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Papers by Padmanaban Mohanan

Journal of Ginseng Research, 2023

BackgroundThe genus Panax in the Araliaceae family has been used as traditional medicinal plants ... more BackgroundThe genus Panax in the Araliaceae family has been used as traditional medicinal plants worldwide and is known to biosynthesize ginsenosides and phytosterols. However, genetic variation between Panax species has influenced their biosynthetic pathways is not fully understood.MethodsSimultaneous analysis of transcriptomes and metabolomes obtained from adventitious roots of two tetraploid species (Panax ginseng and P. quinquefolius) and two diploid species (P. notoginseng and P. vietnamensis) revealed the diversity of their metabolites and related gene expression profiles.ResultsThe transcriptome analysis showed that 2,3-OXIDOSQUALENE CYCLASEs (OSCs) involved in phytosterol biosynthesis are upregulated in the diploid species, while the expression of OSCs contributing to ginsenoside biosynthesis is higher in the tetraploid species. In agreement with these results, the contents of dammarenediol-type ginsenosides were higher in the tetraploid species relative to the diploid species.ConclusionThese results suggest that a whole-genome duplication event has influenced the triterpene biosynthesis pathway in tetraploid Panax species during their evolution or ecological adaptation. This study provides a basis for further efforts to explore the genetic variation of the Panax genus.

Compendium of plant genomes, 2021

Panax ginseng, a medicinally important perennial herb, has been widely used as the medicinal plan... more Panax ginseng, a medicinally important perennial herb, has been widely used as the medicinal plant in South East Asian countries such as Korea, Japan, and China. Various pharmacological efficacies of ginseng are mainly correlated to its unique triterpenoid saponins ginsenosides. The backbone of ginsenosides are synthesized as triterpene saponins via the isoprenoid pathway, and the various ginsenosides are formed by glycosyl transferases. Ginseng plant has a long life cycle over 4-6 years, and it is frequently exposed to environmental stresses during this long-term cultivation. Therefore, ginseng needs to activate an array of defense mechanisms controlled by defense-related genes to confer the enhanced resistance with minimal fitness cost. Infection, wound, irradiation, and other abiotic stresses induce several defense-related genes. Such responses are caused by salicylates and jasmonates,

Horticulture Environment and Biotechnology, Jul 11, 2019

Panax ginseng and Panax quinquefolius are important herbs with similar benefits and medicinal pro... more Panax ginseng and Panax quinquefolius are important herbs with similar benefits and medicinal properties. Both possess ginsenosides, the pharmacologically active components, abundantly in their roots. In an attempt to establish ginseng cultivars with improved agronomical characters and higher ginsenoside content, reciprocal interspecific hybrids were generated by artificial pollination of emasculated flowers. Three elite cultivars of P. ginseng, types Jakyung, Chungkyung, and Hwangsook, and two new lines, 0837 and 0702, were used to assess interspecific hybrid potential. Among F 1 reciprocal hybrids generated using line 0837 as either the maternal or paternal parent showed excellent agronomical traits and hybrid vigor. They showed strong root phenotypes with many lateral and fine roots resulting in 10 to 20% higher ginsenoside content compared to the parental lines. Among the major ginsenosides, pharmacologically active Re, Rb1, Rb2, and F1 were enriched. They accounted for 70% of the protopanaxadiol ginsenosides in 0837/Pq and Pq/0837 crosses, displaying strong maternal dominance. Thus, based on our results, we conclude that P. ginseng line 0837 is superior compared to already existing lines for performing interspecific hybridization with promising outcomes in root quality and ginsenoside content.

Plant Biotechnology Reports, May 14, 2019

The traditional oriental medicine bokbunja, prepared from immature berries of Rubus coreanus is u... more The traditional oriental medicine bokbunja, prepared from immature berries of Rubus coreanus is used as an anti-oxidant, diuretic, and cure for impotence. The bokbunja wine made from fermented fruits of bokbunja has been used as a functional food as well. However, the usage of bokbunja has been problematic over the years due to the abundance of mistakenly identified berries such as Rubus chingii, Rubus crataegifolius, and Rubus occidentalis. Thus, here we developed a method for the molecular differentiation of Rubus species as well as the authentication bokbunja from other Rubus species. We screened several sequences from the chloroplast DNA of these species and found that the rpl16 region was polymorphic for R. coreanus and R. occidentalis, while the trnG-trnS intergenic spacer region was polymorphic for R. chingii and R. crataegifolius. Species-specific primers were designed and a multiplex PCR was performed by combining the markers at the rpl16 and trnG-trnS regions. Amplicons of 686 bp for R. coreanus and 478 bp for R. occidentalis were produced by the primers 5′ Rcor or 5′ Rocci, respectively, with 3′ rpl16; whereas, amplicons of 389 bp for Rubus crataegifolius and 180 bp for R. chingii were produced by 5′ Rcra or 5′ trnG-trnS, respectively, and 3′ Rcra/Rchi. The deduced molecular markers were utilized to authenticate the bokbunja products and demonstrated that the majority of bokbunja samples from the markets were adulterant berries. Hence, our results indicate that the produced molecular markers can serve as an effective tool to authenticate bokbunja.

Molecular Biology Reports, May 19, 2020

Cytochrome P450 monooxygenase 704B (CYP704B), a member of the CYP86 clan, was found to be needed ... more Cytochrome P450 monooxygenase 704B (CYP704B), a member of the CYP86 clan, was found to be needed in Arabidopsis and rice to biosynthesize precursors of sporopollenin through oxidizing fatty acids. In the present study, we cloned and characterized a CYP704B gene in Panax ginseng, named PgCYP704B1. It shared high sequence identity (98-99%) with CYP704 of Arabidopsis, Theobroma cacao, and Morus notabilis. The phylogenetic comparison of ginseng and higher plants between the members of CYP86 clan revealed that ginseng CYP704 was categorized as a group of CYP704B with dicot plants. The expression of PgCYP704B1 is low in the stem, leaf, and fruit, and high in flower buds, particularly detected in the young gametic cell and tapetum layer of the developing anther. Arabidopsis plants overexpressing PgCYP704B1 improved plant biomass such as plant height, siliques and seed number and size. A cytological observation by transverse and longitudinal semi-thin sections of the siliques cuticles revealed that the cell length increased. Furthermore a chemical analysis showed that PgCYP704B1ox lines increased their cutin monomers contents in the siliques. Our results suggest that PgCYP704B1 has a conserved role during male reproduction for fatty acid biosynthesis and its overexpression increases cutin monomers in siliques that eventually could be used for seed production.

Russian Journal of Genetics, Nov 1, 2017

Panax ginseng Meyer is one of the important medicinal plants in the world, particularly in Asian ... more Panax ginseng Meyer is one of the important medicinal plants in the world, particularly in Asian countries. Ginseng encounters many stress exposure during its long cultivation period. However, the molecular mechanism of stress resistance is still poorly understood in spite of its importance. In this study, pathogenesis-related protein 6 (PR6), also called proteinase inhibitor (PI), was isolated from ginseng embryogenic callus, named PgPR6. The small size of PR6, containing an open reading frame of 219 bp encoding 72 amino acids, the typical characteristic of PR6 protein, shares the highest sequence similarity to PR6 of Theobroma cacao (69% identity). Sequence and structural analysis indicated that PgPR6 belongs to class Kunitz-type PI family. This is the first report pertaining to the identification of PR6 gene from the P. ginseng genome. The high-level expression of PgPR6 was observed in root as revealed by quantitative real-time PCR. The temporal expression analysis demonstrated that PgPR6 expression was highly up-regulated by signaling molecules, heavy metals, mechanical wounding, chilling, salt, sucrose, and mannitol stress, indicating that PgPR6 may play an important role in the molecular defense response of ginseng to a various range of environmental stresses.

Journal of Ginseng Research, 2017

Background: Korean ginseng (Panax ginseng) is a well-known medicinal plant of Oriental medicine t... more Background: Korean ginseng (Panax ginseng) is a well-known medicinal plant of Oriental medicine that is still in practice today. Until now, a total of 11 Korean ginseng cultivars with unique features to Korean ginseng have been developed based on the pure-line-selection method. Among them, a new cultivar namely G-1 with different agricultural traits related to yield and content of ginsenosides, was developed in 2012. Methods: The aim of this study was to distinguish the new ginseng cultivar G-1 by identifying the unique single-nucleotide polymorphism (SNP) at its 45S ribosomal DNA and Panax quinquefolius region than other Korean ginseng cultivars using multiplex amplification-refractory mutation systemepolymerase chain reaction (ARMS-PCR). Results: A SNP at position of 45S ribosomal DNA region between G-1, P. quinquefolius, and the other Korean ginseng cultivars was identified. By designing modified allele-specific primers based on this site, we could specifically identified G-1 and P. quinquefolius via multiplex PCR. The unique primer for the SNP yielded an amplicon of size 449 bp in G-1 cultivar and P. quinquefolius. This study presents an effective method for the genetic identification of the G-1 cultivar and P. quinquefolius. Conclusion: The results from our study shows that this SNP-based approach to identify the G-1 cultivar will be a good way to distinguish accurately the G-1 cultivar and P. quinquefolius from other Korean ginseng cultivars using a SNP at 45S ribosomal DNA region.

Plant Cell Reports, Nov 17, 2017

Your article is protected by copyright and all rights are held exclusively by Springer-Verlag Gmb... more Your article is protected by copyright and all rights are held exclusively by Springer-Verlag GmbH Germany, part of Springer Nature. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com".

Applied Biological Chemistry, Feb 26, 2023

Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Pa... more Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Panax ginseng. Although light quality affects ginsenoside content, little is known about the underlying genetic and regulatory mechanisms. Additionally, the correlation between the adaptability of ginseng to shade and ginsenoside biosynthesis remains poorly understood. In the present study, transcriptome analysis of ginseng seedlings using RNA sequencing revealed that the expression of ginsenoside biosynthesis genes, including PgHMGR, PgFPS, PgSS, and PgUGT , was enhanced in shade conditions but downregulated by red light, indicating that far-red light might play an essential role in ginsenoside production. Further, gene expression analysis in adventitious roots and 2-year-old plants using qRT-PCR showed that the light quality-mediated expression patterns of ginsenoside genes varied with tissue and age. However, unlike the transcriptome, there was no difference in the total ginsenoside content in seedlings among various light conditions. Nevertheless, the amount of major protopanaxadiol-type ginsenosides increased under shade and red light conditions. Unlike seedlings and adventitious roots, there was a decrease in the expression of PgHMGR, PgFPS, PgSS, and PgDDS in 2-year-old plants, along with an increase in the ginsenoside content, under far-red light. Taken together, our findings suggest that far-red light is an important environmental factor for ginsenoside biosynthesis and diversification and provide information that can improve the quality of ginseng produced for medicinal purposes.

WORLD SCIENTIFIC eBooks, Nov 27, 2022

Fitoterapia, Oct 1, 2019

Gynostemma pentaphyllum is a traditional oriental medicinal herb used as tea since ancient time. ... more Gynostemma pentaphyllum is a traditional oriental medicinal herb used as tea since ancient time. Among Gynostemma species, G. pentaphyllum has more active chemical components and better therapeutic effect. It is used to cure depression, diabetes, anxiety, hyperlipidemia, fatigue, immunity, cancer, and oxidative stress. Overexploitation of G. pentaphyllum for its medicinal benefits has been on a rise, due to which they are adulterated or mistakenly identified with other members of Gynostemma species. Hence, we used chloroplast universal regions such as ycf3, accD, petD, psbB and their polymorphism to distinguish G. pentaphyllum from other Gynostemma species. By using the species-specific primers derived from the above regions, we established a multiplex allelespecific PCR for the authentication of G. pentaphyllum from other species. Thus the PCR reaction produced unique amplicons of size 244 bp and 438 bp for G. pentaphyllum amplified by the primers flanking ycf3, and accD regions respectively. While a 607 bp, and 787 bp amplicons from the primers targeting psbB, and petD regions distinguished G. longipes, G. burmanicum, and G. pubescens species. Moreover, these primers were successful to analyze the dried tea samples of Gynostemma as well. Thus, the developed molecular markers could authenticate different Gynostemma species as well as its products thereby preventing the mistaken-identity of this medicinal herb.

Journal of Ginseng Research, Apr 1, 2018

Ginseng has gained its popularity as an adaptogen since ancient days because of its triterpenoid ... more Ginseng has gained its popularity as an adaptogen since ancient days because of its triterpenoid saponins, known as ginsenosides. These triterpenoid saponins are unique and classified as protopanaxatriol and protopanaxadiol saponins based on their glycosylation patterns. They play many protective roles in humans and are under intense research as various groups continue to study their efficacy at the molecular level in various disorders. Ginsenosides Rb1 and Rg1 are the most abundant ginsenosides present in ginseng roots, and they confer the pharmacological properties of the plant, whereas ginsenoside Rg3 is abundantly present in Korean Red Ginseng preparation, which is highly known for its anticancer effects. These ginsenosides have a unique mode of action in modulating various signaling cascades and networks in different tissues. Their effect depends on the bioavailability and the physiological status of the cell. Mostly they amplify the response by stimulating phosphotidylinositol-4,5-bisphosphate 3-kinase/protein kinase B pathway, caspase-3/caspase-9-mediated apoptotic pathway, adenosine monophosphateactivated protein kinase, and nuclear factor kappa-light-chain-enhancer of activated B cells signaling. Furthermore, they trigger receptors such as estrogen receptor, glucocorticoid receptor, and N-methyl-Daspartate receptor. This review critically evaluates the signaling pathways attenuated by ginsenosides Rb1, Rg1, and Rg3 in various tissues with emphasis on cancer, diabetes, cardiovascular diseases, and neurodegenerative disorders.

Plants, Jul 14, 2022

This article is an open access article distributed under the terms and conditions of the Creative... more This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY

Journal of Plant Biology, Jan 21, 2023

Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the ki... more Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the king of oriental medicine, which has been practiced since ancient times in East Asian countries and globally in the modern era. Ginseng is used as an adaptogen, and research shows that it has several pharmacological benefits for various ailments such as cancer, inflammation, diabetes, and neurological symptoms. The pharmacological benefits of ginseng are attributed to the triterpenoid saponin ginsenosides found throughout the Panax ginseng species, which are abundant in its root and are found exclusively in P. ginseng and Panax quinquefolius. Recently, with the completion of the entire ginseng genome sequencing and the construction of the ginseng genome database, it has become possible to access information about many genes newly predicted to be involved in ginsenoside biosynthesis. This review briefly summarizes the current progress in ginseng genome analysis and genes involved in ginsenoside biosynthesis, proposing directions for functional studies of the predicted genes related to ginsenoside production and its regulation.

Scientific Reports, Dec 3, 2019

Root rot caused by Ilyonectria mors-panacis is a devastating fungal disease leading to defect in ... more Root rot caused by Ilyonectria mors-panacis is a devastating fungal disease leading to defect in root quality and causes reduced yield during the perennial life cycle of Panax ginseng Meyer. this indicates the imperative need to understand the molecular basis of disease development and also to enhance tolerance against the fungus. With this idea, the protective effect of silicon (supplied as silica nanoparticles) in P. ginseng root rot pathosystem and its molecular mechanism was investigated in the current study. We have tested different concentrations of silicon (Si) to disease-infected ginseng and found that long term analysis (30 dpi) displayed a striking 50% reduction in disease severity index upon the treatment of Si. Expectedly, Si had no direct degradative effect against the pathogen. Instead, in infected roots it resulted in reduced expression of PgSWEET leading to regulated sugar efflux into apoplast and enhanced tolerance against I. mors-panacis. in addition, under diseased condition, both protopanaxadiol (PPD) and protopanaxatriol (PPT) type ginsenoside profile in roots were higher in Si treated plants. This is the first report indicating the protective role of Si in ginseng-root rot pathosystem, thereby uncovering novel features of ginseng mineral physiology and at the same time, enabling its usage to overcome root rot. Panax ginseng Meyer is an Oriental medicinal adaptogen and ginsenosides are the major pharmacologically active components of ginseng, which is proved to be effective against various diseases 1. It is perennial in nature and the transition from vegetative to reproductive phase occurs at the third year and the accumulation of ginsenosides in roots increases with age 2. Ginseng root rot caused by the fungus, Ilyonectria mors-panacis is one of the devastating diseases which initially infects the root tip and then proceeds until the crown. In addition, replanting results in infection of new plants. Hence, there is an imperative need to design strategies to overcome ginseng root rot. Younger age (~2 years) of the plant, acidic soil (pH 5.5-6.0), soil temperature (18-20 °C), high iron content are the major factors that promote the occurrence of root rot 3,4. Pathogen invades a plant to acquire nutrients which are majorly sugars, to support their growth and replication 5. Understanding the molecular signaling events during plant-pathogen interaction is of great importance to establish strategies to overcome the pathogen. The plant defense system initially detects the pathogen, followed by the activation of the appropriate signal cascades. The downstream defense responses especially the crucial role of hormonal pathways such as SA (Salicylic acid), JA (Jasmonic acid) and Ethylene (ET) mediated pathways are well established. Periodic global transcriptome analyses by RNAseq revealed that JA and ET are majorly activated in ginseng-root rot pathosystem 6. Previously, JA had been demonstrated to influence the triterpenoid pathway in ginseng and PgSE2 was found to influence phytosterol biosynthesis 7,8. Phytosterols are essential component of the plasma membrane that determines its rigidity/fluidity. Certain pathogens have the ability to modify the composition of the phytosterols in the plasma membrane to alter nutrient efflux 9,10 .

Panax ginseng is an important medicinal plant containing many pharmacologically valuable ginsenos... more Panax ginseng is an important medicinal plant containing many pharmacologically valuable ginsenosides, the biosynthesis of which is regulated by complex metabolic pathways. In vitro tissue culture is an alternative way to produce ginseng root biomass and metabolites. The hairy root production via transformation using root inducing plasmid (Ri plasmid) of Agrobacterium rhizogenes would be a useful metabolic engineering tool for P. ginseng. Here, we investigated the transformation efficiency and biomass production in five ginseng genotypes. Of these genotypes, the ‘Yunpoong’ cultivar, and the local landrace ‘Ganghwa’ had the highest transformation efficiencies of 66.11% and 65.00%, respectively. The biomass production of transgenic hairy roots was 1.5–2.1 times faster than that of non-transgenic adventitious roots without hormone supplementation. Various ginsenosides such as Rg1, Rf, Rh1, Rb1, Rb2, Rd, F2, and Rg3, were present at similar or higher levels in the hairy roots compared w...

Journal of Plant Biology

Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the ki... more Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the king of oriental medicine, which has been practiced since ancient times in East Asian countries and globally in the modern era. Ginseng is used as an adaptogen, and research shows that it has several pharmacological benefits for various ailments such as cancer, inflammation, diabetes, and neurological symptoms. The pharmacological benefits of ginseng are attributed to the triterpenoid saponin ginsenosides found throughout the Panax ginseng species, which are abundant in its root and are found exclusively in P. ginseng and Panax quinquefolius. Recently, with the completion of the entire ginseng genome sequencing and the construction of the ginseng genome database, it has become possible to access information about many genes newly predicted to be involved in ginsenoside biosynthesis. This review briefly summarizes the current progress in ginseng genome analysis and genes involved in ginsenoside biosynthesis, proposing directions for functional studies of the predicted genes related to ginsenoside production and its regulation.

Applied Biological Chemistry

Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Pa... more Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Panax ginseng. Although light quality affects ginsenoside content, little is known about the underlying genetic and regulatory mechanisms. Additionally, the correlation between the adaptability of ginseng to shade and ginsenoside biosynthesis remains poorly understood. In the present study, transcriptome analysis of ginseng seedlings using RNA sequencing revealed that the expression of ginsenoside biosynthesis genes, including PgHMGR, PgFPS, PgSS, and PgUGT, was enhanced in shade conditions but downregulated by red light, indicating that far-red light might play an essential role in ginsenoside production. Further, gene expression analysis in adventitious roots and 2-year-old plants using qRT-PCR showed that the light quality-mediated expression patterns of ginsenoside genes varied with tissue and age. However, unlike the transcriptome, there was no difference in the total ginsenoside cont...

WORLD SCIENTIFIC eBooks, Nov 27, 2022

Journal of Ginseng Research

Journal of Ginseng Research, 2023

BackgroundThe genus Panax in the Araliaceae family has been used as traditional medicinal plants ... more BackgroundThe genus Panax in the Araliaceae family has been used as traditional medicinal plants worldwide and is known to biosynthesize ginsenosides and phytosterols. However, genetic variation between Panax species has influenced their biosynthetic pathways is not fully understood.MethodsSimultaneous analysis of transcriptomes and metabolomes obtained from adventitious roots of two tetraploid species (Panax ginseng and P. quinquefolius) and two diploid species (P. notoginseng and P. vietnamensis) revealed the diversity of their metabolites and related gene expression profiles.ResultsThe transcriptome analysis showed that 2,3-OXIDOSQUALENE CYCLASEs (OSCs) involved in phytosterol biosynthesis are upregulated in the diploid species, while the expression of OSCs contributing to ginsenoside biosynthesis is higher in the tetraploid species. In agreement with these results, the contents of dammarenediol-type ginsenosides were higher in the tetraploid species relative to the diploid species.ConclusionThese results suggest that a whole-genome duplication event has influenced the triterpene biosynthesis pathway in tetraploid Panax species during their evolution or ecological adaptation. This study provides a basis for further efforts to explore the genetic variation of the Panax genus.

Compendium of plant genomes, 2021

Panax ginseng, a medicinally important perennial herb, has been widely used as the medicinal plan... more Panax ginseng, a medicinally important perennial herb, has been widely used as the medicinal plant in South East Asian countries such as Korea, Japan, and China. Various pharmacological efficacies of ginseng are mainly correlated to its unique triterpenoid saponins ginsenosides. The backbone of ginsenosides are synthesized as triterpene saponins via the isoprenoid pathway, and the various ginsenosides are formed by glycosyl transferases. Ginseng plant has a long life cycle over 4-6 years, and it is frequently exposed to environmental stresses during this long-term cultivation. Therefore, ginseng needs to activate an array of defense mechanisms controlled by defense-related genes to confer the enhanced resistance with minimal fitness cost. Infection, wound, irradiation, and other abiotic stresses induce several defense-related genes. Such responses are caused by salicylates and jasmonates,

Horticulture Environment and Biotechnology, Jul 11, 2019

Panax ginseng and Panax quinquefolius are important herbs with similar benefits and medicinal pro... more Panax ginseng and Panax quinquefolius are important herbs with similar benefits and medicinal properties. Both possess ginsenosides, the pharmacologically active components, abundantly in their roots. In an attempt to establish ginseng cultivars with improved agronomical characters and higher ginsenoside content, reciprocal interspecific hybrids were generated by artificial pollination of emasculated flowers. Three elite cultivars of P. ginseng, types Jakyung, Chungkyung, and Hwangsook, and two new lines, 0837 and 0702, were used to assess interspecific hybrid potential. Among F 1 reciprocal hybrids generated using line 0837 as either the maternal or paternal parent showed excellent agronomical traits and hybrid vigor. They showed strong root phenotypes with many lateral and fine roots resulting in 10 to 20% higher ginsenoside content compared to the parental lines. Among the major ginsenosides, pharmacologically active Re, Rb1, Rb2, and F1 were enriched. They accounted for 70% of the protopanaxadiol ginsenosides in 0837/Pq and Pq/0837 crosses, displaying strong maternal dominance. Thus, based on our results, we conclude that P. ginseng line 0837 is superior compared to already existing lines for performing interspecific hybridization with promising outcomes in root quality and ginsenoside content.

Plant Biotechnology Reports, May 14, 2019

The traditional oriental medicine bokbunja, prepared from immature berries of Rubus coreanus is u... more The traditional oriental medicine bokbunja, prepared from immature berries of Rubus coreanus is used as an anti-oxidant, diuretic, and cure for impotence. The bokbunja wine made from fermented fruits of bokbunja has been used as a functional food as well. However, the usage of bokbunja has been problematic over the years due to the abundance of mistakenly identified berries such as Rubus chingii, Rubus crataegifolius, and Rubus occidentalis. Thus, here we developed a method for the molecular differentiation of Rubus species as well as the authentication bokbunja from other Rubus species. We screened several sequences from the chloroplast DNA of these species and found that the rpl16 region was polymorphic for R. coreanus and R. occidentalis, while the trnG-trnS intergenic spacer region was polymorphic for R. chingii and R. crataegifolius. Species-specific primers were designed and a multiplex PCR was performed by combining the markers at the rpl16 and trnG-trnS regions. Amplicons of 686 bp for R. coreanus and 478 bp for R. occidentalis were produced by the primers 5′ Rcor or 5′ Rocci, respectively, with 3′ rpl16; whereas, amplicons of 389 bp for Rubus crataegifolius and 180 bp for R. chingii were produced by 5′ Rcra or 5′ trnG-trnS, respectively, and 3′ Rcra/Rchi. The deduced molecular markers were utilized to authenticate the bokbunja products and demonstrated that the majority of bokbunja samples from the markets were adulterant berries. Hence, our results indicate that the produced molecular markers can serve as an effective tool to authenticate bokbunja.

Molecular Biology Reports, May 19, 2020

Cytochrome P450 monooxygenase 704B (CYP704B), a member of the CYP86 clan, was found to be needed ... more Cytochrome P450 monooxygenase 704B (CYP704B), a member of the CYP86 clan, was found to be needed in Arabidopsis and rice to biosynthesize precursors of sporopollenin through oxidizing fatty acids. In the present study, we cloned and characterized a CYP704B gene in Panax ginseng, named PgCYP704B1. It shared high sequence identity (98-99%) with CYP704 of Arabidopsis, Theobroma cacao, and Morus notabilis. The phylogenetic comparison of ginseng and higher plants between the members of CYP86 clan revealed that ginseng CYP704 was categorized as a group of CYP704B with dicot plants. The expression of PgCYP704B1 is low in the stem, leaf, and fruit, and high in flower buds, particularly detected in the young gametic cell and tapetum layer of the developing anther. Arabidopsis plants overexpressing PgCYP704B1 improved plant biomass such as plant height, siliques and seed number and size. A cytological observation by transverse and longitudinal semi-thin sections of the siliques cuticles revealed that the cell length increased. Furthermore a chemical analysis showed that PgCYP704B1ox lines increased their cutin monomers contents in the siliques. Our results suggest that PgCYP704B1 has a conserved role during male reproduction for fatty acid biosynthesis and its overexpression increases cutin monomers in siliques that eventually could be used for seed production.

Russian Journal of Genetics, Nov 1, 2017

Panax ginseng Meyer is one of the important medicinal plants in the world, particularly in Asian ... more Panax ginseng Meyer is one of the important medicinal plants in the world, particularly in Asian countries. Ginseng encounters many stress exposure during its long cultivation period. However, the molecular mechanism of stress resistance is still poorly understood in spite of its importance. In this study, pathogenesis-related protein 6 (PR6), also called proteinase inhibitor (PI), was isolated from ginseng embryogenic callus, named PgPR6. The small size of PR6, containing an open reading frame of 219 bp encoding 72 amino acids, the typical characteristic of PR6 protein, shares the highest sequence similarity to PR6 of Theobroma cacao (69% identity). Sequence and structural analysis indicated that PgPR6 belongs to class Kunitz-type PI family. This is the first report pertaining to the identification of PR6 gene from the P. ginseng genome. The high-level expression of PgPR6 was observed in root as revealed by quantitative real-time PCR. The temporal expression analysis demonstrated that PgPR6 expression was highly up-regulated by signaling molecules, heavy metals, mechanical wounding, chilling, salt, sucrose, and mannitol stress, indicating that PgPR6 may play an important role in the molecular defense response of ginseng to a various range of environmental stresses.

Journal of Ginseng Research, 2017

Background: Korean ginseng (Panax ginseng) is a well-known medicinal plant of Oriental medicine t... more Background: Korean ginseng (Panax ginseng) is a well-known medicinal plant of Oriental medicine that is still in practice today. Until now, a total of 11 Korean ginseng cultivars with unique features to Korean ginseng have been developed based on the pure-line-selection method. Among them, a new cultivar namely G-1 with different agricultural traits related to yield and content of ginsenosides, was developed in 2012. Methods: The aim of this study was to distinguish the new ginseng cultivar G-1 by identifying the unique single-nucleotide polymorphism (SNP) at its 45S ribosomal DNA and Panax quinquefolius region than other Korean ginseng cultivars using multiplex amplification-refractory mutation systemepolymerase chain reaction (ARMS-PCR). Results: A SNP at position of 45S ribosomal DNA region between G-1, P. quinquefolius, and the other Korean ginseng cultivars was identified. By designing modified allele-specific primers based on this site, we could specifically identified G-1 and P. quinquefolius via multiplex PCR. The unique primer for the SNP yielded an amplicon of size 449 bp in G-1 cultivar and P. quinquefolius. This study presents an effective method for the genetic identification of the G-1 cultivar and P. quinquefolius. Conclusion: The results from our study shows that this SNP-based approach to identify the G-1 cultivar will be a good way to distinguish accurately the G-1 cultivar and P. quinquefolius from other Korean ginseng cultivars using a SNP at 45S ribosomal DNA region.

Plant Cell Reports, Nov 17, 2017

Your article is protected by copyright and all rights are held exclusively by Springer-Verlag Gmb... more Your article is protected by copyright and all rights are held exclusively by Springer-Verlag GmbH Germany, part of Springer Nature. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com".

Applied Biological Chemistry, Feb 26, 2023

Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Pa... more Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Panax ginseng. Although light quality affects ginsenoside content, little is known about the underlying genetic and regulatory mechanisms. Additionally, the correlation between the adaptability of ginseng to shade and ginsenoside biosynthesis remains poorly understood. In the present study, transcriptome analysis of ginseng seedlings using RNA sequencing revealed that the expression of ginsenoside biosynthesis genes, including PgHMGR, PgFPS, PgSS, and PgUGT , was enhanced in shade conditions but downregulated by red light, indicating that far-red light might play an essential role in ginsenoside production. Further, gene expression analysis in adventitious roots and 2-year-old plants using qRT-PCR showed that the light quality-mediated expression patterns of ginsenoside genes varied with tissue and age. However, unlike the transcriptome, there was no difference in the total ginsenoside content in seedlings among various light conditions. Nevertheless, the amount of major protopanaxadiol-type ginsenosides increased under shade and red light conditions. Unlike seedlings and adventitious roots, there was a decrease in the expression of PgHMGR, PgFPS, PgSS, and PgDDS in 2-year-old plants, along with an increase in the ginsenoside content, under far-red light. Taken together, our findings suggest that far-red light is an important environmental factor for ginsenoside biosynthesis and diversification and provide information that can improve the quality of ginseng produced for medicinal purposes.

WORLD SCIENTIFIC eBooks, Nov 27, 2022

Fitoterapia, Oct 1, 2019

Gynostemma pentaphyllum is a traditional oriental medicinal herb used as tea since ancient time. ... more Gynostemma pentaphyllum is a traditional oriental medicinal herb used as tea since ancient time. Among Gynostemma species, G. pentaphyllum has more active chemical components and better therapeutic effect. It is used to cure depression, diabetes, anxiety, hyperlipidemia, fatigue, immunity, cancer, and oxidative stress. Overexploitation of G. pentaphyllum for its medicinal benefits has been on a rise, due to which they are adulterated or mistakenly identified with other members of Gynostemma species. Hence, we used chloroplast universal regions such as ycf3, accD, petD, psbB and their polymorphism to distinguish G. pentaphyllum from other Gynostemma species. By using the species-specific primers derived from the above regions, we established a multiplex allelespecific PCR for the authentication of G. pentaphyllum from other species. Thus the PCR reaction produced unique amplicons of size 244 bp and 438 bp for G. pentaphyllum amplified by the primers flanking ycf3, and accD regions respectively. While a 607 bp, and 787 bp amplicons from the primers targeting psbB, and petD regions distinguished G. longipes, G. burmanicum, and G. pubescens species. Moreover, these primers were successful to analyze the dried tea samples of Gynostemma as well. Thus, the developed molecular markers could authenticate different Gynostemma species as well as its products thereby preventing the mistaken-identity of this medicinal herb.

Journal of Ginseng Research, Apr 1, 2018

Ginseng has gained its popularity as an adaptogen since ancient days because of its triterpenoid ... more Ginseng has gained its popularity as an adaptogen since ancient days because of its triterpenoid saponins, known as ginsenosides. These triterpenoid saponins are unique and classified as protopanaxatriol and protopanaxadiol saponins based on their glycosylation patterns. They play many protective roles in humans and are under intense research as various groups continue to study their efficacy at the molecular level in various disorders. Ginsenosides Rb1 and Rg1 are the most abundant ginsenosides present in ginseng roots, and they confer the pharmacological properties of the plant, whereas ginsenoside Rg3 is abundantly present in Korean Red Ginseng preparation, which is highly known for its anticancer effects. These ginsenosides have a unique mode of action in modulating various signaling cascades and networks in different tissues. Their effect depends on the bioavailability and the physiological status of the cell. Mostly they amplify the response by stimulating phosphotidylinositol-4,5-bisphosphate 3-kinase/protein kinase B pathway, caspase-3/caspase-9-mediated apoptotic pathway, adenosine monophosphateactivated protein kinase, and nuclear factor kappa-light-chain-enhancer of activated B cells signaling. Furthermore, they trigger receptors such as estrogen receptor, glucocorticoid receptor, and N-methyl-Daspartate receptor. This review critically evaluates the signaling pathways attenuated by ginsenosides Rb1, Rg1, and Rg3 in various tissues with emphasis on cancer, diabetes, cardiovascular diseases, and neurodegenerative disorders.

Plants, Jul 14, 2022

This article is an open access article distributed under the terms and conditions of the Creative... more This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY

Journal of Plant Biology, Jan 21, 2023

Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the ki... more Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the king of oriental medicine, which has been practiced since ancient times in East Asian countries and globally in the modern era. Ginseng is used as an adaptogen, and research shows that it has several pharmacological benefits for various ailments such as cancer, inflammation, diabetes, and neurological symptoms. The pharmacological benefits of ginseng are attributed to the triterpenoid saponin ginsenosides found throughout the Panax ginseng species, which are abundant in its root and are found exclusively in P. ginseng and Panax quinquefolius. Recently, with the completion of the entire ginseng genome sequencing and the construction of the ginseng genome database, it has become possible to access information about many genes newly predicted to be involved in ginsenoside biosynthesis. This review briefly summarizes the current progress in ginseng genome analysis and genes involved in ginsenoside biosynthesis, proposing directions for functional studies of the predicted genes related to ginsenoside production and its regulation.

Scientific Reports, Dec 3, 2019

Root rot caused by Ilyonectria mors-panacis is a devastating fungal disease leading to defect in ... more Root rot caused by Ilyonectria mors-panacis is a devastating fungal disease leading to defect in root quality and causes reduced yield during the perennial life cycle of Panax ginseng Meyer. this indicates the imperative need to understand the molecular basis of disease development and also to enhance tolerance against the fungus. With this idea, the protective effect of silicon (supplied as silica nanoparticles) in P. ginseng root rot pathosystem and its molecular mechanism was investigated in the current study. We have tested different concentrations of silicon (Si) to disease-infected ginseng and found that long term analysis (30 dpi) displayed a striking 50% reduction in disease severity index upon the treatment of Si. Expectedly, Si had no direct degradative effect against the pathogen. Instead, in infected roots it resulted in reduced expression of PgSWEET leading to regulated sugar efflux into apoplast and enhanced tolerance against I. mors-panacis. in addition, under diseased condition, both protopanaxadiol (PPD) and protopanaxatriol (PPT) type ginsenoside profile in roots were higher in Si treated plants. This is the first report indicating the protective role of Si in ginseng-root rot pathosystem, thereby uncovering novel features of ginseng mineral physiology and at the same time, enabling its usage to overcome root rot. Panax ginseng Meyer is an Oriental medicinal adaptogen and ginsenosides are the major pharmacologically active components of ginseng, which is proved to be effective against various diseases 1. It is perennial in nature and the transition from vegetative to reproductive phase occurs at the third year and the accumulation of ginsenosides in roots increases with age 2. Ginseng root rot caused by the fungus, Ilyonectria mors-panacis is one of the devastating diseases which initially infects the root tip and then proceeds until the crown. In addition, replanting results in infection of new plants. Hence, there is an imperative need to design strategies to overcome ginseng root rot. Younger age (~2 years) of the plant, acidic soil (pH 5.5-6.0), soil temperature (18-20 °C), high iron content are the major factors that promote the occurrence of root rot 3,4. Pathogen invades a plant to acquire nutrients which are majorly sugars, to support their growth and replication 5. Understanding the molecular signaling events during plant-pathogen interaction is of great importance to establish strategies to overcome the pathogen. The plant defense system initially detects the pathogen, followed by the activation of the appropriate signal cascades. The downstream defense responses especially the crucial role of hormonal pathways such as SA (Salicylic acid), JA (Jasmonic acid) and Ethylene (ET) mediated pathways are well established. Periodic global transcriptome analyses by RNAseq revealed that JA and ET are majorly activated in ginseng-root rot pathosystem 6. Previously, JA had been demonstrated to influence the triterpenoid pathway in ginseng and PgSE2 was found to influence phytosterol biosynthesis 7,8. Phytosterols are essential component of the plasma membrane that determines its rigidity/fluidity. Certain pathogens have the ability to modify the composition of the phytosterols in the plasma membrane to alter nutrient efflux 9,10 .

Panax ginseng is an important medicinal plant containing many pharmacologically valuable ginsenos... more Panax ginseng is an important medicinal plant containing many pharmacologically valuable ginsenosides, the biosynthesis of which is regulated by complex metabolic pathways. In vitro tissue culture is an alternative way to produce ginseng root biomass and metabolites. The hairy root production via transformation using root inducing plasmid (Ri plasmid) of Agrobacterium rhizogenes would be a useful metabolic engineering tool for P. ginseng. Here, we investigated the transformation efficiency and biomass production in five ginseng genotypes. Of these genotypes, the ‘Yunpoong’ cultivar, and the local landrace ‘Ganghwa’ had the highest transformation efficiencies of 66.11% and 65.00%, respectively. The biomass production of transgenic hairy roots was 1.5–2.1 times faster than that of non-transgenic adventitious roots without hormone supplementation. Various ginsenosides such as Rg1, Rf, Rh1, Rb1, Rb2, Rd, F2, and Rg3, were present at similar or higher levels in the hairy roots compared w...

Journal of Plant Biology

Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the ki... more Panax ginseng is a medicinal plant belonging to the Araliaceae family. Ginseng is known as the king of oriental medicine, which has been practiced since ancient times in East Asian countries and globally in the modern era. Ginseng is used as an adaptogen, and research shows that it has several pharmacological benefits for various ailments such as cancer, inflammation, diabetes, and neurological symptoms. The pharmacological benefits of ginseng are attributed to the triterpenoid saponin ginsenosides found throughout the Panax ginseng species, which are abundant in its root and are found exclusively in P. ginseng and Panax quinquefolius. Recently, with the completion of the entire ginseng genome sequencing and the construction of the ginseng genome database, it has become possible to access information about many genes newly predicted to be involved in ginsenoside biosynthesis. This review briefly summarizes the current progress in ginseng genome analysis and genes involved in ginsenoside biosynthesis, proposing directions for functional studies of the predicted genes related to ginsenoside production and its regulation.

Applied Biological Chemistry

Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Pa... more Ginsenosides are the most valuable and pharmacologically active triterpenoid saponins found in Panax ginseng. Although light quality affects ginsenoside content, little is known about the underlying genetic and regulatory mechanisms. Additionally, the correlation between the adaptability of ginseng to shade and ginsenoside biosynthesis remains poorly understood. In the present study, transcriptome analysis of ginseng seedlings using RNA sequencing revealed that the expression of ginsenoside biosynthesis genes, including PgHMGR, PgFPS, PgSS, and PgUGT, was enhanced in shade conditions but downregulated by red light, indicating that far-red light might play an essential role in ginsenoside production. Further, gene expression analysis in adventitious roots and 2-year-old plants using qRT-PCR showed that the light quality-mediated expression patterns of ginsenoside genes varied with tissue and age. However, unlike the transcriptome, there was no difference in the total ginsenoside cont...

WORLD SCIENTIFIC eBooks, Nov 27, 2022

Journal of Ginseng Research