Effects of erect panicle genotype and environment interactions on rice yield and yield components

The dense and erect panicle (EP) genotype conferred by DEP1 has been widely used in the breeding of high-yield Chinese japonica rice varieties.However,the breeding value of the EP genotype has rarely been determined at the plant population level.Therefore,the effects of the interaction of EP genotype and the environment at different locations and times on rice yield and its various components were investigated in this study.Two sets of near-isogenic lines (NILs)of EP and non-EP (NEP) genotypes with Liaojing 5 (LG5) and Akitakomachi (AKI) backgrounds were grown in the field in 2016 and 2017 in Shenyang,China,and Kyoto,Japan.In 2018,these sets were grown only in Kyoto,Japan.The average yields of the EP and NEP genotypes were 6.67 and 6.13 t ha^(-1)for the AKI background,and 6.66 and 6.58 t ha^(-1)for the LG5 background,respectively.The EP genotype positively affected panicle number (PN) and grain number per square meter (GNPM),mostly resulting in a positive effect on harvest index (HI).In contrast,the EP genotype exerted a negative effect on thousand-grain weight (KGW).The ratio of the performance of the EP genotype relative to the NEP genotype in terms of yield and total biomass correlated positively with mean daily solar radiation during a 40-day period around heading.These results indicate that the effectiveness of the EP genotype depends on the availability of solar radiation,and the effect of this genotype is consistently positive for sink formation,conditional in terms of source capacity,and positive in a high-radiation environment.

Rice DENSE AND ERECT PANICLE 2 is essential for determining panicle outgrowth and elongation

The architecture of the panicle, including grain size and panicle morphology, directly determines grain yield. Panicle erectness, which is selected for achieving ideal plant arehitecture in the northern part of China, has drawn increasing attention of rice breeders. Here, dense and erect panicle 2 （dep2） mutant, which shows a dense and erect panicle phenotype, was identified. DEP2 encodes a plant-specific protein without any known functional domain. Expression profiling of DEP2 revealed that it is highly expressed in young tissues, with most abundance in young panicles. Morphological and expression analysis indicated that mutation in DEP2 mainly affects the rapid elongation of rachis and primary and secondary branches, but does not impair the initiation or formation of panicle primordia. Further analysis suggests that decrease of panicle length in dep2 is caused by a defect in cell proliferation during the exponential elongation of panicle. Despite a more compact plant type in the dep2 mutant, no significant alteration in grain production was found between wild type and dep2 mutant. Therefore, the study of DEP2 not only strengthens our understanding of the molecular genetic basis of panicle architecture but also has important implications for rice breeding.

Description of Aphelenchoides besseyi from Abnormal Rice with ‘Small Grains and Erect Panicles’ Symptom in China

The abnormal rice with small grains and erect panicles were found on a large scale in China, which showed shortend rice panicle and decreased number of grains in comparison with normal rice, and the grain was small and black-brown, and some of them were distorted, while the flag leaf was normal. A kind of nematode of Aphelenchoides was isolated from the grains of rice variety Wuyujing 3 which performed ＇small grains and erect panicles＇ symptom. There were 2014 nematodes in one hundred grains infected, and up to 74 in single grain, 92 percent of the grains tested had nematodes in the infested panicles. The diagnosis characters of nematode include lateral fields about one-fourth as wide as body, with 4 incisures. The terminus bears a mucro of diverse shape with 3-4 pointed processes. The female post-vulval uterine sac extends less than 50% of distance from vulva to anus, no sperm in it. Oocytes usually arrange in 2-4 rows. The male spicules have a moderately developed rostrum. Morphological measurements showed it to be conspecific with Aphelenchoides besseyi Christie, 1942.

Erect panicle super rice varieties enhance yield by harvest index advantages in high nitrogen and density conditions

The erect panicle （Ep） type is an important characteristic for japonica super rice in Northeast China and plays a significant role in enhancing yield. The Ep type is considered to be a genetic ideotype resource to the japonica super rice group by virtue of its agronomic advantages such as grain number per panicle and biomass. This study addresses the effects of nitrogen and planting density conditions on yielding performance regarding panicle type （PT） using the recombinant inbred line （RIL） population derived from the cross between an Ep variety Liaogeng 5 and non-Ep variety Wanlun 422. The genetics underlying the Ep type proved to be robust not only for panicle-type optimization but also plant height, panicle length, flag leaf length and seed density. We also found that regardless of nitrogen and density, correlation between harvest index （HI） and plant height was not significant in Ep type whatever the nitrogen and density. The application of Ep type provides a potential strategy for yield improvement by increasing biomass through HI maintainable in rice.

Effect of Cultural Measures on Nutrient Contellts in Rice Plants with Erect Panicles

Field experiments were carried out with rice variety of Shennong 91 of short culms and erect panicles to study nutrient contents in high-yielding rice plants and to increase rice yield by appropriate fertilization.Nitrogen, phosphorus, potassium, magnesium, zinc, manganese, iron and copper contents in rice plants varied with different treatment factors. The relationship between the nutrient contents and treatment factors could be simulated using a multiple quadratic equation. The nutrient contents in plants should be appropriate for high-yielding rice. If the mean nutrient content in rice plants producing 11 t ha-1 or more of grain (μj) was set as the standard value and the standard deviation (σj) was set as the range of variation, the nutrient content in high-yielding rice plants should be μj±1.99σj. Rice leaves were sensitive to the nutrient elements. Heavy nitrogen dressing increased the content of nitrogen in rice plants. Sparse transplanting also increased nitrogen content. Improper application of nitrogen, phosphorus and potassium could affect the nutrient contents and decrease the grain yield.

Contribution and Prospect of Erect Panicle Type to Japonica Super Rice

Over the past 30 years,super rice played an important role in boosting rice yield.The phenotype of erect panicle(EP)architecture controlled by dense and erect panicle 1(dep1)is the typical characteristic of super rice,and the phenotype has been used in rice breeding for nearly a century.In this review,the molecular genetic basis of EP phenotype,and mechanism of how dep1 affects rice carbon,nitrogen metabolism and grain quality(grain shape and taste quality)were discussed.In addition,we discussed the possible improvement strategies of rice yield and quality.This review provides a quick overview of the whole process for rice quality formation,as well as suggestions and ideas for future research on rice quality improvement.

Genetical and Physiological Basis of Plant Type Model of Erect and Large Panicle Japonica Super Rice in Northern China

The historical changes in rice yields across China were explored. The physiological mechanisms and genetic basis of the erect and large panicle super-high-yield plant type model for breeding japonica super rice were analyzed mainly on the panicle type, number of large vascular bundles （LVB） in the panicle neck, and the panicle type index （PTI）. In the production point of view, we suggested that, for the breeding of super-high-yield japonica rice, the erect panicle types with more LVB numbers in the panicle neck and superior upper grains in the secondary branches would be the key factors. The information has potential significance in the rice breeding and productivity not only in China but also throughout the rice production areas of the world.

Improving grain appearance of erect-panicle japonica rice cultivars by introgression of the null gs9 allele

The panicle architecture and grain size of rice affect not only grain yield but also grain quality,especially grain appearance.The erect-panicle(EP)trait controlled by the qpe9-1/dep1 allele has been widely used in high-yielding japonica rice breeding,but usually accompanied with moderate appearance of milled rice.The null gs9 allele shows a good potential for improving grain shape and appearance.However,GS9 and qPE9-1/DEP1 loci are tightly linked,and their interaction is unclear,which obviously restricts their utilization in modern rice breeding.In the present study,comparative analyses of protein and mRNA levels revealed that GS9 and qPE9-1 function independently.Three nearisogenic lines(NILs)carrying various allelic combinations of these two loci,NIL(gs9/qpe9-1),NIL(GS9/qPE9-1)and NIL(gs9/qPE9-1),in the EP japonica cultivar 2661(GS9/qpe9-1)background were developed for genetic interaction analysis.GS9 and qPE9-1 had additive effects on determining grain size,and the null gs9 allele could decrease grain chalkiness and improve grain appearance without affecting plant and panicle architecture in EP japonica cultivars.Additionally,introgression lines(ILs)developed in another released EP japonica cultivar Wuyujing 27(WYJ27)background showed the same additive effect and the feasibility of utilizing the gs9 allele to improve grain appearance quality in high-yielding EP cultivars.This study provides an effective strategy for rice breeders to improve rice grain appearance in EP japonica and related cultivars.

Effects of Aphelenchoides besseyi Infection on Yield Traits of Different Rice Varieties (Lines)

[Objective] This study aimed to reveal the effects of Aphelenchoides besseyi infection on different rice varieties（lines）.[Method] By field observation and indoor phenotypic investigation,four conventional japonica rice varieties（lines） and japonica rice restorer line R161 under natural onset conditions were observed and analyzed.[Result] After being infected by A.besseyi,different rice varieties（lines）exhibited various symptoms.Specifically,Ning 1707,Ning 1818,Zhendao 88 and Nanjing 9108 had withered leaf tips and exhibited the symptoms of ＂small grains and erect panicles＂;japonica rice restorer line R161 only had withered leaf tips without symptoms of ＂small grains and erect panicles＂,and the withering symptoms occurred in flag leaf tip,whole flag leaf and top second leaf,respectively.After being infected by A.besseyi,all the experimental materials could sprout normally,but their plant height,panicle length,seed-setting rate and 1 000-grain weight were affected to varying degrees.In addition,after being infected by A.besseyi,various symptomatic tissues of R161 exerted different effects on rice yield.Especially,panicles with withered and twisted whole flag leaf were most affected.[Conclusion] This study provided the basis for further exploration of the damages of A.besseyi infection to rice and development of corresponding control measures.

Introgression of qPE9-1 allele,conferring the panicle erectness,leads to the decrease of grain yield per plant in japonica rice(Oryza sativa L.)

Panicle architecture is closely related to yield formation. The qPE9-1 gene has been proved to be widely used in high-yield rice cultivar developments, conferring erect panicle character in japonica rice. Recently, qPE9-1 has been successfully cloned; however, the genetic effect on grain yield per plant of the erect panicle allele qPE9-1 is controversial yet. In the present study, a drooping panicle parent Nongken 57, carrying qpe9-1 allele, was used as recurrent parent to successively backcross to a typical erect panicle line from the double haploid （DH） population （Wuyunjing 8/Nongken 57）, which was previously shown to carry qPE9-1 allele. Thus a pair of near-isogenic lines （NILs） was developed. The comparison of agronomic traits between the NILs showed that, when qpe9-1 was replaced by qPE9-1, the panicle architecture was changed from drooping to erect; moreover, the panicle length, plant height, 1000-grain weight and the tillers were significantly decreased, consequently resulting in the dramatic decrease of grain yield per plant by 30%. Therefore, we concluded that the qPE9-1 was a major factor controlling panicle architecture, and qPE9-1 had pleiotropic nature, with negative effects on grain yield per plant. This result strongly suggests that the erect panicle allele qPEg-1 should be used together with other favorable genes in the high-yield breeding practice. In addition, the effect of qPE9-1 on eating and cooking quality was also discussed in the present study.

Dynamic Development of Aphelenchoides besseyi on Rice Plant by Artificial Inoculation in the Greenhouse

A study was done on the relationship between Aphelenchoides besseyi and the symptoms of small grains and erect panicles. This is an important rice disease in Jiangsu Province, China. A. besseyi was extracted from small grains and erect panicles and cultured artificially, and then inoculated into bud and leaf sheaths of seedlings of two Oryza sativa cultivars, namely Zhendao 2 and Wuyunjing 7 in a greenhouse. The effect on rice growth, in particular the small grains and erect panicles, was revealed by the extent of the disease, seed expansion stages, nematode load, and nematode mortality. In contrast to healthy seedlings, the height, length, and the numbers of spikelets of unhealthy panicles of Zhendao 2 were decreased by 6.7, 16.4, and 13.5%, respectively. Before anthesis, nematodes were attracted to the leaf sheath and apical meristem, nematode load increased by 40%; after anthesis, nematodes occurred in spikelets principally and the number increased by 90.8%. The percentages of infected seeds and nematode load were highest in plump seeds and lowest in empty seeds. Nematode mortality on grain with normal endosperm was lower than seeds with abnormal endosperm. Results indicated that A. besseyi was the pathogen in rice with the symptoms of small grains and erect panicles. Wuyunjing 7 manifests only the small grains and erect panicles symptoms and not the symptoms of leaf white-tip. These symptoms of small grains and erect panicles are new symptom records for the disease caused by A. besseyi on rice.

Effects of Row-Spacing on Canopy Structure and Yield in Different Plant Type Rice Cultivars

Two japonica rice varieties, Longjing 20 （more tillers and curved panicle type, MCP） and Longjing 21 （few tillers and half erect panicle type, FEP）, were used to study the effects of row-spacing on canopy structure, morphological characteristics and yield. The results showed that the percentage of productive tiller reduced first, and increased afterwards as row-spacing increasing. The relationship between row spacing and the percentage of productive tiller fitted a quadratic regression. The effects of row spacing on leaf area index （LAI） at later tillering stage and the highest stem number per square meter also followed a quadratic regression relationship with increasing first and then reducing. The effects of row-spacing on primary branch were larger than the secondary branch in Longjing 20. However, the trend in Longjing 21 was opposite. The relationship between row spacing and seed setting rate of the secondary branch or panicle was negatively correlated. An extreme significant negative correlation was obtained between seed setting rate of secondary branch in Longjing 20. There was no significant positive correlation between row-spacing and yield in Longjing 20 （R2=0.68）. However, the negative correlation between row-spacing and yield of Longjing 21 was extremely significant （R2=–0.96＊＊）. The canopy structure of MCP was more sensitive to row-spacing. The positive correlation between row spacing and the length of the flag leaf （R2=0.89＊＊）, the width of the flag leaf （R2=0.85＊）, the length of the last internode （R2=0.85＊）, the length of the last 2nd internode （R2=0.96＊＊） or the length of the panicle （R2=0.91＊＊） was significant or extremely significant in Longjing 20, but not in Longjing 21. The wider row-spacing promoted the accumulation of the dry matter of panicle, stem and leaf and the yield formation in MCP. The best row-spacing in Longjing 20 was 30 cm. For Longjing 21, the narrower row-spacing was better. The best row-spacing of it was 21 cm. These results suggested that improved the population environment of MCP or the utilization of the free space in the field of FEP could be reached either by wider row-spacing or narrow row-spacing.