水稻种子胚形态与胚乳组成对稻米留胚特性的影响

doi:10.16819/j.1001-7216.2023.220809

摘要/Abstract

摘要：

【目的】稻米留胚能够提高稻米营养品质，但稻米留胚特性与胚性状、胚乳成分的关系等相关生物学机制并不明确。因此，有必要深入研究不同类型稻米品种留胚率差异及其影响因素。【方法】选取粳稻中软米型和普通型两类典型留胚材料及非留胚材料为试材，首先切片观察两类品种籽粒胚发育特点，继而对比表观直链淀粉含量(apparent amylose content, AAC)和离层果胶含量，最后以外源细胞分裂素(cytokinin, CTK)调控，比较胚性状、胚乳组成及留胚率差异。【结果】与非留胚材料相比，软米型和普通型留胚材料具有胚内幅和外幅较大，胚发育速度快，胚体积较大的特点；籽粒充实度好，粒形更宽、厚；同时离层果胶含量也较高。外施CTK后，软米型留胚材料中后期胚发育速度加快，普通型材料胚发育减缓；两类材料的AAC、果胶含量、胚比重及留胚率均提高。【结论】除胚性状对两类品种留胚率均有明显影响外，胚乳组成主要影响软米型材料留胚特性，而粒形是影响普通型材料留胚特性的重要因素。CTK则通过影响籽粒不同性状对不同类型材料留胚率进行调控。这为后续高留胚水稻品种的培育和栽培调控提供了理论依据和指导方向。

关键词: 留胚率, 胚发育, 胚乳成分, 粒形, 细胞分裂素

Abstract:

【Objective】The embryo of embryo-remaining rice has high nutritional value and ensures the taste of rice. However, related biological mechanisms, such as the relationship among embryo-remaining characteristics, embryo traits, and endosperm components remains unclear. Therefore, it is necessary to further study the difference of embryo-remaining rate among different types of rice varieties and its influencing factors.【Method】Therefore, typical embryo-retaining and non-embryo-remaining varieties were selected from soft japonica and common rice as experimental materials to investigate rice embryo-remaining characteristics. Here, the embryo development characteristics of the two types of varieties were observed by slicing; then, apparent amylose content (AAC) and abscission pectin contents were compared. Finally, the differences in embryo-remaining rate, embryo traits and endosperm composition were studied under external cytokinin (CTK) application.【Result】Compared with non-embryo-remaining varieties, embryo-remaining soft japonica varieties and common rice had larger inner and outer embryo width, faster embryo development, large embryo volume, better grain plumpness, wider and thicker grain shape and higher abscission pectin content. After the application of CTK, the development speed of the soft rice varieties was accelerated in the middle and late stages of development, while the embryo development of the common varieties slowed down. The AAC, pectin content, embryo weight percentage and embryo-remaining rate of the two types were increased.【Conclusion】In general, apart from embryo traits, endosperm composition mainly affected the embryo-remaining characteristics of soft rice varieties, whereas grain shape was an important factor influencing the embryo-remaining characteristics of common varieties. CTK regulates the embryo-remaining rate of different types of rice varieties by influencing different traits of rice seeds. These results will lay a new theoretical basis for the breeding and cultivation regulation of high embryo-remaining rice varieties.

Key words: embryo-remaining rate, embryo development, endosperm composition, grain shape, cytokinin

马兆惠, 石一涵, 程海涛, 宋文雯, 路连吉, 刘仁广, 吕文彦. 水稻种子胚形态与胚乳组成对稻米留胚特性的影响[J]. 中国水稻科学, 2023, 37(3): 265-275.

MA Zhaohui, SHI Yihan, CHENG Haitao, SONG Wenwen, LU Lianji, LIU Renguang, LÜ Wenyan. Effects of Embryo Morphology and Endosperm Composition on Embryo-remaining Characteristics in Rice[J]. Chinese Journal OF Rice Science, 2023, 37(3): 265-275.

图/表 8

参考文献 41

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时期 Period	处理 Treatment	材料 Material	内幅 Inner width/μm	胚长 Embryo length/μm	外幅 Outer width/μm	胚面积 Area/μm²
花后9 d 9 days after flowering (DAF)	CK	YJRY1	56.34±1.40 d	424.23±2.65 c	82.47±0.02 c	49105.01±2104.97 c
		1419-8	36.11±0.19 g	258.21±1.47 f	68.07±0.13 d	19954.39±345.44 d
		2045-5	40.99±0.58 f	249.13±2.11 g	53.95±2.43 e	18986.12±539.21 d
		19-3	89.64±1.50 a	513.19±4.52 b	160.25±1.42 a	83755.56±2479.67 b
	H	YJRY1	84.22±1.45 b	660.09±1.46 a	142.81±7.84 b	102768.61±5854.01 a
		1419-8	33.84±0.99 g	205.09±0.47 h	67.04±2.13 d	12899.56±408.27 e
		2045-5	52.62±1.59 e	267.64±4.06 e	73.02±1.39 d	23286.60±1340.44 d
		19-3	60.01±2.51 c	342.42±2.08 d	143.12±1.25 b	47501.99±800.05 c
花后12 d 12 days after flowering (DAF)	CK	YJRY1	155.96±2.69 a	781.79±4.54 a	243.50±3.97 a	204859.15±4693.79 a
		1419-8	82.84±1.19 f	407.71±6.19 f	91.96±0.18 f	55642.64±1324.35 e
		2045-5	110.06±7.88 d	683.42±17.16 b	131.83±5.35 d	113832.48±3559.10 d
		19-3	91.31±3.51 e	605.43±15.06 e	169.63±5.38 bc	113111.26±1822.26 d
	H	YJRY1	129.04±2.31 c	663.43±3.88 c	242.38±14.35 a	175852.57±1232.70 b
		1419-8	136.10±2.74 b	635.14±6.00 d	160.63±4.84 c	129999.88±4063.04 c
		2045-5	78.45±0.10 f	418.05±3.84 f	110.74±3.95 e	52124.91±114.01 e
		19-3	110.70±2.44 d	616.158±5.62 de	181.32±4.63 b	117309.32±532.26 d
花后30 d 30 days after flowering (DAF)	CK	YJRY1	164.85±23.44 cd	932.45±74.40 b	258.85±25.49 ab	266226.18±36850.46 bc
		1419-8	180.27±19.28 bc	966.58±17.01 ab	229.17±18.67 cd	286081.75±33558.01 ab
		2045-5	163.53±17.54 cd	927.89±62.79 b	221.33±30.02 cd	250100.33±30413.79 c
		19-3	191.12±2.44 ab	947.01±5.62 b	205.77±4.63 cd	255104.21±532.26 c
	H	YJRY1	190.00±25.78 ab	1002.21±46.36 a	263.29±8.15 a	299395.46±33433.78 a
		1419-8	208.64±27.04 a	1011.39±26.63 a	233.47±34.12 bc	298319.09±25404.60 a
		2045-5	153.52±17.51 b	923.88±61.48 b	216.31±17.43 cd	242886.23±24922.29 c
		19-3	165.71±19.80 cd	946.25±15.97 b	202.24±21.67 d	247532.44±30002.13 c
时期 Period	处理 Treatment	材料 Material	内幅增速 Growing speed of inner width /(μm⁻²·d⁻¹)	胚长增速 Growing speed of embryo length /(μm⁻²·d⁻¹)	外幅增速 Growing speed of outer width /(μm⁻²·d⁻¹)	胚面积增速 Growing speed of embryo area/(μm⁻²·d⁻¹)
9DAF-12DAF	CK	YJRY1	33.21±0.48 a	119.19±0.79 b	53.68±1.32 a	51918.05±862.99 a
		1419-8	23.02±2.44 b	144.76±5.07 a	25.96±0.97 c	31615.45±1007.57 c
		2045-5	15.58±0.33 c	49.84±1.57 d	7.96±0.02 e	11896.08±326.31 e
		19-3	0.56±0.33 e	30.75±1.16 e	3.13±2.27 f	9785.23±734.58 f
	H	YJRY1	14.94±0.29 c	1.11±0.81 f	33.19±1.23 b	24361.32±734.58 d
		1419-8	8.61±0.50 d	50.14±0.07 d	12.57±0.86 d	9612.77±408.81 f
		2045-5	34.08±0.26 a	143.35±2.01 a	31.20±1.08 b	39033.44±918.77 b
		19-3	16.89±0.02 c	91.25±1.18 c	12.73±0.97 d	23269.11±89.27 d
9DAF-12DAF	CK	YJRY1	0.49±0.06 d	8.36±0.81 f	0.85±0.16 f	3409.28±177.23 e
		1419-8	2.97±0.20 c	13.56±0.46 e	4.97±0.96 bc	7570.44±121.62 d
		2045-5	5.41±0.18 ab	31.02±0.52 a	7.62±0.27 a	12802.17±799.14 a
		19-3	5.52±0.34 a	18.99±0.62 cd	2.01±0.47 d	7888.50±7.09 d
	H	YJRY1	3.39±0.83 c	18.82±0.87 cd	1.16±0.28 de	6863.49±258.68 d
		1419-8	4.17±0.19 abc	28.10±2.23 b	5.86±0.61 b	10597.85±434.95 b
		2045-5	4.03±1.37 bc	20.90±0.80 c	4.05±0.22 c	9351.07±130.49 c
		19-3	3.06±0.71 c	18.34±0.25 d	1.16±0.12 de	7234.62±1196.18 d

时期 Period	处理 Treatment	材料 Material	内幅 Inner width/μm	胚长 Embryo length/μm	外幅 Outer width/μm	胚面积 Area/μm²
花后9 d 9 days after flowering (DAF)	CK	YJRY1	56.34±1.40 d	424.23±2.65 c	82.47±0.02 c	49105.01±2104.97 c
		1419-8	36.11±0.19 g	258.21±1.47 f	68.07±0.13 d	19954.39±345.44 d
		2045-5	40.99±0.58 f	249.13±2.11 g	53.95±2.43 e	18986.12±539.21 d
		19-3	89.64±1.50 a	513.19±4.52 b	160.25±1.42 a	83755.56±2479.67 b
	H	YJRY1	84.22±1.45 b	660.09±1.46 a	142.81±7.84 b	102768.61±5854.01 a
		1419-8	33.84±0.99 g	205.09±0.47 h	67.04±2.13 d	12899.56±408.27 e
		2045-5	52.62±1.59 e	267.64±4.06 e	73.02±1.39 d	23286.60±1340.44 d
		19-3	60.01±2.51 c	342.42±2.08 d	143.12±1.25 b	47501.99±800.05 c
花后12 d 12 days after flowering (DAF)	CK	YJRY1	155.96±2.69 a	781.79±4.54 a	243.50±3.97 a	204859.15±4693.79 a
		1419-8	82.84±1.19 f	407.71±6.19 f	91.96±0.18 f	55642.64±1324.35 e
		2045-5	110.06±7.88 d	683.42±17.16 b	131.83±5.35 d	113832.48±3559.10 d
		19-3	91.31±3.51 e	605.43±15.06 e	169.63±5.38 bc	113111.26±1822.26 d
	H	YJRY1	129.04±2.31 c	663.43±3.88 c	242.38±14.35 a	175852.57±1232.70 b
		1419-8	136.10±2.74 b	635.14±6.00 d	160.63±4.84 c	129999.88±4063.04 c
		2045-5	78.45±0.10 f	418.05±3.84 f	110.74±3.95 e	52124.91±114.01 e
		19-3	110.70±2.44 d	616.158±5.62 de	181.32±4.63 b	117309.32±532.26 d
花后30 d 30 days after flowering (DAF)	CK	YJRY1	164.85±23.44 cd	932.45±74.40 b	258.85±25.49 ab	266226.18±36850.46 bc
		1419-8	180.27±19.28 bc	966.58±17.01 ab	229.17±18.67 cd	286081.75±33558.01 ab
		2045-5	163.53±17.54 cd	927.89±62.79 b	221.33±30.02 cd	250100.33±30413.79 c
		19-3	191.12±2.44 ab	947.01±5.62 b	205.77±4.63 cd	255104.21±532.26 c
	H	YJRY1	190.00±25.78 ab	1002.21±46.36 a	263.29±8.15 a	299395.46±33433.78 a
		1419-8	208.64±27.04 a	1011.39±26.63 a	233.47±34.12 bc	298319.09±25404.60 a
		2045-5	153.52±17.51 b	923.88±61.48 b	216.31±17.43 cd	242886.23±24922.29 c
		19-3	165.71±19.80 cd	946.25±15.97 b	202.24±21.67 d	247532.44±30002.13 c
时期 Period	处理 Treatment	材料 Material	内幅增速 Growing speed of inner width /(μm⁻²·d⁻¹)	胚长增速 Growing speed of embryo length /(μm⁻²·d⁻¹)	外幅增速 Growing speed of outer width /(μm⁻²·d⁻¹)	胚面积增速 Growing speed of embryo area/(μm⁻²·d⁻¹)
9DAF-12DAF	CK	YJRY1	33.21±0.48 a	119.19±0.79 b	53.68±1.32 a	51918.05±862.99 a
		1419-8	23.02±2.44 b	144.76±5.07 a	25.96±0.97 c	31615.45±1007.57 c
		2045-5	15.58±0.33 c	49.84±1.57 d	7.96±0.02 e	11896.08±326.31 e
		19-3	0.56±0.33 e	30.75±1.16 e	3.13±2.27 f	9785.23±734.58 f
	H	YJRY1	14.94±0.29 c	1.11±0.81 f	33.19±1.23 b	24361.32±734.58 d
		1419-8	8.61±0.50 d	50.14±0.07 d	12.57±0.86 d	9612.77±408.81 f
		2045-5	34.08±0.26 a	143.35±2.01 a	31.20±1.08 b	39033.44±918.77 b
		19-3	16.89±0.02 c	91.25±1.18 c	12.73±0.97 d	23269.11±89.27 d
9DAF-12DAF	CK	YJRY1	0.49±0.06 d	8.36±0.81 f	0.85±0.16 f	3409.28±177.23 e
		1419-8	2.97±0.20 c	13.56±0.46 e	4.97±0.96 bc	7570.44±121.62 d
		2045-5	5.41±0.18 ab	31.02±0.52 a	7.62±0.27 a	12802.17±799.14 a
		19-3	5.52±0.34 a	18.99±0.62 cd	2.01±0.47 d	7888.50±7.09 d
	H	YJRY1	3.39±0.83 c	18.82±0.87 cd	1.16±0.28 de	6863.49±258.68 d
		1419-8	4.17±0.19 abc	28.10±2.23 b	5.86±0.61 b	10597.85±434.95 b
		2045-5	4.03±1.37 bc	20.90±0.80 c	4.05±0.22 c	9351.07±130.49 c
		19-3	3.06±0.71 c	18.34±0.25 d	1.16±0.12 de	7234.62±1196.18 d

材料 Material	胚比重 Embryo weight percentage/%		留胚率 Embryo retention rate/%		总果胶含量 Total pectin content/%		直链淀粉含量 Apparent amylose content/%
软米型 Soft rice
YJRY 1 1419-8	1.94±0.22 a		85±0.82 a		0.61±0.05 a		4.06±0.08 a
YJRY 1 1419-8	0.42±0.12 b		23±4.50 b		0.50±0.02 b		4.57±0.21 a
普通型Common
2045-5 19-3	1.90±0.11 a		82±3.27 a		0.52±0.05 a		14.24±1.46 a
2045-5 19-3	0.33±0.20 b		26±3.88 b		0.45±0.06 a		10.00±1.26 a
材料 Material	千粒重 Thousand-grain weight/g	长宽比 Length-width ratio		粒长 Grain length/mm		粒宽 Grain width/mm		粒厚 Grain thickness/mm
软米型Soft rice
YJRY 1 1419-8	22.07±0.13 a	1.82±0.01 a		4.99±0.29 a		2.76±0.22 a		2.19±0.07 a
YJRY 1 1419-8	21.46±0.21 b	1.86±0.01 a		5.07±0.31 a		2.73±0.23 a		2.07±0.06 b
普通型Common
2045-5 19-3	22.38±0.25 a	1.84±0.01 b		5.03±0.33 b		2.76±0.22 a		2.11±0.07 a
2045-5 19-3	19.33±0.42 b	1.98±0.04 a		5.16±0.47 a		2.63±0.31 b		1.87±0.05 b

材料 Material	胚比重 Embryo weight percentage/%		留胚率 Embryo retention rate/%		总果胶含量 Total pectin content/%		直链淀粉含量 Apparent amylose content/%
软米型 Soft rice
YJRY 1 1419-8	1.94±0.22 a		85±0.82 a		0.61±0.05 a		4.06±0.08 a
YJRY 1 1419-8	0.42±0.12 b		23±4.50 b		0.50±0.02 b		4.57±0.21 a
普通型Common
2045-5 19-3	1.90±0.11 a		82±3.27 a		0.52±0.05 a		14.24±1.46 a
2045-5 19-3	0.33±0.20 b		26±3.88 b		0.45±0.06 a		10.00±1.26 a
材料 Material	千粒重 Thousand-grain weight/g	长宽比 Length-width ratio		粒长 Grain length/mm		粒宽 Grain width/mm		粒厚 Grain thickness/mm
软米型Soft rice
YJRY 1 1419-8	22.07±0.13 a	1.82±0.01 a		4.99±0.29 a		2.76±0.22 a		2.19±0.07 a
YJRY 1 1419-8	21.46±0.21 b	1.86±0.01 a		5.07±0.31 a		2.73±0.23 a		2.07±0.06 b
普通型Common
2045-5 19-3	22.38±0.25 a	1.84±0.01 b		5.03±0.33 b		2.76±0.22 a		2.11±0.07 a
2045-5 19-3	19.33±0.42 b	1.98±0.04 a		5.16±0.47 a		2.63±0.31 b		1.87±0.05 b

材料 Material	内幅 Inner width	胚长 Embryo length	外幅 Outer width	胚面积 Area	千粒重 Thousand-grain weight	长宽比 Length-width ratio
YJRY 1	0.996**	−0.077	0.988*	−0.991**	0.676	−0.606
1419-8	0.822	0.961*	−0.994**	−0.491	0.353	−0.866
2045-5	−0.608	−0.992**	−0.970*	0.994**	−0.461	0.990**
19-3	1.000**	−0.988*	−0.792	−0.500	0.848	−0.182
材料 Material	粒长 Grain length	粒宽 Grain width	粒厚 Grain thickness	胚比重 Embryo weight percentage	总果胶含量 Total pectin content	直链淀粉含量 Apparent amylose content
YJRY 1	−0.444	0.141	0.097	0.789	0.582	−0.650
1419-8	0.423	0.520	0.775	0.187	0.780	−0.235
2045-5	−0.099	−0.854	−0.657	0.756	−0.569	0.227
19-3	−0.648	−0.945	−0.011	0.844	−0.969*	−0.412