小麦顶端发育动态模式与特征的研究
|
摘要
以顶端发育为主线,对小麦阶段发育与形态发育的生理生态过程进行系统研究和综合分析,有助于揭示小麦顶端发育的动力学,为小麦生育调控及机理模型建立奠定基础。本研究以代表性冬性品种京411(WV)和春性品种扬麦158(SV)为供试基因型,采用分期播种(播期一:EP,9月30日;播期二:MP,10月30日;播期三:LP,3月2日)的方法,于1996—1998年在南京农业大学(南京,32°N)校内农场进行田间试验,定期取样解剖和分析植株的顶端发育进程。结果表明:
1.小麦茎顶端叶原基、苞叶原基、小穗原基分化与播种后累积生长度日(GDD)之间分别具有近S型曲线、常规S型曲线和类半抛物线型S曲线关系,用Logistic数学模型拟合均达极显著水平,并且冬性与春性品种以及主茎与分蘖分化模式基本一致。
2.小麦顶端原基分化热间距(GDD/原基)按叶片—苞叶—小穗次序大幅度降低。冬、春性品种不同顶端原基分化的热间距及苞叶原基、小穗原基分化持续期所需GDD在MP处理中趋于一致,而在EP、LP中表现了明显的基因型差异。不同播期之间苞叶原基分化终止期及小穗原基分化始期的差异以GDD计远远大于苞叶原基分化始期的差异。不同播期生态条件下,春性品种主茎、分蘖叶原基数与苞叶原基数、小穗原基数之间的相关性均极显著,冬性品种表现不显著。但冬、春性品种主茎、分蘖的苞叶原基数与小穗原基数之间的相关均超过极显著水平。
3.播期二冬、春性品种的主茎和T1、T2(主茎第一、二分蘖)小花原基的分化与播后累积GDD的关系均符合抛物曲线(上升段)模式,小花退化符合速降线性模式,可分别用二次曲线和线性方程描述。T3(主茎第三分蘖)以及春性品种EP和冬性品种LP主茎的小花原基分化与退化过程均表现异常。冬、春性品种LP总小花数均极显著低于EP和MP的总小花数,EP和MP之间差异不显著。MP处理品种间总小花数无明显差异,EP和LP品种间差异极显著。此外,不同播期、品种间小花分化始期GDD值差异均极显著;同时冬、春性品种小花分化热间距LP与MP之间的差异也均达极显著水平,但不同品种的变化趋势相反。
4.小麦叶片出生与播后累积GDD的关系遵循两段(速升段、缓升段)线
ABSTRCT A systematic study and comprehensive analysis on phasic development and apex morphogenesis in wheat will help elucidate the dynamic of apex development and support cultural regulation and growth modeling. The present study was conducted on the experiment station of Nanjing Agric. Univ. (32°N ) in two growing seasons from 1996 to 1998. The study included two types of varieties: winter type variety (WV) "Jing 411" and spring type variety (SV) "Yangmai 158" and three sowing dates: Sept. 30 (EP), Oct. 30 (MP), Mar. 2 (LP). By time-course sampling and dissecting, the development progress of apex was observed and analyzed. The major results were as follows.1. Near-S, regular-S and half-parabola-S shape curves could describe accumulated numbers of leaf, bracteal and spikelet primordia in relation to GDD( Growing Degree Days ) after sowing, respectively. The logistic equations of the three types of S curves were all highly significant with R~2 greater than 0.74. The patterns of apex primordium differentiation were similar patterns in winter- and spring-type varieties and on main stem and tillers.2. Thermalchrons of apex primordium differentiation were reduced greatly in the order of leaf, bracteal and spikelet. Thermalchrons of different types of apex primordium differentiation and duration of bracteal and spikelet primordium differentiation in GDD were near the same in winter- and spring-type varieties in MP treatments, but were different significantly in EP and LP treatments. Measured with GDD, differences between different sowing dates for the end of bracteal primordium differentiation and beginning of spikelet primordium differentiation were much greater than for the beginning of bracteal primordium differentiation. Also, leaf primordium number was highly correlated with bracteal and spikelet primordium numbers on main stems and tillers in spring-type variety, but not in winter-type variety. Bracteal primordium number was significantly correlated with spikelet number in the two types of varieties, as well as in main stems and tillers.
3. Parabolic curve (rise stage) and straight line could describe respectively floret differentiation and degeneration in relation to GDD on MS, Tl and T2 (MS for main stem;Tl, T2 for first and second tillers on main stem, respectively) of MP in wheat. The floret differentiation and degeneration models on T3 (third tiller on main stem) of all planting and on MS of EP with spring-type variety and LP with winter-type variety appeared abnormal. Total floret number of LP was less than EP and MP significantly in both varieties. Difference of total floret number between two varieties was not significant in MP, but highly significant in EP and LP. Also, the differences of GDD from sowing to initiation of floret differentiation among different sowing dates and between the varieties were all highly significant, so did the mean thermalchrons of tloret differentiation between MP and LP, but showed an opposite pattern.4. Two-phase linear model equations ( quick rise, slow rise ) divided by glum differentiation stage could describe leaf emergence progress in relation to GDD after sowing in wheat. This pattern was consistent on main stems and tillers of normal development on both winter-and-spring type varieties. The beginning of slow rise phase shifted to an earlier development stage on main stems of winter type varieties of EP and LP, and on T3 of both varieties of EP, MP and LP due to their abnormal development. The thermal rate of leaf emergence on main stem was relatively high and steady during development for winter type variety of MP, and became higher with later sowing in spring type variety. Phyllochrons of EP and MP were higher in winter type variety than in spring type variety, although the difference between the two varieties of LP was not significant.5. Emergence of tiller was mainly regulated by the ecological factors rather than genetic effect in wheat. For winter-type variety, most variable growth duration throughout the entire growing cycle occurred from tillering to jointing. In contrast, timing of single ridge stage was determined by genotypic effect rather than ecological factors. Single ridge to double ridge stage was most sensitive to vernalization progress For spring sowing of winter-type variety, vernalization process could last till floral primordium differentiation stage The corresponding relationships existed between the phenological stages and spike development stages in different sowing dates, although a little variable. Winter-type variety had more ecologically variable leaf primoudia due to its
stronger vernalization requirement. For winter-type variety, the primordium numbers of reproductive organs were not apparently related to the GDD accumulated prior to vernalization completion. Yet for spring-type variety, numbers of different apex primordia including vegetative and reproductive organs were significantly related to the accumulated GDD during the entire phenological stages as well as before anther separation stage.6.Leaf ages (Haun stages) on main stem of winter type variety of MP were significantly less than those of EP and LP at all major spike differentiation stages (MSDS). In spring type variety, leafage of MP were less than those of EP and LP before double ridge stage, but after then, that of LP was the lest. Also, leaf ages of winter type variety of all sowing date treatments were significantly higher than those of spring type variety at all MSDS. Although differences of remaining leaf primordium number between different sowing dates became less with later spike differentiation stages in both winter and spring-type varieties, they were still significant up to stamen and pistil primordium differentiation stage. Remaining leaf primordium numbers of winter and spring-type varieties were very similar at all MSDS. Differences of leaf age indexes on main stems at MSDS between different sowing dates were small in the same variety. But leaf age indexes of winter type variety were significantly higher than those of spring type variety at MSDS.
1.小麦茎顶端叶原基、苞叶原基、小穗原基分化与播种后累积生长度日(GDD)之间分别具有近S型曲线、常规S型曲线和类半抛物线型S曲线关系,用Logistic数学模型拟合均达极显著水平,并且冬性与春性品种以及主茎与分蘖分化模式基本一致。
2.小麦顶端原基分化热间距(GDD/原基)按叶片—苞叶—小穗次序大幅度降低。冬、春性品种不同顶端原基分化的热间距及苞叶原基、小穗原基分化持续期所需GDD在MP处理中趋于一致,而在EP、LP中表现了明显的基因型差异。不同播期之间苞叶原基分化终止期及小穗原基分化始期的差异以GDD计远远大于苞叶原基分化始期的差异。不同播期生态条件下,春性品种主茎、分蘖叶原基数与苞叶原基数、小穗原基数之间的相关性均极显著,冬性品种表现不显著。但冬、春性品种主茎、分蘖的苞叶原基数与小穗原基数之间的相关均超过极显著水平。
3.播期二冬、春性品种的主茎和T1、T2(主茎第一、二分蘖)小花原基的分化与播后累积GDD的关系均符合抛物曲线(上升段)模式,小花退化符合速降线性模式,可分别用二次曲线和线性方程描述。T3(主茎第三分蘖)以及春性品种EP和冬性品种LP主茎的小花原基分化与退化过程均表现异常。冬、春性品种LP总小花数均极显著低于EP和MP的总小花数,EP和MP之间差异不显著。MP处理品种间总小花数无明显差异,EP和LP品种间差异极显著。此外,不同播期、品种间小花分化始期GDD值差异均极显著;同时冬、春性品种小花分化热间距LP与MP之间的差异也均达极显著水平,但不同品种的变化趋势相反。
4.小麦叶片出生与播后累积GDD的关系遵循两段(速升段、缓升段)线
ABSTRCT A systematic study and comprehensive analysis on phasic development and apex morphogenesis in wheat will help elucidate the dynamic of apex development and support cultural regulation and growth modeling. The present study was conducted on the experiment station of Nanjing Agric. Univ. (32°N ) in two growing seasons from 1996 to 1998. The study included two types of varieties: winter type variety (WV) "Jing 411" and spring type variety (SV) "Yangmai 158" and three sowing dates: Sept. 30 (EP), Oct. 30 (MP), Mar. 2 (LP). By time-course sampling and dissecting, the development progress of apex was observed and analyzed. The major results were as follows.1. Near-S, regular-S and half-parabola-S shape curves could describe accumulated numbers of leaf, bracteal and spikelet primordia in relation to GDD( Growing Degree Days ) after sowing, respectively. The logistic equations of the three types of S curves were all highly significant with R~2 greater than 0.74. The patterns of apex primordium differentiation were similar patterns in winter- and spring-type varieties and on main stem and tillers.2. Thermalchrons of apex primordium differentiation were reduced greatly in the order of leaf, bracteal and spikelet. Thermalchrons of different types of apex primordium differentiation and duration of bracteal and spikelet primordium differentiation in GDD were near the same in winter- and spring-type varieties in MP treatments, but were different significantly in EP and LP treatments. Measured with GDD, differences between different sowing dates for the end of bracteal primordium differentiation and beginning of spikelet primordium differentiation were much greater than for the beginning of bracteal primordium differentiation. Also, leaf primordium number was highly correlated with bracteal and spikelet primordium numbers on main stems and tillers in spring-type variety, but not in winter-type variety. Bracteal primordium number was significantly correlated with spikelet number in the two types of varieties, as well as in main stems and tillers.
3. Parabolic curve (rise stage) and straight line could describe respectively floret differentiation and degeneration in relation to GDD on MS, Tl and T2 (MS for main stem;Tl, T2 for first and second tillers on main stem, respectively) of MP in wheat. The floret differentiation and degeneration models on T3 (third tiller on main stem) of all planting and on MS of EP with spring-type variety and LP with winter-type variety appeared abnormal. Total floret number of LP was less than EP and MP significantly in both varieties. Difference of total floret number between two varieties was not significant in MP, but highly significant in EP and LP. Also, the differences of GDD from sowing to initiation of floret differentiation among different sowing dates and between the varieties were all highly significant, so did the mean thermalchrons of tloret differentiation between MP and LP, but showed an opposite pattern.4. Two-phase linear model equations ( quick rise, slow rise ) divided by glum differentiation stage could describe leaf emergence progress in relation to GDD after sowing in wheat. This pattern was consistent on main stems and tillers of normal development on both winter-and-spring type varieties. The beginning of slow rise phase shifted to an earlier development stage on main stems of winter type varieties of EP and LP, and on T3 of both varieties of EP, MP and LP due to their abnormal development. The thermal rate of leaf emergence on main stem was relatively high and steady during development for winter type variety of MP, and became higher with later sowing in spring type variety. Phyllochrons of EP and MP were higher in winter type variety than in spring type variety, although the difference between the two varieties of LP was not significant.5. Emergence of tiller was mainly regulated by the ecological factors rather than genetic effect in wheat. For winter-type variety, most variable growth duration throughout the entire growing cycle occurred from tillering to jointing. In contrast, timing of single ridge stage was determined by genotypic effect rather than ecological factors. Single ridge to double ridge stage was most sensitive to vernalization progress For spring sowing of winter-type variety, vernalization process could last till floral primordium differentiation stage The corresponding relationships existed between the phenological stages and spike development stages in different sowing dates, although a little variable. Winter-type variety had more ecologically variable leaf primoudia due to its
stronger vernalization requirement. For winter-type variety, the primordium numbers of reproductive organs were not apparently related to the GDD accumulated prior to vernalization completion. Yet for spring-type variety, numbers of different apex primordia including vegetative and reproductive organs were significantly related to the accumulated GDD during the entire phenological stages as well as before anther separation stage.6.Leaf ages (Haun stages) on main stem of winter type variety of MP were significantly less than those of EP and LP at all major spike differentiation stages (MSDS). In spring type variety, leafage of MP were less than those of EP and LP before double ridge stage, but after then, that of LP was the lest. Also, leaf ages of winter type variety of all sowing date treatments were significantly higher than those of spring type variety at all MSDS. Although differences of remaining leaf primordium number between different sowing dates became less with later spike differentiation stages in both winter and spring-type varieties, they were still significant up to stamen and pistil primordium differentiation stage. Remaining leaf primordium numbers of winter and spring-type varieties were very similar at all MSDS. Differences of leaf age indexes on main stems at MSDS between different sowing dates were small in the same variety. But leaf age indexes of winter type variety were significantly higher than those of spring type variety at MSDS.
引文
1.金善宝.中国小麦生态.北京:科学出版社,1991:31-157
2.李存东,曹卫星.小麦阶段发育的生理生态特征评述.南京农业大学学报.1997,20(2):17-21
3.张锦熙等.小麦“叶龄指标促控法”的研究.中国农业科学,1981(2):1—13
4.张锦熙,刘锡山,阎润涛.小麦冬春品种类型及各生育阶段主茎叶数与穗分化进程变异规律的研究.中国农业科学,1986(2):27—35
5.凌启鸿等.叶龄余数在稻穗分化进程鉴定中的应用价值 中国农业科学,1980(4):1—11
6.苗果园等.小麦品种温光效应与主茎叶数的关系.作物学报,1992,18(5):321-329
7.曹广才等.小麦主茎总叶数的变异.作物学报.1990,16(1):73—82
8.曹卫星,江海东.小麦温光反应与发育进程的模拟.南京农业大学学报,1996,19(1):9-16
9. Cao W., and D. N. Moss. Temperature and daylength interaction on phyllochron in wheat and barley. Crop Sci. 1989, 29: 1046-1048
10. Cao W., and D. N. Moss. Sensitivity of winter wheat phyllochron to environmental changes. Agron. J. 1994, 86: 63-66
11. Cao W., and D. N. Moss. Modeling phasic development in wheat: a conceptual integration of physiological components. Journal of Agric. Sci. 1997, 129, 163-172
12. Jame Y. W. et al. Modeling Follar stages of two contrasting spring wheat varieties growth on the Canadian prairies. Modeling for crop-climate-soil-pest system and its applications in sustainable crop production. Published by JAAS. Nanjing, China. 1998, 53-60
13. Jamieson P. D. et al. Prediction of leaf appearance in wheat: a question of temperature. Field Crops Res. 1995, 41;35-44
14. Kirby E. J. M. Analysis of leaf, stem and ear growth in wheat from terminal spikelet to anthesis. Field Crop Res. 1988, 18: 127-140
15. Krenzer E. G., T. L. Nipp, and R. W. McNew. Winter wheat mainstem leaf appearance and tiller formation vs. Moisture treatment. Agron. J. 1991, 83: 663-667
16. McMaster G S, J A Morgan, and W W Wilhelm. Simulating winter wheat spike development and growth. Agric. and Forest Meteor. 1992, 60: 193-220
17. Slafer G A, and H M Rawson. Sensitivity of wheat phasic development to major environmental factors: A re-examination of some assumptions made by physiologists and modellers. Aust J Plant Physiol, 1994, 21: 293-426
2.李存东,曹卫星.小麦阶段发育的生理生态特征评述.南京农业大学学报.1997,20(2):17-21
3.张锦熙等.小麦“叶龄指标促控法”的研究.中国农业科学,1981(2):1—13
4.张锦熙,刘锡山,阎润涛.小麦冬春品种类型及各生育阶段主茎叶数与穗分化进程变异规律的研究.中国农业科学,1986(2):27—35
5.凌启鸿等.叶龄余数在稻穗分化进程鉴定中的应用价值 中国农业科学,1980(4):1—11
6.苗果园等.小麦品种温光效应与主茎叶数的关系.作物学报,1992,18(5):321-329
7.曹广才等.小麦主茎总叶数的变异.作物学报.1990,16(1):73—82
8.曹卫星,江海东.小麦温光反应与发育进程的模拟.南京农业大学学报,1996,19(1):9-16
9. Cao W., and D. N. Moss. Temperature and daylength interaction on phyllochron in wheat and barley. Crop Sci. 1989, 29: 1046-1048
10. Cao W., and D. N. Moss. Sensitivity of winter wheat phyllochron to environmental changes. Agron. J. 1994, 86: 63-66
11. Cao W., and D. N. Moss. Modeling phasic development in wheat: a conceptual integration of physiological components. Journal of Agric. Sci. 1997, 129, 163-172
12. Jame Y. W. et al. Modeling Follar stages of two contrasting spring wheat varieties growth on the Canadian prairies. Modeling for crop-climate-soil-pest system and its applications in sustainable crop production. Published by JAAS. Nanjing, China. 1998, 53-60
13. Jamieson P. D. et al. Prediction of leaf appearance in wheat: a question of temperature. Field Crops Res. 1995, 41;35-44
14. Kirby E. J. M. Analysis of leaf, stem and ear growth in wheat from terminal spikelet to anthesis. Field Crop Res. 1988, 18: 127-140
15. Krenzer E. G., T. L. Nipp, and R. W. McNew. Winter wheat mainstem leaf appearance and tiller formation vs. Moisture treatment. Agron. J. 1991, 83: 663-667
16. McMaster G S, J A Morgan, and W W Wilhelm. Simulating winter wheat spike development and growth. Agric. and Forest Meteor. 1992, 60: 193-220
17. Slafer G A, and H M Rawson. Sensitivity of wheat phasic development to major environmental factors: A re-examination of some assumptions made by physiologists and modellers. Aust J Plant Physiol, 1994, 21: 293-426