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Background: Changing climate is causing erratic rainfall and prolonged drought periods, thus posing serious threats to crop productivity. Owing to severity of drought events, it is imperative to take proactive measures to enhance the resilience of drought sensitive crops like rice. Therefore, the present study was carried out to improve the drought stress tolerance in rice through gamma amino butyric acid (GABA) application., Methods: The experiment was included four GABA concentrations i.e., 0 mM as control, 1 mM, 1.5 mM, and 2 mM, two water levels i.e., 100% and 50% field capacity (referred as FC100 for well-watered and FC50 for drought conditions, respectively), and two fragrant rice cultivars i.e., Super Basmati and Basmati-515., Results: The findings unveiled a comprehensive improvement in various parameters with GABA application in fragrant rice under both well-watered (FC100) and water-limited (FC50) conditions, compared to the control. Specifically, GABA induced enhancements were observed in plant height, root length, fresh weight, dry weight, total soluble protein content, and total free amino acid content across both cultivars. Moreover, GABA application significantly improved peroxidase (POD) and catalase (CAT) enzyme activities, alongside elevating anthocyanin levels, while concurrently reducing H 2 O 2 contents in both FC100 and FC50 treatments. Furthermore, the positive impact of GABA extended to morphological traits, with notable increases in panicle length, total tillers and productive tillers per hill, branch and grain numbers per panicle, and 1000-grain weight for Super Basmati and Basmati 515 cultivars under both water regimes, compared to Ck. Similarly, the grain yield increased by 31.01% and 27.32% under FC100 and 36.85% and 27.71% under FC50 in Super Basmati and Basmati-515, respectively, in response to GABA application, compared to Ck. Additionally, principal component analysis (PCA) revealed significant variances attributed to Dim1 and Dim2, with 86.1% and 4.0% of the variance, respectively, across three bi-plots encompassing rice cultivars, water levels, and GABA treatments. Notably, all tested indices, except for H 2 O 2 and non-productive tillers per hill, exhibited positive correlations amongst themselves and with rice yield, further emphasizing the beneficial effects of GABA application on fragrant rice under well-watered and drought conditions., Conclusions: GABA significantly improved fragrant rice performance under both well-watered (FC100) and water-limited (FC50) conditions. Moreover, integrating GABA application into rice cultivation practices could not only improve the crop resilience to drought stress but also potentially benefiting the future food and nutritional security globally. However, however; further research is needed to understand the cellular and molecular mechanisms of the functionality of GABA in fragrant rice, particularly under drought conditions., (© 2024. The Author(s).)

Ultrasonic seed (US) treatment could alter seed germination mechanism, however, US induced alterations in morph-physiological attributes and yield of fragrant rice were rarely reported. In the present study, the seeds of three fragrant rice cultivars viz., Xiangyaxiangzhan, Meixiangzhan 2, Ruanhuayou 6100 and one non-fragrant rice viz., Wufengyou 615 were exposed to ultrasonic waves at 20-40 kHz for 1.5 min (T) whereas the seeds without exposure were taken as control (CK). Results showed that US treatment caused minor cracks on seed surface while improved seed germination rate (1.79 %-11.09 %) and 3-indoleacetic acid (IAA) (3.36 %-46.91 %). Furthermore, peroxidase (POD) activity and methionine sulfoxide reductase activity was increased by 29.15 %-74.13 % and 11.26 %-20.87 %, respectively; however, methionine sulfoxide reductase related protein repairing gene MSRA4 was down-regulated by 17.93 % -41.04 % under T, compared to CK. Besides, US treatment also improved soluble protein in flag leaf (0.92 %-40.79 %), photosynthesis (3.37 %-16.46 %), biomass (5.17 %-31.87 %), as well as 2-acetyl-1-pyrroline content (4.77 %-15.48 %) in rice grains. In addition, multivariate analysis showed that the dry weight at the maturity stage were significantly related to the POD, glutathione reductase (GR) activity, IAA, and abscisic acid (ABA) content while germination rate was positively related to the GR activity, ABA content, and yield, but which were negatively related to the IAA and gibberellic acid content., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Author(s). Published by Elsevier B.V. All rights reserved.)

Background: The use of liquid fertilizer is an effective measure to increase rice yield and nitrogen use efficiency. There has been a lack of information regarding the effects on the grain yield, biomass accumulation, and nutrient uptake in late-season indica fragrant rice of split fertilizer application and of nitrogen management in liquid fertilizer application., Results: A 2-year field experiment was carried out during 2019 and 2020 with two fragrant rice cultivars grown under differing fertilizer management treatments. Results showed that the fertilization treatments affected the grain yield, yield components, biomass accumulation, and nutrient accumulation significantly. The mean nitrogen recovery efficiency with liquid fertilizer management was greater than in a control treatment corresponding to a practice commonly used by farmers (H2). The effects of nitrogen metabolism enzymes in the leaves of both rice cultivars were stronger with liquid fertilizer treatments than with H2. Grain yield was positively associated with the effective panicle number, spikelets per panicle, dry matter accumulation, N and K accumulation, and the nitrogen metabolism enzymes., Conclusions: Optimized liquid fertilizer management increases biomass accumulation, nitrogen utilization efficiency, and nitrogen metabolism. It stabilizes yields and increases the economic benefits of late-season indica fragrant rice. © 2023 Society of Chemical Industry., (© 2023 Society of Chemical Industry.)

Rice is a staple food for more than half of the world's population, and rice fragrance is a key quality attribute which is highly desired by consumers and attracts premium prices in the international market. There are around 200 volatile compounds involved in rice fragrance, but 2-acetyl-1-pyrroline (2-AP) has been considered a master regulator of aroma in fragrant rice. Consequently, efforts were made to increase the 2-AP contents in the grain by managing agronomical practices or by using modern functional genomic tools, which successfully converted nonfragrant cultivars to fragrant rice. Furthermore, environmental factors were also reported to influence the 2-AP contents. However, a comprehensive analysis of 2-AP biosynthesis in response to agro-management practices, environmental factors, and the application of functional genomic tools for fragrant rice production was missing. In this Review, we summarize how micro/macronutrients, cultivation practices, amino acid precursors, growth regulators, and environmental factors, such as drought, salinity, light, and temperature, influence the 2-AP biosynthesis to modulate the aroma of fragrant rice. Furthermore, we also summarized the successful conversion of nonfragrant rice cultivars to fragrant rice using modern gene editing tools, such as RNAi, TALENS, and CRISPR-Cas9. Finally, we discussed and highlighted the future perspective and challenges related to the aroma of fragrant rice.

Plant mineral nutrition substantially affects the growth, yield and quality of rice, whereas nitrogen (N) application contributes significantly in this regard. Undoubtedly, N application improves rice aroma biosynthesis; however, the molecular mechanism underlying the regulation of grain 2-acetyl-1-pyrroline (2-AP) biosynthesis in the presence of nitrogen application at the booting stage has remained largely unexplored. The present study examined the effects of three N levels, i.e., 0 g per pot (N0), 0.43 g per pot (N1) and 0.86 g per pot (N2) on intermediates, enzymes and genes involved in 2-AP biosynthesis, as well as on the yield of two fragrant rice cultivars viz , Meixiangzhan2 and Xiangyaxiangzhan. N was additionally applied at the booting stage. The results depicted that the levels of precursor, such as proline, and the activity of enzymes involved in 2-AP biosynthesis, such as Δ1-pyrroline-5-carboxylate synthetase (P5CS) and diamine oxidase (DAO), and P5CS1 gene expression were comparatively higher under N1 than N0 in both fragrant rice cultivars. Moreover, the N2 treatment increased the grain panicle -1 , filled grain percentage and grain yield of both rice cultivars, while the grain yield of Meixiangzhan2 and Xiangyaxiangzhan was increased by 15.87% and 12.09%, respectively, under N2 compared to N1 treatment. Hence, 0.43 g per pot of N showed positive performances in yield and aroma accumulation in fragrant rice and should be further employed in the practice and production for better cultivation in the rice market.

Alternate wetting and drying (AWD) has been recognized as a water-saving technology in rice production systems; however, pre- and post-flowering AWD could induce changes in yield, quality and aroma biosynthesis in fragrant rice. In the present study, two fragrant rice cultivars (Guixiangzhan and Nongxiang-18) were subjected to AWD till soil water potential reached -25 to -30 kPa during vegetative stage (VS), reproductive stage (RS), and both stages (VS + RS). The AWD did not affect net photosynthesis and gas exchange significantly, while malondialdehyde (MDA), H 2 O 2 and electrolyte leakage (EL) were higher than in control plants. The AWD treatments variably affected soluble sugars, proline and protein accumulation as well as the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), ascorbate peroxidase (APX), and reduced glutathione (GSH) contents. Moreover, filled grain percentage and 1000-grain weight in AWD treatments were found to be statistically similar (p > 0.05) to control, except grains panicle -1 under AWD-VS + RS that was reduced by 11% and 14% for Guixiangzhan and Nongxiang-18, respectively. On average, yield and related attributes in Guixiangzhan remained higher than in Nongxiang-18. In addition, the grain aroma volatile (2-acetyl-1-pyrroline, 2-AP) content increased by 8.79%, 14.45%, and 6.87% and 7.95%, 14.02%, and 5.04% under AWD-VS, AWD-RS, and AWD-VS + RS treatments, for Guixiangzhan and Nongxiang-18, respectively. Overall, AWD treatments, either at VS or RS, could promote rice aroma in terms of accumulation of 2AP, which might be linked with enhanced endogenous proline contents (a precursor for 2AP biosynthesis) without any severe consequences on rice yield and quality., (© 2022 Scandinavian Plant Physiology Society.)

Application of hydrogen-rich water (HRW) could improve plant growth under stress conditions; however, its effects on early growth and related physiological functions of fragrant rice under cadmium (Cd) and lead (Pb) toxicity are unknown. The present study was composed of two HRW treatments, i.e., H0: without H 2 and HRW: with 500ppb H 2 gas, three treatments with two metals, i.e., M0: without heavy metal, Cd: 75 μmol l -1 of Cd, Pb: 750 μmol l -1 of Pb applied to two popular fragrant rice cultivars, i.e., Yuxiangyouzhan and Xiangyaxiangzhan. The growth and physio-biochemical attributes of fragrant rice at 5 and 10 days after treatment (DAT) were assessed. Results showed that HRW treatment promoted the growth of rice seedlings in terms of increased dry biomass of shoot, root, and the whole seedling at 10 DAT. Moreover, HRW also improved early growth of fragrant rice under Pb stress with substantial increase in fresh and dry weight of roots and the whole seedling at 5 and 10 DAT. Application of HRW slightly alleviated the root inhibition caused by Cd toxicity in rice seedlings at 10 DAT. In addition, antioxidant activities, i.e., catalase (CAT) and peroxidase (POD), were increased with HRW application at 10 DAT while decreased root Pb and Cd contents of both rice cultivars. Overall, HRW alleviated the inhibitory effects of Cd and Pb toxicity by regulating the antioxidant defense response in growing rice plants., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The involvement of zinc (Zn) in terms of aroma formation has been rarely investigated. This study shows that the regulation of 2-acetyl-1-pyrroline (2AP) biosynthesis was evaluated in two different rice cultivars under foliar Zn application. The results showed that the 2AP and Zn contents in leaves and grains were improved substantially under foliar Zn application. The 2AP content was positively related to the expression P5CS2 gene, contents of proline, 1-pyrroline, and Δ1-pyrroline-5-carboxylate (P5C), and the activity of pyrroline-5-carboxylate synthase (P5CS) under Zn application in fragrant rice. Multiple transcription factors (TFs) were differently expressed, such as bZIPs , NACs , and MYBs , to play a role under Zn treatments in fragrant rice, suggesting the crucial role of 46 differently expressed TFs in 2AP improvements in fragrant rice. Furthermore, this study showed that the optimal foliar Zn application at a concentration of 30 mg L -1 could increase the 2AP content of aromatic rice and keep the yield stable or increase the yield. TFs were involved in regulating to promote the 2AP formation in aromatic rice under the foliar Zn application. However, the relationship between 2AP biosynthesis pathway genes and TFs in fragrant rice remains to be further studied.

Rice cultivation under cadmium (Cd) contaminated soil often results in reduced growth with excess grain Cd concentrations. A pot experiment was conducted to assess the potential of ultrasonic seed treatment to alleviate Cd stress in rice. Seeds of two aromatic rice cultivars i.e., Xiangyaxiangzhan and Meixiangzhan 2 and two non-aromatic rice cultivars i.e., Huahang 31 and Guangyan 1 were exposed to ultrasonic waves for 1.5 min in 20-40 KHz mixing frequency. The experimental treatments were comprised of untreated seeds (U0) and ultrasonic treated seeds (U1) transplanted in un-contaminated soil (H0) and Cd-contaminated soil (H1). Results revealed that Cd contents and Cd accumulation in grain in U1 were 33.33-42.31% and 12.86-57.58% lower than U0 for fragrant rice cultivars under H1. Meanwhile, biomass production was higher in U1 than U0 under H0 and better yield was assessed in U1 for all cultivars under H1. The activity of peroxidase (POD) in flag leaves was increased by 8.28-115.65% for all cultivars while malondialdehyde (MDA) contents were significantly decreased in U1 compared with U0 under H0. Conclusively, ultrasonic treatment modulated Cd distribution and accumulation in different parts while improved physiological performance as well as yield and grain quality of rice under Cd contaminated conditions., (Copyright © 2021. Published by Elsevier Inc.)

Cloudy weather with low light is more common during rice growing season of South China which often leads reduced yield and aroma formations in fragrant rice. However, exogenous γ-aminobutyric acid (GABA) application could enhance the 2-acetyl-1-pyrroline (2AP) accumulations and yield of fragrant rice under low light conditions. Field and pot experiments were conducted with three fragrant rice cultivars i.e., Basmati and Yuxiangyouzhan (indica), and Yungengyou 14 (japonica) that were grown under three different treatments i.e., normal light + GABA 0 mg L -1 (CK), low light + GABA 0 mg L -1 (T1), and low light + GABA 250 mg L -1 (T2). The results revealed that the grain 2AP contents were increased by 14.67-34.83% and up to 29.34% under T1 and T2 treatments in pot and field experiments, respectively, as compared with CK. The T1 and T2 treatments improved aroma owing to regulation in the accumulation of micronutrients i.e., Na, Mn, and Fe and enzyme activities involved in 2AP biosynthesis. The grain yield was substantially reduced in T1 as compared with T2 treatment for all rice cultivars. On the other hand, GABA application improved the grain yield under low light conditions by regulating the plant growth, and related physiological and biochemical attributes in all rice cultivars. Thus, GABA could balance low light-induced 2AP content and grain yield by modulating morphological and yield related attributes as well as physio-biochemical responses of fragrant rice., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

Background: Cadmium (Cd) is amongst the most toxic heavy metals that severely affects crop growth, whereas application of nanoparticles (NPs) to negate the toxic effects of heavy metals could be an effective management approach. In the present study, the seeds of two fragrant rice varieties i.e., Yuxiangyouzhan and Xiangyaxiangzhan under normal and Cd stress conditions i.e., 0 and 100 mg L - 1 applied with four levels of ZnO NPs i.e., 0, 25, 50, and 100 mg L - 1 ., Results: Seed priming with ZnO NPs had no significant effect on the seed germination (p > 0.05) however, it substantially improved the seedling growth and other related physiological attributes under the Cd stress. The mean fresh weight of the shoot, and whole seedling was increased by 16.92-27.88% and by 16.92-27.88% after ZnO NPs application. The root fresh weight, root-shoot length was also substantially improved under ZnO NPs treatment. Moreover, application of ZnO NPs induced modulations in physiological and biochemical attributes e.g., the superoxide dismutase (SOD) activity in root and shoot, the peroxidase (POD) activity and metallothionein contents in root were increased under low levels of ZnO NPs. The α-amylase and total amylase activity were improved by ZnO NPs application under Cd Stress. Besides, modulation in Zn concentration and ZnO NPs uptake in the seedling were detected. The metabolomic analysis indicated that various pathways such as alanine, aspartate and glutamate metabolism, phenylpropanoid biosynthesis, and taurine and hypotaurine metabolism were possibly important for rice response to ZnO NPs and Cd., Conclusion: Overall, application of ZnO NPs substantially improved the early growth and related physio-biochemical attributes in rice. Our findings provide new insights regarding the effects of ZnO NPs on seed germination, and early growth of rice, and its potential applications in developing crop resilience against Cd contaminated soils.

The cytochrome P450 (CYP) is a large and complex eukaryotic gene superfamily with enzymatic activities involved in several physiological and regulatory processes. As an objective, an in-silico genome-wide DNA methylation (5mC) analysis was performed in rice ( Oryza sativa cv. Zhonghua11), and the epigenetic role of CYPs in two abiotic stresses was observed. Being a stable representative mark, DNA-methylation alters the gene expression under stressful environmental conditions. Rice plants under salinity and drought stresses were analyzed through MeDIP-chip hybridization, and 14 unique genes of the CYP family were identified in the rice genome with varying degrees of methylation. The gene structure, promoter sequences, and phylogenetic analysis were performed. Furthermore, the responses of CYPs to various abiotic stresses, including salinity, drought, and cold were revealed. Similarly, the expression profile of potential CYPs was also investigated under various phytohormone stresses, which revealed the potential involvement of CYPs to hormone regulations. Overall, the current study provides evidence for CYP's stress regulation and fundamental for further characterization and understanding their epigenetic roles in gene expression regulation and environmental stress regulation in higher plants.

Salinity negatively effects the growth and productivity of crop plants; however, the effects of hydrogen rich water (HRW) on the early growth of fragrant rice under salinity stress are rarely investigated. In present study, two HRW treatments: foliar application (F-HRW) and irrigation (I-HRW) were applied on the two fragrant rice cultivars, Yuxiangyouzhan and Xiangyaxiangzhan, grown under normal and salt stress conditions, i.e., 0 and 150 mmol NaCl L -1 , respectively. Plants without HRW application were grown as control (CK). Results showed that the dry weight per unit plant height (mg cm -1 ) was increased by 12.6% and 23.0% in F-HRW and I-HRW, respectively under salt stress as compared with CK. Application of HRW, regardless of the application method, modulated the antioxidant activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) while reduced malondialdehyde (MDA) contents under salt stress. Moreover, significant and positive relations were observed among total dry weight and shoot dry weight, dry weight per unit plant height, SOD and CAT activity in root. Overall, F-HRW application modulated the early growth and related physiological attributes in fragrant rice under salt stress whereas I-HRW was found to mitigate salt stress. Novelty statement : Involvement of endogenous H 2 in plants for regulating various physiological functions is of great importance to stimulate and/or activate the antioxidant defense responses against oxidative stress; however, there is a lack of research in this aspect. The present study investigated the effects of hydrogen rich water (HRW) on the growth and physiological attributes of two fragrant rice cultivars grown under salt-stress. It was noteworthy to find that application of HRW either foliar application or irrigation improved the morphological characters, i.e., dry weight per unit plant height and enhanced the activities of antioxidants, i.e., peroxidase, superoxide dismutase and catalase whilst decreased the malonaldehyde content. Overall, the application of HRW modulates plant growth and physiological attributes in fragrant rice cultivars under salt-stress conditions. This study will be helpful in improving the early growth and/or stand establishment of fragrant rice nursery under saline conditions.

Increasing evidence shows that cadmium (Cd) toxicity causes severe perturbations on growth performance, physio-biochemical and molecular processes in crop plants. Molybdenum (Mo), an essential trace element, plays key roles in oxidative stress tolerance of higher plants. Hence, the present study has been conducted to investigate the possible role of Mo in alleviating Cd-induced inhibitions in two fragrant rice cultivars namely Guixiangzhan (GXZ) and Meixiangzhan-2 (MXZ-2). The results revealed that Mo application enhanced the plant dry biomass by 73.24% in GXZ and 58.09% in MXZ-2 under Cd stress conditions, suggesting that Mo supplementation alleviated Cd-induced toxicity effects in fragrant rice. The enhanced Cd-tolerance in fragrant rice plants prompted by Mo application could be ascribed to its ability to regulate Cd uptake and reduce Cd-induced oxidative stress as evident by lower hydrogen peroxide levels, electrolyte leakage and malondialdehyde contents in Cd-stressed plants. The ameliorative role of Mo against Cd-toxicity also reflected through its protection to the photosynthetic pigments, proline and soluble protein. Mo also induced antioxidant defense systems via maintaining higher contents of glutathione and ascorbate as well as enhancing the ROS-detoxifying enzymes such as catalase, peroxidase, superoxide dismutase and ascorbate peroxidase activities and up-regulating transcript abundance in both fragrant rice cultivars under Cd stress. Conclusively, Mo-mediated modulation of Cd toxicity in fragrant rice was through restricting Cd uptake, maintaining photosynthetic performance and alleviating oxidative damages via the strong anti-oxidative defense systems; however, GXZ cultivar is comparatively more Cd tolerant and Mo-efficient as evident from the less growth inhibition and biomass reduction as well as enhanced Mo-induced Cd stress tolerance and less oxidative damage than MXZ-2 fragrant rice cultivar.

Nitrate reductase is a nitric oxide (NO) induced enzyme in plants, NO acts as a signaling molecule under ZnO NPs-induced stress whereas melatonin (N-acetyl-5-methoxytryptamine) could improve morpho-physiological attributes of plants under adverse conditions. In present study, seedlings of two rice genotypes differed regarding nitrate reductase activities i.e., transgenic 'NR' and wild type 'WT' were applied with two melatonin levels i.e., 0, 10 μΜ regarded as M0, M10, respectively and three levels of ZnO NPs i.e., 0, 50, 500 mg L -1 regarded as ZnO NPs0, ZnO NPs50 and ZnO NPs500, respectively. Results revealed that melatonin application substantially increased the dry biomass accumulation of both rice genotypes under all ZnO NPs levels. The root growth, mineral absorption as well as the antioxidant responses were also improved by melatonin application under ZnO NPs stress. The interactive effects of melatonin and genotype on plant growth, antioxidant responses and mineral contents i.e., Zn, Na, Fe and Mn were also found significant under different ZnO NPs stress. Furthermore, total plant dry weight was significantly correlated with the leaf dry weight, root volume, catalase (CAT) activity in leaves, Na accumulation in stem sheath and Fe accumulation in root under both M0 and M10 treatments. Moreover, the comparative transcriptome analysis identified key genes which were responsible for melatonin and NO-induced modulations in plant growth under ZnO NPs stress. Overall, melatonin could improve the morphological growth of the rice plants by modulating root-shoot characteristics, antioxidant activities and mineral uptake in root and shoot of rice., Competing Interests: Declaration of competing interest The authors declare no competing interests., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Fragrant rice is a high-valued quality rice type which is gaining much popularity over the globe due to its better cooking qualities and special aromatic characteristics. Selenium (Se) and silicon (Si) could improve the growth and yield of rice; however, the combine effects of Se and Si (Se-Si treatments) on rice grain quality, aroma and lodging in fragrant rice were rarely investigated. The pot and field experiments were conducted with two fragrant rice cultivars i.e., Xiangyaxiangzhan and Yuxiangyouzhan, grown under three Se levels i.e., 0, 120, and 240 mg kg -1 of soil (for pot experiment) and 0, 300, and 600 kg ha -1 (for field experiment) regarded as LSe, MSe and HSe, respectively and two Si levels i.e., 0 and 60 mg kg -1 of soil (for pot experiment) and 0 and 150 kg ha -1 (for field experiment) regarded as -Si and +Si, respectively. Results depicted that the Se-Si treatments regulated head rice yield, grain yield and yield related traits and the HSe+Si treatment sustainably improved the grain yield and head rice yield by regulating plant growth, antioxidant response and malondialdehyde (MDA) contents in fragrant rice. The Se-Si treatments also improved the grain 2AP contents owing to regulation in the proline, pyrroline-5-carboxylate (P5C) and γ-aminobutyric acid (GABA) contents. Besides, Se-Si treatments also regulated the grain quality attributes and influenced the plant Se contents. Moreover, the Si mitigated Se-induced lodging resulted from changes in the lodging parameters i.e., lodging index, fresh weight per tiller, pushing resistance force, plant height and bending moment. Overall, the Se and Si application improved the grain yield and regulated the dry weight accumulation, antioxidant attributes and quality attributes. Meanwhile, the Si application mitigated the negative effect of Se-induced lodging in fragrant rice., Competing Interests: Declaration of competing interests The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Lead (Pb) affects the growth and productivity of rice negatively through soil-Pb-plant interactions. A pot experiment was conducted to assess the Pb uptake and its distributive pattern in different fragrant rice cultivars i.e., Meixiangzhan-2 (MXZ-2), Xiangyaxiangzhan (XYXZ), Guixiangzhan (GXZ), Basmati-385 (B-385), and Nongxiang-18 (NX-18) and relationship of Pb uptake with grain Pb-contents and rice yield and related traits i.e., productive tillers hill -1 , spikelets panicle -1 , filled grain percentage, and 1000-grain weight. Lead as Pb(NO 3 ) 2 was applied at 0 (control), 400 (Pb1), 800 (Pb2) and 1200 mg kg -1 (Pb3) of soil in solution form. Results showed that all rice cultivars accumulated different concentration of Pb, with the highest in roots and the lowest in grains. The translocation factors (TF) from stems to leaves were higher than root to stems and leaves to ears/grains. The grain Pb contents have significant positive associations with ear Pb contents and TF of Pb from leaves to ear at reproductive stage. Moreover, relative changes (RC) in grain yields have significant correlations with the RC in some yield attributes i.e., productive tillers per hill, spikelets per panicle, and filled grain percentage under Pb toxicity. Overall, grain Pb contents are affected mainly by the transference of leaves Pb contents to ears/grains (an index to determine final grain Pb contents) whereas RC in rice yields suggested its relationships with some agronomic attributes of fragrant rice under Pb stress., Competing Interests: Declaration of competing interest No conflict of interest exist among authors., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Cadmium (Cd) toxicity has detrimental effects on plant metabolism and yield formation. This study examined the effects of Cd stress in rice and the possible role of calcium (Ca) in mitigating oxidative damage caused by Cd in two fragrant rice cultivars, i.e., Guixiangzhan and Meixiangzhan 2. The experimental treatments were composed of various Ca and Cd levels as individual, i.e., Ca at 2.5 and 5.0 mg/kg soil (Ca1 and Ca2, respectively), Cd at 50 and 100 mg/kg soil (Cd50 and Cd100, respectively), and combined, i.e., Ca1+Cd50, Ca1+Cd100, Ca2+Cd50, and Ca2+Cd100. Plants without Ca and Cd application were taken as control (CK). Results showed that Cd stress led to a substantial decline in the photosynthetic pigments, i.e., Chl a, Chl b, and carotenoids, while enhanced oxidative damage in terms of increased levels of hydrogen peroxide (H 2 O 2 ) and malondialdehyde (MDA) and electrolyte leakage (EL) in both rice cultivars. Moreover, Cd stress hampered the activities of enzymatic antioxidants, i.e., superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), with lowest antioxidant activities were recorded at Cd100. The overall trend (lowest to highest) for antioxidant activities across treatments was recorded as Cd100 < Ca2+Cd100 < Cd50 < Ca1+Cd100 < CK < Ca1 < Ca1+Cd50 < Ca2+Cd50 < Ca2. Similarly, Ca amendment improved the proline, soluble protein, and soluble sugar contents in both rice cultivars under Cd stress condition. Comparing Ca2 with CK, the yield and related components, i.e., number of panicles, spikelets per panicle, seed setting rate, 1000 grain weight, and grain yield, were found to increase by 13.08, 2.39, 4.03, 5.86, and 27.53% for Guixiangzhan and 16.48, 5.19, 6.87, 15.44, and 51.16% for Meixiangzhan, respectively. Furthermore, Cd contents in roots, stems, leaves, and grains increased with increased Cd concentration applied and reduced with Ca amendment. The Cd contents in grains for all Ca+Cd levels are statistically at par with each other and significantly lower (P < 0.05) than those for individual Cd application. Hence, Ca amendment can be an appropriate approach to ameliorate the toxic effects of Cd in crops grown under Cd-contaminated soils.

Background: A mixed-cropping system that enhances farmland biodiversity has the potential to improve grain yield and quality; however, the impacts of growing different rice cultivars simultaneously has been rarely investigated. In the present study, five popular rice cultivars were selected and ten mixture combinations were made according to the growth period, plant height, grain yield and quality, and pest and disease resistance. Seedlings of the five cultivars and ten mixture combinations (mixed-sowing of the seeds in an equal ratio, then mixed-transplanting and finally mixed-harvesting) were grown in plastic pots in a glasshouse during the early and late growing seasons in 2016., Results: Compared with the mono-cropping systems in the early and late growing seasons in 2016 (paired t-test), the mixed-cropping systems increased the rice leaf photosynthetic rate, soil plant analysis development (SPAD) index and total aboveground dry weight. Moreover, mixed-cropping systems improved the number of spikelets per panicle, seed-setting rate, and grain weight per pot and harvest index by 19.52% and 5.77%, 8.53% and 4.41%, 8.31% and 4.61%, and 10.26% and 6.98% in the early and late growing seasons, respectively. In addition, mixed-cropping systems reduced chalky rice rate and chalkiness degree by 33.12% and 43.42% and by 30.11% and 48.13% in the early and late growing seasons, respectively., Conclusion: The SPAD indexes and photosynthetic rates enhanced at physiology maturity in mixed-cropping systems may result in higher grain yield and better grain quality. In general, it was found that mixed-cropping with different rice cultivars has the potential for increasing grain yield and improving grain quality. © 2018 Society of Chemical Industry., (© 2018 Society of Chemical Industry.)

Factors affecting rice aroma biosynthesis have been well documented previously, however the molecular mechanism lies behind the regulations in grain 2-acetyl-1-pyrroline (2AP) biosynthesis under alternate wetting and drying (AWD) remained largely unexplored. Present study investigated the effects of three irrigation regimes i.e., conventional irrigation (CI), alternate wetting and moderate drying (WMD), and alternate wetting and severe drying (WSD) on the yield, quality traits, intermediates, enzyme activities and genes involved in 2-acetyl-1-pyrroline biosynthesis in two fragrant rice cultivars viz, Meixiangzhan2 and Xiangyaxiangzhan. Results revealed that the levels of intermediates such as Δ1-pyrroline-5-carboxylate (P5C) and Δ1-pyrroline, and the activity of enzymes such as proline dehydrogenase (PRODH), Δ1-pyrroline-5-carboxylate synthetase (P5CS), diamine oxidase (DAO), and gene expressions of PRODH, P5CS2 and DAO were comparatively higher under AWD than CI in both aromatic rice cultivars. The levels of gamma-aminobutyric acid (GABA), betaine aldehyde dehydrogenase 2 (BADH2) and BADH2 gene were lower that together led to enhanced 2-AP contents in rice grains. Moreover, WMD irrigation improved yield and yield characters, while WSD irrigation reduced yield and quality traits of rice. Overall, up-regulation of P5C and Δ1-pyrroline and down-regulation of GABA under AWD treatments resulted in enhanced 2AP biosynthesis in both rice cultivars. Evaluation and adoption of AWD (within safe limits) at field level could be an alternative option to conventional flooded rice to get better yield and quality., (Copyright © 2018. Published by Elsevier Masson SAS.)

Lead (Pb) is a highly toxic metal that damages normal plant metabolism and often results in reduced crop growth and yield. To measure whether or not ultrasonic seed (US) treatment alleviates Pb stress in rice, the seeds of two rice cultivars, i.e., Guangyan1 and Huahang31, were exposed to 30 min of ultrasonic vibration and then the seedlings were transplanted to Pb-contaminated soil. Results revealed that the Pb contents in roots, stems, leaves, panicles (at heading), and brown rice (at maturity) were lower in US treatment than control (CK; without US treatment); the trend for accumulation of Pb in different plant parts was recorded as: root ˃ stems ˃ leaves ˃ panicles ˃ brown rice in both rice cultivars. Overall, the Pb contents in the brown rice of Huahang31 were higher than those in Guangyan1. Moreover, the contents of proline and soluble protein as well as the activities of superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX) were significantly higher while the malondialdehyde (MDA) contents were significantly reduced in US treatment compared to CK. The effective panicle number, seed set percentage, grains per panicle, 1000-grain weight, and grain yield were increased by 8.9%, 5.3%, 6.8%, 4.4%, and 26.6% in Guangyan1 and 3.2%, 5.7%, 5.7%, 5.0%, and 24.2% in Huahang31, respectively in US treatment compared to CK. Hence, seed treatment with ultrasonic waves could improve rice performance and reduce brown rice Pb accumulation under Pb-polluted soils.

Rice cultivation in lead (Pb) polluted soils often leads to high Pb contents in rice grains. The present study investigated the dynamics of Pb uptake under different water regimes in two fragrant rice cultivars i.e., Guixiangzhan and Nongxiang-18. Results revealed that water dynamics regulated the antioxidant activities in both rice cultivars under Pb toxicity. Compared to continuous ponding (CP), taken as control, alternate wetting and drying (AWD) reduced the Pb contents in roots, stems, leaves, and grains up to 17%, 41%, 22%, and 52% in Guixiangzhan and 23%, 19%, 17%, and 37% in Nongxiang-18, respectively. Furthermore, AWD-treatments reduced paddy yield from 11% to 21% in Guixiangzhan and 11-33% in Nongxiang-18 under Pb toxicity. In conclusion, Pb loadings in fragrant rice can be regulated by effective water management and/or by controlling irrigation water at different growth stages. Special control measures or management is required to cultivate the rice in metal(loid)s polluted soils., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Present study was planned to assess the yield and quality responses, plant physio-biochemical characters and Pb distribution pattern in two aromatic rice cultivars viz., Guixiangzhan (GXZ) and Nongxiang-18 (NX-18) under four different Pb-levels viz., control (0), low (400), medium (800) and high (1200) mg kg -1 of soil. Results revealed that Pb toxicity increased H 2 O 2 , lipid peroxidation and electrolyte leakage while inhibited photosynthetic pigments production, but such increment was higher in NX-18 than GXZ. Furthermore, Pb toxicity variably affected protein, proline and soluble sugars and the activities of enzymatic antioxidants viz., superoxide dismutase (SOD), peroxidases (POD), catalases (CAT) and ascorbate peroxidases (APX) and non-enzymatic anti-oxidants viz., reduced glutathione (GSH) and oxidized glutathione (GSSG) in both rice cultivars. The reductions in osmolyte accumulation and antioxidant activities were more severe in NX-18 than GXZ. Pb toxicity severely reduced yield and quality related attributes and plant biomass accumulation; however the reductions were more apparent NX-18 than GXZ. Furthermore, NX-18 accumulated less Pb proportions in roots (∼84%), and transferred more towards shoot, leaves, ears (at panicle heading (PH)) and grains (at maturity (MAT)) than GXZ (∼91% root Pb proportions). Hence, the ability of GXZ to retain higher Pb contents in roots and less towards upper plant parts, higher osmolyte accumulation and antioxidant activities and less yield reduction as compare to NX-18 might be an adaptive response of GXZ under Pb toxicity., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Lead (Pb) affects plant growth and its related physio-biochemical functions negatively. The present study investigated the responses of five different fragrant rice cultivars viz., Meixiangzhan (MXZ-2), Xiangyaxiangzhan (XYXZ), Guixiangzhan (GXZ), Basmati-385 (B-385), and Nongxiang-18 (NX-18) to four different Pb concentrations viz., 0, 400, 800 and 1200 μM. Results depicted that Pb toxicity significantly (P < 0.05) reduced the plant height, tillering ability and biomass accumulation by causing oxidative damage to rice plants; nonetheless, a significant variation was found in the sensitivity of rice cultivars to Pb toxicity. Soluble sugars increased significantly only at 1200 μM in GXZ and 800 μM in B-385, whilst the maximum reductions in protein contents were observed at 1200 μM Pb for all rice cultivars. Proline contents were reduced for XYXZ and NX-18 at Pb1200 μM. Activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) as well as reduced glutathione (GSH) and oxidized glutathione (GSSG) showed differential behavior among Pb treatments and rice cultivars. Among rice cultivars, GXZ showed better antioxidative defense system under Pb toxicity compared with all other cultivars. For all rice cultivars, the trend for Pb accumulation was recorded as: roots > stems > leaves. Furthermore, significant but negative correlations among Pb uptake and plant height (r = -0.79), tillers per plant (r = -0.91) and plant dry biomass (r = -0.81) were recorded for all rice cultivars whereas the values of translocation factor (TF) from stems to leaves were higher than roots to stems. In sum, Pb reduced the early growth and caused physio-biochemical changes in all rice cultivars, nonetheless, GXZ proved better able to tolerate Pb stress than all other rice cultivars under study., (Copyright © 2017 Elsevier Masson SAS. All rights reserved.)

Micro-nutrient application is essential for normal plant growth while a little is known about manganese (Mn)-induced regulations in morpho-physiological attributes, aroma formation and enzyme involved in 2-acetyl-1-pyrroline (2-AP) biosynthesis in aromatic rice. Present study aimed to examine the influence of four levels of Mn i.e., Mn1 (100 mg MnSO4 pot(-1)), Mn2 (150 mg MnSO4 pot(-1)), Mn3 (200 mg MnSO4 pot(-1)), and Mn4 (250 mg MnSO4 pot(-1)) on the growth, yield formation, quality characters, rice aroma and enzyme involved in 2-acetyl-1-pyrroline biosynthesis in two fragrant rice cultivars i.e., Meixiangzhan and Nongxiang 18. Pots without Mn application were served as control (Ck). Each pot contained 15 kg of soil. Effects on agronomic characters, quality attributes, 2-AP contents and enzymes involved in 2-AP biosynthesis have been studied in early and late season rice. Results depicted that Mn improved rice growth, yield and related characters, and some quality attributes significantly. It further up-regulated proline, pyrroline-5-carboxylic acid (P5C) (precursors of 2-AP), soluble proteins and activities of proline dehydrogenase (ProDH), Δ(1) pyrroline-5-carboxylic acid synthetase (P5CS) ornithine aminotransferase (OAT) that led to enhanced 2-AP production in rice grains. Moreover, higher Mn levels resulted in increased grain Mn contents in both rice cultivars. Along with growth and yield improvement, Mn application significantly improved rice aromatic contents. Overall, Nongxiang 18 accumulated more 2-AP contents than Meixiangzhan in both seasons under Mn application. This study further explored the importance of Mn in rice aroma formation and signifies that micro-nutrients can play significant roles in rice aroma synthesis; however, intensive studies at molecular levels are still needed to understand the exact mechanisms of Mn to improve rice aroma formation., (Copyright © 2016 Elsevier Masson SAS. All rights reserved.)

Aromatic rice is highly prized by consumers worldwide due to its special aromatic character. 2-acetyl-1-pyrroline (2-AP) is considered to be the single most important volatile compound responsible for aroma in aromatic rice. The present study demonstrated the effects of 2-AP, zinc (Zn) and lanthanum (La) on the 2-AP concentration of detached aromatic rice panicles in vitro. Detached panicles from three well-known aromatic cultivars, Guixiangzhan, Pin14, and Pin 15, were cultured separately in basic culture medium supplemented with 2-AP, Zn and La, and 2-AP concentrations were assessed at 7 and 14 days after culture (DAC). The results show that supplementation of 2-AP, Zn and La in the basic culture medium significantly increases the accumulation of proline. 2-AP concentration and the activity of proline dehydrogenase (ProDH) were also increased in rice grains. Zn concentrations were also found to be higher when Zn was added to the basic culture medium, and La concentrations in grains were too low to be measured. Additionally, grain 2-AP concentrations were significantly and positively correlated with proline concentrations, ProDH activities in grains and 2-AP in culture medium. In summary, higher grain 2-AP concentrations might be due to Zn- and La-induced increases in proline concentrations and ProDH activities, as well as the direct uptake and transportation of 2-AP from the culture medium. Furthermore, application of both Zn and La might be helpful for improving aroma formation in rice. However, interactions of both these elements with the complex process of 2-AP formation remain to be explored.

Lead (Pb) is a major environmental pollutant that affects plant morpho-physiological and biochemical attributes. Its higher levels in the environment are not only toxic to human beings but also harmful for plants and soil microbes. We have reviewed the uptake, translocation, and accumulation mechanisms of Pb and its toxic effects on germination, growth, yield, nutrient relation, photosynthesis, respiration, oxidative damage, and antioxidant defense system of rice. Lead toxicity hampers rice germination, root/shoot length, growth, and final yield. It reduces nutrient uptake through roots, disrupts chloroplastic ultrastructure and cell membrane permeability, induces alterations in leaves respiratory activities, produces reactive oxygen species (ROS), and triggers some enzyme and non-enzymatic antioxidants (as defense to oxidative damage). In the end, biochar amendments and phytoremediation technologies have been proposed as soil remediation approaches for Pb tainted soils.