园艺学报 ›› 2023, Vol. 50 ›› Issue (7): 1455-1466.doi: 10.16420/j.issn.0513-353x.2022-0699
刘春洋, 彭朝凤, 程世平*(), 姚鹏强, 耿喜宁, 谢丽华
收稿日期:
2022-09-19
修回日期:
2023-03-26
出版日期:
2023-07-25
发布日期:
2023-07-26
通讯作者:
基金资助:
LIU Chunyang, PENG Chaofeng, CHENG Shiping*(), YAO Pengqiang, GENG Xining, XIE Lihua
Received:
2022-09-19
Revised:
2023-03-26
Published:
2023-07-25
Online:
2023-07-26
摘要:
以5年生‘凤丹’牡丹(Paeonia ostii‘Fengdan’)为试验材料,使用“树木非离体枝芽加热处理装置”(ZL200610113448.X)施加高温处理诱导产生2n雌配子。大孢子母细胞减数分裂进程与花蕾的外部形态和小孢子发育进程存在时序相关性,当花蕾发育至“大风铃”期前后,小孢子发育至单核花粉期后3 ~ 4 d,40 ℃高温处理花蕾4 h,自然授粉后代群体中可获得高比率(2.45%)三倍体植株。
刘春洋, 彭朝凤, 程世平, 姚鹏强, 耿喜宁, 谢丽华. 高温诱导‘凤丹’牡丹2n雌配子创制三倍体[J]. 园艺学报, 2023, 50(7): 1455-1466.
LIU Chunyang, PENG Chaofeng, CHENG Shiping, YAO Pengqiang, GENG Xining, XIE Lihua. Creation of Triploid Germplasm in Paeonia ostii‘Fengdan’Through 2n Female Gametes Inducing with High Temperature Treatment[J]. Acta Horticulturae Sinica, 2023, 50(7): 1455-1466.
发育时期 Development stage | 花蕾形态 Morphology of flower bud | 小孢子发育 Development progress of microspore | 大孢子发育 Development progress of megaspore | 胚珠形态 Morphology of ovule | 染色体加倍处理 High temperature treatment |
---|---|---|---|---|---|
花粉母细胞 减数分裂期 Meiosis stage of the pollen mother cells | 介于大、小风铃期之间 Between big wind-bell and small wind -bell | 进入减数分裂期 In meiosis period | — | 胚珠原基分化出珠柄,珠心开始形成 Funiculus was formed by ovule primordium, nucellus formation initially | 小孢子染色体加倍 Chromosomes doubling of microspore |
3 ~ 5 d后(花粉母细胞四分体至单核花粉期) Uni-nucleate microspore stage from tetrad stage after 3 ~ 5 d | 接近大风铃期 Approximately in big wind-bell stage | 单核花粉形成, 可见四分体 Uni-nucleate pollen formation with visible tetrad | 减数分裂前期 Ⅰ Prophase I of meiosis | 珠被隆起,珠心顶端与两侧珠被围成120 ~ 150°角,整体呈元宝形 Integument expanding,120-150° angle was formed by the tip cells of nucellus and two sides of integument, as a ingot | 大孢子染色体加倍开始 The beginning of chromosomes doubling of Macrospore |
单核花粉形成3 ~ 5 d后 After 3 ~ 5 d of Uni-nucleate pollen formation | 大风铃期稍后 After big wind-bell stage | 单核靠边期 Late uni-nucleate microspore | 减数分裂末期Ⅰ至前期Ⅱ From anaphase I to prophaseⅡ | 珠被半包被,珠心顶端与两侧珠被围成150 ~ 180°角 Nucellus was half covered by integument,150-180° angle was formed by the tip cells of nucellus and two sides of integument | 大孢子染色体加倍接近结束 Nearby the end of chromosomes doubling of Macrospore |
表1 ‘凤丹’牡丹花蕾形态与大、小孢子发育进程之间的对应关系
Table 1 Relationship between flower bud morphology and development process of megaspore and microspore in Paeonia ostii‘Fengdan’
发育时期 Development stage | 花蕾形态 Morphology of flower bud | 小孢子发育 Development progress of microspore | 大孢子发育 Development progress of megaspore | 胚珠形态 Morphology of ovule | 染色体加倍处理 High temperature treatment |
---|---|---|---|---|---|
花粉母细胞 减数分裂期 Meiosis stage of the pollen mother cells | 介于大、小风铃期之间 Between big wind-bell and small wind -bell | 进入减数分裂期 In meiosis period | — | 胚珠原基分化出珠柄,珠心开始形成 Funiculus was formed by ovule primordium, nucellus formation initially | 小孢子染色体加倍 Chromosomes doubling of microspore |
3 ~ 5 d后(花粉母细胞四分体至单核花粉期) Uni-nucleate microspore stage from tetrad stage after 3 ~ 5 d | 接近大风铃期 Approximately in big wind-bell stage | 单核花粉形成, 可见四分体 Uni-nucleate pollen formation with visible tetrad | 减数分裂前期 Ⅰ Prophase I of meiosis | 珠被隆起,珠心顶端与两侧珠被围成120 ~ 150°角,整体呈元宝形 Integument expanding,120-150° angle was formed by the tip cells of nucellus and two sides of integument, as a ingot | 大孢子染色体加倍开始 The beginning of chromosomes doubling of Macrospore |
单核花粉形成3 ~ 5 d后 After 3 ~ 5 d of Uni-nucleate pollen formation | 大风铃期稍后 After big wind-bell stage | 单核靠边期 Late uni-nucleate microspore | 减数分裂末期Ⅰ至前期Ⅱ From anaphase I to prophaseⅡ | 珠被半包被,珠心顶端与两侧珠被围成150 ~ 180°角 Nucellus was half covered by integument,150-180° angle was formed by the tip cells of nucellus and two sides of integument | 大孢子染色体加倍接近结束 Nearby the end of chromosomes doubling of Macrospore |
图1 ‘凤丹’牡丹大孢子母细胞减数分裂关键时期与花粉发育进程的对应关系 a ~ c:花粉发育时期,其中a为单核花粉初期,b、c为花粉单核早中期至单核靠边期;d ~ g:雌配子发育时期,其中d为细线期,e为粗线期,f为末期Ⅰ,g为前期Ⅱ。
Fig. 1 The relationship between the critical meiosis stage of megaspore mother cell and pollen development of Paeonia ostii‘Fengdan’ a-c:Pollen development process;a:Early stage of mononuclear pollen;b,c:Early ormid-uninucleate stage to mid-late uninucleate stage of microspore;d-g:Megaspore development process;d:Leptonema;e:Pachytene;f:Telophase Ⅰ;g:Prophase Ⅱ.
单核花粉产生后天数/d Days after formation of mononuclear pollen | 处理温度/℃ Treatment temperature | 处理时间/h Treatment duration | 处理花蕾数 Flower bud number | 收获种子数 Seed number | 单果结实数 Seed number per follicle | 成苗数 Seedling number | 三倍体株数 Triploid number | 三倍体得率/% Rate of triploid |
---|---|---|---|---|---|---|---|---|
1 | 38 | 2 | 18 | 395 | 21.94 | 205 | 0 | |
4 | 17 | 340 | 20.00 | 160 | 0 | |||
6 | 19 | 275 | 14.47 | 141 | 0 | |||
40 | 2 | 18 | 351 | 19.50 | 186 | 0 | ||
4 | 20 | 376 | 18.80 | 192 | 0 | |||
6 | 17 | 175 | 10.29 | 91 | 0 | |||
42 | 2 | 16 | 295 | 18.44 | 160 | 0 | ||
4 | 20 | 256 | 12.80 | 126 | 0 | |||
6 | 18 | 156 | 8.67 | 80 | 0 | |||
44 | 2 | 17 | 176 | 10.35 | 83 | 0 | ||
4 | 19 | 93 | 4.89 | 49 | 0 | |||
6 | 15 | 40 | 2.67 | 18 | 0 | |||
2 | 38 | 2 | 22 | 463 | 21.05 | 246 | 0 | |
4 | 18 | 365 | 20.28 | 183 | 0 | |||
6 | 19 | 247 | 13.00 | 126 | 1 | 0.79 | ||
40 | 2 | 17 | 361 | 21.24 | 190 | 0 | ||
4 | 19 | 366 | 19.26 | 172 | 1 | 0.58 | ||
6 | 20 | 216 | 10.80 | 115 | 0 | |||
42 | 2 | 22 | 420 | 19.09 | 225 | 0 | ||
4 | 22 | 271 | 12.32 | 133 | 0 | |||
6 | 20 | 171 | 8.55 | 90 | 0 | |||
44 | 2 | 18 | 173 | 9.61 | 83 | 0 | ||
4 | 16 | 82 | 5.13 | 40 | 0 | |||
6 | 16 | 43 | 2.69 | 26 | 0 | |||
3 | 38 | 2 | 19 | 396 | 20.84 | 216 | 0 | |
4 | 18 | 386 | 21.44 | 199 | 0 | |||
6 | 21 | 217 | 10.33 | 110 | 1 | 0.91 | ||
40 | 2 | 20 | 408 | 20.40 | 191 | 0 | ||
4 | 15 | 278 | 18.53 | 151 | 2 | 1.32 | ||
6 | 18 | 189 | 10.50 | 107 | 1 | 0.93 | ||
42 | 2 | 17 | 304 | 17.88 | 145 | 1 | 0.69 | |
4 | 16 | 206 | 12.88 | 112 | 0 | |||
6 | 19 | 162 | 8.53 | 90 | 0 | |||
44 | 2 | 19 | 170 | 8.95 | 88 | 0 | ||
4 | 17 | 83 | 4.88 | 45 | 0 | |||
6 | 20 | — | — | — | 0 | |||
4 | 38 | 2 | 19 | 400 | 21.05 | 211 | 0 | |
4 | 19 | 366 | 19.26 | 191 | 0 | |||
6 | 20 | 254 | 12.70 | 136 | 1 | 0.74 | ||
40 | 2 | 21 | 405 | 19.29 | 210 | 1 | 0.48 | |
4 | 15 | 286 | 19.07 | 144 | 2 | 1.39 | ||
6 | 18 | 291 | 16.17 | 144 | 1 | 0.69 | ||
42 | 2 | 16 | 308 | 19.25 | 161 | 1 | 0.62 | |
4 | 18 | 196 | 10.89 | 100 | 0 | |||
6 | 17 | 176 | 10.35 | 101 | 0 | |||
44 | 2 | 15 | 116 | 7.73 | 55 | 0 | ||
4 | 16 | 73 | 4.56 | 39 | 0 | |||
6 | 21 | 39 | 1.86 | 20 | 0 | |||
5 | 38 | 2 | 16 | 368 | 23.00 | 200 | 0 | |
4 | 18 | 343 | 19.06 | 165 | 0 | |||
6 | 19 | 246 | 12.95 | 130 | 0 | |||
40 | 2 | 21 | 425 | 20.24 | 205 | 0 | ||
4 | 20 | 383 | 19.15 | 183 | 1 | 0.55 | ||
6 | 19 | 206 | 10.84 | 111 | 1 | 0.90 | ||
42 | 2 | 19 | 348 | 18.32 | 184 | 0 | ||
4 | 19 | 261 | 13.74 | 140 | 0 | |||
6 | 18 | 167 | 9.28 | 75 | 0 | |||
44 | 2 | 16 | 149 | 9.31 | 60 | 0 | ||
4 | 17 | 89 | 5.24 | 43 | 0 | |||
6 | 19 | 29 | 1.53 | 18 | 0 | |||
对照Control | 30 | 736 | 24.53 | 369 | 0 | |||
总计Total | 1 128 | 15 565 | — | 7 969 | 15 | - |
表2 不同高温诱导‘凤丹’牡丹雌配子加倍选育三倍体的诱导率(2019年)
Table 2 Triploid production of megaspore chromosome doubling by high temperature induction at different developmental stages,temperatures, and durations in Paeonia ostii‘Fengdan’
单核花粉产生后天数/d Days after formation of mononuclear pollen | 处理温度/℃ Treatment temperature | 处理时间/h Treatment duration | 处理花蕾数 Flower bud number | 收获种子数 Seed number | 单果结实数 Seed number per follicle | 成苗数 Seedling number | 三倍体株数 Triploid number | 三倍体得率/% Rate of triploid |
---|---|---|---|---|---|---|---|---|
1 | 38 | 2 | 18 | 395 | 21.94 | 205 | 0 | |
4 | 17 | 340 | 20.00 | 160 | 0 | |||
6 | 19 | 275 | 14.47 | 141 | 0 | |||
40 | 2 | 18 | 351 | 19.50 | 186 | 0 | ||
4 | 20 | 376 | 18.80 | 192 | 0 | |||
6 | 17 | 175 | 10.29 | 91 | 0 | |||
42 | 2 | 16 | 295 | 18.44 | 160 | 0 | ||
4 | 20 | 256 | 12.80 | 126 | 0 | |||
6 | 18 | 156 | 8.67 | 80 | 0 | |||
44 | 2 | 17 | 176 | 10.35 | 83 | 0 | ||
4 | 19 | 93 | 4.89 | 49 | 0 | |||
6 | 15 | 40 | 2.67 | 18 | 0 | |||
2 | 38 | 2 | 22 | 463 | 21.05 | 246 | 0 | |
4 | 18 | 365 | 20.28 | 183 | 0 | |||
6 | 19 | 247 | 13.00 | 126 | 1 | 0.79 | ||
40 | 2 | 17 | 361 | 21.24 | 190 | 0 | ||
4 | 19 | 366 | 19.26 | 172 | 1 | 0.58 | ||
6 | 20 | 216 | 10.80 | 115 | 0 | |||
42 | 2 | 22 | 420 | 19.09 | 225 | 0 | ||
4 | 22 | 271 | 12.32 | 133 | 0 | |||
6 | 20 | 171 | 8.55 | 90 | 0 | |||
44 | 2 | 18 | 173 | 9.61 | 83 | 0 | ||
4 | 16 | 82 | 5.13 | 40 | 0 | |||
6 | 16 | 43 | 2.69 | 26 | 0 | |||
3 | 38 | 2 | 19 | 396 | 20.84 | 216 | 0 | |
4 | 18 | 386 | 21.44 | 199 | 0 | |||
6 | 21 | 217 | 10.33 | 110 | 1 | 0.91 | ||
40 | 2 | 20 | 408 | 20.40 | 191 | 0 | ||
4 | 15 | 278 | 18.53 | 151 | 2 | 1.32 | ||
6 | 18 | 189 | 10.50 | 107 | 1 | 0.93 | ||
42 | 2 | 17 | 304 | 17.88 | 145 | 1 | 0.69 | |
4 | 16 | 206 | 12.88 | 112 | 0 | |||
6 | 19 | 162 | 8.53 | 90 | 0 | |||
44 | 2 | 19 | 170 | 8.95 | 88 | 0 | ||
4 | 17 | 83 | 4.88 | 45 | 0 | |||
6 | 20 | — | — | — | 0 | |||
4 | 38 | 2 | 19 | 400 | 21.05 | 211 | 0 | |
4 | 19 | 366 | 19.26 | 191 | 0 | |||
6 | 20 | 254 | 12.70 | 136 | 1 | 0.74 | ||
40 | 2 | 21 | 405 | 19.29 | 210 | 1 | 0.48 | |
4 | 15 | 286 | 19.07 | 144 | 2 | 1.39 | ||
6 | 18 | 291 | 16.17 | 144 | 1 | 0.69 | ||
42 | 2 | 16 | 308 | 19.25 | 161 | 1 | 0.62 | |
4 | 18 | 196 | 10.89 | 100 | 0 | |||
6 | 17 | 176 | 10.35 | 101 | 0 | |||
44 | 2 | 15 | 116 | 7.73 | 55 | 0 | ||
4 | 16 | 73 | 4.56 | 39 | 0 | |||
6 | 21 | 39 | 1.86 | 20 | 0 | |||
5 | 38 | 2 | 16 | 368 | 23.00 | 200 | 0 | |
4 | 18 | 343 | 19.06 | 165 | 0 | |||
6 | 19 | 246 | 12.95 | 130 | 0 | |||
40 | 2 | 21 | 425 | 20.24 | 205 | 0 | ||
4 | 20 | 383 | 19.15 | 183 | 1 | 0.55 | ||
6 | 19 | 206 | 10.84 | 111 | 1 | 0.90 | ||
42 | 2 | 19 | 348 | 18.32 | 184 | 0 | ||
4 | 19 | 261 | 13.74 | 140 | 0 | |||
6 | 18 | 167 | 9.28 | 75 | 0 | |||
44 | 2 | 16 | 149 | 9.31 | 60 | 0 | ||
4 | 17 | 89 | 5.24 | 43 | 0 | |||
6 | 19 | 29 | 1.53 | 18 | 0 | |||
对照Control | 30 | 736 | 24.53 | 369 | 0 | |||
总计Total | 1 128 | 15 565 | — | 7 969 | 15 | - |
图2 ‘凤丹’牡丹高温诱导大孢子染色体加倍后代的倍性水平检测 a和b:二倍体和三倍体植株嫩叶流式细胞仪检测;c和d:二倍体和三倍体植株体细胞染色体数;e:1年生三倍体植株(箭头)。
Fig. 2 Ploidy level detection of offspring derived from megaspore chromosome doubling with high temperature in Paeonia ostii‘Fengdan’ a,b:Flow cytometric detection of young leaves from diploid and triploid;c-d:Somatic chromosome number of diploid and triploid;e:One-year old triploid offspring(arrowhead)
单核花粉产生后天数/d Days after formation of mononuclear pollen | 处理花蕾数 Flower bud number | 收获种子数 Seed number | 单果结实数 Seed number per follicle | 成苗数 Seedling number | 三倍体株数 Triploid number | 三倍体得率/% Rate of triploid |
---|---|---|---|---|---|---|
1 | 30 | 583 | 19.43 | 286 | 1 | 0.35 |
2 | 30 | 531 | 17.70 | 250 | 3 | 1.20 |
3 | 30 | 539 | 17.97 | 245 | 6 | 2.45 |
4 | 30 | 564 | 18.80 | 247 | 4 | 1.62 |
5 | 30 | 554 | 18.47 | 273 | 0 | 0 |
对照Control | 50 | 1130 | 22.60 | 603 | 0 | 0 |
总计Total | 200 | 3 901 | - | 1 904 | 14 |
表3 High temperature treatment condition verification of megaspore chromosome doubling by 40 ℃ at different developmental stages in Paeonia ostii‘Fengdan’(2021年)
Table 3
单核花粉产生后天数/d Days after formation of mononuclear pollen | 处理花蕾数 Flower bud number | 收获种子数 Seed number | 单果结实数 Seed number per follicle | 成苗数 Seedling number | 三倍体株数 Triploid number | 三倍体得率/% Rate of triploid |
---|---|---|---|---|---|---|
1 | 30 | 583 | 19.43 | 286 | 1 | 0.35 |
2 | 30 | 531 | 17.70 | 250 | 3 | 1.20 |
3 | 30 | 539 | 17.97 | 245 | 6 | 2.45 |
4 | 30 | 564 | 18.80 | 247 | 4 | 1.62 |
5 | 30 | 554 | 18.47 | 273 | 0 | 0 |
对照Control | 50 | 1130 | 22.60 | 603 | 0 | 0 |
总计Total | 200 | 3 901 | - | 1 904 | 14 |
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