玉米氮高效品种选育及根系形态对低氮反应的遗传分析
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摘要
农田氮肥大量投入易造成环境污染、经济效益低及氮肥利用率下降。选育低氮下得到较高产量的氮高效品种是缓解这一问题的重要途径之一。有关氮效率基因差异的报道较多，但氮高效育种实践较少。根系在氮素吸收中起重要作用，对田间全生育期中的动态研究较少，对根系性状的遗传研究相对不足。在本课题组前期氮高效基因型筛选工作的基础上，本研究开展了氮高效杂交种选育工作，利用不同氮效率杂交种在田间研究根系与氮吸收及氮效率的关系。同时，利用有根系显著差异的自交系及其组合，在温室进行砂培试验，研究玉米根系对低氮胁迫的反应基因型差异及其遗传规律，以评价根系在氮高效育种中的重要性及可行性。
通过两个氮水平下的多年多点田间选育，以农大108为对照，得到了一个绿熟型双高效的杂交组合NE1。通过连续4年多点试验，无论在低氮条件下(不施氮肥)，还是在高氮条件下，增产幅度在5～11％左右，显著提高了玉米新品种的耐低氮能力和氮肥利用率，为我国玉米氮营养高效育种开辟了一条切实可行的途径。同时获得一些以NE2为代表的氮低效杂交组合，还鉴定出3个在氮高效性状上有高配合力的自交系。
以NE1(高效)、NE2(低效)及另外两个杂交种农大108(ND108，高效)和四单19(SD19，低效)为材料，研究探讨了NE1等氮高效品种的干物质累积、氮素累积、根系特点及不同氮水平对它们的影响。结果表明，在生长前期，氮高效品种的干物质及氮素累积并未表现出优势。氮高效品种高产的主要原因在于其生长后期对氮素的吸收。两个氮高效基因型又各有特点，ND108主要依靠其较高的后期氮素吸收能力；NE1后期的氮吸收也较强，同是具有较高的氮转移率。与此相对应，氮高效品种在根系的生物量及根系长度方面具有一定的优势，而优势主要体现在生长后期。
玉米对缺氮的一致反应为根冠比的增加，但根系性状本身的变化存在显著基因型差异。总体来看，短期低氮导致根重、总根长的增加。根长的增加是以侧根长的增加为主，轴根总长度变化不大，但轴根数减少，平均轴根长增加。杂交组合的根系反应普遍较自交系敏感，根系的调节能力强于自交系，但是具体的反应与其亲本具有很大的关系，亲本间基因互作会使杂交组合的根系对低氮的反应呈现截然不同的变化。
苗期玉米根系对低氮反应的一般配合力、特殊配合力都存在显著的基因型差异，而且不同的基因型在氮胁迫下的反应也不尽相同。氮胁迫可以影响根系的遗传，表现为根系性状的广义遗传力下降，狭义遗传力上升。高氮下根干重、总根长、侧根长和轴根数和茎叶干重等性状以非加性遗传为主。在低氮胁迫下，根干重、总根长、侧根长和轴根长等性状以加性遗传为主。
综合以上研究，在氮高效育种中，应该注重对根系性状的选择，特别是生长后期根系的生物量、根长等性状可以作为田间筛选的参考指标。选择具有较大根重、总根长、侧根长等性状的自交系作为亲本，可能得到这些性状具有优势的杂交种。
Excessive nitrogen (N) fertilizer input in arable land may result in environmental pollutions, low income for farmers and low N use efficiency. Development of N efficient cultivars may be one of the ways to resolve such problems. Genotypic difference in N efficiency has been reported in many references, but breeding for N-efficient cultivar is still rare. Roots are important for N uptake, but few studies on the role of root in N uptake have been conducted in field conditions. And, less is known about how root traits are inherited at N stress. On the basis of selection for N-efficient inbred lines, the current study concentrated on breeding N-efficient maize hybrids. The role of roots in N uptake and yield formation of selected hybrids in the field was investigated in two N levels. Furthermore, response to low N stress of roots of both hybrids and their parent lines was compared to understand the physiological and genetic basis for root response to low N stress and possibility for root as a trait to breed for
N-cfficient hybrids.
Taking ND108 as a check, some N-efficient hybrids were developed among which NE1 is the most efficient one. Some other hybrids were proven to be N-inefficient. Three inbred lines with high combining ability in N efficiency were identified.
Taking NE1 (N-efficient), ND108 (N-efficient), NE1 (N-inefficient) and SD19 (N-inefficient) as materials, biomass accumulation, N accumulation, root dry weight and root length were investigated under two N levels to understand the basis of N efficiency. It was found that N efficient hybrids did not show any advantages in N uptake and biomass accumulation at the early stage before silking. Efficient N uptake after silking was the charateristic of N-efficient hybrids. Between the two N-efficient hybrids, ND108 accumulated large portion of N after silking, with less translocation of pre-silking N into grains. While in NE1, both N uptake after silking and translocation of pre-silking N uptake contributed to its high N accumulation and yield. N-efficient hybrids had higher root biomass and root length, which might be the reason for its efficient N uptake.
Although the large genotypic difference, low N generally suppresses shoot growth, increases root to shoot ratio with or without increasing root biomass of maize. Maize plants respond to N deficiency by increasing total root length and altering root architecture by increasing the elongation of individual axial roots, enhancing lateral root growth, but reducing the number of axial root. Inbreds showed weaker responses in root biomass and other root parameters than their crosses. Interaction between two parent inbreds had strong influence on the response of their cross to N stress.
The genotypic variations of general combining ability (GCA) and specific combining ability (SCA) on root traits were observed under high and low nitrogen levels (HN and LN). N stress exerted an influence on the genetic ability of root traits, with decreasing of broad genetic ability and increasing of narrow genetic ability. The genetic of most root traits such as root weight, total root length and lateral root length were controlled by non-additive effect at HN, but by additive effect at LN.
It is suggested that the root traits at later growth stage might be one of the selection criteria in selection for N-efficient cultivars. A big root system in a cross can be obtained by selecting parents lines
with large root weight, total root length, total length of lateral roots.
通过两个氮水平下的多年多点田间选育，以农大108为对照，得到了一个绿熟型双高效的杂交组合NE1。通过连续4年多点试验，无论在低氮条件下(不施氮肥)，还是在高氮条件下，增产幅度在5～11％左右，显著提高了玉米新品种的耐低氮能力和氮肥利用率，为我国玉米氮营养高效育种开辟了一条切实可行的途径。同时获得一些以NE2为代表的氮低效杂交组合，还鉴定出3个在氮高效性状上有高配合力的自交系。
以NE1(高效)、NE2(低效)及另外两个杂交种农大108(ND108，高效)和四单19(SD19，低效)为材料，研究探讨了NE1等氮高效品种的干物质累积、氮素累积、根系特点及不同氮水平对它们的影响。结果表明，在生长前期，氮高效品种的干物质及氮素累积并未表现出优势。氮高效品种高产的主要原因在于其生长后期对氮素的吸收。两个氮高效基因型又各有特点，ND108主要依靠其较高的后期氮素吸收能力；NE1后期的氮吸收也较强，同是具有较高的氮转移率。与此相对应，氮高效品种在根系的生物量及根系长度方面具有一定的优势，而优势主要体现在生长后期。
玉米对缺氮的一致反应为根冠比的增加，但根系性状本身的变化存在显著基因型差异。总体来看，短期低氮导致根重、总根长的增加。根长的增加是以侧根长的增加为主，轴根总长度变化不大，但轴根数减少，平均轴根长增加。杂交组合的根系反应普遍较自交系敏感，根系的调节能力强于自交系，但是具体的反应与其亲本具有很大的关系，亲本间基因互作会使杂交组合的根系对低氮的反应呈现截然不同的变化。
苗期玉米根系对低氮反应的一般配合力、特殊配合力都存在显著的基因型差异，而且不同的基因型在氮胁迫下的反应也不尽相同。氮胁迫可以影响根系的遗传，表现为根系性状的广义遗传力下降，狭义遗传力上升。高氮下根干重、总根长、侧根长和轴根数和茎叶干重等性状以非加性遗传为主。在低氮胁迫下，根干重、总根长、侧根长和轴根长等性状以加性遗传为主。
综合以上研究，在氮高效育种中，应该注重对根系性状的选择，特别是生长后期根系的生物量、根长等性状可以作为田间筛选的参考指标。选择具有较大根重、总根长、侧根长等性状的自交系作为亲本，可能得到这些性状具有优势的杂交种。
Excessive nitrogen (N) fertilizer input in arable land may result in environmental pollutions, low income for farmers and low N use efficiency. Development of N efficient cultivars may be one of the ways to resolve such problems. Genotypic difference in N efficiency has been reported in many references, but breeding for N-efficient cultivar is still rare. Roots are important for N uptake, but few studies on the role of root in N uptake have been conducted in field conditions. And, less is known about how root traits are inherited at N stress. On the basis of selection for N-efficient inbred lines, the current study concentrated on breeding N-efficient maize hybrids. The role of roots in N uptake and yield formation of selected hybrids in the field was investigated in two N levels. Furthermore, response to low N stress of roots of both hybrids and their parent lines was compared to understand the physiological and genetic basis for root response to low N stress and possibility for root as a trait to breed for
N-cfficient hybrids.
Taking ND108 as a check, some N-efficient hybrids were developed among which NE1 is the most efficient one. Some other hybrids were proven to be N-inefficient. Three inbred lines with high combining ability in N efficiency were identified.
Taking NE1 (N-efficient), ND108 (N-efficient), NE1 (N-inefficient) and SD19 (N-inefficient) as materials, biomass accumulation, N accumulation, root dry weight and root length were investigated under two N levels to understand the basis of N efficiency. It was found that N efficient hybrids did not show any advantages in N uptake and biomass accumulation at the early stage before silking. Efficient N uptake after silking was the charateristic of N-efficient hybrids. Between the two N-efficient hybrids, ND108 accumulated large portion of N after silking, with less translocation of pre-silking N into grains. While in NE1, both N uptake after silking and translocation of pre-silking N uptake contributed to its high N accumulation and yield. N-efficient hybrids had higher root biomass and root length, which might be the reason for its efficient N uptake.
Although the large genotypic difference, low N generally suppresses shoot growth, increases root to shoot ratio with or without increasing root biomass of maize. Maize plants respond to N deficiency by increasing total root length and altering root architecture by increasing the elongation of individual axial roots, enhancing lateral root growth, but reducing the number of axial root. Inbreds showed weaker responses in root biomass and other root parameters than their crosses. Interaction between two parent inbreds had strong influence on the response of their cross to N stress.
The genotypic variations of general combining ability (GCA) and specific combining ability (SCA) on root traits were observed under high and low nitrogen levels (HN and LN). N stress exerted an influence on the genetic ability of root traits, with decreasing of broad genetic ability and increasing of narrow genetic ability. The genetic of most root traits such as root weight, total root length and lateral root length were controlled by non-additive effect at HN, but by additive effect at LN.
It is suggested that the root traits at later growth stage might be one of the selection criteria in selection for N-efficient cultivars. A big root system in a cross can be obtained by selecting parents lines
with large root weight, total root length, total length of lateral roots.
引文
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