Nelumbonaceae

Nelumbo nucifera

Bibliography

  1. 桑子芳, 周明全, 胡中立, 廖冬青, 何建君, and 洪贤清, “Method for Breeding Lotus Root Varieties,” May-2005. https://patents.google.com/patent/CN1202705C/en.
    The present invention discloses a method for breeding lotus root varieties, which comprises the steps that resource collection is firstly carried out to obtain parent plant lines; secondly, continuous hybridization is carried out, thirdly, continuous selection is carried out for selecting variety lines; fourthly, good variety lines are enlarged and bred so as to be convenient for large-region test. The present invention has the advantages of compact experimental process, convenient operation, stability and reliability, and the obtained good variety lines have the advantages of high yield and low cost and enhance the production level of lotus roots.
  2. 闵 睫, “The Physiological and Biochemical Changes of Lotus (Nelumbo) Flowers in Natural State and Vase,” Botanical Research, vol. 09, pp. 31–38, Jan. 2020. doi: 10.12677/BR.2020.91004.
  3. 李良俊 et al., “Study on carbohydrate metabolism of lotus root during overwintering in the field,” 中国蔬菜, vol. 1, no. 4, pp. 11–13. http://www.cnveg.org/CN/abstract/abstract9276.shtml.
  4. N. S. Abd Rasid, M. N. Naim, H. Che Man, N. F. Abu Bakar, and M. N. Mokhtar, “Evaluation of Surface Water Treated with Lotus Plant; Nelumbo Nucifera,” Journal of Environmental Chemical Engineering, vol. 7, no. 3, p. 103048, Jun. 2019. doi: 10.1016/j.jece.2019.103048.
    The potential of Nelumbo nucifera in treating contaminated surface water was investigated in terms of biochemical oxygen demand (BOD), chemical oxygen demand (COD), turbidity, and nitrate reduction. Batch type lab-scale container cultivated with N. nucifera was exposed to the contaminated surface water for 30 days. Nitrate (NO3−) adsorption and pH level were monitored continuously to identify the plant survival and to avoid any additional contaminants into the samples such as plant decay. For comparison, water lily, Nymphaea, was prepared using the same experimental setup. After 30 days of phytoremediation, the BOD and COD values of the treated water using N. nucifera was significantly reduced to 97.1% and 55%, respectively, due to the unique gas transport mechanism that thermodynamically drive O2 gas from leaves at the water surface to the buried rhizomes located in the anoxic sediments. When treated with Nymphaea, the BOD value in water decreased by 64.5% and the COD value increased by 50.5%. The results indicate that N. nucifera was able to remove the organic contaminants from the surface water by supplying adequate amount of 0.2–2.1 mL/min O2 gas to increase the microbial activities from the control condition.
  5. G. G. (G. G. Ainslie and W. B. (W. B. Cartwright, Biology of the Lotus Borer (Pyrausta Penitalis Grote). Washington, D.C. : U.S. Dept. of Agriculture, 1922. http://archive.org/details/biologyoflotusbo1076ains.
    Caption title; "July, 1922."; "Professional paper."; Bibliography: p. 13-14
  6. S. Al-Hamdani and D. A. Francko, “Effect of Light and Temperature on Photosynthesis, Elongation Rate and Chlorophyll Content of Nelumbo Lutea (Willd.) Pers. Seedlings,” Aquatic Botany, vol. 44, no. 1, pp. 51–58, Dec. 1992. doi: 10.1016/0304-3770(92)90080-3.
    Petiolar photosynthetic carbon assimilation (PCA) rates, elongation rates, and chlorophyll (Chl) content were assessed in seedlings of Nelumbo lutea (Willd.) Pers. grown in aseptic liquid culture under variable photon flux density (30–910 μmol m−2s−1) and temperature (3–30°C) conditions. When plants were grown under standard conditions (23°C, 100 μmol m−2s−1 photon flux density) and exposed to short-term changes in light and temperature regimes, maximum PCA (approximately 650 μmol Cg−1 Chl a h−1) occurred at 20°C and a photon flux density of 500 μmol m−2s−1. Longer term growth experiments (10 day) conducted under a variety of temperature: illumination regimes demonstrated that at all temperatures PCA rates increased curvilinearly up to a photon flux density of 560 μmol m−2s−1, then decreased gradually as light intensity increased. At all photon flux densities, the highest seedling PCA was obtained at 20°C. Petiole elongation rates increased up to 560 μmol m−2s−1 and declined at higher photon flux densities. Overall elongation rates were greatest at 30°C, followed by 20°C and 10°C. Chlorophyll content decreased as photon flux density increased in seedlings grown at 30°C and 20°C, but at 10°C increased up to 350 μmol m−2s−1 and then declined at higher photon flux densities. The collective data suggest that PCA by elongating petioles, may contribute to seedling growth, even under relatively cool, dark conditions near the sediment-water interphase.
  7. S. Arunyanart and M. Chaitrayagun, “Induction of Somatic Embryogenesis in Lotus (Nelumbo Nucifera Geartn.),” Scientia Horticulturae, vol. 105, no. 3, pp. 411–420, Jul. 2005. doi: 10.1016/j.scienta.2005.01.034.
    Callus induction and somatic embryogenesis of lotus (Nelumbo nucifera Gaertn.) cv. Satabankacha were studied. Callus was initiated by culturing bud, cotyledon, and young leaf explants on Murashige and Skoog (MS) (1962) medium containing a combination of 0, 4, 8 and 10μM 2,4-dichlorophenoxy acetic acid (2,4-D) and 0, 1, 2 and 3μM 6-furfuryl amino purine (kinetin) or substituting 0, 0.5 and 1μM benzyladenine (BA) for kinetin. Bud explants cultured on medium containing 4μM 2,4-D and 1μM BA gave the best callus growth. For somatic embryogenesis, the calli initiated on MS medium containing a combination of 4, 6, 8 and 10μM 2,4-D and 1μM BA and subsequently transferred to media containing 2–4μM 2,4-D and 0 or 0.5μM BA produced the most somatic embryos. When cultures were 12-week-old, callus produced on medium with 6μM 2,4-D and 1μM BA showed the best growth for somatic embryo regeneration. When transferred to a medium with 2μM 2,4-D and 0.5μM BA somatic embryos were produced from 33% of the calli. Embryos developed to the stage proembryo within 4 weeks and formed globular, heart, torpedo and mature embryos within 16 weeks.
  8. S. Arunyanart and S. Soontronyatara, “Mutation Induction by γ and X-Ray Irradiation in Tissue Cultured Lotus,” Plant Cell, Tissue and Organ Culture, vol. 70, no. 1, pp. 119–122, Jul. 2002. doi: 10.1023/A:1016021627832.
    Mutations of tissue cultured lotus were induced by treating plantlets with either acute γ-rays at doses of 0, 2, 3, 4, 5 or 6 krad or X-rays at doses of 0, 1, 2, 3, 4 or 5 krad. The 2-krad dose of either γ- or X-ray treatments resulted in a 50% survival rate. The use of γ- and X-rays to induce mutation in lotus resulted in 21 altered characteristics. Mutants from 1- and 2-krad of either γ or X-rays had long secondary roots and numerous adventitious roots. These mutants also exhibited good shoot growth and healthy rhizome development. Most plants treated with 3–5 krad of either γ- or X-rays exhibited abnormal characteristics including vitrification, chlorosis, deformed petioles and in addition had inhibited growth of lateral buds, secondary roots and rhizomes. All plants treated with 6 krad of γ-rays died within 4 weeks. Control plants had stoma lengths of 2.56 μm and cytological analysis of the root tips confirmed the diploid chromosome number of 16. Two groups of aneuploid cells were achieved using irradiation at doses of 3 and 4 krad of either γ- or X-ray. Chromosome numbers were 2n=18 and 20 with associated stoma lengths of 3.43 and 4.34 μm, respectively. Abnormal stomata (cyclocytic and deformity) were observed in plants treated with 4 krad of γ-ray.
  9. W. Barthlott and C. Neinhuis, “Purity of the Sacred Lotus, or Escape from Contamination in Biological Surfaces,” Planta, vol. 202, no. 1, pp. 1–8, Apr. 1997. doi: 10.1007/s004250050096.
    The microrelief of plant surfaces, mainly caused by epicuticular wax crystalloids, serves different purposes and often causes effective water repellency. Furthermore, the adhesion of contaminating particles is reduced. Based on experimental data carried out on microscopically smooth (Fagus sylvatica L., Gnetum gnemon L., Heliconia densiflora Verlot, Magnolia grandiflora L.) and rough water-repellent plants (Brassica oleracea L., Colocasia esculenta (L.) Schott., Mutisia decurrens Cav., Nelumbo nucifera Gaertn.), it is shown here for the first time that the interdependence between surface roughness, reduced particle adhesion and water repellency is the keystone in the self-cleaning mechanism of many biological surfaces. The plants were artificially contaminated with various particles and subsequently subjected to artificial rinsing by sprinkler or fog generator. In the case of water-repellent leaves, the particles were removed completely by water droplets that rolled off the surfaces independent of their chemical nature or size. The leaves of N. nucifera afford an impressive demonstration of this effect, which is, therefore, called the “Lotus-Effect” and which may be of great biological and technological importance.
  10. W. Barthlott, C. Neinhuis, R. Jetter, T. Bourauel, and M. Riederer, “Waterlily, Poppy, or Sycamore: On the Systematic Position of Nelumbo,” Flora, vol. 191, no. 2, pp. 169–174, Apr. 1996. doi: 10.1016/S0367-2530(17)30709-0.
    The systematic position of Nelumbo, originally described as Nymphaea, turnes out to be more confusing than ever. A new set of data derived from high resolution scanning electron microscopy and chemistry of epicuticular waxes is presented. It is compared to results obtained from investigations of several families of Magnoliidae, Ranunculidae, and Hamamelidae. The traditional opinion that Nelumbo is closely related to Nymphaeales or even Magnoliales is rejected. The occurrence of tubular wax crystals containing a significant amount of noncosan-10-ol supports a position near to Ranunculiflorae, if not to “lower” hamamelids (Hamamelidaceae, Daphniphyllaceae, Cercidiphyllaceae). An unique new feature of Nelumbo is the occurrence of aporphin alkaloids (nuciferin, nornuciferin) in cuticular waxes. These alkaloids, well known from the tissues of Nelumbo, are common in Ranunculiflorae, whereas Nymphaeales are characterised by sesquiterpen alkaloids.
  11. C. Bei-sheng, “Changes of Carbohydrates During Lotus Rhizome Development of Nelumbo Nucifera Gaertn,” undefined, 2003. https://www.semanticscholar.org/paper/CHANGES-OF-CARBOHYDRATES-DURING-LOTUS-RHIZOME-OF-Bei-sheng/88bb0d96bc349ee99500d61b4030a5aedd6c9f43.
    The changes of dry materials, starch, reducing sugar, total soluble sugar and soluble protein contents during lotus rhizome development of Nelumbo nucifera Gaertn were studied. The changes of dry materials, starch, reducing sugar, total soluble sugar and soluble protein contents during lotus rhizome development of Nelumbo nucifera Gaertn were studied. The results were as follows: the contents of dry materials, starch, soluble sugar and soluble protein had a significant increase during the development of lotus rhizome. Reducing sugar levels increased at early stage and decreased promptly at middle stage, then it kept a steady level until the end stage of rhizome development. The main storage nutrition of lotus rhizome was starch.
  12. L. Bharathi Priya, C.-Y. Huang, R.-M. Hu, B. Balasubramanian, and R. Baskaran, “An Updated Review on Pharmacological Properties of Neferine—A Bisbenzylisoquinoline Alkaloid from Nelumbo Nucifera,” Journal of Food Biochemistry, vol. 45, no. 12, p. e13986, 2021. doi: 10.1111/jfbc.13986.
    Phytochemicals have recently received a lot of recognition for their pharmacological activities such as anticancer, chemopreventive, and cardioprotective properties. In traditional Indian and Chinese medicine, parts of lotus (Nelumbo nucifera) such as lotus seeds, fruits, stamens, and leaves are used for treating various diseases. Neferine is a bisbenzylisoquinoline alkaloid, a major component from the seed embryos of N. nucifera. Neferine is effective in the treatment of high fevers and hyposomnia, as well as arrhythmia, platelet aggregation, occlusion, and obesity. Neferine has been found to have a variety of therapeutic effects such as anti-inflammatory, anti-oxidant, anti-hypertensive, anti-arrhythmic, anti-platelet, anti-thrombotic, anti-amnesic, and negative inotropic. Neferine also exhibited anti-anxiety effects, anti-cancerous, and chemosensitize to other anticancer drugs like doxorubicin, cisplatin, and taxol. Induction of apoptosis, autophagy, and cell cycle arrest are the key pathways that underlying the anticancer activity of neferine. Therefore, the present review summarizes the neferine biosynthesis, pharmacokinetics, and its effects in myocardium, cancer, chemosensitizing to cancer drug, central nervous system, diabetes, inflammation, and kidney diseases. Practical applications Natural phytochemical is gaining medicinal importance for a variety of diseases like including cancer, neurodegenerative disorder, diabetes, and inflammation. Alkaloids and flavonoids, which are abundantly present in Nelumbo nucifera have many therapeutic applications. Neferine, a bisbenzylisoquinoline alkaloid from N. nucifera has many pharmacological properties. This present review was an attempt to compile an updated pharmacological action of neferine in different disease models in vitro and in vivo, as well as to summarize all the collective evidence on the therapeutic potential of neferine.
  13. B. Bhushan, Y. C. Jung, and K. Koch, “Self-Cleaning Efficiency of Artificial Superhydrophobic Surfaces,” Langmuir, vol. 25, no. 5, pp. 3240–3248, Mar. 2009. doi: 10.1021/la803860d.
    The hierarchical structured surface of the lotus (Nelumbo nucifera, Gaertn.) leaf provides a model for the development of biomimetic self-cleaning surfaces. On these water-repellent surfaces, water droplets move easily at a low inclination of the leaf and collect dirt particles adhering to the leaf surface. Flat hydrophilic and hydrophobic, nanostructured, microstructured, and hierarchical structured superhydrophobic surfaces were fabricated, and a systematic study of wettability and adhesion properties was carried out. The influence of contact angle hysteresis on self-cleaning by water droplets was studied at different tilt angles (TA) of the specimen surfaces (3° for Lotus wax, 10° for n-hexatriacontane, as well as 45° for both types of surfaces). At 3° and 10° TA, no surfaces were cleaned by moving water applied onto the surfaces with nearly zero kinetic energy, but most particles were removed from hierarchical structured surfaces, and a certain amount of particles were captured between the asperities of the micro- and hierarchical structured surfaces. After an increase of the TA to 45° (larger than the tilt angles of all structured surfaces), as usually used for industrial self-cleaning tests, all nanostructured surfaces were cleaned by water droplets moving over the surfaces followed by hierarchical and microstructures. Droplets applied onto the surfaces with some pressure removed particles residues and led to self-cleaning by a combination of sliding and rolling droplets. Geometrical scale effects were responsible for superior performance of nanostructured surfaces.
  14. A. J. Blaylock and R. S. Seymour, “Diaphragmatic Nets Prevent Water Invasion of Gas Canals in Nelumbo Nucifera,” Aquatic Botany, vol. 67, no. 1, pp. 53–59, May 2000. doi: 10.1016/S0304-3770(99)00087-X.
    This study measured pore size in the net-like diaphragms of the lotus (Nelumbo nucifera Gaertn.) and the water pressures necessary to break their menisci. Mean pore radii were 7.0±0.3μm (95% confidence interval) in the rhizome and 19.0±0.5μm in the petiole. Nets in the rhizomes could prevent internal flooding in water depths up to 2m, which corresponds approximately to the depths at which natural stands of the lotus grow. Nets in the emergent petioles could withstand only 0.5m of water pressure, but there was no relationship between pore radius and the proximity of a net to the rhizome. Their role may be to reduce the kinetic energy of incoming water.
  15. R. Buathong, K. Saetiew, S. Phansiri, N. Parinthawong, and S. Arunyanart, “Tissue Culture and Transformation of the Antisense DFR Gene into Lotus (Nelumbo Nucifera Gaertn.) through Particle Bombardment,” Scientia Horticulturae, vol. 161, pp. 216–222, Sep. 2013. doi: 10.1016/j.scienta.2013.06.040.
    The effects of explant and plant growth regulators on callus induction and plant regeneration and particle bombardment-induced transformation were examined in lotus (Nelumbo nucifera Gaertn.) cv. Buntharik. The apical buds from embryos (∼3mm) formed an embryogenic callus when cultured on MS medium supplemented with 40μM 1-napthaleneactic acid (NAA) and 0.5μM 1-phenyl-3-(1,2,3-thiadiazol-5-yl)-urea (TDZ) for 8 weeks. All of the embryogenic calluses were transferred to MS medium supplemented with 0, 40, 50 and 60μM 6-benzyladenine (BA) and the highest number of shoots was achieved in the medium supplemented with 50μM BA after 8 weeks of culture. The transformation of lotus using a particle bombardment device was examined with the shoot clusters from embryogenic apices. The pCAMBIA2301anti-DFR plasmid contained β-glucuronidase (GUS) as a reporter gene, neomycin phosphotransferase (NPTII) as a selectable marker, and antisense dihydroflavonol 4-reductase (anti-DFR) as the gene of interest. The particle bombardment procedure with a helium gas pressure of 1100psi and a target distance of 6cm yielded the highest number of blue spots (4.8 spots/shoot cluster). However, stable transformations, as confirmed by polymerase chain reaction and reverse transcription polymerase chain reaction analysis, were only achieved using a helium gas pressure of 1100psi and a target distance of 9cm. Our work illustrates that the stable transformation of lotus is possible, and we suggest that this methodology can be used to target and modify specific charactics of this plant.
  16. J. Cao, Q. Jin, J. Kuang, Y. Wang, and Y. Xu, “Regulation of Flowering Timing by ABA-NnSnRK1 Signaling Pathway in Lotus,” International Journal of Molecular Sciences, vol. 22, no. 8, p. 3932, Jan. 2021. doi: 10.3390/ijms22083932.
    The lotus produces flower buds at each node, yet most of them are aborted because of unfavorable environmental changes and the mechanism remains unclear. In this work, we proposed a potential novel pathway for ABA-mediated flower timing control in the lotus, which was explored by combining molecular, genetic, transcriptomic, biochemical, and pharmacologic approaches. We found that the aborting flower buds experienced extensive programmed cell death (PCD). The hormonal changes between the normal and aborting flower buds were dominated by abscisic acid (ABA). Seedlings treated with increasing concentrations of ABA exhibited a differential alleviating effect on flower bud abortion, with a maximal response at 80 μM. Transcriptome analysis further confirmed the changes of ABA content and the occurrence of PCD, and indicated the importance of PCD-related SNF1-related protein kinase 1 (NnSnRK1). The NnSnRK1-silenced lotus seedlings showed stronger flowering ability, with their flower:leaf ratio increased by 40%. When seedlings were treated with ABA, the expression level and protein kinase activity of NnSnRK1 significantly decreased. The phenotype of NnSnRK1-silenced seedlings could also be enhanced by ABA treatment and reversed by tungstate treatment. These results suggested that the decline of ABA content in lotus flower buds released its repression of NnSnRK1, which then initiated flower bud abortion.
  17. S. Chen, H. Zhang, Y. Liu, J. Fang, and S. Li, “Determination of Lotus Leaf Alkaloids by Solid Phase Extraction Combined with High Performance Liquid Chromatography with Diode Array and Tandem Mass Spectrometry Detection,” Analytical Letters, vol. 46, no. 18, pp. 2846–2859, Dec. 2013. doi: 10.1080/00032719.2013.816960.
    Lotus (nelumbo) is a traditional Chinese medical herb, and alkaloids are responsible for its pharmaceutical activities. Therefore, a rapid, validated method to determine alkaloids in lotus leaves is useful for its quality control. The goal of this work was to develop and validate an analytical method for the identification and quantitation of the bioactive alkaloids of extracts from lotus leaf and classification of lotus germplasm based on alkaloid concentrations. Solid-phase extraction was used prior to high-performance liquid chromatography coupled with photodiode array detection at 272 nm and tandem electrospray ionization triple quadrupole mass spectrometry for analysis. Nuciferine and O-nornuciferine were the dominant compounds in lotus used for seed production and some used for flower production. However, anonaine, roemerine, and N-nornuciferine were abundant only in some lotus flowers. All the lotuses were classified into four groups by principal component analysis of alkaloid content: 10 flower and three seed lotuses characterized by high nuciferine and O-nornuciferine content; 12 flower lotuses with high N-nornuciferine, roemerine, and anonaine content; two flower lotuses with a relatively high content of all five alkaloids; and cultivars with a low alkaloid content and without dominant alkaloids.
  18. K.-L. Chen and R. Kirschner, “Fungi from Leaves of Lotus (Nelumbo Nucifera),” Mycological Progress, vol. 17, no. 1, pp. 275–293, Jan. 2018. doi: 10.1007/s11557-017-1324-y.
    In spite of the self-cleaning property of its leaves called the lotus effect, leaves of lotus (Nelumbo nucifera) provide a habitat for an unknown fungal diversity. The aim of this study was to detect and identify fungi from leaves of N. nucifera, including ectophytic, parasitic and endophytic fungi, in Taiwan using different collection strategies, as well as morphological and diverse molecular markers established in the different systematic groups of fungi. Among ectophytic and parasitic fungi, a new species of Dissoconium and of Pseudocercospora are described, respectively. Phyllosticta nelumbonis Sawada is transferred to Diaporthe. Among plant parasitic fungi, Erysiphe takamatsui and Ps. nymphaeacea are recorded in Taiwan for the first time. Euryale is recorded as a new host genus for Ps. nymphaeacea. The basidiomycetous yeast Fereydounia khargensis is recorded for the first time from living plants and in East Asia. Endophytic fungi from lotus were studied for the first time. From 1002 plant segments, 476 endophytic isolates were produced in culture, comprising 33 typical terrestrial species mainly belonging to the genera Colletotrichum (mainly C. siamense), Diaporthe (D. tulliensis and D. ueckerae) and Fusarium (F. solani species 6, hitherto known from clinical samples), as well as to Xylariaceae, but no Ingoldian fungi. Most isolates were from leaf laminas (71%) compared to those from petioles (29%). From this observation, we conclude that the fungi of the aquatic lotus plant appear to have terrestrial origin and, after dispersal by wind and in spite of the lotus effect, may enter the plant from the lamina. Only three species isolated as endophytes were also found as ectophytic or parasitic fungi.
  19. Y. Chen, R. Zhou, X. Lin, K. Wu, X. Qian, and S. Huang, “ISSR Analysis of Genetic Diversity in Sacred Lotus Cultivars,” Aquatic Botany, vol. 89, no. 3, pp. 311–316, Oct. 2008. doi: 10.1016/j.aquabot.2008.03.006.
    In this study, inter-simple sequence repeats (ISSR) markers were applied to assess genetic diversity and genetic relationships of 92 cultivars of sacred lotus (Nelumbo nucifera Gaertn.), one of the most famous flowers in China. Our results showed that sacred lotus exhibited a low level of genetic diversity (percentage of polymorphic bands, PPB=55.8%), which may result from its asexual mode of reproduction and long-term artificial selection. Clustering analyses indicated that these cultivars could be divided into two clades. Most cultivars of Chinese lotus species origin were included in one clade, and one cultivar of American lotus species origin was nested in the other clade. The hybrid cultivars from hybridization between the two subspecies were interspersed in these two clades. Seven cultivars native to Thailand formed a distinct subclade among the cultivars of Chinese lotus species origin. Genetic differentiation between two subspecies, and between cultivars from Thailand and other cultivars could be attributed to geographic isolation. The monophyly of three cultivars of Sanshui Winter Lotus and their closest relationships to Chinese lotus species origin suggests that they might have a common origin and may consist completely or mainly of genetic material from N. nucifera subsp. nucifera.
  20. J. Chen et al., “Regulations and Mechanisms of 1-Methylcyclopropene Treatment on Browning and Quality of Fresh-Cut Lotus (Nelumbo Nucifera Gaertn.) Root Slices,” Postharvest Biology and Technology, vol. 185, p. 111782, Mar. 2022. doi: 10.1016/j.postharvbio.2021.111782.
    The quality of fresh-cut lotus root slices is greatly affected by enzymatic browning. In this study, we investigated the effects of 1-methylcyclopropene treatment on browning and quality deterioration of lotus root slices. When fresh-cut lotus root slices were immersed in 0.1 mg L−1 1-MCP for 1 h and stored at 10 ℃ for twelve days, microbial growth, respiration, total phenolic content, phenylalanine ammonia lyase activity, polyphenol oxidase activity, and soluble quinone levels in fresh-cut lotus root slices were suppressed. Furthermore, 1-MCP treatment improved the antioxidant capacities, peroxidase activities, superoxide dismutase activities, catalase activities, and DPPH free radical scavenging rates in fresh-cut lotus root slices. In addition, 1-MCP treatment inhibited the production of reactive oxide species, which delayed quality deterioration of fresh-cut lotus root slices. Based on transcriptomics and real-time quantitative PCR results, gene expression levels of NnmetK2, NnACO, NnETR2, NnEIN3, and NnERF1 were up-regulated while expression levels of NnEBF1 were down-regulated. These findings indicate that 1-MCP inhibits ethylene biosynthesis and signal transduction, which suppresses a series of ethylene-induced physiological and biochemical reactions. Based on our findings, we propose a new strategy for inhibiting enzymatic browning in fresh-cut lotus root slices.
  21. G. Chen, M. Zhu, and M. Guo, “Research Advances in Traditional and Modern Use of Nelumbo Nucifera: Phytochemicals, Health Promoting Activities and Beyond,” Critical Reviews in Food Science and Nutrition, vol. 59, no. sup1, pp. S189–S209, Jun. 2019. doi: 10.1080/10408398.2018.1553846.
    Nelumbo nucifera, or sacred lotus, has been valuable for us to use as vegetable, functional food, and herb medicine for over 2,000 years. The purpose of this article is to systematically review the traditional/modern uses, chemical compositions and pharmacological activities on different parts of N. nucifera. Traditionally, this plant has been used to treat chronic dyspepsia, hematuria, insomnia, nervous disorders, cardiovascular diseases, and hyperlipidemia. Now, phytochemical investigations on different parts of N. nucifera have indicated a wide spectrum of at least 255 constituents belonging to different chemical groups, including proteins, amino acids, polysaccharides, starch, flavonoids, alkaloids, essential oils, triterpenoids, steroids, and glycosides. Meanwhile, its pharmacological activities, including anti-obesity, antioxidant, anti-inflammatory, cardiovascular, hepatoprotective, hypoglycemic, hypolipidemic, antitumor, memory-improving and antiviral activities, have also been reviewed, together with its applications in health food industry and clinic uses of its single plant or herbal formulae. Herein, this review will provide state-of-the-art overview on its traditional and modern uses in food industry and medicines, together with the comprehensive profiles of phytochemicals, and health promoting bioactivities of this valuable plant. In addition, the new perspectives and future challenges in the research on lotus are also outlined.
  22. L. Cheng, S. Li, J. Yin, L. Li, and X. Chen, “Genome-Wide Analysis of Differentially Expressed Genes Relevant to Rhizome Formation in Lotus Root (Nelumbo Nucifera Gaertn),” PLOS ONE, vol. 8, no. 6, p. e67116, Jun. 2013. doi: 10.1371/journal.pone.0067116.
    Lotus root is a popular wetland vegetable which produces edible rhizome. At the molecular level, the regulation of rhizome formation is very complex, which has not been sufficiently addressed in research. In this study, to identify differentially expressed genes (DEGs) in lotus root, four libraries (L1 library: stolon stage, L2 library: initial swelling stage, L3 library: middle swelling stage, L4: later swelling stage) were constructed from the rhizome development stages. High-throughput tag-sequencing technique was used which is based on Solexa Genome Analyzer Platform. Approximately 5.0 million tags were sequenced, and 4542104, 4474755, 4777919, and 4750348 clean tags including 151282, 137476, 215872, and 166005 distinct tags were obtained after removal of low quality tags from each library respectively. More than 43% distinct tags were unambiguous tags mapping to the reference genes, and 40% were unambiguous tag-mapped genes. From L1, L2, L3, and L4, total 20471, 18785, 23448, and 21778 genes were annotated, after mapping their functions in existing databases. Profiling of gene expression in L1/L2, L2/L3, and L3/L4 libraries were different among most of the selected 20 DEGs. Most of the DEGs in L1/L2 libraries were relevant to fiber development and stress response, while in L2/L3 and L3/L4 libraries, major of the DEGs were involved in metabolism of energy and storage. All up-regulated transcriptional factors in four libraries and 14 important rhizome formation-related genes in four libraries were also identified. In addition, the expression of 9 genes from identified DEGs was performed by qRT-PCR method. In a summary, this study provides a comprehensive understanding of gene expression during the rhizome formation in lotus root.
  23. L. Cheng, H. Liu, R. Jiang, and S. Li, “A Proteomics Analysis of Adventitious Root Formation after Leaf Removal in Lotus (Nelumbo Nucifera Gaertn.),” Zeitschrift für Naturforschung C, vol. 73, no. 9-10, pp. 375–389, Sep. 2018. doi: 10.1515/znc-2018-0011.
    The formation of adventitious roots (ARs) is an important process for lotus ( Nelumbo nucifera ), which does not have a well-formed main root. In lotus, the removal of leaves above the waterline significantly promoted AR formation, while the removal of leaves below the waterline inhibited AR formation. Proteins were identified using isobaric tags for relative and absolute quantization technique. The number of proteins decreased with increasing sequencing coverage, and most of the identified proteins had fewer than 10 peptides. In the A1/A0 and A2/A1 stages, 661 and 154 proteins showed increased abundance, respectively, and 498 and 111 proteins showed decreased abundance, respectively. In the B1/B0 and B2/B1 stages, 498 and 436 proteins showed increased abundance, respectively, and 358 and 348 proteins showed decreased abundance, respectively. Among the proteins showing large differences in abundance, 17 were identified as being related to AR formation. Proteins involved in the glycolytic pathway and the citrate cycle showed differences in abundance between the two types of leaf removal. The transcriptional levels of nine genes encoding relevant proteins were assessed by quantitative polymerase chain reaction. The results of this study illustrate the changes in metabolism after different types of leaf removal during AR formation in lotus.
  24. H. Choi, H.-T. Kim, B. E. Nam, Y. J. Bae, and J. G. Kim, “Effect of Initial Planting on Vegetation Establishment in Different Depth Zones of Constructed Farm Ponds,” Restoration Ecology, vol. n/a, no. n/a, p. e13488, Jul. 2021. doi: 10.1111/rec.13488.
    Farm ponds can be constructed and maintained to improve biodiversity and ecosystem functions in agricultural landscapes. Understanding the initial establishment process of vegetation in different water depths and effects of initial planting on this process is important for pond restoration. In the present study, two experimental farm ponds with or without initial planting were monitored for 5 years after their construction. These two ponds maintained high nutrients over 5 years from the beginning. The unplanted pond showed a rapid vegetation establishment and had more species and higher percent vegetation cover in 2–3 years than the reference pond. Planting accelerated this process by 1 year. Most of the early established species appeared to have originated from seed banks and/or seed rains. The species and functional diversity tended to decrease consistently since pond construction in the planted pond but increase initially and then decrease in the unplanted pond. The time when species composition between the unplanted pond and the planted pond became similar was delayed with the increase of water depth. Four years after pond construction, species composition was similar between the two ponds at all water depths. These results suggest that vegetation can establish quickly in farm ponds with fertile conditions and abundant seed sources through natural recolonization. The restoration approach for a farm pond should be decided considering the allowed time to success and the water depth. If the main goal of farm pond restoration is to improve species diversity, management practices may be more important than planting.
  25. P. Chu et al., “Proteomic and Functional Analyses of Nelumbo Nucifera Annexins Involved in Seed Thermotolerance and Germination Vigor,” Planta, vol. 235, no. 6, pp. 1271–1288, Jun. 2012. doi: 10.1007/s00425-011-1573-y.
    Annexins are multifunctional proteins characterized by their capacity to bind calcium ions and negatively charged lipids. Although there is increasing evidence implicating their importance in plant stress responses, their functions in seeds remain to be further studied. In this study, we identified a heat-induced annexin, NnANN1, from the embryonic axes of sacred lotus (Nelumbo nucifera Gaertn.) using comparative proteomics approach. Moreover, the expression of NnANN1 increased considerably in response to high-temperature treatment. Quantitative real-time PCR (qRT-PCR) revealed that the transcripts of NnANN1 were detected predominantly during seed development and germination in sacred lotus, implicating a role for NnANN1 in plant seeds. Ectopic expression of NnANN1 in Arabidopsis resulted in enhanced tolerance to heat stress in transgenic seeds. In addition, compared to the wild-type seeds, transgenic seeds ectopically expressing NnANN1 exhibited improved resistance to accelerated aging treatment used for assessing seed vigor. Furthermore, transgenic seeds showed enhanced peroxidase activities, accompanied with reduced lipid peroxidation and reduced ROS release levels compared to the wild-type seeds. Taken together, these results indicate that NnANN1 plays an important role in seed thermotolerance and germination vigor.
  26. J. W. H. Dacey, “Knudsen-Transitional Flow and Gas Pressurization in Leaves of Nelumbo,” Plant Physiology, vol. 85, no. 1, pp. 199–203, Sep. 1987. doi: 10.1104/pp.85.1.199.
  27. K. L. DeGroft and D. A. Francko, “Effect of Freezing on Germination of Nelumbo Lutea (Willd.) Pers. Seeds,” Journal of Freshwater Ecology, vol. 11, no. 3, pp. 373–376, Sep. 1996. doi: 10.1080/02705060.1996.9664460.
    We tested the ability of cold (4°C) and freezing (-20°C) treatments to scarify Nelumbo lutea seeds and thus enhance germination-frequency. Cold/freezing temperatures alone did not weaken seed coats enough to promote germination. Time-dependent seed germination curves and germination frequencies were statistically similar at all temperature treatments. The data suggest that freeze-thaw events in natural sediments may be insufficient to induce N. lutea germination.
  28. X. Deng et al., “Analysis of Isoquinoline Alkaloid Composition and Wound-Induced Variation in Nelumbo Using HPLC-MS/MS,” Journal of Agricultural and Food Chemistry, vol. 64, no. 5, pp. 1130–1136, Feb. 2016. doi: 10.1021/acs.jafc.5b06099.
    Alkaloids are the most relevant bioactive components in lotus, a traditional herb in Asia, but little is known about their qualitative and quantitative distributions. Here, we report on the alkaloid composition in various lotus organs. Lotus laminae and embryos are rich in isoquinoline alkaloids, whereas petioles and rhizomes contain trace amounts of alkaloids. Wide variation of alkaloid accumulation in lamina and embryo was observed among screened genotypes. In laminae, alkaloid accumulation increases during early developmental stages, reaches the highest level at full size stage, and then decreases slightly during senescence. Vegetative and embryogenic tissues accumulate mainly aporphine-type and bisbenzylisoquinoline-type alkaloids, respectively. Bisbenzylisoquinoline-type alkaloids may be synthesized mainly in lamina and then transported into embryo via latex through phloem translocation. In addition, mechanical wounding was shown to induce significant accumulation of specific alkaloids in lotus leaves.
  29. X. Deng et al., “The Establishment of an Efficient Callus Induction System for Lotus (Nelumbo Nucifera),” Plants, vol. 9, no. 11, p. 1436, Nov. 2020. doi: 10.3390/plants9111436.
    The lotus (Nelumbo nucifera) is one of the most popular aquatic plants in Asia, and has emerged as a novel model for studying flower and rhizome development, and primary and secondary metabolite accumulation. Here, we developed a highly efficient callus induction system for the lotus by optimizing a series of key factors that affect callus formation. The highest efficient callus production was induced on immature cotyledon and embryo explants grown on Murashige and Skoog (MS) basal medium containing an optimized combination of 3 mg/L 2,4-dichlorophenoxyacetic acid (2,4-D) and 0.5 mg/L 6-benzylaminopurine (6-BA). In addition, lotus callus induction was proven to be influenced by lotus genotypes, light conditions, the developmental stages of explants and the time of explant sampling. Collecting immature cotyledons from seeds of the genotype “Shilihe 1”, at 9 days post pollination, and to culture the explants in darkness, are proposed as the optimum conditions for lotus callus induction. Interestingly, highly efficient callus induction was also observed in explants of immature embryo derived aseptic seedlings; and a small amount of lotus benzylisoquinoline alkaloid (BIA) and obvious expression of BIA biosynthetic genes were detected in lotus callus.
  30. J. Deng, M. Li, L. Huang, M. Yang, and P. Yang, “Genome-Wide Analysis of the R2R3 MYB Subfamily Genes in Lotus (Nelumbo Nucifera),” Plant Molecular Biology Reporter, vol. 34, no. 5, pp. 1016–1026, Oct. 2016. doi: 10.1007/s11105-016-0981-3.
    MYB superfamily is one of the most abundant transcription factors in plants, which are involved in many physiological and biochemical processes. Among them, R2R3 MYB might be the most widely studied subfamily. Here, we characterized the gene structures, expressional patterns, and potential functions of 116 R2R3 MYB genes in a genome-wide manner in lotus (Nelumbo nucifera). Compared with those in Arabidopsis and grape, the lotus R2R3 MYB genes are conserved in exon’s number and length. Most of them constitute three exons and two introns with the first two exons being highly conserved in their length. According to the structure of exon and DNA-binding domain-imperfect repeat (R), these R2R3 MYB genes were classified into four groups. The expression of 13 candidate MYB genes that are related to flavonoid biosynthesis was analyzed in lotus different tissues. One gene expresses highly in all tissues and might positively regulate common flavonoid biosynthesis. Two genes might positively regulate anthocyanin and proanthocyanidin biosynthesis, respectively, resulting in the pigmentation of flower and seed. While five members that are similar to Zm38 are expressed lowly in different tissues. This genome-wide analysis of R2R3 MYB genes in lotus may provide more comprehensive insights on the structure and function of this gene family, and hence contribute a lot to plant’s genetic engineering in improving the colorization of flowers or other tissues.
  31. X. Deng et al., “Investigation of Benzylisoquinoline Alkaloid Biosynthetic Pathway and Its Transcriptional Regulation in Lotus,” Horticulture Research, vol. 5, no. 1, pp. 1–16, Jun. 2018. doi: 10.1038/s41438-018-0035-0.
    Lotus predominantly accumulates benzylisoquinoline alkaloids (BIAs), but their biosynthesis and regulation remain unclear. Here, we investigated structural and regulatory genes involved in BIA accumulation in lotus. Two clustered CYP80 genes were identified to be responsible for the biosynthesis of bis-BIAs and aporphine-type BIAs, respectively, and their tissue-specific expression causes divergence in alkaloid component between leaf and embryo. In contrast with the common (S)-reticuline precursor for most BIAs, aporphine alkaloids in lotus leaf may result from the (S)-N-methylcoclaurine precursor. Structural diversity of BIA alkaloids in the leaf is attributed to enzymatic modifications, including intramolecular C–C phenol coupling on ring A and methylation and demethylation at certain positions. Additionally, most BIA biosynthetic pathway genes show higher levels of expression in the leaf of high-BIA cultivar compared with low-BIA cultivar, suggesting transcriptional regulation of BIA accumulation in lotus. Five transcription factors, including three MYBs, one ethylene-responsive factor, and one basic helix–loop–helix (bHLH), were identified to be candidate regulators of BIA biosynthesis in lotus. Our study reveals a BIA biosynthetic pathway and its transcriptional regulation in lotus, which will enable a deeper understanding of BIA biosynthesis in plants.
  32. J. Deng et al., “Systematic Qualitative and Quantitative Assessment of Anthocyanins, Flavones and Flavonols in the Petals of 108 Lotus (Nelumbo Nucifera) Cultivars,” Food Chemistry, vol. 139, no. 1, pp. 307–312, Aug. 2013. doi: 10.1016/j.foodchem.2013.02.010.
    Petal colour is one of the major characteristics that determine the ornamental value of lotus. To assess the contribution of different flavonoids to this character, composition and content of anthocyanins, flavones and flavonols were analysed through high performance liquid chromatography coupled with photodiode array detection tandem electrospray ionisation triple quad mass spectrometry in 108 lotus cultivars with red, pink, yellow, white and red/white pied petal colours. Totally, five anthocyanins and fourteen flavones and flavonols were detected and quantified. In general, the yellow, white and pied species hardly contained any anthocyanins; red cultivars contain more than pink cultivars. Among the five anthocyanins, malvidin 3-O-glucoside was the most abundant one in all the cultivars that contain anthocyanin. The fourteen flavones and flavonols belonged to four groups based on their aglycones. Except for the yellow cultivars, kaempferol-derivatives were the most abundant one. These data might be helpful in lotus breeding for different colours.
  33. Y. Diao et al., “Nuclear DNA C-Values in 12 Species in Nymphaeales,” Caryologia, vol. 59, no. 1, pp. 25–30, Jan. 2006. doi: 10.1080/00087114.2006.10797894.
    Nuclear DNA C-values are the basic data of species and used in a strikingly wide range of biological fields. Genome size of the most species belonging to the three families Nulembonaceae, Cabombaceae and Nym-phaeaceae in Nymphaeales were not assessed, especially none of the species in Cabombaceae was reported so far. In present research, flow cytometry was used to assess the genome size of 12 species belonging to the three families in Nymphaeales and a standard squash technique to count the chromosome numbers of the tested materials. In the tested different species, 2C nuclear DNA contents ranged from 1.55 to 8.11 pg, which is more than fivefold variation. And chromosome numbers varied from 2n=16 to 2n=72. Differences between the species or among populations within a species were also recorded by statistical analysis. The tested species are all small genomes except two species of Victoria. The ploidies did not matched with their DNA contents very well in Nymphaea. Significant differences were found between the species of Nymphaea, between those of Victoria and among wild populations of Nelumbo nicifera., while no obvious differences were found between the species of Nelumbo and among the cultivars of Euryale ferox as well as among those of Brasenia schreberi, respectively.
  34. G. Dieringer, R. Leticia Cabrera, and M. Mottaleb, “Ecological Relationship between Floral Thermogenesis and Pollination in Nelumbo Lutea (Nelumbonaceae),” American Journal of Botany, vol. 101, no. 2, pp. 357–364, 2014. doi: 10.3732/ajb.1300370.
    • Premise of study: Floral thermogenesis is an unusual floral trait with a well-documented physiological process, and yet, there is limited understanding of how this trait influences plant reproduction. The current study was undertaken to gain a better understanding of how floral thermogenesis in Nelumbo lutea impacts pollinator attraction and consequent plant reproduction. • Methods: We conducted field studies on floral thermogenesis and thermoregulation, flower sexual development, floral visitation patterns, breeding system, pollen transfer dynamics, and floral scent production. • Key results: The most abundant visitors to the thermoregulatory flowers included the Phoridae (Diptera), Chrysomelidae (Coleoptera), and Hymenoptera. Chrysomelid beetles, particularly Diabrotica, were frequent visitors to both first-day female- and second-day bisexual-phase flowers, while phorid flies were most common in bisexual-phase flowers. Pollen transfer experiments indicated that Diabrotica was equally effective in depositing pollen on stigmas, as were the less frequent, but pollen-loaded halictid bees. • Conclusions: Flowers received a taxonomically wide assemblage of floral visitors and appear adapted to attract beetles, primarily Chrysomelidae and medium-sized bees. This study is the first to provide strong support that beetles can comprise the dominant portion of floral visitors and are as effective in pollen transfer as bees. Thermogenesis aids in dispersing the main floral scent component—1,4-dimethoxybenzene—attracting both chrysomelids and bees, while thermoregulation causes chrysomelid beetles to actively seek out new flowers for evening residence. This search behavior likely results in chrysomelids affecting cross-pollination.
  35. Y. F. Ding, H. Y. Cheng, and S. Q. Song, “Changes in Extreme High-Temperature Tolerance and Activities of Antioxidant Enzymes of Sacred Lotus Seeds,” Science in China Series C: Life Sciences, vol. 51, no. 9, pp. 842–853, Sep. 2008. doi: 10.1007/s11427-008-0107-8.
    Sacred lotus (Nelumbo nucifera Gaertn. ‘Tielian’) seed is long-lived and extremely tolerant of high temperature. Water content of lotus and maize seeds was 0.103 and 0.129 g H2O [g DW] −1, respectively. Water content, germination percentage and fresh weight of seedlings produced by surviving seeds gradually decreased with increasing treatment time at 100°C. Germination percentage of maize (Zea mays L. ‘Huangbaogu’) seeds was zero after they were treated at 100°Cfor 15 min and that of lotus seeds was 13.5% following the treatment at 100°C for 24 h. The time in which 50% of lotus and maize seeds were killed by 100°C was about 14.5 h and 6 min, respectively. With increasing treatment time at 100°C, relative electrolyte leakage of lotus axes increased significantly, and total chlorophyll content of lotus axes markedly decreased. When treatment time at 100°C was less than 12 h, subcellular structure of lotus hypocotyls remained fully intact. When treatment time at 100°C was more than 12 h, plasmolysis gradually occurred, endoplasmic reticulum became unclear, nuclei and nucleoli broke down, most of mitochondria swelled, lipid granules accumulated at the cell periphery, and organelles and plasmolemma collapsed. Malondialdehyde (MDA) content of lotus axes and cotyledons decreased during 0 −12 h of the treatment at 100°C and then increased. By contrast, the MDA content of maize embryos and endosperms increased during 5–10 min of the treatment at 100°C and then decreased slightly. For lotus seeds: (1) activities of superoxide dismutase (SOD) and glutathione reductase (GR) of axes and cotyledons and of catalase (CAT) of axes increased during the early phase of treatment at 100°C and then decreased; and (2) activities of ascorbate peroxidase (APX) and dehydroascorbate reductase (DHAR) of axes and cotyledons and of CAT of cotyledons gradually decreased with increasing treatment time at 100°C. For maize seeds: (1) activities of SOD and DHAR of embryos and endosperms and of GR of embryos increased during the early phase of the treatment at 100°C and then decreased; and (2) activities of APX and CAT of embryos and endosperms and of GR of endosperms rapidly decreased with increasing treatment time at 100°C. With decrease in seed germination, activities of SOD, APX, CAT, GR and DHAR of axes and cotyledons of lotus seeds decreased slowly, and those of embryos and endosperms of maize seeds decreased rapidly.
  36. Dingding Cao, Zhongyuan Lin, Longyu Huang, Rebecca Njeri Damaris, Ming Li, and Pingfang Yang, “A CONSTANS-LIKE Gene of Nelumbo Nucifera Could Promote Potato Tuberization,” Planta, vol. 253, no. 3, pp. 65–65, Mar. 2021. doi: 10.1007/s00425-021-03581-9.
    MAIN CONCLUSION: CONSTANS-LIKE 5 of Nelumbo nucifera is capable of promoting potato tuberization through CONSTANS-FLOWERING LOCUS T and gibberellin signaling pathways with a probable association with lotus rhizome enlargement. Lotus (Nelumbo nucifera) is an aquatic plant that is affiliated to the Nelumbonaceace family. It is widely used as an ornamental, vegetable, and medicinal herb with its rhizome being a popular vegetable. To explore the molecular mechanism underlying its rhizome enlargement, we conducted a systematic analysis on the CONSTANS-LIKE (COL) gene family, with the results, indicating that this gene plays a role in regulating potato tuber expansion. These analyses included phylogenetic relationships, gene structure, and expressional patterns of lotus COL family genes. Based on these analyses, NnCOL5 was selected for further study on its potential function in lotus rhizome formation. NnCOL5 was shown to be located in the nucleus, and its expression was positively associated with the enlargement of lotus rhizome. Besides, the overexpression of NnCOL5 in potato led to increased tuber weight and starch content under short-day conditions without changing the number of tubers. Further analysis suggested that the observed tuber changes might be mediated by affecting the expression of genes in CO–FT and GA signaling pathways. These results provide valuable insight in understanding the functions of COL gene as well as the enlargement of lotus rhizome.
  37. T. C. M. V. Do, T. D. Nguyen, H. Tran, H. Stuppner, and M. Ganzera, “Analysis of Alkaloids in Lotus (Nelumbo Nucifera Gaertn.) Leaves by Non-Aqueous Capillary Electrophoresis Using Ultraviolet and Mass Spectrometric Detection,” Journal of Chromatography A, vol. 1302, pp. 174–180, Aug. 2013. doi: 10.1016/j.chroma.2013.06.002.
    Lotus leaves are a popular remedy in Asia to treat obesity, insomnia and mental impairment; alkaloids are considered most relevant for bioactivity. In this study the first CE method for the determination of all major alkaloids ((−)-nuciferine, (−)-nornuciferine, (−)-caaverine, (−)-armepavine, (+)-norarmepavine, (+)-isoliensinine and (+)-pronuciferine) in Nelumbo nucifera leaves was developed. The optimum buffer showed to be a solution of 100mM ammonium acetate in a mixture of methanol, acetonitrile and water, also containing 0.6% acid acetic. Applied voltage, temperature and detection wavelength were 25kV, 30°C and 225nm; the required analysis time was 15min. By CE–MS all standard compounds could be assigned in positive ESI mode, and two minor alkaloids were tentatively identified (n-nornuciferine and roemerine). A mixture of water, methanol and acetic acid served as sheath liquid in these experiments. The NACE assay was fully validated and utilized to analyze Lotus leaf samples collected in different parts of Vietnam. Respective results revealed significant qualitative and quantitative differences depending on growing area and season. Yet, in all samples (−)-nuciferine (0.34–0.63%), (−)-armepavine (0.13–0.20%), and (+)-isoliensinine (0.06–0.19%) were the most dominant alkaloids. The total alkaloid content varied from 0.72 to 1.41%.
  38. C. Dong, M. Yang, H. Wang, and J. Mi, “Identification and Expression Analyses of Two Lotus (Nelumbo Nucifera) Dehydrin Genes in Response to Adverse Temperatures, ABA and IAA Treatments,” Biologia, vol. 72, no. 7, pp. 745–752, Jul. 2017. doi: 10.1515/biolog-2017-0080.
    Dehydrins (DHNs) play imperative role in the stress tolerance of high plants. In this study, the YSK2-type DHN (NnDHNl) and SK3-type DHN (NnDHN2) were investigated. The putative NnDHN1 and NnDNH2 shared conserved S-segment as well as K-segment, with distinct isoelectric points, kinase selectivities and number of functional motifs. Phylogenetic analysis revealed that NnDHNs were grouped with YnSKn- and SKn-type of DHNs in high plants. Additionally, NnDHN1 mRNA was only expressed in leaves and embryo. NnDHN2 mRNA was found in embryo, leaves, rhizome and leafstalk. NnDHNs mRNAs were not detected in root of N. nucifera. Real-time PCR assay demonstrated that the levels of NnDHN1 and NnDHN2 mRNAs were significantly elevated during cold temperature, while both were relatively stable in response to high temperature. Moreover, NnDHN2 mRNA was significantly increased after treatment with abscisic acid (ABA) or indole-3-acetic acid (IAA), while NnDHN1 mRNA was induced by ABA treatment with little change in response to IAA. These results illustrate two NnDHNs with partially overlapping function which exhibit different stress responsiveness.
  39. R. D. Doyle and R. M. Smart, “Potential Use of Native Aquatic Plants for Long-Term Control of Problem Aquatic Plants in Guntersville Reservoir, Alabama (Report 2),” US Army Corps of Engineers, Guntersville Reservoir, Alabama, A-93-6, Nov. 1995.
    Although some species of aquatic plants present significant management problems, most species benefit the environment by stabilizing sediments, improving water clarity, removing excess nutrients and pollutants from the water, moderating shoreline erosion, and providing quality food and habitat for fish, waterfowl, and other organisms. However, aquatic plants that form dense, thick canopies or mats at the water surface can negatively impact the ecological and economic values of a water body. These growth forms prevent full utilization of the water body’s potential by restricting exchange of gases between water and air, resulting in degraded water quality and poor habitat, and by interfering with navigational and recreational uses of the lake or reservoir. In Guntersville Reservoir, these growth forms are exhibited by both the weedy, normative aquatic macrophyte Myriophyllwn spicatum (Eurasian watermilfoil) and the nuisance, albeit native, mat-forming cyanobacteria Lyngbya wollei Gyngbya) (Doyle and Smart 1993a)...
  40. H. J. Ensikat, P. Ditsche-Kuru, C. Neinhuis, and W. Barthlott, “Superhydrophobicity in Perfection: The Outstanding Properties of the Lotus Leaf,” Beilstein Journal of Nanotechnology, vol. 2, no. 1, pp. 152–161, Mar. 2011. doi: 10.3762/bjnano.2.19.
    Beilstein Journal of Nanotechnology
  41. 二米炊烟ErmiChuiyan, “【ErmiChuiyan】Lotus Root: Flowers / Leaves / Root / Different Cooking Methods...” Sep-2020. https://www.youtube.com/watch?v=0_qY3sX7_C4.
    「※Please click "cc" to choose your subtitle language, which is on the "Settings" at the bottom right corner of the video.」 Although the lotus root grows in the mud, it’s a treasure. Its flowers, leaves, rhizomes, fruits, can be made into different dishes. Although the lotus has faded in autumn, the lotus root has matured. Made with fresh lotus root, lotus seed, and lotus leaves. Lotus Root Chicken, Lotus Root Fried, Lotus Root Pork Rib Soup. Osmanthus honey with lotus root, Tremella lotus seed soup, each of them is a rare delicacy. —————————————————— Please subscribe to my channel if you like my videos:https://goo.gl/gaNrqE (If you like my video, don’t forget to subscribe ~) Please click the link if you’d like to contribute subtitles to my video:https://goo.gl/FVRwfU —————————————————— #二米炊煙 #LotusRoot #蓮藕 #藕 #cooking #美食 #菜譜 #做飯 #鄉村生活 #ermichuiyan #haimétkhói #Chinesefood #masak #howto #foodrecipe
  42. M. Esterhuizen and Y. J. Kim, “Effects of Polypropylene, Polyvinyl Chloride, Polyethylene Terephthalate, Polyurethane, High-Density Polyethylene, and Polystyrene Microplastic on Nelumbo Nucifera (Lotus) in Water and Sediment,” Environmental Science and Pollution Research, Oct. 2021. doi: 10.1007/s11356-021-17033-0.
    Plastic waste is recognised as hazardous, with the risk increasing as the polymers break down in nature to secondary microplastics or even nanoplastics. The number of studies reporting on the prevalence of microplastic in every perceivable niche and bioavailable to biota is dramatically increasing. Knowledge of the ecotoxicology of microplastic is advancing as well; however, information regarding plants, specifically aquatic macrophytes, is still lacking. The present study aimed to gain more information on the ecotoxicological effects of six different polymer types as 4 mm microplastic on the morphology (germination and growth) and the physiology (catalase and glutathione S-transferase activity) of the rooted aquatic macrophyte, Nelumbo nucifera. The role of sediment was also considered by conducting all exposure both in a sediment-containing and sediment-free exposure system. Polyvinyl chloride and polyurethane exposures caused the highest inhibition of germination and growth compared to the control. However, the presence of sediment significantly decreased the adverse effects. Catalase activity was increased with exposure to polyvinyl chloride, polyurethane, and polystyrene, both in the presence and absence of sediment but more so in the sediment-free system. Glutathione S-transferase activity was significantly increased with exposure to polypropylene, polyvinyl chloride, and polyethylene terephthalate in the sediment-free system and exposure to polyethylene terephthalate and polyurethane in the absence of sediment. There was no clear correlation between the morphological and physiological effects observed. Further studies are required to understand the underlying toxicity mechanism of microplastics.
  43. P. P. Ferrer-Gallego, F. Boisset, and C. E. Jarvis, “Typification of the Sacred Lotus Nelumbo Nucifera (Nelumbonaceae),” TAXON, vol. 64, no. 1, pp. 156–159, 2015. doi: 10.12705/641.2.
    The typification of the sacred lotus, Nelumbo nucifera Gaertner (≡ Nymphaea nelumbo L.; Nelumbonaceae), is discussed. The designation of the nomenclatural type is based on an assessment of Linnaeus’s original material and the literature cited in the protologue. The name is lectotypified using a specimen from Paul Hermann’s herbarium at BM.
  44. D. A. Francko, “Studies on Nelumbo Lutea (Willd.) Pers. I. Techniques for Axenic Liquid Seed Culture,” Aquatic Botany, vol. 26, pp. 113–117, Jan. 1986. doi: 10.1016/0304-3770(86)90009-4.
    Several techniques for the sterile liquid germination and cultivation of Nelumbo lutea (Willd.) Pers. seeds were evaluated. Surface disinfestation with ethanol, hypochlorite and detergent washes did not eliminate bacterial or fungal contamination of cultures upon seedling germination. Sequential washings in ethanol/hypochlorite and two incubations in sterile media containing antibiotics (streptomycin sulfate and penicillin G) and a fungicide (Captan) induced sterility in ca. 34% of cultures, while maintaining >98% germination rates in inoculated seeds. Seedlings elongated and differentiated normally in sterile culture.
  45. Z. Gao, H. Li, X. Yang, P. Yang, J. Chen, and T. Shi, “Biased Allelic Expression in Tissues of F1 Hybrids between Tropical and Temperate Lotus (Nelumbo Nuicfera),” Plant Molecular Biology, vol. 106, no. 1, pp. 207–220, May 2021. doi: 10.1007/s11103-021-01138-8.
    The genome-wide allele-specific expression in F1 hybrids from the cross of tropical and temperate lotus unveils how cis-regulatory divergences affect genes in key pathways related to ecotypic divergence.
  46. H. Godwin and E. H. Willis, “The Viability of Lotus Seeds (Nelumbium Nucifera, Gaertn.),” New Phytologist, vol. 63, no. 3, pp. 410–412, 1964. doi: 10.1111/j.1469-8137.1964.tb07391.x.
  47. A. Goel, S. C. Sharma, and A. N. Sharga, “The Conservation of the Diversity of Nelumbo (Lotus) at the National Botanical Research Institute, Lucknow, India,” Botanic Gardens Conservation News, vol. 3, no. 6, pp. 52–54, 2001. https://www.jstor.org/stable/24798441.
  48. R. Gowthami, N. Sharma, R. Pandey, and A. Agrawal, “A Model for Integrated Approach to Germplasm Conservation of Asian Lotus (Nelumbo Nucifera Gaertn.),” Genetic Resources and Crop Evolution, vol. 68, no. 4, pp. 1269–1282, Apr. 2021. doi: 10.1007/s10722-021-01111-w.
    Asian lotus (Nelumbo nucifera Gaertn.) is one of the most aesthetic ornamental aquatic indigenous species, endowed with unique biological and nutritional traits. The seeds are icon of eternity due to remarkable longevity, known to survive in nature up to 1300 years. Leaves fend off dirt and water by a phenomenon of superhydrophobicity and flowers generate heat by thermoregulation to enhance pollination success. Hence, lotus is an extremely important model system to investigate the process of aging in plants and develop designer seeds with high vigour, viability and longevity. The whole plant is nutritionally rich with great medicinal value. While scientific attention on various aspects ranging from basic botany to genome sequencing in lotus is increasing worldwide, India has somewhat lagged behind in taking due scientific, agronomic and conservation initiatives. At present, the lotus is facing habitat loss because of global environmental change, vanishing of wetlands, over-exploitation from its natural habitat, and lack of concerted management efforts. Possible conservation strategies are proposed with a view to draw the attention of policy makers and stakeholders for prioritizing this species in various biodiversity-related programs of the country, besides promoting it as a potential food and nutraceutical species.
  49. N. M. Grant et al., “Distribution of Thermogenic Activity in Floral Tissues of Nelumbo Nucifera,” Functional Plant Biology, vol. 37, no. 11, pp. 1085–1095, Oct. 2010. doi: 10.1071/FP10024.
    Thermogenesis in Nelumbo nucifera (Gaertn.) has been known to scientists for many years; however, the extent of heating by different floral parts remains unclear. We present evidence that the receptacle, stamens and petals produce heat independently, and that the source of heating in these tissues is most likely the alternative oxidase (AOX). The temperatures of the receptacle, petals and stamens were significantly higher than non-thermogenic leaf tissue. After removal from the pedicel, the receptacle retained the most heat (8.1 ± 1.9°C above non-thermogenic tissue temperature) and the petals the least (2.8 ± 4.2°C), with the stamens intermediate. High AOX protein levels and flux through the AOX pathway (in all tissues) during the thermogenic period are consistent with AOX being the mechanism used for thermogenesis. Lipids and carbohydrates were investigated as possible substrates for thermogenesis. There was little change in total lipids during floral development; however, soluble carbohydrate levels decreased by 70% with the onset of thermogenesis. These sugars may fuel thermogenesis in the stamens. The localisation of AOX protein in the various floral parts and the evolutionary significance of its heating role are discussed.
  50. W. Große, “Pressurised Ventilation in Floating-Leaved Aquatic Macrophytes,” Aquatic Botany, vol. 54, no. 2, pp. 137–150, Jul. 1996. doi: 10.1016/0304-3770(96)01041-8.
    Gas through-flow by pressurised ventilation is well known in floating-leaved aquatic plants. The ventilation system operates via different structural features in different species: being through rhizomes in Nuphar species, is restricted to whorls in Nymphoides, or even individual leaves in Nelumbo. The convective through-flow in these aquatic plants is the result of a gas-pumping system, commonly powered by solar radiation, with air influx and pressurisation of the youngest, just emerged leaves along positive humidity and temperature gradients between leaf aerenchyma and the ambient atmosphere. It improves the internal aeration of submerged plant organs, has high physiological significance to the plant, and confers beneficial effects on the ecosystem. Since the acquisition of gas through-flow systems is not restricted to closely related plant species, but can be correlated with special ecosystemal conditions, it should be seen as a special adaptation to plant survival under oxygen shortage stress.
  51. C. Gu et al., “Characterization of Genes Encoding Granule-Bound Starch Synthase in Sacred Lotus Reveals Phylogenetic Affinity of Nelumbo to Proteales,” Plant Molecular Biology Reporter, vol. 31, no. 5, pp. 1157–1165, Oct. 2013. doi: 10.1007/s11105-013-0605-0.
    Starch is one of the main components of lotus rhizome and seed, and has a high potential influence on their eating and processing quality. However, little is known about the molecular mechanism underlying starch synthesis in lotus plants. Here, we report a genomewide identification and characterization of genes encoding granule-bound starch synthase in sacred lotus. Two GBSS genes, designated NnGBSSII-1 and NnGBSSII-2, have been identified in the genome of lotus variety “China Antique.” NnGBSSII-1 is expressed in all tested tissues, including leaf, flower, leafstalk, rhizome, and seed, whereas NnGBSSII-2 is a pseudogene and its transcripts have not been detected in any tissues. Overall, the expression level of NnGBSSII-1 gene is positively correlated with amylose accumulation in lotus rhizome and seed. Moreover, a phylogenetic analysis shows that GBSS genes from N. nucifera and N. lutea are clustered together, and they have a closer relationship with GBSS genes from Proteaceae. This result supports the hypothesis that Nelumbo is taxonomically placed in the lotus family, Nelumbonaceae, under the order Proteales. Our results will be helpful for future genetic improvement of starch quality in rhizome and lotus seed.
  52. S. Gui et al., “Improving Nelumbo Nucifera Genome Assemblies Using High-Resolution Genetic Maps and BioNano Genome Mapping Reveals Ancient Chromosome Rearrangements,” The Plant Journal, vol. 94, no. 4, pp. 721–734, 2018. doi: 10.1111/tpj.13894.
    Genetic and physical maps are powerful tools to anchor fragmented draft genome assemblies generated from next-generation sequencing. Currently, two draft assemblies of Nelumbo nucifera, the genomes of ‘China Antique’ and ‘Chinese Tai-zi’, have been released. However, there is presently no information on how the sequences are assembled into chromosomes in N. nucifera. The lack of physical maps and inadequate resolution of available genetic maps hindered the assembly of N. nucifera chromosomes. Here, a linkage map of N. nucifera containing 2371 bin markers [217 577 single nucleotide polymorphisms (SNPs)] was constructed using restriction-site associated DNA sequencing data of 181 F2 individuals and validated by adding 197 simple sequence repeat (SSR) markers. Additionally, a BioNano optical map covering 86.20% of the ‘Chinese Tai-zi’ genome was constructed. The draft assembly of ‘Chinese Tai-zi’ was improved based on the BioNano optical map, showing an increase of the scaffold N50 from 0.989 to 1.48 Mb. Using a combination of multiple maps, 97.9% of the scaffolds in the ‘Chinese Tai-zi’ draft assembly and 97.6% of the scaffolds in the ‘China Antique’ draft assembly were anchored into pseudo-chromosomes, and the centromere regions along the pseudo-chromosomes were identified. An evolutionary scenario was proposed to reach the modern N. nucifera karyotype from the seven ancestral eudicot chromosomes. The present study provides the highest-resolution linkage map, the optical map and chromosome level genome assemblies for N. nucifera, which are valuable for the breeding and cultivation of N. nucifera and future studies of comparative and evolutionary genomics in angiosperms.
  53. S. Gui et al., “The Mitochondrial Genome Map of Nelumbo Nucifera Reveals Ancient Evolutionary Features,” Scientific Reports, vol. 6, no. 1, p. 30158, Jul. 2016. doi: 10.1038/srep30158.
    Nelumbo nucifera is an evolutionary relic from the Late Cretaceous period. Sequencing the N. nucifera mitochondrial genome is important for elucidating the evolutionary characteristics of basal eudicots. Here, the N. nucifera mitochondrial genome was sequenced using single molecule real-time sequencing technology (SMRT) and the mitochondrial genome map was constructed after de novo assembly and annotation. The results showed that the 524,797-bp N. nucifera mitochondrial genome has a total of 63 genes, including 40 protein-coding genes, three rRNA genes and 20 tRNA genes. Fifteen collinear gene clusters were conserved across different plant species. Approximately 700 RNA editing sites in the protein-coding genes were identified. Positively selected genes were identified with selection pressure analysis. Nineteen chloroplast-derived fragments were identified and seven tRNAs were derived from the chloroplast. These results suggest that the N. nucifera mitochondrial genome retains evolutionarily conserved characteristics, including ancient gene content and gene clusters, high levels of RNA editing and low levels of chloroplast-derived fragment insertions. As the first publicly available basal eudicot mitochondrial genome, the N. nucifera mitochondrial genome facilitates further analysis of the characteristics of basal eudicots and provides clues of the evolutionary trajectory from basal angiosperms to advanced eudicots.
  54. H. B. Guo, “Cultivation of Lotus (Nelumbo Nucifera Gaertn. Ssp. Nucifera) and Its Utilization in China,” Genetic Resources and Crop Evolution, vol. 56, no. 3, pp. 323–330, May 2009. doi: 10.1007/s10722-008-9366-2.
    Lotus (Nelumbo nucifera Gaertn. ssp. nucifera), one of 12 aquatic species used as vegetable, has been cultivated for more than 2,000 years, and now has been widely cultivated in almost all provinces in China. The largest area under cultivation of lotus is located in the regions surrounding mid-down Yangtse River, including Hubei, Jiangsu, Zhejiang, Anhui, Jiangxi and Hunan provinces. According to different purposes or morphological differences, the Chinese lotus (N. nucifera ssp. nucifera) is usually classified into three types: rhizome lotus, seed lotus and flower lotus. Rhizome lotus is mainly cultivated in Hubei, Jiangsu, Anhui and Zhejiang provinces; Seed lotus in Jiangxi, Fujian and Hunan, and flower lotus in Wuhan, Hubei province, and Beijing. Up to the year 2002, a total of 572 lotus accessions (including landraces, cultivars and breeding lines) were conserved in National Garden of Aquatic Vegetable, Wuhan, Hubei province, including those collections from 153 counties in 18 provinces, and lines bred by breeders. Out of these accessions, 310 were rhizome lotus which contains 201 landraces and 109 breeding lines; 229 were flower lotus including 172 cultivars and 57 breeding lines; and the rest 33 were seed lotus with 18 cultivars.
  55. H. B. Guo, S. M. Li, J. Peng, and W. D. Ke, “Genetic Diversity of Nelumbo Accessions Revealed by RAPD,” Genetic Resources and Crop Evolution, vol. 54, no. 4, pp. 741–748, Jun. 2007. doi: 10.1007/s10722-006-0025-1.
    Total 65 lotus accessions in genus Nelumbo mainly collected from China, were subjected to random amplified polymorphic DNA (RAPD) markers to estimate the genetic diversity and to test the genetic basis of the relationships between morphotypes and molecular markers. Seventeen primers generated a total of 195 highly reproducible and discernible loci, among which 173 were polymorphic. Percent polymorphism varied from 66.7 to 100 with an average of 88.72, and five primers out of them, OPC05, OPG10, OPN20, OPP09 and OPS17, showed 100% polymorphism. A relatively high genetic diversity was detected among all the samples with the similarity coefficient values ranging from 0.45 to 0.85, and Nei’s gene diversity (h) 0.30, and Shannon index (I) 0.46. The UPGMA dendrogram clustered 65 accessions in four clusters and the clustering pattern showed two groups, N. nucifera ssp. nucifera and those accessions related to the American lotus, and some special cultivars, landraces, hybrids and the American lotus. Principal Coordinate Analysis (PCA) further indicated that the genetic diversity of Nelumbo accessions was not evenly distributed, instead, was presented by a clustered distribution pattern. Similar to the results revealed by the dendrogram, two main groups representing the two subspecies of N. nucifera, as well as some special landraces, cultivars of Chinese lotus, the Japanese lotus and hybrids out of the two groups were obtained. Neither the UPGMA dendrogram nor the PCA analysis exhibited strict relationship with geographic distribution and morphotypes among the accessions.
  56. S. C. Gupta, G. S. Paliwal, and R. Ahuja, “The Stomata of Nelumbo Nucifera: Formation, Distribution and Degeneration,” American Journal of Botany, vol. 55, no. 3, pp. 295–301, 1968. doi: 10.1002/j.1537-2197.1968.tb07379.x.
    Contrary to earlier reports, well-organized but fewer stomata develop on the lower surface of the leaves of Nelumbo nucifera Willd. during aerial growth. The stomata, however, become obliterated by the readjustment of neighboring epidermal cells. During initial stages of degeneration the guard cells show irregularly thickened walls, disintegrated nuclei, and highly vacuolated cytoplasm. Such abnormal features finally lead to the disappearance of stomata from the lower surface of leaves. The ontogeny, structure and distribution of stomata on leaves, perianth lobes, stamens, receptacles and carpels are described. The stomata are haplocheilic in development and are anomocytic (ranunculaceous) at maturity. The concept of a meristemoid and the significance of this study in taxonomy and phylogeny are discussed.
  57. T. F. Hall and W. T. Penfound, “The Biology of the American Lotus, Nelumbo Lutea (Wild.) Pers,” The American Midland Naturalist, vol. 31, no. 3, pp. 744–758, 1944. doi: 10.2307/2421417.
    1. The American lotus, Nelumbo lutea (Willd.) Pers. is a perennial, emergent, aquatic herb with fibrous, adventitious roots, slender rhizomes, elongated tubers, erect and floating leaves, large fragrant, yellow flowers, and an obconic toms containing acorn-like fruits. 2. The roots are characterized by a large, aerenchymous cortex; the rhizome, tubers, petioles, and peduncles are distinguished by a small cortex, scattered bundles, and four to eight large air tubes; the midveins of the leaf are characterized by upper and lower masses of collenchyma, closed bundles, and air tubes in a groundwork of parenchyma. 3. The rapid colonization of new areas by lotus is accomplished by the elongated rhizomes and to a less extent by tubers and fruits. Complete inundation of lotus colonies for a continuous period of two weeks or dewatering for one month destroyed all the leaves and flowers and many of the rhizomes and tubers. 4. The American lotus provides favorable environmental conditions for the development of malaria mosquitoes, but it can be controlled effectively by recurrent cutting of the leaves where the water is relatively turbid.
  58. Y.-C. Han, C.-Z. Teng, S. Zhong, M.-Q. Zhou, Z.-L. Hu, and Y.-C. Song, “Genetic Variation and Clonal Diversity in Populations of Nelumbo Nucifera (Nelumbonaceae) in Central China Detected by ISSR Markers,” Aquatic Botany, vol. 86, no. 1, pp. 69–75, Jan. 2007. doi: 10.1016/j.aquabot.2006.09.007.
    To obtain accurate estimates of population structure for purposes of conservation planning for wild lotus (Nelumbo nucifera Gaertn.) in central China, genetic diversity among and within six populations, and clonal diversity within another two populations of the species were analyzed. The genetic diversity was high (percentage of polymorphic bands, PPB=90.0%; Shannon’s information index, I=0.383±0.234) at the species level, but low within individual study populations (PPB=35.8%; Shannon’s information index I=0.165±0.241). The mean coefficient of gene differentiation (Gst) was 0.570, indicating that 43.0% of the genetic diversity resided within the population. Analysis of molecular variance (AMOVA) indicated that 50.47% of the genetic diversity among the study populations was attributed to geographical location while 12.3% was attributed to differences in their habitats. An overall value of mean estimated number of gene flow (Nm=0.377) indicated that there was limited gene flow among the sampled populations. The level of clonal diversity found within the populations was considerably high (Simpson’s diversity index, D=0.985) indicating that clonal diversity contributes to a major extent to the overall genetic variation in the genetic structure of N. nucifera. On the basis of the high Gst and D values detected in this study we recommend that any future conservation plans for this species should be specifically designed to include those representative populations with the highest genetic variation for both in situ conservation and germplasm collection expeditions.
  59. Y.-C. Han, C.-Z. Teng, G. R. Wahiti, M.-Q. Zhou, Z.-L. Hu, and Y.-C. Song, “Mating System and Genetic Diversity in Natural Populations of Nelumbo Nucifera (Nelumbonaceae) Detected by ISSR Markers,” Plant Systematics and Evolution, vol. 277, no. 1, pp. 13–20, Jan. 2009. doi: 10.1007/s00606-008-0096-x.
    In this study, outcrossing rates and genetic diversity in natural populations of Nelumbo nucifera were investigated. The estimated multilocus outcrossing rate (tm) based on 28 ISSR loci was over 90%. Analysis of genetic diversity revealed that this index was high at the species level (Hs = 0.325, I = 0.514), but low within the individual study populations (Hs = 0.148, I = 0.212). Gst-B was 0.547 and Nm was 0.414. The results of AMOVA indicated that 54.6% of the variation was due to the difference between the regions and 45.4% to the variation within the region. Although the populations were predominantly outcrossing, most of the genetic diversity was attributed to geographical effects instead of their habitats because low sexual recruitment and clonal growth deeply reduced the genetic diversity within the populations. On the basis of the high tm, Gst-B and low Nm values, we recommended that any future conservation plans should include both in situ conservation and germplasm collection.
  60. V. Hayes, E. L. Schneider, and S. Carlquist, “Floral Development of Nelumbo Nucifera (Nelumbonaceae),” International Journal of Plant Sciences, vol. 161, no. S6, pp. S183–S191, Nov. 2000. doi: 10.1086/317577.
    The floral development of Nelumbo nucifera was compared with that noted in previous studies of Nelumbo, Nymphaeaceae, and other basal angiosperms. Important features include developmental evidence of only two sepals, development of an androecial ring meristem, and an apocarpous gynoecium composed of ascidiate carpels that become embedded in an expanded receptacle. Secretory papillate trichomes cover the stigma and line the stylar canal. The unique apocarpous gynoecium, which lacks a conical residual floral apex, and a greatly expanded receptacle distinguish Nelumbonaceae from the Nymphaeaceae, as does the distinctive androecial ring. Nelumbo is characterized by polysymmetric floral development, with some organs originating spirally (petals) and some in simultaneous whorls (stamens and carpels). This pattern of floral development, as well as the pattern of carpel closure by secretion, is common in several paleoherbs and eudicots and indicates phylogenetic affinity between Nelumbonaceae and basal angiosperms. Because of its unique floral development and anatomy, Nelumbo appears to be an isolated member of the eudicot clade.
  61. P. Hongpakdee and S. Ruamrungsri, “Enhanced Flowering of Sacred Lotus (Nelumbo Nucifera Gaertn.) by Extending the Photoperiod with Supplemental Lighting Techniques,” Acta Horticulturae, no. 1171, pp. 47–52, Sep. 2017. doi: 10.17660/ActaHortic.2017.1171.7.
    Sacred lotus (Nelumbo nucifera Gaertn.) is one of the most popular cut flowers in Thailand and other Asian countries, commonly used for religious purposes. However, basic knowledge regarding the control of flowering is relatively lacking. The effects of photoperiod on growth of N. nucifera were investigated using a completely randomized design experiment with six day-length conditions: 1) 11 h for 2 months, 2) 11 h for 1 month and then to 13 h for 1 month, 3) 11 h for 1 month and then 15 h for 1 month, 4) 13 h for 1 month and then 11 h for 1 month, 5) 13 h for 2 months and 6) 13 h for 1 month and then 15 h for 1 month. Each pot was planted with 90 stem cuttings, filled to 2/3 of the pot level with natural clay soil and then filled to 3/4 of the pot level with tap water after planting. The stolons were grown under natural conditions (ambient temperature 30/18°C with an 11 h day length in mild winter from January-February 2013). Night interruption after 6:00 pm was used to induce a long day-length condition (13 and 15 h). None of photoperiod conditions affected the total number of leaves. However, extending the photoperiod from 11 to 15 h and from 13 to 15 h and a constant photoperiod of 13 h produced the largest leaves. Shortened photoperiods from 13 to 11 h decreased the flowering percentage and number of flowers. Extended photoperiods from 11 to 13 h, 11 to 15 h and 13 to 15 h only increased the flowering percentage compared with that of constant photoperiods of 11 and 13 h. Nevertheless, the extended photoperiod delayed the date of visible flowering.
  62. P. Hongpakdee, C. Samranyat, and S. Ruamrungsri, “Propagation of Sacred Lotus (Nelumbo Nucifera Gaertn.) by Stolon Cutting with Active Bud and Different Nodes Number,” Acta Horticulturae, no. 1263, pp. 233–240, Nov. 2019. doi: 10.17660/ActaHortic.2019.1263.30.
    Sacred lotus is one of the valuable economic cut flowers in Thailand and Asian countries. The growers usually propagate by cutting stolon segment, since this technique allows cloning to supply uniform planting stock. They however consider that only longer stolon cutting with many nodes and active buds would be suitable as propagules. To clarify this condition, the effect of vegetative propagation technique by node number on plant growth and flowering in ’Sattabutsaya’ sacred lotus was conducted with 3 stolon cutting types i.e., stolon cutting with 1, 2 and 3 nodes, respectively, using a completely randomized design. All active stolons were planted singly in 1.20×1.00 m circular concrete pots, filled with 1/3 clay soil and 2/3 tap water under natural conditions (May to August 2016). It was found that increasing node number of cutting material increased in total leaf number at 6 weeks after planting (WAP). None of the stolon cutting treatments affected total leaf area, new internode length, flower stalk length and flower length at 3 months after planting. Nevertheless, planting sacred lotus with single node stolon cuttings gave the lowest new stolon length, flower number, percentage of flowering and delay more 2-3 weeks of flowering. Triple nodes of propagated stolon cutting gave the best result in plant dry weight components (flower, pod, new stolon and fibrous root). Planting sacred lotus stolon cuttings with 2 or 3 nodes did not affect total leaf number, new stolon length and girth and visible flowering time. It could be concluded that stolon cuttings with at least two nodes for propagative material is sufficient for creating plant stock for flowering lotus production.
  63. J. Hu et al., “Comparative Analysis of Genetic Diversity in Sacred Lotus (Nelumbo Nucifera Gaertn.) Using AFLP and SSR Markers,” Molecular Biology Reports, vol. 39, no. 4, pp. 3637–3647, Apr. 2012. doi: 10.1007/s11033-011-1138-y.
    The sacred lotus (Nelumbo nucifera Gaertn.) is an aquatic plant of economic and ornamental importance in China. In this study, we developed twenty novel sacred lotus SSR markers, and used AFLP and SSR markers to investigate the genetic diversity and genetic relationships among 58 accessions of N.nucifera including 15 seed lotus, 12 rhizome lotus, 24 flower lotus and 7 wild lotus. Our results showed that sacred lotus exhibited a low level of genetic diversity, which may attribute to asexual reproduction and long-term artificial selection. A dendrogram based on both AFLP and SSR clustering data showed that: (1) the seed lotus accessions and rhizome lotus accessions were distinctly clustered into different groups, which indicated the significant genetic differentiation between them. This may be attributed to the two modes of reproduction and lack of genetic exchange; (2) the accessions of Thailand wild lotus were separated from other wild lotus accessions. This implied that the Thailand lotus might be genetically differentiated from other wild lotuses. In addition, Mantel test conducted gave highly significant correlation between AFLP-SSR data and each of the AFLP and SSR ones, with the values of r = 0.941 and r = 0.879, respectively, indicating the higher efficiency of the combination of these techniques (AFLP and SSR) in estimation and validation of the genetic diversity among the accession of sacred lotus. This knowledge of the genetic diversity and genetic relatedness of N. nucifera is potentially useful to improve the current strategies in breeding and germplasm conservation to enhance the ornamental and economic value of sacred lotus.
  64. J. Hu, S. Gui, Z. Zhu, X. Wang, W. Ke, and Y. Ding, “Genome-Wide Identification of SSR and SNP Markers Based on Whole-Genome Re-Sequencing of a Thailand Wild Sacred Lotus (Nelumbo Nucifera),” PLOS ONE, vol. 10, no. 11, p. e0143765, Nov. 2015. doi: 10.1371/journal.pone.0143765.
    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.
  65. J. Hu, J. Jin, Q. Qian, K. Huang, and Y. Ding, “Small RNA and Degradome Profiling Reveals miRNA Regulation in the Seed Germination of Ancient Eudicot Nelumbo Nucifera,” BMC Genomics, vol. 17, no. 1, p. 684, Aug. 2016. doi: 10.1186/s12864-016-3032-4.
    MicroRNAs (miRNAs) play important roles in plant growth and development. MiRNAs and their targets have been widely studied in model plants, but limited knowledge is available concerning this small RNA population and their targets in sacred lotus (Nelumbo nucifera Gaertn.).
  66. L. Huang, M. Li, D. Cao, and P. Yang, “Genetic Dissection of Rhizome Yield-Related Traits in Nelumbo Nucifera through Genetic Linkage Map Construction and QTL Mapping,” Plant Physiology and Biochemistry, vol. 160, pp. 155–165, Mar. 2021. doi: 10.1016/j.plaphy.2021.01.020.
    Lotus (Nelumbo nucifera) is a perennial aquatic plant with great value in ornamentation, nutrition, and medicine. Being a storage organ, lotus rhizome is not only used for vegetative reproduction, but also as a popular vegetable in Southeast Asia. Rhizome development, especially enlargement, largely determines its yield and hence becomes one of the major concerns in rhizome lotus breeding and cultivation. To obtain the genetic characteristic of this trait, and discover markers or genes associated with this trait, an F2 population was generated by crossing between temperate and tropical cultivars with contrasting rhizome enlargement. Based on this F2 population and Genotyping-by-Sequencing (GBS) technique, a genetic map was constructed with 1475 bin markers containing 12,113 SNP markers. Six traits associated with rhizome yield were observed over 3 years. Quantitative trait locus (QTL) mapping analysis identified 22 QTLs that are associated with at least one of these traits, among which 9 were linked with 3 different intervals. Comparison of the genes located in these three intervals with our previous transcriptomic data showed that light and phytohormone signaling might contribute to the development and enlargement of lotus rhizome. The QTLs obtained here could also be used for marker-assisted breeding of rhizome lotus.
  67. L. Huang et al., “Whole Genome Re-Sequencing Reveals Evolutionary Patterns of Sacred Lotus (Nelumbo Nucifera),” Journal of Integrative Plant Biology, vol. 60, no. 1, pp. 2–15, 2018. doi: 10.1111/jipb.12606.
    Sacred lotus (Nelumbo nucifera or lotus) is an important aquatic plant in horticulture and ecosystems. As a foundation for exploring genomic variation and evolution among different germplasms, we re-sequenced 19 individuals from three cultivated temperate lotus subgroups (rhizome, seed and flower lotus), one wild temperate lotus subgroup (wild lotus), one tropical lotus group (Thai lotus) and an outgroup (Nelumbo lutea). Through genetic diversity and polymorphism analysis by non-missing SNP sites widely distributed in the whole genome, we confirmed that wild and Thai lotus exhibited greater differentiation with a higher genomic diversity compared to cultivated lotus. Rhizome lotus had the lowest genomic diversity and a closer relationship to wild lotus, whereas the genomes of seed and flower lotus were admixed. Genes in energy metabolism process and plant immunity evolved rapidly in lotus, reflecting local adaptation. We established that candidate genes in genomic regions with significant differentiation associated with temperate and tropical lotus divergence always exhibited highly divergent expression pattern. Together, this study comprehensive and credible interpretates important patterns of genetic diversity and relationships, gene evolution, and genomic signature from ecotypic differentiation of sacred lotus.
  68. M.-H. Im, B.-W. Kim, Y.-S. Park, S.-Y. Yang, C.-E. Song, and B.-G. Heo, “Effects of Scarification, Temperature and Sulfuric Acid Treatments on Seed Germination of White Lotus (Nelumbo nucifera),” Korean Journal of Plant Resources, vol. 25, no. 1, pp. 7–13, 2012. doi: 10.7732/kjpr.2012.25.1.007.
    연 종자의 발아 특성을 구명하기 위해 전남 무안산 백련 종자를 이용해 종피의 파상부위, 발아 온도, 황산용액 처리 농도 및 시간별에 따른 종자 무게 변화와 발아율을 조사하였다. 종자를 파상하지 않고 파종한 것은 15일째가 되어도 전혀 발아가 되지 않았다. 그러나 기부를 파상하여 }25\^{∘}C{에서 발아를 시킨 것은 6일 만에 100%가 발아 되어 가장 우수한 결과를 보였다. 종자를 물에 침적시켰을 때 물에 뜬 것(0.90 g)과 가라앉은 것(1.18 g) 모두 발아 되었으나 침적된 종자를 }25\^{∘}C{에서 발아 시켰을 때 발아세가 가장 좋았다. 황산처리는 80% 용액에서 40-160분간 침적처리 했을때 6일째에 100%가 발아해 가장 우수한 발아세를 나타내었다. 종자 채취 후 0개월, 12개월 및 24개월 된 것을 파종한 결과 발아세와 발아율에 차이가 없었다. 위와 같은 결과는 연의 종자번식에 도움이 될 것으로 사료된다. This study was conducted to determine the effects of scarification temperature, and sulfuric acid treatments on seed germination of white lotus collected from the Muan districts, Jeonnam in Korea. Without scarification, white lotus seeds were not germinated at all at 15 days after seeding. However, seeds sacrificed at basal parts showed 100% germination rate at }25\^{∘}C{ 6 days after seeding. All the seeds floated (0.90 g) and soaked (1.18 g) in the water were completely germinated. Especially, the lotus seeds soaked in the water at }25\^{∘}C{ showed high germination rate. Seeds treated with 80% sulfuric acid for 40-160 hours were germinated completely within 6 days after seeding. No difference in seed germination rate of white lotus stored up to 0, 12 and 24 months after harvest was observed. Overall results would be useful means for propagation and production of white lotus.
  69. W. Imsabai and W. G. van Doorn, “Effects of Auxin, Gibberellin, and Cytokinin on Petal Blackening and Flower Opening in Cut Lotus Flowers (Nelumbo Nucifera),” Postharvest Biology and Technology, vol. 75, pp. 54–57, Jan. 2013. doi: 10.1016/j.postharvbio.2012.05.015.
    Cut lotus flowers (Nelumbo nucifera Gaertn. cv. Saddabutra), which are sold as closed buds, fail to open and show rapid petal blackening when placed in vase water. We investigated the effect on bud opening and petal blackening of treatments with an auxin, a gibberellic acid, and two cytokinins. Continuous treatment of cut flowers placed in an aqueous solution containing ≥0.1mM naphthylacetic acid (NAA) hastened petal blackening and resulted in stem curvature, but lower concentrations (0.01–10μM) had no effect. Depending on the experiment, continuous treatment with 0.03–0.45mM of the gibberellin GA3 delayed petal blackening by 0.5–1.5d (controls lasted 4d), but in experiments during the hot/rainy season (May–September) GA3 had no effect. At 25–100μM the cytokinin benzyladenine (6-benzylaminopurine; BA) delayed petal blackening by about 1.0d. Similarly, the cytokinin thidiazuron (TDZ) delayed petal blackening by about 1.0d, at 1.25–2.5μM. Pulse treatments had similar or better effects. A 3–12h pulse treatment with 0.45mM GA3 or with 10μM TDZ delayed the time to petal blackening by 1.1–2.3d. However, none of these treatments promoted bud opening. It is concluded, nonetheless, that a pulse treatment with GA3 or TDZ seems promising for practice.
  70. W. Imsabai, S. Ketsa, and W. G. van Doorn, “Role of Ethylene in the Lack of Floral Opening and in Petal Blackening of Cut Lotus (Nelumbo Nucifera) Flowers,” Postharvest Biology and Technology, vol. 58, no. 1, pp. 57–64, Oct. 2010. doi: 10.1016/j.postharvbio.2010.04.007.
    Lotus flowers (Nelumbo nucifera) are cut at the bud stage, kept dry and brought to a temple as part of a religious tradition. When placed in water, even if this is done immediately after cutting, the flowers fail to open and show rapid petal blackening. We tested the role of ethylene in these processes. The rate of ethylene production by the cut flowers (which had been held dry for 2h after harvest and were then placed in water) transiently increased, with a maximum 9–15h after harvest. This was accompanied by an increase in ACC synthase activity, while the ACC oxidase activity remained unchanged. Exogenous ethylene, applied at 0.1–10μL/L, for 3h prior to vase life or continuously during vase life, had no effect on petal blackening. Continuous inclusion in the vase water of ethephon, a compound that releases ethylene, stimulated the rate of ethylene production, and accelerated petal blackening. Treatment with 1-methylcyclopropene (1-MCP), an inhibitor of the ethylene receptor, reduced the rate of ethylene production and delayed initial petal blackening by about 2d. None of these treatments had an effect on flower opening. Two partial cDNAs were isolated from the petals, one encoding a 1-aminocyclopropane-1-carboxylic acid (ACC) synthase (Nn-ACS) and the other an ACC oxidase (Nn-ACO). The transcript abundance of both genes had increased by 12h after the onset of vase life. 1-MCP decreased the transcript abundance of both genes, whereas ethephon increased it. It is concluded that cut lotus flowers placed in water show an increase in the rate of ethylene production, apparently due to increased ACS activity and the expression of one or more ACS genes. Ethylene seems part of the causal chain leading to early petal blackening, but it is not the cause of the lack of flower opening.
  71. F. Ishizuna and N. Tsutsumi, “Flower Bud Formation of Sacred Lotus (Nelumbo Nucifera Gaertn.): A Case Study of ‘Gyozankouren’ Grown in a Container,” HortScience, vol. 49, no. 4, pp. 516–518, Apr. 2014. doi: 10.21273/HORTSCI.49.4.516.
    The genus Nelumbo consists of two species, N. nucifera and N. lutea. N. nucifera is an ornamental and edible plant that is widely cultivated. Earlier studies of sacred lotus (N. nucifera) flowers focused mainly on morphology, phyllotaxis, leaf arrangements, and flower development. During the growing season, sacred lotus produces one foliage leaf at each node. Flower buds emerge from the abaxial side of the basal part of the foliage leaf. However, the number of blooming flowers is much less than the number of foliage leaves. Little is known concerning flower bud formation during lotus plant development. This is the first experimental study to reveal that every node has one flower bud even in the dormant shoot apex and that most of the formed flower buds aborted in the course of floral development. Our results suggest that flower bud formation of sacred lotus is independent of daylength. On the other hand, whether a formed bud reaches blooming may depend on environmental factors.
  72. M. R. Islam, Y. Zhang, Z.-Z. Li, H. Liu, J.-M. Chen, and X.-Y. Yang, “Genetic Diversity, Population Structure, and Historical Gene Flow of Nelumbo Lutea in USA Using Microsatellite Markers,” Aquatic Botany, vol. 160, p. 103162, Jan. 2020. doi: 10.1016/j.aquabot.2019.103162.
    American lotus (Nelumbo lutea Willd.) is a popular plant with ornamental, medicinal, and culinary value. Studies related to the genetic diversity and population structure of N. lutea are limited. We sampled 326 individuals from 19 populations of wild N. lutea across its distribution range in the USA and genotyped them using 12 microsatellite markers. A total of 274 alleles were amplified by 12 microsatellites with a mean of 22.83. We observed a low level of genetic diversity for American lotus (Ae\,= 2.40 and HE\,= 0.459) in the USA. The analysis of molecular variance (AMOVA) exhibited closer variation among populations (42.00%) than within populations (58.00%) of N. lutea, with strong genetic differentiation observed (FST\,= 0.419). There was no significant correlation between genetic and geographic distances. The Bayesian analysis and neighbor-joining dendrogram displayed strong population structure. A very low level of historical gene flow (NM\,= 0.073) was observed among the populations. Fourteen of the 19 populations showed a demographic bottleneck, indicating the vulnerability of the American lotus populations. We thus proposed priority population sites for the conservation of N. lutea via both ex-situ and in-situ techniques.
  73. G. K. Jaganathan, D. Song, W. Liu, Y. Han, and B. Liu, “Relationship between Seed Moisture Content and Acquisition of Impermeability in Nelumbo Nucifera (Nelumbonaceae),” Acta Botanica Brasilica, vol. 31, pp. 639–644, 2017-Oct-Dec. doi: 10.1590/0102-33062017abb0188.
    ABSTRACT Seeds of Nelumbo nucifera do not imbibe water, and thus have physical dormancy (PY). However, a proportion of seeds are permeable to water, and so we hypothesized that variation in moisture content is a reason for the development of both permeable and impermeable seeds. The permeable proportion of seeds present in a lot collected from Suzhou, China, was separated using an imbibition test. The permeable proportion had an average moisture content of 15.6 %, compared with 8.5 % for impermeable seeds. Drying permeable seeds above silica gel to 10 % and 8 % f. wb., resulted in 77 and 100 % impermeable seeds, respectively, compared with no impermeable seeds at 15 % moisture content. Dried to 10 % moisture content, and incubated above water in an airtight container, 46 % of the seeds reverse impermeability. Permeable seeds with 15 % moisture content maintained above LiCl2 (RH=70 %) did not develop impermeability after three months of storage. The seeds dried to 6 % moisture content and stored above water in an airtight container showed no increase in moisture. Based on these results, we conclude that there is a strong relationship between moisture content and the onset of impermeability in this species.
  74. T. A. N. G. C.-Q. JI Hong-Wei and T. A. N. G. C.-Q. JI Hong-Wei, “Photosystem Development in Dark-grown Lotus (Nelumbo nucifera) Seedlings (in English),” Journal of Integrative Plant Biology, vol. 43, no. 11, p. 1129, Nov. 2001. https://www.jipb.net/CN/abstract/abstract25697.shtml.
    $article.zhaiYao_CN.replaceAll(
  75. R. Z. Jia, R. Ming, and Y. J. Zhu, “Genome-Wide Analysis of Nucleotide-Binding Site (NBS) Disease Resistance (R) Genes in Sacred Lotus (Nelumbo Nucifera Gaertn.) Reveals Their Transition Role During Early Evolution of Land Plants,” Tropical Plant Biology, vol. 6, no. 2, pp. 98–116, Sep. 2013. doi: 10.1007/s12042-013-9122-4.
    Nucleotide-binding site (NBS) containing genes comprise the largest class in identified plant resistance genes. A total of 137 NBS class resistance genes were identified from the newly sequenced sacred lotus genome (Nelumbo nucifera Gaertn.) through a reiterative computational sequence analysis. Three distinct groups of NBS-encoding genes were classified: 5 with Toll/interleukin-1 receptor homology (TIR) domain at N-terminal (TIR-NBS [-LRR (leucine-rich repeat)]), 37 with CC (coiled coil) domain (CC-NBS [-LRR]), and 95 with neither TIR nor CC at N-terminal (NBS [-LRR]). Sequence analysis revealed high divergence of NBS-LRR genes in sacred lotus. The result of cluster and syntenic analysis of NBS genes suggested a duplication and recombination event, which is consistent with the correspondent result of whole genome analysis. In addition, we also identified 52 NBS genes which have a putative NACHT domain embedded in the NBS domains. This characteristic has only been reported in animals, fungi and bacteria, suggesting that NACHT and NBS domains shared a similar ancient origin; and sacred lotus NBS (NACHT) genes may represent a transition role during the early evolution of disease resistance in land plants.
  76. Jiao Deng, Juanjuan Li, Mengyue Su, Zhongyuan Lin, Lei Chen, and Pingfang Yang, “A bHLH Gene NnTT8 of Nelumbo Nucifera Regulates Anthocyanin Biosynthesis,” Plant physiology and biochemistry, vol. 158, pp. 518–523, Jan. 2021. doi: 10.1016/j.plaphy.2020.11.038.
    Lotus is an important aquatic ornamental plant, whose flower color is one of the key horticultural traits that determines its ornamental value. Previous studies revealed that anthocyanins largely determined the red color of lotus flower, which are also the main component that has beneficial effects on human health. However, the regulation mechanism of flower pigmentation in lotus flower remains unclear. In the present study, in order to further understand the regulatory mechanism underlying the anthocyanin biosynthesis, a bHLH gene NnTT8 was characterized to be phylogenetically close to AtTT8 and the bHLH proteins from other plant species that have been indicated to be involved in the positive regulation of anthocyanin biosynthesis. Complementation analysis in Arabidopsis tt8 mutant showed that NnTT8 could function similarly to AtTT8 in regulating anthocyanin and proanthocyanin biosynthesis. An MYB transcription factor capable of interacting with NnTT8 was also characterized from lotus. The identification of a bHLH transcription factor playing regulatory roles in anthocyanin biosynthesis is crucial, as it might help to obtain more in-depth insight into the coloration of lotus and help in breeding high anthocyanin content lotus variety that can be explored for lotus flower beverages.
  77. Q. Jin et al., “Interactions between Ethylene, Gibberellin and Abscisic Acid in Regulating Submergence Induced Petiole Elongation in Nelumbo Nucifera,” Aquatic Botany, vol. 137, pp. 9–15, Jan. 2017. doi: 10.1016/j.aquabot.2016.11.002.
    Nelumbo nucifera (Lotus) responds to complete submergence with increasing petiole elongation which allows leaves to emerge above the water surface thereby restoring contact with the atmosphere. We carried out a detailed study of this growth response under submergence, to elucidate the role of several phytohormones. Under submergence, petiole elongation of lotus was mainly caused by cell division on the apical part of petiole. Exogenous phytohormone application studies and endogenous ethylene measurements revealed that ethylene, abscisic acid (ABA), and gibberellin (GA) were all involved in regulating petiole elongation under submergence. In response to submergence, ethylene accumulated first, which then initiated elongation growth of lotus petiole. An exogenously applied ethylene releasing compound, ethrel, and its precursor, 1-aminocyclopropane-1-carboxylate (ACC) promoted petiole elongation under submergence. An inhibitor of ethylene action (silver thiosulphate, STS) could partially block petiole elongation under submergence. Exogenous GA treatment caused a similar promotion in petiole elongation. Lotus seedlings treated with different concentrations of paclobutrazol (PAC, a GA inhibitor) induced a dose-dependent decrease of petiole elongation under submergence. In contrast, ABA treatment eliminated submergence-induced petiole elongation completely, while its biosynthesis inhibitor, Tungstate, and GA could reverse the inhibitory effect of ABA. However, the effect of ethrel was not affected by Tungstate. Meanwhile, an ABA biosynthesis inhibitor enhanced the effect of GA and ethrel. In summary, under submergence, increased ethylene altered the balance between a growth-inhibiting hormone (ABA) and a growth-promoting (GA) and contributed to the submergence-induced petiole elongation in lotus.
  78. J. A. Jones, “Overcoming Delayed Germination of Nelumbo Lutea,” Botanical Gazette, vol. 85, no. 3, pp. 341–343, May 1928. doi: 10.1086/333846.
    1. In spite of the presence of a large number of small pores in the seed coat, the seeds of Nelumbo lutea did not absorb water and germinate, even after eighteen months’ soaking at room temperature. 2. When the seed coats were broken without injuring the embryos, the seeds germinated without exception. 3. Seeds may be prepared for germination by treatment for five hours with concentrated sulphuric acid, following by thorough washing in tap water and later by drying on a screen to eliminate immediate germination. Seeds so treated may be stored by the commercial grower and shipped dry as ordered.
  79. F. K, “The Lotus Seed, Nelumbo Nucifera Gaertn. as a New Host Plant of the Adzuki Bean Weevil, Callosobruchus Chinensis L. (Coleoptera:Bruchidae).,” Applied Entomology and Zoology, vol. 22, no. 3, pp. 388–389, 1987. https://jglobal.jst.go.jp/en/detail?JGLOBAL_ID=200902016956684272.
    Article “The lotus seed, Nelumbo nucifera Gaertn. as a new host plant of the adzuki bean weevil, Callosobruchus chinensis L. (Coleoptera:Bruchidae).” Detailed information of the J-GLOBAL is a service based on the concept of Linking, Expanding, and Sparking, linking science and technology information which hitherto stood alone to support the generation of ideas. By linking the information entered, we provide opportunities to make unexpected discoveries and obtain knowledge from dissimilar fields from high-quality science and technology information within and outside JST.
  80. W. Kai and Z. Zhiwei, “Occurrence of Arbuscular Mycorrhizas and Dark Septate Endophytes in Hydrophytes from Lakes and Streams in Southwest China,” International Review of Hydrobiology, vol. 91, no. 1, pp. 29–37, 2006. doi: 10.1002/iroh.200510827.
    In this study, the colonization of arbuscular mycorrhizas (AM) and dark septate endophytes (DSE) in 140 specimens of 32 hydrophytes collected from four lakes and four streams in southwest China were investigated. The arbuscular mycorrhizal fungi (AMF) and DSE colonization in these hydrophytes were rare. Typical AM structures were observed in one of the 25 hydrophytic species collected in lakes and six of the 17 species collected in streams. Spores of 10 identified AMF species and an unidentified Acaulospora sp. were isolated from the sediments. The identified AMF came from the four genera, Acaulospora, Gigaspora, Glomus and Scutellospora . Glomus and G. mosseae were the dominant genus and species respectively in these aquatic environments. The presence of DSE in hydrophytes was recorded for the first time. DSE occurred in one of the 25 hydrophyte species collected in lakes and three of the 17 species collected in streams. (© 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)
  81. M. Katori, K. Nomura, and K. Yoneda, “Propagation of Flowering Lotus (Nelumbo Nucifera Gaertn) by Rhizome Straps, without Enlarged Rhizomes,” Japanese Journal of Tropical Agriculture, vol. 46, no. 3, pp. 195–197, 2002. doi: 10.11248/jsta1957.46.195.
  82. A. Khatfan, Z. Li, L.-Q. Chen, N. Riablershirun, V. Sathornviriyapong, and N. Juntawong, “Pollen Viability, Germination, and Seed Setting of Nelumbo Nucifera,” ScienceAsia, vol. 40, no. 6, p. 384, 2014. doi: 10.2306/scienceasia1513-1874.2014.40.384.
  83. \relax Y. A. Khrolenko, M. S. Yatsunskaya, T. \relax Y. Gorpenchenko, and T. A. Bezdeleva, “Development of Nelumbo Komarovii Grossh. (Nelumbonaceae) from Seeds under Artificial Conditions,” Inland Water Biology, vol. 12, no. 1, pp. 18–25, Jun. 2019. doi: 10.1134/S1995082919050122.
    The development of Nelumbo komarovii Grossh from seeds under artificial conditions has been studied. Five phases of N. komarovii morphogenesis are identified: the seed, the primary (main) shoot, the main-shoot system and anchoring shoots, curtain, and tuberiform structure. The phases of morphogenesis correlate with periods of ontogenesis and age-related stages of individuals.
  84. M.-J. Kim, W. Nelson, C. A. Soderlund, and D. R. Gang, “Next-Generation Sequencing-Based Transcriptional Profiling of Sacred Lotus ‘China Antique,’” Tropical Plant Biology, vol. 6, no. 2, pp. 161–179, Sep. 2013. doi: 10.1007/s12042-013-9130-4.
    Rhizomes are underground stems that serve various purposes including vegetative propagation, invasion of new territory, and bioactive compound synthesis and storage. An important rhizomatous plant is sacred lotus (Nelumbo nucifera), which is prized in Asia as a medicine and a food. RNA-seq and total transcriptome analysis of rhizomes and other lotus tissues was applied to identify genes involved in rhizome growth, development and metabolism. Root, petiole, rhizome internode, and leaf tissues were used for single-read RNA-seq analysis. Two whole transcriptome paired-end read libraries from rhizome apical tip and elongation zone tissues were also generated in order to survey gene expression profiles. In this analysis, 22,803 genes were expressed: 20,476 in rhizome apical meristem and elongation zone, 17,171 in rhizome internode, 16,656 in leaf, 19,457 in root, and 16,845 in petiole. Gene ontology (GO) analysis indicated that “other membrane”, “nucleotide binding”, and “other cellular processes” were highly represented in the expressed genes. A total of 231 genes displayed rhizome-specific expression including several transcription factors, protein kinases, cytochromes P450 and a sulfate transporter. GOseq analysis showed that genes in the “molecular function” GO category and several genes related to cell proliferation based on KEGG IDs were preferentially up-regulated in rhizome tissue. In addition, 1,251 possible exon-skipping events were observed in 1,149 gene models. These results provide valuable insight into gene expression profiles and regulation in sacred lotus, and the identified rhizome-specific genes provide insight into important processes involved in the biology and development of sacred lotus rhizomes.
  85. O. G. Koren, M. S. Yatsunskaya, and O. V. Nakonechnaya, “Low Level of Allozyme Polymorphism in Relict Aquatic Plants of the Far East Nelumbo Komarovii Grossh. and Euryale Ferox Salisb.,” Russian Journal of Genetics, vol. 48, no. 9, pp. 912–919, Sep. 2012. doi: 10.1134/S1022795412060075.
    Using allozyme analysis, genetic variation of two relict aquatic plants from Primorsky krai, Komarov lotus (Nelumbo komarovii Grossh.) and Gorgon plant (Euryale ferox Salisb.), was examined. The absence of allozyme variation in the Primorye populations of Nelumbo komarovii along with low polymorphism level in the population of Euryale ferox (P95 = 7.69; A = 1.07; Ho = 0.072; He = 0.038) was demonstrated. Since the data for the species examined are reported for the first time ever, the phenotypes and genetic interpretation of the enzyme systems tested are presented. The isoenzyme profiles of N. komarovii were compared with the data reported for N. nucifera from China. The absence of allozyme variation in N. komarovii, along with extremely low level of variation revealed for E. ferox, is discussed in association with the evolutionary histories of these species, their dispersal after the Pleistocene-Holocene cooling, and survival on this territory in range boundaries.
  86. S. S. Kreunen and J. M. Osborn, “Pollen and Anther Development in Nelumbo (Nelumbonaceae),” American Journal of Botany, vol. 86, no. 12, pp. 1662–1676, Dec. 1999. doi: 10.2307/2656664.
    The Nelumbonaceae are a small family of aquatic angiosperms comprising Nelumbo nucifera and Nelumbo lutea. Historically, the genus has been considered to be closely related to Nymphaeales, however new systematic work has allied Nelumbo with lower eudicots, particularly Platanus. In recent years, studies of pollen development have contributed greatly to the understanding of phylogenetic relationships, but little has been known about these events in Nelumbo. In this paper, pollen and anther development are morphologically described for the first time in N. lutea. A comprehensive ontogenetic sequence is documented, including the sporogenous tissue, microspore mother cell, tetrad, free spore, and mature pollen grain stages. The deposition of a microspore mother cell coat and callose wall, the co-occurrence of both tetrahedral and tetragonal tetrads, the formation of a primexine in tetrads, and primexine persistence into the late free spore stage are shown. The majority of exine development occurs during the free spore stage with the deposition of a tectate-columellate ectexine, a lamellate endexine, and an unusual granular layer below and intermixed with the endexine lamellae. A two-layered intine forms rapidly during the earliest mature pollen stage. Major events of anther development documented include the degradation of a secretory-type tapetum during the free spore stage and the rapid formation of U-shaped endothecial thickenings in the mature pollen grain stage. The majority of mature pollen grains are tricolpate, however less common monosulcate and diaperturate grains also develop. Co-occurring aperture types in Nelumbo have been suggested to be an important transition in angiosperm aperture number. However, aperture variability in Nelumbo may be correlated with the lateness of aperture ontogeny in the genus, which occurs in the early free spore stage. This character, as well as other details of pollen and anther ontogeny in Nelumbo, are compared to those of Nymphaeales and Platanus in an effort to provide additional insight into systematic and phylogenetic relationships. Although Nelumbo is similar to both groups in several characters, the ontogenetic sequence of the genus is different in many ways.
  87. N. Kubo, M. Hirai, A. Kaneko, D. Tanaka, and K. Kasumi, “Classification and Diversity of Sacred and American Nelumbo Species: The Genetic Relationships of Flowering Lotus Cultivars in Japan Using SSR Markers,” Plant Genetic Resources, vol. 7, no. 3, pp. 260–270, Dec. 2009. doi: 10.1017/S1479262109356580.
    The water lotus, genus Nelumbo, contains two species, the sacred (Nelumbo nucifera) and American lotuses (Nelumbo lutea). Hundreds of flowering lotus cultivars are currently known. However, their classification is unclear. For the classification of Nelumbo cultivars, in addition to 35 simple sequence repeat (SSR) markers recently developed, we have developed 17 and 16 of new Nelumbo SSR markers from SSR-enriched genomic libraries and expressed sequence tag (EST) data, respectively. Out of these 68 SSRs, along with SSRs recently published by others, 52 showed clear polymorphisms in 98 Nelumbo samples. A total of 300 alleles were observed, ranging from 2 to 11 alleles per locus, with an average of 5.77. Alleles specific for the American lotus-derived cultivars and a cluster of the American lotus-derived cultivars on a neighbour-joining tree confirmed genetic differences between N. lutea and N. nucifera. In addition, a possible differentiation between Chinese and Japanese cultivars was also suggested. Parentage analysis using the SSR markers confirmed four known parentages and predicted currently-unknown parentages of six cultivars. The present data have demonstrated that site-specific, co-dominant SSR markers enable more accurate classification, identification and comparison of Nelumbo species.
  88. M. Kumar, S. Chikara, M. K. Chand, and A. K. Bhatnagar, “Accumulation of Lead, Cadmium, Zinc, and Copper in the Edible Aquatic Plants Trapa Bispinosa Roxb. and Nelumbo Nucifera Gaertn,” Bulletin of Environmental Contamination and Toxicology, vol. 69, no. 5, pp. 0649–0654, Nov. 2002. doi: 10.1007/s00128-002-0110-x.
  89. J. Kunitomo et al., “Alkaloids of Nelumbo Nucifera,” Phytochemistry, vol. 12, no. 3, pp. 699–701, Mar. 1973. doi: 10.1016/S0031-9422(00)84467-2.
    The alkaloids of leaves of Nelumbo nucifera Gaertn. were examined using combined GLC-MS. The occurrence of four new alkaloids, dehydroroemerine (XII), dehydronuciferine (XI), dehydroanonaine (XIII) and N-methylisococlaurine (III) were revealed, besides the known roemerine (V), nuciferine (VI), anonaine (VII), pronuciferine (IV), N-nornuciferine (VIII), nornuciferine (IX), amepavine (I) and N-methylcoclaurine (II).
  90. H. Kurashita et al., “Chemical and Microbial Characteristics of Blackening Disease in Lotus (Nelumbo Nucifera Gaertn.) Caused by Hirschmanniella Diversa Sher,” Agronomy, vol. 11, no. 12, p. 2517, Dec. 2021. doi: 10.3390/agronomy11122517.
    The lotus (Nelumbo nucifera Gaertn.) is widely cultivated in Asia, but a blackening disease in the lotus tuber, called “kurokawa-senchu-byo”, is a serious problem caused by the Hirschmanniella diversa Sher plant-parasitic nematode. To effectively control the disease, we must elucidate the blackening mechanisms; therefore, in this study, we performed a soil chemical analysis and an evaluation of the disease level in the lotus cultivation fields, identified the chemical components of the black spots on the lotus surface, and performed a 16S rRNA gene-based microbial community analysis of the black spots. Using linear regression analysis, a positive linear relationship with a strong correlation between the damage index values and fertilizer components such as P2O5 was observed. As a result of scanning electron microscopy and energy-dispersive X-ray spectroscopy analysis, phosphorus (P) and iron (Fe) were found to be concentrated in the black spots of the lotus tubers. Furthermore, we found that the concentrations of P and Fe in the black spots were 1.5- and 2.7-fold higher, respectively, than those found in the healthy parts of the lotus tubers. A 16S rRNA gene analysis revealed that dissimilatory Fe(III)-reducing bacteria (DIRB) were predominant in the black spots, suggesting that these bacteria are important to the formation of P and Fe compounds in the black spots.
  91. W. La-ongsri, C. Trisonthi, and H. Balslev, “Management and Use of Nelumbo Nucifera Gaertn. in Thai Wetlands,” Wetlands Ecology and Management, vol. 17, no. 4, pp. 279–289, Aug. 2009. doi: 10.1007/s11273-008-9106-6.
    Management and use of Nelumbo nucifera Gaertn.—the lotus plant—was studied in 58 wetlands distributed throughout Thailand. Although traditionally harvested in extractive systems depending on natural wetlands, N. nucifera is now increasingly being managed. Two hundred eighty informants mentioned 20 different uses, mainly for food, medicine, and religious rites, in both subsistence and cash economies. The uses of N. nucifera appeared to be rather generalized throughout the country even if informants in the northern and central regions knew of more uses and ways of commercializing its products, possibly related to the abundance of wetlands and natural stand in those regions, and maybe also cultural differences.
  92. H.-L. Li, “Classification and Phylogeny of Nymphaeaceae and Allied Families,” The American Midland Naturalist, vol. 54, no. 1, pp. 33–41, 1955. doi: 10.2307/2422174.
  93. Y. Li, N. Awasthi, N. Nosova, and J.-X. Yao, “Comparative Study of Leaf Architecture and Cuticles of Nelumbo Changchangensis from the Eocene of Hainan Island, China, and the Two Extant Species of Nelumbo (Nelumbonaceae),” Botanical Journal of the Linnean Society, vol. 180, no. 1, pp. 123–137, 2016. doi: 10.1111/boj.12361.
    Fossil leaves of Nelumbo changchangensis, collected from the Eocene of Hainan Island, China, were studied and compared with those of the extant species of Nelumbo, N. nucifera Gaertn. and N. lutea Willd. The fossil leaves have all the specialized features of extant Nelumbo in leaf architecture, except that the organization of the areolae looks much more irregular than that of extant Nelumbo. Comparisons of the cuticle and epicuticular ultrastructure indicate that: (1) N. changchangensis resembles N. nucifera in that anticlinal cell walls of the lower epidermis are straight along the major veins and near leaf bases and are shallowly undulate with U- to V-shaped undulations inside the areolae; (2) N. changchangensis differs from N. lutea in that anticlinal cell walls of the lower epidermis of the latter are deeply undulate with U-, V- to reversed Ω-shaped undulations inside the areolae; and (3) epicuticular wax crystals are more densely distributed on the leaves of N. changchangensis and N. nucifera than they are in N. lutea. These findings shed significant light on the cuticle differentiation of fossil and extant Nelumbo species. The morphometric comparisons indicate that almost all the synapomorphies of extant Nelumbo were already present by the Eocene, © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180, 123–137.
  94. J. Li et al., “Comprehensive Analysis and Functional Studies of WRKY Transcription Factors in Nelumbo Nucifera,” International Journal of Molecular Sciences, vol. 20, no. 20, p. 5006, Jan. 2019. doi: 10.3390/ijms20205006.
    The WRKY family is one of the largest transcription factor (TF) families in plants and plays central roles in modulating plant stress responses and developmental processes, as well as secondary metabolic regulations. Lotus (Nelumbo nucifera) is an aquatic crop that has significant food, ornamental and pharmacological values. Here, we performed an overview analysis of WRKY TF family members in lotus, and studied their functions in environmental adaptation and regulation of lotus benzylisoquinoline alkaloid (BIA) biosynthesis. A total of 65 WRKY genes were identified in the lotus genome and they were well clustered in a similar pattern with their Arabidopsis homologs in seven groups (designated I, IIa-IIe, and III), although no lotus WRKY was clustered in the group IIIa. Most lotus WRKYs were functionally paired, which was attributed to the recently occurred whole genome duplication in lotus. In addition, lotus WRKYs were regulated dramatically by salicilic acid (SA), jasmonic acid (JA), and submergence treatments, and two lotus WRKYs, NnWRKY40a and NnWRKY40b, were significantly induced by JA and promoted lotus BIA biosynthesis through activating BIA biosynthetic genes. The investigation of WRKY TFs for this basal eudicot reveals new insights into the evolution of the WRKY family, and provides fundamental information for their functional studies and lotus breeding.
  95. Z. Li, X. Liu, R. W. Gituru, N. Juntawong, M. Zhou, and L. Chen, “Genetic Diversity and Classification of Nelumbo Germplasm of Different Origins by RAPD and ISSR Analysis,” Scientia Horticulturae, vol. 125, no. 4, pp. 724–732, Jul. 2010. doi: 10.1016/j.scienta.2010.05.005.
    In this study, RAPD and ISSR markers were used to investigate the genetic diversity and genetic relationships among different germplasm of Nelumbo including 70 Chinese ornamental cultivars, 7 wild Thai genotypes, 2 Nelumbo lutea genotypes and 8 hybrids of Nelumbo nucifera and N. lutea. High genetic diversities of 96.4% and 91.2% respectively were detected in the Nelumbo accessions using RAPD and ISSR markers. A dendrogram based on both RAPD and ISSR clustering data indicated that: (1) the genotypes of N. nucifera and N. lutea from different geographical origins were clustered into different groups. This indicated significant genetic differentiation attributed to extensive periods of geographical isolation and lack of gene exchange; (2) the Thai wild genotypes were separated from Chinese genotypes. This indicated genetic divergence between germplasm from Southeast Asia and that from China. Geographical location appears to have affected genetic diversity due to adaptation of the plants to the different environments. A new Southeastern Asia Lotus category is suggested as an addition to the current lotus cultivars classification system; (3) data on three morphological traits (namely: plant size, petal shape and flower color), showed that only the data on plant size was consistent with the dendrogram constructed from molecular data. This finding suggests that using data on genetic relationships in combination with morphological characteristics would serve to improve the classification system of lotus cultivars currently in use. The finding of previously unknown germplasm in this study indicated the potential of RAPD and ISSR techniques in identifying and managing lotus resources. Both marker techniques are potentially useful in improving the current strategies in breeding and germplasm conservation to enhance the ornamental and economic value of lotus.
  96. C. Li, H.-bo Mo, D.-ke Tian, Y.-xian Xu, J. Meng, and K. Tilt, “Genetic Diversity and Structure of American Lotus (Nelumbo Lutea Willd.) in North America Revealed from Microsatellite Markers,” Scientia Horticulturae, vol. 189, pp. 17–21, Jun. 2015. doi: 10.1016/j.scienta.2015.03.026.
    American lotus (Nelumbo lutea Willd.) was an important food resource for native American people and continues to be a popular ornamental plant worldwide. However, its habitats have been suffering from destruction by both natural and human’s activities. Little is known about the genetic diversity of this species. In this study, genetic diversity and structure of American lotus in North America were evaluated based on seven populations using 19 microsatellite markers. Sixty alleles were obtained from 19 loci in seven populations. The number of effective alleles (NE=1.18), Shannon’s information index (I=0.17), expected heterozygosity (HE=0.11) indicated a low level of genetic diversity in populations. High level of genetic differentiation among populations was reflected by FST=0.81 and AMOVA analysis. UPGMA dendrogram and STRUCTURE analysis clustered the seven populations into two clusters or gene pools. Population of Ocheechobee Lake in Florida formed an independent cluster or gene pool and is separated from other populations. Populations from Ontario and Maryland likely originated from individuals surviving floods or were introduced not long ago from other places. Further study is necessary based on more extensive sampling from other distribution regions and countries to better understand genetic diversity of American lotus.
  97. Z.-Z. Li et al., “Genetic Diversity Comparisons of Wild Populations of Nelumbo Nucifera (Nelumbonaceae) in Russia and China Using Microsatellite Markers,” Plant Systematics and Evolution, vol. 306, no. 5, p. 79, Aug. 2020. doi: 10.1007/s00606-020-01708-1.
    The wild sacred lotus Nelumbo nucifera is considered as an endangered aquatic plant in China and Russia. To date, the assessments and comparisons of the genetic diversity of wild populations of N. nucifera in Russia and China are very limited. Here, we sampled 15 wild populations of N. nucifera, including 208 individuals from these two key areas and genotyped using 13 microsatellite markers. Our results revealed 154 multilocus genotypes (MLGs) among individuals. Excluding the NS049 Locus that showed significant null alleles, we further assessed the genetic diversity of 154 MLGs in our study. The Chinese populations exhibited slightly lower genetic diversity compared to the Russian ones. Overall, the study showed low levels of genetic diversity (HO\,= 0.174 and HE\,= 0.251) with substantial genetic differentiation (FST\,= 0.547) in N. nucifera populations. We found a greater genetic variation residing among populations (54.74%) and a significant correlation (R2\,= 0.8756, P\,= 0.01) between genetic and geographic distances. In Bayesian analyses, we found that the populations were clustered into three distinct genetic pools (K\,= 3) supported by principal coordinates analysis (PCoA) and neighbor-joining (NJ) tree. We observed low gene flow between the genetic pools (NM\,= 0.2649). Nine of the 15 populations showed recent bottlenecks, demonstrating the N. nucifera populations were vulnerable or endangered in Russia and China. Therefore, based on the available genetic variations and the levels of threats, we recommended important population sites for the preservation of N. nucifera through complimentary conservation approaches.
  98. M. Li, I. Hameed, D. Cao, D. He, and P. Yang, “Integrated Omics Analyses Identify Key Pathways Involved in Petiole Rigidity Formation in Sacred Lotus,” International Journal of Molecular Sciences, vol. 21, no. 14, p. 5087, Jan. 2020. doi: 10.3390/ijms21145087.
    Sacred lotus (Nelumbo nucifera Gaertn.) is a relic aquatic plant with two types of leaves, which have distinct rigidity of petioles. Here we assess the difference from anatomic structure to the expression of genes and proteins in two petioles types, and identify key pathways involved in petiole rigidity formation in sacred lotus. Anatomically, great variation between the petioles of floating and vertical leaves were observed. The number of collenchyma cells and thickness of xylem vessel cell wall was higher in the initial vertical leaves’ petiole (IVP) compared to the initial floating leaves’ petiole (IFP). Among quantified transcripts and proteins, 1021 and 401 transcripts presented 2-fold expression increment (named DEGs, genes differentially expressed between IFP and IVP) in IFP and IVP, 421 and 483 proteins exhibited 1.5-fold expression increment (named DEPs, proteins differentially expressed between IFP and IVP) in IFP and IVP, respectively. Gene function and pathway enrichment analysis displayed that DEGs and DEPs were significantly enriched in cell wall biosynthesis and lignin biosynthesis. In consistent with genes and proteins expressions in lignin biosynthesis, the contents of lignin monomers precursors were significantly different in IFP and IVP. These results enable us to understand lotus petioles rigidity formation better and provide valuable candidate genes information on further investigation.
  99. H. Li et al., “Nelumbo Genome Database, an Integrative Resource for Gene Expression and Variants of Nelumbo Nucifera,” Scientific Data, vol. 8, no. 1, p. 38, Jan. 2021. doi: 10.1038/s41597-021-00828-8.
    Sacred lotus (Nelumbo nucifera, or lotus) is one of the most widely grown aquatic plant species with important uses, such as in water gardening and in vegetable and herbal medicine. A public genomic database of lotus would facilitate studies of lotus and other aquatic plant species. Here, we constructed an integrative database: the Nelumbo Genome Database (NGD, http://nelumbo.biocloud.net). This database is a collection of the most updated lotus genome assembly and contains information on both gene expression in different tissues and coexpression networks. In the NGD, we also integrated genetic variants and key traits from our 62 newly sequenced lotus cultivars and 26 previously reported cultivars, which are valuable for lotus germplasm studies. As applications including BLAST, BLAT, Primer, Annotation Search, Variant and Trait Search are deployed, users can perform sequence analyses and gene searches via the NGD. Overall, the valuable genomic resources provided in the NGD will facilitate future studies on population genetics and molecular breeding of lotus.
  100. Y. Li, T. Smith, P. Svetlana, J. Yang, J.-H. Jin, and C.-S. Li, “Paleobiogeography of the Lotus Plant (Nelumbonaceae: Nelumbo) and Its Bearing on the Paleoclimatic Changes,” Palaeogeography, Palaeoclimatology, Palaeoecology, vol. 399, pp. 284–293, Apr. 2014. doi: 10.1016/j.palaeo.2014.01.022.
    The historical reconstruction of the origin and dispersal of plant taxa in space and time facilitates a better understanding of their modern distribution patterns. However, most studies of paleobiogeography have focused on terrestrial plants, and the distribution changes of aquatic plants are less well understood. Here we study the lotus plant Nelumbo (Nelumbonaceae), an aquatic perennial herb, with a disjunctive distribution across East, South and Southeast Asia-North Australia and North America. The reproductive organs of Nelumbo changchangensis He et Jin from the Eocene of Hainan, China are supplementarily described. Analysis of the spatial and temporal distributions of Nelumbo in the geologic past indicates that the genus first occurs in mid-latitude area of Laurasia in the Early Cretaceous, then becomes widespread in North America and Eurasia and expands into South America during the Late Cretaceous, and reaches its maximum northern limit during the Eocene. The genus persists and thrives in North America and Eurasia until the Pliocene. The Pleistocene ice age causes the extinction of Nelumbo in Europe and central Asia, and its populations in North American and Asia are also restricted to refuges of lower latitude. Like the terrestrial plants Metasequoia (Cupressaceae) and Nordenskioeldia (Trochodendraceae), the fluctuations of Nelumbo distribution ranges are also linked to climatic changes in the Cenozoic. The cooling climate and increasing seasonality in the Eocene of East Asia may favor the origin of tubers and the differentiating of the ecotypes in lotus, which allow the deciduous type to survive in cold winters.
  101. Y. Li, Y.-shun Zhang, S. Wei, L. Liu, and Y. Chen, “Recovery of Antioxidant Gene Expression in Sacred Lotus (Nelumbo Nucifera Gaertn.) Embryonic Axes Enhances Tolerance to Extreme High Temperature,” 2012. doi: 10.5897/AJB12.678.
    It was shown that viability of seeds did not severely decline after exposure to 90°C for 24 h and the recovery of antioxidant gene expression enhanced tolerance to extreme high temperature stress in sacred lotus. Sacred lotus ( Nelumbo nucifera Gaertn.) seed is long-living and have various stress-resistance characteristics. We investigated the protecting mechanisms of lotus seeds against extreme high temperature by comparison of expression patterns of antioxidant genes in embryonic axes between exposure and non-exposure to extreme high temperature. It was shown that viability of seeds did not severely decline after exposure to 90°C for 24 h. Germination and growth were inhibited and H 2 O 2 was accumulated at high level in the lotus embryonic axes germinated after heat treatment. Transcriptional levels of superoxide dismutase (SOD), ascorbate peroxidase (APX), peroxidase (POD), glutathione peroxidase (GPX) and thioredoxin-dependent peroxidase (TPX) encoding genes were induced to rise at late germination stage. Transcriptional levels of APX, POD, GPX and alternative oxidase (AOX) encoding genes were also immediately stimulated and up-regulated after heat treatment. These results suggest that the embryonic axes of sacred lotus maintain a protective and recovery mechanism from heat damage during and after exposure to extreme high temperature. Furthermore, the recovery of antioxidant gene expression enhanced tolerance to extreme high-temperature stress in sacred lotus. Keywords: Antioxidant gene, high temperature, seed germination, Nelumbo nucifera Gaertn
  102. Y. Li, P. Svetlana, J. Yao, and C. Li, “A Review on the Taxonomic, Evolutionary and Phytogeographic Studies of the Lotus Plant (Nelumbonaceae: Nelumbo),” Acta Geologica Sinica - English Edition, vol. 88, no. 4, pp. 1252–1261, 2014. doi: 10.1111/1755-6724.12287.
    Nelumbo Adans. (Nelumbonaceae) is an important member of the early-diverging eudicots. It contains two extant species: N. nucifera Gaertn. (the Sacred lotus), distributed in Asia and Australia and N. lutea Willd. (the American lotus), occurring in North America. This paper reviews the taxonomic, evolutionary and phytogeographic studies of the genus Nelumbo, and also raises scientific questions about it in further paleobotanic research. There are about 30 fossil species of Nelumbo established since the Early Cretaceous. Based on fossil studies, the ancestors of the extant N. nucifera and N. lutea are respectively considered to be N. protospeciosa from the Eocene to Miocene of Eurasia, and N. protolutea from the Eocene of North American. However, molecular systematic studies indicate that N. nucifera and N. lutea are probably split from a common ancestor during the Late Miocene to Early Pliocene, or even the Pleistocene, rather than separate relicts from extinct species on different continents. The characters of lotus stomatal development, seedling morphology as well as its flowering, pollination and fertilization in air reveal that it evolves from the land plants. Fossil data of Nelumbo indicates that the genus first occurs in mid-latitude area of Laurasia in the Early Cretaceous, then becomes widespread in North America and Eurasia and expands into Africa and South America during the Late Cretaceous; the genus probably colonizes the Indian Subcontinent from Asia during the Early Eocene after the collision of India and the Asian plates; the genus becomes extinct in Europe, but survives in Asia and North America during the Quaternary Ice Age, and later forms the present East Asia and North Australia-North America disjunctive distribution.
  103. J. Li, T. Shi, L. Huang, D. He, T. M. Nyong’A, and P. Yang, “Systematic Transcriptomic Analysis Provides Insights into Lotus (Nelumbo Nucifera) Seed Development,” Plant Growth Regulation, vol. 86, no. 3, pp. 339–350, Dec. 2018. doi: 10.1007/s10725-018-0433-1.
    Seed size is one of the most important determinants of yield in lotus (Nelumbo nucifera). The cotyledon, which is responsible for nutrient storage in the mature seed, is the major factor affecting seed size in this economically important crop. Here, transcriptome analysis on cotyledons were performed during the rapid expansion stage of two lotus cultivars with different seed size and yield, China Antique and Jianxuan-17, at 9, 12, and 15 days after pollination (DAP). We identified 22,549 genes, including 2414 novel genes. Among them, 8437 genes were differentially expressed between CA and JX from 9 to 15 DAP. Gene ontology analysis suggested that these DEGs were significantly enriched in cell proliferation and gene expression. Dozens of DEGs are involved in brassinosteroids (BRs) biosynthesis and signaling pathway. Nine genes controlling seed size by cell number and size were candidate genes regulating lotus seed size. There was a notable difference in the expression patterns and level of starch-synthesis genes between two cultivars. NNU_20629 and NNU_05331 were likely responsible for the difference in starch accumulation between CA and JX, which might lead to their different yield. Pairwise comparisons of our transcriptome data provide insights into lotus seed development, which could facilitate projects aimed at breeding lotus with improved traits.
  104. C. libao, Z. minrong, H. Zhubing, L. huiying, and L. Shuyan, “Comparative Transcriptome Analysis Revealed the Cooperative Regulation of Sucrose and IAA on Adventitious Root Formation in Lotus (Nelumbo Nucifera Gaertn),” BMC Genomics, vol. 21, no. 1, p. 653, Sep. 2020. doi: 10.1186/s12864-020-07046-3.
    In China, lotus is an important cultivated crop with multiple applications in ornaments, food, and environmental purification. Adventitious roots (ARs), a secondary root is necessary for the uptake of nutrition and water as the lotus principle root is underdeveloped. Therefore, AR formation in seedlings is very important for lotus breeding due to its effect on plant early growth. As lotus ARs formation was significantly affected by sucrose treatment, we analyzed the expression of genes and miRNAs upon treatment with differential concentrations of sucrose, and a crosstalk between sucrose and IAA was also identified.
  105. C. Libao et al., “Gene Expression Profiling Reveals the Effects of Light on Adventitious Root Formation in Lotus Seedlings (Nelumbo Nucifera Gaertn.),” BMC Genomics, vol. 21, no. 1, p. 707, Oct. 2020. doi: 10.1186/s12864-020-07098-5.
    Lotus is an aquatic horticultural crop that is widely cultivated in most regions of China and is used as an important off-season vegetable. The principal root of lotus is degenerated, and adventitious roots (ARs) are irreplaceable for plant growth. We found that no ARs formed under darkness and that exposure to high-intensity light significantly promoted the development of root primordia. Four differential expression libraries based on three light intensities were constructed to monitor metabolic changes, especially in indole-3-acetic acid (IAA) and sugar metabolism.
  106. C. Libao, H. yuyan, L. Huiying, J. Runzhi, and L. Shuyan, “Transcriptomic Analysis Reveals Ethylene’s Regulation Involved in Adventitious Roots Formation in Lotus (Nelumbo Nucifera Gaertn.),” Acta Physiologiae Plantarum, vol. 41, no. 6, p. 97, May 2019. doi: 10.1007/s11738-019-2895-9.
    Adventitious roots (ARs) play an irreplaceable role in the uptake of water and nutrients due to under-developed principle root in plants. The process of ARs formation is affected by plant hormone. In this study, by employing High-Throughout Tag-sequencing Technique and ELISA method, we analyzed of the transcriptome and indole-3-acetic acid (IAA) content to monitor the changes of metabolism regulated by ethylene signaling in lotus. Exogenous application of ethephon (ethylene precursor) dramatically accelerated ARs development, and while restrained by 1-methylcyclopropene (1-MCP, the ethylene perception inhibitor), indicating the crucial role ethylene in ARs development. Transcriptomic analysis showed that both treatment of ethephon and 1-MCP dramatically altered the expression of numerous genes. In total, transcriptional expressions of 694 genes were induced and 554 genes were suppressed in ETH/CK0 stages compared with MCP/CK0 stages. Most of these up-regulated genes exhibited the one-three folds changes. In ETH/MCP libraries, we found nine and five genes involved in the metabolism or transcriptional responses to ethylene and IAA, and fourteen genes, which were considered to NAC, bHLH, AP2-EREBP, MYB, LOB, bHLH and bZIP families, respectively, exhibited an increase in transcriptional level. In addition, an enhanced mRNA levels of seven genes [1-aminocyclopropane-1-carboxylate oxidase (ACO), leucine-rich repeat receptor, pectinesterase, pyruvate decarboxylase, ethylene oxide synthase, respiratory burst oxidase homolog protein and xyloglucan endotransglucosylase] relevant to ARs formation were detected in was detected in ETH/MCP libraries. Furthermore, we found that IAA content was obviously decreased after applications were detected on ethephon and 1-MCP. However, the decreased IAA level in 1-MCP treatment was more pronounced than that in ethephon treatment, and kept a low level during the whole periods of ARs development. Taken together, our findings provided a comprehensive understanding of ethylene’s regulation during ARs formation in lotus seedlings.
  107. Z. Lin et al., “Genome-Wide DNA Methylation Profiling in the Lotus (Nelumbo Nucifera) Flower Showing Its Contribution to the Stamen Petaloid,” Plants, vol. 8, no. 5, p. 135, May 2019. doi: 10.3390/plants8050135.
    DNA methylation is a vital epigenetic modification. Methylation has a significant effect on the gene expression influencing the regulation of different physiological processes. Current studies on DNA methylation have been conducted on model plants. Lotus (Nelumbo nucifera) is a basic eudicot exhibiting variations during development, especially in flower formation. DNA methylation profiling was conducted on different flower tissues of lotuses through whole genome bisulfite sequencing (WGBS) to investigate the effects of DNA methylation on its stamen petaloid. A map of methylated cytosines at the single base pair resolution for the lotus was constructed. When the stamen was compared with the stamen petaloid, the DNA methylation exhibited a global decrease. Genome-wide relationship analysis between DNA methylation and gene expression identified 31 different methylation region (DMR)-associated genes, which might play crucial roles in floral organ formation, especially in the stamen petaloid. One out of 31 DMR-associated genes, NNU_05638 was homolog with Plant U-box 33 (PUB33). The DNA methylation status of NNU_05638 promoter was distinct in three floral organs, which was confirmed by traditional bisulfite sequencing. These results provide further insights about the regulation of stamen petaloids at the epigenetic level in lotus.
  108. Z. Lin, D. Cao, R. N. Damaris, and P. Yang, “Genome-Wide Identification of MADS-Box Gene Family in Sacred Lotus (Nelumbo Nucifera) Identifies a SEPALLATA Homolog Gene Involved in Floral Development,” BMC Plant Biology, vol. 20, no. 1, p. 497, Oct. 2020. doi: 10.1186/s12870-020-02712-w.
    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.
  109. Z. Lin, C. Zhang, D. Cao, R. N. Damaris, and P. Yang, “The Latest Studies on Lotus (Nelumbo Nucifera)-an Emerging Horticultural Model Plant,” International Journal of Molecular Sciences, vol. 20, no. 15, p. 3680, Jul. 2019. doi: 10.3390/ijms20153680.
    Lotus (Nelumbo nucifera) is a perennial aquatic basal eudicot belonging to a small family Nelumbonaceace, which contains only one genus with two species. It is an important horticultural plant, with its uses ranging from ornamental, nutritional to medicinal values, and has been widely used, especially in Southeast Asia. Recently, the lotus obtained a lot of attention from the scientific community. An increasing number of research papers focusing on it have been published, which have shed light on the mysteries of this species. Here, we comprehensively reviewed the latest advancement of studies on the lotus, including phylogeny, genomics and the molecular mechanisms underlying its unique properties, its economic important traits, and so on. Meanwhile, current limitations in the research of the lotus were addressed, and the potential prospective were proposed as well. We believe that the lotus will be an important model plant in horticulture with the generation of germplasm suitable for laboratory operation and the establishment of a regeneration and transformation system.
  110. Z. Lin, R. N. Damaris, T. Shi, J. Li, and P. Yang, “Transcriptomic Analysis Identifies the Key Genes Involved in Stamen Petaloid in Lotus (Nelumbo Nucifera),” BMC Genomics, vol. 19, no. 1, p. 554, Jul. 2018. doi: 10.1186/s12864-018-4950-0.
    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.
  111. D. K. Liscombe, B. P. MacLeod, N. Loukanina, O. I. Nandi, and P. J. Facchini, “Evidence for the Monophyletic Evolution of Benzylisoquinoline Alkaloid Biosynthesis in Angiosperms,” Phytochemistry, vol. 66, no. 11, pp. 1374–1393, Jun. 2005. doi: 10.1016/j.phytochem.2005.04.029.
    Benzylisoquinoline alkaloids (BIAs) consist of more than 2500 diverse structures largely restricted to the order Ranunculales and the eumagnoliids. However, BIAs also occur in the Rutaceae, Lauraceae, Cornaceae and Nelumbonaceae, and sporadically throughout the order Piperales. Several of these alkaloids function in the defense of plants against herbivores and pathogens – thus, the capacity for BIA biosynthesis is expected to play an important role in the reproductive fitness of certain plants. Biochemical and molecular phylogenetic approaches were used to investigate the evolution of BIA biosynthesis in basal angiosperms. The occurrence of (S)-norcoclaurine synthase (NCS; EC 4.2.1.78) activity in 90 diverse plant species was compared to the distribution of BIAs superimposed onto a molecular phylogeny. These results support the monophyletic origin of BIA biosynthesis prior to the emergence of the eudicots. Phylogenetic analyses of NCS, berberine bridge enzyme and several O-methyltransferases suggest a latent molecular fingerprint for BIA biosynthesis in angiosperms not known to accumulate such alkaloids. The limited occurrence of BIAs outside the Ranunculales and eumagnoliids suggests the requirement for a highly specialized, yet evolutionarily unstable cellular platform to accommodate or reactivate the pathway in divergent taxa. The molecular cloning and functional characterization of NCS from opium poppy (Papaver somniferum L.) is also reported. Pathogenesis-related (PR)10 and Bet v 1 major allergen proteins share homology with NCS, but recombinant polypeptides were devoid of NCS activity.
  112. F. Liu et al., “Characterization of Genomic Variation from Lotus (Nelumbo Adans.) Mutants with Wide and Narrow Tepals,” Horticulturae, vol. 7, no. 12, p. 593, Dec. 2021. doi: 10.3390/horticulturae7120593.
    Compared with rose, chrysanthemum, and water lily, the absence of short-wide and long-narrow tepals of ornamental lotus (Nelumbo Adans.) limits the commercial value of flowers. In this study, the genomes of two groups of lotus mutants with wide-short and narrow-long tepals were resequenced to uncover the genomic variation and candidate genes associated with tepal shape. In group NL (short for N. lutea, containing two mutants and one control of N. lutea), 716,656 single nucleotide polymorphisms (SNPs) and 221,688 insertion-deletion mutations (Indels) were obtained, while 639,953 SNPs and 134,6118 Indels were obtained in group WSH (short for ‘Weishan Hong’, containing one mutant and two controls of N. nucifera ‘Weishan Hong’). Only a small proportion of these SNPs and Indels was mapped to exonic regions of genome: 1.92% and 0.47%, respectively, in the NL group, and 1.66% and 0.48%, respectively, in the WSH group. Gene Ontology (GO) analysis showed that out of 4890 (NL group) and 1272 (WSH group) annotated variant genes, 125 and 62 genes were enriched (Q < 0.05), respectively. Additionally, in the Kyoto Encyclopedia of Genes and Genomes (KEGG) database, 104 genes (NL group) and 35 genes (WSH group) were selected (p < 0.05). Finally, there were 306 candidate genes that were sieved to determine the development of tepal shape in lotus plants. It will be an essential reference for future identification of tepal-shaped control genes in lotus plants. This is the first comprehensive report of genomic variation controlling tepal shape in lotus, and the mutants in this study are promising materials for breeding novel lotus cultivars with special tepals.
  113. R. Liu et al., “Comparative Physiological Analysis of Lotus (Nelumbo Nucifera) Cultivars in Response to Salt Stress and Cloning of NnCIPK Genes,” Scientia Horticulturae, vol. 173, pp. 29–36, Jun. 2014. doi: 10.1016/j.scienta.2014.04.032.
    Saline or salt water in the ocean accounts for 96.5% of total fresh water resource in the planet. Salinity is a global social and economic problem which severely inhibited plant growth and development. Utilization of marginal salt affected land and/or water resource becomes increasingly important because of the explosion of world population and climate change. In this study, salt stress resistance of fifteen N. nucifera cultivars was firstly evaluated. The results showed that Welcoming Guests was the most resistant cultivar, while Hunan Lotus was the most sensitive one. Resistant cultivar Welcoming Guests accumulated significant higher amount MDA and proline than Hunan Lotus prior to salt stress treatment, indicating Welcoming Guests was pre-conditioned to salt stress. Salt sensitive lotus cultivar exhibited relative lower antioxidant enzyme activities and higher reactive oxygen species accumulation than resistant one after salt treatment. Since calcineurin B-like protein interaction protein kinase (CIPK)/SALT OVERLY SENSITIVE2 gene family played essential roles during plant salt stress response, three NnCIPK genes were successfully cloned in this study. Phylogenetic analysis showed that these genes were high homologous to Arabidopsis and grape CIPK genes. Expression level analysis indicated that NnCIPK6 was highly induced by NaCl treatment in resistant cultivar, while expression levels of NnCIPK14s showed fluctuation in susceptible cultivar after salt treatment. These results partially characterized mechanisms of lotus salt stress resistance and provided useful information for utilization of lotus cultivars in salt water.
  114. Z. Liu et al., “Construction of a High-Density, High-Quality Genetic Map of Cultivated Lotus (Nelumbo Nucifera) Using next-Generation Sequencing,” BMC Genomics, vol. 17, no. 1, p. 466, Jun. 2016. doi: 10.1186/s12864-016-2781-4.
    The sacred lotus (Nelumbo nucifera) is widely cultivated in China for its edible rhizomes and seeds. Traditional plant breeding methods have been used to breed cultivars with increased yields and quality of rhizomes and seeds with limited success. Currently, the available genetic maps and molecular markers in lotus are too limited to be useful for molecular genetics based breeding programs. However, the development of next-generation sequencing (NGS) technologies has enabled large-scale identification of single-nucleotide polymorphisms (SNPs) for genetic map construction. In this study, we constructed an SNP-based high-density genetic map for cultivated lotus using double digest restriction site-associated DNA sequencing (ddRADseq).
  115. A. Liu, D. Tian, Y. Xiang, and H. Mo, “Effects of Biochar on Growth of Asian Lotus (Nelumbo Nucifera Gaertn.) and Cadmium Uptake in Artificially Cadmium-Polluted Water,” Scientia Horticulturae, vol. 198, pp. 311–317, Jan. 2016. doi: 10.1016/j.scienta.2015.11.030.
    To further understand the benefits of biochar in crop production and remediation of heavy metal pollution, the effects of biochar on the growth of Nelumbo nucifera ‘Taikong 36’ and cadmium (Cd) accumulation in plant tissues were evaluated in the artificially Cd-polluted water-soil. The results showed that plant biomass significantly increased with the proportion (0–32%) of biochar in the soil mix. However, the impacts of biochar on plant physiological indicators were not clear. Compared with the control, the addition of 32% biochar significantly decreased Cd content in rhizomes, petioles, and leaves of N. nucifera ‘Taikong 36’ by 69, 81, 55%, respectively. Correspondingly, the bioaccumulation coefficient of Cd was reduced, by a maximum of 71%, and the Cd transfer coefficient from underground to aboveground tissues increased up to 1.3 fold. The bioaccumulation coefficient of Cd in rhizomes was significantly lower than that of leaves, indicating that biochar mitigated the effect of heavy metal ions and reduced the accumulation of Cd in the edible parts of lotus. Therefore, biochar has potential in green agriculture production and remediation of heavy-metal polluted water and soil due to its positive effects: improving plant growth and reducing heavy metal pollution.
  116. F. Liu et al., “Effects of Parent Species Type, Flower Color, and Stamen Petaloidy on the Fruit-Setting Rate of Hybridization and Selfing in Lotus (Nelumbo),” Aquatic Botany, vol. 172, p. 103396, Jun. 2021. doi: 10.1016/j.aquabot.2021.103396.
    Understanding the relationship of fruit-setting rate (FSR) with species type, flower color, and stamen petaloidy (tepal number) of the parents in the crossing and selfing of lotus (Nelumbo) can improve breeding efficiency. In this study, 193 artificial crosses with 1.932 flowers and selfing (artificial and non-interference selfing) with 1.120 flowers were carried out in 218 lotus accessions. There were no differences in the cross FSR between Asian lotus (Nelumbo nucifera) and American lotus (N. lutea), but both of their cross FSRs were significantly higher than that (7.9 %) of Asian-American hybrids. The selfing FSRs were highest in American lotus, followed by Asian lotus and Asian-American hybrids. Regardless of whether they were used as the female or male parents in crosses, lotus accessions with pink tepals had higher FSRs than those with white tepals. While the yellow-flowered American lotus obtained the highest FSRs among all flower colors in both of selfing. As the tepal number increased, the FSRs of cross and artificial selfing did not continuously decrease. Cultivars with 91–120 tepals had the highest cross FSR in the double-tepalled lotus, regardless of whether they were used as the male or female parents. Therefore, the cross and selfing FSRs of lotus were associated with parent species type, flower color, and stamen petaloidy. Generally, the results of this study provide useful insights into parent selection, cross design, and sample sizes for reducing costs and improving efficiency in lotus breeding.
  117. J. Liu, B. Dong, Y. Cui, W. Zhou, and F. Liu, “An Exploration of Plant Characteristics for Plant Species Selection in Wetlands,” Ecological Engineering, vol. 143, p. 105674, Jan. 2020. doi: 10.1016/j.ecoleng.2019.105674.
    Plants are a major component of constructed wetlands (CWs) but arriving at a reasonable choice of plants is a challenge. In this study, surface flow CWs planted with eight different species of plants were investigated to explore the relationship between plant characteristics, including density, height, leaf area index, evapotranspiration (ET) and plant coefficient (the ratio of the actual plant ET to a reference ET), and the treatment performance of these plants in purifying water. Of the five plant characteristics, only the plant coefficient was found to be negatively correlated with kv (first-order removal rate) for nitrogen and phosphorus. The interpretation of this relationship requires caution, as the negative relationship may be caused by the water consumed by ET in the long term. The relationship between ETc (actual ET of a plant), plant coefficient and the removal load explain the two contradictory effects of ET on the removal load. Considering treatment performance and plant coefficient, the plant most recommended for use in CWs is Iris sibirica and that which is the least recommended is Nelumbo nucifera Gaertn. The combination of treatment performance and plant coefficient provides a robust approach to the selection of plants for areas with a high ET.
  118. Y. Liu, S.-shuang Ma, S. A. Ibrahim, E.-hu Li, H. Yang, and W. Huang, “Identification and Antioxidant Properties of Polyphenols in Lotus Seed Epicarp at Different Ripening Stages,” Food Chemistry, vol. 185, pp. 159–164, Oct. 2015. doi: 10.1016/j.foodchem.2015.03.117.
    In this study, polyphenols from lotus seed epicarp (PLSE) at three different ripening stages were purified by column chromatography and identified by RP-HPLC and HPLC–ESI-MS2. The antioxidant activities of PLSE were also investigated. We found that the contents of PLSE at the green ripening stage, half ripening stage and full ripening stage are 13.08%, 10.95% and 6.73% respectively. The levels of catechin, epicatechin, hyperoside, and isoquercitrin in PLSE at the three different ripening stages were different. Moreover, the amounts of catechin and epicatechin decreased, while the contents of hyperoside and isoquercitrin increased as the seed ripened. We found that PLSE at three different ripening stages had good scavenging abilities on DPPH and ABTS+ radicals. However, the scavenging ability decreased with maturation. Our results may be valuable with regard to the utilization of lotus seed epicarp as a functional food material.
  119. Y. Liu, P. Chaturvedi, J. Fu, Q. Cai, W. Weckwerth, and P. Yang, “Induction and Quantitative Proteomic Analysis of Cell Dedifferentiation during Callus Formation of Lotus (Nelumbo Nucifera Gaertn.Spp. Baijianlian),” Journal of Proteomics, vol. 131, pp. 61–70, Jan. 2016. doi: 10.1016/j.jprot.2015.10.010.
    Lotus is an aquatic plant with high nutritional, ornamental and medical values. Its callus formation is crucial for germplasm innovation by genetic transformation. In this study, embryogenic callus was successfully induced on appropriate medium using cotyledons at 12days after pollination as explants. To dissect cellular dedifferentiation and callus formation processes at the proteome level, cotyledons before and tissues from 10 to 20days after induction were sampled for shotgun proteomic analysis. By applying multivariate statistics 91 proteins were detected as differentially regulated, and sorted into 6 functional groups according to MapMan ontology analysis. Most of these proteins were implicated in various metabolisms, demonstrating that plant cells underwent metabolism reprogramming during callus induction. 14.3% proteins were associated with stress and redox, indicating that the detached explants were subjected to a variety of stresses; 13.2% were cell and cell wall-related proteins, suggesting that these proteins played important roles in rapid cell division and proliferation. Some proteins were further evaluated at the mRNA levels by quantitative reverse transcription PCR analysis. In conclusion, the results contributed to further deciphering of molecular processes of cellular dedifferentiation and callus formation, and provided a reference data set for the establishment of transgenic transformation in lotus.
  120. Q. Liu, D. Zhang, F. Liu, M. Qin, and D. Tian, “Micropropagation of Nelumbo Nucifera ‘Weishan Hong’ through Germfree Mature Embryos,” In Vitro Cellular & Developmental Biology - Plant, vol. 55, no. 3, pp. 305–312, Jun. 2019. doi: 10.1007/s11627-019-09984-4.
    Asian lotus (Nelumbo nucifera Gaertn.) is an economically important aquatic plant that is widely cultivated in Asia. It is mainly propagated by dividing rhizomes, which maintains genotype stability but is also laborious and time-consuming. An efficient in vitro method that ensures genetic stability is highly desirable. An optimized method was developed to produce intact and sterile mature embryos by soaking fruits in sterile water containing 2% (v/v) plant preservative mixture. Using mature embryos as explants, an improved micropropagation procedure was established in N. nucifera ‘Weishan Hong.’ The growth patterns of explants differed with the types and concentrations of cytokinins. Media containing 0.5 to 2.0 mg L−1 kinetin were suitable for elongation of unexpanded stems, while media with 1.0 to 2.0 mg L−1 6-benzylaminopurine (BAP) the stems were shorter. Higher concentrations of about 3.0 mg L−1 BAP promoted shoot cluster formation. Liquid–solid, double-layer media promoted plantlet growth. The well-rooted plantlets could normally grow after being transplanted. Lotus is known to be recalcitrant to regeneration from callus. Apices of mature embryos, aseptic plantlets, and young embryos formed callus successfully when the medium was supplemented with 2,4-dichlorophenoxyacetic acid and thidiazuron. Callus morphology differed with respect to the source. However, only a few of the calluses developed axillary shoots.
  121. Z. Liu et al., “Molecular Cloning and Expression Profile of Class E Genes Related to Sepal Development in Nelumbo Nucifera,” Plants, vol. 10, no. 8, p. 1629, Aug. 2021. doi: 10.3390/plants10081629.
    The lotus (Nelumbo Adans.) is an important aquatic plant with ornamental, medicinal and edible values and cultural connotations. It has single-, semi-double-, double- and thousand-petalled types of flower shape and is an ideal material for developmental research of flower doubling. The lotus is a basal eudicot species without a morphological difference between the sepals and petals and occupies a critical phylogenetic position in flowering plants. In order to investigate the genetic relationship between the sepals and petals in the lotus, the class E genes which affect sepal formation were focused on and analyzed. Here, SEPALLATA 1(NnSEP1) and its homologous genes AGAMOUS-LIKE MADS-BOXAGL9 (NnAGL9) and MADS-BOX TRANSCRIPTION FACTOR 6-like (NnMADS6-like) of the class E gene family were isolated from the flower buds of the Asian lotus (Nelumbo nucifera Gaertn.). The protein structure, subcellular localization and expression patterns of these three genes were investigated. All three genes were verified to locate in the nucleus and had typical MADS-box characteristics. NnSEP1 and NnMADS6-like were specifically expressed in the sepals, while NnAGL9 was highly expressed in the petals, suggesting that different developmental mechanisms exist in the formation of the sepals and petals in the lotus. The significant functional differences between NnSEP1, NnMADS6-like and NnAGL9 were also confirmed by a yeast two-hybrid assay. These results expand our knowledge on the class E gene family in sepal formation and will benefit fundamental research on the development of floral organs in Nelumbo.
  122. Y. Luo et al., “Bioaccumulation of Heavy Metals in the Lotus Root of Rural Ponds in the Middle Reaches of the Yangtze River,” Journal of Soils and Sediments, vol. 17, no. 10, pp. 2557–2565, Oct. 2017. doi: 10.1007/s11368-017-1692-6.
    The subject of this study is the sediment and wild lotus plants in unmanaged ponds, near Yichang City, contaminated by heavy metals. The objective is to determine the extent and frequency of heavy metal accumulation by lotus root in the ponds of rural areas and its significance to food safety and human health.
  123. S. Luo, H. Hu, L. Zhang, H. Zhou, and P. Li, “Sugars in Postharvest Lotus Seeds Were Modified by 6-Benzylaminopurine Treatment through Altering Related Enzymes Involved in Starch-Sucrose Metabolism,” Scientia Horticulturae, vol. 221, pp. 73–82, Jul. 2017. doi: 10.1016/j.scienta.2017.03.044.
    To investigate the effect of 6-benzylaminopurine (6-BA) treatment on starch-sucrose metabolism in postharvest lotus seeds, the lotus pods were immersed with 20mgL−1 6-BA and stored at 25±1°C for 8days. Our results revealed that 6-BA treatment not only delayed the browning of lotus pods and seeds, but also inhibited the respiration rate of lotus pods. Compared with controls, 6-BA treatment enhanced the deposition of starch, which was supported by the decreased amylase level. Contrary to the alteration of starch content, glucose and fructose contents in lotus seeds were reduced by 6-BA treatment through inhibiting the production of hexose from sucrose and starch degradations. However, the sucrose content in 6-BA treated group was higher than that in control groups until 6th day of storage, which was resulted from enhanced sucrose phosphate synthase (SPS) level and inhibited activities of synthase-cleavage (SS-cleavage) and invertase by 6-BA treatment. These results indicated that 6-BA treatment could maintain the starch and sucrose contents of lotus seeds by affecting related-synthetic or degrading enzymes involved in starch-sucrose metabolism, and alleviate postharvest senescence of lotus pods and seeds.
  124. C. Ma et al., “Purification and Characterization of Aporphine Alkaloids from Leaves of Nelumbo Nucifera Gaertn and Their Effects on Glucose Consumption in 3T3-L1 Adipocytes,” International Journal of Molecular Sciences, vol. 15, no. 3, pp. 3481–3494, Mar. 2014. doi: 10.3390/ijms15033481.
    Aporphine alkaloids from the leaves of Nelumbo nucifera Gaertn are substances of great interest because of their important pharmacological activities, particularly anti-diabetic, anti-obesity, anti-hyperlipidemic, anti-oxidant, and anti-HIV’s activities. In order to produce large amounts of pure alkaloid for research purposes, a novel method using high-speed counter-current chromatography (HSCCC) was developed. Without any initial cleanup steps, four main aporphine alkaloids, including 2-hydroxy-1-methoxyaporphine, pronuciferine, nuciferine and roemerine were successfully purified from the crude extract by HSCCC in one step. The separation was performed with a simple two-phase solvent system composed of n-hexane-ethyl acetate-methanol-acetonitrile-water (5:3:3:2.5:5, v/v/v/v/v). In each operation, 100 mg crude extracts was separated and yielded 6.3 mg of 2-hydroxy-1-methoxyaporphine (95.1% purity), 1.1 mg of pronuciferine (96.8% purity), 8.5 mg of nuciferine (98.9% purity), and 2.7 mg of roemerine (97.4%) respectively. The chemical structure of four aporphine alkaloids are identified by means of electrospray ionization MS (ESI-MS) and nuclear magnetic resonance (NMR) analysis. Moreover, the effects of four separated aporphine alkaloids on insulin-stimulated glucose consumption were examined in 3T3-L1 adipocytes. The results showed that 2-hydroxy-1-methoxyaporphine and pronuciferine increased the glucose consumption significantly as rosiglitazone did.
  125. Y. Maeda, T. Minamikawa, and B.-M. Xu, “Metabolic Activities in Germinated Ancient Lotus Seeds,” Journal of Experimental Botany, vol. 47, no. 297, pp. 577–582, 1996. https://www.jstor.org/stable/23695348.
    Seeds of Taizi lotus (Nelumbo nucifera Gaertn cv. Taizi) were unearthed in a western suburb of Beijing in 1984, and determined to be 580 ± 70-years-old by radiocarbon dating. Pretreatment with concentrated sulphuric acid for 6 h promoted 75% of the seeds tested to germinate. Over 3 weeks of incubation after the addition of water, storage starch, total soluble sugar and globulin in the cotyledons decreased whereas albumin and soluble α-amino nitrogen increased. These changes in cotyledonary components were similar to the changes seen in four present-day Indian lotus varieties, although the content of reducing sugar in Taizi lotus was significantly lower. The polypeptide compositions in extracts from the cotyledons and embryonic axis of Taizi seeds, as analysed by SDS—PAGE, were similar to those of present-day Jiangxi seeds. Taizi lotus also showed the ability to incorporate radioactive leucine into the trichloroacetic acid (TCA)-insoluble fraction of a cotyledonary extract during the 24 h post-imbibition period. The incorporation was inhibited when seeds of Taizi lotus were allowed to rehydrate for 24 h in the presence of α-amanitin or cycloheximide.
  126. N. Mahmad, R. Mat Taha, R. Othman, A. Saleh, N. A. Hasbullah, and H. Elias, “Effects of NAA and BAP, Double-Layered Media, and Light Distance on In Vitro Regeneration of Nelumbo Nucifera Gaertn. (Lotus), an Aquatic Edible Plant,” The Scientific World Journal, vol. 2014, pp. 1–8, 2014. doi: 10.1155/2014/745148.
    In vitro direct regeneration of Nelumbo nucifera Gaertn. was successfully achieved from immature explants (yellow plumule) cultured on a solid MS media supplemented with combinations of 0.5 mg/L BAP and 1.5 mg/L NAA which resulted in 16.00 ± 0.30 number of shoots per explant and exhibited a new characteristic of layered multiple shoots, while normal roots formed on the solid MS basal media. The double-layered media gave the highest number of shoots per explant with a ratio of 2\,: 1 (liquid to solid) with a mean number of 16.67 ± 0.23 shoots per explant with the formation of primary and secondary roots from immature explants. In the study involving light distance, the tallest shoot (16.67 ± 0.23 mm) obtained from the immature explants was at a light distance of 200 mm from the source of inflorescent light (1000 lux). The plantlets were successfully acclimatized in clay loam soil after 8 months being maintained under in vitro conditions.
  127. N. Mahmad, “Germination, Regeneration and Pigment Detection in Nelumbo Nucifera Gaertn. (Pink Asian Lotus) / Noraini Binti Mahmad,” Master's thesis, University of Malaya, Kuala Lumpur, 2012. http://pendeta.um.edu.my/client/default/search/results?qu=Germination%2C+regeneration+and+pigment+detection+in+nelumbo+nucifera+gaertn&te=.
    In vivo and in vitro germination of three types of seeds including matured (black), immature or fruits (green) and young (yellow) of Nelumbo nucifera Gaertn. (pink Asian lotus) were done. The black and green seeds both gave the highest responses to technique of scarified with medium-sand paper and germinated in liquid (tap water) and solid (MS basal media) substrates. The black seeds with germination rate of 67.01±0.28% in vivo and 78.35±0.61% in vitro, while, green seed with 75.12±0.16% in vivo and 100.00% in vitro germination. The highest shoot length in seeds without cotyledon for black and green seed with 28.00±0.55 mm and 26.00±0.16 mm, respectively. The highest shoot length was in media with pH 5.5 for both black seed and green seeds and with 64.03±0.02 mm and 32.55±0.04 mm, respectively. The maximum root length for both black and green seeds was 14 mm on the 14th day with 3 green shoots and 6 white primary roots. Whilst, maximum root length for yellow seeds were on the 18th day with 3 green shoots and 6 white primary roots. For storage purposes, the optimum concentration for the formation of encapsulation matrix was 3.0% sodium alginate (NaC6H7O6). Encapsulated explants were soaked in 100 mM calcium chloride dehydrate (CaCl2.2H2O) solution for 30 minutes. Through sodium alginate encapsulated method (4±1°C), seeds germination reduced from 100% (day 1) to 53.3±1.2% (day 90), while, through frozen whole seeds method (-20±1°C), 100% germination rate until 60 days in storage. After 90 days in storage, the germination rate was still high with 93.3±0.6%. Direct regeneration of Nelumbo nucifera Gaertn. were successfully achieved from green seed explants cultured on solid MS media supplemented with combinations of 1.5 mg/l BAP and 0.5 mg/l NAA with 10.33 ±0.23 shoots per explant (true-to-type), and with 3.67 ±0.31 roots per explant. Direct regeneration of Nelumbo nucifera Gaertn. were successfully achieved from yellow seed explants cultured on solid MS media supplemented with combinations of 0.5 mg/l BAP and 1.5 mg/l NAA with 16.00±0.30 shoots per explant, with new characteristics of layered multiple shoots. Roots were formed on solid MS basal media. Some formation of abnormal shoots (pinkish, red and oval leaf) occurred from green seed explants on solid MS media supplemented with combinations of 1.0 mg/l BAP and 2.5 mg/l NAA, 2.5 mg/l BAP and 2.5 mg/l NAA, 1.5 mg/l BAP and 2.0 mg/l NAA and 1.5 mg/l BAP and 2.5 mg/l NAA. Furthermore, the highest shoots per explant for green seed explants was in 8.8mg/l MS powder (double strength) with mean 19.03±0.05 shoots per explant. The highest shoots per explants for yellow seed explants was on 4.4 mg/l MS powder (full strength) with mean 16.06±0.06 shoots per explant. Media with pH 5.5 resulted in the highest height of shoots for green and yellow seed explants with mean 12.04±0.7 mm and 16.03±0.30 mm, respectively. While, the highest height of shoots for green seed explants with 9.41±1.11 mm in 250.00 mm light distance. The highest height of shoots with 16.67±0.23 mm for yellow seed explants in 200.00 mm light distance. Solid MS basal media was optimum for root formation within 4 weeks for green seed explants and after 24 weeks for yellow seed explants on solid MS media supplemented with 0.5mg/l BAP and 1.5mg/l NAA. In double layer media, the highest number of shoots per explant was both in ratio liquid to solid 2:1 with mean 16.67±0.23 number of shoots per explant with formation of primary and secondary roots for explants from yellow seeds with formation of layered multiple shoots, while mean 9.00±0.15 number of shoots per explant for green seeds. Extraction from lotus stamen was analyzed through HPLC system. Pigment detected was found to have the total carotenoid content of 526.96 ± 0.52 μg/g DW, whereas for individual carotenoid β-carotene (460 ± 10.28 μg/g DW) was found with a relatively high concentration and neoxanthin (39.26 ± 0.82 μg/g DW) was found in lower concentrations. One unknown carotenoid also was detected. For coating experiments, at room temperature with acidic conditions (pH 1-6), yellow pigment changed to the darker colour, yellow-brown, while, at strong alkaline condition (pH 12-14), the colour change to yellow-green. The optimum yellow colour was at pH 8-10. The highest glossiness was in 1.0ml/30.0ml v/v of pigment-resin solution with mean of 74.67±0.33°. Yellow pigment in 20% PMMA (resin) was as stable as pigment-resin added with 1% tartaric acid, coated on glass slides. In contrast, the addition of 1% citric acid reduced the carotenoid color. Extraction for antimicrobial activity from in vivo and in vitro rolled leaves showed no inhibition zone to all tested bacteria Staphylococcus aureus, Pseudomonas aeruginosa ,Bacillus cereus and E.coli. However, the highest inhibition zone was from in vivo sample for both on fungi, Fusarium sp. and Trichoderma sp. with mean 9.0±0.1 mm and with 7.3±02 mm diameters, respectively. Inhibition zone for in vitro samples were 8 times lower than in vivo with mean of 2.0±0.4 mm on Fusarium sp. and 1.1±0.3 mm on Trichoderma sp. Soil samples collected from Tasik Chini with mean pH of 3.97±0.02, while the optimum survival for acclimatization plantlets was on black clay loam (mean pH 6.03±0.29) that commercialized at all the nurseries in Malaysia. The most acidic soils were collected from Tasik Chini with the lowest survival rate (31.34±0.77%) for green seed and 11.11±0.51% for yellow seeds. The highest survival rate was in black clay loam for both plantlets from green and yellow seeds with 83.01±0.23% and 69.22±0.43%, respectively. The highest plantlets height was obtained from plant acclimatized under full sunlight exposure with 10 times from shady exposure. Acclimatized plantlets showed the same character as mother plants, even though at the beginning of transferred with layered multiple shoots (new character was formed in vitro only).
  128. A. Marmur, “The Lotus Effect:\,Superhydrophobicity and Metastability,” Langmuir, vol. 20, no. 9, pp. 3517–3519, Apr. 2004. doi: 10.1021/la036369u.
    To learn how to mimic the Lotus effect, superhydrophobicity of a model system that resembles the Lotus leaf is theoretically discussed. Superhydrophobicity is defined by two criteria:\,a very high water contact angle and a very low roll-off angle. Since it is very difficult to calculate the latter for rough surfaces, it is proposed here to use the criterion of a very low wet (solid−liquid) contact area as a simple, approximate substitute for the roll-off angle criterion. It is concluded that nature employs metastable states in the heterogeneous wetting regime as the key to superhydrophobicity on Lotus leaves. This strategy results in two advantages: (a) it avoids the need for high steepness protrusions that may be sensitive to breakage and (b) it lowers the sensitivity of the superhydrophobic states to the protrusion distance.
  129. J. Masuda, Y. Ozaki, I. Miyajima, and H. Okubo, “Ethylene Is Not Involved in Rhizome Transition to Storage Organ in Lotus (Nelumbo Nucifera),” Journal of the Faculty of Agriculture, Kyushu University, vol. 55, no. 2, pp. 231–232, Oct. 2010. doi: 10.5109/18834.
  130. J.-ichiro Masuda, Y. Ozaki, M. Hiramatsu, K. Sakai, J. Kim, and H. Okubo, “Plasticity of Morphology in Subterranean Stems in Long and Short Day in the Lotus (Nelumbo Nucifera),” The Horticulture Journal, vol. 87, no. 1, pp. 80–88, 2018. doi: 10.2503/hortj.OKD-082.
    Effects of photoperiod and temperature on rhizome enlargement (dormancy induction) and accompanied dormancy depth were investigated in this study. Nine-day-old seedlings were transplanted from 26 July at 1 week intervals, and they were grown under a natural photoperiod for 5 weeks in an unheated greenhouse in Fukuoka Prefecture, Japan. Although subterranean stems elongated in the plants grown until 30 August or 6 September, enlarged rhizomes were formed in those grown until 13 September or 20 September. It was revealed from these results that the lotus recognizes a natural photoperiod after 6 September as a short day. When 9 treatments of day length combinations (LD0+SD8–LD8+SD0) were applied to the seedlings, the plants grown under short day after long day treatment of 0 (LD0+SD8), 1 (LD1+SD7), 2 (LD2+SD6), 3 (LD3+SD5), 4 (LD4+SD4), 5 (LD5+SD3), 6 (LD6+SD2), or 7 (LD7+SD1) weeks formed enlarged rhizomes from the fifth, fifth, sixth, seventh, eighth, tenth, twelfth, and fourteenth internodes, respectively. Although photoperiodic treatment in the first week was different between LD0+SD8 and LD1+SD7 treatments, subterranean stems began to enlarge from the same internode (fifth internode) in both treatments. This indicates that photoperiod treatments for the first week do not affect morphology of subterranean stems. Seven treatments of day length combinations (LD2+SD0+LD6–LD2+SD6+LD0) were applied to seedlings after long day treatment for 2 weeks. Enlarged subterranean stems were observed in the plants grown under short day for 6 weeks (LD2+SD6+LD0), but not in those under long day for 6 weeks (LD2+SD0+LD6). On the other hand, subterranean stems elongated again after rhizome enlargement under a subsequent long day following 1 (LD2+SD1+LD5), 2 (LD2+SD2+LD4), 3 (LD2+SD3+LD3), or 4 (LD2+SD4+LD2) weeks of short day. This clarified that morphogenesis in subterranean stems is completely dependent on photoperiod. Further, it is expected that such growth resumption may be attributed to a weak dormant state in the enlarged rhizome. The enlarged rhizomes were exposed to natural low temperatures to examine environmental factors for deepening dormancy. Rhizomes sprouted in all treatments irrespective of exposure to low temperatures when they were transferred to ideal conditions. Rapid growth in leaves and subterranean stems was particularly observed by exposure to low temperature. It was suggested that low temperature is an environmental factor for releasing dormancy, but not for deepening dormancy. It is proposed from these results that subterranean stem growth is completely dependent on photoperiod, and that enlarged rhizomes show weak dormancy.
  131. J.-ichiro Masuda, Y. Ozaki, and H. Okubo, “Rhizome Transition to Storage Organ Is under Phytochrome Control in Lotus (Nelumbo Nucifera),” Planta, vol. 226, no. 4, pp. 909–915, Sep. 2007. doi: 10.1007/s00425-007-0536-9.
    We examined photoperiodic response of lotus (Nelumbonucifera) rhizome morphogenesis (its transition to a storage organ) by using seed-derived plants. Rhizome enlargement (increase in girth) was brought about under 8, 10 and 12 h photoperiods, whereas the rhizomes elongated under 13 and 14 h photoperiods. Rhizomes elongated under 14 h light regimes supplied as 8 h of natural light plus 6 h supplemental hours of white, yellow or red light, but similar treatments with supplemental blue, green or far red light, caused enlargement in girth of the rhizomes. A 2 h interruption of the night with white, yellow or red light, in plants entrained to 8 h photoperiod brought rhizome elongation, whereas 2 h-blue, green or far red light night breaks still resulted in rhizome increase in girth. The inhibitory effect of a red (R) light night break on rhizome increase in girth was reversed by a far-red (FR) light given immediately afterwards. Irradiation with R/FR/R inhibited the rhizome increase in girth. FR light irradiation following R/FR/R irradiation cancelled the effect of the last R light irradiation. It was demonstrated that the critical photoperiod for rhizome transition to storage organ is between 12 and 13 h photoperiod. It was also evident that the optimal light quality range for interruption of dark period (night break) is between yellow and red light and that a R/FR reversible reaction is observed. From these results, we propose that phytochrome plays an important role in photoperiodic response of rhizome increase in girth in lotus. This is the first report on phytochrome-dependent morphogenesis of storage organs in rhizomous plants.
  132. J. Masuda, S. Yoshimizu, Y. Ozaki, and H. Okubo, “Rhythmic Response of Rhizome Growth to Light-Break in Lotus (Nelumbo Nucifera),” Journal of the Faculty of Agriculture, Kyushu University, vol. 52, no. 1, pp. 35–38, Feb. 2007. doi: 10.5109/9277.
    We examined the response of rhizome growth to red light–break under different short daylengths in lotus (Nelumbo nucifera) seedlings. Maximum inhibitory response of rhizome enlargement to light–break under 10, 8 and 4 hr daylengths occurred 10, 8–10 and 12–14 hrs after the beginning of the dark period, respectively. It was found that rhythmic response to light–break is involved in rhizome growth of lotus.
  133. J.-I. Masuda, T. Urakawa, Y. Ozaki, and H. Okubo, “Short Photoperiod Induces Dormancy in Lotus (Nelumbo Nucifera),” Annals of Botany, vol. 97, no. 1, pp. 39–45, Jan. 2006. doi: 10.1093/aob/mcj008.
    BACKGROUND AND AIMS: Lotus (Nelumbo nucifera) has been cultivated as an ornamental and food plant in Japan for more than 1000 years. As large areas are required for its cultivation (approximately 2 m2 per plant), physiological research, such as into the effect of environmental factors on dormancy, has not been well studied until recently. In this paper, seedlings were used to examine environmental factors affecting dormancy induction. METHODS: In a first experiment, seeds were sown from 6 April to 6 October at 2-month intervals, and cultivated for 2 months in an unheated greenhouse. In a second experiment, seeds were prepared for germination on 16 November and 16 May and the seedlings were grown at 25 or 30 degrees C under natural daylength in phytotron growth rooms. After 1 month, the seedlings were cultivated at 20, 25 or 30 degrees C for a further month. The number of leaves and rhizome branches on the main stem were counted, and growth of rhizomes on the main stem was calculated using a rhizome enlargement index (= maximum internode diameter/internode length) after 2 months of culture in both experiments. KEY RESULTS: Rhizomes elongated without enlargement when the seeds were sown in April and June. Sowing the seeds in August and October resulted in rhizome enlargement from the tenth and fifth internodes, respectively. Rhizomes enlarged in the November-sowing but elongated in the May-sowing irrespective of temperature treatments under natural daylength in the phytotron rooms. The seedlings cultivated from May at 25-30 degrees C for 2 months had more leaves, and more rhizome branches and nodes than those cultivated from November. CONCLUSIONS: Short days led to induced dormancy in lotus.
  134. P. G. D. Matthews and R. S. Seymour, “Anatomy of the Gas Canal System of Nelumbo Nucifera,” Aquatic Botany, vol. 85, no. 2, pp. 147–154, Aug. 2006. doi: 10.1016/j.aquabot.2006.03.002.
    Nelumbo nucifera (Gaertn.) grows by extending a creeping rhizome through anaerobic sediments. Nodes form at intervals along the rhizome, each producing a single leaf, and gas canals channel air from the leaves throughout the petioles and rhizomes. The gas flow pathway was mapped by casting the canals in growing shoots with silicone and by blowing air through complexes of rhizomes and petioles. Air from a leaf flows to a rhizome through one of two petiolar canal pairs, joining with the lowermost of three canal pairs in the rhizome through a chamber in the node. The lowermost canal pair links these nodal chambers along the length of a rhizome, allowing air from a node to flow both forward, toward a growing shoot, and backward, toward preceding leaves. These linked chambers also connect with the middle pair of canals on their proximal side, enabling flow to proceed backward along the rhizome to an adjacent node. A chamber in the next node then diverts the flow into the upper canal pair. This pair leads to a third node and chamber from which the air vents to the atmosphere through the second petiolar canal pair. Thus, pressurised air from one leaf must flow backward through two nodes before it returns to the atmosphere. Forward flow also ventilates a shoot’s growing tip, with air from the lowermost canal pair entering a chamber in the developing node which, as described above, connects with the middle canal. This allows the air to reverse direction at the tip and enter the vent flow pathway.
  135. P. G. D. Matthews and R. S. Seymour, “Stomata Actively Regulate Internal Aeration of the Sacred Lotus Nelumbo Nucifera,” Plant, Cell & Environment, vol. 37, no. 2, pp. 402–413, 2014. doi: 10.1111/pce.12163.
    The sacred lotus Nelumbo nucifera (Gaertn.) possesses a complex system of gas canals that channel pressurized air from its leaves, down through its petioles and rhizomes, before venting this air back to the atmosphere through large stomata found in the centre of every lotus leaf. These central plate stomata (CPS) lie over a gas canal junction that connects with two-thirds of the gas canals within the leaf blade and with the larger of two discrete pairs of gas canals within the petiole that join with those in the rhizome. It is hypothesized that the lotus actively regulates the pressure, direction and rate of airflow within its gas canals by opening and closing these stomata. Impression casting the CPS reveal that they are open in the morning, close at midday and reopen in the afternoon. The periodic closure of the CPS during the day coincides with a temporary reversal in airflow direction within the petiolar gas canals. Experiments show that the conductance of the CPS decreases in response to increasing light level. This behaviour ventilates the rhizome and possibly directs benthic CO2 towards photosynthesis in the leaves. These results demonstrate a novel function for stomata: the active regulation of convective airflow.
  136. A. McDonald, “A Botanical Perspective on the Identity of Soma (Nelumbo Nucifera Gaertn.) Based on Scriptural and Iconographic Records,” Economic Botany, vol. 58, no. 1, pp. S147–S173, Dec. 2004. doi: 10.1663/0013-0001(2004)58[S147:ABPOTI]2.0.CO;2.
    An examination of the mythic and artistic records of India and Southeast Asia indicates that the famous psychotropic of the ancient Aryans was the eastern lotus, Nelumbo nucifera. Vedic epithets, metaphors, and myths that describe the physical and behavioral characteristics of the ‘soma’ plant as a sun, serpent, golden eagle, arrow, lightning bolt, cloud, phallic pillar, womb, chariot, and immortal navel, relate individually or as a whole to the eastern lotus. Since most Hindu and Buddhist gods and goddesses trace their origins from the Vedas and have always shared close symbolic associations with Nelumbo, there is reason to believe the divine status of this symbolic plant derives from India’s prehistoric past.
  137. C. McGrath, “Fertilizers Affect Water and Substrate EC, pH, and Nutritional Concentrations for Nelumbo Production,” Doctor of Philosophy, Auburn University, Auburn, Alabama, 2012. https://etd.auburn.edu//handle/10415/3088.
    Sacred Lotus (Nelumbo nucifera Gaertn.) is an aquatic, herbaceous perennial considered to be one of the most valuable plants in the world. Each part of lotus is consumed throughout Asia for food or used for medicinal purposes. Effects of fertilizer type (conventional, organic, or no fertilizer), fertility rate, and water depth on water and substrate electrical conductivity (EC), pH, nitrate-nitrogen, and ammonium-nitrogen concentration were evaluated in greenhouse and outdoor studies to determine effect on potential lotus growth. All fertilizers influenced water and substrate EC, pH, and nutritional concentration. According to substrate analysis, EC rates were above recommended levels. Both organic treatments resulted in high sodium levels and the organic Nature Safe treatment resulted in higher levels of most macronutrients by termination of all studies. Results indicated increased water volumes led to reduced nutrient concentration and availability. All measured parameters decreased with increased water depths due to greater water volume and dilution factors and researchers determined a water depth of 15.2 cm (6 in) resulted in satisfactory EC levels for lotus production. There would be no additional benefit in maintaining shallower or greater depths. EC is a strong factor influencing lotus growth and with shallower depths, EC could rise close to toxic levels as was revealed in the organic Medina Growin’ Green treatment. Toxic EC level of 1.0 mS•cm-1 was surpassed with increasing rates to 1.3 kg•m-3N among both conventional and organic treatments. Under greenhouse conditions with moderate temperatures, researchers determined 0.6 kg•m-3N was a potentially acceptable rate to target for the fertilizers tested for outdoor production. The rate resulted in toxic levels of soluble salts for some fertilizers and required removal and replacement of plants, substrate, and fertilizer; adjusting the rate to 0.4 kg•m-3N. A rate of 0.44 kg•m-3N resulted in acceptable EC levels for all fertilizers trialed and tested. More research needs to be conducted to determine the interactions, cause and effect of the many variables on specific fertilizer nutrient release to target a satisfactory level to maximize growth while minimizing any potential crop damage due to an increase in EC to toxic levels.
  138. T. Meelaph et al., “Coregulation of Biosynthetic Genes and Transcription Factors for Aporphine-Type Alkaloid Production in Wounded Lotus Provides Insight into the Biosynthetic Pathway of Nuciferine,” ACS Omega, vol. 3, no. 8, pp. 8794–8802, Aug. 2018. doi: 10.1021/acsomega.8b00827.
    Lotus (Nelumbo nucifera Gaertn.) contains various bioactive compounds, with benzylisoquinoline alkaloids (BIAs) as one of the major groups. The biosynthetic pathways of two major bioactive BIAs in this plant, nuciferine and N-nornuciferine, are still not clear. Therefore, several genes related to BIA biosynthesis were searched from the lotus database to study the role of key genes in regulating these pathways. In this study, the expression profiles of NCS, CNMT, 6OMT, CYP80G2, and WRKY TFs were investigated in mechanically wounded lotus leaves. It was found that the accumulation of nuciferine and N-nornuciferine significantly increased in the mechanically wounded lotus leaves in accordance with the relative expression of putative CYP80G2 and one WRKY transcription factor (NNU_24385), with the coregulation of CNMT. Furthermore, the role of methyltransferase-related genes in this study suggested that methylation of the isoquinoline nucleus to yield a methylated-BIA structure may occur at the N position before the O position. Altogether, this study provides improved understanding of the genes regulating BIA biosynthesis under stressed conditions, which could lead to improvements in BIA production from the commercial lotus.
  139. N. Mehta, P. Ekta, P. Patani, V. Patani, and B. Shah, “Nelumbo Nucifera (Lotus): A Review on Ethanobotany, Phytochemistry and Pharmacology,” vol. 1, pp. 152–167, Jun. 2013. doi: 10.30750/ijpbr.1.4.26.
    Nelumbo nucifera Gaertn (Nymphaeaceae), a perennial aquatic plant, has been used as a medicinal herb in China and India. It has been recorded in the most famous medicinal book in China for more than 400 years. Different part of plant (leaves, seeds, flower, and rhizome) can be used in traditional system of medicine. In traditional system of medicine, the different parts of plant is reported to possess beneficial effects as in for the treatment of pharyngopathy, pectoralgia, spermatorrhoea, leucoderma, smallpox, dysentery, cough, haematemesis, epistaxis, haemoptysis, haematuria, metrorrhagia, hyperlipidaemia, fever, cholera, hepatopathy and hyperdipsia. Following the traditional claims for the use of N.nucifera as cure of numerous diseases considerable efforts have been made by researchers to verify it’s utility through scientific pharmacological screenings. The pharmacological studies have shown that N.nucifera posseses various notable pharmacological activities like amti-ischemic, antioxidant, anticancer, antiviral, antiobesity, lipolytic, hypocholestemic, antipyretic, hepatoprotective, hypoglycaemic, antidiarrhoeal, antifungal, antibacterial, antiinflammatory and diuretic activities. A wide variety of phytoprinciples have been isolated from the plant. The present review is an effort to consolidate traditional, ethnobotanic, phytochemical and pharmacological information available on N.nucifera
  140. L. Mei, F. cheng Xin, X. Na, and L. Ming-yuan, “The Adaption Expressed in Ontogeny of Nelumbo Nucifera Root,” Bulletin of Botanical Research, vol. 19, no. 4, p. 469, Dec. 1999. http://bbr.nefu.edu.cn/EN/.
    The genesis and development of Nelumbo nucifera root and root system i...
  141. Y. Mekbib et al., “The Level of Genetic Diversity and Differentiation of Tropical Lotus, Nelumbo Nucifera Gaertn. (Nelumbonaceae) from Australia, India, and Thailand,” Botanical Studies, vol. 61, no. 1, p. 15, May 2020. doi: 10.1186/s40529-020-00293-3.
    Nelumbo nucifera Gaertn., a perennial aquatic macrophyte species, has been cultivated in several Asian countries for its economic importance, and medicinal uses. Two distinct ecotypes of the species are recognized based on the geographical location where the genotypes are adapted, i.e., tropical lotus and temperate lotus. The genetic diversity levels and differentiation of the tropical lotus from poorly studied geographic regions still remain unclear. Here, the population genetic diversity and structure of 15 tropical lotus populations sampled from the previous understudied natural distribution ranges, including India, Thailand, and Australia, were assessed using nine polymorphic SSR markers.
  142. I. M. Menéndez-Perdomo and P. J. Facchini, “Benzylisoquinoline Alkaloids Biosynthesis in Sacred Lotus,” Molecules, vol. 23, no. 11, p. 2899, Nov. 2018. doi: 10.3390/molecules23112899.
    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.
  143. I. M. Menéndez-Perdomo and P. J. Facchini, “Isolation and Characterization of Two O-Methyltransferases Involved in Benzylisoquinoline Alkaloid Biosynthesis in Sacred Lotus (Nelumbo Nucifera),” Journal of Biological Chemistry, vol. 295, no. 6, pp. 1598–1612, Feb. 2020. doi: 10.1074/jbc.RA119.011547.

    Benzylisoquinoline alkaloids (BIAs) are a major class of plant metabolites with many pharmacological benefits. Sacred lotus (Nelumbo nucifera) is an ancient aquatic plant of medicinal value because of antiviral and immunomodulatory activities linked to its constituent BIAs. Although more than 30 BIAs belonging to the 1-benzylisoquinoline, aporphine, and bisbenzylisoquinoline structural subclasses and displaying a predominant R-enantiomeric conformation have been isolated from N. nucifera, its BIA biosynthetic genes and enzymes remain unknown. Herein, we report the isolation and biochemical characterization of two O-methyltransferases (OMTs) involved in BIA biosynthesis in sacred lotus. Five homologous genes, designated NnOMT1–5 and encoding polypeptides sharing >40% amino acid sequence identity, were expressed in Escherichia coli. Functional characterization of the purified recombinant proteins revealed that NnOMT1 is a regiospecific 1-benzylisoquinoline 6-O-methyltransferase (6OMT) accepting both R- and S-substrates, whereas NnOMT5 is mainly a 7-O-methyltransferase (7OMT), with relatively minor 6OMT activity and a strong stereospecific preference for S-enantiomers. Available aporphines were not accepted as substrates by either enzyme, suggesting that O-methylation precedes BIA formation from 1-benzylisoquinoline intermediates. K\textsubscriptm values for NnOMT1 and NnOMT5 were 20 and 13 μm for (R,S)-norcoclaurine and (S)-N-methylcoclaurine, respectively, similar to those for OMTs from other BIA-producing plants. Organ-based correlations of alkaloid content, OMT activity in crude extracts, and OMT gene expression supported physiological roles for NnOMT1 and NnOMT5 in BIA metabolism, occurring primarily in young leaves and embryos of sacred lotus. In summary, our work identifies two OMTs involved in BIA metabolism in the medicinal plant N. nucifera.

  144. Z. Meng et al., “Chromosome Nomenclature and Cytological Characterization of Sacred Lotus,” Cytogenetic and Genome Research, vol. 153, no. 4, pp. 223–231, 2017. doi: 10.1159/000486777.
    Sacred lotus is a basal eudicot plant that has been cultivated in Asia for over 7,000 years for its agricultural, ornamental, religious, and medicinal importance. A notable characteristic of lotus is the seed longevity. Extensive endeavors have been devoted to dissect its genome assembly, including the variety China Antique, which germinated from a 1,300-year-old seed. Here, cytogenetic markers representing the 10 largest megascaffolds, which constitute approximately 70% of the lotus genome assembly, were developed. These 10 megascaffolds were then anchored to the corresponding lotus chromosomes by fluorescence in situ hybridization using these cytogenetic markers, and a set of chromosome-specific cytogenetic markers that could unambiguously identify each of the 8 chromosomes was generated. Karyotyping was conducted, and a nomenclature based on chromosomal length was established for the 8 chromosomes of China Antique. Comparative karyotyping revealed relatively conserved chromosomal structures between China Antique and 3 modern cultivars. Interestingly, significant variations in the copy number of 45S rDNA were detected between China Antique and modern cultivars. Our results provide a comprehensive view on the chromosomal structure of sacred lotus and will facilitate further studies and the genome assembly of lotus.
  145. J. Mevi-Schutz and W. Grosse, “The Importance of Water Vapour for the Circulating Air Flow through Nelumbo Nucifera,” Journal of Experimental Botany, vol. 39, no. 9, pp. 1231–1236, 1988. doi: 10.1093/jxb/39.9.1231.
    Mevi-Schutz, J. and Grosse, W. 1988. The importance of water vapour for the circulating air flow through Nelumbo nucifera.—J. exp. Bot. 39: 1231-1236.
  146. J. Mevi-Schutz and W. Grosse, “A Two-Way Gas Transport System in Nelumbo Nucifera,” Plant, Cell & Environment, vol. 11, no. 1, pp. 27–34, 1988. doi: 10.1111/j.1365-3040.1988.tb01773.x.
    The aquatic vascular plant Nelumbo nucifera Gaertn. is able to improve its oxygen supply to the submerged and buried organs by a thermo-osmotic gas transport. Investigations with tracer gas and oxygen measurements have shown that thermo-osmotic gas transport exists in N. nucifera when there is a temperature difference between the lacunar air of the leaves and the surrounding atmosphere. The gas transport was increased by up to 935% when a temperature difference of 2.9 ± 1.0 K was detected. Lacunar pressure of up to 166 ± 44 Pa was measured in both young and old leaves. In contrast to the flow-through ventilation system recently described for Nuphar lutea and Nymphoides peltata, a two-way flow in separate air canals in the petioles of both young and old Nelumbo leaves may carry oxygen-rich air down to the rhizome and excess air back to the atmosphere. Anatomical investigations have shown that, in Nelumbo, the two largest air canals of the petiole end directly under the mesh system of the centre plate. These large air canals are proposed to be predominant in the upward flow of air in sunlight. The other air canals of the petiole veer into the leaf blade well below the centre plate. The gas flow system through fresh leaves may carry as much as 10.3 ± 4.5 cm3 air per minute to the buried rhizome.
  147. W. C. Meyer, “Dormancy and Growth Studies of the American Lotus, Nelumbo Lutea,” Plant Physiology, vol. 5, no. 2, pp. 225–234, Apr. 1930. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC440211/.
    Images null
  148. R. E. Miller, J. R. Watling, S. A. Robinson, R. E. Miller, J. R. Watling, and S. A. Robinson, “Functional Transition in the Floral Receptacle of the Sacred Lotus (Nelumbo Nucifera): From Thermogenesis to Photosynthesis,” Functional Plant Biology, vol. 36, no. 5, pp. 471–480, May 2009. doi: 10.1071/FP08326.
    The receptacle of the sacred lotus is the main source of heat during the thermogenic stage of floral development. Following anthesis, it enlarges, greens and becomes a fully functional photosynthetic organ. We investigated development of photosynthetic traits during this unusual functional transition. There were two distinct phases of pigment accumulation in receptacles. Lutein and photoprotective xanthophyll cycle pigments accumulated first with 64 and 95% of the maximum, respectively, present before anthesis. Lutein epoxide comprised 32% of total carotenoids in yellow receptacles, but declined with development. By contrast, more than 85% of maximum total chlorophyll, β-carotene and Rubisco were produced after anthesis, and were associated with significant increases in maximum electron transport rates (ETR) and photochemical efficiency (Fv/Fm). Leaves and mature receptacles had similar Rubisco content and ETRs (>200 μmol m−2 s−1), although total chlorophyll and total carotenoid contents of leaves were significantly higher than those of green receptacles. Receptacle δ13C before anthesis was similar to that of leaves; consistent with leaf photosynthesis being the source of C for these tissues. In contrast, mature receptacles had significantly lower δ13C than leaves, suggesting that 14–24% of C in mature receptacles is the result of refixation of respired CO2.
  149. T. Min et al., “Transcription Profiles Reveal the Regulatory Synthesis of Phenols during the Development of Lotus Rhizome (Nelumbo Nucifera Gaertn),” International Journal of Molecular Sciences, vol. 20, no. 11, p. 2735, Jan. 2019. doi: 10.3390/ijms20112735.
    Lotus (Nelumbo nucifera Gaertn) is a wetland vegetable famous for its nutritional and medicinal value. Phenolic compounds are secondary metabolites that play important roles in the browning of fresh-cut fruits and vegetables, and chemical constituents are extracted from lotus for medicine due to their high antioxidant activity. Studies have explored in depth the changes in phenolic compounds during browning, while little is known about their synthesis during the formation of lotus rhizome. In this study, transcriptomic analyses of six samples were performed during lotus rhizome formation using a high-throughput tag sequencing technique. About 23 million high-quality reads were generated, and 92.14% of the data was mapped to the reference genome. The samples were divided into two stages, and we identified 23,475 genes in total, 689 of which were involved in the biosynthesis of secondary metabolites. A complex genetic crosstalk-regulated network involved in the biosynthesis of phenolic compounds was found during the development of lotus rhizome, and 25 genes in the phenylpropanoid biosynthesis pathway, 18 genes in the pentose phosphate pathway, and 30 genes in the flavonoid biosynthesis pathway were highly expressed. The expression patterns of key enzymes assigned to the synthesis of phenolic compounds were analyzed. Moreover, several differentially expressed genes required for phenolic compound biosynthesis detected by comparative transcriptomic analysis were verified through qRT-PCR. This work lays a foundation for future studies on the molecular mechanisms of phenolic compound biosynthesis during rhizome formation.
  150. R. Ming et al., “Genome of the Long-Living Sacred Lotus (Nelumbo Nucifera Gaertn.),” Genome Biology, vol. 14, no. 5, p. R41, May 2013. doi: 10.1186/gb-2013-14-5-r41.
    Sacred lotus is a basal eudicot with agricultural, medicinal, cultural and religious importance. It was domesticated in Asia about 7,000 years ago, and cultivated for its rhizomes and seeds as a food crop. It is particularly noted for its 1,300-year seed longevity and exceptional water repellency, known as the lotus effect. The latter property is due to the nanoscopic closely packed protuberances of its self-cleaning leaf surface, which have been adapted for the manufacture of a self-cleaning industrial paint, Lotusan.
  151. V. Mishra, “Accumulation of Cadmium and Copper from Aqueous Solutions Using Indian Lotus (Nelumbo Nucifera),” AMBIO: A Journal of the Human Environment, vol. 38, no. 2, pp. 110–112, Mar. 2009. doi: 10.1579/0044-7447-38.2.110.
    Publishing interdisciplinary research addressing the scientific, social, economic, and cultural factors that influence the condition of the human environment.
  152. S. Mitchell, “Aquatic Plant Potting Soil Recipe,” Greater Washington Aquatic Plant Association. Jun-2009. https://gwapa.org/wordpress/articles/aquatic_plant_potting_soil_recipe/.
    Sherry Mitchell maintains a beautiful 2000 gallon pond with several varieties of water lilies. To pot her lilies, and other aquatic plants, Sherry uses this recipe to meet their nutrition requirements.
  153. K. Miyake, “Some Physiological Observations on Nelumbo Nucifera, Gærtn,” Shokubutsugaku Zasshi, vol. 12, no. 141, pp. en85–en101, 1898. doi: 10.15281/jplantres1887.12.85.
  154. C. F. Moro et al., “Immature Seed Endosperm and Embryo Proteomics of the Lotus (Nelumbo Nucifera Gaertn.) by One-Dimensional Gel-Based Tandem Mass Spectrometry and a Comparison with the Mature Endosperm Proteome,” Proteomes, vol. 3, no. 3, pp. 184–235, Aug. 2015. doi: 10.3390/proteomes3030184.
    Lotus (Nelumbo nucifera Gaertn.) seed proteome has been the focus of our studies, and we have recently established the first proteome dataset for its mature seed endosperm. The current study unravels the immature endosperm, as well as the embryo proteome, to provide a comprehensive dataset of the lotus seed proteins and a comparison between the mature and immature endosperm tissues across the seed’s development. One-dimensional gel electrophoresis (SDS-PAGE) linked with tandem mass spectrometry provided a protein inventory of the immature endosperm (122 non-redundant proteins) and embryo (141 non-redundant proteins) tissues. Comparing with the previous mature endosperm dataset (66 non-redundant proteins), a total of 206 non-redundant proteins were identified across all three tissues of the lotus seed. Results revealed some significant differences in proteome composition between the three lotus seed tissues, most notably between the mature endosperm and its immature developmental stage shifting the proteins from nutrient production to nutrient storage.
  155. C. F. Moro et al., “Unraveling the Seed Endosperm Proteome of the Lotus (Nelumbo Nucifera Gaertn.) Utilizing 1DE and 2DE Separation in Conjunction with Tandem Mass Spectrometry,” PROTEOMICS, vol. 15, no. 10, pp. 1717–1735, 2015. doi: 10.1002/pmic.201400406.
    Nelumbo nucifera (Gaertn.) or lotus, is an aquatic plant native to India, and presently consumed as food mainly in China and Japan. Lotus is also widely used in Indian and Chinese traditional medicine. Extracts from different parts of the lotus plant have been reported to show diverse biological activities—antioxidant, free radical scavenging, anti-inflammatory, and immunomodulatory. Despite this, little work has been done in isolating and identifying proteins responsible for these activities, or yet importantly to establish a lotus proteome. The aim of our group is to develop a proteome catalog of the lotus plant, starting with its seed, the nutrient rich food source. In this present study, the seed endosperm—most abundant in proteins, and main nutrient storage tissue—was targeted for protein extraction by testing five different extraction protocols, followed by their proteomic analyses using complementary 1DE and 2DE approaches in conjunction with MS/MS. The inventory of 66 nonredundant proteins obtained by 1DE-MS and the 30 obtained by 2DE-MS provides the first catalog of the lotus seed endosperm, where the most abundant protein functions were in categories of metabolic activities related to carbohydrate metabolism and nutrient storage.
  156. P. K. Mukherjee, D. Mukherjee, A. K. Maji, S. Rai, and M. Heinrich, “The Sacred Lotus (Nelumbo Nucifera)– Phytochemical and Therapeutic Profile,” Journal of Pharmacy and Pharmacology, vol. 61, no. 4, pp. 407–422, 2009. doi: 10.1211/jpp.61.04.0001.
    Objectives Nelumbo nucifera Gaertn. (Nymphaeaceae), also known as sacred lotus, is a well known medicinal plant. This article reviews the traditional uses, phytochemistry and therapeutic reports on different parts of N. nucifera viz. the seeds, rhizomes, leaves and flowers. This review also describes various compounds isolated from different parts of this plant and the therapeutic benefits derived from those phytoconstituents. Key findings There are several therapeutic benefits of this plant for which different parts are used. The extracts of rhizomes, seeds, flowers and leaves have been reported to have varied therapeutic potential. Several bioactive compounds have been derived from these plant parts belonging to different chemical groups, including alkaloids, flavonoids, glycosides, triterpenoid, vitamins etc., which all have their own therapeutic impact. Thus, the pharmacological effects and various active ingredients of different parts of N. nucifera are well understood. Summary In this review we explore the current pharmaceutical, phytochemical and pharmacological knowledge about this well known plant species as well as several emerging aspects for research on N. nucifera.
  157. T. NAGASHIMA, “On the Flowering of Tuusonji-Lotus (Nelumbo Nucifera) Preserved over 800 Years,” Research bulletin, no. 32, pp. 1–17, Mar. 2001. https://ci.nii.ac.jp/naid/110000483265.
  158. T. Nagashima, “The Leaves Formed in the First Node of Main Stem Growing out of a Terminal Bud of the Seed-Rhizome of Indian Lotus, Nelumbo nucifera GAERTN,” Journal of the Japanese Society for Horticultural Science, vol. 46, no. 2, pp. 201–210, 1977. doi: 10.2503/jjshs.46.201.
    The Indian Lotus, Nelumbo nucifera GAERTN. has two kinds of leaves. They are the standing leaf that stands at the surface of the water and the floating leaf that rises to the surface of the water. The first leaf which is formed in the first node of the main stem growing out of a terminal bud of the seed-rhizome (a storage stem) becomes the standing leaf, or the floating leaf. What conditions will make the first leaf in the first node of the main stem the standing leaf, or the floating one?The present investigation is aimed at the effect of the amount of applied fertilizer (zero, standard, and double) and shading treatment with cheesecloth (roof type shading and tunnel type shading) on these leaves. The result may be summarized by the following points:1. Both at the control and at the roof type shading, the first leaves formed in the first node of the main stem have become the standing leaves. But by the amount of the standard and double fertilizer applied with the tunnel type shading, they have become the floating leaves respectively, where the relative humidity was remarkably higher than other treatment.2. The thickening grade of the cell wall, that composes the hypodermis in the petiole of the first leaves of the main stem, was more remarkable in the standing leaves than in the floating leaves. The Phloroglucin Hydrochloric Acid Reaction was in proportion to the thickening grade of the cell wall; namely in the remarkable thickening in the petiole of the standing leaf the Phloroglucin Hydrochloric Acid Reaction was remarkable. Slight thickening at the petiole of the floating leaf was not remarkable.3. The thickening grade of the fibrous cell, that composes the hypodermis in the petiole of the first leaves of the main stem, was more remarkable in the standing leaf than in the floating leaf. The morphology of the fibrous cell in the petiole was almost the same as in both the standing leaf and floating leaf. The thickening of the fibrous cell in the standing leaf was remarkable and in the floating leaf it was slight. It was found that the histological difference of the standing leaf and floating leaf depended upon the thickening grade of the fibrous cells.
  159. S. Nakamura et al., “Alkaloid Constituents from Flower Buds and Leaves of Sacred Lotus (Nelumbo Nucifera, Nymphaeaceae) with Melanogenesis Inhibitory Activity in B16 Melanoma Cells,” Bioorganic & Medicinal Chemistry, vol. 21, no. 3, pp. 779–787, Feb. 2013. doi: 10.1016/j.bmc.2012.11.038.
    Methanolic extracts from the flower buds and leaves of sacred lotus (Nelumbo nucifera, Nymphaeaceae) were found to show inhibitory effects on melanogenesis in theophylline-stimulated murine B16 melanoma 4A5 cells. From the methanolic extracts, a new alkaloid, N-methylasimilobine N-oxide, was isolated together with eleven benzylisoquinoline alkaloids. The absolute stereostructure of the new alkaloid was determined from chemical and physicochemical evidence. Among the constituents isolated, nuciferine, N-methylasimilobine, (−)-lirinidine, and 2-hydroxy-1-methoxy-6a,7-dehydroaporphine showed potent inhibition of melanogenesis. Comparison of the inhibitory activities of synthetic related alkaloids facilitated characterization of the structure-activity relationships of aporphine- and benzylisoquinoline-type alkaloids. In addition, 3–30μM nuciferine and N-methylasimilobine inhibited the expression of tyrosinase mRNA, 3–30μM N-methylasimilobine inhibited the expression of TRP-1 mRNA, and 10–30μM nuciferine inhibited the expression of TRP-2 mRNA.
  160. O. V. Nakonechnaya and M. S. Yatsunskaya, “Genetic and Genotypic Variation of Nelumbo Komarovii Grossh,” Russian Journal of Genetics, vol. 54, no. 7, pp. 816–824, Jul. 2018. doi: 10.1134/S1022795418070116.
    Genetic and genotypic variabilities of a rare relict aquatic plant Nelumbo komarovii Grossh. from six natural populations of Primorsky krai (Russia) were assessed using isozyme analysis. The highest genetic and genotypic diversities were observed in the Razdol’noe and Okeanskaya populations, which can be explained by cross-pollination participation in the seed formation for maintenance of the Razdol’noe population size and by sowing seeds from different habitats for the Okeanskaya population. Low polymorphism at the population level (P = 8.97, HO = 0.055, HE = 0.039) is determined by the history of existence of the species from the Tertiary Period as well as by the influence of gene drift. Clonal renewal plays a decisive role in the propagation of N. komarovii in most natural populations.
  161. Q. V. Nguyen, D. Hicks, Rural Industries Research and Development Corporation (Australia), and Asian Foods Research and Development, Exporting Lotus to Asia: An Agronomic and Physiological Study : A Report for the Rural Industries Research and Development Corporation. Barton, A.C.T.: RIRDC, 2001.
  162. S. A. Nichols, “The Interaction between Biology and the Management of Aquatic Macrophytes,” Aquatic Botany, vol. 41, no. 1-3, p. 28, 1991. http://pubs.er.usgs.gov/publication/70016365.
    ’Management’ refers to controlling nuisance aquatic species and to restoring or restructing aquatic plant communities. Producing stable, diverse, aquatic plant communities containing a high percentage of desirable species is a primary management goal. There are a variety of techniques including harvesting, herbicides, water-level fluctuation, sediment alteration, nutrient limitation, light alteration, and biological controls which can be used for managing macrophytes. These techniques are briefly reviewed along with discussions of biological considerations important to the efficacy of the technique and the environmental impacts of the technique. There is a growing interest in restoring and restructing aquatic plant communities. Techniques for community restoration are discussed as are emerging management technologies using growth regulators and bioengineering. New management technologies will probably be limited by costs and environmental impacts. In the near future, better macrophyte management will come through better planning and more effective use of present technology. The challenge is to make current planning and management techniques more effective through increased biological inputs. The potential for biological input ranges from subcellular biology to species biology, to community and ecosystem biology. Some information needs are identified. ?? 1991.
  163. M. Nishimura-Tagui et al., “Case of Anaphylaxis Due to Lotus Root,” The Journal of Dermatology, vol. 47, no. 6, pp. e227–e228, 2020. doi: 10.1111/1346-8138.15329.
  164. Y. Nisikado and K. Watanabe, “On the Rhizome Rot of Lotus, Nelumbo Nucifera Gaertn.,caused by a New Fusarium, F. Bulbigenum Wr. Nelumbicolum Nis. Et Wat.,” Berichte des Ohara Instituts für landwirtschaftliche Forschungen, vol. 10, no. 1, pp. 1–8, 1953. http://ousar.lib.okayama-u.ac.jp/49907.
  165. “Blue Nelumbo Nucifera Seeds | World Seed Supply.” Nov-2014. https://www.worldseedsupply.com/product/nelumbo-nucifera-blue-blue-lotus-seeds/.
    Offering several colors of lotus and waterlily seeds to choose from
  166. “Evaluation of Sacred Lotus (Nelumbo Nucifera Gaertn.) as an Alternative Crop for Phyto-Remediation - ProQuest.” . https://www.proquest.com/openview/0f64854f2d27fcd8da26bfc4689b8283/1?pq-origsite=gscholar&cbl=18750.
    Explore millions of resources from scholarly journals, books, newspapers, videos and more, on the ProQuest Platform.
  167. C. NODA and J. TATSUMI, “Morphology and Function of Apparatus Found on the Surface of Central Plate of Leaf Blade in Nelumbo Nucifera,” 日本作物学会東海支部会報, no. 126, pp. 19–21, 1998. https://ci.nii.ac.jp/naid/110001796286.
  168. S. Nohara and T. Tsuchiya, “Effects of Water Level Fluctuation on the Growth of Nelumbo Nucifera Gaertn. in Lake Kasumigaura, Japan,” Ecological Research, vol. 5, no. 2, pp. 237–252, 1990. doi: 10.1007/BF02346994.
    Investigations were made of the growth of Nelumbo nucifera, an aquatic higher plant, in a natural stand in Lake Kasumigaura. A rise of 1.0 m in the water level after a typhoon in August 1986 caused a subsequent decrease in biomass of N. nucifera from the maximum of 291 g d.w. m−2 in July to a minimum of 75 g d.w. m−2. The biomass recovered thereafter in shallower regions. The underground biomass in October tended to increase toward the shore. The total leaf area index (LAI) is the sum of LAI of floating leaves and emergent leaves. The maximum total LAI was 1.3 and 2.8 m2 m−2 in 1986 and 1987, respectively. LAI of floating leaves did not exceed 1 m2 m−2. The elongation rates of the petiole of floating and emergent leaves just after unrolling were 2.6 and 3.4 cm day−1, respectively. The sudden rise in water level (25 cm day−1) after the typhoon in August 1986 caused drowning and subsequent decomposition of the mature leaves. Only the young leaves were able to elongate, allowing their laminae to reach the water surface. The fluctuation in water level, characterized by the amplitude and duration of flooding and the time of flooding in the life cycle, is an important factor determining the growth and survival of N. nucifera in Lake Kasumigaura.
  169. S. Nohara, “Growth of the Indian Lotus (Nelumbo Nucifera GAERTN.) and the Influence of Tuber Density on Foliage Structure and Biomass,” Japanese Journal of Limnology (Rikusuigaku Zasshi), vol. 57, no. 3, pp. 235–243, 1996. doi: 10.3739/rikusui.57.235.
    The growth of Nelumbo nucifera GAERTN. was investigated at an artificial stand in outdoor concrete ponds. The ratio of above- to below-ground biomass (T/R ratio) ranged from 0.3 to 3. Although N. nucifera is a perennial, the reproductive organ (tuber) begins to form later in the growing period than in annual plants; in addition, the life span of the tuber is less than one year. Maximum relative growth rate was 0.037 g⋅g-1⋅day-1 in July.The leaf area index of floating leaves did not exceed 1 m2⋅m-2.The influence of initial tuber density on the foliage structure and below-ground biomass of N. nucifera were investigated. The initial death rate at the time of lamina rolling increased with leaf density. Control of the density of floating leaves was done twice ; once at the early stage of lamina rolling, and again at the late stage of the fully expanded leaf. The below-ground biomass in the following spring did not correlate with initial tuber size. When the tuber biomass was under about 400 g dry weight⋅m-2, net production of below-ground biomass of the N. nucifera stand seems to have attained a plus value. N. nucifera exhibited typical pioneer characteristics in an environment of frequent disturbances such as flooding, i.e., a high T/R ratio, high vegetative propagation by stolons and a short life span of the tuber.
  170. S. Nohara and M. Kimura, “Growth Characteristics of Nelumbo Nucifera Gaertn. in Response to Water Depth and Flooding,” Ecological Research, vol. 12, no. 1, p. 11, Apr. 1997. doi: 10.1007/BF02523605.
    Three experiments on the effects of water depth and flooding onNelumbo nucifera Gaertn. were made in the artificial environment of concrete ponds. First, plants were harvested in autumn after growing under seven different water levels ranging from 0.2–3 m The number of floating leaves, the total number of leaves and the leaf area index of emergent leaves were greatest in the tanks at 0.5 m depth. The petiole dry weight per unit length of emergent leaves and the ratio of aboveground to belowground biomass rose with increasing water depth up to 2 m. In contrast, that of floating leaves was constant at about 10 mg dry weight cm−1. The proportion of biomass in tubers fell from 20% at 0.2 m to 6% at 2 m. Second, petiole elongation responses to the amplitude of flooding were investigated in early summer. The maximum rate of petiole elongation was 25 cm per day at 2.4 m water depth. This was the maximum depth at whichN. nucifera could grow. No petioles could elongate from 3 m to 5 m depth. Finally, the effects of timing of flooding on growth were investigated. At the end of growing season, the belowground biomass of plants in the flooding treatment in late summer was smallest among the flooding treatment plants (P<0.05), and was most severe when flooding occurred in this season. Based on the results of these experiments, the growth characteristics ofN. nucifera in relation to petiole elongation, biomass allocation, and flooding tolerance were discussed.
  171. I. Ohga, “A Comparison of the Life Activity of Century–Old and Recently Harvested Indian Lotus Fruits,” American Journal of Botany, vol. 13, no. 10, pp. 760–765, 1926. doi: 10.1002/j.1537-2197.1926.tb05909.x.
  172. I. Ohga, “On the Fertilization of Nelumbo Nucifera,” Cytologia, vol. FujiiJubilaei, no. 2, pp. 1033–1035, 1937. doi: 10.1508/cytologia.FujiiJubilaei.1033.
  173. I. Ohga, “The Germination of Century-Old and Recently Harvested Indian Lotus Fruits, with Special Reference to the Effect of Oxygen Supply,” American Journal of Botany, vol. 13, no. 10, pp. 754–759, 1926. doi: 10.1002/j.1537-2197.1926.tb05908.x.
  174. Y. Orimoto and M. Takei, “Influence of reducing nitrogen application rate with coated fertilizer in combination with irrigation water saving on yield of east Indian Lotus (Nelumbo nucifera) and nitrogen effluent,” Japanese Journal of Soil Science and Plant Nutrition (Japan), 2007. doi: 10.20710/dojo.78.4_363.
    East Indian Lotus (Nelumbo nucifera) is mainly cultivated in the Lake Kasumigaura and Kitaura Basin in Ibaraki Prefecture, and the effluent nitrogen from the fields is considered one of the causes of nutrient enrichment of the lakes. Two experimental plots were set up to develop a method for reducing the nitrogen effluent: 1) a conventional plot, where 240 kg/ha/year of nitrogen divided into three portions of 80 kg/ha a basal dressing with 2 top-dressings-was applied and an irrigation gate was controlled manually to keep the soil surface flooded throughout the year to prevent rhizome rot; 2) a low-input plot, where the nitrogen rate was reduced to 29% of the conventional rate with a single application of coated fertilizer as the basal dressing, and the irrigation gate was controlled automatically with an auto-irrigator for water saving. The yield and nitrogen balance of each plot were examined. The yield of East Indian Lotus at the low-input plot was almost equal to that at the conventional plot. The net nitrogen input to the low-input plot during the crop period (341 d) was 210 kg/ha, 30% less than that to the conventional plot. Thus, the effluent nitrogen from the low-input plot during the crop period was 68 kg/ha, 41% less than that from the conventional plot. The decrease in effluent nitrogen at the low-input plot was due mainly to the decrease in the amount of surface drainage as an effect of irrigation water saving.
  175. W. Orozco-Obando et al., “PHYTO-REMEDIATION OF NUTRIENT RUN-OFF FROM A RECIRCULATING AQUACULTURE SYSTEM USING LOTUS (NELUMBO NUCIFERA GAERTN.) MAY 06, 2012,” p. 26.
  176. Y. Pan, G. Han, Z. Mao, Y. Zhang, J. Huang, and L. Qu, “The Anatomy of Lotus Fibers Found in Petioles of Nelumbo Nucifera,” Aquatic Botany, vol. 95, no. 2, pp. 167–171, Aug. 2011. doi: 10.1016/j.aquabot.2011.05.002.
    Lotus fibers are the isolated helical secondary cell wall thickenings from tracheary elements of lotus (Nelumbo nucifera Gaertn) petioles. In this study the anatomical characteristics of lotus petioles and microstructures of tracheary elements were studied using light microscopy (LM) and scanning electron microscopy (SEM). The results show that vascular bundles of lotus petioles are scattered throughout ground tissue. Their tracheary elements are of various sizes and there are several patterns of secondary wall thickening present. However, only secondary thickening in a ribbon-like helical pattern can be drawn out from the petiole to form lotus fibers for subsequent utilization. Study of the microstructure of the tracheary elements reveals that there are two pit structures present in the end walls in addition to pits with intact pit membranes: those with porose or web-like remnants pit membrane and those that lack pit membranes. This is an indication of the transitional stage between tracheids and vessel elements. This study provides supportive evidence that lotus fibers are found in both helically thickened tracheids and helically thickened primitive vessels.
  177. L. Pan, X. Wang, J. Jin, X. Yu, and J. Hu, “Bioinformatic Identification and Expression Analysis of Nelumbo Nucifera microRNA and Their Targets,” Applications in Plant Sciences, vol. 3, no. 9, p. 1500046, 2015. doi: 10.3732/apps.1500046.
    Premise of the study: Sacred lotus (Nelumbo nucifera) is a perennial aquatic herbaceous plant of ecological, ornamental, and economic importance. MicroRNAs (miRNAs) play an important role in plant development. However, reports of miRNAs and their role in sacred lotus have been limited. Methods: Using the homology search of known miRNAs with genome and transcriptome contig sequences, we employed a pipeline to identify miRNAs in N. nucifera. We also predicted the targets of these miRNAs. Results: We found 106 conserved miRNAs in N. nucifera, and 456 of their miRNA targets were annotated. Quantitative real-time PCR (qRT-PCR) analysis revealed the different expression levels of the 10 selected conserved miRNAs in tissues of young leaves, stems, and flowers of N. nucifera. Negative correlation of expression level between five miRNAs and their target genes was also revealed. Discussion: Combining bioinformatics and experiment analysis, we identified the miRNAs in N. nucifera. The results can be used as a workbench for further investigation of the roles of miRNAs in N. nucifera.
  178. L. Pan et al., “Genetic Diversity and Differentiation of Lotus (Nelumbo Nucifera) Accessions Assessed by Simple Sequence Repeats,” Annals of Applied Biology, vol. 159, no. 3, pp. 428–441, 2011. doi: 10.1111/j.1744-7348.2011.00509.x.
    Nelumbo nucifera (lotus) is a perennial aquatic crop of substantial economical and ecological importance. Currently, the evaluation of the genetic variation of lotus germplasm accessions using codominant simple sequence repeat (SSR) markers is significant, and it is essential for understanding the population structure of N. nucifera. Here we report the genetic diversity and differentiation of 92 N. nucifera accessions (82 cultivated varieties and 10 wild lotus) using 50 polymorphic SSR markers. A total of 195 alleles were detected, with an average of 3.9 alleles/locus. The mean polymorphic information content (PIC) and the mean expected heterozygosity were 0.43 and 0.50, respectively. The genetic relationships among accessions were estimated using an unweighted pair-group method with arithmetic average (UPGMA) cluster and principal coordinate analysis (PCA). Both methods revealed that the lotus accessions from China and those from its adjacent Asian countries formed a single cluster, respectively. The cultivated varieties were correlated with their major characteristics in cultivation (the seed, rhizome and flower type) rather than their geographic distribution. On the basis of the Bayesian model-based analyses, two genetically distinct groups (the seed lotus group and the rhizome lotus group) were generated, with a strong differentiation between them (FST = 0.57). The seed lotus group exhibited higher genetic diversity than did the rhizome lotus group. The results herein indicated that the current levels of genetic diversity and differentiation between the lotuses have been greatly influenced by artificial selection.
  179. L. Pan et al., “Isolation and Characterization of Microsatellite Markers in the Sacred Lotus (Nelumbo Nucifera Gaertn.),” Molecular Ecology Notes, vol. 7, no. 6, pp. 1054–1056, 2007. doi: 10.1111/j.1471-8286.2007.01774.x.
    The sacred lotus (Nelumbo nucifera Gaertn.) is an aquatic plant of economic and ornamental importance in China. From an (AG)n-enriched genomic library, 24 microsatellites were isolated and identified by using the (fast isolation by the AFLP of sequences containing repeats) FIASCO protocol. Eleven loci showed polymorphism with two to six alleles per locus. These markers yielded 42 alleles in a survey of 32 accessions of the sacred lotus. Eleven effective primer pairs of simple sequence repeats were designed and will be used as genetic markers to evaluate the fine-scale population structure of the sacred lotus in the future.
  180. K. R. Paudel and N. Panth, “Phytochemical Profile and Biological Activity of Nelumbo Nucifera,” Evidence-Based Complementary and Alternative Medicine, vol. 2015, pp. 1–16, 2015. doi: 10.1155/2015/789124.
    Nelumbo nucifera Gaertn. (Nymphaeaceae) is a potential aquatic crop grown and consumed throughout Asia. All parts of N. nucifera have been used for various medicinal purposes in various systems of medicine including folk medicines, Ayurveda, Chinese traditional medicine, and oriental medicine. Many chemical constituents have been isolated till the date. However, the bioactive constituents of lotus are mainly alkaloids and flavonoids. Traditionally, the whole plant of lotus was used as astringent, emollient, and diuretic. It was used in the treatment of diarrhea, tissue inflammation, and homeostasis. The rhizome extract was used as antidiabetic and anti-inflammatory properties due to the presence of asteroidal triterpenoid. Leaves were used as an effective drug for hematemesis, epistaxis, hemoptysis, hematuria, and metrorrhagia. Flowers were used to treat diarrhea, cholera, fever, and hyperdipsia. In traditional medicine practice, seeds are used in the treatment of tissue inflammation, cancer and skin diseases, leprosy, and poison antidote. Embryo of lotus seeds is used in traditional Chinese medicine as Lian Zi Xin, which primarily helps to overcome nervous disorders, insomnia, and cardiovascular diseases (hypertension and arrhythmia). Nutritional value of lotus is as important as pharmaceutical value. These days’ different parts of lotus have been consumed as functional foods. Thus, lotus can be regarded as a potential nutraceutical source.
  181. R. E. Paull, A. Carroll, and N. J. Chen, “Lotus Cell Walls and the Genes Involved in Its Synthesis and Modification,” Tropical Plant Biology, vol. 6, no. 2, pp. 152–160, Sep. 2013. doi: 10.1007/s12042-013-9129-x.
    The lotus genome (Nelumbo nucifera (Gaertn.)) lacks the paleo-triplication found in other eudicots and has evolved remarkably slowly with fewer nucleotide mutations. It is thought to have greater retention of duplicated genes than other angiosperms. We evaluated the potential genes involved in cell wall synthesis and its modification, and ethylene synthesis and response. In many cell wall transferases and hydrolases families, lotus had fewer members in most families when compared to Arabidopsis. Lotus had similar or fewer members in each family as found in poplar, grape and papaya. The exceptions were in the sialyl and beta-glucuronsyl transferases where similar number were found as in the core eudicots. Lotus had similar numbers of polygalacturonase and pectin methyl esterases as found in Arabidopsis but fewer in all other hydrolases families. For starch degradation, lotus had only two alpha amylases predicted genes versus eight to ten in other eudicots, with similar numbers of beta amylase genes predicted. Lotus also had less than half the number of genes predicted for the enzymes involved in lignin and tannin synthesis compared to Arabidopsis. The stress plant growth regulator ethylene’s synthesis, reception and response predicted genes were fewer in lotus than other eudicots. Only two ethylene receptor genes were predicted in lotus with five reported for Arabidopsis and six for tomato. Our analysis does not supports the conclusion that this species has greater retention of duplicated genes though our data does support the conclusion that lotus split occurred at the base of the eudicots.
  182. R. Pearl, “Variation in the Number of Seeds of the Lotus,” The American Naturalist, vol. 40, no. 479, pp. 757–768, Nov. 1906. doi: 10.1086/278678.
  183. W. Ping-he, C. Wei-pei, and C. Rui-yang, “Study on the Karyotype Analysis of Nymphaeaceae and Its Taxonomic Position,” Journal of Systematics and Evolution, vol. 32, no. 4, p. 293, Jul. 1994. https://www.jse.ac.cn/EN/.
    The present paper reports the karyotypes of six species in the family...
  184. D. A. Priestley and M. A. Posthumus, “Extreme Longevity of Lotus Seeds from Pulantien,” Nature, vol. 299, no. 5879, pp. 148–149, Sep. 1982. doi: 10.1038/299148a0.
    In 1923 Ohga1 first brought to general scientific attention the existence of a cache of viable seeds of East Indian lotus (Nelumbo nucifera Gaertn.), contained within an ancient lakebed deposit at Pulantien in southern Manchuria. From the geological and historical evidence available, Ohga suggested a possible age in excess of 400 years1,2. Libby3 radiocarbon dated some of Ohga’s material at 1040±210 years BP. This evidence of great antiquity for viable seeds has been controversial4–6. The main hindrance to the resolution of the problem has been the paucity of available Pulantien seeds following the dissipation of Ohga’s original collection7. Recently we have received four Pulantien seeds, three of which were viable, albeit lacking in vigour. The lipids of the unimbibed seeds were examined and found to be still highly polyunsaturated, suggesting that they had undergone little atmospheric autoxidation. Radiocarbon dating of one of the viable seeds suggested a probable age of about 466 years at the time of germination. This is the oldest viable seed for which an age has been directly determined.
  185. H. Qi, F. Yu, R. N. Damaris, and P. Yang, “Metabolomics Analyses of Cotyledon and Plumule Showing the Potential Domestic Selection in Lotus Breeding,” Molecules, vol. 26, no. 4, p. 913, Jan. 2021. doi: 10.3390/molecules26040913.
    Lotus (Nelumbo nucifera) seeds are widely consumed as functional food or herbal medicine, of which cotyledon (CL) is the main edible part, and lotus plumule (LP) is commonly utilized in traditional Chinese medicine. However, few studies have been conducted to investigate the chemical components of CL and LP in dry lotus seeds, not to mention the comparison between wild and domesticated varieties. In this study, a widely targeted metabolomics approach based on Ultra Performance Liquid Chromatography-electrospray ionization-Tandem mass spectrometry (UPLC-ESI-MS/MS) was utilized to analyze the metabolites in CL and LP of China Antique (“CA”, a wild variety) and Jianxuan-17 (“JX”, a popular cultivar). A total of 402 metabolites were identified, which included flavonoids (23.08% to 27.84%), amino acids and derivatives (14.18–16.57%), phenolic acids (11.49–12.63%), and lipids (9.14–10.95%). These metabolites were classified into ten clusters based on their organ or cultivar-specific characters. Most of these metabolites were more abundant in LP than in CL for both varieties, except for metabolites belonging to organic acids and lipids. The analysis of differentially accumulated metabolites (DAMs) demonstrated that more than 25% of metabolites detected in our study were DAMs in CL and LP comparing “JX” with “CA”, most of which were less abundant in “JX”, including 35 flavonoids in LP, 23 amino acids and derivatives in CL, 7 alkaloids in CL, and 10 nucleotides and derivatives in LP, whereas all of 11 differentially accumulated lipids in LP were more abundant in “JX”. Together with the fact that the seed yield of “JX” is much higher than that of “CA”, these results indicated that abundant metabolites, especially the functional secondary metabolites (mainly flavonoids and alkaloids), were lost during the process of breeding selection.
  186. K. Qingdong, F. Xinfa, Y. Baoguo, Y. Yuanying, and K. Weidong, “A Preliminary Study on Lotus Breeding Method,” Acta Horticulturae, no. 402, pp. 278–282, Jul. 1995. doi: 10.17660/ActaHortic.1995.402.45.
    Lqtus is a kind of plant with vigorus capacity of propagation. They may be propagated with asexual method by bud and sexual method by seed. Dur to a diversity of lotus propagation and pecularity of its reproductive organ, so wer have adopted the various methods in order to improve its quantitative and qualitative characteristics. Systematic selection from natural mixed population. Selection from progeny of terminal bud propagation. Selection from progeny of natural crossing seedling. Selection from progeny of artificial crossing seedling. We can fix the haterosis of lotus hybrid by asexual propagation. So we don’t need many generations for purifing the progeny by the self crossing or preparing hybrid seed by artificial cross annually. Thus, this is a effective way for lotus breeding. We listed some characteristics of hybrid(F1) in this paper and introduce the five new varieties which was bred by the above methods. Lotus is a kind of plant with vigrous capacity of Propagation, espicially for this asexual propagation. In fact, the lotus is a rhizome, the terminal bud of lotus rhizome is a very shortened shoot. There is a bud axis inside terminal bud and a growing point on its top. The growing point can be differentiated into leaf meristem and floral meristem and axile bud meristem successfully. In keeping pace with growth of bud axis, a leaf or floral meristem will be developed into leaves or flowers. A axile bud will be growing up to a lotus rhizome. Manu lotus rhizomes are growing up parallel under earth. The terminal bud of main rhizome can be also differentiated into many branching rhizome. Every bud of branching rhizomes can be also differentiated and growed up independently, The few terminal internodes of each rhizomes can be expanded to a new lotus in an optimum circumstances. Thus the lotus can be propagated generation after generation. This is the main process of lotus by asexual propagation. Lotus may be also propagated by sexual method. The process of sexual propagation may be achieved by the way of blooming, pollinating, fertilization and fruiting. Due to a diversity of lotus propagation method and pecularity of its reproductive organ, we have adopted the following various methods in order to improve the lotus variety.
  187. S. Riya et al., “Role of Leaves in Methane Emission from Sacred Lotus (Nelumbo Nucifera),” Aquatic Botany, vol. 163, p. 103203, Apr. 2020. doi: 10.1016/j.aquabot.2020.103203.
    Sacred lotus (Nelumbo nucifera) is known to vent pressurized air in the leaf blade through stomata on the central plate of the leaf via the rhizome. This study measured diurnal CH4 emissions from lotus-cultivated in tanks during July and August 2016. CH4 emissions from the central plate of leaves of different age were also evaluated. Average daily CH4 fluxes of lotus-cultivated tanks were 23.1\,± 5.6 and 11.4\,± 6.0 mg C m−2 h−1 during July and August, respectively. Both the diurnal pattern of flow rate and the CH4 concentration of gas emitted from the central plate were similar to those of lotus and other wetland plants. However, the diurnal pattern of CH4 flux of younger leaves was dissimilar to that of tanks with cultivated lotus. Younger leaves exhibited a higher flow rate and greater concentration of CH4 than older leaves. This implies that pressurized air in older leaves is emitted through younger leaves, which contrasts with other plants that emit gas from older leaves or columns via pressurization.
  188. M. G. Ryon et al., “Technique for Rapid Establishment of American Lotus in Remediation Efforts,” Native Plants Journal, vol. 14, no. 1, pp. 33–38, Mar. 2013. doi: 10.3368/npj.14.1.33.
    A technique for increasing the establishment rate of American lotus (Nelumbo lutea Willd. [Nelumbonaceae]) and simplifying planting was developed as part of a pond remediation project. Lotus propagation techniques typically require scarification of the seeds, germination in heated water, and planting in nursery containers. Then mature (approximately 1 y) nursery-grown stock is transferred to the outplanting site, or scarified seeds are simply broadcast applied to the outplanting site. Mature plants should grow more quickly but can be sensitive to handling, require more time to plant, and cost more. Scarified seeds are easier to plant and are inexpensive but have a lag time in growth, can fail to germinate, and can be difficult to site precisely. We developed an intermediate technique using small burlap bags that makes planting easier, provides greater germination success, and avoids lag time in growth. Data on survival and growth from experiments using mature stock, scarified seeds, and bag lotus demonstrate that bag lotus grow rapidly in a variety of conditions, have a high survival rate, can be processed and planted easily and quickly, and are suitable for a variety of remediation projects.
  189. J. Sayre, “Propagation Protocol for American Lotus ( Nelumbo Lutea Willd.),” Native Plants Journal, vol. 5, no. 1, pp. 14–17, 2004. doi: 10.1353/npj.2004.0017.
    The Native Plants Journal 5.1 (2004) 14-17
  190. E. L. Schneider and J. D. Buchanan, “Morphological Studies of the Nymphaeaceae. Xi. the Floral Biology of Nelumbo Pentapetala,” American Journal of Botany, vol. 67, no. 2, pp. 182–193, 1980. doi: 10.1002/j.1537-2197.1980.tb07640.x.
    The floral biology of Nelumbo pentapetala (Walter) Fernald, the American lotus, native to Texas, was investigated. Anthesis occurs over three consecutive days with flowers opening each morning and closing around noon. First-day flowers are protogynous with the perianth parts partially expanded so that pollen-covered insects which are attracted by floral color and the intense “fruity” odor (diffused with the aid of increased floral temperature) are directed on to the flattened receptacle (= carpellary receptacle) from which the receptive stigmas protrude, thus accomplishing pollination. During the second morning anther dehiscence begins and insects which visit and forage within the flower become covered with pollen and typically crawl over the still receptive stigmas achieving “facilitated” self-pollination (indirect autogamy). By mid-morning of the second day the stigmas dry and become non-receptive to pollen. During the third day of anthesis perianth and staminal parts quickly abscise and over the period of a few weeks the receptacle and enclosed fruits mature. In most populations studied, Hymenoptera (e.g., Lusioglossum spp., and Apis mellifera) were the most abundant and effective pollinators. In some populations, however, Coleoptera (e.g., Chauliognathus) were also numerous and effective pollinators. It is suggested that the overall floral structure (e.g., large numbers of stamens, masses of pollen, staminal appendages) are adaptations which facilitate the pollination of Nelumbo by beetles.
  191. E. L. Schneider and S. Carlquist, “Vessels in Nelumbo (Nelumbonaceae),” American Journal of Botany, vol. 83, no. 9, pp. 1101–1106, 1996. doi: 10.1002/j.1537-2197.1996.tb13889.x.
    Vessels have been previously reported in the roots (but not rhizomes) of Nelumbo on the basis of light microscopy. We have reinvestigated vessel occurrence in Nelumbo by means of scanning electron microscopy (SEM). On the basis of these studies, vessels are characteristic of root metaxylem. Pores in primary walls of pits of end walls of these vessels are various in size, but feature incomplete lysis of pit membranes, often with residual webs or threads of primary wall material, much as in vessels of primitive woody dicotyledons. In addition, we newly report occurrence of vessels in metaxylem of rhizomes; pores in these vessels are smaller and more confined than in those of roots. The present study offers not only data of possible use in determining the phylogenetic position of Nelumbonaceae, but also contributes evidence that vessel origin in Nelumbonaceae relates to habit and ecology. We conclude that organographic distribution of vessels in Nelumbo follows the patterns seen in monocotyledons, which, like Nymphaeaceae and other aquatics, have sympodial architecture.
  192. D. C. Seo et al., “Nutrient Uptake and Release in Ponds under Long-Term and Short-Term Lotus (Nelumbo Nucifera) Cultivation: Influence of Compost Application,” Ecological Engineering, vol. 36, no. 10, pp. 1373–1382, Oct. 2010. doi: 10.1016/j.ecoleng.2010.06.015.
    Uptake and release of nutrients from ponds used for lotus cultivation were measured in ponds under short-term (1 yr) cultivation with compost application (pond I) and under long-term (20 yr) cultivation without compost application (pond II). Total inflow loads of TN (irrigation water, rainfall and compost) during lotus cultivation period in ponds I and II were 72.3 and 34.3kgha−1 182day−1, respectively. TN removal rates in ponds I and II were 77.3 and 49.8% of total inflow load, respectively. Major removal mechanisms of TN were attributed to microbial processes and uptake by lotus. The total outflow loads (infiltration and runoff) of TN during the lotus cultivation period were 13.9kgha−1 182day−1 (19.2% of total inflow TN load) for pond I, and 11.3kgha−1 182day−1 (32.9% of total inflow TN load) for pond II. For TP the total inflow loads (irrigation water, rainfall and compost) during lotus cultivation in ponds I and II were 80.8 and 1.9kgha−1 182day−1, respectively. TP removal rates in ponds I and II were 84.9 and −274.1% of total input, respectively. Phosphorus removal was attributed to lotus uptake and soil adsorption. The total outflow loads (infiltration and runoff) of TP during lotus cultivation period were 10.1kgha−1 182day−1 (12.5% of total inflow TP load) for pond I, and 6.6kgha−1 182day−1 (355.6% of total inflow TP load) for pond II. TN and TP in runoff from pond I (with compost) was higher than that in pond II (without compost), showing that TN and TP in runoff were strongly influenced by compost addition. Therefore, in order to satisfy established water-quality standards, the amount of compost used in lotus cultivation should be evaluated.
  193. R. S. Seymour, K. Ito, Y. Umekawa, P. D. G. Matthews, and S. A. Pirintsos, “The Oxygen Supply to Thermogenic Flowers,” Plant, Cell & Environment, vol. 38, no. 4, pp. 827–837, 2015. doi: 10.1111/pce.12454.
    Thermogenic flowers produce heat by intense respiration, and the rates of O2 consumption (Ṁo2) in some species can exceed those of all other tissues of plants and most animals. By exposing intact flowers to a range of O2 pressures (Po2) and measuring Ṁo2, we demonstrate that the highest respiration rates exceed the capacity of the O2 diffusive pathway and become diffusion limited in atmospheric air. The male florets on the inflorescence of Arum concinnatum have the highest known mass-specific Ṁo2 and can be severely diffusion limited. Intact spadices of Japanese skunk cabbage Symplocarpus renifolius are diffusion limited in air only when Ṁo2 is maximal, but not at lower levels. True flowers of the sacred lotus Nelumbo nucifera and the appendix of Arum concinnatum are never diffusion limited in air. Ṁo2 – Po2 curves are evaluated quantitatively with the ‘Regulation Index’, a new tool to measure dependence of Ṁo2 on ambient Po2, as well as the conventional ‘Critical Po2’. The study also includes measurements of Po2 within thermogenic tissues with O2-sensitive fibre optics, and reveals that the diffusion pathway is complicated and that O2 can be provided not only from the surface of the tissues but also from the pith of the flower’s peduncle.
  194. R. S. Seymour and P. Schultze–Motel, “Physiological Temperature Regulation by Flowers of the Sacred Lotus,” Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, vol. 353, no. 1371, pp. 935–943, Jun. 1998. doi: 10.1098/rstb.1998.0258.
    Flowers of the sacred lotus, Nelumbo nucifera Gaertn. (Nelumbonaceae) are thermogenic and physiologically thermoregulatory. The 42 g flowers remain between 30–36°C during a 2 to 4–day period despite fluctuations in environmental temperatures between about 10–45°C. As the ambient temperature drops, the flowers increase heat production in proportion. Temperature regulation apparently occurs at a cellular level, by a steep, reversible thermal inhibition of respiration at flower temperatures above 30°C. There was a marked time lag between change in flower temperature and compensatory response, suggesting regulation through a biochemical feedback mechanism rather than structural changes in enzymes or membranes. By oxidizing carbohydrate, the flowers produce up to 1 W, with about half of the heat coming from the 8.5 g carpellary receptacle. The period of temperature regulation begins before petal opening and continues through the period of stigma receptivity. Temperature regulation may reward insect pollinators with a warm, equable environment, or it possibly enhances and coordinates flower development.
  195. R. S. Seymour, “Scaling of Heat Production by Thermogenic Flowers: Limits to Floral Size and Maximum Rate of Respiration,” Plant, Cell & Environment, vol. 33, no. 9, pp. 1474–1485, 2010. doi: 10.1111/j.1365-3040.2010.02190.x.
    Effect of size of inflorescences, flowers and cones on maximum rate of heat production is analysed allometrically in 23 species of thermogenic plants having diverse structures and ranging between 1.8 and 600 g. Total respiration rate (, µmol s−1) varies with spadix mass (M, g) according to in 15 species of Araceae. Thermal conductance (C, mW °C−1) for spadices scales according to C = 18.5M0.73. Mass does not significantly affect the difference between floral and air temperature. Aroids with exposed appendices with high surface area have high thermal conductance, consistent with the need to vaporize attractive scents. True flowers have significantly lower heat production and thermal conductance, because closed petals retain heat that benefits resident insects. The florets on aroid spadices, either within a floral chamber or spathe, have intermediate thermal conductance, consistent with mixed roles. Mass-specific rates of respiration are variable between species, but reach 900 nmol s−1 g−1 in aroid male florets, exceeding rates of all other plants and even most animals. Maximum mass-specific respiration appears to be limited by oxygen delivery through individual cells. Reducing mass-specific respiration may be one selective influence on the evolution of large size of thermogenic flowers.
  196. Sharda Dhadse, Shahrukh Nawaj Alam, and M. Mallikarjuna Rao, “Development of Nutrient Rich Biofertilizer by Co-Vermistabilization of Aquatic Weeds Using Herbal Pharmaceutical Wastewater along with Sediment of Lake,” Bioresource technology reports, vol. 13, p. 100633-, Feb. 2021. doi: 10.1016/j.biteb.2021.100633.
    Six aquatic weeds species namely Hydrilla verticilata (L.f.) Royle, Ceratophyllum demursum L., Nelumbo nucifera Gaerth., Ludwigia palustris L., Pistia stratioles L., and Eicchornia crassipis along with Eudrilus eugeniae (earthworm sp.) were selected for vermistabilization. Aquatic sediment, cow dung and microbes were taken in ratio 2:1:1 allowed up to 45 days for stabilization, the study claimed, newly developed vermicompost is a novel kind with a total NPK of 3.32%, 2.03%, 0.62% with enriched microbial flora. Pistia stratioles L. shown potential growth on plants.
  197. Z. Sharip, A. T. A. Zaki, and S. Zakaria, “Flooding Effects on the Population Dynamics of Cabomba Furcata and Nelumbo Nucifera in a Shallow Floodplain Wetland,” Wetlands, vol. 34, no. 4, pp. 713–723, Aug. 2014. doi: 10.1007/s13157-014-0536-z.
    Cabomba furcata is an introduced species that has been naturalized in Malaysia and is becoming invasive in a shallow wetland - Lake Chini, Pahang - displacing the dominance of the native Nelumbo nucifera. The water level structure, which has reduced flooding, plays a vital role in structuring species dominance in this shallow alluvial wetland of the Pahang River floodplain. Biomass sampling focusing on floating-leaved N. nucifera and submerged C. furcata was conducted from September 2009 to March 2011 to determine its association with flooding and season. We found that the non-native C. furcata exhibits no clear seasonal patterns in biomass compared to the native N. nucifera. However, the C. furcata biomass varied with the lake’s water level, which fluctuates with flooding from the Pahang River. High amplitude flooding affected the composition of both C. furcata and N. nucifera. Long-term hydrological and flood data indicate the variability of flooding into the wetland system. The paper concludes by noting the need for concerted sustainable management strategies to be coordinated and managed in addressing Cabomba’s threats.
  198. B. R. Sharma, L. N. S. Gautam, D. Adhikari, and R. Karki, “A Comprehensive Review on Chemical Profiling of Nelumbo Nucifera: Potential for Drug Development,” Phytotherapy Research, vol. 31, no. 1, pp. 3–26, 2017. doi: 10.1002/ptr.5732.
    Nelumbo nucifera, also known as sacred lotus, has primarily been used as food throughout the Asian continent, and its medicinal values have been described in Ayurvedic and Traditional Chinese Medicine. The purpose of this study is to systematically characterize the chemical profiling and pharmacological activities of N. nucifera. Herein, we critically reviewed and analysed the phytochemical and pharmacological reports of N. nucifera. Our search for the keyword ‘Nelumbo nucifera pharmacology’ in all databases reported in Web of Science yielded 373 results excluding reviews and abstracts in document types. Two hundred and forty-three spectrum natural compounds from different parts of N. nucifera belonging to diverse chemical groups, including alkaloids, flavonoids, glycosides, terpenoids, steroids, fatty acids, proteins, minerals, and vitamins have been reported. In addition, distinct pharmacological activities, mainly against cancer, microbial infection, diabetes, inflammation, atherosclerosis, and obesity, have been associated with crude extracts, fractions, and isolated compounds. This review highlights potential use of neferine, liensinine, isoliensinine, and nuciferine in clinical trials. In depth, mechanism of the potential chemical entities from N. nucifera via structure activity relationship needs to be explored to guarantee the stability and safety for the clinical use. Copyright © 2016 John Wiley & Sons, Ltd.
  199. S. C. Sharma and A. K. Goel, “Environmental Degradation and Ex-Situ Conservation of Nelumbo Nucifera,” in Environmental Stress: Indication, Mitigation and Eco-Conservation, M. Yunus, N. Singh, and L. J. de Kok, Eds. Dordrecht: Springer Netherlands, 2000, pp. 405–410. doi: 10.1007/978-94-015-9532-2_35.
    Plants have always played a significant role in day to day human life. The aquatic bodies and wetlands comprise important components of the natural ecosystems. During the last five decades, aquatic habitats are deteriorating rapidly due to extensive demographic pressure, urbanisation, invasion of aquatic weeds and increased inflow of industrial effluents causing environmental degradation and serious threat to natural ecosystems. Soil washed down from denuded slopes and fields, is choking the wetlands and rivers at a very fast rate. As a result, the fragile ecosystems, such as tropical rain forests, lakes and reservoirs, mangroves and coastal areas, which constitute the hotspots of biodiversity are endangered. Global warming is also one of the root cause for destabilizing the equilibrium of the wetland ecosystems.
  200. M. F. Shaw, “A Microchemical Study of the Fruit Coat of Nelumbo Lutea,” American Journal of Botany, vol. 16, no. 5, pp. 259–276, 1929. doi: 10.1002/j.1537-2197.1929.tb09480.x.
  201. S. A. Sheikh, “Ethno-Medicinal Uses and Pharmacological Activities of Lotus (Nelumbo Nucifera),” Journal of Medicinal Plant Studies, vol. 2, no. 6, pp. 42–46, 2014.
    Nelumbo nucifera is grown in many parts of the globe including India for its medicinal and nutritional value. In Kashmir, the plant grows naturally in the lakes and its stem is being extensively used in many famous Kashmiri cuisines. In addition, its fruits and seeds are also consumed, but to a lesser extent. Many studies have established a wide range of the pharmacological activities of this plant. The current review highlights the importance of Nelumbo nucifera in traditional medicines and its pharmacological activities.
  202. J. Shen-Miller et al., “Centuries-Old Viable Fruit of Sacred Lotus Nelumbo Nucifera Gaertn Var. China Antique,” Tropical Plant Biology, vol. 6, no. 2, pp. 53–68, Sep. 2013. doi: 10.1007/s12042-013-9125-1.
    During the Sino-Japanese conflict of the 1920s, Japanese botanist Ichiro Ohga was presented single-seeded fruit of Nelumbo nucifera var. China Antique collected by a local farmer from a dry lakebed in Northeast China (then, “Manchuria”). Ohga studied the fruit and published his findings. Years later, we tested the germination of Nelumbo fruit from the same locality. The oldest seed sprouted, having a germination time of ~3 days, was radiocarbon dated to be ~1300 years old. These cold- and drought-tolerant seeds exhibited shoot-before-root emergence and a primary green plumule capable of “dim-light” photosynthesis. Such traits and the notable long-term viability of the fruit spurred the interest of Ray Ming, University of Illinois that has now led to the sequencing of the Nelumbo genome. Analyses of this genome may provide insight into the biochemistry of Nelumbo on wax-biosynthesis genes, and application of aging-related thermostable proteins to the extension of seed-life and improvement of food quality of economic crops. Here, we review the history of these long-lived Nelumbo fruit, and their occurrence, discovery, collection, propagation, and methods of seedling care. The robust impermeable wax- and suberin-covered pericarp is a major factor contributing to their remarkable longevity. New findings are presented on the modern and 459- and 464-year-old pericarp anatomy, impermeability to water, and whole fruit and pericarp mechanical properties, and comparison of the mode of fruit weight-gain during imbibition and germination time relative to fruit maturity.
  203. J. Shen-Miller, M. B. Mudgett, J. W. Schopf, S. Clarke, and R. Berger, “Exceptional Seed Longevity and Robust Growth: Ancient Sacred Lotus from China,” American Journal of Botany, vol. 82, no. 11, pp. 1367–1380, 1995. doi: 10.1002/j.1537-2197.1995.tb12673.x.
    A 1,288 ± 271-yr-old (1,350 ± 220 yr BP, radiocarbon age) seed of Sacred Lotus (Nelumbo nucifera Gaertn.) from an ancient lake bed at Pulantien, Liaoning Province, China, has been germinated and subsequently radiocarbon dated. This is the oldest demonstrably viable and directly dated seed ever reported, the preserved relict of one of the early crops of lotus cultivated by Buddhists at Pulantien after introduction of the religion into the region prior to 372 A.D. A small portion of the dry pericarp of a second lotus fruit from the same locale has been dated as being 332 ± 135-yr-old (270 ± 60 yr BP, radiocarbon age) by accelerator mass spectroscopy at the Lawrence Livermore National Laboratory. This polycentenarian seed not only germinated but is still growing (since March, 1994). Of six old lotus fruits tested, two-thirds germinated, almost all in fewer than 4 d, as rapidly as fruits harvested from the progeny of Pulantien Sacred Lotus plants (under cultivation by the National Park Service in Washington, DC), and more rapidly than fresh fruits of Yellow Lotus [N. lutea (Willd.) Pers.]. Growth of the old lotus is robust: rhizome formation and leaf emergence at rhizome nodes are more rapid than those of the Pulantien progeny, although the leaf width is smaller. Activity of the protein-repair enzyme L-isoaspartyl methyltransferase in the old lotus seed is persistent during germination and is as robust as that in the progeny, and the degree of aspartyl racemization in proteins of the two groups of plants is minimal and essentially identical. The six dated ancient Sacred Lotus fruits range in age from 95 to 1,288 yr (with a mean age of 595 ± 380 yr), evidently reflecting their production, deposition, and preservation at varying times during the intervening millennium.
  204. J. Shen-Miller et al., “Long-Living Lotus: Germination and Soil Gamma-Irradiation of Centuries-Old Fruits, and Cultivation, Growth, and Phenotypic Abnormalities of Offspring,” American Journal of Botany, vol. 89, no. 2, pp. 236–247, Feb. 2002. doi: 10.3732/ajb.89.2.236.
    Sacred lotus (Nelumbo nucifera) has been cultivated as a crop in Asia for thousands of years. An ∼1300-yr-old lotus fruit, recovered from an originally cultivated but now dry lakebed in northeastern China, is the oldest germinated and directly (14)C-dated fruit known. In 1996, we traveled to the dry lake at Xipaozi Village, China, the source of the old viable fruits. We identified all of the landmarks recorded by botanist Ichiro Ohga some 80 yr ago when he first studied the deposit, but found that the fruits are now rare. We (1) cataloged a total of 60 lotus fruits; (2) germinated four fruits having physical ages of 200-500 yr by (14)C dating; (3) measured the rapid germination of the old fruits and the initially fast growth and short dormancy of their seedlings; (4) recorded abnormal phenotypes in their leaves, stalks, roots, and rhizomes; (5) determined γ-radiation of ∼2.0 mGy/yr in the lotus-bearing beds; and (6) measured stratigraphic sequences of the lakebed strata. The total γ-irradiation of the old fruits of 0.1-3 Gy (gray, the unit of absorbed dosage defined as 1 joule/kg; 1 Gy = 100 rad), evidently resulting in certain of the abnormal phenotypes noted in their seedlings, represents the longest natural radiobiology experiment yet recorded. Most of the lotus abnormalities resemble those of chronically irradiated plants exposed to much higher irradiances. Though the chronic exposure of the old fruits to low-dose γ-radiation may be responsible in part for the notably weak growth and mutant phenotypes of the seedlings, it has not affected seed viability. All seeds presumably repair cellular damage before germination. Understanding of repair mechanisms in the old lotus seeds may provide insight to the aging process applicable also to other organisms.
  205. J. Shen-Miller, “Sacred Lotus, the Long-Living Fruits of China Antique,” Seed Science Research, vol. 12, no. 3, pp. 131–143, Sep. 2002. doi: 10.1079/SSR2002112.
    In the West, lotus (Nelumbo nucifera Gaertn.) is relatively little known. However, for more than 3000 years, lotus plants have been cultivated as a crop in Far-East Asia, where they are used for food, medicine and play a significant role in religious and cultural activities. Holder of the world’s record for long-term seed viability (1300 years) is a lotus fruit (China Antique) from Xipaozi, Liaoning Province, China. Five offspring of this variety, from 200–500-year-old fruits (14C dates) collected at Xipaozi, have recently been germinated, and are the first such seedlings to be raised from directly dated fruits. The fruits at Xipaozi, preserved in a dry ancient lakebed, have been exposed to low-dose γ-radiation for hundreds of years (having an accumulated soil irradiation of 0.1–1.0 Gy). Offspring from these old fruits show abnormalities that resemble those in various modern seedlings irradiated at much higher doses. Although these lotus offspring are phenotypically abnormal, the viability of old seeds was evidently not affected by accumulated doses of up to 3 Gy. Growth characteristics of first- and second-year lotus offspring of these fruits, products of the longest-term radiobiological experiment on record, are summarized here (rapid early growth, phenotypic abnormalities, lack of vigour, poor rhizome development and low photosynthetic activity during second-year growth). Aspects of their chromosomal organization, phenotype and physiology (rapid recovery from stress, heat-stable proteins, protein-repair enzyme) are discussed. Important unsolved problems are suggested to elicit interest among members of the seed science community to the study of old fruits recently collected at Xipaozi, with particular emphasis on aspects of ageing and repair.
  206. J. Shen-Miller et al., “Thermal-Stable Proteins of Fruit of Long-Living Sacred Lotus Nelumbo Nucifera Gaertn Var. China Antique,” Tropical Plant Biology, vol. 6, no. 2, pp. 69–84, Sep. 2013. doi: 10.1007/s12042-013-9124-2.
    Single-seeded fruit of the sacred lotus Nelumbo nucifera Gaertn var. China Antique from NE China have been shown to remain viable for as long as ~1,300 years, determined by direct radiocarbon-dating, and to have a germination rate of 84 %. The pericarp, a fruit tissue that encloses the single seeds of Nelumbo, is one of the major factors contributing to fruit longevity. Proteins that are heat stable and have a protective function are equally important to such centuries-long seed viability. We document proteins of Nelumbo fruit that are able to withstand heating, 32 % of which remained soluble in the 110 °C-treated embryo axis of a 549-year-old fruit and 76 % retained fluidity in its cotyledons. The genome of Nelumbo has recently been published and annotated. The amino-acid sequences of 11 “thermal proteins” (soluble at 100 °C) of modern Nelumbo embryo axes and cotyledons, identified by mass spectrometry, Western blot and bioassay, are assembled and aligned with those of an archaeal hyperthermophile Methancaldococcus jannaschii (“Mj,” an anaerobic methanogen having a growth optimum of 85 °C) and with those of five mesophile angiosperms. These thermal proteins have roles in protection and repair under stress. More than half (55 %) of the durable Nelumbo thermal proteins are present in the archaean Mj, indicating their ancient history. One Nelumbo protein-repair enzyme exhibits activity at 100 °C, having a heat-tolerance higher than the comparable enzyme of Arabidopsis. A list of 30 sequenced but unassembled thermal proteins of Nelumbo is appended.
  207. J. Sheng, X. Li, and D. Zhang, “Gibberellins, Brassinolide, and Ethylene Signaling Were Involved in Flower Differentiation and Development in Nelumbo Nucifera,” Horticultural Plant Journal, Jun. 2021. doi: 10.1016/j.hpj.2021.06.002.
    Hormones play important roles in vegetative and reproductive processes; however, the regulatory roles of hormones in Nelumbo nucifera (Lotus) growth and development are unclear. In this study, nine types of endogenous hormones, including gibberellins (GA1, GA3, and GA4), indole-3-acetic acid (IAA), brassinolide (BR), ethylene (ETH), jasmonic acid (JA), abscisic acid (ABA), and zeatin (ZT) were detected in dormant shoot tips, vegetative shoot tips, developing leaf buds, and developing flower buds of lotus. The results indicated that GA, ETH, and BR signaling can promote vegetative and reproductive development of lotus. GA signaling regulates plant height and stimulates flower bud differentiation. GA levels were the highest in the flower buds; exogenous GA3+4 increased plant height by approximately 90%, increased flower quantity by nearly 40%, and advanced flowering by 4 d. Suppressing GA biosynthesis using paclobutrazol decreased plant height and flower quantity by 38% and 87.1%, respectively, and delayed flowering by 15.6 d. ETH signaling has positive regulatory effects on vegetative growth and flower development. The ETH concentration in the developing leaf buds was at least 50% higher than that in other samples. Ethephon spraying led to remarkable increases in plant height and leaf thickness and extended the flowering duration. BR signaling acts as a growth promoter during vegetative and reproductive development in lotus. The highest BR levels were detected in the vegetative shoot tips. External application of 28-epihomobrassinolide resulted in growth-promoting phenotypes including longer scapes, thicker leaves, and prolonged flowering.
  208. T. Shi, K. Wang, and P. Yang, “The Evolution of Plant microRNAs: Insights from a Basal Eudicot Sacred Lotus,” The Plant Journal, vol. 89, no. 3, pp. 442–457, 2017. doi: 10.1111/tpj.13394.
    microRNAs (miRNAs) are important noncoding small RNAs that regulate mRNAs in eukaryotes. However, under which circumstances different miRNAs/miRNA families exhibit different evolutionary trajectories in plants remains unclear. In this study, we sequenced the small RNAs and degradome from a basal eudicot, sacred lotus (Nelumbo nucifera or lotus), to identify miRNAs and their targets. Combining with public miRNAs, we predicted 57 pre-eudicot miRNA families from different evolutionary stages. We found that miRNA families featuring older age, higher copy and target number tend to show lower propensity for miRNA family loss (PGL) and stronger signature of purifying selection during divergence of temperate and tropical lotus. Further analyses of lotus genome revealed that there is an association between loss of miRNA families in descendent plants and in duplicated genomes. Gene dosage balance is crucial in maintaining those preferentially retained MIRNA duplicates by imposing stronger purifying selection. However, these factors and selection influencing miRNA family evolution are not applicable to the putative MIRNA-likes. Additionally, the MIRNAs participating in lotus pollen–pistil interaction, a conserved process in angiosperms, also have a strong signature of purifying selection. Functionally, sequence divergence in MIRNAs escalates expression divergence of their target genes between temperate and tropical lotus during rhizome and leaf growth. Overall, our study unravels several important factors and selection that determine the miRNA family distribution in plants and duplicated genomes, and provides evidence for functional impact of MIRNA sequence evolution.
  209. S.-Y. Shou, L.-X. Miao, W.-S. Zai, X.-Z. Huang, and D.-P. Guo, “Factors Influencing Shoot Multiplication of Lotus (Nelumbo Nucifera),” Biologia plantarum, vol. 52, no. 3, pp. 529–532, Sep. 2008. doi: 10.1007/s10535-008-0103-7.
    Effect of plant growth regulators, explant size, season of explant collection, temperature (20, 25 and 30 °C) and photoperiod on in vitro lotus (Nelumbo nucifera Gaertn.) shoot formation and growth were examined. Shoots formation was greatly influenced by growth regulators, explant size and season of explant collection. The maximum number of shoots were induced from bud explants on Murashige and Skoog (MS) medium containing 4.44 µM benzyladenine (BA) + 0.54 µM α-naphthalene acetic acid (NAA). Explants formed by bud of one expanded and one unexpanded leaf, which was collected in spring gave encouraging results of shoot production. Higher temperature favoured shoot induction and subsequent growth was much better at 25 °C compared to that at 20 and 30 °C.
  210. Shouhui Dai, Jing Qiu, and Min Wang, “Concentration-Dependent Enantioselective Accumulation of Chiral Polychlorinated Biphenyls in Nelumbo Nucifera Gaertn. Root from Contaminative Sediment,” Environmental science and pollution research, vol. 28, no. 22, pp. 27878–27884, Jun. 2021. doi: 10.1007/s11356-021-12530-8.
    Nelumbo nucifera Gaertn. (lotus) roots were collected from contaminated sediments which were artificially adding different concentrations of chiral polychlorinated biphenyls (PCBs) to investigate the effect of concentration on the accumulation characteristics and chiral signatures of PCBs in lotus root during its growth period of 150 days. Under high PCB exposure concentration, the biota-sediment accumulation factors (BSAFs) of PCBs 91, 95, and 136 in the lotus root were up to 0.25–0.46 and 8.10–10.5 times higher than those under low-exposure concentration (0.024–0.052). The BSAFs of PCBs 149, 176, and 183 under high-exposure concentration were up to 0.24–0.44, while they were undetected at low concentration. The significant difference observed in the BSAFs based on different concentrations indicates that the lotus root accumulation efficiency toward chiral PCBs increases with the contaminate concentration. Although the (–)-enantiomers of PCBs 91, 95, and 136 were all preferentially accumulated in lotus root under two exposure concentrations, the extent of the preferential accumulation of (–)-PCB 95 decreased with increasing exposure concentration throughout the whole growth period (30–150 days). In addition, the (–)-enantiomers of PCBs 91 and 136 also showed the same tendency during most of the growth period. Conclusively, the exposure concentrations are an important influence factor on the enantioselective accumulation of chiral PCBs in lotus root.
  211. J. R. Snow, “Establishment and Competitive Ability of Nelumbo Lutea in Relation to Myriophyllum Spicatum,” Thesis or Dissertation, University of North Texas, 2000. https://digital.library.unt.edu/ark:/67531/metadc2694/.
    Limitations from reduced light and increasing water depth on Nelumbo lutea seedlings were determined in tank experiments. Survival was high in all tested light levels. Total biomass increased significantly with increasing light. Biomass allocation shifted significantly to root production between 3 and 6 weeks in the 10 and 24% levels. Survival decreased with increasing planting depth, and biomass of survivors reduced significantly between 0.5, 1.0, and 1.5 m depths. Nelumbo lutea and Myriophyllum spicatum populations were monitored for one season in a 0.7 ha pond to track changes in species dominance. Myriophyllum spicatum dominated early, and N. lutea dominated from July through October, suppressing M. spicatum at all depths. Competitive interactions between N. lutea and M. spicatum were investigated for two seasons in a container experiment situated within a pond. Where established, N. lutea dominated in the presence of M. spicatum. However, N. lutea could not be established in depths greater than 1 meter.
  212. S. H. Sohmer, “The Name of the American Nelumbo,” TAXON, vol. 24, no. 4, pp. 491–493, 1975. doi: 10.2307/1219504.
    Although the binomial Nelumbo pentapetala (Walter) Fernald was published in 1934 as the correct name for the American Nelumbo, it has not been generally accepted. The combination is based on Nymphaea pentapetala Walter, a nomenclaturally legitimate, although inappropriate, name. The rejection of Fernald’s combination on the grounds that it was based on a monstrosity does not seem pertinent. The type for the taxon is Walter’s description as no trace has been found of the specimen upon which Walter based his name and description.
  213. Y. Sun et al., “DIA-Based Quantitative Proteomics Reveals the Protein Regulatory Networks of Floral Thermogenesis in Nelumbo Nucifera,” International Journal of Molecular Sciences, vol. 22, no. 15, p. 8251, Jan. 2021. doi: 10.3390/ijms22158251.
    The sacred lotus (Nelumbo nucifera) can maintain a stable floral chamber temperature between 30 and 35 °C when blooming despite fluctuations in ambient temperatures between about 8 and 45 °C, but the regulatory mechanism of floral thermogenesis remains unclear. Here, we obtained comprehensive protein profiles from receptacle tissue at five developmental stages using data-independent acquisition (DIA)-based quantitative proteomics technology to reveal the molecular basis of floral thermogenesis of N. nucifera. A total of 6913 proteins were identified and quantified, of which 3513 differentially abundant proteins (DAPs) were screened. Among them, 640 highly abundant proteins during the thermogenic stages were mainly involved in carbon metabolism processes such as the tricarboxylic acid (TCA) cycle. Citrate synthase was identified as the most connected protein in the protein-protein interaction (PPI) network. Next, the content of alternative oxidase (AOX) and plant uncoupling protein (pUCP) in different tissues indicated that AOX was specifically abundant in the receptacles. Subsequently, a protein module highly related to the thermogenic phenotype was identified by the weighted gene co-expression network analysis (WGCNA). In summary, the regulation mechanism of floral thermogenesis in N. nucifera involves complex regulatory networks, including TCA cycle metabolism, starch and sucrose metabolism, fatty acid degradation, and ubiquinone synthesis, etc.
  214. S.-S. Sun, P. F. Gugger, Q.-F. Wang, and J.-M. Chen, “Identification of a R2R3-MYB Gene Regulating Anthocyanin Biosynthesis and Relationships between Its Variation and Flower Color Difference in Lotus (Nelumbo Adans.),” PeerJ, vol. 4, p. e2369, Sep. 2016. doi: 10.7717/peerj.2369.
    The lotus (Nelumbonaceae: Nelumbo Adans.) is a highly desired ornamental plant, comprising only two extant species, the sacred lotus (N. nucifera Gaerten.) with red flowers and the American lotus (N. lutea Willd.) with yellow flowers. Flower color is the most obvious difference of two species. To better understand the mechanism of flower color differentiation, the content of anthocyanins and the expression levels of four key structural genes (e.g., DFR, ANS, UFGT and GST) were analyzed in two species. Our results revealed that anthocyanins were detected in red flowers, not yellow flowers. Expression analysis showed that no transcripts of GST gene and low expression level of three UFGT genes were detected in yellow flowers. In addition, three regulatory genes (NnMYB5, NnbHLH1 and NnTTG1) were isolated from red flowers and showed a high similarity to corresponding regulatory genes of other species. Sequence analysis of MYB5, bHLH1 and TTG1 in two species revealed striking differences in coding region and promoter region of MYB5 gene. Population analysis identified three MYB5 variants in Nelumbo: a functional allele existed in red flowers and two inactive forms existed in yellow flowers. This result revealed that there was an association between allelic variation in MYB5 gene and flower color difference. Yeast two-hybrid experiments showed that NnMYB5 interacts with NnbHLH1, NlbHLH1 and NnTTG1, and NnTTG1 also interacts with NnbHLH1 and NlbHLH1. The over-expression of NnMYB5 led to anthocyanin accumulation in immature seeds and flower stalks and up-regulation of expression of TT19 in Arabidopsis. Therefore, NnMYB5 is a transcription activator of anthocyanin synthesis. This study helps to elucidate the function of NnMYB5 and will contribute to clarify the mechanism of flower coloration and genetic engineering of flower color in lotus.
  215. H. Sun et al., “Transcriptome Analysis Provides Strategies for Postharvest Lotus Seeds Preservation,” Postharvest Biology and Technology, vol. 179, p. 111583, Sep. 2021. doi: 10.1016/j.postharvbio.2021.111583.
    The rapid deteriorative quality is a major factor that currently limits storage and transport of fresh lotus seeds. However, the physiological changes and molecular mechanisms of lotus seeds during postharvest storage remains poorly understood. Here, physiological and RNA-sequencing analyses were conducted on the postharvest seeds of seed-lotus cultivar ‘Jianxuan 17’. A rapid increase in starch and protein content was observed, while soluble sugar content was continuously decreased during postharvest storage, which could explain increased hardness and reduced sweetness of lotus seeds. Transcriptome analysis identified a total of 3148 differentially expressed genes (DEGs), and functional enrichment analysis showed six pathways that included starch and sucrose metabolism were commonly enriched in all comparison groups. Most DEGs involved in energy metabolic pathways, such as glycolysis and tricarboxylic acid cycle were down-regulated. Altered starch and soluble sugar contents were associated with significant changes in activity of enzymes involved in starch and sucrose metabolism. In addition, the content of plant hormones including, auxin (IAA), jasmonoyl-isoleucine (JA-Ile) and salicylic acid (SA), increased in lotus seeds during postharvest storage, and the activation of signaling transduction pathways were demonstrated at transcriptional level. These results provide not only valuable gene expression dataset for investigating molecular mechanism underlying changes during postharvest storage, but also is a useful reference for developing further preservation technology of fresh lotus seeds.
  216. K. Takagi, Y. Harazono, S.-ichi Noguchi, A. Miyata, M. Mano, and M. Komine, “Evaluation of the Transpiration Rate of Lotus Using the Stem Heat-Balance Method,” Aquatic Botany, vol. 85, no. 2, pp. 129–136, Aug. 2006. doi: 10.1016/j.aquabot.2006.03.006.
    To reveal the mechanism of transpiration by hydrophytes in the field, it is necessary to evaluate the transpiration rate without the effect of the evaporation from the water surface. In order to test the suitability for evaluating the transpiration rate of lotus (Nelumbo nucifera Gaertn.) leaves in the field, stem heat-balance method was applied and the obtained sap-flow rate was compared with the transpiration rate measured by weighing and with the overall canopy evapotranspiration rate by means of the eddy covariance technique. The transpiration rate estimated with the sap-flow measurements showed good agreement with that obtained from the weighing method. Lotus has many air canals in its petiole to carry oxygen-rich air to the rhizome and methane- and carbon dioxide-rich air back to the atmosphere, but there was little effect of the mass flow of air through these canals on the sap-flow rates. In the field observations, the canopy evapotranspiration rate (0.28mmh−1 at maximum) was nearly equal to the sum of the transpiration rate from all sunlit leaves (0.30mmh−1), and the contribution of the transpiration from shaded leaves and evaporation from the water surface was considered to be minor in the seasons when the leaves were fully developed. Evaluation of bulk leaf conductance revealed that the conductance in the leaf boundary layer of lotus could be low (ca. 0.23molm−2s−1) because of its large leaf area. The low conductance in the leaf boundary layer would increase leaf temperature, which, in turn, would generate air circulation within the plant’s ventilation system. Because there was a linear relationship between transpiration rate and the leaf-to-air vapor-pressure deficit, with no apparent maximum, high vapor-pressure deficits (3.4kPa at maximum) did not appear to cause significant stomatal closure in lotus plants. The stomata of lotus leaves play a role as air inlets to carry oxygen-rich air to the rhizome, so their low sensitivity would help to increase air intake.
  217. M. Takagi, M. Goto, D. Wari, M. Saito, R. N. Perry, and K. Toyota, “Screening of Nematicides against the Lotus Root Nematode, Hirschmanniella Diversa Sher (Tylenchida: Pratylenchidae) and the Efficacy of a Selected Nematicide under Lotus Micro-Field Conditions,” Agronomy, vol. 10, no. 3, p. 373, Mar. 2020. doi: 10.3390/agronomy10030373.
    In Japan, Hirschmanniella diversa is an important pest in lotus cultivation in paddy fields and only lime nitrogen is registered for its control. Therefore, additional nematicides are required to control the nematode. The objective of this study was to screen for an effective nematicide. Fourth-stage juveniles and adults of H. diversa sampled from a lotus field were tested in in vitro solution experiments against 37 pesticides that are registered for the pest control of crops in Japan. Carbamate-based benfuracarb, organophosphate-based fenthion, nereistoxin-based cartap hydrochloride and cyanamide showed nematicidal effects against H. diversa. Benfuracarb at 1 μg/mL showed a nematostatic effect on H. diversa in an agar plate assay. Further, H. diversa treated with benfuracarb did not resume activity 7 days post nematicide treatment when transferred to distilled water. Benfuracarb was tested in micro-field experiments, in which H. diversa density and lotus tuber damage levels were monitored. Results showed that benfuracarb reduced H. diversa densities in the roots during the cultivation period in 2012 and consistently reduced damage levels during a five year study period. Thus, benfuracarb is recommended as an effective nematicide to be used for H. diversa control in lotus cultivation.
  218. N.-J. Teng, Y.-L. Wang, C.-Q. Sun, W.-M. Fang, and F.-D. Chen, “Factors Influencing Fecundity in Experimental Crosses of Water Lotus (Nelumbo Nucifera Gaertn.) Cultivars,” BMC Plant Biology, vol. 12, no. 1, p. 82, Jun. 2012. doi: 10.1186/1471-2229-12-82.
    Breeding programs for the water lotus (Nelumbo nucifera) are hampered by an inability to account for variation in seed set associated with crosses between different cultivars. We studied seed set in two reciprocal crosses between lotus cultivars (‘Guili’\,×\,‘Aijiangnan’ and ‘Molingqiuse’\,×\,‘Qinhuaiyanzhi’) to obtain insights into factors that govern fecundity in these experimental hybrids. Pollen viability, stigma receptivity and embryo development were compared for each hybrid and reciprocal cross.
  219. D. Tian, “Container Production and Post -Harvest Handling of Lotus (Nelumbo) and Micropropagation of Herbaceous Peony (Paeonia),” Doctor of Philosophy, Auburn University, Auburn, Alabama, 2008. https://www.proquest.com/openview/99a0807aa08b42703feaf4d91b4cd4c6/1?pq-origsite=gscholar&cbl=18750.
    Explore millions of resources from scholarly journals, books, newspapers, videos and more, on the ProQuest Platform.
  220. H.-L. Tian, J.-H. Xue, J. Wen, G. Mitchell, and S.-L. Zhou, “Genetic Diversity and Relationships of Lotus (Nelumbo) Cultivars Based on Allozyme and ISSR Markers,” Scientia Horticulturae, vol. 116, no. 4, pp. 421–429, May 2008. doi: 10.1016/j.scienta.2008.02.011.
    Genetic diversity and genetic relationships of lotus (Nelumbo Adanson) cultivars were evaluated using allozyme and ISSR markers. The samples used covered 11 accessions of possible hybrids between Nelumbo nucifera and Nelumbo lutea and 92 accessions of N. nucifera including 69 flower lotus, 13 rhizome lotus, 5 seed lotus and 5 wild lotus. For allozyme studies, a total of 31 alleles at 23 loci of 18 enzyme systems were detected of which 5 (21.7%) loci Aat, Idh, Mdh-2, Pgd, Sod were polymorphic. The loci of Aat and Idh included two alleles, Mdh-2, Pgd and Sod included three alleles. Eighteen genotypes were detected with the 13 alleles of the 5 polymorphic loci. The parameters of average allele number, observed heterozygosity, expected heterozygosity and Shannon information index of 92 N. nucifera samples were 1.35±0.71, 0.06±0.21, 0.05±0.14, 0.10±023, respectively. Thirteen ISSR primers generated 93 loci, of which 37.63% were polymorphic across all samples. The percentage of polymorphic loci, average allele number, expected heterozygosity and Shannon information index of 92 N. nucifera samples were 26.67%, 1.30±0.46, 0.10±0.18 and 0.15±0.25, respectively for the ISSR data. The ‘Bottleneck effect’ and rapid propagation of clones after the ice ages may explain the low genetic diversity of lotus. The dendrograms based on ISSR and allozymes were not congruent. Based on the ISSR data, the 103 samples were divided into the N. nucifera group (Group I), and the group containing inter-specific hybrids between N. nucifera and N. lutea (Group II). The flower lotus, rhizome lotus, and seed lotus each has multiple sources of origin. Plant size, a criterion commonly used in the classification of cultivars of lotus, is not correlated with genetic variation. Flower color is correlated with the cultivar classification to some degree, but its variation is complex in the hybrids.
  221. D. Tian, K. M. Tilt, J. L. Sibley, F. Dane, and F. M. Woods, “Response of Lotus (Nelumbo Sp.) to Container Soil Volume,” Journal of Environmental Horticulture, vol. 27, no. 2, pp. 79–84, Jun. 2009. doi: 10.24266/0738-2898-27.2.79.
    The effect of soil volume on containerized lotus (Nelumbo) production has been underreported. American lotus (Nelumbo lutea Willd.) and three cultivars (‘Embolene’, ‘98 Seed’ and ‘Karizma’) of Asian lotus (N. nucifera Gaertn.) were investigated for growth response to container soil volume in this study. Electrical conductivity, pH, plant growth indices, and plant nutritional content were influenced by container soil volume. Differences in some plant growth indices were significant between treatments with ½ and higher (½ and ¾) container height soil (CHS) in 21 or 29 liter (#5 or #7) containers. However, plant growth indices were generally not different between treatments with ½ and ¾ CHS. Lotus planted in containers with ¼ CHS usually produced the greatest plant height and underground fresh weight, while the largest number of propagules often occurred in containers with ½ or ¾ CHS. The highest number of emerging leaves was observed in plants with ¼ or ½ CHS treatments, with no significant difference in emerging leaf number between lotus grown in containers with ½ and ¾ CHS. Flower number generally decreased as soil level increased. The ¼ and ½ CHS were more efficient than ¾ CHS for lotus production in containers.
  222. D. Tian, K. M. Tilt, J. L. Sibley, F. M. Woods, and F. Dane, “Response of Lotus (Nelumbo Nucifera Gaertn.) to Planting Time and Disbudding,” HortScience, vol. 44, no. 3, pp. 656–659, Jun. 2009. doi: 10.21273/HORTSCI.44.3.656.
    Lotus (Nelumbo) is a highly valued plant with a long history for vegetable, ornamental, and medicinal use. Little information is available on the effects of planting time on performance of lotus, especially when grown in containers. The objectives of this study were to find a suitable planting time and to determine best management practices that are of importance for container lotus production. Effects of planting time and disbudding on plant growth indices in southeast Alabama were evaluated in a container production system for the ornamental lotus, N. nucifera ‘Embolene’. Results indicated that plant growth indices were little influenced by different planting dates in March, but were much influenced by planting dates with a difference over a month between February and May. Plants potted and placed outdoors in March and April performed best, and lotus planted in the greenhouse in February and planted outdoors in February and May performed worst. Flower number was not largely influenced by the planting time, but flowering characteristics, especially the flowering peaks, were different among treatments. Planting lotus outdoors between March and May produced the largest return. Influence of planting time on plant growth indices of lotus appeared to be explained by effects of growth-season climate conditions after planting. Disbudding had no impact on plant height but significantly increased underground fresh weight and the number of propagules. Therefore, disbudding should be considered a best management practice to maximize the yield of rhizomes or propagules. Positive linear, quadratic, or cubic relationships were detected among emerging leaf number, underground fresh biomass, and propagule number. Based on the regression models, the yield of lotus rhizomes or propagules can be predicted by the number of emerging leaves. This research provided a guide for nurseries, researchers, and collectors to select the best time to plant lotus outdoors.
  223. D. Tian and K. Tilt, “’Zhizun Qianban’: A Recognition of an Obscure Lotus (Nelumbo) Cultivar with an Interesting Legend,” HortScience, vol. 46, no. 7, pp. 1044–1045, Jul. 2011. doi: 10.21273/HORTSCI.46.7.1044.
    "‘Zhizun Qianban’: A Recognition of an Obscure Lotus (Nelumbo) Cultivar with an Interesting Legend" published on Jul 2011 by American Society for Horticultural Science.
  224. Ting Min et al., “The Effects of Different Temperatures on the Storage Characteristics of Lotus (Nelumbo Nucifera G.) Root,” Food chemistry, vol. 348, p. 129109-, Jun. 2021. doi: 10.1016/j.foodchem.2021.129109.
    Lotus root (Nelumbo nucifera G.) is a high economic value crop in the world. In this study, the storage characteristics (color, sensory, texture, and fatty acids) of lotus root (“Elian No.5″) were evaluated at different harvest periods (September 2018, October 2018, November 2018, December 2018, and January 2019). Moreover, the storage characteristics were evaluated after the short- term and long-term storage of lotus root at 4 °C and 20 °C. The hardness of lotus root significantly decreased at both temperatures (4 °C and 20 °C) during the first 3 days of storage. In contrast, the decrease in hardness delayed at 4 °C (beyond 3 days of storage). Further, genes related to hardness at different storage temperatures were identified using the RNA-seq and qRT-PCR. The results of this study provide a reference for lotus root storage and a basis for the molecular breeding of longterm-storable lotus root.
  225. Y. Toma, R. Iwamoto, K. Inayoshi, N. Nagasaki, and H. Ueno, “Evaluation of the Growth of Lotus (Nelumbo Nucifera), Yield and Quality of Lotus Root, and Soil Nutrient Dynamics on Shallow Soil Cultivation System,” Japanese Journal of Soil Science and Plant Nutrition, vol. 86, pp. 89–97, Apr. 2015.
    Much effort for harvesting rhizome of lotus has been required on the lotus cultivation. In our study, growth of lotus, yield and quality of rhizome, soil nutrient dynamics and irrigation were evaluated on the shallow soil cultivation system. Chemical (C) and organic (O) fertilizer application plots were set up at the pool installed 10cm depth of soil in 2011 and 2012. Nitrogen (N) and phosphorus application rate were same between two plots, while potassium (K) application in O plot was only 30 of that in C plot due to low concentration of K in organic fertilizer. Yields of rhizome (3.15 to 3.93 kg m 2) were larger than that in conventional cultivation, and there was no difference in the yield between the plots. In C plot, rhizome of 100200 g was tended to larger than in O plot might be due to much K application in C plot. That is supported by the result more than 50 of K in lotus was concentrated in the rhizome. Color of the rhizome was almost same between two plots. However, color of the rhizome in 2013 was significantly white compared with that in 2012, because soil iron oxide, which was causal material for brown color of rhizome, might be reduced during the winter. Cation exchange capacity, soil total carbon and N concentration, and exchangeable K concentration reduced 1455 after two cultivations. This showed that soil nutrient reduced by plant uptake should be supplied by organic matter or other materials. Irrigation amount of water during the cultivation was approximately 1,000 mm. However, water should be supplied when evapotranspiration was greater than precipitation, especially in August and September. This study showed that high yield and quality of lotus rhizome was promising at the soil shallow cultivation system, though management for maintaining soil fertility would be required.
  226. N. T. Q. Trang, H. T. K. Avtar, V. T. M. Huong, and D. T. Long, “In Vitro Propagation of Red Lotus (Nelumbo Nucifera Gaertn) - An Aquatic Edible Plant in Vietnam,” Agricultural Science Digest - A Research Journal, no. Of, Oct. 2020. doi: 10.18805/ag.D-257.
    Background: Lotus is an important aquatic plant with great economic value, not only as an ornamental flower but also as a source of herbal medicine. In general, lotus is usually propagated through the rhizome or tuber but the normal propagation rate is very low and it often depends on the quality of water environment and the weather conditions where it is grown. Lotus propagation by tissue culture has many predominant advantages compared with the traditional propagation methods. The current study aims to study the in vitro propagation of Hue’s Red Lotus, a famous local lotus in Hue city, Vietnam.
  227. T. Tsuchiya and S. Nohara, “Growth and Life Span of the Leaves of Nelumbo Nucifera Gaertn. in Lake Kasumigaura, Japan,” Aquatic Botany, vol. 36, no. 1, pp. 87–95, Dec. 1989. doi: 10.1016/0304-3770(89)90094-6.
    Growth of leaves of Nelumbo nucifera Gaertn. in Lake Kasumigaura was investigated with special emphasis on leaf dynamics. Floating leaves were found throughout the growing period, while emergent leaves were not found until July. However, emergent leaves contributed approximately 75% of the total leaf area index at its seasonal maximum, 2.76 m2 m−2. Mean leaf life span of emergent leaves with thick laminae and stiff petioles was much longer than that of floating leaves (44.5 vs. 17.1 days).
  228. D. Tungmunnithum, S. Renouard, S. Drouet, J.-P. Blondeau, and C. Hano, “A Critical Cross-Species Comparison of Pollen from Nelumbo Nucifera Gaertn. vs. Nymphaea Lotus L. for Authentication of Thai Medicinal Herbal Tea,” Plants, vol. 9, no. 7, p. 921, Jul. 2020. doi: 10.3390/plants9070921.
    "Bau Luang" or Nelumbo nucifera Gaertn. is an aquatic medicinal herb that has been used as a component of traditional medicines, medicinal products, and herbal tea for good health, particularly in Asia. The stamen of N. nucifera is an important part of this medicinal plant that is used in the form of dried and/or powdered stamens for herbal tea as well as the main ingredient of some traditional remedies. However, there is another aquatic herb called "Bau Sai" or Nymphaea lotus L. that is distributed in similar locations. Living plants of these two aquatic species may be classified according to their morphology, but the dried and powdered stamens of these two medicinal species are difficult to distinguish. The major reason of adulteration is the higher price of Bau Luang stamen. As a result, various methods of authentication, such as pollen micromorphology evaluation using scanning electron microscopy (SEM) analysis, bioinformatics analysis of two nuclear and plastic DNA markers, phytochemical stamen profiling, and Fourier transform infrared (FTIR) analysis of stamen plant material authentication from Bau Luang and Bau Sai, have been used in this present research in order to avoid some adulteration and/or misuse between the dried stamens of Bau Luang and Bau Sai. These results showed that the micro-morphology of pollen (size of pollen grain, number of apertures, and surface ornamentation) from the SEM analysis, some phytochemical compounds and the FTIR sporopollenin-to-protein ratio signal analysis are potential tools for authentication and identification of these two medicinal plants from their dried-stamen materials. This model of investigation may also be used to distinguish dried plant material from other problematic plant groups.
  229. T. Ushimaru, S. Kanematsu, M. Katayama, and H. Tsuji, “Antioxidative Enzymes in Seedlings of Nelumbo Nucifera Germinated under Water,” Physiologia Plantarum, vol. 112, no. 1, pp. 39–46, 2001. doi: 10.1034/j.1399-3054.2001.1120106.x.
    Dry seeds of anoxia-tolerant lotus (Nelumbo nucifera Gaertn=Nelumbium speciosum Willd.) have green shoots with plastids containing chlorophyll, so photosynthesis starts even in seedlings germinated under water, namely hypoxia. Here we investigated antioxidative enzyme changes in N. nucifera seedlings responding to oxygen deficiency. The activity of superoxide dismutase (SOD; EC 1.15.1.1), dehydroascorbate reductase (DHAR; EC 1.8.5.1) and glutathione reductase (GR; EC 1.6.4.2) were lower in seedlings germinated under water (submerged condition) in darkness (SD seedlings) than those found in seedlings germinated in air and darkness (AD seedlings). In contrast, ascorbate peroxidase (APX; EC 1.11.1.11) activity was higher in SD seedlings and the activity of catalase (EC 1.11.1.6) and monodehydroascorbate reductase (MDAR; EC 1.6.5.4) in SD seedlings was nearly the same as in AD seedlings. When SD seedlings were exposed to air, the activity of SOD, DHAR and GR increased, while the activity of catalase and MDAR decreased. Seven electrophoretically distinct SOD isozymes were detectable in N. nucifera. The levels of plastidic Cu,Zn-SODs and Fe-SOD in SD seedlings were comparable with those found in AD seedlings, which may reflect the maintenance of green plastids in SD seedlings as well as in AD seedlings. These results were substantially different from those previously found in rice seedlings germinated under water.
  230. P. F. Van Bergen, P. G. Hatcher, J. J. Boon, M. E. Collinson, and J. W. de Leeuw, “Macromolecular Composition of the Propagule Wall of Nelumbo Nucifera,” Phytochemistry, vol. 45, no. 3, pp. 601–610, Jun. 1997. doi: 10.1016/S0031-9422(96)00880-1.
    The macromolecular constituents of the sclerotic propagule wall of Nelumbo nucifera and seed coat of Nymphaea caerulea were studied using scanning electron- and light microscopy in combination with Curiepoint pyrolysis-gas chromatography-mass spectrometry. In addition, the Nelumbo material was analyzed using solid state 13C nuclear magnetic resonance and in-source pyrolysis-mass spectrometry. The sclerotic seed coat of the Nymphaea caerulea revealed the presence of angiosperm lignin-cellulose similar to that found in most sclerotic plant remains. In sharp contrast, the fruit wall plus seed coat of Nelumbo is believed to be composed of a complex of polysaccharides, based on primarily galactose and mannose units, and insoluble tannins, which are suggested to play the same structural role as the lignin-cellulose in the sclerotic seed coat. The distinctive nature of the chemical constituents present in the propagule wall of Nelumbo, supports the systematic distinction of this genus in the separate family Nelumbonaceae. The characteristic chemical composition of the propagule walls of Nelumbo could be an additional factor in favour of a prolonged longevity of these fruits. However, the distinctive composition of polysaccharides and tannins without the presence of lignin is considered to be the main reason for the absence of these propagules in the fossil record, despite their physical resistance.
  231. W. a. M. Van Leeuwen, “A Study of the Structure of the Gynoecium of Nelumbo Lutea (Willd.) Pers.,” Acta Botanica Neerlandica, vol. 12, no. 1, pp. 84–97, 1963. doi: 10.1111/j.1438-8677.1963.tb00109.x.
  232. S. Vogel, “Contributions to the Functional Anatomy and Biology of Nelumbo Nucifera (Nelumbonaceae) I. Pathways of Air Circulation,” Plant Systematics and Evolution, vol. 249, no. 1, pp. 9–25, Oct. 2004. doi: 10.1007/s00606-004-0201-8.
    The thermo-osmotically driven air stream that temporarily ventilates the plant body of Nelumbo nucifera, supplying rhizome and roots with oxygen, takes its way through interconnected canals and caverns. The course of this circulation, starting with the uptake of gas through leaf stomata and ending with its expulsion via the specialized navel (“central plate”) of the leaves, as well as organs regulating the circuit, were examined using anatomical, pneumatic, and silicone casting methods. The aerenchyma of the peltate leaf, closely paralleling the nervature, is divided into four separate domains: two mirroring lateral halves including halves of the central plate, and each half again separated into an adaxial (basal) and abaxial (distal) sector. Gas absorbed by the adaxial sectors flows through a definite pair of pipes down the petiole into three lateral pairs of the eight main tubes of the horizontal shoot. There it passes several nodes with adjoining leaves, where homologous ducts, arranged in series along the shoot, contribute to the gas flux in its basipetal course. A reverse gas flow, confined to another pair of cauline pipes and also arranged in series, enters in the nodes another pair of petiolar pipes that directly lead to the foliar central plates, where it is released through stomatal pores. These pores are three times as large as the laminal stomata and control the gas release by opening and closing. Each abaxial leaf sector is, apart from its ducts leading down the petiole, via a shortcut connected with the upstream pipes supplying the central plate, so that its air circuit, when active, is only intralaminar and does not join the entirety of the system. The question where, and how, the two downstreaming currents merge into the two upstreaming ones remains unresolved. The structure of the two types of air canal diaphragms, petiolar and nodal, are documented by SEM.
  233. S. Vogel, “Contributions to the Functional Anatomy and Biology of Nelumbo Nucifera (Nelumbonaceae) II. Unique Emergent Druses on the Floral Receptacle,” Plant Systematics and Evolution, vol. 249, no. 1, pp. 27–35, Oct. 2004. doi: 10.1007/s00606-004-0202-7.
    Apart from the typical, multifaceted calcium oxalate druses distributed in the mesenchyma of Nelumbo nucifera, another, unique type of druse idioblast exists in this plant. They loosely cover the lateral surface of the obconical flower receptacle. Based on light and SEM microscopy, their structure and ontogeny are described for the first time. The crystal complex consists of a flat, usually six-rayed, sharp-pointed star positioned above, and parallel to, the epidermal surface. The idioblast is supported and elevated by six adjacent columnar epidermal cells alternating with the crystal spikes. The spikes represent individual, lamellate crystals symmetrically arranged around a core of crystal sand. In the ontogeny, six steps can be discerned: 1) Enlargement and dome-like emergence of the idioblast. 2) Appearance of vacuoles below the distal endopolyploid nucleus. 3) The vacuoles coalesce and minute crystal particles collect in the vacuole and aggregate to a cluster. 4) A small six-rayed primary druse forms at a variable position. 5) The druse migrates to the cell top, becomes strictly parallel to the epidermal surface, and while the cell head further expands, a second star complex develops, becoming superimposed on the primary druse, its spikes bulging out the thin cell wall. 6) The cytoplasm vanishes and the proximal cell section becomes compressed by the surrounding columnar cells. Because the star-like bodies are confined to the nutrient-rich receptacle, are freely exposed and easily detached, they represent specialized modifications of the internal druses and apparently serve as a deterrent against herbivory.
  234. S. Vogel and F. Hadacek, “Contributions to the Functional Anatomy and Biology of Nelumbo Nucifera (Nelumbonaceae) III. An Ecological Reappraisal of Floral Organs,” Plant Systematics and Evolution, vol. 249, no. 3, pp. 173–189, Nov. 2004. doi: 10.1007/s00606-004-0203-6.
    The flower of Nelumbo nucifera displays some highly specialized structural details apparently related to its pollination ecology. The functional significance of these structures, however, is still under debate. This paper presents new anatomical and chemical data that contribute to our knowledge of the floral life of the lotus. Special focus is directed on the source of scent and heat production, the role of the staminal appendages, and stigma characteristics. The receptacle, the staminal appendages and connectives are covered with a papillose, osmogenous epithelium and function as osmophores. A fractioned GC-analysis revealed that mainly 1,4-dimethoxybenzene (1,4-DMB) is emitted by these parts, whereas high amounts of C15 – C17-alkanes, n-alkenes, n-alcadienes and n-alkatrienes dominated the volatile blend that was diffusely emitted by the petals. In accordance with the literature, absolute predominance of 1,4-DMB and pentadecane was found, while some differences occurred in minor components. The active source of heat generation, apparently a means to enhance fragrance volatilization, was the receptacle. The compact mesenchyma underlying its upper platform and flanks is glandular and rests on a storage zone replete with starch that becomes exhausted during the metabolic flare-up. Spot measurements confirm the flower’s capacity to maintain a temperature level of ca. 31 °C by compensating the environmental temperature fluctuations. The staminal appendages warm up because they are close to the receptacle; they are not actively thermogenous, as formerly proposed. Their starch load remains nearly unaffected until defloration. The appendages, besides contributing to olfaction, may represent food bodies. Along with the pollen – shed at the male stage of anthesis – the appendages may function as a primary attractant for coleopteran pollinators, an interpretation corroborated by the remaining floral traits, which suggest the cantharophilous syndrome. Published field data, so far only available for the American subspecies, reveal variable mixed assemblages of insect visitors; no feeding on the staminal appendages is recorded. Possibly, the system once involved a partnership that has not persisted during the long history of the genus, which is nowadays attended by opportunists.
  235. A. M. Wagner, K. Krab, M. J. Wagner, and A. L. Moore, “Regulation of Thermogenesis in Flowering Araceae: The Role of the Alternative Oxidase,” Biochimica et Biophysica Acta (BBA) - Bioenergetics, vol. 1777, no. 7, pp. 993–1000, Jul. 2008. doi: 10.1016/j.bbabio.2008.04.001.
    The inflorescences of several members of the Arum lily family warm up during flowering and are able to maintain their temperature at a constant level, relatively independent of the ambient temperature. The heat is generated via a mitochondrial respiratory pathway that is distinct from the cytochrome chain and involves a cyanide-resistant alternative oxidase (AOX). In this paper we have used flux control analysis to investigate the influence of temperature on the rate of respiration through both cytochrome and alternative oxidases in mitochondria isolated from the appendices of intact thermogenic Arum maculatum inflorescences. Results are presented which indicate that at low temperatures, the dehydrogenases are almost in full control of respiration but as the temperature increases flux control shifts to the AOX. On the basis of these results a simple model of thermoregulation is presented that is applicable to all species of thermogenic plants. The model takes into account the temperature characteristics of the separate components of the plant mitochondrial respiratory chain and the control of each process. We propose that 1) in all aroid flowers AOX assumes almost complete control over respiration, 2) the temperature profile of AOX explains the reversed relationship between ambient temperature and respiration in thermoregulating Arum flowers, 3) the thermoregulation process is the same in all species and 4) variations in inflorescence temperatures can easily be explained by variations in AOX protein concentrations.
  236. Z. Wang et al., “Alkaloids from Lotus (Nelumbo Nucifera): Recent Advances in Biosynthesis, Pharmacokinetics, Bioactivity, Safety, and Industrial Applications,” Critical Reviews in Food Science and Nutrition, vol. INVALID_SCITE_VALUE, no. INVALID_SCITE_VALUE, pp. 1–34, Nov. 2021. doi: 10.1080/10408398.2021.2009436.
    Different parts of lotus (Nelumbo nucifera Gaertn.) including the seeds, rhizomes, leaves, and flowers, are used for medicinal purposes with health promoting and illness preventing benefits. The presence of active chemicals such as alkaloids, phenolic acids, flavonoids, and terpenoids (particularly alkaloids) may account for this plant’s pharmacological effects. In this review, we provide a comprehensive overview and summarize up-to-date research on the biosynthesis, pharmacokinetics, and bioactivity of lotus alkaloids as well as their safety. Moreover, the potential uses of lotus alkaloids in the food, pharmaceutical, and cosmetic sectors are explored. Current evidence shows that alkaloids, mainly consisting of aporphines, 1-benzylisoquinolines, and bisbenzylisoquinolines, are present in different parts of lotus. The bioavailability of these alkaloids is relatively low in vivo but can be enhanced by technological modification using nanoliposomes, liposomes, microcapsules, and emulsions. Available data highlights their therapeutic and preventive effects on obesity, diabetes, neurodegeneration, cancer, cardiovascular disease, etc. Additionally, industrial applications of lotus alkaloids include their use as food, medical, and cosmetic ingredients in tea, other beverages, and healthcare products; as lipid-lowering, anticancer, and antipsychotic drugs; and in facial masks, toothpastes, and shower gels. However, their clinical efficacy and safety remains unclear; hence, larger and longer human trials are needed to achieve their safe and effective use with minimal side effects.
  237. X. Wang, S. Gui, L. Pan, J. Hu, and Y. Ding, “Development and Characterization of Polymorphic microRNA-Based Microsatellite Markers in Nelumbo Nucifera (Nelumbonaceae),” Applications in Plant Sciences, vol. 4, no. 1, p. 1500091, 2016. doi: 10.3732/apps.1500091.
    Premise of the study: Polymorphic microRNA (miRNA)–based microsatellite markers were developed to investigate the genetic diversity and population structure of Nelumbo nucifera (Nelumbonaceae). Methods and Results: A total of 485 miRNA-based microsatellites were found from the genomic DNA sequences of N. nucifera. After several rounds of screening, 21 primer pairs flanking di-, tri-, or pentanucleotide repeats were identified that revealed high levels of genetic diversity in four populations with two to five alleles per locus. The observed and expected heterozygosity per locus ranged from 0.000 to 1.000 and from 0.000 to 0.803, respectively. Conclusions: The polymorphic microsatellite markers will be useful for studying the genetic diversity and population structure of N. nucifera.
  238. Y. Wang et al., “Effects of Ethylene Biosynthesis and Signaling on Oxidative Stress and Antioxidant Defense System in Nelumbo Nucifera G. under Cadmium Exposure,” Environmental Science and Pollution Research, vol. 27, no. 32, pp. 40156–40170, Nov. 2020. doi: 10.1007/s11356-020-09918-3.
    Water contamination with cadmium (Cd) is a global environmental problem and its remediation becomes urgent. Phytoremediation using ornamental plants has attracted much attention for its advantages of cost-effectiveness and beautification of the environment. Nelumbo nucifera G. is a popular ornamental aquatic macrophyte with fast growth, large biomass, and high capacities for Cd accumulation and removal. However, information about Cd resistance and defense responses in N. nucifera is rather scarce, which restricts its large-scale utilization for phytoremediation. The phytohormone ethylene plays an important role in plant resistance to Cd stress, but the underlying mechanism remains unclear. In this study, we investigated morphophysiological responses of N. nucifera seedlings to Cd stress, and focused on the effects of ethylene on oxidative damage, Cd accumulation, and antioxidant defense system at the metabolic and transcript levels in leaves under Cd stress. Our results showed that Cd exposure led to leaf chlorosis and necrosis, coupled with an increase in contents of hydrogen peroxide, electrolyte leakage, and malondialdehyde, and decrease in chlorophyll content. Exogenous ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) application intensified Cd-induced stress responses and Cd accumulation, and increased ethylene production by inducing ACC synthase (ACS) gene NnACS. Such enhanced ethylene emission inhibited catalase (CAT), ascorbate peroxidase (APX), and glutathione reductase (GR) activities, and modulated ascorbate (AsA) and glutathione (GSH) accumulation through transcriptional regulation of their respective metabolic genes. After ethylene action inhibitor silver thiosulfate (STS) supplementation, Cd-induced oxidative damage was abolished, and Cd content declined but still at a relatively high level. Blocking of ethylene perception by STS inhibited ethylene biosynthesis; enhanced CAT, APX, and GR activities and their transcript levels; increased AsA accumulation via inducing its biosynthetic genes; but reduced GSH content and NnGSH2 expression level. These results suggest that ethylene biosynthesis and signaling play an important role in N. nucifera response to Cd stress, and maintaining appropriate ethylene level and low ethylene sensitivity could improve its Cd tolerance via efficient antioxidant defenses.
  239. G. F. Wang, T. Tian, J. F. Meng, X. Q. Xiao, and Y. N. Xiao, “First Report of Fusarium Incarnatum Causing Rot Disease on Lotus in China,” Journal of Plant Pathology, vol. 102, no. 2, pp. 595–595, May 2020. doi: 10.1007/s42161-019-00477-2.
  240. Y. Wang, Y. Chen, M. Yuan, Z. Xue, Q. Jin, and Y. Xu, “Flower Color Diversity Revealed by Differential Expression of Flavonoid Biosynthetic Genes in Sacred Lotus,” Journal of the American Society for Horticultural Science, vol. 141, no. 6, pp. 573–582, Nov. 2016. doi: 10.21273/JASHS03848-16.
    Sacred lotus (Nelumbo nucifera) is an important aquatic ornamental plant which contains several diverse flower colors, but the underlying mechanisms of its flower coloration remain unclear. In this study, seven complementary DNA (cDNA) clones of genes involved in flavonoid biosynthesis, including chalcone synthase (CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (F3H), flavonoid 3′-hydroxylase (F3′H), flavonoid 3′,5′-hydroxylase (F3′5′H), dihydroflavonol 4-reductase (DFR), and anthocyanidin synthase (ANS), were isolated and characterized. Moreover, expression patterns of these seven genes and pigment profiles were investigated across four N. nucifera cultivars with different flower colors: Zhongguohongbeijing [ZGH (red)], Xinghuafen [XHF (pink)], Molingqiuse [MLQS (yellow)], and Zhufengcuiying [ZFCY (white)]. Real-time quantitative polymerase chain reaction (qRT-PCR) analysis showed that during flower development, transcripts of early biosynthetic genes (NnCHS, NnCHI, and NnF3H) were abundant at the early stage; noticeably, highest expression of NnCHI in MLQS probably induced abundant anthoxanthin synthesis and displayed yellow. Expression of late biosynthetic genes, especially NnDFR and NnANS, was generally consistent with change patterns of anthocyanins in ZGH and XHF, but NnF3′H was barely detectable in the pink cultivars. Meanwhile, negligible expression of NnDFR and NnANS was detected in MLQS and ZFCY, respectively, which blocked their colored anthocyanin biosynthesis. Spatial expression analysis revealed that most flavonoid biosynthetic genes were highly expressed in floral tissues, rather than leaves. These results suggest that in N. nucifera cultivars with different flower colors, flavonoid biosynthesis is differentially regulated by the expression of these flavonoid biosynthetic genes, among which, NnCHI, NnF3′H, NnDFR, and NnANS are supposed to be critical for pigment accumulation, and therefore, affect different flower coloration.
  241. Y. Wang et al., “The Sacred Lotus Genome Provides Insights into the Evolution of Flowering Plants,” The Plant Journal, vol. 76, no. 4, pp. 557–567, 2013. doi: 10.1111/tpj.12313.
    Sacred lotus (Nelumbo nucifera) is an ornamental plant that is also used for food and medicine. This basal eudicot species is especially important from an evolutionary perspective, as it occupies a critical phylogenetic position in flowering plants. Here we report the draft genome of a wild strain of sacred lotus. The assembled genome is 792 Mb, which is approximately 85–90% of genome size estimates. We annotated 392 Mb of repeat sequences and 36 385 protein-coding genes within the genome. Using these sequence data, we constructed a phylogenetic tree and confirmed the basal location of sacred lotus within eudicots. Importantly, we found evidence for a relatively recent whole-genome duplication event; any indication of the ancient paleo-hexaploid event was, however, absent. Genomic analysis revealed evidence of positive selection within 28 embryo-defective genes and one annexin gene that may be related to the long-term viability of sacred lotus seed. We also identified a significant expansion of starch synthase genes, which probably elevated starch levels within the rhizome of sacred lotus. Sequencing this strain of sacred lotus thus provided important insights into the evolution of flowering plant and revealed genetic mechanisms that influence seed dormancy and starch synthesis.
  242. D. B. Ward, “Nelumbo Lutea, the Correct Name for the American Lotus,” TAXON, vol. 26, no. 2-3, pp. 227–234, 1977. doi: 10.2307/1220557.
    The American lotus has long been known as Nelumbo lutea (Willd.) Pers. An earlier epithet, derived from Nymphaea pentapetala Walt., was once interpreted by Fernald as applying to the American plant, justifying the new name Nelumbo pentapetala (Walt.) Fern.; this name was soon opposed on the ground that it was based on a monstrosity. Although Fernald’s name has not been generally accepted, Sohmer has recently argued that this ground for its rejection is not pertinent. However, a broader consideration of the context in which Walter described his Nymphaea pentapetala, including examination of the other nymphaeoid plants described by him, his probable dependence upon Fraser for part of their collection, and the inadequacy of any present herbarium record, suggests that he may have been describing the Old World sacred lotus, Nelumbo nucifera Gaertn., or a mixture of the sacred lotus and the American lotus, or that his basis, at least in part, was a confused verbal report provided him by Fraser. Nymphaea pentapetala Walt. remains of uncertain application, and nomenclatural stability is best served by the retention of Nelumbo lutea (Willd.) Pers. for the American lotus.
  243. P. S. Williamson and E. L. Schneider, “Nelumbonaceae,” in Flowering Plants · Dicotyledons: Magnoliid, Hamamelid and Caryophyllid Families, K. Kubitzki, J. G. Rohwer, and V. Bittrich, Eds. Berlin, Heidelberg: Springer, 1993, pp. 470–473. doi: 10.1007/978-3-662-02899-5_55.
    Herbaceous aquatic perennial developing horizontal rhizomes and tubers. Roots adventitious, produced at nodes. Phyllomes distributed in sets of three along the rhizome (one foliage leaf, two fleshy cataphylls). Foliage leaves simple, peltate, emergent and floating, producing latex. Petioles of emergent leaves terete, to 2 m in length, bearing prickles. Blades concave, large, 10–100 cm in diameter, orbiculate, entire, bluish green adaxially and remarkably water-repellent. Flowers solitary, perfect, actinomorphic, hypogynous, 10–100 cm across, pink to white or yellowish, elevated above water on terete peduncles up to 2 m in length; floral phyllotaxy spiral; perianth caducous; sepals 2–5; petals ca. 20–30; stamens 200–300, filaments elongate, bearing four introrsely to latrorsely dehiscent anthers, terminated by claw-like, thermogenic appendage. Gynoecium of 2–30 free carpels embedded in the truncate surface of the enlarged obconical, spongy receptacle; each carpel with a distinct, circular stigma with a central canal into the ovary; ovule solitary, ventral-apical, pendulous, anatropous, bitegmic-crassinucellar. Fruits globose or elongate ovoid, with the carpel wall adnate to the testa forming a hard-walled nut, filled by the edible embryo, embedded in an enlarged, dry, sclerified receptacle. Embryo with two thick and fleshy cotyledons surrounding a green plumule, the latter enclosed by a delicate stipule-like sheath. Radicle non-functional. Endosperm minute, helobial, lacking perisperm; seeds exalbuminous.
  244. K. Worarad et al., “Transcriptome Profiling for Pericarp Browning during Long-Term Storage of Intact Lotus Root (Nelumbo Nucifera),” Plant Growth Regulation, vol. 95, no. 2, pp. 207–221, Nov. 2021. doi: 10.1007/s10725-021-00736-2.
    Lotus root (Nelumbo nucifera) is an edible rhizome that the consumption/production has continuously increased as more consumers demand convenient and ready-to-eat foods. However, the processing, storage, and transportation of fresh-cut fruits and vegetables promotes physiological deterioration, such as browning, which leads to a reduction in the value of the product. This study aimed to reduce the browning of lotus root pericarps during long-term storage and long-distance transport, and to clarify the functions of unigenes and browning-associated metabolic pathways using RNA-sequencing techniques. Intact lotus root cv. ‘Kanasumi No.34’ browning decreased after the roots were packed along with an anti-browning solution. Over 200 million short single-end reads were mapped onto the N. nucifera consensus coding sequence set. The significantly differentially expressed genes (DEGs) were identified. Based on the Uniprot, GO, and KEGG databases, secondary metabolism, lipid metabolism, and redox state genes were significantly upregulated in the un-packed and packed with water treatments compared to after harvest sample. Additionally, 16 expected DEGs (e.g., PPO, PAL, POD, CHS, PDCR, and SOD), which are affected by browning development, were differentially regulated in lotus root pericarp. The gene expression data presented in this study will help elucidate the molecular mechanism underlying browning development in intact lotus root during long-term storage. The results may also inform future research on improving the post-harvest shelf life of lotus roots’.
  245. J.-Z. Wu et al., “Evaluation of the Quality of Lotus Seed of Nelumbo Nucifera Gaertn from Outer Space Mutation,” Food Chemistry, vol. 105, no. 2, pp. 540–547, Jan. 2007. doi: 10.1016/j.foodchem.2007.04.011.
    Lotus seeds from outer space mutation, named as No. 36 space lotus seed, have been used for 6 years in Jianou county, Fujian province, China and compared to the native counterpart. The proximate composition (ash, moisture, protein, lipid, alcohol extract, 100-seed weight, carbohydrate) and nutritional components (amino acids, vitamins B1, B2, B6, C, E, phospholipids) of No. 36 space lotus seed and native lotus seed embryos were compared. The results indicate that the profiles of proximate composition and nutritional components of No. 36 space lotus seed and native lotus seed embryos were similar; however, most chemical contents were significantly higher (P<0.05) in the former. This result was also confirmed by the HPLC fingerprint. The quality of No. 36 space lotus seed was better than that of native lotus seed and the results support the use of this seed as a food and a herbal medicine product.
  246. P. Wu, A. Liu, Y. Zhang, K. Feng, S. Zhao, and L. Li, “NnABI4-Mediated ABA Regulation of Starch Biosynthesis in Lotus (Nelumbo Nucifera Gaertn),” International Journal of Molecular Sciences, vol. 22, no. 24, p. 13506, Jan. 2021. doi: 10.3390/ijms222413506.
    Starch is an important component in lotus. ABA is an important plant hormone, which plays a very crucial role in regulating plant starch synthesis. Using ‘MRH’ as experimental materials, the leaves were sprayed with exogenous ABA before the rhizome expansion. The results showed that stomatal conductance and transpiration rate decreased while net photosynthetic rate increased. The total starch content of the underground rhizome of lotus increased significantly. Meanwhile, qPCR results showed that the relative expression levels of NnSS1, NnSBE1 and NnABI4 were all upregulated after ABA treatment. Then, yeast one-hybrid and dual luciferase assay suggested that NnABI4 protein can promote the expression of NnSS1 by directly binding to its promoter. In addition, subcellular localization results showed that NnABI4 encodes a nuclear protein, and NnSS1 protein was located in the chloroplast. Finally, these results indicate that ABA induced the upregulated expression of NnABI4, and NnABI4 promoted the expression of NnSS1 and thus enhanced starch accumulation in lotus rhizomes. This will provide a theoretical basis for studying the molecular mechanism of ABA regulating starch synthesis in plant.
  247. Z. Wu et al., “A Precise Chloroplast Genome of Nelumbo Nucifera (Nelumbonaceae) Evaluated with Sanger, Illumina MiSeq, and PacBio RS II Sequencing Platforms: Insight into the Plastid Evolution of Basal Eudicots,” BMC Plant Biology, vol. 14, no. 1, p. 289, Nov. 2014. doi: 10.1186/s12870-014-0289-0.
    The chloroplast genome is important for plant development and plant evolution. Nelumbo nucifera is one member of relict plants surviving from the late Cretaceous. Recently, a new sequencing platform PacBio RS II, known as ‘SMRT (Single Molecule, Real-Time) sequencing’, has been developed. Using the SMRT sequencing to investigate the chloroplast genome of N. nucifera will help to elucidate the plastid evolution of basal eudicots.
  248. J. Xiao, B. Tian, B. Xie, E. Yang, J. Shi, and Z. Sun, “Supercritical Fluid Extraction and Identification of Isoquinoline Alkaloids from Leaves of Nelumbo Nucifera Gaertn,” European Food Research and Technology, vol. 231, no. 3, pp. 407–414, Jul. 2010. doi: 10.1007/s00217-010-1290-y.
    Isoquinoline alkaloids from leaves of Nelumbo nucifera (N. nucifera) were extracted using supercritical CO2. The effects of the parameters such as the dynamic extraction time, temperature, pressure, various modifiers, and flow rate of the modifier on the yield of nuciferine and the ratio of total isoquinoline alkaloids to the total extract were investigated. Nuciferine content of the extract was determined by high-performance liquid chromatography (HPLC). The results indicated that the yield of nuciferine increased with increases in the dynamic extraction time, pressure, temperature, and flow rate of the modifier. The highest nuciferine yield of 325.54 μg/g was obtained when the extraction was carried out for 2 h at 70 °C under 30 MPa, with 10% (v/v) diethylamine and 1% (v/v) water in methanol as the modifier which kept a flow rate of 1.2 mL/min. The ratio of total isoquinoline alkaloids to the extract was 49.85% at the highest nuciferine yield. Five kinds of isoquinoline alkaloids extracted from N. nucifera leaves were identified by high-performance liquid chromatography combined with ion trap/time-of-flight mass spectrometry (LC/MS-ITTOF). They were Dehydronuciferine, N-nornuciferine, O-nornuciferine, Nuciferine, and Roemerine in the order of retention time.
  249. K. Xie, X. Zhang, L. Yang, and J. Xu, “Breeding of New Varieties Taikong Lotus No.1,No.2,No.3,” ACTA AGRICULTURAE NUCLEATAE SINICA, vol. 18, no. 4, pp. 325–325, 2004. http://caod.oriprobe.com/articles/7635578/Breeding_of_new_varieties_Taikong_Lotus_No_1_No_2_No_3.htm.
  250. K. Xie, X. Zhang, L. Yang, J. Xu, Y. Su, and T. Zhang, “Effects of Ion Implantation on Lotus Seeds from Space Mutation,” ACTA AGRICULTURAE NUCLEATAE SINICA, vol. 18, no. 4, pp. 303–306, 2004. http://caod.oriprobe.com/articles/7635585/EFFECTS_OF_ION_IMPLANTATION_ON_LOTUS_SEEDS_FROM_SPACE_MUTATION.htm.
    3年的试验观察表明,离子注入经太空诱变的白莲种子育出的花型由单瓣变为半重瓣、重瓣花;花色由红色变成粉红、爪红色;莲蓬、莲籽变异出多种形状;始蕾期比对照提前5~21d,末蕾期延迟到9月下旬;单缸现蕾数高达14朵,1m2小区现蕾21朵;单株平均每蓬实粒数最多的为21.6粒,铁莲子最大粒重达2.02g,鲜壳莲子最大粒重达5.0g,上述性状变异是常规育种方法少见的.
  251. K. Xie, L. Yang, X. Zhang, J. Xu, and D. Jiang, “Selection of Lotus from Seeds Re-Onboard in Space Satellite,” ACTA AGRICULTURAE NUCLEATAE SINICA, vol. 18, no. 4, pp. 300–302, 2004. http://caod.oriprobe.com/articles/7635586/SELECTION_OF_LOTUS_FROM_SEEDS_RE_ONBOARD_IN_SPACE_SATELLITE.htm.
    经二次航天搭载后,白莲种子SP1代产生广谱分离,其生物习性、形态特征及经济生状等出现较大变异,其中花型由单瓣变为重瓣或半重瓣,并出现部分种子只长立叶,不开花,部分种子只开花不结蓬、部分种子结蓬后不能正常成熟的现象,也出现种子单缸现蕾9朵,花期始蕾期提前7~22d,种子最大单粒重达1.85g的优良变异,这是采用常规育种很少出现的情况.
  252. J. Xue, L. Zhuo, and S. Zhou, “Genetic Diversity and Geographic Pattern of Wild Lotus (Nelumbo Nucifera) in Heilongjiang Province,” Chinese Science Bulletin, vol. 51, no. 4, pp. 421–432, Feb. 2006. doi: 10.1007/s11434-006-0421-0.
    Based on 47 accessions from Ussuri River Valley, Songhua River Valley and Heilong River Valley together with 2 accessions from Russia and 27 accessions of cultivated lotus from other provinces in China, genetic diversity of wild lotus (Nelumbo nucifera) were revealed using RAPD and ISSR markers. Twenty RAPD primers generated 113 loci, of which 71.68% were polymorphic across all samples. The expected heterozygosity was 0.1583. The percentage of polymorphic loci and expected heterozygosity in the wild lotus were 50.44% and 0.1241, respectively. The parameters of the cultivated lotus were slightly higher, 53.98% and 0.1651 correspondingly. Sixteen ISSR primers produced 90 loci. The percentages of polymorphic loci and expected heterozygosity were 41.11% and 0.0851 at species level, 28.89% and 0.0661 for the wild lotus, and 32.22% and 0.0963 for the cultivated lotus. AMOVA analysis of the wild lotus showed that a small number of variances exist among the 3 river valleys (21.68% for RAPD with Gst=0.1312 and 15.11% for ISSR with Gst=0.1352). The molecular variances of both the wild and the cultivated lotuses came predominantly from within the 3 river valleys and the cultivated samples (73.25% for RAPD and 81.11% for ISSR). Variance components from the wild and the cultivated lotus accounted for 19.17% for RAPD and 13.17% for ISSR, and variations among the valleys and the culta seemed the least important (7.585 for RAPD and 5.725 for ISSR). Neighbor-joining analysis demonstrated that considerable differentiation happened between the wild and the cultivated lotus. The wild lotus at middle reaches of the Songhua River Valley seemed to be the centre of remnants, from which it spread to the Ussuri River Valley and the Heilong River Valley. The very limited genetic diversity suggests that the wild lotus has experienced severe bottleneck effect, founder effect and rebirth effect. Considering its long evolutionary history, scarcity of genetic variations and importance in wetland ecosystems, we appeal to take lawful measures to protect the wild lotus. For conservation purpose, special attention should be paid to the lotus at the middle reaches of the Songhua River Valley.
  253. J.-H. Xue, W.-P. Dong, T. Cheng, and S.-L. Zhou, “Nelumbonaceae: Systematic Position and Species Diversification Revealed by the Complete Chloroplast Genome,” Journal of Systematics and Evolution, vol. 50, no. 6, pp. 477–487, 2012. doi: 10.1111/j.1759-6831.2012.00224.x.
    Nelumbonaceae is a morphologically unique family of angiosperms and was traditionally placed in Nymphaeales; more recently, it was placed in Proteales based on molecular data, or in an order of its own, Nelumbonales. To determine the systematic position of the family and to date the divergence time of the family and the divergence time of its two intercontinentally disjunct species, we sequenced the entire chloroplast genome of Nelumbo lutea and most of the chloroplast genes of N. nucifera. We carried out phylogenetic and molecular dating analyses of the two species and representatives of 47 other plant families, representing the major lineages of angiosperms, using 83 plastid genes. The N. lutea genome was 163 510 bp long, with a total of 130 coding genes and an overall GC content of 38%. No significant structural differences among the genomes of N. lutea, Nymphaea alba, and Platanus occidentalis were observed. The phylogenetic relationships based on the 83 plastid genes revealed a close relationship between Nelumbonaceae and Platanaceae. The divergence times were estimated to be 109 Ma between the two families and 1.5 Ma between the two Nelumbo species. The estimated time was only slightly longer than the age of known Nelumbo fossils, suggesting morphological stasis within Nelumbonaceae. We conclude that Nelumbonaceae holds a position in or close to Proteales. We further conclude that the two species of Nelumbo diverged recently from a common ancestor and do not represent ancient relicts on different continents.
  254. J. Xue, S. Wang, and S.-L. Zhou, “Polymorphic Chloroplast Microsatellite Loci in Nelumbo (Nelumbonaceae),” American Journal of Botany, vol. 99, no. 6, pp. e240–e244, 2012. doi: 10.3732/ajb.1100547.
    • Premise of the study: To study population genetics, phylogeography, and hybridization of Nelumbo (Nelumbonaceae), chloroplast microsatellite markers were developed. • Methods and Results: Seventeen microsatellite loci were identified from the chloroplast genomes of N. nucifera and N. lutea. Polymorphisms were assessed in three populations of N. nucifera and one population of N. lutea. Nine loci were found to be polymorphic in N. nucifera, and all 17 loci were found to be polymorphic in N. lutea. In N. nucifera, the number of alleles per locus ranged from two to six, and the unbiased haploid diversity per locus ranged from 0.198 to 0.790. In N. lutea, the number of alleles ranged from two to four, and the unbiased haploid diversity per locus ranged from 0.245 to 0.694. • Conclusions: The identified chloroplast simple sequence repeat markers will be useful for the study of genetic diversity, phylogeography, and identification of Nelumbo cultivars.
  255. T.-H. Yang and C.-M. Chen, “On the Alkaloids of Nelumbo Nucifera Gaertn. Studies on the Alkaloids of Loti Embryo,” Journal of the Chinese Chemical Society, vol. 17, no. 4, pp. 235–242, 1970. doi: 10.1002/jccs.197000030.
    A new tertiary tetrahydroisoquinoline alkaloid, methylcorypalline (V) and biscoclaurine base, neferine (II) were isolated from a Formosan lotus embryo (embryo of the seed of Nelumbo nucifera Gaertn.) (Nymphaeaceae), besides the earlier reported isoliensinine (I) and lotusine (IV). Characterization of this new base, methylcorypalline (I), mp. 58–59° (n-hexane), , by spectral data and direct comparison with synthetic sample prepared from vanillin have proved to be 6,7-dimethoxy-2-methyl-1,2,3,4-tetrahydroisoquinoline (V). Methylcorypalline (V) is the first instance occurring in nature having coronary dilator action.
  256. M. Yang, Y. Han, L. Xu, J. Zhao, and Y. Liu, “Comparative Analysis of Genetic Diversity of Lotus (Nelumbo) Using SSR and SRAP Markers,” Scientia Horticulturae, vol. 142, pp. 185–195, Jul. 2012. doi: 10.1016/j.scienta.2012.05.021.
    The lotus genus, Nelumbo Adans., comprises two species, N. nucifera Gaertn. and N. lutea (Willd.) Pers. N. nucifera is distributed in Asia and northern Australia, and is cultivated as a commercial crop for the showy flowers, seeds and rhizomes. N. lutea is distributed in the Americas and has been used to create interspecific hybrids. The present study was conducted to assess the genetic diversity and relatedness for accessions of N. nucifera, N. lutea and interspecific hybrids using simple sequence repeat (SSR) and sequence-related amplified polymorphism (SRAP) markers for comparative analysis. Forty-three Nelumbo accessions were genotyped using 38 SSR and 16 SRAP primer pairs. The SSR markers yielded 161 polymorphic loci with an average polymorphism information content (PIC) of 0.575, whereas the 16 SRAP primer pairs generated 119 polymorphic alleles with an average PIC of 0.741. Dendrograms constructed with the UPGMA method from the SSR and SRAP data showed similar clusters with some exceptions. Two major groups were identified: group I included N. nucifera and four hybrids, and revealed complex genetic relationships among flower, seed and rhizome lotus; group II included N. lutea and nine hybrids, and confirmed the interspecific genetic distinction between N. nucifera and N. lutea. The results of UPGMA clustering were consistent with those of the PCA and population structure analysis. Mantel’s test indicated there was good concordance between the SSR and SRAP data (r=0.732), and highly significant correlations between combined SSR–SRAP and separate SSR (r=0.935) and SRAP (r=0.926) data. Overall, both types of molecular marker were shown to be valuable tools for the analysis of genetic relationships, but combination of the SSR and SRAP data was more informative for assessment of genetic diversity of Nelumbo. These findings provide a basis for future use of these molecular markers in the genetic analysis of Nelumbo.
  257. M. Yang, L. Zhu, L. Xu, C. Pan, and Y. Liu, “Comparative Transcriptomic Analysis of the Regulation of Flowering in Temperate and Tropical Lotus (Nelumbo Nucifera) by RNA-Seq,” Annals of Applied Biology, vol. 165, no. 1, pp. 73–95, 2014. doi: 10.1111/aab.12119.
    The switch from vegetative to reproductive growth is a major developmental transition in flowering plants, which depends on the balanced expression of the genes within a complex network that is controlled by both environmental and endogenous factors. Molecular regulation of flower development has been investigated extensively in model plants, particularly Arabidopsis. However, little is known about the mechanisms that regulate flowering in lotus. To analyse the molecular regulation of flowering in lotus, comparative transcript profiling was performed at two stages of bud development – the initial developmental stage (T1) and the fast developing stage (T2) – in the lotus cultivars ‘BG’ (B, temperate lotus) and ‘WR1’ (W, tropical lotus). A total of 140 504 368 high-quality 100-bp reads were obtained and aligned against the lotus reference genome. Of the 23 361 genes assembled, at least 88% of these transcripts were detected in each sample. These genes were significantly enriched in 40 Gene Ontology terms and 236 Kyoto Encyclopedia of Genes and Genomes pathways. Further comparisons of the transcripts in the four libraries revealed that 1808, 1330, 785, 702, 1954, and 2050 genes were differentially expressed between BT1 and BT2, WT1 and WT2, BT1 and WT1, BT2 and WT2, BT1 and WT2 and BT2 and WT1 samples, respectively. Analysis of the four libraries identified 147 lotus flowering-time genes homologous to genes that control flowering-time pathways in other plants. Differential regulation of the COP1, CCA1, LHY, CO-LIKE, VIN3, GAI, and FT genes, which participate in the photoperiod, vernalization and gibberellic acid pathways, suggested that they might control the early flowering of lotus. The extensive transcriptome dataset should provide a foundation for comparative gene expression studies on the regulation of flowering in lotus. Comparative transcriptomic analysis detected several differentially expressed genes and potential candidate genes required for early flowering in lotus. These results provide new insight into the molecular mechanisms that regulate flowering in lotus.
  258. M. Yang et al., “Digital Gene Expression Analysis Provides Insight into the Transcript Profile of the Genes Involved in Aporphine Alkaloid Biosynthesis in Lotus (Nelumbo Nucifera),” Frontiers in Plant Science, vol. 8, p. 80, 2017. doi: 10.3389/fpls.2017.00080.
    The predominant alkaloids in lotus leaves are aporphine alkaloids. These are the most important active components and have many pharmacological properties, but little is known about their biosynthesis. We used digital gene expression (DGE) technology to identify differentially-expressed genes (DEGs) between two lotus cultivars with different alkaloid contents at four leaf development stages. We also predicted potential genes involved in aporphine alkaloid biosynthesis by weighted gene co-expression network analysis (WGCNA). Approximately 335 billion nucleotides were generated; and 94% of which were aligned against the reference genome. Of 22 thousand expressed genes, 19,000 were differentially expressed between the two cultivars at the four stages. Gene Ontology (GO) enrichment analysis revealed that catalytic activity and oxidoreductase activity were enriched significantly in most pairwise comparisons. In Kyoto Encyclopedia of Genes and Genomes (KEGG) analysis, dozens of DEGs were assigned to the categories of biosynthesis of secondary metabolites, isoquinoline alkaloid biosynthesis, and flavonoid biosynthesis. The genes encoding norcoclaurine synthase (NCS), norcoclaurine 6-O-methyltransferase (6OMT), coclaurine N-methyltransferase (CNMT), N-methylcoclaurine 3′-hydroxylase (NMCH), and 3′-hydroxy-N-methylcoclaurine 4′-O-methyltransferase (4′OMT) in the common pathways of benzylisoquinoline alkaloid biosynthesis and the ones encoding corytuberine synthase (CTS) in aporphine alkaloid biosynthetic pathway, which have been characterized in other plants, were identified in lotus. These genes had positive effects on alkaloid content, albeit with phenotypic lag. The WGCNA of DEGs revealed that one network module was associated with the dynamic change of alkaloid content. Eleven genes encoding proteins with methyltransferase, oxidoreductase and CYP450 activities were identified. These were surmised to be genes involved in aporphine alkaloid biosynthesis. This transcriptomic database provides new directions for future studies on clarifying the aporphine alkaloid pathway.
  259. X. Yang et al., “Evolutionarily Conserved Function of the Sacred Lotus (Nelumbo Nucifera Gaertn.) CER2-LIKE Family in Very-Long-Chain Fatty Acid Elongation,” Planta, vol. 248, no. 3, pp. 715–727, Sep. 2018. doi: 10.1007/s00425-018-2934-6.
    Identification of NnCER2 and NnCER2-LIKE from Nelumbo nucifera, which are required for the very-long-chain fatty acid elongation, provides new evidence that CER2 proteins are evolutionarily conserved across the eudicots.
  260. M. Yang et al., “Genetic Linkage Maps for Asian and American Lotus Constructed Using Novel SSR Markers Derived from the Genome of Sequenced Cultivar,” BMC Genomics, vol. 13, no. 1, p. 653, Nov. 2012. doi: 10.1186/1471-2164-13-653.
    The genus Nelumbo Adans. comprises two living species, N. nucifera Gaertan. (Asian lotus) and N. lutea Pers. (American lotus). A genetic linkage map is an essential resource for plant genetic studies and crop improvement but has not been generated for Nelumbo. We aimed to develop genomic simple sequence repeat (SSR) markers from the genome sequence and construct two genetic maps for Nelumbo to assist genome assembly and integration of a genetic map with the genome sequence.
  261. M. Yang, F. Liu, Y. Han, L. Xu, N. Juntawong, and Y. Liu, “Genetic Diversity and Structure in Populations of Nelumbo from America, Thailand and China: Implications for Conservation and Breeding,” Aquatic Botany, vol. 107, pp. 1–7, May 2013. doi: 10.1016/j.aquabot.2013.01.001.
    Lotus (Nelumbo Adans.) is an important ornamental plant and a commercial crop that is distributed widely in Asia and northern Australia. Wild lotus is suffering from habitat loss because of global environmental change and intensive human activities. Conservation of wild lotus requires protection of genetic resources. To assess the genetic diversity of wild lotus worldwide, 83 individuals from populations in America, Thailand, and China were analyzed with simple sequence repeat (SSR) markers. A total of 145 alleles were identified with 36 SSR markers in all accessions. Analysis of molecular variance (AMOVA) showed that the higher genetic diversity was observed among populations (88%) than within populations (12%). A high degree of genetic differentiation was identified among the 11 populations (average FST=0.42). Low genetic diversity was revealed within each population, which possibly resulted from predominant asexual reproduction of this species. Mantel’s test demonstrated that genetic differentiation among populations was congruent with geographical distance (r=0.89, P<0.001), which indicated that isolation-by-distance was responsible for the observed genetic structure of the populations of Nelumbo analyzed. Cluster, principal coordinate and STRUCTURE analyses revealed that the American populations (Nelumbo lutea Pers.) were genetically distinct from those from Thailand and China (Nelumbo nucifera Gaertn.), and confirmed the genetic distinction between N. nucifera and N. lutea. Moreover, the Thai genotypes were separated from the Chinese genotypes, which represented two ecotypes of N. nucifera. Given the high level of genetic differentiation among populations, every population should be considered for conservation. In situ preservation is the preferred strategy to conserve the natural genetic diversity and evolutionary processes of Nelumbo, but should be supplemented by ex situ conservation.
  262. M. Yang, L. Zhu, L. Xu, and Y. Liu, “Population Structure and Association Mapping of Flower-Related Traits in Lotus (Nelumbo Adans.) Accessions,” Scientia Horticulturae, vol. 175, pp. 214–222, Aug. 2014. doi: 10.1016/j.scienta.2014.06.017.
    Association mapping was used to identify candidate molecular markers associated with four flower-related traits-flower color, petal number per flower, flowering time, and flowering duration-in a collection of lotus (Nelumbo Adans.) accessions evaluated over two years. Significant phenotypic variations (P<0.001) were detected for petal number, flowering time, and flowering duration. Use of SSR, SRAP, and AFLP markers identified 423 polymorphic loci in the population. The population structure and principal coordinates analyses based on the three kinds of marker identified three clearly distinct subpopulations, which in general corresponded to Nelumbo nucifera, Nelumbo lutea, and their hybrids. Pairwise linkage disequilibrium (LD) among the 423 markers in the population varied from 0 to 1, with a mean of 0.034. The majority of pairwise LD (96%) were r2<0.2, and less than 1% of LD was r2>0.5, indicating a low level of LD between the markers. Using the general linear model considering population structure, association mapping identified several significant markers (P<0.01) that were correlated with the four flower traits across two years. The contributions of these markers to phenotypic variation ranged from 5% to 16%. Some significant associations were repeated for petal number, flowering time, and flowering duration over the two years, and three markers were associated with both flowering time and flowering duration. Of the three kinds markers used, AFLP was the highest efficient marker in association analysis. Our study suggested that lotus might be amenable to association mapping. Candidate marker loci associated with these traits highlighted regions to be targeted in future association studies, and provided useful information for dissecting genetic variation in traits of interest for lotus improvement.
  263. M. Yang, L. Xu, Y. Liu, and P. Yang, “RNA-Seq Uncovers SNPs and Alternative Splicing Events in Asian Lotus (Nelumbo Nucifera),” PLOS ONE, vol. 10, no. 4, p. e0125702, Apr. 2015. doi: 10.1371/journal.pone.0125702.
    RNA-Seq is an efficient way to comprehensively identify single nucleotide polymorphisms (SNPs) and alternative splicing (AS) events from the expressed genes. In this study, we conducted transcriptome sequencing of four Asian lotus (Nelumbo nucifera) cultivars using Illumina HiSeq2000 platform to identify SNPs and AS events in lotus. A total of 505 million pair-end RNA-Seq reads were generated from four cultivars, of which 86% were mapped to the lotus reference genome. Using the four sets of data together, a total of 357,689 putative SNPs were identified with an average density of one SNP per 2.2 kb. These SNPs were located in 1,253 scaffolds and 15,016 expressed genes. A/G and C/T were the two major types of SNPs in the Asian lotus transcriptome. In parallel, a total of 177,540 AS events were detected in the four cultivars and were distributed in 64% of the expressed genes of lotus. The predominant type of AS events was alternative 5’ first exon, which accounted for 41.2% of all the observed AS events, and exon skipping only accounted for 4.3% of all AS. Gene Ontology analysis was conducted to analyze the function of the genes containing SNPs and AS events. Validation of selected SNPs and AS events revealed that 74% of SNPs and 80% of AS events were reliable, which indicates that RNA-Seq is an efficient approach to uncover gene-associated SNPs and AS events. A large number of SNPs and AS events identified in our study will facilitate further genetic and functional genomics research in lotus.
  264. T.-H. Yang, C.-M. Chen, C.-S. Lu, and C.-L. Liao, “Studies on the Alkaloids of Lotus Receptacle,” Journal of the Chinese Chemical Society, vol. 19, no. 3, pp. 143–147, 1972. doi: 10.1002/jccs.197200020.
    From lotus receptacle (seed pod of Nelumbo nucifera Gaertn., four alkaloids, nuciferine (I), N-nornuciferine (VIII), oxoushinsunine (IX) and N-norarmepavine (X) were isolated. Identification of these bases were carried out by their spectral data and direct comparison with authentic samples. It is interesting that oxoushinsunine (IX), a cytotoxic alkaloid shows tumor inhibitory activity against nasopharynx carcinoma reported recently by D. Warthen et al13). The biscoclaurine (IV and V) and quaternary water soluble base (VI) only occures in the embryo. Comparison their alkaloids distribution in each part of Formosan lotus was listed in the Table 1.
  265. M. Yang et al., “Transcriptomic Analysis of the Regulation of Rhizome Formation in Temperate and Tropical Lotus (Nelumbo Nucifera),” Scientific Reports, vol. 5, no. 1, p. 13059, Aug. 2015. doi: 10.1038/srep13059.
    Rhizome is the storage organ of lotus derived from modified stems. The development of rhizome is a complex process and depends on the balanced expression of the genes that is controlled by environmental and endogenous factors. However, little is known about the mechanism that regulates rhizome girth enlargement. In this study, using RNA-seq, transcriptomic analyses were performed at three rhizome developmental stages—the stolon, middle swelling and later swelling stage —in the cultivars ‘ZO’ (temperate lotus with enlarged rhizome) and ‘RL’ (tropical lotus with stolon). About 348 million high-quality reads were generated and 88.5% of the data were mapped to the reference genome. Of 26783 genes identified, 24069 genes were previously predicted in the reference and 2714 genes were novel transcripts. Moreover, 8821 genes were differentially expressed between the cultivars at the three stages. Functional analysis identified that these genes were significantly enriched in pathways carbohydrate metabolism and plant hormone signal transduction. Twenty-two genes involved in photoperiod pathway, starch metabolism and hormone signal transduction were candidate genes inducing rhizome girth enlargement. Comparative transcriptomic analysis detected several differentially expressed genes and potential candidate genes required for rhizome girth enlargement, which lay a foundation for future studies on molecular mechanisms underlying rhizome formation.
  266. Yanrong Fu et al., “Genetic Diversity of the Wild Asian Lotus (Nelumbo Nucifera) from Northern China,” Horticultural plant journal, vol. 7, no. 5, pp. 488–500, Sep. 2021. doi: 10.1016/j.hpj.2021.04.005.
    Asian lotus (Nelumbo nucifera Gaertn.) is an aquatic plant with ornamental, cultural, economic, and ecological values. China has abundant germplasm resources of Asian lotus. However, in many areas, the wild Asian lotuses have been destroyed and the germplasms is now facing extinction. In addition, the knowledge of the genetic diversity of the wild Asian lotus in China is poor. To identify and protect the germplasms of Chinese wild Asian lotus, eleven genomic-SSR primers, three EST-SSR primers, and three chloroplast DNA primers were used to investigate the genetic diversity among 69 samples of wild Asian lotus from 25 locations in northern China. The genetic diversity of 27 samples of wild Asian lotus from southern China and other countries, the ancient Asian lotus, Asian lotus cultivars from China, and Asian-American hybrids was also compared. The genetic diversity of the wild Asian lotus from northern China was characterized as medium, and the mean values of observed heterozygosity (Hₒ) and expected heterozygosity (Hₑ) were 0.087 and 0.552, respectively. Based on a UPGMA dendrogram and STRUCTURE analysis, the wild Asian lotus samples in northern China were divided into three groups. The wild Asian lotus samples from northern China contained 16 haplotypes. The Nei’s genetic distance between the wild Asian lotus samples from the Songhua River basin and the Liao River basin in northeastern China was relatively small, and these germplasms might be relatively primitive compared to those from other regions. This study provides essential information regarding the genetic diversity of the wild Asian lotus resources in northern China, and provides a basis for further analysis of population-level genetic evolution through high-throughput sequencing.
  267. L. Yao, “Study on the ABA Content and SOD Activity in Ancient Lotus and Modern Lotus Seeds,” undefined, 2000. https://www.semanticscholar.org/paper/Study-on-the-ABA-Content-and-SOD-Activity-in-Lotus-Yao/aec2bf92438afa87eb9b218e8c38e8bd18731abd.
    The mechanism of longevity in the ancient lotus seeds (Taizi lotus seed_aged 580±70 years found at Tai Zhou Wu Village of Beijing City in 1984) is focused on, while ABA content in dry peel (peel and seed coat) of both lots are much higher than those in cotyledons and in embryos respectively. This study focused attention on the mechanism of longevity in the ancient lotus seeds (Taizi lotus seed_aged 580±70 years found at Tai Zhou Wu Village of Beijing City in 1984). The modern one is closely related Harbin Lotus of seed_aged 8 years. The content of abscisic acid (ABA) and activity of superoxide dismutase (SOD) of both lotus seeds were measured. ABA content in dry peel (peel and seed coat)of both lotus are much higher than those in cotyledons and in embryos respectively. Content of ABA in the Tai_Zi Lotus was 26% lower than that in Harbin Lotus. The difference of SOD activities in cotyledons of both lotus at imbibition and seedling stage was little, while the SOD activities in embryos and shoots of ancient lotus at the above two stages were 50% or 27% higher than those in Harbin lotus. Possible Physiological role of those phenomena is discussed.
  268. K. Yasuo and M. Shoko, “Deposition and Phytotoxicity of Fungicides Added Spreaders to the Leaves of East Indian Lotus.,” 徳島県立農業試験場試験研究報告, no. 29, pp. 28–36, 1993. https://jglobal.jst.go.jp/en/detail?JGLOBAL_ID=200902140893606325.
    Article “Deposition and phytotoxicity of fungicides added spreaders to the leaves of east indian lotus.” Detailed information of the J-GLOBAL is a service based on the concept of Linking, Expanding, and Sparking, linking science and technology information which hitherto stood alone to support the generation of ideas. By linking the information entered, we provide opportunities to make unexpected discoveries and obtain knowledge from dissimilar fields from high-quality science and technology information within and outside JST.
  269. Y. Yi, C. K. Lin, and J. S. Diana, “Recycling Pond Mud Nutrients in Integrated Lotus–Fish Culture,” Aquaculture, vol. 212, no. 1, pp. 213–226, Sep. 2002. doi: 10.1016/S0044-8486(02)00022-4.
    An experiment was conducted in nine 200-m2 fertilized earthen ponds at the Asian Institute of Technology, Thailand, during January–September 2000. This experiment was designed to assess the recovery of nutrients from pond mud by lotus (Nelumbo nucifera), to assess pond mud characteristics after lotus–fish co-culture, and to compare fish growth with and without lotus integration. There were three treatments in triplicate: (A) lotus–tilapia co-culture; (B) tilapia alone; (C) lotus alone. Seedlings (0.39±0.09 kg) of Thai lotus variety were transplanted to ponds of the treatments with lotus (treatments A and C) at a density of 25 seedlings pond−1, while sex-reversed all-male Nile tilapia (Oreochromis niloticus) fingerlings (8.6–10.3 g) were stocked at two fish per square meter in ponds of the treatments with tilapia (treatments A and B) when the water depth had been increased to 50 cm due to increasing lotus height. Ponds stocked with tilapia (treatments A and B) were fertilized weekly with urea and triple super phosphate (TSP) at a rate of 4 kg nitrogen (N) and 1 kg phosphorus (P)/ha/day after tilapia stocking. There was no fertilization in ponds of the lotus alone treatment. Lotus co-cultured with tilapia or cultured alone in ponds was able to effectively remove nutrients from old pond mud. Annual nutrient losses from mud in a 1-ha pond was about 2.4 ton N, and 1 ton P, among which about 300 kg N and 43 kg P were incorporated in lotus biomass. There were no significant differences in lotus growth performance between the lotus–tilapia and lotus alone treatments, while Nile tilapia cultured alone grew significantly better than when co-cultured with lotus. The present experiment has demonstrated the effectiveness of nutrient removal from old pond mud by lotus and the feasibility of rotation and co-culture of lotus and Nile tilapia. Both systems can recycle nutrients effectively within ponds and are environmentally friendly culture systems.
  270. M. J. Yoo, P. S. Soltis, and D. E. Soltis, “Expression of Floral MADS‐Box Genes in Two Divergent Water Lilies: Nymphaeales and Nelumbo,” International Journal of Plant Sciences, vol. 171, no. 2, pp. 121–146, Feb. 2010. doi: 10.1086/648986.
    To provide insights into the floral developmental genetics of Nymphaeales (water lilies), we investigated the expression patterns of floral organ identity genes in three genera: Cabomba, Nuphar, and Nymphaea. Additionally, because of the superficial floral similarity between Nymphaea and the early‐diverging eudicot Nelumbo, we conducted the same experiments in the latter taxon. We focused on gene expression associated with (1) perianth differentiation in Nymphaeales, (2) the transition of petaloid staminodes to stamens in Nymphaea, and (3) organ identity in Nymphaea and Nelumbo. In Cabomba, the expression patterns of B‐class gene homologues fit the “sliding boundaries” model, with B‐class gene expression in sepals and petals. In contrast, Nuphar and Nymphaea exhibit broad B‐class gene expression that extends across all floral organs (i.e., “fading borders” model). The gene expression patterns observed suggest that the innermost petals of Nymphaea originated from petaloid staminodes. Also, despite the morphological differences between flowers of Nymphaea and Nelumbo and the phylogenetic distance between these genera, floral gene expression patterns are nearly identical. Last, we infer that either the “out‐of‐male” hypothesis or the “mostly male” hypothesis, both of which specify derivation of floral parts from male structures, might apply to the ancestors of Nymphaeales.
  271. Q. Yu-zhu, “Industry Analysis and Development Strategy on Double-Season Nelumbo Nucifera Gaertn in Liujiang County,” undefined, 2007. https://www.semanticscholar.org/paper/Industry-analysis-and-development-strategy-on-in-Yu-zhu/2d3eb7340f503b7783bb2eaa46047a91e1f96a70.
    The double-season Nelumbo nucifera Gaertn industry is one of the advantaged industries for market with higher economic benefit in Liujiang County and the main restriction factors on development were analyzed. The double-season Nelumbo nucifera Gaertn is one of the advantaged industries for market with higher economic benefit in Liujiang County.In the paper,the advantage of double-season Nelumbo nucifera Gaertn industry and the main restriction factors on development were analyzed.According to the current status,the strategy on further developing the double-season Nelumbo nucifera Gaertn industry in Liujiang was put forward to providing the theory basis on developing the industry and continuously improving the development of Nelumbo nucifera Gaertn industry in Liujiang County.
  272. T. Yuan, Q. Wang, W. Li, C. Guo, T. Zhang, and J. Liu, “Water Quality, Nutrient Budgets and Growth Performance in Yellow Catfish (Pelteobagrus Fulvidraco Richardson) and Lotus (Nelumbo Nucifera Gaertn) Co-Culture Systems,” Aquaculture Research, vol. 50, no. 10, pp. 3050–3059, Oct. 2019. doi: 10.1111/are.14264.
    Abstract An experiment was conducted to determine the growth efficiency of lotus (Nelumbo nucifera Gaertn) on nutrient capture in farming systems of yellow catfish (Pelteobagrus fulvidraco Richardson). Three treatments, a control treatment of N. nucifera alone (T1), a treatment of N. nucifera-P. fulvidraco co-culture (T2) and a treatment of P. fulvidraco alone (T3), were conducted in triplicate. Except pH, conductivity and chemical oxygen demand (CODMn), most of the water quality parameters were significantly lowered by N. nucifera cultivation. Total nitrogen and phosphorus concentrations in the T2 system were 62.2% and 71.6% higher than those in the T1 system respectively; and 31.8% and 59.2% lower than those in the T3 system respectively. The nitrogen and phosphorus release rates were 41.2% and 36.8% in the T2 system, respectively, and 95.6% and 99.1% in the T3 system respectively. 7.31%?21.4% of nitrogen and 3.35%?15.9% of phosphorus were unaccounted. N. nucifera can effectively remove nutrients from water and sediments and may promote the outflow of nutrients. No significant difference in fish or N. nucifera growth performance was observed among the systems. This study indicates that the co-culture of N. nucifera with P. fulvidraco is an optimal culture system that can increase food production and reduce waste discharge.
  273. W. Zhang et al., “Characterization of Flower-Bud Transcriptome and Development of Genic SSR Markers in Asian Lotus (Nelumbo Nucifera Gaertn.),” PLOS ONE, vol. 9, no. 11, p. e112223, Nov. 2014. doi: 10.1371/journal.pone.0112223.
    Background Asian lotus (Nelumbo nucifera Gaertn.) is the national flower of India, Vietnam, and one of the top ten traditional Chinese flowers. Although lotus is highly valued for its ornamental, economic and cultural uses, genomic information, particularly the expressed sequence based (genic) markers is limited. High-throughput transcriptome sequencing provides large amounts of transcriptome data for promoting gene discovery and development of molecular markers. Results In this study, 68,593 unigenes were assembled from 1.34 million 454 GS-FLX sequence reads of a mixed flower-bud cDNA pool derived from three accessions of N. nucifera. A total of 5,226 SSR loci were identified, and 3,059 primer pairs were designed for marker development. Di-nucleotide repeat motifs were the most abundant type identified with a frequency of 65.2%, followed by tri- (31.7%), tetra- (2.1%), penta- (0.5%) and hexa-nucleotide repeats (0.5%). A total of 575 primer pairs were synthesized, of which 514 (89.4%) yielded PCR amplification products. In eight Nelumbo accessions, 109 markers were polymorphic. They were used to genotype a sample of 44 accessions representing diverse wild and cultivated genotypes of Nelumbo. The number of alleles per locus varied from 2 to 9 alleles and the polymorphism information content values ranged from 0.6 to 0.9. We performed genetic diversity analysis using 109 polymorphic markers. A UPGMA dendrogram was constructed based on Jaccard’s similarity coefficients revealing distinct clusters among the 44 accessions. Conclusions Deep transcriptome sequencing of lotus flower buds developed 3,059 genic SSRs, making a significant addition to the existing SSR markers in lotus. Among them, 109 polymorphic markers were successfully validated in 44 accessions of Nelumbo. This comprehensive set of genic SSR markers developed in our study will facilitate analyses of genetic diversity, construction of linkage maps, gene mapping, and marker-assisted selection breeding for lotus.
  274. C.-Y. Zhang and M. Guo, “Comparing Three Different Extraction Techniques on Essential Oil Profiles of Cultivated and Wild Lotus (Nelumbo Nucifera) Flower,” Life, vol. 10, no. 9, p. 209, Sep. 2020. doi: 10.3390/life10090209.
    Essential oil components of Nelumbo nucifera flowers from cultivated and wild lotus samples were analyzed and compared using three different extraction techniques, i.e., headspace extraction (HE), steam distillation (SD) and solvent extraction (SE), coupled with GC-MS. Forty-two peaks in the GC-MS analysis were identified as essential oil components extracted by the three methods from N. nucifera flower. The major essential oil components extracted by SD method were found to be Z,Z-10,12-hexadecadienal and E-14-hexadecenal with relative contents of 16.3% and 16.7%, respectively, which is different from that of SE method, i.e., n-hexadecanoic acid and Z,Z-9,12-octadecadienoic acid accounting for 25.8% and 26.8%, respectively. HE method demonstrated a possibility to be used as an in situ and simplest method for extracting the essential oil components from raw materials. By adding a small amount of glycerinum onto the surface of the air-dried flower sample as a solvent trap in the HE method, the volatility of the essential oil components was found to increase by two times for the first time, which could be further utilized to improve the extraction efficiency and the recovery of the essential oil components from other materials for more applications. In addition, the comparison of essential oil components between cultivated and wild samples showed that they differed only in the chemical contents but not in chemical components. This will be a comprehensive report on the chemical information of the essential oil components of N. nucifera flower and provide guidance for its further exploration as high value-added products in the food and healthcare industries.
  275. D. Zhang et al., “Histological and Cytological Characterization of Anther and Appendage Development in Asian Lotus (Nelumbo Nucifera Gaertn.),” International Journal of Molecular Sciences, vol. 20, no. 5, p. 1015, Jan. 2019. doi: 10.3390/ijms20051015.
    The lotus (Nelumbo Adans.) is a perennial aquatic plant with important value in horticulture, medicine, food, religion, and culture. It is rich in germplasm and more than 2000 cultivars have been cultivated through hybridization and natural selection. Microsporogenesis and male gametogenesis in the anther are important for hybridization in flowering plants. However, little is known about the cytological events, especially related to the stamen, during the reproduction of the lotus. To better understand the mechanism controlling the male reproductive development of the lotus, we investigated the flower structure of the Asian lotus (N. nucifera). The cytological analysis of anther morphogenesis showed both the common and specialized cytological events as well as the formation of mature pollen grains via meiosis and mitosis during lotus anther development. Intriguingly, an anatomical difference in anther appendage structures was observed between the Asian lotus and the American lotus (N. lutea). To facilitate future study on lotus male reproduction, we categorized pollen development into 11 stages according to the characterized cytological events. This discovery expands our knowledge on the pollen and appendage development of the lotus as well as improving the understanding of the species differentiation of N. nucifera and N. lutea.
  276. L. Zhang et al., “The Lotus NnFTIP1 and NnFT1 Regulate Flowering Time in Arabidopsis,” Plant Science, vol. 302, p. 110677, Jan. 2021. doi: 10.1016/j.plantsci.2020.110677.
    In higher plants, floral signals are mainly collected and transduced to FLOWERING LOCUS T (FT) in Arabidopsis and its orthologues. The movement of FT from leaves to the shoot apical meristem (SAM) is partially mediated by FT-INTERACTING PROTEIN1 (FTIP1). Although the functions of OsFTIP1 in rice and DOFTIP1 in orchid in FT transport have also been investigated, the FTIP1 homologue in lotus (Nelumbo nucifera Gaertn.), a type of horticultural plant with high economic and cultural value, has not been isolated, and the mechanism of NnFT1 transport has not been explored. Here, we revealed that NnFTIP1 mediates the transport of NnFT1 in ectopic transgenic lines in Arabidopsis. Overexpression of NnFTIP1 in the ftip1-1 background rescued the late flowering phenotype of ftip1-1, indicating that NnFTIP1 has a conserved function as FTIP1. NnFTIP1 and NnFT1 share similar tissue expression patterns and subcellular localization. NnFTIP1 and NnFT1 interact both in vitro and in vivo. In addition, NnFTIP1 affects NnFT1 transport from leaves to the SAM. Furthermore, we found that NnUOF8, a MYB-like transcription factor, directly regulates the expression of NnFTIP1. Our results suggest that the functions of FTIP1 and FT are conserved during evolution in flowering plants.
  277. Y. Zhang, H. Wu, X. Yu, F. Chen, and J. Wu, “Microscopic Observations of the Lotus Leaf for Explaining the Outstanding Mechanical Properties,” Journal of Bionic Engineering, vol. 9, no. 1, pp. 84–90, Mar. 2012. doi: 10.1016/S1672-6529(11)60100-5.
    The leaf of lotus (Nelumbo nucifera) exhibits exceptional ability to maintain the opening status even under adverse weather conditions, but the mechanism behind this phenomenon is less investigated. In this paper, lotus leaves were investigated using environmental scanning electron microscopy in order to illustrate this mechanism. The macro-observations show that the primary veins are oriented symmetrically from leaf center and then develop into fractal distribution, with net-shaped arrangement of the side veins. Further micro-observations show that all the veins are composed of honeycomb micro-tubes viewed from cross section, the inner of micro-tubes are patterned with extended closed-hexagons from vertical section. Different positions of leaf possess diverse mechanical properties by size variation of diameter and inner hexagons of veins, which is theoretically analyzed by building a regular honeycomb model. Specifically, the central area of lotus tends to be stiffer while its margin be softer. These special distribution and composition of the veins mainly account for the distinct behavior of lotus.
  278. Y. Zhang et al., “Nutritional Composition, Physiological Functions and Processing of Lotus (Nelumbo Nucifera Gaertn.) Seeds: A Review,” Phytochemistry Reviews, vol. 14, no. 3, pp. 321–334, Jun. 2015. doi: 10.1007/s11101-015-9401-9.
    Nelumbo nucifera Gaertn. has a relatively wide geographical distribution and biological diversity; various lotus parts have excellent food and medicinal values. Lotus seeds, which are currently the oldest known plant seeds, contain many functional ingredients. They can be eaten raw or cooked, and are often added to foods as ingredients or supplements. Many naturally occurring ingredients isolated from lotus seeds are certified to be multiple functional compounds, such as polyphenols, protein, polysaccharides. Proteins and carbohydrates are the main nutrients of lotus seeds. Low fat content and good proportion of amino acids confer to lotus seeds unique nutritional values that have attracted increasing attention around the world: multiple studies have assessed the functional components of lotus seeds. The bioactivity of ingredients from lotus seeds in vitro and in vivo include antioxidant activity, hypoglycemic, immunomodulatory, antibacterial, anti-inflammatory, analgesic effects as well as gastrointestinal regulation. Lotus seeds show prospective application in function food area and traditional medicine research. Furthermore, structure–activity relationship of functional compounds from lotus seeds will attracts much more interests in recent years. This work briefly reviews the nutrition composition, physiological functions and processing methods of lotus seeds, describing the impact of the latter on nutrient preservation. In addition, this review addresses the recent progresses made in this area and discusses the potential applications and limitations.
  279. X. Zhao, J. Shen, K. J. Chang, and S. H. Kim, “Comparative Analysis of Antioxidant Activity and Functional Components of the Ethanol Extract of Lotus (Nelumbo Nucifera) from Various Growing Regions,” Journal of Agricultural and Food Chemistry, vol. 62, no. 26, pp. 6227–6235, Jul. 2014. doi: 10.1021/jf501644t.
    The variations in antioxidant activity and concentration of functional components in the ethanol extracts of lotus seeds and rhizomes based on the growing region and dryness were investigated. Free radical scavenging activity, total phenolic and flavonoid content, and concentration of several specific flavonoids and alkaloids in the ethanol extracts of lotus were measured. Antioxidant activity and its correlative total phenolic content varied characteristically depending on the growing region and dryness. High-perfomance liquid chromatography analysis showed that the ethanol extracts of lotus seeds from Vietnam (Ho Chi Minh City), raw rhizomes from Korea (Siheung), and dried rhizomes from Japan (Nigata) had the greatest specific flavonoid content. The ethanol extracts of seeds from China (Hubei), raw rhizomes from Japan (Nigata), and dried rhizomes from Korea (Siheung) had the greatest specific alkaloid content. Astragaline, rutin, isoquercetin, nuciferine, dauricine, isoliensinine, and neferine were identified in lotus rhizomes for the first time in this study.
  280. M. Zhao et al., “Detection of Highly Differentiated Genomic Regions Between Lotus (Nelumbo Nucifera Gaertn.) With Contrasting Plant Architecture and Their Functional Relevance to Plant Architecture,” Frontiers in Plant Science, vol. 9, p. 1219, 2018. doi: 10.3389/fpls.2018.01219.
    The lotus (Nelumbo nucifera Gaertn.) is one of the most economically and ornamentally important perennial aquatic plants. Plant architecture is an important trait for lotus classification, cultivation, breeding, and applications. In this study, traits representing plant architecture were measured in 390 lotus germplasms for 3 years. According to the phenotypic distribution, 21 large architecture (LA) and 22 small architecture (SA) germplasms exhibiting extreme phenotypes were selected as representatives of plant architecture. Microscopy analyses revealed that LA lotuses possessed far more vertical cells and longer cell lengths than SA lotuses, and there was a closer linear relationship between vertical cell number and plant architecture than cell length and plant architecture. Furthermore, based on whole genome re-sequencing data from 10 LA and 10 SA lotus germplasms, fixation index (FST) genome scan identified 11.02 Mb of genomic regions that were highly differentiated between the LA and SA lotus groups. Chi-square test revealed that 17,154 single nucleotide polymorphisms (SNPs) and 1,554 insertions and deletions (InDels) showed distinct allelic distribution between the LA and SA lotus groups within these regions. A total of 126 variants with distinct allelic distribution in the highly differentiated region were predicted to cause amino acid changes in 60 genes. Among the 41 genes with functional annotation, the expression patterns of six genes involved in cell division and cell wall construction were confirmed using quantitative reverse-transcription PCR (qRT-PCR). In addition, 34 plant architecture-associated InDel markers were developed and verified in the remaining 11 LA and 12 SA lotus plant architecture representatives. This study identified promising functional markers and candidates for molecular breeding and will facilitate further elucidation of the genetic mechanisms underlying plant architecture in the lotus.
  281. S. Zhao et al., “The Effect of Nitrogen Fertilizer on Rhizome Quality and Starch Physicochemical Properties in Nelumbo Nucifera,” Agronomy, vol. 12, no. 4, p. 794, Apr. 2022. doi: 10.3390/agronomy12040794.
    Optimal nitrogen (N) supply significantly increases the starch content, components, and yield of Nelumbo nucifera. However, the underlying transcriptional mechanism and starch accumulation under dose-dependent nitrogen fertilizer are poorly understood. In this study, we found that the optimal nitrogen fertilizer (N2, 30 kg/667 m2) was more beneficial to improve the stomatal conductance (Gs), leaf intercellular CO2 concentration (Ci), transpiration rate (Tr), net photosynthetic rates (Pn), chlorophyll content, starch content, and plot yield. What is more, N2-fertilizer treatment induced a higher number of starch granule, AP2 content, and RVA curve peaks. Then, the transcriptomic analyses performed in control (CK) and N2-fertilizer treatment (N2) showed that the expressions of many differentially expressed genes (DEGs) were significantly induced by N2. KEGG and GO enrichment analysis showed that these DEGs were significantly enriched in biosynthesis of secondary metabolites, phenylpropanoid biosynthesis, carbon metabolism, carbon fixation in photosynthetic organisms, plant hormone signal transduction, and starch and sucrose metabolisms, suggesting that nitrogen fertilizer induced alterations of photosynthesis- and starch accumulation-related gene expression profiles. Finally, six photosynthesis-related genes and fourteen starch synthesis-related genes were confirmed to be required for starch accumulation in the Nelumbo nucifera development. qPCR analysis of six starch accumulation-related genes demonstrated the accuracy of the transcriptome. Hence, our study provides valuable resource for future studies on molecular mechanisms underlying starch accumulation in Nelumbo nucifera rhizome under N-fertilizer treatment.
  282. L.-J. Zheng et al., “Antioxidant Activity of Lotus (Nelumbo Nucifera Gaertn.) Receptacles of Eleven Cultivars Grown in China,” Journal of Medicinal Plants Research, vol. 6, no. 10, pp. 1902–1911, Mar. 2012. doi: 10.5897/JMPR11.1373.
    Receptacles of 11 lotus (Nelumbo nuciferaGaertn) cultivars [ShiLiHe 1 (L-1), Taikong 1 (L-2), GanLisn 62 (L-3), DaHongLian (L-4), Taikong 3 (L-5), JianXuan 17 (L-6), CunSanLian (L-7), JianLian (L-8), Taikong 36 (L-9), Zajiaolian 8236 (L-10) and Jianjibaihualian (L-11)] were analyzed for the contents of total phenolic, flavonoid and proanthocyanidin, and tested for antioxidant activity by some different assays. Extracts from L-8, L-9, L-10 and L-11 showed higher total phenolic, flavonoid, proanthocyanidin contents and antioxidant activities, which were measured by the 2,2-diphenyl-1-picrylhydrazyl (DPPH), diammonium salt (ABTS), superoxide anion radical scavenging activities and total antioxidant activity assays, than the other cultivars. Moreover, the total phenolic, flavonoid and proanthocyanidin contents correlates well with the antioxidant activities as measured by the DPPH, ABTS, superoxide anion scavenging activity, reducing power, and total antioxidant assays, but does not correlate with metal chelating activity andβ-carotene bleaching activity. Key words:Lotus (Nelumbo nucifera), phenolic, flavonoid, proanthocyanidin, antioxidant activity.
  283. M. Zhou et al., “Identification and Comparison of Anti-Inflammatory Ingredients from Different Organs of Lotus Nelumbo by UPLC/Q-TOF and PCA Coupled with a NF-κB Reporter Gene Assay,” PLOS ONE, vol. 8, no. 11, p. e81971, Nov. 2013. doi: 10.1371/journal.pone.0081971.
    Lotus nelumbo (LN) (Nelumbo nucifera Gaertn.) is an aquatic crop that is widely distributed throughout Asia and India, and various parts of this plant are edible and medicinal. It is noteworthy that different organs of this plant are used in traditional herbal medicine or folk recipes to cure different diseases and to relieve their corresponding symptoms. The compounds that are contained in each organ, which are named based on their chemical compositions, have led to their respective usages. In this work, a strategy was used to identify the difference ingredients and screen for Nuclear-factor-kappaB (NF-κB) inhibitors with anti-inflammatory ability in LN. Seventeen main difference ingredients were compared and identified from 64 samples of 4 different organs by ultra-performance liquid chromatography that was coupled with quadrupole/time of flight mass spectrometry (UPLC/Q-TOF-MS) with principal component analysis (PCA). A luciferase reporter assay system combined with the UPLC/Q-TOF-MS information was applied to screen biologically active substances. Ten NF-κB inhibitors from Lotus plumule (LP) extracts, most of which were isoquinoline alkaloids or flavone C-glycosides, were screened. Heat map results showed that eight of these compounds were abundant in the LP. In conclusion, the LP extracts were considered to have the best anti-inflammatory ability of the four LN organs, and the chemical material basis (CMB) of this biological activity was successfully validated by multivariate statistical analysis and biological research methods.
  284. Y. Zhou et al., “NnHSP17.5, a Cytosolic Class II Small Heat Shock Protein Gene from Nelumbo Nucifera, Contributes to Seed Germination Vigor and Seedling Thermotolerance in Transgenic Arabidopsis,” Plant Cell Reports, vol. 31, no. 2, pp. 379–389, Feb. 2012. doi: 10.1007/s00299-011-1173-0.
    In plants, small heat shock proteins (sHSPs) are unusually abundant and diverse proteins involved in various abiotic stresses, but their functions in seed vigor remain to be fully explored. In this study, we report the isolation and functional characterization of a sHSP gene, NnHSP17.5, from sacred lotus (Nelumbo nucifera Gaertn.) in seed germination vigor and seedling thermotolerance. Sequence alignment and phylogenetic analysis indicate that NnHSP17.5 is a cytosolic class II sHSP, which was further supported by the cytosolic localization of the NnHSP17.5-YFP fusion protein. NnHSP17.5 was specifically expressed in seeds under normal conditions, and was strongly up-regulated in germinating seeds upon heat and oxidative stresses. Transgenic Arabidopsis seeds ectopically expressing NnHSP17.5 displayed enhanced seed germination vigor and exhibited increased superoxide dismutase activity after accelerated aging treatment. In addition, improved basal thermotolerance was also observed in the transgenic seedlings. Taken together, this work highlights the importance of a plant cytosolic class II sHSP both in seed germination vigor and seedling thermotolerance.
  285. Z.-L. Zhou et al., “Polyphenol Oxidase Activity in Harvest Rhizome of Lotus (Nelumbo Nucifera Gaertn. Ssp. Nucifera) and Its Relationship with Morphological Characteristics,” Scientia Horticulturae, vol. 179, pp. 85–90, Nov. 2014. doi: 10.1016/j.scienta.2014.09.001.
    Rhizome of lotus (Nelumbo nucifera Gaertn. ssp. nucifera) has become the most popular aquatic vegetables in the world. An important problem for its storage and process is easily browning and discoloration, which has been proved to mainly result from polyphenol oxidase (PPO) activity. Harvest rhizomes collected from 56 varieties including wild individuals, landraces and cultivars were investigated for their PPO activities as well as 19 morphological characteristics. Several correlations were found to exist in PPO activity values among different parts of all rhizome knots. Four out of 19 morphological characteristics were associated with PPO activity in different parts or knots. It was observed that the PPO activity was uniformly distributing neither in different parts within a knot nor in all knots. The PPO activity in heart part was lower than that in the periphery except for a special case that the 4th knot of red flower lotus rhizome. The PPO activity in former three knots with less number of stoma on skin of rhizome (SA) was lower than it in the same ones with more SA ones. The PPO activity in red-flower-lotus rhizomes was lower than that in white-flower plants. Our results also provide some clues for lotus rhizome processing. For red flower varieties, those with a few knot number of main stem of rhizome and the former three knots can be selected in priority, while the heart part of former four knots in white flower variety should be preferred. The periphery of former four knots in white flower variety that contained higher PPO activity would be beneficial for pest and disease resistance and for breeding medicinal and functional cultivars related with different phenolic compounds.
  286. H.-huan Zhu, J.-xiang Yang, C.-han Xiao, T.-yu Mao, J. Zhang, and H.-yan Zhang, “Differences in Flavonoid Pathway Metabolites and Transcripts Affect Yellow Petal Colouration in the Aquatic Plant Nelumbo Nucifera,” BMC Plant Biology, vol. 19, no. 1, p. 277, Jun. 2019. doi: 10.1186/s12870-019-1886-8.
    The Asia lotus (Nelumbo nucifera Gaertn.) is an ornamental aquatic plant with high economic value. Flower colour is an important ornamental trait, with much of N. nucifera breeding focusing on its yellow flowers. To explore the yellow flower colouration mechanism in N. nucifera, we analysed its pigment constituents and content, as well as gene expression in the flavonoid pathway, in two N. nucifera cultivars.
  287. Y. Zhu, J. Lu, J. Wang, F. Chen, F. Leng, and H. Li, “Regulation of Thermogenesis in Plants: The Interaction of Alternative Oxidase and Plant Uncoupling Mitochondrial Protein,” Journal of Integrative Plant Biology, vol. 53, no. 1, pp. 7–13, 2011. doi: 10.1111/j.1744-7909.2010.01004.x.
    Thermogenesis is a process of heat production in living organisms. It is rare in plants, but it does occur in some species of angiosperm. The heat is generated via plant mitochondrial respiration. As possible involvement in thermogenesis of mitochondrial factors, alternative oxidases (AOXs) and plant uncoupling mitochondrial proteins (PUMPs) have been well studied. AOXs and PUMPs are ubiquitously present in the inner membrane of plant mitochondria. They serve as two major energy dissipation systems that balance mitochondrial respiration and uncoupled phosphorylation by dissipating the H+ redox energy and proton electrochemical gradient (ΔμH+) as heat, respectively. AOXs and PUMPs exert similar physiological functions during homeothermic heat production in thermogenic plants. AOXs have five isoforms, while PUMPs have six. Both AOXs and PUMPs are encoded by small nuclear multigene families. Multiple isoforms are expressed in different tissues or organs. Extensive studies have been done in the area of thermogenesis in higher plants. In this review, we focus on the involvement and regulation of AOXs and PUMPs in thermogenesis.
  288. Y. Zou et al., “Small RNA and Transcriptome Sequencing Reveals miRNA Regulation of Floral Thermogenesis in Nelumbo Nucifera,” International Journal of Molecular Sciences, vol. 21, no. 9, p. 3324, Jan. 2020. doi: 10.3390/ijms21093324.
    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.