Search Results (1,125)

Silicon (Si) has been proven to enhance the stress resistance of rice, but its effect on the lodging resistance of rice under dry cultivation (DCR) is still unclear. The purpose of this experiment is to clarify the appropriate amount of silicon fertilizer for DCR to resist lodging and to elucidate how it coordinates lodging resistance and yield. This experiment used the ‘Suigeng 18’ cultivar as the material and set six silicon fertilizers (SiO₂) with dosages of 0, 15, 30, 45, 60, and 75 kg·ha⁻¹ (Si0, Si1, Si2, Si3, Si4, Si5). Analyze the effects and key indicators of silicon on lodging resistance of DCR from the perspectives of plant weight distribution, stem structure and composition, and root architecture. The results showed that the Si3 treatment had the highest yield and the lowest lodging index (LI). Si3 increases the weight of the upper three leaves and 4–5 internodes, thereby promoting panicle weight and yield. An increase of 13.38% in 2/3PWSI (weight of the 4th–5th stems and upper three leaves/weight of the 1st–3rd stems and lower leaves) can reflect the promoting effect of silicon on stem and leaf development near the panicle. Si3 reduces the GA/IAA value, shortens the length of the second internode, and increases the diameters of the major and minor axes, thereby increasing culm thickness and section modulus (SM), achieving the effect of “short and thick”. Si3 also increases the content of silicon and non-structural carbohydrates (NSCs) in the second internode, and increases lignin and cellulose content by upregulating the expression levels of CAD7, PAL, COMT, and CesA4 genes, thereby increasing fullness and flexural strength (M), achieving “short, thick, and strong” and reducing LI. The 38.95% reduction in IFL (second internode length/fullness) reflects the positive effect of silicon on the “short, thick, and strong” stem. In the underground part, adding silicon reduces the CTK/IAA value of roots, and increases root length, root tip number, root surface area, and root weight. The key to coordinating the lodging resistance and yield of DCR with appropriate silicon dosage is to reduce the IFL in the second internode and increase 2/3 PWSI and root growth. The key to DCR and breeding is to focus on the relationship between basal internode length and fullness, as well as stem and leaf growth near the panicle. Full article

Cytokinins (CKs) play important functions in plant growth and development and in response to adversity stress. However, little is known about the role CK plays in rice grain quality. We hypothesized that exogenous cytokinins could improve rice grain quality by regulating physiological traits and genes related to starch synthesis. Therefore, we exogenously applied different concentrations of kinetin (KT), an exogenous CK, during the grain-filling period. Our results show that all the different concentrations of exogenous KT treatments resulted in a significant increase in thousand-grain weight. In particular, chalkiness and chalky grain rate were significantly reduced, and gel consistency (GC) content and alkali spreading value (ASV) were significantly increased in 10⁻⁸ M KT treatment. Meanwhile, the exogenous application of 10⁻⁸ M KT positively affected the transcription of some starch synthesis-related genes, which was in contrast to the 10⁻⁵ M KT treatment. In conclusion, the exogenous application of appropriate concentrations of KT during the grain-filling period can ultimately affect rice grain quality by regulating the changes in the relevant indicators, such as appearance quality (AQ) and eating and cooking qualities (ECQ). Full article

Previous studies have shown that a one-time application of polymer-coated urea (PCU) can increase rice yield and nitrogen (N) uptake. However, the connection between rice root morphology and physiological traits and grain yield and N absorption has still not been well understood. The objective of this study was to explore whether one-time application of PCU could enhance shoot growth, improve plant physiological activity, and ultimately boost rice yield and NUE by optimizing root morphological and physiological traits. In this study, a super-large-panicle indica-japonica hybrid rice variety, Yongyou1540, was cultivated under three N treatments during 2022 and 2023: (1) 0N, throughout the entire growth period, no N fertilizer was applied; (2) LFP, local farmers’ N management practices were followed, using urea as the N source, and N fertilizer management was carried out according to the local farmers’ customary fertilization practices; and (3) PCU, a one-time application of PCU was performed at one day before transplanting. PCU is a controlled-release fertilizer in which urea granules are coated with a synthetic polymer layer; it has been widely used in rice cultivation. In both LFP and PCU treatments, N was applied at a rate of 200 kg N ha⁻¹. PCU is a type of controlled-release fertilizer in which urea granules are coated with a layer of synthetic polymer. Compared to LFP, PCU significantly improved several root morphological traits, including increased deep-root proportion and specific root length (SRL), throughout the entire growth period; increased root length and root length density at heading and maturity; and increased root biomass growth rate from jointing to heading and reduced reduction rate after heading. Additionally, PCU enhanced root oxidative activity (ROA) and increased zeatin and zeatin riboside (Z+ZR) content in both roots and root bleeding sap at the middle and late grain-filling stages. Furthermore, PCU markedly increased the flag-leaf net photosynthetic rate, Z+ZR content in leaves, and activities of key enzymes involved in sucrose-to-starch conversion in grains during the middle and late grain-filling stages. Correlation analysis indicated that root and shoot biomass growth rate showed a significant positive correlation before heading, and that root biomass reduction rate was significantly negatively correlated with shoot biomass growth rate after heading. ROA and Z+ZR content in both roots and root bleeding sap were significantly associated with flag-leaf photosynthetic rate, Z+ZR content in leaves, and the activities of key enzymes involved in the sucrose-to-starch conversion in grains. On average, PCU increased rice yield by 10.0% and agronomic NUE by 46.2%, compared to LFP. These findings suggest that PCU could optimize root morphological and physiological traits, and thereby promote shoot growth, enhance physiological activity, and ultimately increase both rice yield and NUE. Further research could also investigate the potential for combining PCU with other agronomic practices to enhance both rice yield and NUE. Full article

Rice (Oryza sativa L.) is a staple crop for nearly half of the global population and one of China’s most extensively cultivated cereals. Heading date, a critical agronomic trait, determines the regional and seasonal adaptability of rice varieties. In this study, a series of mutants (elh5 to elh12) exhibiting extremely late heading under both long-day (LD) and short-day (SD) conditions were identified from an ethyl methanesulfonate (EMS) mutant library. Using MutMap and map-based cloning, the causative gene was identified as a novel allele of Ehd2/OsID1/RID1/Ghd10. Functional validation through CRISPR/Cas9 knockout and complementation assays confirmed its role in regulating heading. The elh6 mutation was found to cause intron retention due to alternative splicing. Ehd2 encodes a Cys-2/His-2-type zinc finger transcription factor with an IDD domain and transcriptional activity in yeast. Its expression peaks in developing leaves before heading and spikes during reproductive conversion. In elh6 mutants, delayed heading resulted from downregulating the Ehd1-Hd3a pathway genes. Salinity stress significantly hampers rice growth and productivity. Transcriptomic analysis of elh10 and ZH8015 seedlings exposed to salt stress for 24 h identified 5150 differentially expressed genes (DEGs) at the seedling stage, predominantly linked to stress response pathways. Ehd2 was revealed as a modulator of salt tolerance, likely through the regulation of ion transport, enzyme activity, and antioxidant systems. This study establishes Ehd2 as a pivotal factor in promoting heading while negatively regulating salt tolerance in rice. Full article

Botanical dietary supplements are widely used, but issues of authenticity, consistency, safety, and efficacy that complicate their poorly understood mechanism of action have prompted questions and concerns in the popular and scientific literature. Black cohosh (Actaea racemosa L., syn. Cimicifuga racemosa, Nutt., Ranunculaceae) is a multicomponent botanical therapeutic used as a popular remedy for menopause and dysmenorrhea and explored as a treatment in breast and prostate cancer. However, its use and safety are controversial. A. racemosa tissues contain the bioactive serotonin analog N-methylserotonin, which is thought to contribute to the serotonergic activities of black cohosh–containing preparations. A. racemosa has several TDC-like genes hypothesized to encode tryptophan decarboxylases (TDCs) converting L-tryptophan to tryptamine, a direct serotonin precursor in plants. Expression of black cohosh TDC1, TDC2, and TDC3 in Saccharomyces cerevisiae resulted in the production of tryptamine. TDC1 and TDC3 had approximately fourfold higher activity than TDC2, which was attributable to a variable Cys/Ser active site residue identified by site-directed mutagenesis. Co-expression in yeast of the high-activity black cohosh TDCs with the next enzyme in serotonin biosynthesis, tryptamine 5-hydroxylase (T5H), from rice (Oryza sativa) resulted in the production of serotonin, whereas co-expression with low-activity TDCs did not, suggesting that TDC activity is a rate-limiting step in serotonin biosynthesis. Two T5H-like sequences were identified in A. racemose, but their co-expression with the high-activity TDCs in yeast did not result in serotonin production. TDC expression was detected in several black cohosh tissues, and phytochemical analysis using LC-MS revealed several new tryptamines, including tryptamine and serotonin, along with N-methylserotonin and, interestingly, N-N-dimethyl-5-hydroxytryptamine (bufotenine), which may contribute to hepatotoxicity. Incubation of A. racemosa leaves with tryptamine and N-methyltryptamine resulted in increased concentrations of serotonin and N-methylserotonin, respectively, suggesting that methylation of tryptamine precedes hydroxylation in the biosynthesis of N-methylserotonin. This work indicates a significantly greater variety of serotonin derivatives in A. racemosa than previously reported. Moreover, the activities of the TDCs underscore their key role in the production of serotonergic compounds in A. racemosa. Finally, it is proposed that tryptamine is first methylated and then hydroxylated to form the black cohosh signature compound N-methylserotonin. Full article

Grain chalkiness adversely affects rice quality, and the positional variation of grain chalkiness within a rice panicle presents a substantial obstacle to quality improvement in China. However, the molecular mechanism underlying this variation is unclear. This study conducted a genetic and physiological analysis of grains situated at distinct positions (upper, middle, and bottom primary branches of the rice panicle, denoted as Y1, Y2, and Y3) within a rice panicle using the Yangdao 6 variety. The results indicated that the percentage of chalky grains (PCG) in Y1 was the highest, i.e., 17.12% and 52.18% higher than that of Y2 and Y3, respectively. Y2 exhibited the highest degree of grain chalkiness (DGC), attributable to its greater area of endosperm chalkiness (AEC) than the others. Y3 demonstrated the lowest PCG and DGC. Additionally, Y1 and Y2 were characterized by lower amylose and protein contents, as well as looser starch granule morphology, in comparison to Y3. Compared with Y3, both the average and maximum filling rates of Y1 and Y2 increased markedly; however, the active filling duration was notably reduced by 7.10 d and 5.56 d, respectively. The analysis of genomic expression levels indicated an enrichment of starch and sucrose metabolism in Y1-vs.-Y2, Y2-vs.-Y3, and Y1-vs.-Y3, with 7 genes (5 up-regulated and 2 down-regulated), 53 genes (12 up-regulated and 41 down-regulated), and 12 genes (2 up-regulated and 10 down-regulated) in the Y1-vs.-Y2, Y2-vs.-Y3, and Y1-vs.-Y3. The majority of these genes were down-regulated, linking metabolic activity to grain filling and contributing to the occurrence of grain chalkiness in rice panicles. In conclusion, the metabolic processes associated with sucrose and starch play a crucial role in regulating grain filling and the formation of chalkiness in rice. Full article

Water deficit affects rice growth, development, and yield. Knowledge of genetic diversity for water deficit tolerance, as well as the genetic architecture that is responsible for this trait, can accelerate rice cultivars’ improvement. In this study, different tools were applied to assess genetic diversity and identify genome regions associated with shoot and root traits in rice germplasm grown under water deficit at an early vegetative stage. A panel of 177 rice genotypes grown under water deficit was evaluated for root length (RL), root dry weight (RDW), shoot length (SL), and shoot dry weight (SDW). Genetic diversity was investigated using means grouping and principal component analysis. For the genome-wide association study, a general linear model was applied, using RL, RDW, SL, and SDW phenotypic data converted into Best Linear Unbiased Prediction (BLUPs); genotypic data (1185 single nucleotide polymorphism—SNPs-loci); and population structure. Overall, little genetic diversity was observed, but genotypes with a higher water deficit tolerance were identified. Several significant SNPs were mapped, 81, 5, 53, and 41 for RL, RDW, SL, and SDW, respectively. Among the identified genes, there are those encoding kinases, proteins involved in phytohormone and cell wall metabolism, and Cytochrome P450. The obtained results provide insight into genetic diversity and the genetic architecture of water deficit tolerance, which will be useful in improving this trait in rice grown in Brazil. Full article

Salt stress is a vital environmental stress that severely limits plant growth and productivity. Prohexadione-calcium (Pro-Ca) has been extensively studied to regulate plant growth, development, and stress responses. However, the constructive role of Pro-Ca in alleviating damages and enhancing rice tillers’ morph-physiological characteristics under salt stress remains largely unknown. The results showed that Pro-Ca significantly improved Changmaogu’s (CMG’s) productive tillering rate and the total yield per plant by 17.1% and 59.4%, respectively. At tillering stage, the results showed that Pro-Ca significantly improved the morph-physiological traits, i.e., leaf area, and photosynthetic traits of the rice variety with salt tolerance, under NaCl stress. Pro-Ca significantly increased the seedling index of the main stem and tiller by 10.3% and 20.0%, respectively. Pro-Ca significantly increased the chlorophyll a (chl a), chlorophyll b (chl b) and carotenoid contents by 32.8%, 58.4%, and 33.2%, respectively under NaCl stress. Moreover, Pro-Ca significantly enhanced the net photosynthetic rate (A) by 25.0% and the non-photochemical (NPQ) by 9.0% under NaCl stress. Furthermore, the application of Pro-Ca increased the activities of antioxidant enzymes by 7.5% and 14.7% in superoxide dismutase (SOD), 6.76% and 18.0% in peroxidase (POD), 26.4% and 58.5% in catalase (CAT), 11.0% and 15.9% in ascorbate peroxidase (APX), and Pro-Ca reduced the membrane damage index by 10.8% and 2.19% in malondialdehyde (MDA) content, respectively, for main stem and tiller leaves under NaCl stress. Pro-Ca significantly enhanced the soluble protein content of the main stem and tiller leaves by 2.60% and 6.08%, respectively. The current findings strongly suggested that exogenous application of Pro-Ca effectively alleviated the adverse impact of NaCl stress on the main stem and tillers by enhancing the photosynthetic capacity and antioxidant enzyme activity, and ultimately increased the productive tillering rate and grain yield. Full article

Background/Objectives: Cold stress poses a significant threat to Asian rice cultivation, disrupting important physiological processes crucial for seedling establishment and overall plant growth. It is, thus, crucial to elucidate genetic pathways involved in cold stress tolerance response mechanisms. Methods: We mapped OsUBC7, a Radiation-sensitive 6 (RAD6)-type homolog of rice, to a low-temperature seedling survivability (LTSS) QTL and used genomics, molecular genetics, and physiological assays to assess its role in plant resilience against low-temperature stress. Results: OsUBC7 is cold responsive and has higher expression levels in cold-tolerant japonica than cold-sensitive indica. Overexpression of OsUBC7 enhances LTSS of indica and freezing tolerance of Arabidopsis, increases levels of soluble sugars and chlorophyll A, boosts leaf development after cold exposure, and increases leaf cell numbers and plants size, but it does not affect membrane stability after cold stress exposure. Additionally, OsUBC7 has a positive role for germinability in the presence of salt and for flowering and yield-related traits. The OsUBC7 protein physically interacts with the developmental stage-specific and histone-modifying E3 ligases OsRFPH2-12 and OsHUB1/2, respectively, and potential target genes such as cell cycle dependent kinases were identified. Conclusions: OsUBC7 might contribute to cold resilience by regulating sugar metabolism to provide energy for promoting cellular homeostasis restoration after cold stress exposure via new cell growth, particularly in leaf cells crucial for photosynthesis and metabolic activity, possibly by interacting with cell cycle regulating proteins. Overall, the present study suggests that OsUBC7 may be involved in plant development, reproduction, and stress adaptation, and contributes to a deeper understanding of rice plant cold stress tolerance response mechanisms. OsUBC7 may be a promising candidate for improving crop productivity and resilience to stressful environments. Full article

Plant A/T-rich sequence- and zinc-binding protein (PLATZ) is a type of plant-specific zinc-dependent DNA-binding protein that binds to A/T-rich DNA sequences. This family is essential for plant growth, development, and stress response. In this study, 15 OsPLATZs were identified in the rice genome with complete PLATZ-conserved domains by CD-search, similar to those found in angiosperms. Multi-species phylogenetic analysis showed that PLATZs were conserved in photosynthetic organisms, and an evolutionary branch unique to angiosperms was identified among members of the PLATZ family. Fifteen OsPLATZs were represented by five groups, each with distinct characteristics. An analysis of protein structures and sequence motifs showed that OsPLATZs were similar within groups, but varied between them. The expression profile and qRT-PCR results showed that OsPLATZs had distinct expression patterns in different tissues, with some responding to stress induction. Most of the OsPLATZs localized to the nuclei, and were predicted to bind to DNA sequences by AlphaFold3, suggesting that they likely function as conventional transcription factors. We also identified OsPLATZ1, a caryopsis-specific gene that regulates grain filling and caryopsis development in rice. This research lays the foundation for exploring the structural diversity, evolutionary traits, expression profile, and possible roles of PLATZ transcription factors in rice. Full article

Background/Objectives: The balanced regulation of innate immunity plays essential roles in rhizobial infection and the establishment and maintenance of symbiosis. The evolutionarily conserved cell death suppressor Bax inhibitor-1 plays dual roles in nodule symbiosis, providing a valuable clue in balancing immunity and symbiosis, while it remains largely unexplored in the legume Lotus japonicus. Methods/Results: In the present report, the BI-1 gene family of L. japonicus was identified and characterized. We identified 6 BI-1 genes that translate into peptides containing 240–255 amino acids with different structural characteristics and isoelectric points. We performed phylogenetic analyses and detected evolutionary conservation and divergence among BI-1 proteins from L. japonicus, Glycine max, Medicago truncatula, Arabidopsis thaliana, and Oryza sativa. Expression profiles among different roots indicated that the inoculation of MAFF303099 significantly increased the expression of most of the L. japonicus BI-1 family genes. We down-regulated the transcripts of LjBI-1a by RNA interference and observed that LjBI-1a promotes nodulation and nodule formation. Conclusions: These discoveries shed light on the functions of BI-1 genes in L. japonicus, and simultaneously emphasize the potential application of LjBI-1a in enhancing the symbiotic nitrogen fixation ability of legumes. Full article

The Na⁺/H⁺ antiporter (NHX) gene family plays a pivotal role in plant salt tolerance in regulating intracellular Na⁺ and H⁺ homeostasis. In this study, seven candidate OsNHX genes (OsNHX1 to OsNHX7) were identified in the rice genome and classified into three phylogenetic clusters (Vac, Endo, and PM) based on their predicted subcellular localization. Five OsNHX gene pairs (OsNHX1/OsNHX2, OsNHX1/OsNHX3, OsNHX1/OsNHX4, OsNHX2/OsNHX6, and OsNHX5/OsNHX6) were found to have arisen from dispersed duplication events and exhibited purifying selection, indicating functional conservation. Analysis of cis-regulatory elements (CREs) revealed a diverse range of elements associated with tissue-specific expression, hormone signaling, and stress responses, particularly to dehydration and salinity. Notably, CREs associated with tissue/organelle-specific expression and stress responses were the most abundant, suggesting a potential role for OsNHX genes in regulating growth, development, and stress tolerance in rice. Importantly, expression profiling revealed that OsNHX1, OsNHX2, OsNHX3, and OsNHX5 were upregulated under salt stress, with significantly higher expression levels in the salt-tolerant rice cultivar Pokkali compared to the salt-sensitive cultivar IR64. Our findings provide a comprehensive analysis of the evolutionary, structural, and functional features of the OsNHX gene family and highlights their critical role in rice salt tolerance, offering insights into potential applications for crop improvement. Full article

Seed storability is a crucial agronomic trait and indispensable for the safe storage of rice seeds and grains. Nevertheless, the metabolite mechanisms governing Indica rice seed storability under natural conditions are still poorly understood. Methods: Therefore, the seed storage tolerance of global rice core germplasms stored for two years under natural aging conditions were identified, and two extreme groups with different seed storabilities from the Indica rice group were analyzed using the UPLC-MS/MS metabolomic strategy. Results: Our results proved that the different rice core accessions showed significant variability in storage tolerance, and the metabolite analysis of the two Indica rice pools exhibited different levels of storability. A total of 103 differentially accumulated metabolites (DAMs) between the two pools were obtained, of which 38 were up-regulated and 65 were down-regulated, respectively. Further analysis disclosed that the aging-resistant rice accessions had higher accumulation levels of flavonoids, terpenoids, phenolic acids, organic acids, lignans, and coumarins while exhibiting lower levels of lipids and alkaloids compared to the storage-sensitive rice accessions. The Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway enrichment analysis indicated that several biosynthesis pathways were involved in the observed metabolite differences, including alpha-linolenic acid metabolism, butanoate metabolism, and propanoate metabolism. Notably, inhibition of the linolenic acid metabolic pathway could enhance seed storability. Additionally, increased accumulations of organic acids, such as succinic acid, D-malic acid, and methylmalonic acid, in the butanoate and propanoate metabolisms were identified as a beneficial factor for seed storage. Conclusions: These new findings will deepen our understanding of the underlying mechanisms governing rice storability. Full article

Rice blast disease, caused by Magnaporthe oryzae (M. oryzae), is a significant threat to global rice production. Conventional methods for disease management face limitations, emphasizing the importance of sustainable alternatives. In this study, two rice cultivars with different blast resistant abilities, the susceptible variety CO39 and the resistant variety Pi4b, were used as materials to study the effects of Piriformospora indica (Pi) on the resistance to M. oryzae infection and rice growth. The in vitro tests revealed no direct antagonistic interaction between Pi and M. oryzae. However, the in vivo experiments showed that Pi promoted plant growth by increasing root and shoot length, chlorophyll content, and nitrogen uptake, particularly in CO39 during pathogen infection. Pi inoculation also significantly reduced disease severity, which was indicated by smaller lesion areas and shorter lesion lengths in both cultivars but a more pronounced effect in CO39. This occurred due to the decreasing levels of MDA and the modulating activity of antioxidant enzymes in Pi-inoculated rice plants. At the early stage of M. oryzae infection, the expression of the ethylene signaling gene OsEIN2 and the gibberellin biosynthesis gene OsGA20ox1 in Pi-inoculated CO39 decreased but significantly increased in both rice cultivars at the later stage. The reverse was found for the pathogenesis-related (PR) genes OsPR10 and OsPBZ1 and the blast-resistant genes OsBRG1, OsBRG2, and OsBRW1, suggesting early growth suppression for rice resilience to blast followed by a later shift back to growth. Meanwhile, Pi inoculation increased OsCesA9 expression in rice to strengthen cell walls and establish the primary defense barrier against M. oryzae and upregulated the expression of OsNPR1 without a significant difference in CO39 but downregulated it in Pi4b to activate PR genes to enhance plant resistance. In summary, these results underscore the potential of Pi as a sustainable biological control agent for rice blast disease, which is particularly beneficial for blast-susceptible rice cultivars. Full article

Chalkiness in rice is adversely affected by high temperatures during the flowering and grain-filling stages. Potassium (K) is essential for improving grain quality and heat resilience. The effects of split application K fertilizer on rice chalkiness under high temperatures during the flowering and grain-filling stages were investigated in this study. Four treatments, including ambient temperatures with basal K fertilizer (AT-K1), high temperatures with basal K fertilizer (HT-K1), high temperatures with 70% K pre-transplanting and 30% K at the heading stage (HT-K2), and high temperatures with 30% K pre-transplanting and 70% K at the heading stage (HT-K3), were conducted. The results revealed that the chalky grain rate and chalkiness degree were reduced by 9.2–13.72% and 12.16–19.91%, respectively, by the split application of K fertilizer through effectively modulating the sucrose-to-starch conversion process in the rice grains, relative to the single basal application of K fertilizer under high temperatures. Specifically, the split application of K fertilizer reduced the enzymatic activities of SuSy, ADPGase, and SBE by 3.17–34.20% at 5–10 DAA, and GBSS and SSS by 6.48–13.50% at 5 DAA, but enhanced them by 5.50–54.90% from 15 DAA and 2.07–97.10% from 10 DAA. Similarly, the gene expression levels of enzymes involved in this process were decreased by 3.52–24.12% at 5 DAA but increased by 8.61–30.00% at 20 DAA by the split application of K fertilizer. This modulation led to a retardation in the excessive accumulation of starch during the early grain-filling stage but a higher starch accumulation rate during the middle and later stages, combined with a longer duration of starch accumulation, ultimately resulting in higher starch accumulation and reduced rice chalkiness. These results suggest that the application of K fertilizer during the heading stage is effective in compensating the deterioration of rice chalkiness by high temperatures. Full article