小麦产量和氮素吸收利用特性的改良特征及生理基础
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摘要
小麦是世界上最重要的粮食作物之一,提高小麦产量潜力和氮素吸收利用效率是解决全球粮食安全和保护生态环境的重要手段。品种改良是提高小麦产量和氮素吸收利用效率的重要措施之一,明确小麦品种产量和氮素吸收利用特性改良的规律及其生理基础对于阐释小麦产量形成机理和高效利用氮素机制及确定未来小麦高产高效育种目标均具有重要的意义。本研究选用20世纪50年代以来长江中下游冬麦区不同年代育成或引进的32个具有广泛代表性的小麦品种,采用大田试验在3个施氮处理(OkgNha-1,112.5kgNha-1和225kgNha-1)下研究了小麦品种改良过程中产量、主要农艺性状和氮素吸收利用特性的变化规律及生理基础。并在前期研究的基础上选用5个具有广泛代表性的小麦品种,采用大田和盆栽试验进一步探讨了不同年代小麦品种根系形态与生理特性、植株衰老特性和光合特性的差异及其与产量和氮素吸收利用的关系。主要研究结果如下：
1小麦产量和主要农艺性状的改良特征及生理基础
小麦籽粒产量随品种改良进程逐步提高,产量的演变特征在不同施氮条件下差异显著。产量年遗传进展在N225处理(53.8kgha-1yr-1)显著大于N112.5(43.7kgha-1yr-1)和NO (32.4kg ha-1yr-1)处理。小麦品种改良过程中穗粒数、千粒重和收获指数显著增加,而单位面积穗数从50年代至80年代显著下降,随后保持稳定。现代小麦品种对氮肥的响应增强,增施氮肥显著提高了小麦产量、单位面积穗数、穗粒数,但降低了千粒重；提高了开花期叶面积、叶面积指数和单穗产量,但降低了粒叶比,说明氮肥对优化小麦源库关系和提高产量具有重要的作用。同时也说明品种的氮肥效应在品种改良评价过程中应加以考虑。小麦品种改良过程中产量提高主要来源于以下几个方面：
(1)单穗产量和收获指数的提高：在品种改良过程中单穗产量和收获指数显著增加,而生物产量在60年代降低,随后没有明显的变化。说明产量的增加主要来源于单穗产量和收获指数的增加而与生物产量没有明显的关系。比较收获指数和产量构成因素对产量的贡献发现,收获指数增加对产量贡献最大,在高氮处理千粒重对籽粒的贡献大于穗粒数,在低氮处理穗粒数大于千粒重,总体来看穗粒数和千粒重对产量的贡献几乎相同。穗粒数的增加来源于小穗结实率的提高,而与小穗数没有明显的关系。千粒重的提高来源于化后物质积累的增加和灌浆速率的提高。
(2)植株形态的改良：小麦品种改良过程中株高从50年代至80年代显著降低,80年代以后保持相对稳定,基部节间在株高中的比例下降,而穗和穗下茎的比例升高,同时穗下颈在穗下节间的比例下降,说明旗叶和穗的相对位置升高。同时随着品种改良小麦叶片增大、叶面积指数提高、粒叶比增加。说明品种改良改善了植株形态、优化了源库关系,有利于提高抗倒伏能力和优化穗和旗叶的受光姿态,为小麦光合能力的提高奠定了基础。
(3)叶片光合功能的改善：随着小麦品种改良小麦功能叶叶面积、叶面积持续期(LAD)、叶绿素含量、光合速率和光合高值持续期(PAD)显著提高,为产量提高提供了物质和能量来源。现代小麦品种实际光量子效率(ΦPSII)和最大光能转化效率(Fv/Fm)的下降速率较慢,表明品种改良提高了小麦花后光能利用能力和防御光抑制的能力,延缓了光合系统的衰老,是小麦产量提高的生理基础。
(4)植株抗氧化能力的增强：小麦旗叶SOD.CAT和POD活性随品种育成年代显著提高,MDA含量显著降低,进一步表明品种改良通过提高小麦的活性氧清除能力,从而维持和延长LAD和PAD,为提高光合能力和获得高产奠定了生理基础。
(5)灌浆速率的提高：现代小麦品种籽粒灌浆速率显著提高,最大灌浆时期提前,对提高籽粒产量和抵御后期灾害具有重要的意义。早期品种籽粒灌浆期库源比显著高于现代品种,说明品种改良降低了籽粒灌浆过程中源的限制性,使源库关系更加平衡。
(6)物质积累转运的增加：品种改良过程中小麦拔节前和开花后物质积累量、群体生长速率显著提高,而拔节至开花阶段显著降低。花前干物质转运量、转运率和贡献率均随年代推进显著提高。说明品种改良协调了小麦不同生育阶段物质积累,平衡了花前和花后干物质对籽粒的贡献。品种改良降低了功能叶比叶重,提高了功能叶物质转运量和物质输出能力,表明现代品种资源获取能力和生产力增强,为产量的增加奠定了物质基础。
综上：适当降低并稳定单位面积穗数和生物产量,提高单穗产量和收获指数是品种改良的重要特征。提高拔节前营养生长、增加花后干物质积累和花前物质转运是小麦产量改良的生物学基础。稳定叶面积指数,提高花后光合能力,在增加花后物质积累的基础上提高花前物质转运从而提高单穗产量是今后高产育种的重要目标。
2小麦氮素吸收利用特性的改良特征及生理基础.
品种改良过程中小麦氮素吸收利用效率逐步提高,主要由于以下形态和生理过程的变化引起的：
(1)氮素积累转运特性的变化：品种改良过程中小麦旗叶可溶性蛋白含量、硝酸还原酶(NR)活性和谷氨酰胺合成酶(GS)活性显著提高,说明品种改良提高了小麦植株氮同化能力,为氮积累量的增加和氮效率的提高奠定了生理基础。氮效率的提高与花前氮素积累转运特性的改良关系密切。品种改良过程中小麦花前氮积累量、各器官的氮转运能力和拔节至开花阶段积累率显著提高,氮肥吸收效率和氮肥农学效率与开花前氮积累量和氮转运量显著正相关,叶片的转运量和转运率提高对氮效率的贡献最大。因此,增加花前氮积累量和拔节至开花阶段氮积累比例,提高营养器官氮素的转运能力是品种氮效率提高的重要原因。
(2)根系形态生理的变化：品种改良提高了小麦拔节至开花阶段根系干物质积累量、生长速率和花后根系干物质转运量；增加了根系总根长、根系表面积、根体积、060cm各土层根重密度和上层根的分布比例。现代小麦品种根系活力显著提高,同时根系可溶性蛋白含量、SOD活性显著提高,MDA含量显著降低,说明品种改良改善了根系形态、延缓了根系的衰老,为产量和氮素吸收利用效率的提高奠定了生理基础。
(3)光合面积和光合能力的提高：叶面积和光合能力提高与氮吸收利用效率关系密切。功能叶叶面积和叶面积持续期(LAD)与氮素籽粒生产效率(NYE)、氮肥吸收效率(NUpE)、氮素生理效率(NPE)显著正相关,最大光合速率与NUpE、NPE显著正相关,单位氮叶面积持续期(LADN)和单位氮光合持续期(PADN)与NFY和NHI显著正相关,与NUpE显著负相关。说明光合面积和光合能力的改良能拉动植株对氮的吸收,提高LADN和PADN可增加营养器官氮素向籽粒的转运,同时也说明LADN和PADN可作为氮转运和氮效率改良的评价指标。
综上所述,品种改良通过提高花前NR、GS活性提高了植株的氮同化能力,从而增加了花前氮素积累量和拔节至开花阶段氮积累率；通过提高光合面积、光合能力、根系总根长、根系吸收面积、根系活力和根系氮同化能力提高了植株氮素吸收能力；通过增加氮素在叶片中的分配,提高了花前氮转运量。因此,提高NR、GS舌性和叶片光合能力、提高根系吸收面积和根系活力、增加开花期氮素在叶片中的分配是品种氮效率改良的生理基础,也是未来氮高效品种选育的重要目标。
Wheat is an important food crop worldwide. Genetic improvements have contributed much to wheat production since the1960s. Verifying the evolution of agronomic traits and the physiological basis of grain yield will facilitate breeders and agronomists in developing new wheat cultivars, with the aim of stable and high yields. Thirty-three wheat cultivars, bred or widely planted in the Yangtze River Basin from1950to2005, were grown in field experiments under three N rates (0,112.5, and225kg N ha-1) from2007to2011in Nanjing, China. And grain yield, main agronomic traits and physiological basis of nitrogen uptake during the wheat cultivars genetic improvement process were mensurated. Thereafter five represent culitvars were selected to further explore the relationship among root morphology and physiology, plant aging properties, properties of source and sink and nitrogen uptake use relation and the response of nitrogen fertilizer by field and pot experiment. The main results were summarized as follows:
1Characteristics of genentic improvements on grain yield and agronomic traits of winter wheat and their physiological basis
Wheat grain yield increased gradually with the genetic improvement process, the differences of the evolution characteristics on grain yield under different nitrogen rates were significant. Grain yield also increased with increased N fertilization rate; the annual increment was higher at225N (53.8kg ha-1year-1) than at112.5N (43.7kg ha-1year-1) and ON (32.4kg ha-1year-1), indicating that modern improved cultivars were more sensitive to increased N fertilizer application. Kernel number per spike and TKW increased linearly with genetic development of cultivars. Leaf area, LAI, and Pn increased significantly with increasing nitrogen fertilizer. Grain yield, spikes per unit land, and kernels per spike increased significantly with increasing nitrogen fertilizer, but TKW and HI decreased, indicating that nitrogen plays an important role to improve the grain yield and optimize the source-sink relationships of wheat. The grain yields increases were mainly related to the aspects as the follows:
(1) The increase of spike yield and harvest index. Spike yield and HI increased linearly with cultivar development from1950s to2000s. Biomass decreased significantly from1950s to1960s but did not changed significantly in cultivars developed since the1960s. This indicates that the improved grain yield of the wheat cultivars was mainly related to improve spike yield and HI, not biomass. The contribution of HI was higher than kernels and TKW to grain yield. The contribution of TKW to grain yield was higher than kernels under high nitrogen rates (N225), but lower at N112.5and NO. In the whole, the kernels and TKW almost had the same contribution to grain yield. The grain number increase was mainly related to the increase of spikelet fertility, but not spikelet number, and the grain weight increase was mainly related to dry matter accumulation and grain filling rate.
(2) The improvement of plant type. Cultivar height decreased with cultivar development from the1950s to the1980s, and remained relatively stable in subsequent cultivars. The proportion of the length of the top internode to total plant height increased with cultivar development from the1950s to the1980s and thereafter fell, while the length of the basal internode (BI) maintained a shortening trend. The proportion of the length of the spike to whole plant length and the ratio of neck-panicle node (NPN) length to top internode (T1) length decreased with cultivar development, indicating that the relative height of the spike and flag leaf increased, improving the position and light interception of the spike and flag leaf.
(3) The improvement of photo synthetic function. Leaf area per culm, leaf area index (LAI) chlorophyll content, Pn and photosynthetic activity duration (PAD) increased with cultivar development, and the ΦPSII and Fv/Fm declining rate of modern cultivars were slower than earlier cultivars, the results showed that cultivars improvement increased the light energy use ability at after-anthesis and defense photoinhibition ability, delaying the aging of photosynthetic system, is the physiological basis of wheat source enhanced, which was the physiological basis of wheat source imcrease.
(4) The improvement of plant antioxidant capacity. SOD, CAT and POD activity of flag leaf were improved significantly with the cultivars genetic progress, and the MDA content decreased significantly, which further showed that cultivars improvement improved the clearance ability of O2-, in order to maintain and extend the LAD and PAD, to establish the physiological basis for high yield.
(5) The improvement of the grain filling rate. The grain filling rate of modern wheat cultivars increased significantly, the maximum grain filling period in advance of great significance to improve grain yield and to resist the post-disaster. Which were higher significantly than the early cultivars of the grain filling stage of library source than modern cultivars, nitrogen application significantly reduces the source-sink ratio, indicating that the improved cultivars to reduce the grain filling restrictive, so that a more balanced source-sink relationship.
(6) The increase of dry matter accumulation and translocation. Genetic improvement increased the dry matter accumulation and growth rate in the growth stage of emergence to jointing and anthesis to maturity whereas reduced contribution of post-anthesis accumulated dry matter to grains (CPA). Grain yield was significantly positive associated with contribution of pre-anthesis translocation to grains (CPT), dry matter accumulation and growth rate from emergence to jointing and anthesis to maturity, whereas significantly negative with CPA, dry matter accumulation and growth rate from jointing to anthesis. Dry matter production capacity and production efficiency were improved during wheat cultivar improvement, yet dry matter accumulation at different growth stage were coordinated and the contribution of pre-anthesis and post-anthesis accumulation to grains became more balance.
In conclusion, breeding for high yield should be related to improvement in kernels per spike and TKW per unit land and increased sink-source ratios with a feasible LAI, and N fertilizer management should be considered during breeding for higher yields. Improving the pre-jointing vegetative growth post-anthesis accumulation and CPT are important physiological basis for wheat grain yield enhancement, which will be a key improvement target for high yield wheat breeding in the future.
2Characteristics of genentic improvements on nitrogen uptke and utilization of winter wheat and their physiological basis
The nitrogen uptake and utilization efficiency increased gradually during the cultivars genetic imprvement, and the increase of nitrogen utilization efficiency were mainly due to the improvement of the morphological and physiological process as the following:
(1) The improvement of N accumulation and translocation. Soluble protein content, nitrate reductase (NR) and glutamine synthase (GS) activity of wheat flag leaf increased significantly with cultivars genetic progress, indicating that cultivars genetic improvement improved the assimilation nitrogen and ammonium nitrate ability of the plant. The pre-anthesis nitrogen accumulation, nitrogen translocation of each organs increased significantly, nitrogen residual of plant vegetative organs in maturation decrease and grain nitrogen accumulation were improved, which showed that the improvement of nitrogen efficiency were closely related to pre-anthesis nitrogen accumulation, distribution and translocation enhancement. The nitrogen accumulation of different wheat cultivars were jointing to anthesis stage in peak, along with aging propulsion, wheat flowering stage to jointing stage accumulation rate significantly increased. Modern cultivars leaf nitrogen distribution and translocation rate increased significantly. Therefore, increasing the pre-anthesis nitrogen accumulation and improving nitrogen distribution in leaves are the important characteristics of nitrogen efficiency improved, also the important breeding nitrogen efficiency index.
(2) The improvement of root type and root physiology. The root dry matter accumulation, growth rate and after-anthesis dry matter translocation from jointing to flowering stage were improved during the cultivars genetic improvement. And the total root length, root surface area and root volume have gradually improvement with cultivar genetic progress, wheat cultivar improvement enhanced distribution proportion about the root weight of0to60cm soil and root density in0-20cm of the soil. Along with the incubation of cultivars genetic progress, wheat root soluble protein content, SOD activity improved significantly and significantly reduce the MDA content, root activity was improved significantly explaining that genetics improvement delayed the root of aging, to improve yield and nitrogen uptake efficiency. Nitrogen efficiency was significantly and positively associated with total root length, root surface area, root volume, NR, GS and SOD activity, was significantly and negatively associated with root MDA content at flowering time, indicating that increasing total root length, and root surface root volume, improving NR, GS and SOD activity, reducing the MDA content played an important role on the nitrogen efficiency.
(3) The improvement of photosynthetic area and photosynthetic ability. The leaf area duration per unit nitrogen (LADN) and photosynthetic activity duration per unit nitrogen (PADN) increased linearly with cultivars genetic progress, and there were different expression in different nitrogen treatment under the different leaves, showed that the enhanced source that could pull plant nitrogen-uptake, meanwhile that nitrogen fertilizer plays an important regulated role of the source-sink balance and nitrogen uptake. The maximum effective leaf area was significantly and positively associated with nitrogen grain yield efficiency (NYE), nitrogen uptake efficiency (NUpE), nitrogen physiological efficiency (NPE), the biggest photosynthetic rate was significantly and positively associated with NRE and NPE, PAD was significantly and positively associated with NYE and NPE, yield per-spike was significantly and positively associated with NYE, NAE, NUpE and NPE. LADN and PADN was significantly and positively associated with NFY and NH, while was significantly and negatively associated with NUpE. Indicating that improve LADN and PADN could improve nitrogen translocation from vegetative organs to grain, also explaining LADN and PADN could be used as nitrogen translocation and nitrogen efficiency evaluation index.
In conclusion, cultivars improvement increased NR and GS activity of flag leaf at pre-antheshis to improve nitrogen assimilation ability of the plant, thereby increasd the amount of nitrogen accumulation and nitrogen accumulation rato of jointing to anthesis stage; increased the photosynthetic area, photosynthetic capacity, root, total root length, root absorption area, root activity and root nitrogen assimilation improved the nitrogen uptake capacity; increasing the distribution of nitrogen in the leaves, the nitrogen translocation of pre-anthesis. Therefore, the improvement of the NR GS activity and leaf photosynthetic capacity, root absorption area and root activity, to increase the distribution of nitrogen in the leaves are the important physiological basis for wheat N efficiency enhancement, which will be a key improvement target for high N efficiency wheat breeding in the future.
1小麦产量和主要农艺性状的改良特征及生理基础
小麦籽粒产量随品种改良进程逐步提高,产量的演变特征在不同施氮条件下差异显著。产量年遗传进展在N225处理(53.8kgha-1yr-1)显著大于N112.5(43.7kgha-1yr-1)和NO (32.4kg ha-1yr-1)处理。小麦品种改良过程中穗粒数、千粒重和收获指数显著增加,而单位面积穗数从50年代至80年代显著下降,随后保持稳定。现代小麦品种对氮肥的响应增强,增施氮肥显著提高了小麦产量、单位面积穗数、穗粒数,但降低了千粒重；提高了开花期叶面积、叶面积指数和单穗产量,但降低了粒叶比,说明氮肥对优化小麦源库关系和提高产量具有重要的作用。同时也说明品种的氮肥效应在品种改良评价过程中应加以考虑。小麦品种改良过程中产量提高主要来源于以下几个方面：
(1)单穗产量和收获指数的提高：在品种改良过程中单穗产量和收获指数显著增加,而生物产量在60年代降低,随后没有明显的变化。说明产量的增加主要来源于单穗产量和收获指数的增加而与生物产量没有明显的关系。比较收获指数和产量构成因素对产量的贡献发现,收获指数增加对产量贡献最大,在高氮处理千粒重对籽粒的贡献大于穗粒数,在低氮处理穗粒数大于千粒重,总体来看穗粒数和千粒重对产量的贡献几乎相同。穗粒数的增加来源于小穗结实率的提高,而与小穗数没有明显的关系。千粒重的提高来源于化后物质积累的增加和灌浆速率的提高。
(2)植株形态的改良：小麦品种改良过程中株高从50年代至80年代显著降低,80年代以后保持相对稳定,基部节间在株高中的比例下降,而穗和穗下茎的比例升高,同时穗下颈在穗下节间的比例下降,说明旗叶和穗的相对位置升高。同时随着品种改良小麦叶片增大、叶面积指数提高、粒叶比增加。说明品种改良改善了植株形态、优化了源库关系,有利于提高抗倒伏能力和优化穗和旗叶的受光姿态,为小麦光合能力的提高奠定了基础。
(3)叶片光合功能的改善：随着小麦品种改良小麦功能叶叶面积、叶面积持续期(LAD)、叶绿素含量、光合速率和光合高值持续期(PAD)显著提高,为产量提高提供了物质和能量来源。现代小麦品种实际光量子效率(ΦPSII)和最大光能转化效率(Fv/Fm)的下降速率较慢,表明品种改良提高了小麦花后光能利用能力和防御光抑制的能力,延缓了光合系统的衰老,是小麦产量提高的生理基础。
(4)植株抗氧化能力的增强：小麦旗叶SOD.CAT和POD活性随品种育成年代显著提高,MDA含量显著降低,进一步表明品种改良通过提高小麦的活性氧清除能力,从而维持和延长LAD和PAD,为提高光合能力和获得高产奠定了生理基础。
(5)灌浆速率的提高：现代小麦品种籽粒灌浆速率显著提高,最大灌浆时期提前,对提高籽粒产量和抵御后期灾害具有重要的意义。早期品种籽粒灌浆期库源比显著高于现代品种,说明品种改良降低了籽粒灌浆过程中源的限制性,使源库关系更加平衡。
(6)物质积累转运的增加：品种改良过程中小麦拔节前和开花后物质积累量、群体生长速率显著提高,而拔节至开花阶段显著降低。花前干物质转运量、转运率和贡献率均随年代推进显著提高。说明品种改良协调了小麦不同生育阶段物质积累,平衡了花前和花后干物质对籽粒的贡献。品种改良降低了功能叶比叶重,提高了功能叶物质转运量和物质输出能力,表明现代品种资源获取能力和生产力增强,为产量的增加奠定了物质基础。
综上：适当降低并稳定单位面积穗数和生物产量,提高单穗产量和收获指数是品种改良的重要特征。提高拔节前营养生长、增加花后干物质积累和花前物质转运是小麦产量改良的生物学基础。稳定叶面积指数,提高花后光合能力,在增加花后物质积累的基础上提高花前物质转运从而提高单穗产量是今后高产育种的重要目标。
2小麦氮素吸收利用特性的改良特征及生理基础.
品种改良过程中小麦氮素吸收利用效率逐步提高,主要由于以下形态和生理过程的变化引起的：
(1)氮素积累转运特性的变化：品种改良过程中小麦旗叶可溶性蛋白含量、硝酸还原酶(NR)活性和谷氨酰胺合成酶(GS)活性显著提高,说明品种改良提高了小麦植株氮同化能力,为氮积累量的增加和氮效率的提高奠定了生理基础。氮效率的提高与花前氮素积累转运特性的改良关系密切。品种改良过程中小麦花前氮积累量、各器官的氮转运能力和拔节至开花阶段积累率显著提高,氮肥吸收效率和氮肥农学效率与开花前氮积累量和氮转运量显著正相关,叶片的转运量和转运率提高对氮效率的贡献最大。因此,增加花前氮积累量和拔节至开花阶段氮积累比例,提高营养器官氮素的转运能力是品种氮效率提高的重要原因。
(2)根系形态生理的变化：品种改良提高了小麦拔节至开花阶段根系干物质积累量、生长速率和花后根系干物质转运量；增加了根系总根长、根系表面积、根体积、060cm各土层根重密度和上层根的分布比例。现代小麦品种根系活力显著提高,同时根系可溶性蛋白含量、SOD活性显著提高,MDA含量显著降低,说明品种改良改善了根系形态、延缓了根系的衰老,为产量和氮素吸收利用效率的提高奠定了生理基础。
(3)光合面积和光合能力的提高：叶面积和光合能力提高与氮吸收利用效率关系密切。功能叶叶面积和叶面积持续期(LAD)与氮素籽粒生产效率(NYE)、氮肥吸收效率(NUpE)、氮素生理效率(NPE)显著正相关,最大光合速率与NUpE、NPE显著正相关,单位氮叶面积持续期(LADN)和单位氮光合持续期(PADN)与NFY和NHI显著正相关,与NUpE显著负相关。说明光合面积和光合能力的改良能拉动植株对氮的吸收,提高LADN和PADN可增加营养器官氮素向籽粒的转运,同时也说明LADN和PADN可作为氮转运和氮效率改良的评价指标。
综上所述,品种改良通过提高花前NR、GS活性提高了植株的氮同化能力,从而增加了花前氮素积累量和拔节至开花阶段氮积累率；通过提高光合面积、光合能力、根系总根长、根系吸收面积、根系活力和根系氮同化能力提高了植株氮素吸收能力；通过增加氮素在叶片中的分配,提高了花前氮转运量。因此,提高NR、GS舌性和叶片光合能力、提高根系吸收面积和根系活力、增加开花期氮素在叶片中的分配是品种氮效率改良的生理基础,也是未来氮高效品种选育的重要目标。
Wheat is an important food crop worldwide. Genetic improvements have contributed much to wheat production since the1960s. Verifying the evolution of agronomic traits and the physiological basis of grain yield will facilitate breeders and agronomists in developing new wheat cultivars, with the aim of stable and high yields. Thirty-three wheat cultivars, bred or widely planted in the Yangtze River Basin from1950to2005, were grown in field experiments under three N rates (0,112.5, and225kg N ha-1) from2007to2011in Nanjing, China. And grain yield, main agronomic traits and physiological basis of nitrogen uptake during the wheat cultivars genetic improvement process were mensurated. Thereafter five represent culitvars were selected to further explore the relationship among root morphology and physiology, plant aging properties, properties of source and sink and nitrogen uptake use relation and the response of nitrogen fertilizer by field and pot experiment. The main results were summarized as follows:
1Characteristics of genentic improvements on grain yield and agronomic traits of winter wheat and their physiological basis
Wheat grain yield increased gradually with the genetic improvement process, the differences of the evolution characteristics on grain yield under different nitrogen rates were significant. Grain yield also increased with increased N fertilization rate; the annual increment was higher at225N (53.8kg ha-1year-1) than at112.5N (43.7kg ha-1year-1) and ON (32.4kg ha-1year-1), indicating that modern improved cultivars were more sensitive to increased N fertilizer application. Kernel number per spike and TKW increased linearly with genetic development of cultivars. Leaf area, LAI, and Pn increased significantly with increasing nitrogen fertilizer. Grain yield, spikes per unit land, and kernels per spike increased significantly with increasing nitrogen fertilizer, but TKW and HI decreased, indicating that nitrogen plays an important role to improve the grain yield and optimize the source-sink relationships of wheat. The grain yields increases were mainly related to the aspects as the follows:
(1) The increase of spike yield and harvest index. Spike yield and HI increased linearly with cultivar development from1950s to2000s. Biomass decreased significantly from1950s to1960s but did not changed significantly in cultivars developed since the1960s. This indicates that the improved grain yield of the wheat cultivars was mainly related to improve spike yield and HI, not biomass. The contribution of HI was higher than kernels and TKW to grain yield. The contribution of TKW to grain yield was higher than kernels under high nitrogen rates (N225), but lower at N112.5and NO. In the whole, the kernels and TKW almost had the same contribution to grain yield. The grain number increase was mainly related to the increase of spikelet fertility, but not spikelet number, and the grain weight increase was mainly related to dry matter accumulation and grain filling rate.
(2) The improvement of plant type. Cultivar height decreased with cultivar development from the1950s to the1980s, and remained relatively stable in subsequent cultivars. The proportion of the length of the top internode to total plant height increased with cultivar development from the1950s to the1980s and thereafter fell, while the length of the basal internode (BI) maintained a shortening trend. The proportion of the length of the spike to whole plant length and the ratio of neck-panicle node (NPN) length to top internode (T1) length decreased with cultivar development, indicating that the relative height of the spike and flag leaf increased, improving the position and light interception of the spike and flag leaf.
(3) The improvement of photo synthetic function. Leaf area per culm, leaf area index (LAI) chlorophyll content, Pn and photosynthetic activity duration (PAD) increased with cultivar development, and the ΦPSII and Fv/Fm declining rate of modern cultivars were slower than earlier cultivars, the results showed that cultivars improvement increased the light energy use ability at after-anthesis and defense photoinhibition ability, delaying the aging of photosynthetic system, is the physiological basis of wheat source enhanced, which was the physiological basis of wheat source imcrease.
(4) The improvement of plant antioxidant capacity. SOD, CAT and POD activity of flag leaf were improved significantly with the cultivars genetic progress, and the MDA content decreased significantly, which further showed that cultivars improvement improved the clearance ability of O2-, in order to maintain and extend the LAD and PAD, to establish the physiological basis for high yield.
(5) The improvement of the grain filling rate. The grain filling rate of modern wheat cultivars increased significantly, the maximum grain filling period in advance of great significance to improve grain yield and to resist the post-disaster. Which were higher significantly than the early cultivars of the grain filling stage of library source than modern cultivars, nitrogen application significantly reduces the source-sink ratio, indicating that the improved cultivars to reduce the grain filling restrictive, so that a more balanced source-sink relationship.
(6) The increase of dry matter accumulation and translocation. Genetic improvement increased the dry matter accumulation and growth rate in the growth stage of emergence to jointing and anthesis to maturity whereas reduced contribution of post-anthesis accumulated dry matter to grains (CPA). Grain yield was significantly positive associated with contribution of pre-anthesis translocation to grains (CPT), dry matter accumulation and growth rate from emergence to jointing and anthesis to maturity, whereas significantly negative with CPA, dry matter accumulation and growth rate from jointing to anthesis. Dry matter production capacity and production efficiency were improved during wheat cultivar improvement, yet dry matter accumulation at different growth stage were coordinated and the contribution of pre-anthesis and post-anthesis accumulation to grains became more balance.
In conclusion, breeding for high yield should be related to improvement in kernels per spike and TKW per unit land and increased sink-source ratios with a feasible LAI, and N fertilizer management should be considered during breeding for higher yields. Improving the pre-jointing vegetative growth post-anthesis accumulation and CPT are important physiological basis for wheat grain yield enhancement, which will be a key improvement target for high yield wheat breeding in the future.
2Characteristics of genentic improvements on nitrogen uptke and utilization of winter wheat and their physiological basis
The nitrogen uptake and utilization efficiency increased gradually during the cultivars genetic imprvement, and the increase of nitrogen utilization efficiency were mainly due to the improvement of the morphological and physiological process as the following:
(1) The improvement of N accumulation and translocation. Soluble protein content, nitrate reductase (NR) and glutamine synthase (GS) activity of wheat flag leaf increased significantly with cultivars genetic progress, indicating that cultivars genetic improvement improved the assimilation nitrogen and ammonium nitrate ability of the plant. The pre-anthesis nitrogen accumulation, nitrogen translocation of each organs increased significantly, nitrogen residual of plant vegetative organs in maturation decrease and grain nitrogen accumulation were improved, which showed that the improvement of nitrogen efficiency were closely related to pre-anthesis nitrogen accumulation, distribution and translocation enhancement. The nitrogen accumulation of different wheat cultivars were jointing to anthesis stage in peak, along with aging propulsion, wheat flowering stage to jointing stage accumulation rate significantly increased. Modern cultivars leaf nitrogen distribution and translocation rate increased significantly. Therefore, increasing the pre-anthesis nitrogen accumulation and improving nitrogen distribution in leaves are the important characteristics of nitrogen efficiency improved, also the important breeding nitrogen efficiency index.
(2) The improvement of root type and root physiology. The root dry matter accumulation, growth rate and after-anthesis dry matter translocation from jointing to flowering stage were improved during the cultivars genetic improvement. And the total root length, root surface area and root volume have gradually improvement with cultivar genetic progress, wheat cultivar improvement enhanced distribution proportion about the root weight of0to60cm soil and root density in0-20cm of the soil. Along with the incubation of cultivars genetic progress, wheat root soluble protein content, SOD activity improved significantly and significantly reduce the MDA content, root activity was improved significantly explaining that genetics improvement delayed the root of aging, to improve yield and nitrogen uptake efficiency. Nitrogen efficiency was significantly and positively associated with total root length, root surface area, root volume, NR, GS and SOD activity, was significantly and negatively associated with root MDA content at flowering time, indicating that increasing total root length, and root surface root volume, improving NR, GS and SOD activity, reducing the MDA content played an important role on the nitrogen efficiency.
(3) The improvement of photosynthetic area and photosynthetic ability. The leaf area duration per unit nitrogen (LADN) and photosynthetic activity duration per unit nitrogen (PADN) increased linearly with cultivars genetic progress, and there were different expression in different nitrogen treatment under the different leaves, showed that the enhanced source that could pull plant nitrogen-uptake, meanwhile that nitrogen fertilizer plays an important regulated role of the source-sink balance and nitrogen uptake. The maximum effective leaf area was significantly and positively associated with nitrogen grain yield efficiency (NYE), nitrogen uptake efficiency (NUpE), nitrogen physiological efficiency (NPE), the biggest photosynthetic rate was significantly and positively associated with NRE and NPE, PAD was significantly and positively associated with NYE and NPE, yield per-spike was significantly and positively associated with NYE, NAE, NUpE and NPE. LADN and PADN was significantly and positively associated with NFY and NH, while was significantly and negatively associated with NUpE. Indicating that improve LADN and PADN could improve nitrogen translocation from vegetative organs to grain, also explaining LADN and PADN could be used as nitrogen translocation and nitrogen efficiency evaluation index.
In conclusion, cultivars improvement increased NR and GS activity of flag leaf at pre-antheshis to improve nitrogen assimilation ability of the plant, thereby increasd the amount of nitrogen accumulation and nitrogen accumulation rato of jointing to anthesis stage; increased the photosynthetic area, photosynthetic capacity, root, total root length, root absorption area, root activity and root nitrogen assimilation improved the nitrogen uptake capacity; increasing the distribution of nitrogen in the leaves, the nitrogen translocation of pre-anthesis. Therefore, the improvement of the NR GS activity and leaf photosynthetic capacity, root absorption area and root activity, to increase the distribution of nitrogen in the leaves are the important physiological basis for wheat N efficiency enhancement, which will be a key improvement target for high N efficiency wheat breeding in the future.
引文
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