Scientia Agricultura Sinica ›› 2022, Vol. 55 ›› Issue (3): 451-466.doi: 10.3864/j.issn.0578-1752.2022.03.003

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Geographic Differentiation and Evolution of Photo-Thermal Comprehensive Responses of Growth-Periods in Global Soybeans

JIANG FenFen(),SUN Lei,LIU FangDong,WANG WuBin,XING GuangNan,ZHANG JiaoPing,ZHANG FengKai,LI Ning,LI Yan,HE JianBo(),GAI JunYi()   

  1. Soybean Research Institute, Nanjing Agricultural University/MARA National Center for Soybean Improvement/MARA Key Laboratory of Biology and Genetic Improvement of Soybean (General)/State Key Laboratory for Crop Genetics and Germplasm Enhancement/Jiangsu Collaborative Innovation Center for Modern Crop Production, Nanjing 210095
  • Received:2021-08-13 Accepted:2021-10-11 Online:2022-02-01 Published:2022-02-11
  • Contact: JianBo HE,JunYi GAI E-mail:2019201036@njau.edu.cn;hjbxyz@gmail.com;sri@njau.edu.cn

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Abstract:

【Objective】 As a short-day (SD) and thermophilic plant, soybean is sensitive to photo-thermal(day length, temperature) conditions. The sensitivity of soybeans to photo-thermal response is an important domestication and adaptability trait of soybean. Under natural conditions, geographical location and/or sowing season are two important ecological factors that determine the differentiation of wild and cultivated soybeans, and they work together to regulate the growth and development of soybeans through environmental factors such as day length and temperature. Therefore, the study of the geographical and seasonal differentiation of photo-thermal comprehensive response characteristics during growth periods of soybeans may help soybean introduction and breeding for adaption. 【Method】 A total of 1 519 representative world wild and cultivated soybeans were selected and tested with two-year spring seeding and summer seeding field trials at Dangtu, Anhui Province. The difference in growth period between sowing seasons was used to evaluate the photo-thermal comprehensive response sensitivity (PTCRS) of each soybean accession, and to study the photo-temperature response characteristics of the growth and development stages of various geographic and ecological soybeans.【Result】 (1) The photothermal response characteristics of soybeans existed throughout the period of growth and development. (2) With the migration of wild soybeans from south to north, the PTCRS of the days from sowing to flowering (DSF) and days from sowing to maturity (DSM) decreased, the PTCRS of the days from flowering to maturity (DFM) increased, and the photothermal response type changed from the front-sensitive and post-insensitive to the front-insensitive and post-sensitive, and the photothermal response of DSM is sensitive. (3) With the domestication of wild soybeans to cultivated soybeans, the PTCRS of DSF and DSM decreased by 20% and 16%, respectively, and relatively small changes were observed for the PTCRS of DFM. The main photothermal response type changed from the front-sensitivity and post-insensitive to the front-insensitive and post-sensitive and the front-insensitive and post-insensitive. (4) The PTCRS of DSM of summer-autumn (SA) and spring (SP) sowing type soybeans both show gradual decrease from south to north. The geographical differentiation of PTCRS of DSF and DFM of SA and SP is different that when migrate from south to north, the PTCRS of DSF of SA decreased, and the PTCRS of DFM of SA first increased and then decreased, and the PTCRS of DSF of SP there was no significant change, and the PTCRS of DFM of SP decreased. (5) With the Huang-Huai and Yangtze River Valleys and South China as the origin center of cultivated soybeans, the PTCRS of DSF, DFM and DSM decreased significantly when spreading north to Northeast China, Russian Far East and Southern Sweden. The PTCRS of DSF and DSM decreased when spreading east to Korean Peninsula and Japan Island and west to Northern North America, Southern North America and the Central and South America, but no obvious change was observed for the PTCRS of DFM. When cultivated soybeans spread south to Southeast Asia, South Asia and Africa, the PTCRS of DSF and DSM increased, and there was no significant change in PTCRS of DFM. (6) Comparing the PTCRS between different ecotypes in the same eco-region, the PTCRS of DSF, DFM and DSM of SP was the smallest, and the PTCRS of DSF of wild soybeans was stronger than that of SA, and the PTCRS of DFM of wild soybeans was weaker than that of SA, and there was no significant difference between the PTCRS of DSM of wild soybeans and SA. Comparison of PTCRS between different geographic and sowing-seasonal eco-type of soybeans, PTCRS of DSF: Southern wild soybeans is the most sensitive, followed by wild soybeans in the Yangtze River Valleys and SA in the Southern, followed by Huang-Huai wild soybeans and SA in the Yangtze River Valleys, and the remaining geo-ecotypes have no significant differences, all of which are relatively insensitive; PTCRS of DFM: SA in the Yangtze River Valleys is the most sensitive, followed by the Northeast and Huang-Huai wild soybeans and the Southern and Huang-Huai SA, and the remaining geo-ecotypes have relatively small differences, all of which are relatively insensitive; PTCRS of DSM: there is no significant difference between wild soybeans and SA in the Southern and the Yangtze River Valleys, all of which are sensitive, followed by Huang-Huai wild soybeans, followed by Northeast wild soybeans and Huang-Huai SA, and the PTCRS of SP is the smallest, and it decreases significantly with the increase of latitude.【Conclusion】 The photo-thermal comprehensive conditions determined by geography and sowing season are important factors affecting soybean growth and development. Differentiation of response to photo-thermal comprehensive conditions existed in wild and cultivated soybeans of different geography and sowing season ecological types. With cultivated soybeans spread from Huang-Huai and Changjiang River Valleys and South China to geographical regions of different latitudes in the world, different changes were observed for the photo-thermal comprehensive response during growth period. Sensitivity to photo-thermal during sowing to maturity is the original trait of soybean, and the summer-autumn sowing type soybeans in the Middle and Lower Yangtze Valleys may be the most cultivated type with this wild primitive trait.

Key words: cultivated soybean (Glycine max (L.) Merr.), wild soybean (Glycine soja Sieb. and Zucc.), growth period traits, geographic and sowing-seasonal photo-thermal eco-response

Table 1

Geographic distribution of tested materials"

栽培大豆地理分布
Geographic region of cultivated soybean		 中国大陆野生和栽培大豆生态区分布
Eco-region of Chinese mainland wild and cultivated soybean
地理亚群
Geo-subpopulation		 地理区域
Geo-region		 熟期组
Maturity group		 材料数
No. of materials		 类型
Type		 生态区
Eco-region		 熟期组
Maturity group		 材料数
No. of materials
O 中国黄淮HCHN Ⅰ-Ⅴ 226 野生大豆
WA G. soja		 Ⅳ 68
中国南方SCHN 0-Ⅸ 542 Ⅲ 76
中国台湾TCHN Ⅴ-Ⅷ 11 Ⅱ 71
A 中国东北NCHN 000-Ⅳ 173 Ⅰ 52
俄罗斯远东RUFE 000-Ⅲ 38 合计Sum 267
瑞典南部SSWE 000 17 栽培春大豆
SP G. max		 Ⅳ 0-Ⅴ 168
B 朝鲜半岛KORP 0-Ⅵ 20 Ⅲ 0-Ⅳ 33
日本岛JPAN 0-Ⅶ 38 Ⅰ 000-Ⅳ 173
C 东南亚SEAS Ⅴ-Ⅹ 25 合计Sum 374
南亚SASI Ⅳ-Ⅹ 15 栽培夏秋大豆
SA G. max		 Ⅳ Ⅳ-Ⅸ 178
非洲AFRI Ⅵ-Ⅹ 11 Ⅲ Ⅲ-Ⅷ 163
D 北美北部NNAM 000-Ⅴ 69 Ⅱ Ⅰ-Ⅵ 226
北美南部SNAM Ⅳ-Ⅹ 37 合计Sum 567
中南美CSAM Ⅴ-Ⅹ 30 总计Total 1208
总计Total 1252

Table 2

The descriptive statistics of growth period related traits and PTCRS"

性状
Trait		 环境
Env.		 组距
Class interval		 材料数
No. of materials		 均值a
Mean		 变幅
Range		 遗传变
异系数
GCV (%)		 遗传率
h2(%)
生育前期
DSF (d)		 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100 100-110 110-120 120-130 130-140 140-150
18SUS 278 384 310 241 185 94 27 1519 45.4 20-88 31.9 99.6
19SUS 211 461 284 241 192 80 24 1493 45.4 22-85 31.8 99.3
18SPS 26 330 389 187 61 96 124 76 89 69 57 11 1515 62.4 25-136 45.5 99.5
19SPS 7 200 304 303 138 125 119 53 84 70 47 28 1 1479 65.7 27-143 40.5 99.0
生育后期
DFM (d)
32-40 40-48 48-56 56-64 64-72 72-80 80-88 88-96 96-104 104-112 112-120 120-128 128-136 136-144
18SUS 2 121 431 480 380 95 9 1518 59.4 38-86 13.5 95.2
19SUS 14 199 603 526 114 3 1459 54.8 33-80 10.8 85.8
18SPS 19 132 140 152 114 133 175 199 144 129 85 55 16 2 1495 80.7 35-139 28.7 95.4
19SPS 4 57 156 252 247 230 232 158 71 15 4 1426 72.2 32-117 19.9 92.5
全生育期
DSM (d)
66-76 76-86 86-96 96-106 106-116 116-126 126-136 136-146 146-156 156-166 166-176 176-186 186-196 196-206
18SUS 52 254 198 321 224 235 203 30 1 1518 104.8 66-149 17.0 99.1
19SUS 16 177 379 394 260 169 63 1 1459 100.3 66-137 13.0 97.2
18SPS 46 148 44 114 18 122 136 53 121 86 273 189 146 5 1501 142.9 71-201 25.6 98.4
19SPS 2 29 74 121 216 136 82 103 209 206 143 86 30 2 1439 137.2 75-198 20.7 98.2
光温综合反
应敏感性
PTCRS (%)		 0-10 10-20 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100 100-110 110-120 120-130 130-140
生育前期
DSF		 Meana 78 584 306 91 118 81 83 88 68 20 1517 32.6 0-99 69.0 95.0
生育后期
DFM		 Meana 296 234 186 158 164 182 126 92 52 13 1 1 1505 35.2 0-113 64.0 82.6
全生育期
DSM		 Meana 108 82 84 132 122 80 83 92 230 271 178 40 4 1 1507 66.1 0-131 46.6 90.2

Table 3

Multi-environment joint analysis of variance of growth period related traits"

变异来源
Source		 生育前期 DSF 生育后期 DFM 全生育期 DSM
自由度
DF		 均方
MS		 F值
F value		 自由度
DF		 均方
MS		 F值
F value		 自由度
DF		 均方
MS		 F值
F value
环境 Env. 3 364996.00 1951.42** 3 384088.00 989.83** 3 1370239.00 3307.50**
环境内区组 Blk (Env.) 5 61.78 10.12** 5 196.98 9.05** 5 146.96 7.65**
基因型 Geno 1518 3582.12 27.37** 1517 1059.90 4.90** 1517 4151.92 14.39**
基因型×环境 Geno×Env. 4484 134.95 22.11** 4377 226.48 10.41** 4396 302.02 15.73**
误差 Error 7171 6.10 6470 21.76 6575 19.20

Table 4

Average growth period of soybeans of different ecotypes"

群体
Population		 生态区
Eco-region		 生育前期DSF(d) 生育后期DFM(d) 全生育期DSM(d)
春播均值a SPS-mean 夏播均值b SUS-mean 春播均值 SPS-mean 夏播均值SUS-mean 春播均值 SPS-mean 夏播均值SUS-mean
WA Ⅳ 119.9±15.32 76.7±8.11 54.9±9.34 49.5±3.90 174.8±9.93 126.2±7.38
Ⅲ 97.4±23.97 65.4±10.23 67.2±18.40 49.4±4.34 164.3±8.51 114.9±8.63
Ⅱ 70.3±24.37 51.9±13.30 72.6±17.74 50.5±5.16 142.7±28.16 102.3±11.41
Ⅰ 47.0±15.10 40.3±10.22 78.3±18.49 52.3±5.10 125.2±25.52 92.5±10.82
G. soja 86.1±33.39 59.8±16.91 67.7±18.33 50.3±4.72 153.6±26.79 110.0±15.51
SA Ⅳ 92.8±20.10 62.8±9.28 81.5±13.28 60.7±4.85 174.3±15.38 123.6±9.39
Ⅲ 72.9±15.50 53.7±6.97 93.3±11.39 62.3±4.37 166.0±13.50 116.0±7.11
Ⅱ 50.4±13.13 40.6±7.28 82.8±14.21 62.1±5.71 133.1±21.35 102.5±9.89
总计c Total 70.2±24.15 51.3±12.31 85.4±14.08 61.7±5.12 155.4±25.51 113.0±12.75
SP Ⅳ 52.1±9.64 41.9±5.77 72.1±16.47 55.7±5.26 124.2±21.69 97.6±9.26
Ⅲ 48.5±10.61 38.7±6.87 63.0±15.05 52.8±6.28 111.7±22.08 91.5±11.28
Ⅰ 39.4±10.93 32.0±7.38 61.1±15.36 59.3±5.35 100.1±23.45 90.9±10.53
总计d Total 45.9±12.00 37.0±8.17 66.2±16.67 57.1±5.80 112.0±25.27 94.0±10.55
G. max-CHNM 60.5±23.45 45.7±12.91 77.8±17.84 59.9±5.85 138.1±33.11 105.5±15.13
O 65.4±22.82 48.8±11.84 81.7±16.16 60.1±5.99 146.8±28.57 108.9±14.01
A 38.7±9.73 31.3±6.69 60.1±14.71 58.8±5.50 98.6±21.70 89.8±10.14
B 49.9±16.18 40.0±9.30 81.6±24.04 64.0±8.81 131.4±31.84 103.8±14.43
C 86.5±23.73 60.6±11.98 83.4±14.63 61.7±4.63 167.4±21.90 121.9±12.11
D 51.6±17.18 40.5±10.44 86.5±16.93 67.2±5.44 138.0±28.70 107.7±13.48
G. max-WLD 59.1±23.41 44.8±13.15 78.3±18.55 60.9±6.43 137.2±33.40 105.5±15.46
总计e Total 63.9±27.43 47.4±15.01 76.4±18.95 59.0±7.38 140.0±32.93 106.3±15.56
CK 70.3 48.3 98 62.8 168.3 111

Table 5

Analysis of variance of PTCRS at different growth stages"

变异来源
Source		 生育前期光温综合反应敏感性 PTCRSDSF 生育后期光温综合反应敏感性 PTCRSDFM 全生育期光温综合反应敏感性 PTCRSDSM
自由度 DF 均方 MS F值 F value 自由度 DF 均方 MS F值 F value 自由度 DF 均方 MS F值 F value
年份 Year 1 6829.90 390.39** 1 20936.10 299.87** 1 1646.88 24.39**
基因型 Geno 1516 339.68 19.42** 1504 386.65 5.54** 1506 656.92 9.73**
误差 Error 1461 17.50 1378 69.82 1394 67.53

Table 6

Correlation between PTCRSs and growth periods"

相关系数
Correlation coefficient		 PTCRSDSF PTCRSDFM PTCRSDSM DSFSp DFMSp DSMSp
PTCRSDSF
PTCRSDFM -0.19**
PTCRSDSM 0.60** 0.67**
DSFSp 0.97** -0.03 0.70**
DFMSp -0.20** 0.94** 0.62** -0.06*
DSMSp 0.72** 0.50** 0.96** 0.83** 0.51**

Fig. 1

Comparison of PTCRS in the growth stages (A: DSF, B: DFM, C: DSM) of soybeans in various geographical ecotypes of China and (D) the growth stages of the cultivated soybeans in different regions of the worldPTCRSDSF: Photothermal comprehensive response sensitivity during from sowing to flowering; PTCRSDFM: Photothermal comprehensive response sensitivity during from flowering to maturity; PTCRSDSM: Photothermal comprehensive response sensitivity during from sowing to maturity. Ⅳ: Central South、Southwest and South China soybean eco-region, Ⅲ: Middle and Lower Changjiang Valley soybean eco-region, Ⅱ: Huang-Huai-Hai soybean eco-region, Ⅰ: Northern soybean eco-region; O: The materials come from Huang-Huai River Valleys (HCHM) and Changjiang River Valleys and its south (SCHM) and China's Taiwan (TCHN). A: The materials are from Northeast China (NCHM), Far-East of Russia (RUFE) and southern Sweden (SSWE). B: The materials are from Korea Peninsular (KORP) and Japan islands (JPAN). C: The materials are from Southeast Asia (SEAS), South Asia (SASI) and Africa (AFRI). D: The materials are from northern North America (NNAM), southern North America (SNAM) and Central and South America (CSAM). G. soja: Glycine soja Sieb. and Zucc., SA: Summer-autumn sowing type soybeans, SP: Spring sowing type soybeans. “x” in the figure represents the average value of subgroup, the top and bottom represents the maximum and minimum values respectively, the shape represents distribution of the subgroup materials and the width of the shape is distribution frequency. The same as below"

Fig. 2

Comparison of PTCRS of wild soybean accessions and cultivated soybean accessions in mainland ChinaG. soja: Glycine soja Sieb. and Zucc.; G. max: Glycine max (L.) Merr.; DSF: Days from sowing to flowering; DFM: Days from flowering to maturity; DSM: Days from sowing to maturity. *** and * indicates there are extremely significant (P≤0.01) and significant difference(P≤0.05) between G. soja and G. max, respectively. The same as below"

Table 7

Comparison of PTCRS in different growth stages of different geographical ecotype soybeans (%)"

群体
Population		 生态区
Eco-region		 生育前期 DSF 生育后期 DFM 全生育期 DSM
均值
Mean		 变幅
Range		 Duncan group 均值
Mean		 变幅
Range		 Duncan group 均值
Mean		 变幅
Range		 Duncan group
WA Ⅳ 79.4 11.1-94.4 D 11.4 0.0-62.1 A 88.4 59.2-106.1 F
Ⅲ 59.6 3.9-96.8 C 32.5 0.0-99.9 CD 91.5 71.9-105.6 F
Ⅱ 35.9 3.5-80.6 B 41.2 0.0-88.3 DE 75.7 0.0-103.5 E
Ⅰ 17.3 0.4-62.9 A 47.3 0.0-87.1 E 62.6 1.3-105.3 D
G. soja 50.1 0.4-96.8 32.3 0.0-99.9 80.9 0.0-106.1
SA Ⅳ 56.7 8.8-95.9 C 39.4 0.3-87.3 CDE 95.9 39.3-131.0 F
Ⅲ 37.4 8.5-82.9 B 58.3 19.3-112.6 F 95.3 34.2-124.6 F
Ⅱ 20.8 0.4-90.6 A 40.4 0.0-97.2 CDE 60.6 0.0-116.2 CD
总计c Total 36.9 0.4-95.9 45.3 0.0-112.6 81.6 0.0-131.0
SP Ⅳ 21.4 6.4-82.7 A 31.7 0.0-86.0 C 53.0 18.0-104.4 C
Ⅲ 20.7 10.8-55.8 A 21.2 0.0-75.1 B 40.8 4.7-100.4 B
Ⅰ 16.2 2.3-66.5 A 13.4 0.0-91.1 AB 24.2 0.0-111.0 A
总计d Total 18.9 2.3-82.8 22.3 0.0-91.1 38.7 0.0-111.0
G. max-CHNM 29.7 0.4-95.9 36.1 0.0-112.6 64.5 0.0-131.0
O 32.8 0.4-95.9 C 41.4 0.0-112.6 B 73.7 0.0-131.0 C
A 16.0 2.3-66.5 A 12.5 0.0-91.1 A 23.2 0.0-111.0 A
B 21.3 3.1-79.8 AB 38.1 0.0-90.5 B 57.5 5.7-110.5 B
C 49.7 10.1-99.4 D 39.7 0.0-81.6 B 85.8 10.2-111.4 D
D 23.2 7.3-78.5 B 40.4 0.0-87.1 B 62.6 0.0-114.9 B

Fig. 3

Variations of PTCRS of growth periods in cultivated soybeans in various geographical regions of the world Circle represents distribution of PTCRS of the subgroup materials, and blue indicates the weakest PTCRS, red indicates the strongest PTCRS and the color depth represents the size of PTCRS"

Fig. 4

Comparison of PTCRS of growth periods in wild soybeans, summer-autumn sowing type soybeans and spring sowing type soybeansNS: There is no significant differences (P>0.05) between different types of soybeans, respectively"

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