Research Advances on the Low Phytic Acid Rice Breeding and Their Genetic Physiological Regulation and Environmental Adaptability

doi:10.16819/j.1001-7216.2019.8083 95

Abstract

Abstract:

Breeding variety with improved quality while maintaining or improving yields is one of the primary objectives in rice breeding. Among which, reducing the anti-nutritional factors, such as grain phytic acid content, is an effective strategy to cope with hidden hunger and increase grain bioavailabilities of zinc and iron. In this paper, we reviewed the biosynthesis of phytic acid and the genetic characteristics of related functional genes, the co-regulatory networks of phytic acid synthesis and other physiological metabolism, breeding of low phytic acid (lpa) germplasm resource and their genetic characteristics, agronomic performance and environmental ecological adaptability of lpa mutants, the possible reasons for their agronomic deterioration and ecological adaptation change, and the environmental regulation of grain phytic acid accumulation. Those contents could provide reference for production of lpa rice with suitable agronomic cultivation practices.

Key words: phytic acid, rice (Oryza sativa L.), grain nutrition, genetic regulation, ecological effect

摘要：

提高或维持水稻产量的同时,提高稻米品质已成为目前水稻育种的首要目标之一。其中,通过降低籽粒中植酸等抗营养因子,增加锌、铁生物有效性以提升水稻营养品质,是目前水稻品质改良的一个重要方向。本文主要综述了水稻籽粒中植酸合成的代谢路径、低植酸水稻的筛选及相关功能基因的遗传特点、植酸生理代谢的调控网络、低植酸水稻农艺性状劣变和生态适应性降低的生理原因、籽粒植酸合成的环境调控效应等相关研究进展。可为低植酸水稻品质改良以及栽培调优提供借鉴。

关键词: 植酸, 水稻, 籽粒品质, 遗传调控, 生态效应

CLC Number:

Da SU, Liangquan WU, K Rasmussen Søren, Lujian ZHOU, Fangmin CHENG. Research Advances on the Low Phytic Acid Rice Breeding and Their Genetic Physiological Regulation and Environmental Adaptability[J]. Chinese Journal OF Rice Science, 2019, 33(2): 95-107.

苏达, 吴良泉, SørenKRasmussen, 周庐建, 程方民. 低植酸水稻种质资源筛选、遗传生理调控与环境生态适应性研究进展[J]. 中国水稻科学, 2019, 33(2): 95-107.

Figures/Tables 2

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突变方法 Origin of mutation	基因位点 Locus	低植酸突变体名称 Name of lpa	植酸含量降幅 Reduction in Phytic acid /%	无机磷变化 Pi variation	总磷变化 Total P variation	产量劣变 Yield inferiority	其他 Other phenotypic changes	参考文献 Reference
antisense under Ole18	MIPS (RION1)	--	68-75	增加 (等摩尔)	--	否	植酸变化仅影响胚乳和糊粉层	^[23]
antisense under 35S	MIPS (RION1)	--	--	--	不变	--	--	^[24]
RNAi under GluB-1	MIPS (RION1)	lpa1	17	增加 (等摩尔)	--	--	--	^[26]
RNAi	MIPS	--	--	增加 (等摩尔)	--	--	MIPS基因下调4.59倍;阳离子含量增加, 其中精米中铁含量增加1.6倍;肌醇和抗坏血酸含量降低	^[27]
RNAi under Oleosin 18	IPK1	--	69(T₄代)	增加 (等摩尔)	不变	否	IPK1 转录表达下调3.85倍;胚乳中铁含量增加1.8倍	^[32]
EMS	ITPK	--	46-68	--	--	是	--	^[31]
γ-rays + sodium zide	MIK	Os-lpa-XS110-1	34-64	--	--	是.	23个蛋白表达量增加;肌醇,果聚糖, 半乳糖和半乳糖苷含量增加;低价磷酸肌醇盐未检出	^{[52, 53, 58, 59]}
EMS	MIK	lpa N15-186	34-75	--	--	--	--	^[28]
Gene silence (amiMIK)	MIK	--	37.0-50.7,	增加 3.2-4.8倍	--	--	OsMIK转录表达降低,不同株系总磷含量变化不一致	^[25]
Gene silence (hpMIK)	MIK	--	14.9-50.2	增加 1.8-4.4倍	--	--	OsMIK转录表达降低,不同株系总磷含量变化不一致	^[25]
γ-rays + sodium zide	MRP	Os-lpa-XS110-2	20	--	--	是	肌醇和寡聚糖降低	^{[52, 53, 58]}
γ-rays + sodium zide	MRP	Os-lpa-XS110-3	约100	增加10倍	不变	致死	肌醇含量增加400%;植酸量<0.20 mg/g	^{[34, 52]}
T-DNA insertion	MRP	--	90	增加10倍	不变	致死	--	^[52]
RNAi under Oleosin 18	MRP	--	35.8-71.9	增加 7.5 倍	--	是	脂和核酸中的磷含量降低	^[45]
γ-rays	--	Os-lpa-XQZ-3	--	--	--	致死	--	^[52]
γ-rays	Sultr3;3	--	44	增加 (等摩尔)	--	是	--	^[52]
γ-rays	Sultr3;3	lpa-Z9B-1	45	--	降低	是	--	^{[52, 53]}
γ-rays	Sultr3;3	lpa-MH86-1	43.9-35.2	增加108%-170%	降低	是	糖、小分子籽粒内含物、丝氨酸、赖氨酸、硫、磷酸、 GABA活性增加;植酸、硫酸盐和磷酸盐基因表达量改变	^{[52, 53]}
γ-rays	2-PGK	KBNT lpa1-1	39-71	增加5%-32%	--	减产10%	植酸蛋白体体积减小;低价肌醇磷酸盐增加	^[28,29,30]
⁶⁰Co-γ	2-PGK	DR1331-2	45	增加 (等摩尔)	不变	否	--	^{[29, 30]}
γ-rays	--	lpa1	45	增加 (等摩尔)	不变	否	胚中植酸降幅高于糊粉层;钙, 锰降低、锌增加; 无低价磷酸盐	^{[16, 60]} ^{[61, 62]}
⁶⁰Co-γ+ NaN₃	--	Os-lpa-R6547-3	--	--	--	致死	--	^[52]
EMS	--	Dontokoi176	15	--	增加12%	否	垩白增幅率较低,铁,钼含量降低,其它矿质元素增加	^[63]
EMS	--	Koshihikari2623	45	--	不变	否	垩白增幅率较高,萌发前期活性较低	^[63]
γ-rays	--	Koshihikari4019	36	--	不变	否	垩白增幅率较高	^[63]
Callus culture	--	NC1857	17.1	增加 (等摩尔)	不变	小幅降低	铜、锰、钠、锌增加,钼降低41.9%	^[63]
EMS	--	Dontokoil 76	14.7	增加	增加 22%	小幅降低	增加了43.4%的铜,33.4%的钾,20.4%的镁,11.5%的钠, 39.1%的磷,23.6%的硫,10.8%的二氧化硅,44.2%的锌。而降低铁13.8%,钼17.5%	^[63]
γ-rays	--	Koshihikari49	5.9	增加	增加 9.4%	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari2623	44.7	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari3847 Koshihikari	47.9	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari4019	35.9	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]

突变方法 Origin of mutation	基因位点 Locus	低植酸突变体名称 Name of lpa	植酸含量降幅 Reduction in Phytic acid /%	无机磷变化 Pi variation	总磷变化 Total P variation	产量劣变 Yield inferiority	其他 Other phenotypic changes	参考文献 Reference
antisense under Ole18	MIPS (RION1)	--	68-75	增加 (等摩尔)	--	否	植酸变化仅影响胚乳和糊粉层	^[23]
antisense under 35S	MIPS (RION1)	--	--	--	不变	--	--	^[24]
RNAi under GluB-1	MIPS (RION1)	lpa1	17	增加 (等摩尔)	--	--	--	^[26]
RNAi	MIPS	--	--	增加 (等摩尔)	--	--	MIPS基因下调4.59倍;阳离子含量增加, 其中精米中铁含量增加1.6倍;肌醇和抗坏血酸含量降低	^[27]
RNAi under Oleosin 18	IPK1	--	69(T₄代)	增加 (等摩尔)	不变	否	IPK1 转录表达下调3.85倍;胚乳中铁含量增加1.8倍	^[32]
EMS	ITPK	--	46-68	--	--	是	--	^[31]
γ-rays + sodium zide	MIK	Os-lpa-XS110-1	34-64	--	--	是.	23个蛋白表达量增加;肌醇,果聚糖, 半乳糖和半乳糖苷含量增加;低价磷酸肌醇盐未检出	^{[52, 53, 58, 59]}
EMS	MIK	lpa N15-186	34-75	--	--	--	--	^[28]
Gene silence (amiMIK)	MIK	--	37.0-50.7,	增加 3.2-4.8倍	--	--	OsMIK转录表达降低,不同株系总磷含量变化不一致	^[25]
Gene silence (hpMIK)	MIK	--	14.9-50.2	增加 1.8-4.4倍	--	--	OsMIK转录表达降低,不同株系总磷含量变化不一致	^[25]
γ-rays + sodium zide	MRP	Os-lpa-XS110-2	20	--	--	是	肌醇和寡聚糖降低	^{[52, 53, 58]}
γ-rays + sodium zide	MRP	Os-lpa-XS110-3	约100	增加10倍	不变	致死	肌醇含量增加400%;植酸量<0.20 mg/g	^{[34, 52]}
T-DNA insertion	MRP	--	90	增加10倍	不变	致死	--	^[52]
RNAi under Oleosin 18	MRP	--	35.8-71.9	增加 7.5 倍	--	是	脂和核酸中的磷含量降低	^[45]
γ-rays	--	Os-lpa-XQZ-3	--	--	--	致死	--	^[52]
γ-rays	Sultr3;3	--	44	增加 (等摩尔)	--	是	--	^[52]
γ-rays	Sultr3;3	lpa-Z9B-1	45	--	降低	是	--	^{[52, 53]}
γ-rays	Sultr3;3	lpa-MH86-1	43.9-35.2	增加108%-170%	降低	是	糖、小分子籽粒内含物、丝氨酸、赖氨酸、硫、磷酸、 GABA活性增加;植酸、硫酸盐和磷酸盐基因表达量改变	^{[52, 53]}
γ-rays	2-PGK	KBNT lpa1-1	39-71	增加5%-32%	--	减产10%	植酸蛋白体体积减小;低价肌醇磷酸盐增加	^[28,29,30]
⁶⁰Co-γ	2-PGK	DR1331-2	45	增加 (等摩尔)	不变	否	--	^{[29, 30]}
γ-rays	--	lpa1	45	增加 (等摩尔)	不变	否	胚中植酸降幅高于糊粉层;钙, 锰降低、锌增加; 无低价磷酸盐	^{[16, 60]} ^{[61, 62]}
⁶⁰Co-γ+ NaN₃	--	Os-lpa-R6547-3	--	--	--	致死	--	^[52]
EMS	--	Dontokoi176	15	--	增加12%	否	垩白增幅率较低,铁,钼含量降低,其它矿质元素增加	^[63]
EMS	--	Koshihikari2623	45	--	不变	否	垩白增幅率较高,萌发前期活性较低	^[63]
γ-rays	--	Koshihikari4019	36	--	不变	否	垩白增幅率较高	^[63]
Callus culture	--	NC1857	17.1	增加 (等摩尔)	不变	小幅降低	铜、锰、钠、锌增加,钼降低41.9%	^[63]
EMS	--	Dontokoil 76	14.7	增加	增加 22%	小幅降低	增加了43.4%的铜,33.4%的钾,20.4%的镁,11.5%的钠, 39.1%的磷,23.6%的硫,10.8%的二氧化硅,44.2%的锌。而降低铁13.8%,钼17.5%	^[63]
γ-rays	--	Koshihikari49	5.9	增加	增加 9.4%	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari2623	44.7	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari3847 Koshihikari	47.9	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]
EMS	--	Koshihikari4019	35.9	增加 (等摩尔)	不变	小幅降低	铜,锰,钠和锌增加,钼降低	^[63]