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Grain yield, nitrogen use efficiency and physiological performance of indica/japonica hybrid rice in response to various nitrogen rates
ZHOU Qun, YUAN Rui, ZHANG Wei-yang, GU Jun-fei, LIU Li-jun, ZHANG Hao, WANG Zhi-qin, YANG Jian-chang
2023, 22 (1): 63-79.   DOI: 10.1016/j.jia.2022.08.076
Abstract471)      PDF in ScienceDirect      

Utilizing the heterosis of indica/japonica hybrid rice (IJHR) is an effective way to further increase rice grain yield.  Rational application of nitrogen (N) fertilizer plays a very important role in using the heterosis of IJHR to achieve its great yield potential.  However, the responses of the grain yield and N utilization of IJHR to N application rates and the underlying physiological mechanism remain elusive.  The purpose of this study was to clarify these issues.  Three rice cultivars currently used in rice production, an IJHR cultivar Yongyou 2640 (YY2640), a japonica cultivar Lianjing 7 (LJ-7) and an indica cultivar Yangdao 6 (YD-6), were grown in the field with six N rates (0, 100, 200, 300, 400, and 500 kg ha–1) in 2018 and 2019.  The results showed that with the increase in N application rates, the grain yield of each test cultivar increased at first and then decreased, and the highest grain yield was at the N rate of 400 kg ha–1 for YY2640, with a grain yield of 13.4 t ha–1, and at 300 kg ha–1 for LJ-7 and YD-6, with grain yields of 9.4–10.6 t ha–1.  The grain yield and N use efficiency (NUE) of YY2640 were higher than those of LJ-7 or YD-6 at the same N rate, especially at the higher N rates.  When compared with LJ-7 or YD-6, YY2640 exhibited better physiological traits, including greater root oxidation activity and leaf photosynthetic rate, higher cytokinin content in the roots and leaves, and more remobilization of assimilates from the stem to the grain during grain filling.  The results suggest that IJHR could attain both higher grain yield and higher NUE than inbred rice at either low or high N application rates.  Improved shoot and root traits of the IJHR contribute to its higher grain yield and NUE, and a higher content of cytokinins in the IJHR plants plays a vital role in their responses to N application rates and also benefits other physiological processes. 

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Optimization of nitrogen fertilization improves rice quality by affecting the structure and physicochemical properties of starch at high yield levels
ZHOU Tian-yang, LI Zhi-kang, LI En-peng, WANG Wei-lu, YUAN Li-min, ZHANG Hao, LIU Li-jun, WANG Zhi-qin, GU Jun-fei, YANG Jian-chang
2022, 21 (6): 1576-1592.   DOI: 10.1016/S2095-3119(21)63678-X
Abstract268)      PDF in ScienceDirect      
A major challenge in modern rice production is to achieve the dual goals of high yield and good quality with low environmental costs.  This study was designed to determine whether optimized nitrogen (N) fertilization could fulfill these multiple goals.  In two-year experiments, two high yielding ‘super’ rice cultivars were grown with different N fertilization management regimes, including zero N input, local farmers’ practice (LFP) with heavy N inputs, and optimized N fertilization (ONF).  In ONF, by reducing N input, increasing planting density, and optimizing the ratio of urea application at different stages, N use efficiency and the physicochemical and textural properties of milled rice were improved at higher yield levels.  Compared with LFP, yield and partial factor productivity of applied N (PFP) under ONF were increased (on average) by 1.70 and 13.06%, respectively.  ONF increased starch and amylose content, and significantly decreased protein content.  The contents of the short chains of A chain (degree of polymerization (DP) 6–12) and B1 chain (DP 13–25) of amylopectin were significantly increased under ONF, which resulted in a decrease in the stability of rice starch crystals.  ONF increased viscosity values and improved the thermodynamic properties of starch, which resulted in better eating and cooking quality of the rice.  Thus, ONF could substantially compensate the negative effects caused by N fertilizer and achieve the multiple goals of higher grain quality and nitrogen use efficiency (NUE) at high yield levels.  These results will be useful for applications of high quality rice production at high yield levels.
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Deciphering the morpho–physiological traits for high yield potential in nitrogen efficient varieties (NEVs): A japonica rice case study
ZHU Kuan-yu, YAN Jia-qian, SHEN Yong, ZHANG Wei-yang, XU Yun-ji, WANG Zhi-qin, YANG Jian-chang
2022, 21 (4): 947-963.   DOI: 10.1016/S2095-3119(20)63600-0
Abstract202)      PDF in ScienceDirect      
The use of nitrogen (N)-efficient rice (Oryza sativa L.) varieties could reduce excessive N input without sacrificing yields. However, the plant traits associated with N-efficient rice varieties have not been fully defined or comprehensively explored.  Here, three japonica N-efficient varieties (NEVs) and three japonica N-inefficient varieties (NIVs) of rice were grown in a paddy field under N omission (0 N, 0 kg N ha–1) and normal N (NN, 180 or 200 kg N ha−1) treatments.  Results showed that NEVs exhibited higher grain yield and nitrogen use efficiency (NUE) than NIVs under both treatments, due to improved sink size and filled-grains percentage in the former which had higher root oxidation activity and greater root dry weight, root length and root diameter at panicle initiation (PI), as well as higher spikelet–leaf ratio and more productive tillers during the grain-filling stage.  Compared with NIVs, NEVs also exhibited enhanced N translocation and dry matter accumulation after heading and improved flag leaf morpho–physiological traits, including greater leaf thickness and specific leaf weight and higher contents of ribulose-1,5-bisphosphate carboxylase/oxygenase, chlorophyll, nitrogen, and soluble sugars, leading to better photosynthetic performance.  Additionally, NEVs had a better canopy structure, as reflected by a higher ratio of the extinction coefficient for effective leaf N to the light extinction coefficient, leading to enhanced canopy photosynthesis and dry matter accumulation.  These improved agronomic and physiological traits were positively and significantly correlated with grain yield and internal NUE, which could be used to select and breed N-efficient rice varieties.

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The effects of water and nitrogen on the roots and yield of upland and paddy rice
ZHANG Ya-jie, XU Jing-nan, CHENG Ya-dan, WANG Chen, LIU Gao-sheng, YANG Jian-chang
2020, 19 (5): 1363-1374.   DOI: 10.1016/S2095-3119(19)62811-X
Abstract109)      PDF in ScienceDirect      
It is of great significance to study the root characteristics of rice to improve water and nitrogen (N) use efficiency and reduce environmental pollution.  This study investigated whether root traits and architecture of rice influence grain yield, as well as water and N utilization efficiency.  An experiment was conducted using the upland rice cultivar Zhonghan 3 (a japonica cultivar) and paddy rice cultivar Huaidao 5 (also a japonica cultivar) using three N levels, namely, 2 g urea/pot (low amount, LN), 3 g urea/pot (normal amount, NN), and 4 g urea/pot (high amount, HN), and three soil water potentials (SWPs, namely, well-watered (0 kPa), mildly dried (–20 kPa) and severely dried (–40 kPa).  The results showed that with decreasing SWP, the percentage of upland rice roots increased in the 0–5 cm tillage layer, and decreased in the 5–10 and 10–20 cm tillage layers, whereas paddy rice roots showed the opposite trend.  With increasing amounts of N, the yield of upland and paddy rice increased, and the percentage of root volume ratios of the two rice cultivars in the 0–5 and 5–10 cm tillage layers increased, whereas that in the 10–20 cm tillage layer decreased.  The roots of upland rice are mainly distributed in the 10–20 cm tillage layer, whereas most paddy rice roots are in the 0–5 cm tillage layer.  These results indicate that the combination of −20 kPa SWP and NN in upland rice and 0 kPa SWP and LN in paddy rice promotes the growth of the root system during the middle and late stages, which in turn may decrease the requirements for water and N fertilizer and increase rice yield. 
 
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Changes in starch quality of mid-season indica rice varieties in the lower reaches of the Yangtze River in last 80 years
ZHANG Hao, JING Wen-jiang, XU Jing-ju, MA Bing-ju, WANG Wei-lu, ZHANG Wei-yang, GU Jun-fei, LIU Li-jun, WANG Zhi-qin, YANG Jian-chang
2020, 19 (12): 2983-2996.   DOI: 10.1016/S2095-3119(20)63431-1
Abstract119)      PDF in ScienceDirect      
Rice (Oryza sativa L.) quality depends mainly on the characteristics of starch stored in kernels.  Understanding the changes in starch characteristics in kernels during variety improvement would have great significance to improve rice quality.  This study was designed to investigate the starch characteristics in the kernels and associated physiological traits of indica rice varieties in the lower reaches of the Yangtze River in China in last 80 years.  Eight representative mid-season indica rice varieties were grown in the field.  The results showed that the grain yield was significantly increased with the improvement of varieties and such an increase was mainly attributed to the increase in total number of spikelets.  The tall varieties applied in the 1940s–1950s had higher protein content, relative crystallinity and infrared (IR) ratio of 1 045/1 022 cm–1.  The semi-dwarf varieties applied in the 1980s–1990s had higher gel consistency, amylopectin content, IR ratio of 1 022/995 cm–1, and breakdown value.  With the improvement of varieties, the amylose content, large-sized starch granule number and volume distribution, onset and peak of gelatinization temperature, gelatinization and retrogradation enthalpy, setback value, pasting temperature, viscosity of peak, hot and final, and 1-aminocycopropane-1-carboxylic acid (ACC) concentrations in panicles and root bleeding were gradually decreased, whereas the medium-sized starch granule number and volume distribution, activities of key enzymes in grains, and zeatin (Z)+zeatin riboside (ZR) contents in panicles and root bleeding at grain filling stage were gradually increased.  Correlation analysis showed that starch thermodynamic characteristics were closely related to starch structure and components, key enzymes and hormones.  The results suggest that starch quality was enhanced through the optimization of starch components, structure, thermodynamics, and the regulation of key enzymes in grains and hormones in panicles and root bleedings at grain filling stage during the improvement of mid-season indica rice.
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Optimizing integrative cultivation management improves grain quality while increasing yield and nitrogen use efficiency in rice
ZHANG Hao, HOU Dan-ping, PENG Xian-long, MA Bing-ju, SHAO Shi-mei, JING Wen-jiang, GU Jun-fei, LIU Li-jun, WANG Zhi-qin, LIU Yuan-ying, YANG Jian-chang
2019, 18 (12): 2716-2731.   DOI: 10.1016/S2095-3119(19)62836-4
Abstract202)      PDF in ScienceDirect      
A major challenge in rice (Oryza sativa L.) production is to cope with increasing grain yield and fertilizer use efficiency without compromising grain quality.  This study was designed to determine if optimizing integrative cultivation management in rice could improve grain quality while increase yield and nitrogen use efficiency (NUE).  An indica-japonica hybrid rice cultivar and a japonica rice cultivar were grown in the field, with five cultivation managements including no N application (0 N), local farmer’s practice (LFP), and three optimizing integrative cultivation managements, reducing N rate and increasing plant density (ND), ND+alternate wetting and moderate soil drying irrigation (NDW), and NDW+applying rapeseed cake fertilizer (NDWR).  The results showed that the optimizing integrative cultivation managements could not only increase grain yield, but also enhance NUE compared to LFP.  Compared to LFP, NDWR significantly increased brown, milled, head milled rice rate, ratio of the kernel length to breadth and breakdown value of starch, whereas decreased amylose content, gel consistency, prolamin content, setback value, percentage of chalky kernels, and chalkiness.  The three optimizing integrative cultivation managements increased contents of total proteins, albumin and glutelin, activities of the key enzymes involved in the sucrose-starch conversion in grains, root oxidation activity, and malic and succinic acid concentrations in root exudates during the grain-filling period.  The results suggested that optimizing integrative cultivation managements could improve grain quality meanwhile increase grain yield and NUE by enhancing physiological activities of rice plants.
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Physiological mechanism underlying spikelet degeneration in rice
WANG Zhi-qin, ZHANG Wei-yang, YANG Jian-chang
2018, 17 (07): 1475-1481.   DOI: 10.1016/S2095-3119(18)61981-1
Abstract446)      PDF (716KB)(364)      
The phenomenon of degenerated spikelets is very common in cereals, and considered as a serious physiological defect and a main constraint to grain production.  Understanding the physiological mechanism in which spikelet degeneration occurs would have great significance in enhancing yield potential in grain crops.  Taking rice as an example, the paper reviewed the physiological mechanism underlying spikelet degeneration, with focus on the roles of phytohormones in regulating the process.  There are several hypotheses for the spikelet degeneration, such as resource limitation, self-organization, and primigenic dominance.  However, convincing evidences are not enough to support the assumptions.  Phytohormones including auxins, cytokinins, gibberellins, abscisic acid, and ethylene are involved in regulating spikelet degeneration in cereals.  The new phytohormones of brassinosteroids and polyamines have been observed to suppress spikelet degeneration in rice.  The interactions among or between plant hormones may play a more important role in regulating spikelet degeneration.  However, the information on such interactions is very limited.  Some agronomic practices, especially proper water and nitrogen management, could reduce spikelet degeneration but the mechanism underlying remains unclear.  Further research is needed to understand the cross-talk among/between phytohormones on spikelet degeneration, to reveal the physiological and molecular mechanism in which phytohormones and their interactions regulate the degeneration of spikelets, to exploit approaches to decrease spikelet degeneration and to elucidate their mechanism.
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Grain yield and water use efficiency of super rice under soil water deficit and alternate wetting and drying irrigation
ZHOU Qun, JU Cheng-xin, WANG Zhi-qin, ZHANG Hao, LIU Li-jun, YANG Jian-chang, ZHANG Jian-hua
2017, 16 (05): 1028-1043.   DOI: 10.1016/S2095-3119(16)61506-X
Abstract941)      PDF in ScienceDirect      
This study investigated if super rice could better cope with soil water deficit and if it could have better yield performance and water use efficiency (WUE) under alternate wetting and drying (AWD) irrigation than check rice.  Two super rice cultivars and two elite check rice cultivars were grown in pots with three soil moisture levels, well watered (WW), moderate water deficit (MWD) and severe water deficit (SWD).  Two cultivars, each for super rice and check rice, were grown in field with three irrigation regimes, alternate wetting and moderate drying (AWMD), alternate wetting and severe drying (AWSD) and conventional irrigation (CI).  Compared with that under WW, grain yield was significantly decreased under MWD and SWD treatments, with less reduction for super rice than for check rice.  Super rice had higher percentage of productive tillers, deeper root distribution, higher root oxidation activity, and greater aboveground biomass production at mid and late growth stages than check rice, especially under WMD and WSD.  Compared with CI, AWMD increased, whereas AWSD decreased grain yield, with more increase or less decrease for super rice than for check rice.  Both MWD and SWD treatments and either AWMD or AWSD regime significantly increased WUE compared with WW treatment or CI regime, with more increase for super rice than for check rice.  The results suggest that super rice has a stronger ability to cope with soil water deficit and holds greater promising to increase both grain yield and WUE by adoption of moderate AWD irrigation.
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Performance in Grain Yield and Physiological Traits of Rice in the Yangtze River Basin of China During the Last 60 yr
ZHANG Hao, CHEN Ting-ting, LIU Li-jun, WANG Zhi-qin, YANG Jian-chang , ZHANG Jian-hua
2013, 12 (1): 57-66.   DOI: 10.1016/S2095-3119(13)60205-1
Abstract1963)      PDF in ScienceDirect      
Knowledge on the performance in grain yield and physiological traits is essential to understand the main yield-limiting factor and make strategies for breeding and crop management in rice (Oryza sativa L.). This study investigated the changes in grain yield and associated physiological traits of rice in the Yangtze River Basin of China during the last 60 yr. Thirteen mid-season indica and 12 japonica rice cultivars that were popularly used were grown in the field in 2008 and 2009. The grain yield and yield components, biomass, leaf area, leaf photosynthesis, root oxidation activity, and harvest index were examined. The results showed that grain yield and grain yield per day have progressively increased during the years and such increases are mainly attributed to the expanded sink size as a result of more spikelets per panicle, especially for the case of super rice. Both biomass and harvest index were increased with the improvement of cultivars. Increase in biomass for modern rice cultivars was associated with an enhancement of leaf area and photosynthesis, root dry weight, and root oxidation activity, although the indica super rice cultivars showed a lower leaf photosynthetic rate and root oxidation activity than the semi-dwarf cultivars during the grain filling period. Both indica and japonica super rice cultivars exhibited a low percentage of filled grains, which may limit their great yield potential. All the data suggested that grain yield have been substantially improved during the 60 yr of rice breeding in the Yangtze River Basin. Expanded sink size, increased dry matter production and harvest index, and enhanced leaf area and photosynthesis, root dry weight, and root oxidation activity contribute to the improvement in grain yield. Increase in filling efficiency could realize the great yield potential in super rice.
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Root Morphology and Physiology in Relation to the Yield Formation of Rice
YANG Jian-chang, ZHANG Hao, ZHANG Jian-hua
2012, 12 (6): 920-926.   DOI: 10.1016/S1671-2927(00)8614
Abstract2291)      PDF in ScienceDirect      
Root system is a vital part of plant and regulates many aspects of shoot growth and development. This paper reviews how some traits of root morphology and physiology are related to the formation of grain yield in rice (Oryza sativa L.). Higher root biomass, root oxidation activity, and cytokinin contents in roots are required for achieving more panicle number, more spikelets per panicle, greater grain-filling percentage, and higher grain yield. However, these root traits are not linearly correlated with yield components. When these traits reach very high levels, grain filling and grain yield are not necessarily enhanced. High numbers of mitochondria, Golgi bodies, and amyloplasts in root tip cells benefit root and shoot growth and yield formation. Proper crop management, such as an alternate wetting and moderate soil drying irrigation, can significantly improve ultra-structure of root tip cells, increase root length density and concentration of cytokinins in root bleedings, and consequently, increase grain-filling percentage, grain yield, and water use efficiency. Further studies are needed to investigate the mechanism underlying root-shoot and root-soil interactions for high grain yield, the roles of root-sourced hormones in regulating crop growth and development and the effects of soil moisture and nutrient management on the root architecture and physiology.
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