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Rice Science ›› 2019, Vol. 26 ›› Issue (4): 229-238.DOI: 10.1016/j.rsci.2018.08.008

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  • 收稿日期:2018-03-31 接受日期:2018-08-22 出版日期:2019-07-28 发布日期:2019-04-04

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. [J]. Rice Science, 2019, 26(4): 229-238.

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链接本文: http://www.ricesci.org/CN/10.1016/j.rsci.2018.08.008

               http://www.ricesci.org/CN/Y2019/V26/I4/229

图/表 12

Supplemental Fig. 1 Schematic view of mutant screening for heat tolerance of a large Fast Neutron-treated population and the workflow of the present study, including screening and validation (Ruengphayak et al, 2015). In the first step, the heat tolerance as indicated by spikelet fertility was screened in 10,000 lines at high temperature. In the second step, lines were selected for >80% spikelet fertility (IRRI, 2013) under extreme heat. In the third part, the heat tolerance was validated by an evaluation of the selected JHN lines with rice cultivars.

Supplemental Fig. 1 Schematic view of mutant screening for heat tolerance of a large Fast Neutron-treated population and the workflow of the present study, including screening and validation (Ruengphayak et al, 2015). In the first step, the heat tolerance as indicated by spikelet fertility was screened in 10,000 lines at high temperature. In the second step, lines were selected for >80% spikelet fertility (IRRI, 2013) under extreme heat. In the third part, the heat tolerance was validated by an evaluation of the selected JHN lines with rice cultivars.

Table 1 Temperature (T), relative humidity (RH) and light intensity (LI) for the screening experiments.
Experiment Duration Greenhouse
Day time (6 h) Night time
T (oC) RH (%) LI [µmol/(L·m2·s)] T (oC) RH (%)
Preliminary screening April-December 2012 41.70 ± 1.90 56.52 ± 4.42 1 001.17 29.07 ± 2.28 82.18 ± 4.90
Repeat screening April-May 2013 42.13 ± 1.15 61.51 ± 4.47 929.26 29.41 ± 1.10 86.34 ± 4.85
Validation screening July-August 2013 41.44 ± 1.62 59.10 ± 3.44 997.34 28.35 ± 1.20 84.57 ± 5.86
Experiment Duration In field
Day time (6 h) Night time
T (oC) RH (%) LI [µmol/(L·m2·s)] T (oC) RH (%)
Preliminary screening April-December 2012 35.42 ± 2.90 47.61 ± 7.76 1 667.13 28.68 ± 2.31 80.42 ± 6.49
Repeat screening April-May 2013 35.24 ± 1.16 56.38 ± 5.81 1 468.89 27.97 ± 0.86 85.37 ± 7.01
Validation screening July-August 2013 33.25 ± 1.26 55.37 ± 6.13 1 538.70 27.08 ± 0.95 83.45 ± 8.57

Table 1 Temperature (T), relative humidity (RH) and light intensity (LI) for the screening experiments.

Experiment Duration Greenhouse
Day time (6 h) Night time
T (oC) RH (%) LI [µmol/(L·m2·s)] T (oC) RH (%)
Preliminary screening April-December 2012 41.70 ± 1.90 56.52 ± 4.42 1 001.17 29.07 ± 2.28 82.18 ± 4.90
Repeat screening April-May 2013 42.13 ± 1.15 61.51 ± 4.47 929.26 29.41 ± 1.10 86.34 ± 4.85
Validation screening July-August 2013 41.44 ± 1.62 59.10 ± 3.44 997.34 28.35 ± 1.20 84.57 ± 5.86
Experiment Duration In field
Day time (6 h) Night time
T (oC) RH (%) LI [µmol/(L·m2·s)] T (oC) RH (%)
Preliminary screening April-December 2012 35.42 ± 2.90 47.61 ± 7.76 1 667.13 28.68 ± 2.31 80.42 ± 6.49
Repeat screening April-May 2013 35.24 ± 1.16 56.38 ± 5.81 1 468.89 27.97 ± 0.86 85.37 ± 7.01
Validation screening July-August 2013 33.25 ± 1.26 55.37 ± 6.13 1 538.70 27.08 ± 0.95 83.45 ± 8.57
Table 2 Heat tolerance scoring system in rice (IRRI, 2013).
Score Spikelet fertility Description
1 > 80% Highly tolerant
3 61%-80% Tolerant
5 41%-60% Moderately tolerant
7 11%-40% Susceptible
9 < 11% Highly susceptible

Table 2 Heat tolerance scoring system in rice (IRRI, 2013).

Score Spikelet fertility Description
1 > 80% Highly tolerant
3 61%-80% Tolerant
5 41%-60% Moderately tolerant
7 11%-40% Susceptible
9 < 11% Highly susceptible
Supplemental Table 1 Origin, grain type and special traits of various rice cultivars used during the wet season of 2013.
Cultivar Origin Grain type Special trait
Dawk Pa-yawm Native rice Long Upland rice
Homcholasit RGDU* Long submergence tolerance
Jao Hom Nil RGDU Medium Wild Type
Pathum Thani 1 Rice Department. Long Good quality
Pin Kaset 1 RGDU Long High yield
Pin Kaset 2 RGDU Medium High yield
Pin Kaset 3 RGDU Long High yield
PSL2 Rice Department. Medium Commercial cultivar
RD15 Rice Department. Medium Adaptable to drought
RD33 Rice Department. Long Blast resistance
Suphan Buri 1 Rice Department. Medium Commercial cultivar
Thanyasirin RGDU Medium Waxy rice
A5029 RGDU Long Very long grain
Sinlek RGDU Long Control for high temperature sensitivity
* RGDU; Rice gene discovery unit

Supplemental Table 1 Origin, grain type and special traits of various rice cultivars used during the wet season of 2013.

Cultivar Origin Grain type Special trait
Dawk Pa-yawm Native rice Long Upland rice
Homcholasit RGDU* Long submergence tolerance
Jao Hom Nil RGDU Medium Wild Type
Pathum Thani 1 Rice Department. Long Good quality
Pin Kaset 1 RGDU Long High yield
Pin Kaset 2 RGDU Medium High yield
Pin Kaset 3 RGDU Long High yield
PSL2 Rice Department. Medium Commercial cultivar
RD15 Rice Department. Medium Adaptable to drought
RD33 Rice Department. Long Blast resistance
Suphan Buri 1 Rice Department. Medium Commercial cultivar
Thanyasirin RGDU Medium Waxy rice
A5029 RGDU Long Very long grain
Sinlek RGDU Long Control for high temperature sensitivity
* RGDU; Rice gene discovery unit
Fig. 1. Sensitivity of spikelet fertility to heat stress at 40 ºC to 45 ºC during the reproductive stages.T1, High temperature from panicle initiation to booting stages; T2, High temperature from panicle initiation to harvesting stages; T3, High temperature from booting to harvesting stages; T4, High temperature from flowering to harvesting stages. For the control, the plants were grown outside of the heat chamber under normal conditions. Different lowercase letters indicate significant difference at the 0.05 level. Bars are SE (n =15).

Fig. 1. Sensitivity of spikelet fertility to heat stress at 40 ºC to 45 ºC during the reproductive stages.T1, High temperature from panicle initiation to booting stages; T2, High temperature from panicle initiation to harvesting stages; T3, High temperature from booting to harvesting stages; T4, High temperature from flowering to harvesting stages. For the control, the plants were grown outside of the heat chamber under normal conditions. Different lowercase letters indicate significant difference at the 0.05 level. Bars are SE (n =15).

Fig. 2. Frequency distribution of 10 000 mutant plants and lines selected for spikelet fertility at 40 ºC to 45 ºC daytime temperature during the reproductive stage until harvesting.

Fig. 2. Frequency distribution of 10 000 mutant plants and lines selected for spikelet fertility at 40 ºC to 45 ºC daytime temperature during the reproductive stage until harvesting.

Fig. 3. Selection of rice mutants for heat tolerance based on spikelet fertility. A, Frequency of spikelet fertility in the greenhouse at high temperature (40 ºC to 45 ºC). B, Frequency of spikelet fertility in the field.

Fig. 3. Selection of rice mutants for heat tolerance based on spikelet fertility. A, Frequency of spikelet fertility in the greenhouse at high temperature (40 ºC to 45 ºC). B, Frequency of spikelet fertility in the field.

Table 3 Effects of heat stress on panicle traits in the candidate heat tolerant lines.
Genotype Spikelet fertility (%) Panicle length (cm) Filled grains per panicle Total florets per panicle
Field Greenhouse Field Greenhouse Field Greenhouse Field Greenhouse
M9962 84.3 aA 82.2 abA 20.4 deA 19.9 eA 105.4 aA 75.1 cdB 119.7 aA 104.4 bcB
M3181 79.5 abcdA 75.1 cdeA 22.3 cdA 20.6 deA 73.1 cdeA 53.6 hiB 92.4 cdefA 83.7 fgA
M7988 75.4 deA 74.3 deA 20.5 deB 23.5 bcA 81.7 bcA 53.1 hiB 109.8 abA 83.5 fgB
M8269 76.1 bcdA 73.7 deA 20.3 deA 19.6 eA 54.3 ghjA 50.9 ijA 84.6 efggA 85.7 defgA
M8281 83.0 abA 72.2 deB 20.6 deA 19.6 eA 64.3 efgA 54.4 ghiA 100.4 bcdA 85.7 defgA
M8372 76.6 bcdA 69.8 eB 20.2 deA 19.8 eA 62.7 fghA 40.7 jB 94.9 bcdefA 77.1 gB
M7766 69.6 eA 16.2 hB 24.7 abA 23.3 bcA 73.1 cdeA 9.2 kB 107.9 abA 101.8 bcA
JHN (wild type) 82.3 abcA 39.3 gB 24.9 abA 21.9 cdeA 85.6 bA 40.7 jB 103.0 bcA 99.2 bcdeA
Sinlek (control) 62.1 fA 10.4 hB 25.9 aA 24.1 abcA 70.1 defA 17.8 kB 109.5 abA 108.6 abA
Genotype (G) 2.50* 0.44* 4.37* 4.68*
Treatment (T) 4.94* 1.67* 7.26* 10.54*
G × T 6.98* 2.37 ns 10.27* 14.91
Different lowercase letters followed the values in the column are significantly different at P < 0.05 by the Tukey’s least significant difference (LSD) test. Different uppercase letters in each row for the same trait indicate significant difference at P < 0.05 by the Tukey’s LSD test.

Table 3 Effects of heat stress on panicle traits in the candidate heat tolerant lines.

Genotype Spikelet fertility (%) Panicle length (cm) Filled grains per panicle Total florets per panicle
Field Greenhouse Field Greenhouse Field Greenhouse Field Greenhouse
M9962 84.3 aA 82.2 abA 20.4 deA 19.9 eA 105.4 aA 75.1 cdB 119.7 aA 104.4 bcB
M3181 79.5 abcdA 75.1 cdeA 22.3 cdA 20.6 deA 73.1 cdeA 53.6 hiB 92.4 cdefA 83.7 fgA
M7988 75.4 deA 74.3 deA 20.5 deB 23.5 bcA 81.7 bcA 53.1 hiB 109.8 abA 83.5 fgB
M8269 76.1 bcdA 73.7 deA 20.3 deA 19.6 eA 54.3 ghjA 50.9 ijA 84.6 efggA 85.7 defgA
M8281 83.0 abA 72.2 deB 20.6 deA 19.6 eA 64.3 efgA 54.4 ghiA 100.4 bcdA 85.7 defgA
M8372 76.6 bcdA 69.8 eB 20.2 deA 19.8 eA 62.7 fghA 40.7 jB 94.9 bcdefA 77.1 gB
M7766 69.6 eA 16.2 hB 24.7 abA 23.3 bcA 73.1 cdeA 9.2 kB 107.9 abA 101.8 bcA
JHN (wild type) 82.3 abcA 39.3 gB 24.9 abA 21.9 cdeA 85.6 bA 40.7 jB 103.0 bcA 99.2 bcdeA
Sinlek (control) 62.1 fA 10.4 hB 25.9 aA 24.1 abcA 70.1 defA 17.8 kB 109.5 abA 108.6 abA
Genotype (G) 2.50* 0.44* 4.37* 4.68*
Treatment (T) 4.94* 1.67* 7.26* 10.54*
G × T 6.98* 2.37 ns 10.27* 14.91
Different lowercase letters followed the values in the column are significantly different at P < 0.05 by the Tukey’s least significant difference (LSD) test. Different uppercase letters in each row for the same trait indicate significant difference at P < 0.05 by the Tukey’s LSD test.
Fig. 4. Effects of high temperature stress during the booting to harvesting stages on the spikelet fertility (A) and grain yield per plant (B). Values (Mean ± SE, n = 15) with the same lowercase letters are not significantly different at the 0.05 level.

Fig. 4. Effects of high temperature stress during the booting to harvesting stages on the spikelet fertility (A) and grain yield per plant (B). Values (Mean ± SE, n = 15) with the same lowercase letters are not significantly different at the 0.05 level.

Table 4 Effects of heat stress at the booting stages on the 100-grain weight, filled grains per panicle and panicle weight in different rice cultivars in wet season 2013.
Genotype 100-grain weight (g) DF (%) No. of filled grains per panicle DF (%) Panicle weight (g) DF (%)
In field Greenhouse In field Greenhouse In field Greenhouse
M9962 2.10 uA 1.93 vB 8 101.0 defgA 103.0 defgA - 3.45 fghiA 2.82 jklmB 18.3
M3181 2.49 noA 2.31 qrB 7 68.3 jklmnA 50.9 mnoA 25 2.42 klmnopA 1.65 rsB 31.8
M7988 2.27 rstA 2.09 uB 8 77.3 hijklA 79.9 ghijklA - 2.85 ijklA 2.43 klmnoA 14.7
Thanyasirin 2.57 mnA 2.22 rstB 14 143.2 abA 104.1 defB 27 5.76 aA 3.82 efgB 33.7
M8269 2.44 opA 2.31 qrB 5 74.1 ijklmA 58.2 lmnoA 37 2.75 jklmA 2.34 lmnopqA 14.9
Suphan Buri 1 2.67 jkA 2.43 opB 9 115.0 cdeA 66.2 klmnoB 42 4.80 bcA 2.88 ijklB 40.0
PSL2 3.17 cA 2.30 qrsB 27 92.4 efghiA 60.8 lmoB 33 4.40 cdeA 2.64 jklmnB 40.0
Pathum Thani 1 2.85 ghA 2.46 opB 14 91.0 efghiA 53.1 mnoB 41 3.62 fghA 2.11 nopqrsB 41.7
Jao Hom Nil 2.52 mnoA 2.21 stB 12 85.3 fghijkA 46.6 nopB 45 2.99 hijkA 2.08 nopqrsB 30.4
Pin Kaset 2 2.91 fgA 2.67 jkB 8 134.0 abcA 44.3 opqB 67 5.03 bA 2.54 klmnB 49.5
Pin Kaset 1 3.04 deA 2.86 ghB 6 114.1 cdeA 26.9 pqrB 76 4.50 bcdA 2.06 nopqrsB 54.2
A5029 3.45 aA 2.98 efB 14 87.2 fghikA 20.2 rB 77 4.83 bcA 1.83 pqrsB 62.1
Dawk Payawm 2.77 hiA 2.37 pqB 14 154.3 aA 24.8 pqrB 83 4.90 bcA 2.21 mnopqrB 54.8
Pin Kaset 3 3.12 cdA 2.78 hiB 11 120.0 bcdA 15.4 rB 87 4.71 bcA 1.74 qrsB 63.1
Homcholasit 3.28 bA 2.61 klmB 20 99.0 defghA 22.3 qrB 77 3.70 efgA 1.89 opqrsB 48.9
Sinlek 2.98 efA 2.66 jklB 11 69.6 ijklmnA 17.8 rB 74 3.19 ghijA 1.79 pqrsB 43.9
RD15 2.64 klA 2.48 noB 6 122.0 bcdA 15.2 rB 88 3.90 defA 1.53 sB 60.8
RD33 2.74 ijA 2.18 tuB 20 73.4 ijklmA 7.4 rB 89 2.43 klmnoA 1.59 sB 34.5
Genotype (G) 0.07* 19.9* 2.04*
Treatment (T) 0.02* 5.6* 0.14*
G × T 0.09* 23.9* 0.62*
DF, Percentage decrease from the field.
Different lowercase letters followed the values in the column are significantly different at P < 0.05 by the Tukey’s least significant difference (LSD) test. Different uppercase letters in the each row for the same trait indicate significant difference at P < 0.05 by the Tukey’s LSD test.

Table 4 Effects of heat stress at the booting stages on the 100-grain weight, filled grains per panicle and panicle weight in different rice cultivars in wet season 2013.

Genotype 100-grain weight (g) DF (%) No. of filled grains per panicle DF (%) Panicle weight (g) DF (%)
In field Greenhouse In field Greenhouse In field Greenhouse
M9962 2.10 uA 1.93 vB 8 101.0 defgA 103.0 defgA - 3.45 fghiA 2.82 jklmB 18.3
M3181 2.49 noA 2.31 qrB 7 68.3 jklmnA 50.9 mnoA 25 2.42 klmnopA 1.65 rsB 31.8
M7988 2.27 rstA 2.09 uB 8 77.3 hijklA 79.9 ghijklA - 2.85 ijklA 2.43 klmnoA 14.7
Thanyasirin 2.57 mnA 2.22 rstB 14 143.2 abA 104.1 defB 27 5.76 aA 3.82 efgB 33.7
M8269 2.44 opA 2.31 qrB 5 74.1 ijklmA 58.2 lmnoA 37 2.75 jklmA 2.34 lmnopqA 14.9
Suphan Buri 1 2.67 jkA 2.43 opB 9 115.0 cdeA 66.2 klmnoB 42 4.80 bcA 2.88 ijklB 40.0
PSL2 3.17 cA 2.30 qrsB 27 92.4 efghiA 60.8 lmoB 33 4.40 cdeA 2.64 jklmnB 40.0
Pathum Thani 1 2.85 ghA 2.46 opB 14 91.0 efghiA 53.1 mnoB 41 3.62 fghA 2.11 nopqrsB 41.7
Jao Hom Nil 2.52 mnoA 2.21 stB 12 85.3 fghijkA 46.6 nopB 45 2.99 hijkA 2.08 nopqrsB 30.4
Pin Kaset 2 2.91 fgA 2.67 jkB 8 134.0 abcA 44.3 opqB 67 5.03 bA 2.54 klmnB 49.5
Pin Kaset 1 3.04 deA 2.86 ghB 6 114.1 cdeA 26.9 pqrB 76 4.50 bcdA 2.06 nopqrsB 54.2
A5029 3.45 aA 2.98 efB 14 87.2 fghikA 20.2 rB 77 4.83 bcA 1.83 pqrsB 62.1
Dawk Payawm 2.77 hiA 2.37 pqB 14 154.3 aA 24.8 pqrB 83 4.90 bcA 2.21 mnopqrB 54.8
Pin Kaset 3 3.12 cdA 2.78 hiB 11 120.0 bcdA 15.4 rB 87 4.71 bcA 1.74 qrsB 63.1
Homcholasit 3.28 bA 2.61 klmB 20 99.0 defghA 22.3 qrB 77 3.70 efgA 1.89 opqrsB 48.9
Sinlek 2.98 efA 2.66 jklB 11 69.6 ijklmnA 17.8 rB 74 3.19 ghijA 1.79 pqrsB 43.9
RD15 2.64 klA 2.48 noB 6 122.0 bcdA 15.2 rB 88 3.90 defA 1.53 sB 60.8
RD33 2.74 ijA 2.18 tuB 20 73.4 ijklmA 7.4 rB 89 2.43 klmnoA 1.59 sB 34.5
Genotype (G) 0.07* 19.9* 2.04*
Treatment (T) 0.02* 5.6* 0.14*
G × T 0.09* 23.9* 0.62*
DF, Percentage decrease from the field.
Different lowercase letters followed the values in the column are significantly different at P < 0.05 by the Tukey’s least significant difference (LSD) test. Different uppercase letters in the each row for the same trait indicate significant difference at P < 0.05 by the Tukey’s LSD test.
Fig. 5. Pollen viability (A) and anther dehiscence (B) under field conditions and at high temperature for the six rice genotypes. *, Significant at the 0.05 level.

Fig. 5. Pollen viability (A) and anther dehiscence (B) under field conditions and at high temperature for the six rice genotypes. *, Significant at the 0.05 level.

Fig. 6. Images of six cultivars showing the pollen viability (A1-A6), anther dehiscence (apical and basal pore) (B1-B6) and pollen number on the stigma (C1-C6) under high temperature conditions.

Fig. 6. Images of six cultivars showing the pollen viability (A1-A6), anther dehiscence (apical and basal pore) (B1-B6) and pollen number on the stigma (C1-C6) under high temperature conditions.

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