chr7	宁粳35号/杨和白皮稻F₂	RM6018-RM5508	-	5.00-8.00	[65]
Chr8	龙锦1号/香软米1578F₂	RM152-RM3572	1.83	4.00	[55]
Chr8	香黑糯195/中花紫香糯F₂	RM447-RM7356	5.88	13.80	[60]
Chr8	香黑糯195/中花紫香糯F₂	RM7356-RM1109	5.60	14.37	[60]
Chr9	宁大62/宁粳40号F₂	RM285-RM219	-	11.00	[66]
Chr9	红香1号/松98-131F_2：3家系	RM410-RM566	10.85	24.07	[56]
Chr10	Asominori/IR24RILs	C751B-C148	2.20	7.60	[53]
Chr11	奉新红米/明恢100F₂	RM212-RM144	1.52	7.74	[59]
Chr11	郑山97/明慧RILs	C794-RG118	5.65	18.61	[62]
Chr12	龙锦1号/香软米1578F₂	RM1036-RM1986	1.55	5.21	[55]
Chr12	Madhukar/SwarnaRILs	RM260-RM7102	2.90	34.00	[67]
Chr12	红香1号/松98-131F_2：3家系	RM1300-RM1227	4.50	46.80	[56]

家族 Family	基因 Gene	物种 Species	表达部位 Expression position	表达诱导条件 Expression induction conditions	基因转运功能 Gene transfer function	参考文献 Reference
ZIP家族	OsZIP1	水稻	根、花穗	缺Zn	吸收、转运Zn	[81]
ZIP家族	OsZIP2	水稻	根	缺Zn	吸收、分配Zn	[62,79]
ZIP家族	OsZIP3	水稻	根、叶	缺Zn	分配Zn	[76]
ZIP家族	OsZIP4	水稻	地上部、根	缺Zn	转运Zn	[76]
ZIP家族	OsZIP5	水稻	根	缺Zn、Mn	吸收Zn	[76]
ZIP家族	OsZIP6	水稻	地上部、根	缺Zn、Mn、Fe	吸收、分配Zn	[76]
ZIP家族	OsZIP7	水稻	地上部、根	缺Zn	吸收Zn	[76]
ZIP家族	OsZIP8	水稻	地上部、根	缺Fe	吸收、分配Zn	[76-77]
ZIP家族	OsIRT1	水稻	茎的韧皮部、根	缺Fe	吸收、转运Zn、Fe、Cd	[77]
NRAMP家族	OsNRAMP2	水稻	根	缺Zn	吸收、转运Zn	[82-83]
NRAMP家族	OsNRAMP3	水稻	根	缺Zn	吸收、转运Zn	[82-83]