摘要
我国稻田施氮量大,氮肥利用率低。过大的施氮量和较低的氮肥利用率,增加了氮肥资源消耗和水稻生产成本,还导致河水富营养化、地下水污染等不良的环境反应。因此,提高水稻氮素吸收利用效率对于提高我国种稻效益与生态环境效益,促进稻作生产的可持续发展具有重要现实意义。选育氮素高效吸收利用的品种,充分挖掘水稻自身对氮素吸收利用的潜力是提高水稻产量和氮素吸收利用效率最经济、最有效的途径。水稻产量可理解为吸氮量与氮素籽粒生产效率(吸收到植株体内单位氮素生产的籽粒产量)的乘积,提高吸氮量或提高氮素籽粒生产效率均可显著提高水稻产量水平。从这个角度看,氮高效水稻可以分为氮素高效吸收和氮素高效利用两种类型。关于氮素吸收与水稻其他性状关系的研究已有不少报道,但供试材料多为几个品种,亦有含较多材料的品种试验,但多为籼型水稻。关于不同氮素吸收类型粳稻品种间产量及其构成因素、源库指标、物质生产与分配、氮素吸收利用及根系性状的演变趋势及差异,氮素高效吸收型粳稻品种上述指标有何特点,目前尚不清楚。本研究于2008-2009年在群体水培条件下,以国内外不同年代育成的94个常规粳稻品种为供试材料,测定其抽穗期和成熟期的产量及其构成因素、不同器官的干物重、含氮率、叶面积系数、库容量、根系形态性状、根系活性等,试图明确氮素高效吸收型粳稻品种的基本特点及影响氮素吸收利用的主要因素,以期为粳稻品种氮素高效吸收利用的遗传改良,持续提高粳稻品种的产量水平提供参考依据。前人研究及本研究粳稻品种生育期、株高差异较大,据一些研究表明,生育期、株高可能会对水稻氮素吸收利用产生较大的影响,在生育期、株高分布相对集中的遗传群体中,它们是否会对氮素吸收利用产生明显影响目前尚不清楚。为此,于2010-2011年进一步开展试验,以生育期、株高分布相对集中的染色体单片段代换系群体(Single Segment Substitution Lines, SSSLs)122个株系为材料,分析氮高效吸收型水稻株系的基本特点,分析两个不同水稻群体影响氮素吸收利用的因素是否存在差异,差异多大。主要研究结果如下：
1.供试水稻品种或株系间吸氮量的差异很大。粳稻品种吸氮量的变幅为16.47～48.41g·m-2(2008年)和14.27～37.00g·m2(2009年),吸氮量最大品种为最小品种的2.94倍(2008年)和2.59倍(2009年)；SSSLs吸氮量的变幅为14.25-53.59g·m-2,最大的株系是最小的株系的3.76倍(2010年)。
2.供试水稻品种株高、生育期和全生育期吸氮强度均存在显著差异,但SSSLs的株高、生育期分布较粳稻品种相对集中,生育期更集中。氮高效吸收型水稻株高、全生育期吸氮强度明显大于氮低效吸收型水稻,生育期明显长于(粳稻)或稍长于(SSSLs)氮低效吸收型水稻。生育期、全生育期吸氮强度均是影响水稻氮素吸收的重要因素,相关和通径分析均表明,全生育期吸氮强度对吸氮量的影响大于生育期,SSSLs更突出。
3.不同氮素吸收型水稻产量的差异显著。随着吸氮量增加,粳稻品种产量水平呈显著提高趋势,SSSLs的产量呈先增加后下降趋势。吸氮量与粳稻品种产量呈极显著线性正相关(r2008=0.8096**,r2009=0.7864**),与SSSLs产量呈极显著抛物线型正相关(R201o=0.5916**)。增加吸氮量有利于粳稻品种产量水平的提高,但过大会导致SSSLs的产量下降。
4.在产量构成因素方面,不同氮素吸收型水稻间单位面积穗数、每穗颖花数差异显著,氮素高效吸收型单位面积穗数、每穗颖花数多(粳稻品种)或较多(SSSLs),结实率和千粒重差异较小。在稳定结实率和千粒重的基础上,增加(粳稻品种)或适当(SSSLs)增加穗数、增加每穗粒数,可显著提高吸氮量,进而显著提高水稻的产量水平。
5.氮素高效吸收型粳稻品种抽穗期叶面积系数(高效、有效、总叶面积)、抽穗期单穗叶面积(高效、有效、总叶面积)、成熟期单穗和总叶面积系数、抽穗期绿叶重、单穗和群体库容量均大于氮素低效吸收类型品种,但比叶重无显著优势,吸氮量过大时,使单位叶面积籽粒产量、结实期净同化率下降。氮高效吸收型SSSLs水稻在抽穗期和成熟期总叶面积系数、绿叶重、单穗和群体库容量、结实期净同化率等性状与粳稻品种有相同的趋势,均显著大于氮低效吸收型水稻,且比叶重也大于氮低效吸收型。粳稻品种分析表明,叶面积系数和库容量对总吸氮量均有重要的影响,库容量的作用大于叶面积系数。
6.不同氮素吸收型粳稻品种间干物质生产量存在显著差异,氮素高效吸收型水稻抽穗期、成熟期群、个体干物质生产量显著大于其它类型水稻,经济系数无显著优势(粳稻品种)或小于中等吸收型(SSSLs)。氮素高效吸收型粳稻品种各器官(根、茎鞘叶、穗)干重和单穗重大；氮素高效吸收型SSSLs根干重、茎鞘叶干重大,穗干重、单穗重虽较大但小于中等偏大吸收型SSSLs。粳稻品种相关和通径分析表明：成熟期群体干物质生产量对总吸氮量作用大于经济系数的作用；抽穗前物质生产量、抽穗后物质生产量对成熟期物质生产量均有重要的作用,前者略大于后者；单穗干物质生产量对群体干物质生产量的作用大于单位面积穗数的作用,抽穗前更明显；提高抽穗前后茎鞘叶干重和成熟期穗干重有利于成熟期干物质生产量提高；促进单穗干物质生产量尤其是抽穗前单穗干物质生产量,促进抽穗前后茎鞘叶干重和抽穗后穗干重的提高可显著提高氮高效吸收型品种成熟期群体物质生产量。
7.氮素高效吸收型水稻抽穗期吸氮量显著大于氮素低效吸收型品种。氮高效吸收型粳稻品种各器官(根、茎鞘叶、穗)吸氮量、单穗吸氮量、全生育期(群、个体)吸氮强度均较大,但植株含氮率、氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数较无优势。氮素高效吸收型SSSLs根、茎鞘叶吸氮量表现与粳稻品种有相同的趋势,均大于氮低效吸收型；植株含氮率较大,氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数较小,穗吸氮量虽较大但小于中等偏大型水稻。粳稻品种的通径分析表明,抽穗前吸氮量对总吸氮量影响显著大于抽穗后吸氮量的影响；单穗吸氮量对总吸氮量的影响显著大于穗数；干物质生产量对总吸氮量的影响显著大于植株含氮率
8.不同氮素吸收型水稻品种(株系)间根系性状差异显著。氮素高效吸收型粳稻品种单株根系性状(单株根数、单株根干重、单株不定根总长、单株总吸收面积、单株活跃吸收面积、单株根系活力、最长根长、众数根长)、单条根系形状(单条不定根长、单条不定根粗)、单穗根系性状(单穗根数、单穗根干重、单穗根系活跃吸收面积)、抽穗期冠根比、成熟期冠根比均不同程度地优于氮素低效吸收型粳稻品种,单条不定根干重较大但小于氮素中等偏大吸收型品种,单穗总根长、单穗根系总吸收面积、单穗根粗没有差异。氮素高效吸收型SSSLs单株根干重、单株不定根总长、单株根系总吸收面积、单株根系活跃吸收面积、最长根长、众数根长、抽穗期冠根比与粳稻品种表现相同的趋势,均大于氮低效吸收型水稻,但成熟期冠根比、单株不定根数、单株根系活力或小或无优势。通径分析表明,无论是粳稻品种还是SSSLs,根干重、成熟期冠根比、单株不定根长、单条不定根重是影响粳稻品种吸氮能力的主要根系性状,说明氮高效吸收型株系根系不仅较长,且充实程度也较高。改良根干重、冠根比(抽穗期和成熟期)、单株不定根总长、单条不定根重等性状能显著提高水稻的吸氮量。
The common problem in paddy field of China is the excessive usage of nitrogen fertilizer and low nitrogen utilization efficiency, which caused eutrophication and ground water pollution, meanwhile increased the cost of rice production. In light of this, improving nitrogen uptake and utilization efficiency of rice is very important for sustainable development of rice production. The increase of rice yield could be achieved by exploring genotypic variation in nitrogen uptake and utilization efficiency among rice cultivars and developing new varieties with high nitrogen uptake and utilization efficiency. Rice yield is the product of amount of nitrogen absorption (ANA) and nitrogen use efficiency for grain yield (NUEg), the increase of ANA or NUEg can improve yield, accordingly, rice cultivars with high nitrogen efficiency could be grouped into high nitrogen uptake efficiency and high nitrogen utilization efficiency. Previous reports on the relationship between nitrogen uptake and other traits usually included only a few varieties, or most of them are indica rice varieties. The genotypic variation in conventional japonica rice cultivars of different ANA types on yield and yield components, index of source and sink, dry matter accumulation and distribution, nitrogen distribution, nitrogen uptake efficiency and use efficiency and root traits are not known, and no work has been conducted on characterising traits determing high nitrogen uptake efficiency in conventional japonica rice cultivars. Some studies suggested that, plant height and growth duration have great influence on nitrogen uptake and use efficiency, but it's not clear these two factors affect nitrogen uptake efficiency of rice cultivars with similar plant height and phenology. Ninty four conventional japonica rice cultivars were solution-cultured in2008and2009, respectively. Grain yield, dry matter weight, as well as nitrogen content in individual organs, leaf area index (LAI), sink potential, morphological and physiological traits of root were measured. The objectives of the study were to characterize the traits of japonica rice cultivars with high nitrogen uptake efficiency, in order to provide experimental evidence for genetic improvements of conventional japonica rice cultivars.122Single Segment Substitution Lines(SSSLs) were solution-cultured in2010and2011to characterize lines with different nitrogen uptake efficiency. The similarities and differences between Japonic rice cultivars and SSSLs with similar genetic background were compared on nitrogen uptake effeciency. The main results were as follows:
1. There was a significant difference among all tested cultivars and lines on amount of nitrogen absorption (ANA). The range of ANA in japonica rice cultivars was16.47-48.41g·m-2(2008) and14.27～37.00g·m-2(2009), the biggest ANA was2.94and2.59times of the minimum one in2008and2009, respectively; The range of ANA in SSSLs was14.25-53.59g·m-2, the biggest ANA was3.76times of the minimum one(2009).
2. There was a significant difference in plant height, growth duration and nitrogen uptake intensity among different types of cultivars and SSSLs, while the distribution of growth duration and nitrogen uptake intensity was more concentrated in SSSLs. The plant height, growth duration and nitrogen uptake intensity of high nitrogen uptake efficiency types were superior to those with low uptake efficiency. Plant height and growth duration were the main factors influencing ANA, correlation analysis and direct path analysis showed that, the influence of nitrogen uptake intensity to ANA was bigger than growth duration, especially in SSSLs.
3. There was a significant difference in yield among different types of japonica rice cultivars and SSSLs. With the increase of ANA, The grain yield ofjaponica rice cultivars increased significantly, the yield of SSSLs increased first then declined. Grain yield of japonica rice cultivars was significantly related to ANA(r2008=0.8096**,r2009=0.7864**), while grain yield was significantly parabolic related to ANA(R2010=0.5916**). Enhancing ANA was beneficial to improve the grain yield of japonica rice cultivars, while the grain yield of SSSLs will decline when the ANA was too big.
4. In terms of yield components, there were remarkable differences in panicle number per area and spikelet number per panicle among different uptake efficiency types of conventional japonica rice cultivars and SSSLs. Rice cultivars with high N uptake efficiency showed more panicle number per area and spikelet number per panicle than those with low N uptake efficiency. There was no significant difference in grain filling rate and1000-grain weight both of Japonic rice cultivars and SSSLs. Enhancing panicle number per area and spikelet number per panicle on stable grain filling rate and1000-grain weight, could improve the grain yield of japonica rice cultivars significantly.
5. The traits of rice cultivars with high N uptake efficiency, including LAI (high effective LAI, effective LAI, total LAI) and leaf area per panicle (high effective leaf area per panicle, effective leaf area per panicle, total leaf area per panicle) at heading stage, LAI and leaf area per panicle at ripening stage, green leaf weight in heading stage, Sink potential, as well as sink potential per panicle were bigger than those with low N uptake efficiency. Grain yield per unit leaf area and net assimilation rate would declined when ANA was too big, there was no significant superiority in green leaf weight per unit leaf area among different types. The traits of SSSLs with high N uptake efficiency, including LAI at heading and ripening stage, green leaf weight in heading stage, green leaf weight per unit leaf area, Sink potential and sink potential per panicle, as well as net assimilation rate were bigger than those with low N uptake efficiency. Both the LAI and sink potential had important influence on the ANA, and the influence of sink potential was stronger than the LAI.
6. There was remarkable difference in dry matter accumulation among different types of rice. Dry matter accumulation and dry matter accumulation per panicle at heading and maturity stage with high N uptake efficiency were significantly higher than those with low N uptake efficiency, while there was no significant difference in harvest index. Japonica rice cultivars of high N uptake efficiency types showed higher organic dry matter weight (root, stem, sheath and leaf, panicle) and panicle weight. SSSLs in high nitrogen uptake efficiency types showed bigger dry matter weight of root, dry matter weight of stem, sheath and leaf than other types, while dry matter weight of panicle, single panicle weight, harvest index was smaller than those in medium nitrogen uptake efficiency types. Correlation analysis and direct path analysis showed that:dry matter accumulation at maturity stage had more important effect on ANA than the harvest index. Both of dry matter accumulation before and after heading had important influence on the dry matter accumulation at maturity stage, especially the dry matter accumulation before heading. The dry matter accumulation per single panicle had more important effect on the dry matter accumulation at maturity than the panicle numbers per unit area, especially the dry matter accumulation per single panicle before heading. The improving of dry matter accumulation of leaf sheath and stem and panicle weight at maturity was beneficial to improving dry matter accumulation at maturity stage. The promotion of the dry matter accumulation per single panicle, especially the dry matter accumulation per single panicle before heading, as well as the promotion of dry matter per leaf sheath and stem heading and after heading and dry matter per panicle after heading could significantly improved the dry matter accumulation per population in the N uptake efficiency of japonica rice cultivars.
7. Rice cultivars of high N uptake efficiency types had significantly higher ANA at heading stage than those with low N uptake efficiency. The ANA of organics (root, stem, sheath and leaf, panicle), ANA per panicle, population nitrogen uptake intensity and nitrogen uptake intensity per panicle of rice cultivars with high N uptake efficiency were bigger than those with low N uptake efficiency, while N content, N use efficiency for grain output, N use efficiency for dry matter production and nitrogen harvest index had no advantage. SSSLs of high nitrogen uptake efficiency types showed bigger ANA of root, ANA of stem, sheath and leaf, N content, while N use efficiency for grain output, N use efficiency for dry matter production and nitrogen harvest index were smaller. ANA of panicle of SSSLs with high nitrogen uptake efficiency was smaller than that with medium N uptake efficiency. Direct path analysis showed that:ANA before heading stage had significant effect on the total ANA than the ANA at grain filling stage; ANA of single panicle makes significant greater effect on total ANA than the panicle number; Dry matter accumulation makes significant greater effect on total ANA than N content.
8. There was a significant difference in root traits among different types of japonica rice cultivars and lines. The root traits of rice cultivars with high N uptake efficiency, including root traits per plant (number of adventitious roots, root dry weight, total length of adventitious roots, total root absorptions area, root activity absorptions area, root activity per plant, the longest root length, model root length), single root traits (single adventitious root length, single adventitious root diameter), root traits per panicle (number of adventitious roots, root dry weight, root activity absorptions area per panicle), ratio of shoot to root at heading and ripening stage, were superior to those with low N uptake efficiency more or less. Weight of single adventitious root with high N uptake efficiency was smaller than that with medium N uptake efficiency, while total length of adventitious roots per panicle, total root absorption area per panicle and adventitious root diameter per panicle had no advantage. SSSLs of high nitrogen uptake efficiency types showed bigger root dry weight per plant, total length of adventitious roots per plant, total root absorptions area per plant, root activity absorptions area, the longest root length, model root length and ratio of shoot to root at heading stage, but smaller Shoot to root at ripening stage. There was no advantage in number of adventitious roots per plant and root activity per plant. Further analysis showed that, both of japonica rice cultivars and lines, high ANA could be reached by improving root dry weight per plant, ratio of shoot to root (at heading and ripening stage), number of adventitious roots per plant, Weight of single adventitious root. Root dry weight per plant, the ratio of shoot to root (at heading and ripening stage), the total length of adventitious roots per plant and dry weight per adventitious are the main factors influencing N absorptive capacity.
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