关键词:超级稻; 籽粒灌浆; 产量; 生理指标 Grain-filling Characteristics in Super Rice with Different Panicle Types WEI Ying-Juan, ZHAO Yang, ZOU Ying-Bin* Agronomy College of Hunan Agricultural University, Changsha 410128, China Fund:This study was supported by the China Agriculture Research System (CARS-01-34) and the Scientific Innovation Program for Graduate Students of Hunan Agricultural University (CX2014B296) AbstractThe objective of this study was to ascertain the characteristics of the grain-filling and physiological indexes after heading and the yield components of super rice with different panicle-grain types. An experiment with different sowing date was conducted using different growth duration varieties (Zhunliangyou 527, Liangyoupeijiu, Y-liangyou 1, Yuxiangyouzhan, and Huanghuazhan) under the field conditions to ensure the grain filling was under the similar temperature and solar radiation conditions. The grain yield and its components averaged in three sowing treatments displayed significant difference among tested varieties, of which the highest yield reached 9.47 t ha-1 for Liangyoupeijiu and the lowest was 8.39 t ha-1 for Yuxiangyouzhan. According to the hierarchical cluster analysis based on the grain number per panicle and 1000-grain weight, tested varieties were divided into the big grain type (Zhunliangyou 527), the medium grain type (Liangyoupeijiu and Y-liangyou 1), and the small grain type (Yuxiangyouzhan and Huanghuazhan). There were obvious differences intensity and peak duration of grain filling among the tested varieties. The big grain variety showed quick grain growth at the beginning, high intensity and short peak duration for grain filling during the middle-late periods, being one step filling in the upper, middle and lower branch grains; the medium grain variety showed quicker grain growth at the beginning, smaller intensity and longer peak duration for grain filling during the middle- late periods, with two filling peaks from 12 d to 18 d and from 24 d to 30 d after heading, respectively; the small grain variety showed slow grain growth at the beginning, smaller intensity and shorter peak duration for grain filling during the middle-late periods, which was one step filling in the upper or middle branch grains and two step filling in the lower branch grains. The average grain filling rate closely related to the panicle-grain type of varieties, was 0.68 mg d-1 for big grain variety, 0.48-0.51 mg d-1 for medium varieties, and 0.41-0.47 mg d-1 for small grain varieties. The intensity and duration of the grain-filling also associated with the changes of physiological indexes of flag leaf and roots after heading. At the beginning of the grain-filling, chlorophyll content, chlorophyll a/ b ratio, carotenoids content gradually declined, leaf soluble sugar content, MDA content gradually increased, POD and CAT activities and root activity successively underwent two rise and fall processes. In conclusion, different panicle-grain types super rice have significantly different characteristics of yield components and grain filling. The intensity and the duration of grain-filling depend upon the panicle-grain type of varieties and the branch and grain positions inserted on the panicle, with the same tendency of physiological indexes variation in flag leaf and roots.
2 结果与分析2.1 不同穗型粒型超级稻产量及穗粒结构类型2.1.1 产量及其构成的年份间、播期间及品种间差异 从表1可以看出, 不同穗粒型超级稻的干物质重、收获指数、产量及产量构成因子的品种间差异显著或极显著。除了干物质重和单位面积颖花数的差异不显著外, 不同播期对收获指数、产量及产量构成因子的影响显著或极显著。不同年份间的产量、颖花数、千粒重差异极显著, 但干物质重、收获指数及其他产量构成因子的差异不显著。品种× 年份的互作对产量的影响显著, 对各产量构成因子的影响不显著; 年份× 播期的互作对各产量构成因子的影响显著, 对产量的影响不显著; 品种× 播期的互作对产量、有效穗数、每穗粒数的影响显著, 对其他产量构成因子的影响不显著。 2.1.2 不同播种期条件下的产量表现及生育期变化 从表2可以看出, 不同穗粒型超级稻产量及生育期存在年份间、播期间、品种间差异。不同品种平均产量随播期推迟而增加, 2011年第一、二、三播期分别为9.01、9.27、9.31 t hm-2, 差异不显著; 2012年分别为8.42、8.55、9.04 t hm-2, 处理间差异显著。不同播期平均产量品种间差异显著, 其中以两优培九最高, 2011年、2012年分别为9.92 t hm-2和9.03 t hm-2; 以玉香油占最低, 两年分别为8.44 t hm-2和8.34 t hm-2。表3表明平均产量2011年为9.20 t hm-2, 2012年为8.67 t hm-2, 年度间差异显著。值得注意的是, 同一块稻田种植同一品种, 在不同年份、不同播期的产量差异主要受生长期间, 尤其是灌浆结实期间光照、温度等气候因素影响。表2还表明, 每推迟10 d播种, 播种至抽穗日数缩短1~2 d, 全生育期缩短2~3 d。品种间全生育期差异明显, 以两优培九最长, 为136~141 d, 其余依次为Y两优1号(131~137 d)、准两优527 (127~133 d)、黄华占(122~128 d)、玉香油占(123~125 d)。全生育期年度间稍有差异, 2011年平均为129.3 d, 2012年为131.1 d。 表1 Table 1 表1(Table 1)
表1 不同穗粒型超级稻产量表现、产量构成的年份、播期、品种间差异及其交互作用(F值) Table 1 Variance analysis of yield and its components of supper rice with different panicle-grain types (F-value)
方差来源 Source
df
干物质 Dry matter
收获指数 Harvest index
有效穗数 Effective panicle number
每穗颖花数 Spikelet number per panicle
每平方米 颖花数 Spikelet number per m2
结实率 Grain filling
千粒重 1000-grain weight
产量 Grain yield
年份Year (Y)
1
3.78
0.00
0.04
131.91* *
94.72* *
0.66
14.89* *
22.09* *
播期Sowing date (S)
2
0.99
4.25*
44.82* *
13.56* *
0.16
4.76* *
5.72* *
5.77* *
品种Variety (V)
4
3.23*
4.09* *
98.52* *
61.79* *
40.20* *
13.97* *
1203.23* *
10.66* *
Y× S
2
7.06*
2.20
10.29* *
6.11* *
6.39* *
7.23* *
29.52* *
1.41
Y× V
4
3.12*
0.88
1.48
2.23
1.75
0.71
0.17
3.24*
S× V
8
0.60
6.51* *
4.50* *
4.68* *
2.05
1.53
1.97
3.34* *
Y× S× V
8
2.73*
1.23
1.13
1.32
1.44
0.96
1.02
1.83
* Significant at the 0.05 probability level based on analysis of variance. * * Significant at the 0.01 probability level based on analysis of variance. * 表示差异达到0.05的显著水平, * * 表示差异达到0.01的显著水平。
表1 不同穗粒型超级稻产量表现、产量构成的年份、播期、品种间差异及其交互作用(F值) Table 1 Variance analysis of yield and its components of supper rice with different panicle-grain types (F-value)
表2 Table 2 表2(Table 2)
表2 不同播种期条件下不同穗粒型超级稻品种的产量表现 Table 2 Effect of sowing date on the yield performance of supper rice with different panicle-grain types
播种日期 Sowing date (month/day)
品种 Variety
2011年试验 Field experiment in 2011
2012年试验 Field experiment in 2012
收割产量 Harvest yield (t hm2)
播种-抽穗日数 Sowing- Heading (d)
播种-成熟日数 Sowing- Maturity (d)
收割产量 Harvest yield (t hm2)
播种-抽穗日数 Sowing- Heading (d)
播种-成熟日数 Sowing- Maturity (d)
5/03
ZLY527
9.01 cdef
94
129
8.09 cde
94
133
LYPJ
10.15 ab
99
140
8.90 ab
99
141
YLY-1
8.97 cdef
98
135
7.88 e
98
137
YXYZ
8.49 defg
86
124
8.43 bcde
91
127
HHZ
8.43 efg
87
125
8.78 abc
92
128
平均Average
9.01 A
92.8
130.6
8.42 B
94.8
133.2
5/13
ZLY527
9.73 abc
93
129
8.76 abc
93
132
LYPJ
9.40 abcd
96
137
9.01 ab
99
141
YLY-1
10.04 ab
96
133
8.62 abcde
96
134
YXYZ
7.96 g
86
123
7.91 de
89
124
HHZ
9.23 bcde
87
124
8.46 bcde
90
125
平均Average
9.27 A
91.6
129.2
8.55 B
93.4
131.2
5/23
ZLY527
8.21 fg
91
127
9.03 ab
91
130
LYPJ
10.22 a
96
136
9.18 ab
96
138
YLY-1
10.18 ab
94
132
9.35 a
95
131
YXYZ
8.87 cdefg
84
123
8.69 abcd
86
121
HHZ
9.08 cdef
86
122
8.96 ab
88
125
平均Average
9.31 A
90.2
128.0
9.04 A
91.2
129.0
ZLY527: Zhunliangyou 527; LYPJ: Liangyoupeijiu; YLY-1: Y-liangyou 1; YXYZ: Yuxiangyouzhan; HHZ: Huanghuazhan. Within a column, means followed by different small letters are significantly different at P< 0.05 between cultivars, and those by different capital letters are significantly different at P< 0.05 between sowing dates. ZLY527: 准两优527; LYPJ: 两优培九; YLY-1: Y两优1号; YXYZ: 玉香油占; HHZ: 黄华占。同一列数据后不同小写字母表示品种间差异显著(P< 0.05), 不同大写字母表示播期间差异显著(P< 0.05)。
表2 不同播种期条件下不同穗粒型超级稻品种的产量表现 Table 2 Effect of sowing date on the yield performance of supper rice with different panicle-grain types
表3 播种期对不同穗粒型超级稻产量构成因子的影响 Table 3 Effect of sowing date on yield components of supper rice with different panicle and grain types
播期 Sowing date (month/day)
品种 Variety
2011年试验 Field experiment in 2011
2012年试验 Field experiment in 2012
有效穗数 Panicle number per m2
每穗粒数 Spikelet number per panicle
结实率 Grain filling (%)
千粒重 1000-grain weight (mg)
有效穗数 Panicle number per m2
每穗粒数 Spikelet number per panicle
结实率 Grain filling (%)
千粒重 1000-grain weight (mg)
5/03
ZLY527
250.6 cd
144.4 gh
87.7 a
29.3 a
262.7 defg
106.0 h
75.2 de
28.9 b
LYPJ
214.4 e
241.8 a
77.6 cde
23.8 bc
241.0 i
183.7 a
73.7 de
23.5 d
YLY-1
238.6 de
197.2 cd
76.8 cde
24.4 b
257.5 efgh
156.6 bcde
69.1 e
23.7 d
YXYZ
234.7 de
217.4 bc
80.4 abcde
20.0 ef
252.1 fghi
176.0 ab
77.4 cde
19.1 g
HHZ
307.2 b
147.4 fgh
81.3 abcd
20.4 e
311.9 ab
139.4 defg
84.8 abc
19.7 fg
平均Average
249.1 C
189.6 A
80.8 A
23.6 A
265.0 B
152.3 A
76.0 B
23.0 C
5/13
ZLY527
263.1 c
140.6 h
86.6 ab
29.4 a
249.4 ghi
117.2 gh
89.8 a
29.6 ab
LYPJ
256.7 cd
211.3 bc
74.8 de
22.9 d
251.3 fghi
157.2 bcde
80.3 bcd
23.6 d
YLY-1
257.2 cd
196.5 cd
79.7 abcde
23.3 cd
245.6 hi
150.0 cdef
80.7 bcd
23.6 d
YXYZ
256.4 cd
220.5 ab
78.0 bcde
19.3 fg
237.7 i
163.9 abcd
76.5 de
19.9 fg
HHZ
303.3 b
183.8 de
87.8 a
19.7 ef
300.4 b
137.9 efg
90.7 a
20.5 ef
平均Average
267.3 B
190.5 A
81.4 A
22.9 B
256.9 C
145.2 A
83.6 A
23.4 B
5/23
ZLY527
263.6 c
133.6 h
79.5 abcde
29.0 a
272.3 cd
119.2 gh
84.9 abc
30.2 a
LYPJ
265.3 c
169.5 ef
72.2 e
23.6 bcd
265.4 def
140.9 cdefg
75.7 de
24.1 d
YLY-1
255.3 cd
166.8 efg
77.1 cde
23.8 bc
266.9 cde
165.3 abc
80.6 bcd
25.5 c
YXYZ
297.8 b
194.6 cd
77.8 bcde
18.8 g
280.0 c
163.5 abcde
86.7 ab
20.2 f
HHZ
348.1 a
151.9 fgh
85.2 abc
19.4 fg
325.8 a
126.3 fgh
89.7 a
21.1 e
平均Average
286.0 A
163.3 B
78.4 A
22.9 B
282.1 A
143.0 A
83.5 A
24.2 A
Within a column, means followed by different small letters are significantly different at P< 0.05 between cultivars, and those by different capital letters are significantly different at P< 0.05 between sowing dates. Abbreviations are the same as given in Table 2. 同一列数据后不同小写字母表示品种间差异显著(P< 0.05), 不同大写字母表示播期间差异显著(P< 0.05)。缩写同表2。
表3 播种期对不同穗粒型超级稻产量构成因子的影响 Table 3 Effect of sowing date on yield components of supper rice with different panicle and grain types
图1 不同穗粒型超级稻全穗的籽粒增长动态缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 1 Grain growth dynamics of the whole panicle of supper rice with different panicle-grain types
图2 不同穗粒型超级稻上部、中部、下部枝梗籽粒增长动态缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 2 Grain growth dynamics of upper, middle and lower branches grains of supper rice with different panicle-grain types
图3 不同穗粒型超级稻整穗籽粒灌浆速率变化缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 3 Grain filling rate of the whole panicle of supper rice varieties with different panicle-grain types
表4 Table 4 表4(Table 4)
表4 不同穗粒型超级稻品种的平均灌浆速率 Table 4 Average grain filling rate of supper rice varieties with different panicle-grain types
品种 Variety
2011年灌浆速率 Grain filling rate in 2011 (mg d-1)
2012年灌浆速率 Grain filling rate in 2012 (mg d-1)
上部枝梗 Upper branch
中部枝梗 Middle branch
下部枝梗 Lower branch
整穗 Panicle
上部枝梗 Upper branch
中部枝梗 Middle branch
下部枝梗 Lower branch
整穗 Panicle
平均灌浆速率 Average filling rate
LYPJ
0.54 b
0.46 b
0.45 b
0.48 bc
0.55 c
0.48 b
0.46 b
0.49 bc
YLY-1
0.58 b
0.48 b
0.45 b
0.50 b
0.59 b
0.49 b
0.47 b
0.51 b
ZLY527
0.71 a
0.66 a
0.68 a
0.68 a
0.74 a
0.66 a
0.65 a
0.68 a
YXYZ
0.46 c
0.41 c
0.40 c
0.42 d
0.47 d
0.39 c
0.37 c
0.41 d
HHZ
0.46 c
0.45 bc
0.43 bc
0.45 cd
0.48 d
0.47 b
0.45 b
0.47 c
最大灌浆速率 Maximum filling rate
LYPJ
1.08 b
0.80 c
0.86 bc
0.85 bc
1.26 a
0.81 b
0.97 bc
0.83 b
YLY-1
1.52 a
0.98 b
1.00 b
0.96 b
1.24 a
1.12 a
1.18 ab
1.04 ab
ZLY527
1.65 a
1.22 a
1.52 a
1.26 a
1.46 a
1.12 a
1.36 a
1.19 a
YXYZ
1.12 b
0.76 c
0.69 c
0.79 bc
1.22 a
0.87 ab
0.83 c
0.90 ab
HHZ
1.02 b
0.77 c
0.83 bc
0.74 c
1.23 a
0.80 b
0.95 bc
0.83 b
Within a column, means followed by different letters are significantly different (P< 0.05) between cultivars. Abbreviations are the same as those given in Table 2. 同一列同一指标数据后不同字母表示品种间差异显著(P< 0.05)。缩写同表2。
表4 不同穗粒型超级稻品种的平均灌浆速率 Table 4 Average grain filling rate of supper rice varieties with different panicle-grain types
图4 不同穗粒型超级稻上部、中部、下部枝梗籽粒灌浆速率缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 4 Grain filling rate of upper, middle and lower branches of supper rice with different panicle-grain types
图5 不同穗粒型超级稻抽穗后剑叶叶绿素含量、叶绿素a/b值及类胡萝卜素含量变化缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 5 Variation of carotenoid, chlorophyll content and chlorophyll a/b ratio of the flag leaf after heading in super rice with different panicle-grain types
图6 不同穗粒型超级稻抽穗后剑叶POD、CAT酶活性和可溶性糖含量及MDA含量变化(2012年)缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 6 Variation of POD, CAT activities, soluble sugar content and net photosynthetic rate of the flag leaf after heading in supper rice with different panicle-grain types in 2012
图7 不同穗粒型超级稻抽穗后根系活力变化缩写同表2。Abbreviations are the same as those given in Table 2.Fig. 7 Variation of root activity after heading in supper rice with different panicle-grain types
表5 Table 5 表5(Table 5)
表5 不同穗型超级稻抽穗后剑叶生理指标及根系活力与籽粒增重间的相关系数 Table 5 Correlation coefficient between the grain weight increasing and the flag leaf physiology index and root activity after heading in supper rice with different panicle and grain types
品种 Variety
POD活性 POD activity
CAT活性 CAT activity
可溶糖含量 Soluble sugar content
MDA含量 MDA content
叶绿素含量 Chlorophyll content
叶绿素a/b值 Chlorophyll a/b ratio
类胡萝卜素含量 Carotenoid content
根系α 萘胺氧化力 α -naphthyl amine oxidation of roots
ZLY527
-0.382
-0.681* *
0.461*
0.857* *
-0.822* *
-0.674* *
-0.562* *
-0.697* *
LYPJ
-0.686* *
-0.694* *
0.841* *
0.717* *
-0.845* *
-0.538*
-0.747* *
0.100
YLY-1
-0.536*
-0.676* *
0.654* *
0.741* *
-0.919* *
-0.659* *
-0.768* *
-0.361
YXYZ
0.264
-0.714* *
0.524*
0.883* *
-0.872* *
-0.770* *
-0.649* *
-0.737* *
HHZ
-0.440*
-0.642* *
0.629* *
0.721* *
-0.839* *
-0.307
-0.575* *
-0.328
* Significant at the 0.05 probability level based on analysis of variance. * * Significant at the 0.01 probability level based on analysis of variance. Abbreviations are the same as those given in Table 2. * 表示差异达到0.05的显著水平, * * 表示差异达到0.05的显著水平。缩写同表2。
表5 不同穗型超级稻抽穗后剑叶生理指标及根系活力与籽粒增重间的相关系数 Table 5 Correlation coefficient between the grain weight increasing and the flag leaf physiology index and root activity after heading in supper rice with different panicle and grain types
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表1 不同穗粒型超级稻产量表现、产量构成的年份、播期、品种间差异及其交互作用(F值) Table 1 Variance analysis of yield and its components of supper rice with different panicle-grain types (F-value)
