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Lotus (Nelumbo Adans) is an aquatic perennial plant that flourished during the middle Albian stage. In this study, we characterized the digital gene expression signatures for China Antique lotus under conditions of heat shock stress. Using RNA-seq technology, we sequenced four libraries, specifically, two biological replicates for control plant samples and two for heat stress samples. As a result, 6,528,866 to 8,771,183 clean reads were mapped to the reference genome, accounting for 92-96% total clean reads. A total of 396 significantly altered genes were detected across the genome, among which 315 were upregulated and 81 were downregulated by heat shock stress. Gene ontology (GO) enrichment of differentially expressed genes revealed protein folding, cell morphogenesis and cellular component morphogenesis as the top three functional terms under heat shock stress. Kyoto Encyclopedia of Genes and Genomes (KEGG) analysis led to the identification of protein processing in endoplasmic reticulum, plant-pathogen interactions, spliceosome, endocytosis, and protein export as significantly enriched pathways. Among the upregulated genes, small heat shock proteins (sHsps) and genes related to cell morphogenesis were particularly abundant under heat stress. Data from the current study provide valuable clues that may help elucidate the molecular events underlying heat stress response in China Antique lotus.

Background Flower morphology, a phenomenon regulated by a complex network, is one of the vital ornamental features in Nelumbo nucifera . Stamen petaloid is very prevalent in lotus flowers. However, the mechanism underlying this phenomenon is still obscure. Results Here, the comparative transcriptomic analysis was performed among petal, stamen petaloid and stamen through RNA-seq. Using pairwise comparison analysis, a large number of genes involved in hormonal signal transduction pathways and transcription factors, especially the MADS-box genes, were identified as candidate genes for stamen petaloid in lotus. Conclusions Taken together, these results provide an insight into the molecular networks underlying lotus floral organ development and stamen petaloid.

Background Starch in the lotus seed contains a high proportion of amylose, which endows lotus seed a promising property in the development of hypoglycemic and low-glycemic index functional food. Currently, improving starch content is one of the major goals for seed-lotus breeding. ADP-glucose pyrophosphorylase (AGPase) plays an essential role in regulating starch biosynthesis in plants, but little is known about its characterization in lotus. Results We describe the nutritional compositions of lotus seed among 30 varieties with starch as a major component. Comparative transcriptome analysis showed that AGPase genes were differentially expressed in two varieties (CA and JX) with significant different starch content. Seven putative AGPase genes were identified in the lotus genome ( Nelumbo nucifera Gaertn.), which could be grouped into two subfamilies. Selective pressure analysis indicated that purifying selection acted as a vital force in the evolution of AGPase genes. Expression analysis revealed that lotus AGPase genes have varying expression patterns, with NnAGPL2a and NnAGPS1a as the most predominantly expressed, especially in seed and rhizome. NnAGPL2a and NnAGPS1a were co-expressed with a number of starch and sucrose metabolism pathway related genes, and their expressions were accompanied by increased AGPase activity and starch content in lotus seed. Conclusions Seven AGPase genes were characterized in lotus, with NnAGPL2a and NnAGPS1a , as the key genes involved in starch biosynthesis in lotus seed. These results considerably extend our understanding on lotus AGPase genes and provide theoretical basis for breeding new lotus varieties with high-starch content.

Background Sacred lotus ( Nelumbo nucifera ) is a vital perennial aquatic ornamental plant. Its flower shape determines the horticultural and ornamental values. However, the mechanisms underlying lotus flower development are still elusive. MADS-box transcription factors are crucial in various features of plant development, especially in floral organogenesis and specification. It is still unknown how the MADS-box transcription factors regulate the floral organogenesis in lotus. Results To obtain a comprehensive insight into the functions of MADS-box genes in sacred lotus flower development, we systematically characterized members of this gene family based on the available genome information. A total of 44 MADS-box genes were identified, of which 16 type I and 28 type II genes were categorized based on the phylogenetic analysis. Furthermore, the structure of MADS-box genes and their expressional patterns were also systematically analyzed. Additionally, subcellular localization analysis showed that they are mainly localized in the nucleus, of which a SEPALLATA3 ( SEP3 ) homolog NnMADS14 was proven to be involved in the floral organogenesis. Conclusion These results provide some fundamental information about the MADS-box gene family and their functions, which might be helpful in not only understanding the mechanisms of floral organogenesis but also breeding of high ornamental value cultivars in lotus.

Key message NnNAC100-NnSBEII modules enhance starch content of the rhizome in Nelumbo nucifera Gaertn. Nelumbo nucifera Gaertn. is a popular aquatic vegetable and traditional Chinese medicine whose quality and taste are mainly determined by the starch. Although starch-related genes have been functionally characterized, the regulated mechanism of enzyme (SBE) remains unclear. In this study, we identified and functionally elucidated the functions of NnSBEII and NnNAC100 using transient overexpression of NnSBEII and NnNAC100 in rhizomes of lotus, and it significantly increased the amylopectin content and total starch content. Accordingly, functional complementation assay in defective Arabidopsis also showed that NnSBEII compensated for the low content of starch in the mutant sbe2.2 . In addition, overexpression of NnSBEII and NnNAC100 significantly increased the content of starch in transgenic lines. Consistently, opposite results were observed under the background of repressed NnSBEII and NnNAC100 in rhizomes of lotus. Furthermore, yeast one-hybrid and dual-luciferase assays revealed that NnNAC100 could directly bind to the NnSBEII promoter and promote the expression of NnSBEII . Transient overexpression of NnNAC100 upregulated NnSBEII expression significantly, while the expression level of AtSBE2.2 in transgenic Arabidopsis overexpressing NnNAC100 was higher than that of WT, which indicated that NnNAC100 promoted the synthesis of amylopectin by enhancing the expression of NnSBEII . In addition, we found that NnSBEII could form a complex protein by interacting with soluble starch synthase (NnSS2) to increase the activity of the SBEII enzyme. These results reveal a novel mechanism that the NnNAC100- NnSBEII -NnSBEII/NnSS2 module regulates amylopectin biosynthesis and these will provide insights into the broader implications of the regulation mechanism of starch biosynthesis.

Genomic resources such as single nucleotide polymorphism (SNPs), insertions and deletions (InDels) and SSRs (simple sequence repeats) are essential for crop improvement and better utilization in genetic breeding. However, the resources for the sacred lotus (Nelumbo nucifera Gaertn.) are still limited. In the present study, to dissect large-scale genomic molecular marker resources for sacred lotus, we re-sequenced a Thailand sacred lotus cultivar 'Chiang Mai wild lotus' and compared with the reported lotus genome 'Middle lake wild lotus'. A total of 3,180,059 SNPs, 328, 251 InDels and 14,191 SVs were found between the two genomes. The functional impact analyses of these SNPs indicated that they may be involved in metabolic processes, binding, catalytic activity, etc. Mining the genome sequences for SSRs showed that 191,657 SSRs were identified with a frequency of one SSR per 4.23 kb and 103,656 SSR primer pairs were designed. Furthermore, 14, 502 EST-SSRs were also indentified using the available RNA-seq data in the NCBI. A subset of 150 SSRs (genomic and EST-SSRs) was randomly selected for validation and genetic diversity analysis. The genotypes could be easily distinguished using these SSR markers and the 'Chiang Mai wild lotus' was obviously differentiated from the other Chinese accessions. This study provides considerable amounts of genomic resources and markers for the quantitative trait locus (QTL) identification and molecular selection of the species, which could have a potential role in various applications in sacred lotus breeding.

The sacred lotus (Nelumbo nucifera Gaertn.) can produce heat autonomously and maintain a relatively stable floral chamber temperature for several days when blooming. Floral thermogenesis is critical for flower organ development and reproductive success. However, the regulatory role of microRNA (miRNA) underlying floral thermogenesis in N. nucifera remains unclear. To comprehensively understand the miRNA regulatory mechanism of thermogenesis, we performed small RNA sequencing and transcriptome sequencing on receptacles from five different developmental stages. In the present study, a total of 172 known miRNAs belonging to 39 miRNA families and 126 novel miRNAs were identified. Twenty-nine thermogenesis-related miRNAs and 3024 thermogenesis-related mRNAs were screened based on their expression patterns. Of those, seventeen differentially expressed miRNAs (DEMs) and 1765 differentially expressed genes (DEGs) had higher expression during thermogenic stages. The upregulated genes in the thermogenic stages were mainly associated with mitochondrial function, oxidoreductase activity, and the energy metabolism process. Further analysis showed that miR156_2, miR395a_5, miR481d, and miR319p may play an important role in heat-producing activity by regulating cellular respiration-related genes. This study provides comprehensive miRNA and mRNA expression profile of receptacle during thermogenesis in N. nucifera, which advances our understanding on the regulation of floral thermogenesis mediated by miRNA.

Background Lotus root, a crucial aquatic cash crop, demonstrates primary root atrophy, while developing adventitious roots critically mediate nutrient acquisition and product organ development. Previous investigation reveals the functional involvement of NnWOX1-1 in root development through transgenic Arabidopsis models. Results In this study, we demonstrated nuclear and membrane co-localization of NnWOX1-1 fluorescence signals in tobacco leaf epidermal cells. NnWOX1-1 overexpression significantly enhanced root elongation and increased root primordia formation in transgenic lotus root and Sagittaria trifolia plants. We identified NnDREB2C as a downstream target gene of NnWOX1-1. The 615 bp nucleotide sequence of NnDREB2C encoded a 205-amino acid polypeptide containing a conserved AP2 domain. Subcellular localization assays revealed exclusive nuclear accumulation of NnDREB2C -GFP fusion proteins in tobacco mesophyll cells. NnDREB2C transcription was significantly upregulated under 200 mg/L ethephon treatment and 20 g/L sucrose exposure, while remaining unresponsive to 10 µM indole-3-acetic acid (IAA). Tissue-specific expression analysis showed maximum transcript abundance in leaves, floral organs, and internodal segments, with minimal expression in fruits and petioles. Transgenic manipulation experiments established that NnDREB2C overexpression enhanced the number and length of adventitious root in both lotus and S. trifolia plants. Conversely, RNAi-mediated knockdown lines exhibited significant reductions in the adventitious root development compared to wild-type counterparts. Conclusions Through functional analysis of NnWOX1-1 , NnDREB2C and the measurement of ethylene content in transgenic plants, we demonstrated that NnDREB2C regulated by NnWOX1-1 was involved in the formation of adventitious roots in lotus seedling. Additionally, NnWOX1-1 and NnDREB2C affected synthesis of ethylene which was also an important regulator of adventitious roots.

Sacred lotus (Nelumbo nucifera Gaertn.) is an ancient aquatic plant used throughout Asia for its nutritional and medicinal properties. Benzylisoquinoline alkaloids (BIAs), mostly within the aporphine and bisbenzylisoquinoline structural categories, are among the main bioactive constituents in the plant. The alkaloids of sacred lotus exhibit promising anti-cancer, anti-arrhythmic, anti-HIV, and anti-malarial properties. Despite their pharmacological significance, BIA metabolism in this non-model plant has not been extensively investigated. In this review, we examine the diversity of BIAs in sacred lotus, with an emphasis on the distinctive stereochemistry of alkaloids found in this species. Additionally, we discuss our current understanding of the biosynthetic genes and enzymes involved in the formation of 1-benzylisoquinoline, aporphine, and bisbenzylisoquinoline alkaloids in the plant. We conclude that a comprehensive functional characterization of alkaloid biosynthetic enzymes using both in vitro and in vivo methods is required to advance our limited knowledge of BIA metabolism in the sacred lotus.

Fifteen compounds were extracted and purified from the leaves of Nelumbo nucifera Gaertn. cv. Rosa-plena. These compounds include liriodenine (1), lysicamine (2), (−)-anonaine (3), (−)-asimilobine (4), (−)-caaverine (5), (−)-N-methylasimilobine (6), (−)-nuciferine (7), (−)-nornuciferine (8), (−)-roemerine (9), 7-hydroxydehydronuciferine (10) cepharadione B (11), β-sitostenone (12), stigmasta-4,22-dien-3-one (13) and two chlorophylls: pheophytin-a (14) and aristophyll-C (15). The anti-oxidation activity of the compounds was examined by antiradical scavenging, metal chelating and ferric reducing power assays. The results have shown that these compounds have antioxidative activity. The study has also examined the antiproliferation activity of the isolated compounds against human melanoma, prostate and gastric cancer cells. The results shown that 7-hydroxydehydronuciferine (10) significantly inhibited the proliferation of melanoma, prostate and gastric cancer cells. Together, these findings suggest that leaves of Nelumbo nucifera Gaertn. cv. Rosa-plena are a good resource for obtaining the biologically active substances with antioxidant properties.