水稻高产与养分高效利用栽培技术及其生理基础的研究
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摘要
水稻是我国最大的粮食作物。随着人口的增长和经济的发展,需要不断增加粮食产量。但另一方面,随着产量的增加,需要加大水肥资源的投入。持续提高作物产量是否必须依赖于水肥资源的大量投入?作物高产与资源高效能否协同?这一直是国内外关注的热点,也是学术界仍在争论的重大科学命题。发达国家在这个问题上往往采用环境优先的原则,而我国人多地少、资源紧缺,持续提高作物单产,同时高效利用有限的资源,是农业可持续发展的必由之路。本研究通过栽培技术的集成与优化,设置了不同的栽培模式,分析其对水稻产量、群体生育特性、养分吸收与利用效率、生理性状以及稻米品质的影响。主要结果如下：
1、水稻高产高效的限制因素与水稻高产高效栽培技术
通过调查、实地取证和试验,明确了限制单季水稻高产与氮肥高效利用的主要因素是栽培措施不当,即秧苗素质差、栽插密度过稀、施肥量大、大水漫灌等直接影响了群体质量和造成资源浪费；品种的耐肥性太强和大穗型品种弱势粒充实差也是影响水稻高产高效的一个重要原因。以水稻实地养分管理技术、轻干湿交替灌溉技术为核心技术,以培育壮秧、增密控肥、前氮后移、叶面喷施和增施硅肥等为配套技术,创建集成了水稻高产高效栽培技术体系。
2、水稻高产高效群体生长发育特性
(1)以杂交粳稻常优3号和常优5号为材料,设置了未施氮处理(ON)、当地高产栽培(对照)、高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培等6种不同栽培模式,观察上述栽培模式对水稻产量和群体质量的影响(试验一)。结果表明,高产高效栽培、超高产栽培、超高产高效栽培的产量两品种平均分别达9.9t hm-2、12.0t hm-2、10.9t hm-2和9.3t hm-2,较当地常规高产栽培高出10.1%-41.9%。(2)以扬粳4038、扬稻6号以及连粳6号为材料,设置ON、当地高产栽培以及高产高效栽培3个处理,并观察3个处理对水稻产量及群体质量的影响(试验二)。结果表明,与当地高产栽培相比,高产高效栽培模式下水稻产量平均增加了14.4%。与当地高产栽培相比,高产高效栽培、超高产栽培以及超高产高效栽培等处理的茎蘖成穗率明显提高,叶面积指数和有效叶面积比率增加,叶面积持续期、作物生长率和抽穗至成熟期的干物质积累增加。还提高了水稻粒叶比,改善了源库关系,并提高了根干重和根系伤流量。说明通过高产高效栽培可以改善水稻群体质量,获得更高产量。
3、高产高效栽培水稻的养分吸收与利用效率
试验一的结果表明,与当地高产栽培相比,高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培的氮肥吸收利用率分别提高了50.3%、87.4%、78.5%和44.3%,氮肥农学利用率分别提高了74.7%、91.1%、97.1%和78.3%,高产高效栽培模式、超高产高效栽培模式以及氮高效利用栽培模式的抽穗至成熟期养分运转率也显著提高。不同栽培模式下水稻穗分化至抽穗期的氮、磷、钾的吸收量与产量呈显著正相关。试验二的结果表明,与当地高产栽培相比,高产高效栽培下水稻水分利用效率、氮肥吸收利用率、农学利用率分别增加了35.6%、42.5%和64.1%。上述结果表明通过栽培技术的集成优化,可以大幅度同步提高水稻产量和水肥利用效率。
4、水稻高产高效的生理特性
在试验一中,观察了不同栽培模式对水稻叶片、根系以及籽粒中生理活性的影响。结果表明,与当地高产栽培相比,高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培显著提高了剑叶SPAD值、光合速率和过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)等抗氧化酶活性和根系氧化力,增加了根系和籽粒中细胞分裂素含量以及籽粒中吲哚乙酸的含量。在分蘖期,上述各栽培模式的叶片硝酸还原酶(NR)、谷氨酰胺合成酶(GS)、谷氨酸合成酶(GOGAT)等氮代谢酶活性与当地高产栽培差异不显著,但在穗分化期、抽穗期和成熟期均显著增加。在试验二中,观察了高产高效栽培对水稻植株中细胞分裂素玉米素(Z)+玉米素核苷(ZR)含量、灌浆期剑叶光合特性和根系活性以及籽粒中与蔗糖-淀粉转化相关酶活性的影响。结果表明,与当地高产栽培相比,高产高效栽培提高了水稻灌浆期剑叶光合速率、根系氧化力和籽粒中蔗糖合酶(Susase)和腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)活性,增加了生育中期和后期植株中的Z+ZR含量。说明高产高效栽培通过提高水稻根系和地上部的生理活性,促进高产与水肥的高效利用。
5、高产高效栽培对稻米品质及其米粉中矿质元素含量的影响
观察了试验一中6种栽培模式对水稻稻米品质和精米中矿质元素含量的影响。结果表明,与当地高产栽培相比,超高产栽培和超高产高效栽培显著增加了稻米的整精米率、蛋白质含量和矿质元素在精米中的累积量,增加了稻米的胶稠度、崩解值,降低了垩白度、直链淀粉含量和消减值。说明高产高效栽培可以改善稻米的加工品质、外观品质及营养品质。
以上结果说明,高产高效栽培模式不仅可以显著提高产量,而且可以显著提高水分和养分利用效率。在高产高效栽培模式下,较高的根系活力、叶片光合势和氮代谢酶活性、茎中光合同化物向籽粒的运转率、植株中较高的Z+ZR含量和灌浆期籽粒中较高的蔗糖-淀粉凋谢途径关键酶活性是高产高效栽培模式获取高产与养分高效利用的重要生理原因。水稻实地养分管理技术和轻干湿交替灌溉技术是高产高效栽培模式的两个关键栽培技术。
Rice is the foremost stable food crop in China. With the growth of population and economic development, it is necessary to increase more crop yield. Meanwhile, this increase needs to be accomplished under high input of water and fertilizer resources. It remains debates whether it is necessary to input more water and fertilizer resources for more food, and whether higher grain yield and higher nutrient use efficiency can be coordinately achieved? It is not only always focusing of attention at home and abroad, but also an important scientific proposition in academic circles. In developed contries, the principle is adopted that environment takes for the first place. But, in China, there is less land and a shortage of resources for large population. It is an essential way for agricultural sustainable development to increase crop yield and meanwhile increase resource use efficiency.This study investigated the characteristics of grain yield and popoulation growth and development, nutrient absorption and utilization efficiency, physiological characteristics and rice quality under different cultivation patterns throgh integrating and optimizing cultivation techniques. The main results are as follows:
1. The limiting factors and the cultivation techniques for high-yielding and high nutrient use efficiency in rice
By investigating, collecting evidence and useful information and conducting experiments, the main factors limiting high yielding and high nitrogen use efficiency were made known, i. e., improper cultivation measures, such as poor seedling quality, lower plant denstiny, higher fertilizer amount, and flodding irrigation which could result in poor quality of rice population directly and caused a waste of resources. Meanwhile, the cultivars with strong adaptability of fertilizer and the poor grain filling of inferor spikelets for a large panicle with numberios spikelets were also contributed to the reduction in grain yield and nutrient use efficiency in rice. A cultivation technique system was created and integrated by taking site-specific nutrient management(SSNM) and alternate wetting and moderate drying(WMD) as the two key techniques and some practices as supporting techqiques, including cultivating strong seedlings, increasing densitny, controlling fertilizer application, applying N fertilizer at later stage, spraying foliage fertilizer, and increasing silicon fertilizer application.
2. The characteristics of population growth and development of rice under high-yielding and high nutrient use efficiency cultivation
A high quality of population is necessary to achieve high grain yield in rice, but little information is avaliable on understanding how cultivation patterns affect population quality. In this study,two experiments were conducted.(1) two japonica hybrid rice cultivars Changyou3and Changyou5were grown in the field, with six cultivation patterns including no nitrogen application (ON), local high yielding cultivation (control), cultivation for high yielding and high efficiency(CHYHE), super high yielding cultivation (SHY), cultivation for super high yielding and high efficiency (SHYHE) and cultivation for high use efficiency of N (HUEN)(Experiment1). The results showed that, the average yields of two cultivars under CHYHE, SHY,SHYHE and HUEN were9.9t ha-1,12.01ha-1,10.9t ha-1and9.3t ha-1, respectively, and10.1%-41.9%' higher than these of control.(2) Three rice cultivars Yangjing4038,Yangdao6and Lianjing6were grown in the field, with three treatments including local farmer's practice (LFP), an improved crop management (ICM) and N omission (ON)(Experiment2).The results showed that, compared with LFP, the average grain yields of ICM increased by14.4%. Both experiment1and experiment2showed that, compared with control or LFP, CHYHE, SHY, SHYE and HUEN or ICM increased percentage of productive tillers, leaf area index(LAI) and efficient LAI, leaf area duration(LAD), crop growth rate (CGR), biomass from heading to mature, grain-leaf ratio, root biomass and root bleeding after heading. The results indicate that the population quality of rice could be improved and higher yield could be achieved through high-yielding and high nutrient use efficiency cultivation patterns.
3. Nutrient absorption and utilization efficiency of rice under high-yielding and high nutrient use efficiency cultivation
The objiective of this study was to investigate if a cultivation technique could coordinately increase both grain yield and nutrient use efficiency of rice. In this study, two experiments were conducted. The results of experiment1showed that, compared with those under the control, nitrogen recovery efficiency (NRE) was increased by50.3%,87.4%,78.5%and44.3%, and nitrogen agronomic efficiency(NAE) was increased by74.7%,91.1%,97.1%and78.3%respectively, under CHYHE, SHY, SHYE and HUNE. The percentage of nitrogen (N), phosphors (P), potassium (K) translocation from heading to maturity under CHYHE, SHYE and HUNE were also increased. The accumulation of N, P, K from panicle initation to heading was very significantly and positively correlated with grain yield.
The results of experiment2showed that, the average WUE, NRE and NAE under ICM were increased by35.6%,42.5%and64.1%, respectively, than those under LFP. The results of two experiments indicate that a higher yield and higher WUE and NUE can be achieved through integrating and optimizing cultivatin techniques in rice production.
4. Physiological traits of rice under high-yielding and high nutrient efficiency cultivation
The results from experiment1showed that, compared with the control, CHYHE, SHY, SHYE and HUEN increased SPAD values, photosynthetic rate (Phr) and activities of antioxidant enzymes, catalase (CAT), peroxidase (POD) and superoxide dismutase(SOD), in/of flag leaves, root oxidation activity (ROA), contents of zeatin+zeatin riboside(Z+ZR) in roots and grains and indoleacetic acid (IAA) in grains. The activities of key enzymes involved in nitrogen metabolism, nitrate reductase (NR), glutamine synthetase (GS), and glutamate synthase (GOGAT), showed no significant difference at mid-tillering stage, but were significantly higher than the control at the panicle initation, heading time and maturity stages. Results in experiment2showed that, compared with LFP, ICM increased Phr in flag leaves, ROA, activities of sucrose synthase (SuSase) and adenine diphosphogiucose pyrophosphorylase (AGPase) in garains during grain filling, and contents of Z+ZR in leaves, panicles and roots at mid and late growth stages. The results of two experiments suggest that an enhanced phsiological activties of shoot and root contributes to the increase in grain yield and nutrient use efficiency under the high-yielding and high nutrient use efficiency cultivation patterns.
5. The rice qualities and contents of mineral elements in milled rice under high-yielding and high nutrient use efficiency cultivation
This study was to investigate the formation characteristics of rice quality under high-yielding and high nutrient use efficiency cultivation. In experiment1, the rice qualities and contents of mineral elemnts in milled rice were analyzed under six cultivation patterns. The results showed that, compared with the control, SHY and SHYE significantly increased head rice, contents of protein and mineral elements accumulation in milled rice, and increased the gel consistency and breakdown of rice, and reduced chalkness, amylose and setback. The results indicate that the milling quality, appearance quality and nutrient quality could be improved under the high-yielding and high nutrient use efficiency cultivation.
In summary, a high-yielding and high nutrient use efficiency cultivation pattern could increase not only grain yield but also nutrient use efficiency and water use efffiency. A larger ROA and leaf area duration, higher activities of key enzymes involved in nitrogen metabolism in leaves, more remobilization of pre-stored NSC from stems to the grain, greater content of Z+ZR in plants, and enhanced activities of key enzymes involved in sucrose-to-starch conversion in grains during the grain filling period under the high-yielding and high nutrient use efficiency cultivation pattern may have contributed to achieving the dual goal of increasing grain yield and resource use efficiency. The SSNM and AWMD irrigation were two key techniques in the high-yielding and high nutrient use efficiency cultivation system of rice.
1、水稻高产高效的限制因素与水稻高产高效栽培技术
通过调查、实地取证和试验,明确了限制单季水稻高产与氮肥高效利用的主要因素是栽培措施不当,即秧苗素质差、栽插密度过稀、施肥量大、大水漫灌等直接影响了群体质量和造成资源浪费；品种的耐肥性太强和大穗型品种弱势粒充实差也是影响水稻高产高效的一个重要原因。以水稻实地养分管理技术、轻干湿交替灌溉技术为核心技术,以培育壮秧、增密控肥、前氮后移、叶面喷施和增施硅肥等为配套技术,创建集成了水稻高产高效栽培技术体系。
2、水稻高产高效群体生长发育特性
(1)以杂交粳稻常优3号和常优5号为材料,设置了未施氮处理(ON)、当地高产栽培(对照)、高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培等6种不同栽培模式,观察上述栽培模式对水稻产量和群体质量的影响(试验一)。结果表明,高产高效栽培、超高产栽培、超高产高效栽培的产量两品种平均分别达9.9t hm-2、12.0t hm-2、10.9t hm-2和9.3t hm-2,较当地常规高产栽培高出10.1%-41.9%。(2)以扬粳4038、扬稻6号以及连粳6号为材料,设置ON、当地高产栽培以及高产高效栽培3个处理,并观察3个处理对水稻产量及群体质量的影响(试验二)。结果表明,与当地高产栽培相比,高产高效栽培模式下水稻产量平均增加了14.4%。与当地高产栽培相比,高产高效栽培、超高产栽培以及超高产高效栽培等处理的茎蘖成穗率明显提高,叶面积指数和有效叶面积比率增加,叶面积持续期、作物生长率和抽穗至成熟期的干物质积累增加。还提高了水稻粒叶比,改善了源库关系,并提高了根干重和根系伤流量。说明通过高产高效栽培可以改善水稻群体质量,获得更高产量。
3、高产高效栽培水稻的养分吸收与利用效率
试验一的结果表明,与当地高产栽培相比,高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培的氮肥吸收利用率分别提高了50.3%、87.4%、78.5%和44.3%,氮肥农学利用率分别提高了74.7%、91.1%、97.1%和78.3%,高产高效栽培模式、超高产高效栽培模式以及氮高效利用栽培模式的抽穗至成熟期养分运转率也显著提高。不同栽培模式下水稻穗分化至抽穗期的氮、磷、钾的吸收量与产量呈显著正相关。试验二的结果表明,与当地高产栽培相比,高产高效栽培下水稻水分利用效率、氮肥吸收利用率、农学利用率分别增加了35.6%、42.5%和64.1%。上述结果表明通过栽培技术的集成优化,可以大幅度同步提高水稻产量和水肥利用效率。
4、水稻高产高效的生理特性
在试验一中,观察了不同栽培模式对水稻叶片、根系以及籽粒中生理活性的影响。结果表明,与当地高产栽培相比,高产高效栽培、超高产栽培、超高产高效栽培和氮高效利用栽培显著提高了剑叶SPAD值、光合速率和过氧化物酶(POD)、过氧化氢酶(CAT)、超氧化物歧化酶(SOD)等抗氧化酶活性和根系氧化力,增加了根系和籽粒中细胞分裂素含量以及籽粒中吲哚乙酸的含量。在分蘖期,上述各栽培模式的叶片硝酸还原酶(NR)、谷氨酰胺合成酶(GS)、谷氨酸合成酶(GOGAT)等氮代谢酶活性与当地高产栽培差异不显著,但在穗分化期、抽穗期和成熟期均显著增加。在试验二中,观察了高产高效栽培对水稻植株中细胞分裂素玉米素(Z)+玉米素核苷(ZR)含量、灌浆期剑叶光合特性和根系活性以及籽粒中与蔗糖-淀粉转化相关酶活性的影响。结果表明,与当地高产栽培相比,高产高效栽培提高了水稻灌浆期剑叶光合速率、根系氧化力和籽粒中蔗糖合酶(Susase)和腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)活性,增加了生育中期和后期植株中的Z+ZR含量。说明高产高效栽培通过提高水稻根系和地上部的生理活性,促进高产与水肥的高效利用。
5、高产高效栽培对稻米品质及其米粉中矿质元素含量的影响
观察了试验一中6种栽培模式对水稻稻米品质和精米中矿质元素含量的影响。结果表明,与当地高产栽培相比,超高产栽培和超高产高效栽培显著增加了稻米的整精米率、蛋白质含量和矿质元素在精米中的累积量,增加了稻米的胶稠度、崩解值,降低了垩白度、直链淀粉含量和消减值。说明高产高效栽培可以改善稻米的加工品质、外观品质及营养品质。
以上结果说明,高产高效栽培模式不仅可以显著提高产量,而且可以显著提高水分和养分利用效率。在高产高效栽培模式下,较高的根系活力、叶片光合势和氮代谢酶活性、茎中光合同化物向籽粒的运转率、植株中较高的Z+ZR含量和灌浆期籽粒中较高的蔗糖-淀粉凋谢途径关键酶活性是高产高效栽培模式获取高产与养分高效利用的重要生理原因。水稻实地养分管理技术和轻干湿交替灌溉技术是高产高效栽培模式的两个关键栽培技术。
Rice is the foremost stable food crop in China. With the growth of population and economic development, it is necessary to increase more crop yield. Meanwhile, this increase needs to be accomplished under high input of water and fertilizer resources. It remains debates whether it is necessary to input more water and fertilizer resources for more food, and whether higher grain yield and higher nutrient use efficiency can be coordinately achieved? It is not only always focusing of attention at home and abroad, but also an important scientific proposition in academic circles. In developed contries, the principle is adopted that environment takes for the first place. But, in China, there is less land and a shortage of resources for large population. It is an essential way for agricultural sustainable development to increase crop yield and meanwhile increase resource use efficiency.This study investigated the characteristics of grain yield and popoulation growth and development, nutrient absorption and utilization efficiency, physiological characteristics and rice quality under different cultivation patterns throgh integrating and optimizing cultivation techniques. The main results are as follows:
1. The limiting factors and the cultivation techniques for high-yielding and high nutrient use efficiency in rice
By investigating, collecting evidence and useful information and conducting experiments, the main factors limiting high yielding and high nitrogen use efficiency were made known, i. e., improper cultivation measures, such as poor seedling quality, lower plant denstiny, higher fertilizer amount, and flodding irrigation which could result in poor quality of rice population directly and caused a waste of resources. Meanwhile, the cultivars with strong adaptability of fertilizer and the poor grain filling of inferor spikelets for a large panicle with numberios spikelets were also contributed to the reduction in grain yield and nutrient use efficiency in rice. A cultivation technique system was created and integrated by taking site-specific nutrient management(SSNM) and alternate wetting and moderate drying(WMD) as the two key techniques and some practices as supporting techqiques, including cultivating strong seedlings, increasing densitny, controlling fertilizer application, applying N fertilizer at later stage, spraying foliage fertilizer, and increasing silicon fertilizer application.
2. The characteristics of population growth and development of rice under high-yielding and high nutrient use efficiency cultivation
A high quality of population is necessary to achieve high grain yield in rice, but little information is avaliable on understanding how cultivation patterns affect population quality. In this study,two experiments were conducted.(1) two japonica hybrid rice cultivars Changyou3and Changyou5were grown in the field, with six cultivation patterns including no nitrogen application (ON), local high yielding cultivation (control), cultivation for high yielding and high efficiency(CHYHE), super high yielding cultivation (SHY), cultivation for super high yielding and high efficiency (SHYHE) and cultivation for high use efficiency of N (HUEN)(Experiment1). The results showed that, the average yields of two cultivars under CHYHE, SHY,SHYHE and HUEN were9.9t ha-1,12.01ha-1,10.9t ha-1and9.3t ha-1, respectively, and10.1%-41.9%' higher than these of control.(2) Three rice cultivars Yangjing4038,Yangdao6and Lianjing6were grown in the field, with three treatments including local farmer's practice (LFP), an improved crop management (ICM) and N omission (ON)(Experiment2).The results showed that, compared with LFP, the average grain yields of ICM increased by14.4%. Both experiment1and experiment2showed that, compared with control or LFP, CHYHE, SHY, SHYE and HUEN or ICM increased percentage of productive tillers, leaf area index(LAI) and efficient LAI, leaf area duration(LAD), crop growth rate (CGR), biomass from heading to mature, grain-leaf ratio, root biomass and root bleeding after heading. The results indicate that the population quality of rice could be improved and higher yield could be achieved through high-yielding and high nutrient use efficiency cultivation patterns.
3. Nutrient absorption and utilization efficiency of rice under high-yielding and high nutrient use efficiency cultivation
The objiective of this study was to investigate if a cultivation technique could coordinately increase both grain yield and nutrient use efficiency of rice. In this study, two experiments were conducted. The results of experiment1showed that, compared with those under the control, nitrogen recovery efficiency (NRE) was increased by50.3%,87.4%,78.5%and44.3%, and nitrogen agronomic efficiency(NAE) was increased by74.7%,91.1%,97.1%and78.3%respectively, under CHYHE, SHY, SHYE and HUNE. The percentage of nitrogen (N), phosphors (P), potassium (K) translocation from heading to maturity under CHYHE, SHYE and HUNE were also increased. The accumulation of N, P, K from panicle initation to heading was very significantly and positively correlated with grain yield.
The results of experiment2showed that, the average WUE, NRE and NAE under ICM were increased by35.6%,42.5%and64.1%, respectively, than those under LFP. The results of two experiments indicate that a higher yield and higher WUE and NUE can be achieved through integrating and optimizing cultivatin techniques in rice production.
4. Physiological traits of rice under high-yielding and high nutrient efficiency cultivation
The results from experiment1showed that, compared with the control, CHYHE, SHY, SHYE and HUEN increased SPAD values, photosynthetic rate (Phr) and activities of antioxidant enzymes, catalase (CAT), peroxidase (POD) and superoxide dismutase(SOD), in/of flag leaves, root oxidation activity (ROA), contents of zeatin+zeatin riboside(Z+ZR) in roots and grains and indoleacetic acid (IAA) in grains. The activities of key enzymes involved in nitrogen metabolism, nitrate reductase (NR), glutamine synthetase (GS), and glutamate synthase (GOGAT), showed no significant difference at mid-tillering stage, but were significantly higher than the control at the panicle initation, heading time and maturity stages. Results in experiment2showed that, compared with LFP, ICM increased Phr in flag leaves, ROA, activities of sucrose synthase (SuSase) and adenine diphosphogiucose pyrophosphorylase (AGPase) in garains during grain filling, and contents of Z+ZR in leaves, panicles and roots at mid and late growth stages. The results of two experiments suggest that an enhanced phsiological activties of shoot and root contributes to the increase in grain yield and nutrient use efficiency under the high-yielding and high nutrient use efficiency cultivation patterns.
5. The rice qualities and contents of mineral elements in milled rice under high-yielding and high nutrient use efficiency cultivation
This study was to investigate the formation characteristics of rice quality under high-yielding and high nutrient use efficiency cultivation. In experiment1, the rice qualities and contents of mineral elemnts in milled rice were analyzed under six cultivation patterns. The results showed that, compared with the control, SHY and SHYE significantly increased head rice, contents of protein and mineral elements accumulation in milled rice, and increased the gel consistency and breakdown of rice, and reduced chalkness, amylose and setback. The results indicate that the milling quality, appearance quality and nutrient quality could be improved under the high-yielding and high nutrient use efficiency cultivation.
In summary, a high-yielding and high nutrient use efficiency cultivation pattern could increase not only grain yield but also nutrient use efficiency and water use efffiency. A larger ROA and leaf area duration, higher activities of key enzymes involved in nitrogen metabolism in leaves, more remobilization of pre-stored NSC from stems to the grain, greater content of Z+ZR in plants, and enhanced activities of key enzymes involved in sucrose-to-starch conversion in grains during the grain filling period under the high-yielding and high nutrient use efficiency cultivation pattern may have contributed to achieving the dual goal of increasing grain yield and resource use efficiency. The SSNM and AWMD irrigation were two key techniques in the high-yielding and high nutrient use efficiency cultivation system of rice.
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