, Volume 248, Issue 3, pp 519–544 | Cite as

Recent advances in steroidal saponins biosynthesis and in vitro production

  • Swati Upadhyay
  • Gajendra Singh Jeena
  • Shikha
  • Rakesh Kumar ShuklaEmail author


Main conclusion

Steroidal saponins exhibited numerous pharmacological activities due to the modification of their backbone by different cytochrome P450s (P450) and UDP glycosyltransferases (UGTs). Plant-derived steroidal saponins are not sufficient for utilizing them for commercial purpose so in vitro production of saponin by tissue culture, root culture, embryo culture, etc, is necessary for its large-scale production.

Saponin glycosides are the important class of plant secondary metabolites, which consists of either steroidal or terpenoidal backbone. Due to the existence of a wide range of medicinal properties, saponin glycosides are pharmacologically very important. This review is focused on important medicinal properties of steroidal saponin, its occurrence, and biosynthesis. In addition to this, some recently identified plants containing steroidal saponins in different parts were summarized. The high throughput transcriptome sequencing approach elaborates our understanding related to the secondary metabolic pathway and its regulation even in the absence of adequate genomic information of non-model plants. The aim of this review is to encapsulate the information related to applications of steroidal saponin and its biosynthetic enzymes specially P450s and UGTs that are involved at later stage modifications of saponin backbone. Lastly, we discussed the in vitro production of steroidal saponin as the plant-based production of saponin is time-consuming and yield a limited amount of saponins. A large amount of plant material has been used to increase the production of steroidal saponin by employing in vitro culture technique, which has received a lot of attention in past two decades and provides a way to conserve medicinal plants as well as to escape them for being endangered.


Metabolites P450 UGTs Transcriptome Pathway 


Majority of higher plants and microbes synthesize useful secondary metabolites, which are categorized into alkaloids, terpenoids, polyphenols, polyketides, etc., according to their chemical structures and biosynthetic routes. These compounds protect plants from pathogens, competitors and UV light. They also assist in plant adaptation and most of them have valuable medicinal properties that make them useful in pharmaceutical, agrochemical, flavor and aroma industries. Saponins are important plant secondary metabolites generally known for their multiple pharmacological properties (Williams and Gong 2007).

The word saponin is derived from sapo, i.e., soap, which describes the soapy appearance of saponin when combined with water. Due to their amphiphilic properties and tendency to form the foam, they can be used as surfactant or emulsifying agents (Kime et al. 2015). Structurally, saponins are amphiphilic molecules that are composed of one or more hydrophilic sugar residues and hydrophobic steroidal or triterpenoidal part on the basis of which they are called as steroidal saponin or triterpenoidal saponins. The non-sugar water-insoluble part is called as sapogenin, and on the basis of sugar present in saponin molecules they can be either monodesmosidic (contains only one sugar residue), bidesmosidic (contains two sugar residues) or polydesmosidic (more than two sugar residues) saponins (Lorent et al. 2014). Structural variants of saponin are found in plants due to the presence of different sugar at different position and orientation. The most common sugars residues that are found in saponins are d-glucose, d-galactose, l-rhamnose, l-arabinose, d-glucuronic acid, d-fucose and d-xylose (Lorent et al. 2014).

Saponins are synthesized from 30-carbon linear precursor molecule 2, 3-oxidosqualene, but during the synthesis of steroidal aglycone, there is a loss of three methyl groups that results in the formation of a skeleton having 27 carbon atoms while during synthesis of triterpenoidal aglycone all the 30 carbon atoms retains in its backbone (Moses et al. 2014a, b; Jeena et al. 2017). Steroidal saponins are further classified into two subfamilies, i.e., spirostanol saponins and furostanol saponins. Spirostanol saponins contain a bicyclic spiroacetal moiety at 22nd carbon that involves the steroid E and F rings while furostanol saponins bear a hemiacetal, methyl acetal, or Δ20(22)-unsaturation at this position (Fig. 1) (Challinor and De Voss 2013). Both spirostanol and furostanol saponins have either R or S configuration at 25th carbon position (Masullo et al. 2016). Infrared (IR) spectroscopy has been used to differentiate between spirostanol and furostanol saponins. Spirostanol possesses characteristic absorption peaks at around 980, 920, 900, and 860 cm−1 (Challinor et al. 2012a, b, c). The relative intensities at 920 and 900 cm−1 bands are predictive C-25S and C-25R configuration in spirostanol (Jones et al. 1953) but the detection of the stereochemistry of C-25 in furostanol saponins is more challenging due to the absence of ring configuration as observed in spirostanol saponins. So the conversion of furostanol saponin to the corresponding spirostanol form by either hydrolysis or by enzymatic cleavage of glucose moiety at 26th carbon position for ring closure of its side chain is the most authentic method for prediction of C-25 configuration in furostanol saponin (Inoue et al. 1996). In this review, we will focus on the steroidal saponin-producing plants whose transcriptome sequencing has been done along with we encapsulate the information related to diverse applications of steroidal saponin and its biosynthetic enzymes specifically P450s and UGTs that are involved at later step modification of saponin backbone.
Fig. 1

Structural chemical backbone of steroidal saponins: a furostanol type steroidal saponin b spirostanol type steroidal saponin. Both furostanol and spirostanol type of steroidal saponins are derived from either the 30-carbon linear precursor 2, 3-oxidosqualene (cycloartenol pathway) or 22,26-dihydroxycholesterol (cholesterol pathway) but during the synthesis of their steroidal aglycone loss of three methyl groups results in a 27-carbon backbone. Conversion of furostanol saponin into spirostanol form and vice versa is catalyzed by furostanol glycoside 26-O-β-glucosidase (F26G) and UGT glucosyltransferase enzyme, respectively

The value of steroidal saponins

Steroidal saponins obtained from Dioscorea zingiberensis are widely used for preventing cardiovascular diseases (Qin et al. 2009). In addition to providing cardiovascular protection, steroidal saponin isolated from Ophiopogon japonicus plant also exhibited various other pharmacological activities, such as anticancer, immunomodulation, anti-oxidation, anti-inflammation, cough relief, antimicrobial, and anti-diabetes (Chen et al. 2016). Steroidal saponin constituents from Paris species are used to treat cancer, malignant boil, bleeding and gastritis (Wang et al. 2015a, b, c). They also exhibited antifungal activity (Morrissey and Osbourn 1999) and used as an efficient natural sweetener like glycyrrhizin from licorice roots (Kitagawa 2002). In pharmaceutical industry, they are widely used as raw materials for the production of steroid hormones (Guclu Ustundag and Mazza 2007). Steroidal saponin from Anemarrhena asphodeloides, i.e., timosaponin AIII was found to inhibit the growth of tumor cells and can be considered as a significant compound for the development of the novel anticancerous drug (Wang et al. 2016a, b). Steroidal saponin diosgenin increases bone formation (Folwarczna et al. 2016), executes anti-thrombotic activity by inhibiting the activity of factor VIII and platelet aggregation (Zhang et al. 2013a, b, c), cardioprotective (Jayachandran et al. 2016), suppresses skin inflammation (Kim et al. 2016), have potential to treat liver fibrosis (Xie et al. 2015), useful in the treatment of metabolic disease by regulating cholesterol homeostasis, effective in reversing hyperlipidemia (Fuller and Stephens 2015), anti-inflammatory, improves antioxidant status, inhibits lipid peroxidation (Chen et al. 2015) etc. So this compound can be further used for pharmacotherapy of various diseases.

Spicatoside A from Liriope platyphylla possesses anti-inflammatory activities, anti-asthma activities, anti-osteoclastogenesis activities, memory consolidation activities, neurite outgrowth activities, anti-cancer activities (Ramalingam and Kim 2016) while steroidal saponin from allium species possesses antispasmodic effect, cardioprotective activity, antifungal activity and cytotoxic activity (Sobolewska et al. 2016).

The antiproliferative and cytotoxic activity of steroidal saponins were mainly explored by researchers. It was observed that Paris polyphylla steroidal saponins (PPSS) induce apoptosis and autophagy in human lung cancer cell lines that indicate the anti-cancerous properties of steroidal saponins (He et al. 2015). Additionally, spirostane-type steroidal glycosides from Allium flavum also showed cytotoxicity against a human cancerous cell line (Rezgui et al. 2014). Trillium tschonoskii steroidal saponins have a potential of prevention and treatment of colorectal cancer (Li et al. 2015a, b, c) and they also have the capability to reverse the multidrug resistance and enhance the drug sensitivity in hepatocellular carcinoma (Wang et al. 2013). Steroidal saponin TTB2 from Trillium tschonoskii exerts anticancer effects against Ewing sarcoma cell line through cell cycle arrest at G2/M and S phase (Huang and Zou 2015). Terrestrosin D, PSVII and diosgenin induce apoptosis in prostate, cervical and breast cancer stem cells, respectively (Zhang et al. 2014a, b; Bhuvanalakshmi et al. 2017). Polyphyllin Ι from Paris polyphylla induces cell cycle arrest and ROS dependent autophagy in colorectal cancerous cells.

Anticarcinogenic activity was shown by a variety of steroidal saponins from different plants including Digitalis trojana (Kirmizibekmez et al. 2014), Allium schoenoprasum (Timité et al. 2013), Dioscorea zingiberensis (Tong et al. 2012), Fagonia indica (Waheed et al. 2012), Rhizoma Paridis (Xiao et al. 2012), Solanum violaceum (Yen et al. 2012), Agave sisalana (Chen et al. 2011a, b), Anemarrhena asphodeloides (Kang et al. 2011), Dioscorea bulbifera (Liu et al. 2011), Trigonella foenum-graecum (Kawabata et al. 2011), Paris polyphylla (Zhu et al. 2011), Raphia farinifera (Tapondjou et al. 2015) etc.

Characterized enzymes involved in the biosynthesis of steroidal saponins

Both the cytosolic mevalonate (MVA) pathway and plastidial methylerythritol 4-phosphate (MEP) pathway are involved in the biosynthesis of steroidal saponins. Acetyl Co-A is the precursor of MVA pathway in the cytosol while in the chloroplast, glyceraldehyde-3-phosphate and pyruvate serve as precursor molecules to initiate MEP pathway. Both pathways lead to the formation of dimethylallyl diphosphate (DMAPP) that is used to synthesize squalene via the sequential action of three enzymes, i.e., geranyl diphosphate synthase (GPS), farnesyl diphosphate synthase (FPS) and squalene synthase (SQS). Squalene is then converted into 2, 3-oxidosqualene under the catalysis of squalene epoxidase (SE). The cyclization of 2, 3-oxidosqualene is then carried out by different oxidosqualene cyclases leading to the products that undergo cyclization, hydroxylation and glycosylation reactions ultimately yielding different types of steroidal saponin glycosides.

Some of the enzymes related to saponin biosynthesis have been characterized and most are yet to be characterized. The exact biosynthetic pathway of saponin biosynthesis is not yet established and now it has become an interesting area of research to find out each enzyme of this biosynthetic pathway and signaling molecules involved in the regulation of saponin biosynthesis. Some characterized enzyme of this pathway in different plant species is reviewed and summarized below.

Enzymes of MVA pathway

Enzymes of MVA pathway was characterized in different plants as 3-hydroxy-3-methylglutaryl CoA reductase (HMGR) is the crucial regulatory and rate-limiting enzyme of mevalonate pathway of saponin biosynthesis that catalyzes the irreversible conversion of HMGR to mevalonate. In addition to steroidal saponin, HMGR is also involved in the biosynthesis of phytosterols and sitosterols. By increasing HMGR gene expression, diosgenin content significantly increased in in vitro cultures of Dioscorea zingiberensis (Diarra et al. 2013). Similarly in Medicago truncatula, silencing of Mkb1 (E3 ubiquitin ligase), that control the activity of HMGR results in accumulation of monoglycosylated saponins (Moses et al. 2014a, b). This enzyme was characterized in different plants including Hevea brasiliensis (Chye et al. 1991), potato (Bhattacharyya et al. 1995), wheat (Aoyagi et al. 1993), Camptotheca acuminate (Maldonado-Mendoza et al. 1997), mulberry (Jain et al. 2000), rice (Ha et al. 2001), Capsicum annuum (Ha et al. 2003), Ginkgo biloba (Shen et al. 2006), Corylus avellana (Wang et al. 2007), Euphorbia pekinensi (Cao et al. 2010), Salvia miltiorrhiza (Dai et al. 2011), coffee (Tiski et al. 2011), Glycyrrhiza uralensis (Rong et al. 2011), Withania somnifera (Akhtar et al. 2013), Dendrobium officinale (Zhang et al. 2014a, b), Paris fargesii (Liang et al. 2014), Centella asiatica (Kalita et al. 2015), Arabidopsis (Suzuki et al. 2004), Cyanotis arachnoidea (Wang et al. 2014a, b, c) and Panax ginseng (Kim et al. 2014).

Mevalonate kinase (MK) catalyzes a step in the isoprenoid biosynthetic pathway, which ultimately leads to huge number of compounds that play important roles in plant growth and development (Tang and Newton 2006). This enzyme was characterized in Catharanthus roseus (Schulte et al. 2000) and Bacopa monniera (Kumari et al. 2015). Mevalonate diphosphate decarboxylase enzyme requires ATP and Mg+2 to convert mevalonate 5-diphosphate (MVAPP) into isopentenyl diphosphate (IPP), has been characterized in Bacopa monniera (Abbassi et al. 2015). Isopentenyl diphosphate isomerase (IPP isomerase) is an important enzyme of isoprenoid biosynthesis that converts IPP into DMAPP has been characterized in Tripterygium wilfordii (Tong et al. 2015), Taxus media (Shen et al. 2015) and Camptotheca acuminate (Pan et al. 2008).

Another important enzyme of isoprenoid biosynthesis is FPS that forms farnesyl diphosphate by a head to tail condensation of two molecules of IPP and one molecule of DMAPP (Fig. 2a). FPS can increase terpenoid accumulation in plants as suggested by gene expression analysis (Zhao et al. 2015). Gene expression studies have demonstrated that FPS expression is positively related to the isoprene content in plants. This gene has been characterized in many plants including Bacopa monniera (Vishwakarma et al. 2012), Tripterygium wilfordii (Zhao et al. 2015) and Arabidopsis thaliana (Keim et al. 2012).
Fig. 2

Steroidal saponin biosynthetic pathway: a overview of steroidal saponin biosynthesis in plants that involve both cytosolic mevalonate (MVA) and plastidial methylerythritol 4-phosphate (MEP) pathways. b This figure mainly represents known P450 s, UGTs and glucosidases involved in hydroxylation, oxidation, glycosylation and final cyclization of saponin backbone. Enzyme and species names are italicized. Dashed arrows imply multiple steps in the pathway. DHCR24 delta 24-sterol reductase, DHCR7 7-dehydrocholesterol reductase, CyP51 sterol 14α-demethylase

Enzymes of methylerythritol 4-phosphate (MEP) pathway

Enzymes of MEP pathway was also characterized in different plants like 1-deoxy-d-xylulose 5-phosphate synthase (DXS) enzyme that was the first enzyme of MEP pathway in plastid that catalyzes the formation of 1-deoxy-d-xylulose 5-phosphate (DXP) by condensation of C1 aldehyde group of d-glyceraldehyde 3-phosphate (GAP) and pyruvate. Intramolecular rearrangement and reduction of DXP were catalyzed by DXP reductoisomerase (DXR) that results in the formation of 2-C-methyl-d-erythritol 4-phosphate (MEP). MEP is further converted into 2C-methyl-d-erythritol 2, 4-cyclodiphosphate (MEC) by the activity of MEC synthase enzyme. DXS was characterized in Aquilaria sinensis (Xu et al. 2014), Maize (Cordoba et al. 2011), DXR enzyme was characterized in Ginkgo biloba (Gong et al. 2005), 2C-methyl-d-erythritol 2,4-cyclodiphosphate (MEC) synthase enzyme of MEP pathway was characterized in Taxus media (Jin et al. 2006) and Ginkgo biloba (Kim et al. 2006). In the last step of MEP pathway both IPP and DMAPP is formed and they also interconvertible in another form or isomerized by the activity of IPP isomerase (Fig. 2a). IPP isomerase was characterized in Taxus media (Shen et al. 2015), Tripterygium wilfordii (Tong et al. 2016), Gossypium barbadense (Wang et al. 2009a, b), Arabidopsis thaliana (Phillips et al. 2008), Camptotheca acuminate (Pan et al. 2008), Cinchona robusta (Ramos-Valdivia et al. 1997) etc.

Both phytosterol and saponins are synthesized from the same precursors that are squalene and 2, 3-oxidosqualene (Haralampidis et al. 2002). SQS is a membrane-bound enzyme that catalyzes the initial enzymatic reaction in the biosynthesis of triterpenoids like plant sterols from MVA isoprenoid pathway (Abe et al. 1993). It catalyzes the condensation of two FPP molecules into squalene. SQS was characterized in a wide variety of plant species including Panax ginseng (Kim et al. 2011), Chlorophytum borivilianum (Kalra et al. 2013), Ornithogalum caudatum (Liu et al. 2017), Panax notoginseng (Jiang et al. 2017), Siraitia grosvenorii (Zhao et al. 2017), Salvia miltiorrhiza (Rong et al. 2016) and Magnolia officinalis (Zha et al. 2016).

SE is another rate-limiting enzyme that catalyzes the epoxidation of the double bond of squalene to produce 2,3 oxidosqualene in saponin biosynthesis. This is the first oxidation step in phytosterol and saponin biosynthesis in plants. It was shown that the expression of SE in Eleutherococcus senticosus was affected by endophytic fungi (Xing et al. 2012). SE is one of the important enzymes for biosynthesis of saponins in vitro using synthetic biology approach. Major work related to this enzyme was performed in Panax species that is supposed to be an important source of triterpenoidal saponin. Molecular cloning, expression profiling and characterization of SE were done in Panax notoginseng and it was observed that it is highly expressed in root tissues. Methyl jasmonate (MeJA) treatment increases the expression level of SE in Panax ginseng (Choi et al. 2005). Chitosan induces saponin biosynthesis by increasing the expression level of squalene epoxidase, but it was found that the activity of chitosan and the expression level of SE were inhibited by chitosan inhibitor PD98059.

Two isoforms of SE were identified in Panax ginseng, one is involved in ginsenoside biosynthesis and another one is involved in phytosterol synthesis (Han et al. 2010). Besides Panax species and yeast, SE was also characterized in Euphorbia tirucalli (Uchida et al. 2007), Withania somnifera (Razdan et al. 2013), Arabidopsis (Laranjeira et al. 2015), Gynostemma pentaphyllum (Guo et al. 2016) etc.

Cycloartenol synthase (CAS) is another membrane-bound enzyme as SQS and SE that is involved in phytosterol biosynthesis in a plant system. In arabidopsis, CAS1 is reported to be involved in the biogenesis of plastid (Babiychuk et al. 2008). CAS was also characterized in Withania somnifera (Dhar et al. 2014), Nicotiana (Gas-Pascual et al. 2014), Siraitia grosvenorii (Zhao et al. 2017) etc. In addition to the above-mentioned enzymes, many P450 and glucosyltransferases that were involved in the biosynthesis of steroidal saponin is discussed below.

Later step modification enzymes (P450 and UGT)

Intermediates of the phytosterol pathway are involved in saponin biosynthesis and primarily enzymes that belong to the multigene families of oxidosqualene cyclases (OSCs), P450 s and family 1 of UGTs are supposed to be involved in saponin biosynthesis (Augustin et al. 2011). Both P450s and UGTs are the member of plant supergene families. In plants, during the biosynthesis of diverse plant secondary metabolites like, terpenoids, fatty acids, lignins, hormones, sterols, pigments and defense-related phytoalexins, P450s catalyzes the oxidative reactions (Schuler 1996). P450s and UGTs are considered as a key enzyme, which leads to diversification of natural products in plants. The diverse characteristics of these enzymes make their identification in saponin pathway difficult. Arabidopsis thaliana contains around 246 P450 genes, and it is really difficult to analyze the function of each P450 gene using reverse genetics approach (Nelson 2006). Since these modifications are necessary for making the saponins soluble and active. Researchers are making large-scale experiments and attempts to isolate P450 s and UGTs involved in saponin biosynthetic pathway in plants.

Plant P450 can be divided into two groups that are mainly A-type and non A-type P450. A-type P450 are mainly involved in secondary metabolites biosynthesis while non A-type P450 is a highly diverse group that includes P450 related to lipid or hormone metabolism (Paquette et al. 2000). Animal, microbial and plant P450 differ at their protein level and show sequence homology of 30% or less than 30%. Beside this CyP51 family members are the most conserved P450 among phyla and share sequence homology more than 30–40% (Bak et al. 2011). In the eukaryotic system, members of CyP51 family catalyze 14α-demethylation of sterols.

As reported earlier that cycloartenol is an important intermediate between 2,3 oxidosqualene and obtusifoliol in photosynthetic eukaryotic organisms while in non-photosynthetic eukaryotic organism cycloartenol is not present and lanosterol is derived directly from oxidosqualene (Fabris et al. 2014). Obtusifoliol 14α demethylase gene was characterized from Sorghum bicolor (L.) and showed strict substrate specificity towards obtusifoliol, which belongs to P450 gene family CyP51 (Bak et al. 1997). Wheat sterol 14α demethylase gene (CyP51) was found to complement ERG11 (lanosterol 14-alpha-demethylase) disrupted engineered yeast strain (Cabello-Hurtado et al. 1999). Steroidal saponins are mainly synthesized from cholesterol by oxygenations at C-16, C-22 and C-26 positions (Ohnishi et al. 2009). P450s that were identified in a transcriptome analysis of medicinal plants and supposed to be involved in steroidal saponin biosynthesis is summarized in Fig. 2b.

Glycosylation of saponins is supposed to be the last step in the biosynthesis of saponins. Glycosylation also regulates the biological activities and solubility of these naturally derived saponins (Sawai and Saito 2011). The identification and characterization of these glycosyltransferase enzymes that catalyze the transfer of sugar moiety to these steroidal and terpenoidal backbones will certainly help us to understand the mechanism of the diversity of these saponins and also control their biological activities in plants. Despite of their importance only few glucosyltransferases are identified and characterized. We have compiled some of the recently identified and characterized UGTs along with their function (Fig. 2b).

Presence and absence of sugar residue at specific position allow saponin glycoside to achieve their different biological activity. For example presence of C-26 sugar in avenacosides A and B from oat plant inhibited their antimicrobial activity but the removal of sugar residue from same carbon position required to gain antimicrobial activity in this plant (Osbourn 1996). Similarly, 26 desgluco derivatives of tobacco furostanol saponin B is required to maintain its hemolytic and antifungal activity while its structural homolog had no such kind of activity (Gruenweller et al. 1990). Number and position of sugar moiety attached with saponin molecule also influence its biological activity as incorporation of α-l-rhamnosyl at C-2 of the glucosyl unit of diosgenin leads to a disaccharide saponin with potent cytotoxicity but when l-rhamnose is attached to C-3 or C-4 of d-glucose, the resulting diosgenyl saponins do not exhibit significant cytotoxic activity against HL-60 cells (Perez-Labrada et al. 2012).

Biosynthesis of steroidal saponin can be best studied in solanum species. Wide ranges of steroidal saponins are synthesized in this genus. It was reported that SaGT4A glucosylate steroidal saponins as well as steroidal alkaloids in Solanum aculeatissimum (Kohara et al. 2005) while StSGT a glycosyltransferase from Solanum tuberosum share 75% sequence homology with SaGT4A only glycosylate steroidal alkaloids (Moehs et al. 1997). It was reported that fructose was added to furostan saponins at the position of C6-OH of 26-O-β-d-glucopyranosyl by β-fructosidase enzyme of a microbial strain of Arthrobacter nitroguajacolicus. This kind of fructosylation is possible only with furostan saponins but not with spirostan saponins due to the stereo-hindrance effect (Liu et al. 2015a, b).

Generally, spirostanol saponins are water-insoluble but the attachment of α-l-rhamnopyranosyl group with the glucopyranosyl moiety at the C2-OH of the C3-O-glycoside; increase the solubility of saponin as well as its absorption by the human body. Toruzyme 3.0 L, a kind of cyclodextrin glucanotransferase was used to synthesize steroidal saponins with novel sugar chains to increase their solubility as well as their activity. Nine steroidal saponins with different aglycones and sugar chains were used as substrates to be glucosylated by this cyclodextrin glucanotransferase (Wang et al. 2010). Cantalasaponin I, a spirostanoside from Agave sisalanta was used as substrates to be glucosylated by toruzyme 3.0 L (CGT), and five new glucosylated products were isolated and identified (Kang et al. 2012a, b, c). Beside glycosylation, deglycosylation of steroidal saponin to sapogenin was reported to be carried out by rumen bacteria, especially the cell-associated enzymes fraction of ruminal content (Wang and McAllister 2010). UGTs, as well as glucosidases that were identified in a transcriptome analysis of steroidal saponin-producing plants, were summarized in Fig. 2b.

Newly identified steroidal saponins in last decade

Generally, steroidal saponins were reported in monocot families like Dioscoreaceae, Amaryllidaceae, Smilacaceae, Liliaceae, Agavaceae, Alliaceae, Asparagaceae, Bromeliaceae, Palmae and Scrophulariaceae, etc, and they are accumulated abundantly in crop plants such as yam, alliums, asparagus, fenugreek, yucca and ginseng (Hostettmann and Marston 2005). The presence of steroidal saponin was also reported in certain dicot families like Apocynaceae, Leguminosae, Solanaceae etc. (Moses et al. 2014a, b). Steroidal saponins from different angiosperms were extracted and their structure was elucidated by spectral analysis. Modern spectrometric methods used for the structural analysis of saponins are mass spectrometry (MS), nuclear magnetic resonancy (NMR) and IR. In different organs of the plant there is a considerable variation in the quantity of saponin. Saponin content depends on age, cultivar and geographical distribution of plant (Vincken et al. 2007). Most commonly steroidal saponins are isolated from underground parts of the plant especially roots and rhizomes, and less often from above ground tissues such as leaves, stems, seeds, and fruits (Challinor and De Voss 2013). Cereals and grasses are generally deficient in saponins, with some notable exceptions such as avena species (oats), which accumulate both triterpenoid and steroidal saponins (Osbourn 2003). It is known that over 90 plant families contain steroidal saponins and many new occurrences are still being reported (Hostettmann and Marston 1995). Some recently identified steroidal saponin with their plant part is summarized in Table 1.
Table 1

List of some recently identified, steroidal saponin-producing plants

S. No.

Steroidal saponin


Plant part



Sarsasapogenin M, Sarsasapogenin N

Asparagus officinalis


Huang and Kong (2006)


Ornithosaponins A–D

Ornithogalum thyrsoides


Kuroda et al. (2006)


Three spirostanol and one furostanol saponin

Calamus insignis


Ohtsuki et al. (2006)


Chekiangensosides A and B

Cynanchum chekiangense


Li et al. (2006)


Neosibiricosides A–D

Polygonatum sibiricum


Ahn et al. (2006)


Two furostane type, two cholestane and two spirostane type

Tribulus alatus Del

Aerial parts

Temraz et al. (2006)


Spirostane type steroidal saponin

Agave shrevei


Pereira et al. (2004)


Two steroidal saponins

Allium ursinum L

Underground parts

Sobolewska et al. (2006)


Two steroidal saponins

Capsicum frutescens

Whole Plant

De Lucca et al. (2006)


Timosaponins AI, AII, AIII, and AIV, and timosaponins BI, and BII

Anemarrhena asphodeloides


Nian et al. (2006)


Three steroidal glycosides

Furcraea selloa


El-Sayed et al. (2006)



Solanum lyratum

Whole plant

Sun et al. (2006)


Two new spirostanol saponins

Smilax medica


Sautour et al. (2006)


Racemosides A, B and C

Asparagus racemosus


Mandal et al. (2006)


Atropurosides A–G

Smilacina atropurpurea


Zhang et al. (2006)


Taccaoside C and D

Tacca plantaginea

Whole plant

Liu et al. (2006)


Timosaponin N, E1, E2, O and Purpureagitosid

Anemarrhena asphodeloides


Kang et al. (2006)


Ophiopogonin E

Ophiopogon japonicus


Cheng et al. (2006)


Solanigrosides C–H

Solanum nigrum

Whole plant

Zhou et al. (2006)


A pair of diastereoisomeric steroidal saponins

Tupistra chinensis


Zou et al. (2006)


Steroidal saponins 1, 2, 3 and 4

Smilax china

Whole Plant

Shao et al. (2007)


Ypsilandroside A, B, isoypsilandroside A,B and isoypsilandrogenin

Ypsilandra thibetica

Whole plant

Xie et al. (2006)


Polypunctosides A–D

Polygonatum punctatum


Yang and Yang (2006)


Dioscin, protodioscin and protogracillin

Dioscorea septemloba Thunb


Tan et al. (2006)



Veronica turrilliana

Aerial parts

Kostadinova et al. (2007)


Six new steroidal saponins

Asparagus acutifolius


Sautour et al. (2007)



Yucca smalliana


Jin et al. (2007)


Cynanversicoside A, B, D, G, glaucoside C and glaucogenin C-3-O-beta-d-thevetopyranoside

Radix Cynanchi Atrati

Whole plant

Liang et al. (2007)


Two new steroidal saponins

Dracaena ombet


Moharram and EI-Shenawy (2007)


Filiasparosides A–D

Asparagus filicinus


Zhou et al. (2007)


A pair of stereoisomeric spirostanol saponins and a new cholestane saponin

Paris polyphylla


Zhao et al. (2007)


Elephanoside A, elephanosides B–F, Ys-II, Ys-IV

Yucca elephantipes


Zhang et al. (2008)


Pseudoprotodioscin, methyl protodioscin and dioscin

Dioscorea nipponica Makino


Lin et al. (2007)


Methyl parvifloside, methyl protodeltonin, zingiberensis saponin I, deltonin

Dioscorea villosa


Hayes et al. (2007)


Paridiformoside B

Lysimachia Paridiformis

Whole plant

Xu et al. (2007a, b)


Shatavarins VI–X

Asparagus racemosus


Hayes et al. (2008)


Aspafiliosides E and F

Asparagus filicinus


Zhou and Chen (2008)


Balanitin-6 and balanitin-7

Balanites aegyptiaca


Gnoula et al. (2008)


Ilwensisaponin A and C

Verbascum pterocalycinum var. mutense


Akkol et al. (2007)


Zingiberenin G

Dioscorea zingiberensis


Xu et al. (2007a, 2007b)


Lyconosides Ia, Ib, II, III, and IV

Solanum lycocarpum St. Hil.


Nakamura et al. (2008)


Five steroidal saponins

Allium leucanthum C. Koch


Mskhiladze et al. (2008a, b)


Cesdiurins I–III

Cestrum diurnum L


Fouad et al. (2008)


Ophiofurospiside A

Ophiopogon japonicus (Thunb.) Ker-Gawl


Xu et al. (2008)


Mannioside A

Dracaena mannii

Stem bark

Tapondjou et al. (2008)


Six C-21 steroidal glycosides

Cynanchum auriculatum Royle ex Wight

Root tuber

Peng et al. (2008)


Paris saponin I, Paris saponin V, Paris saponin VI, Paris saponin VII, Paris saponin H

Paris polyphylla Smith var stenophylla Franch

Whole plant

Yin et al. (2008)


Stemucronatoside K

Stephanotis mucronata

Whole plant

Ye et al. (2008)


Three new spirostanol glycosides, a new furostanol glycoside, and a new cholestane glycoside

Clintonia udensis


Matsuo et al. (2008)


Three steroidal saponins

Paris polyphylla Smith


Deng et al. (2008)


Two new steroidal saponin along with curillin G, asparagoside E, asparoside A, asparoside B

Smilax aspera subsp. mauritanica


Belhouchet et al. (2008)


Smilacinoside A, B, C, D and funkioside D, aspidistrin

Smilacina atropurpurea

Dried tender aerial parts

Yang et al. (2009)


Solanolactosides A, B, Torvosides M, N

Solanum torvum

Aerial parts

Lu et al. (2009)


Two new steroidal saponins

Tribulus terrestris


Su et al. (2009)


Polygonoides A and B

Polygonatum sibiricum


Xu et al. (2009a, b, c)


Five new steroidal saponins

Tribulus terrestris


Su et al. (2009)


Eleven steroidal saponins

Trillium erectum


Hayes et al. (2009)


Two new furostanol saponins and one new spirostanol saponin

Paris polyphylla


Zhao et al. (2009)


Seven spirostanol type saponins

Allium leucanthum


Mskhiladze et al. (2008a, 2008b)


Two new steroidal saponins

Allium macrostemon bunge

Dried bulbs

Chen et al. (2009)



Chlorophytum nimonii (Grah) Dalz

Aerial part

Lakshmi et al. (2009)


Three steroidal saponins

Polygonatum odoratum


Wang et al. (2009a, 2009b)


Three new furostanol saponins together with known protodioscin, pseudoprotodioscin and dioscin

Smilax excelsa


Ivanova et al. (2009)


15-hydroxy-pseudoprotodioscin and 15-methoxy-pseudoprotodioscin

Smilax china


Huang et al. (2009)


Padelaosides A and B

Paris delavayi


Zhang et al. (2009a, b)


Ceparosides C and D

Allium cepa L


Yuan et al. (2009)


Five spirostanol saponins, Three furostanol saponins

Agave utahensis

Whole plant

Yokosuka and Mimaki (2009)


Zingiberenin F

Dioscorea zingiberensis


Xu et al. (2009a, 2009b, 2009c)


Ypsilandrosides C–G

Ypsilandra thibetica

Whole plant

Xie et al. (2009)


Eight new spirostanol saponins and three new furostanol saponins

Agave utahensis

Whole plants

Yokosuka et al. (2009)


Shatavaroside A and shatavaroside B together with filiasparoside C

Asparagus racemosus


Sharma et al. (2009)


Yamogenin II

Asparagus officinalis L

Dried Stems

Sun et al. (2010)


Spirostanol saponin, chantrieroside A

Tacca chantrieri


Zhang et al. (2009a, b)


Three new steroidal saponins

Tribulus terrestris L


Liu et al. (2010a, b)


Two new steroidal saponins

Tribulus terrestris L


Liu et al. (2010a, 2010b)


Two steroidal saponin

Hosta sieboldiana

Leaf and Leafstalk

Yada et al. (2010)


Monodesmosidic spirostanol saponi

Agave macroacantha Zucc


Eskander et al. (2010)


Orchidastrosides A–F, and chloromaloside D

Chlorophytum orchidastrum


Acharya et al. (2010)


Timosaponins AIII, BIII, and D

Anemarrhena asphodeloides


Lee et al. (2010)


Tribufurosides D and E

Tribulus terrestris L


Xu et al. (2009a, 2009b, 2009c)


Three new steroid saponin

Tacca integrifolia


Shwe et al. (2010)


Parvifloside, protodeltonin, protodioscin, protogracillin, zingiberensis saponin, deltonin, dioscin and trillin

Dioscorea zingiberensis C.H. Wright


Li et al. (2010)


Two new furostanol saponins

Lilium longiflorum Thunb


Munafo et al. (2010)


Angudracanosides A–F

Dracaena angustifolia

Fresh stems

Xu et al. (2010)


Esculeoside A

Solanum lycopersicum

Ripe Fruit

Nohara et al. (2010)


Filiasparosides E, F, filiasparoside G, asparagusin A filiasparoside A, filiasparoside B, aspafilioside A, aspafilioside B, and filiasparoside C

Asparagus filicinus


Wu et al. (2010)


Fistulosaponins A–F

Allium fistulosum


Lai et al. (2010)


Deistelianosides A, B and arboreasaponins A and B

Dracaena deisteliana and Dracaena arborea

Stem and bark

Kougan et al. (2010)


Ypsilandrosides H–L and a known saponin polyphylloside III

Ypsilandra thibetica

Whole plant

Lu et al. (2010)


Two furostanol saponins

Ruscus ponticus


Napolitano et al. (2010)


Two steroidal saponins

Solanum surattense Burm. f.

Aerial parts

Lu et al. (2011)


Two new steroidal saponins

Agave sisalana


Chen et al. (2011a, 2011b)


Diosbulbisides D and E

Dioscorea bulbifera


Liu et al. (2011)


Lirigramosides A (1) and B

Liriope graminifolia (Linn.)

Underground parts

Wang et al. (2011a, b)


Five steroidal glycosides, including three spirostane, one furostane and one cholestane type saponins

Yucca gloriosa L


Skhirtladze et al. (2011)


Helosides A and B

Chamaelirium luteum


Challinor et al. (2011)


A new steroidal saponin

Ophiopogon japonicus

Dried roots

Qu et al. (2011)


Polyphyllin D

Paris polyphylla


Chan et al. (2011)


Thirteen steroidal saponins

Beaucarnea recurvata


Eskander et al. (2011)


Esculeoside B-5

Solanum lycopersicum

Ripe fruit

Ohno et al. (2011)



Liriope muscari (Decne.) Baily


Ma et al. (2011)


A bidesmosidic steroidal saponins

Yucca schidigera


Kowalczyk et al. (2011)


A new aglycone of amplexicogenin B

Cynanchum amplexicaule

Whole plant

Chen et al. (2011a, 2011b)


Pallidiflosides A, B and C

Fritillaria pallidiflora

Dry bulbs

Shen et al. (2011)


Desmettianosides A and B

Yucca desmettiana


Diab et al. (2012)


Parvifloside, methyl protodeltonin, trigofoenoside A-1, zingiberensis saponin I, deltonin, dioscin and prosapogenin A of dioscin

Dioscorea villosa


Yoon et al. (2012)


Seven spirostane and furostane-type glycosides

Cestrum ruizteranianum


Galarraga et al. (2011)


Timosaponin J, timosaponin K, (25S)-karatavioside C, timosaponin L, and (25S)-officinalisnin-I

Anemarrhena asphodeloides


Kang et al. (2012a, 2012b, 2012c)


Two new furostanol saponins sarsaparilloside B and sarsaparilloside C

Smilax ornata Lem.


Challinor et al. (2012a, 2012b, 2012c)


A new steroidal saponin

Allium ampeloprasum


Adão et al. (2012)


Lycioside A and lycioside B

Lycium barbarum


Wang et al. (2011a, 2011b)


Parisyunnanosides G–I, one new C(21) steroidal glycoside, parisyunnanoside J

Paris polyphylla


Kang et al. (2012a, 2012b, 2012c)


Two new C-21 steroidal glycosides

Cynanchum amplexicaule

Whole plant

Chen et al. (2012)


Dioscins E and F

Dioscorea nipponica


Zhang et al. (2012a, b)


Ophiopogonins H–O

Ophiopogon japonicus


Zhang et al. (2012a, 2012b)


Japonicoside A, japonicoside B and japonicoside C

Smilacina japonica

Dried rhizomes and roots

Liu et al. (2012a, b)


Pallidifloside D, pallidifloside E, pallidifloside G, pallidifloside H and pallidifloside I

Fritillaria pallidiflora Schrenk

Dry bulbs

Shen et al. (2012)


Tupisteroide A–C

Tupistra chinensis


Liu et al. (2012a, 2012b)


Two new steroidal saponins

Tribulus terrestris


Chen et al. (2013)


Pariposides A–D

Paris polyphylla var. yunnanensis


Wu et al. (2012a, b)


15 steroidal saponins

Chamaelirium luteum (false unicorn)


Challinor et al. (2012a, 2012b, 2012c)


Chonglouosides SL-1-SL-6

Paris polyphylla

Stems and Leaves

Qin et al. (2012)


A novel steroidal saponin

Fagonia indica

Aerial Parts

Waheed et al. (2012)


Eleven steroidal saponins

Paris polyphylla


Wu et al. (2012a, 2012b)


Two new furostanol saponins 1 and 2

Ruscus aculeatus L

Underground parts

De Marino et al. (2012)


Stauntosides C–K

Cynanchum stauntonii


Yu et al. (2013)


Ophiopogonin P–S

Ophiopogon japonicus

Tuberous roots

Li et al. (2013)


Two new steroidal saponins

Smilax microphylla

Roots and Rhizome

Lin et al. (2012)


A new isospirostanol-type steroidal saponin

Smilax scobinicaulis

Roots and Rhizome

Zhang et al. (2013a, 2013b, 2013c)


Shatavaroside C

Asparagus racemosus


Sharma et al. (2012)


Avenacosides A and B

Avena sativa L


Pecio et al. (2012)


Seven new steroidal saponins

Lilium brownii var. viridulum


Hong et al. (2012)


Tupichinin A

Tupistra chinensis


Pan et al. (2012)


Pumilum A

Lilium pumilum DC


Zhou et al. (2012)


Three new steroidal saponins

Dracaena marginata


Rezgui et al. (2013)


Nolinospiroside F

Ophiopogon japonicus


Sun et al. (2013)


Three new steroidal compounds

Hosta longipes


Kim et al. (2013)


Four spirostane-type glycosides

Allium schoenoprasum

Whole plant

Timité et al. (2013)


Seven steroidal glycosides

Solanum torvum


Colmenares et al. (2013)


Torvpregnanosides A and B, two pregnane glycosides, and torvoside Q, a 23-keto-spirostanol glycoside

Solanum torvum

Aerial parts

Lee et al. (2013)


Solanolactoside C

Solanum torvum Swartz

Aerial part

Shu et al. (2013)


Anemarnoside A and anemarnoside B

Anemarrhena asphodeloides

Whole plant

Liu et al. (2013)


Three furostanol type saponins

Digitalis trojana

Aerial parts

Kirmizibekmez et al. (2014)


Two new furostanol saponins

Ophiopogon japonicus


Guo et al. (2013)


Two new steroidal saponins

Selaginella uncinata (Desv.)

Whole plant

Zheng et al. (2013)


Magueyosides A–E

Agave offoyana


Pérez et al. (2013)


Three novel cholestane-type steroidal glycosides and two novel spirostane-type steroidal saponins

Polygonatum odoratum


Bai et al. (2014)


Three new steroidal saponins

Ophiopogon japonicus (Thunb.) Ker-Gawl


Ye et al. (2013)


Dioscoreanosides A–K

Dioscorea bulbifera var. sativa


Tapondjou et al. (2013)


Drangustosides A–B

Dracaena angustifolia Roxb

Whole plant

Huang et al. (2013)


Longipetalosides A–C

Tribulus longipetalus

Whole plant

Naveed et al. (2014)


Cambodianosides A–F

Dracaena cambodiana

Whole plant

Shen et al. (2014)


Five oleanane-type saponins

Ganophyllum giganteum

Root bark

Montes et al. (2014)


Three new spirostane-type glycosides

Allium flavum

Whole plant

Rezgui et al. (2014)


Five steroidal saponins

Agave offoyana


Pérez et al. (2014)


Diospreussinosides A–C

Dioscorea preussii


Tabopda et al. (2014)


Riparoside B and timosaponin J

Smilax riparia

Roots and Rhizomes

Wu et al. (2014a, b)


Zingiberenosides A and B

Dioscorea zingiberensis


Zheng et al. (2014)


Two new furostanol saponins

Asparagus cochinchinensis


Zhu et al. (2014)


Cynanosides A–O

Cynanchum atratum


Yan et al. (2014)


Diospreussinosides A–C

Dioscorea preussii


Tabopda et al. (2014)


Six pennogenyl saponins

Paris quadrifolia L


Gajdus et al. (2014)


Seven steroidal saponins

Anemarrhena asphodeloides


Guo et al. (2015)


Polygodosides A–F

Polygonatum odoratum

Fibrous roots

Zhang et al. (2014a, 2014b)



Ophitopogin japonicas

Whole plant

Zeng et al. (2015)


Smilaxchinoside A and smilaxchinoside C

Smilax riparia

Roots and Rhizomes

Wu et al. (2014a, 2014b)


Sixteen steroidal saponin

Tribulus terrestris

Whole plant

Kang et al. (2014)


Timosaponin X and timosaponin Y

Anemarrhena asphodeloides


Yuan et al. (2014)


A new steroidal saponin

Antigonon leptopus

Whole plant

Apaya and Chichioco-Hernandez (2014)


Spicatoside A and spicatoside D

Liriope plathyphylla

Whole Plant

Choi et al. (2015)


Ypsilandroside S and ypsilandroside T

Ypsilandra thibetica

Whole Plant

Si et al. (2014)


A new C21 steroidal saponin

Azadirachta indica


Liu et al. (2014)


Four new spirostane steroidal saponins

Bletilla striata


Wang and Meng (2015)


Parisverticosides A–C

Paris verticillata

Aerial parts

Sun et al. (2014)


Periplocoside P

Periplocae Cortex

Whole plant

Liu et al. (2015a, 2015b)


Two new furostanol saponins and a new spirostanol saponin

Tupistra chinensis

Roots and Rhizomes

Li et al. (2015a, 2015b, 2015c)


Spirostane-type saponins

Dracaena fragrans

Bark, Roots and leaves

Rezgui et al. (2015)


Two new steroidal saponins

Solanum paniculatum L

Aerial parts

Vieira Júnior et al. (2015)


Cambodianoside G

Dracaena cambodiana


Luo et al. (2015)


Polyhydroxy hellebosaponins

Helleborus niger L


Duckstein et al. (2014)


Four new furostanol glycosides

Hosta plantaginea (Lam.) Aschers


Li et al. (2015a, 2015b, 2015c)


Trigoneoside XIIIa, parvifloside, trigoneoside IVa, deltoside, protobioside, lilioglycoside k, zingiberensis newsaponin I, deltonin, prosapogenin A of dioscin, and trillin

Dioscorea zingiberensis


Wang et al. (2014a, 2014b, 2014c)


Eight steroidal saponins

Raphia farinifera

Mesocarp of the fruits

Tapondjou et al. (2015)


Three new steroidal saponins

Helleborus thibetanus

Dried roots and rhizomes

Zhang et al. (2016a, b)


Two new steroidal saponins

Avena sativa

Oat Bran

Yang et al. (2016)


Typaspidosides B–L

Aspidistra typica


Cui et al. (2016)


Chlorodeistelianosides A–D

Chlorophytum deistelianum

Aerial parts

Tabopda et al. (2016)


Esculeosides B-1 and B-2

Solanum lycopersicum

Tomato juice

Nohara et al. (2015)


Pratioside G and H

Polygonatum prattii


Zhang et al. (2016a, 2016b)


Govanoside A

Trillium govanianum


Shafiq-ur-Rahman et al. (2015)


Chonglouosides SL-9-SL-20

Paris polyphylla var. yunnanensis

Stems and leaves

Qin et al. (2016)



Trillium tschonoskii Maxim

Whole plant

Huang and Zou (2015)


Timosaponin B-II

Anemarrhena asphodeloides

Whole plant

Yuan et al. (2015)


Cynawilfosides A–I

Cynanchum wilfordii


Li et al. (2016)


Two new steroidal saponins

Cestrum laevigatum L


Ribeiro et al. (2016)


Taccavietnamosides A–E

Tacca vietnamensis


Yen et al. (2016)


Timosaponin AIII

Anemarrhena asphodeloides Bge


Wang et al. (2016a, 2016b)


Methyl protodioscin

Dioscorea nipponica


Chung et al. (2016)


Padelaosides C–F

Paris delavayi


Liu et al. (2016a, 2016b)


Terrestrinin J–T

Tribulus terrestris

Whole plant

Wang et al. (2016a, 2016b)

Transcriptome analysis of medicinal plants that produce steroidal saponins

The advent of high throughput next-generation sequencing (NGS) has made the transcriptomic analysis of plant possible with increasing speed and affordability. Transcriptome analysis of medicinal plants whose genomic information is not available in public databases helps in the identification of prospective candidate genes involved in the biosynthesis of the secondary metabolic pathway and this will further increase our understanding related to biosynthesis, regulation, and diversity of secondary metabolites. Some of the medicinal plants that were explored to understand the steroidal saponin biosynthesis are discussed below and summarized in Fig. 3.
Fig. 3

Representation of steroidal saponin-producing plant with transcriptome data: diagrammatic representation of steroidal saponin-producing non-model plants with their plant part whose comparative or non-comparative transcriptome sequencing has been performed using NGS platform. Purple star indicate more steroidal saponin accumulation and more number of upregulated transcripts related to steroidal saponin biosynthesis

Dioscorea species

Mainly steroidal saponins are found in the monocot species such as some species of Dioscoreaceae family. Steroidal saponins of this plant species, i.e., diosgenin is used as the raw material for industrial production of steroidal drugs (Lin and Yang 2008). Comparative transcriptome analysis of two high saponin-producing Dioscorea esculenta and Dioscorea cayenensis and a low saponin-producing Dioscorea alata was performed using 454 pyrosequencing to investigate steroidal saponin biosynthesis in Dioscorea species. From transcriptome analysis, it was found that DeF26G transcript was highly expressed in Dioscorea esculenta but not in Dioscorea alata. The biochemical characterization of DeF26G revealed that it was a furostanol glycoside 26-O-β-glucosidase that was involved in the conversion of protodioscin (furostanol saponin) to dioscin (spirostanol saponin). The expression of the DeF26G1 gene in the leaves of Dioscorea esculenta was higher than that in the tubers (Nakayasu et al. 2015). Using Illumina deep sequencing Dioscorea composita was used to analyze the transcriptome of 18-month-old Dioscorea composita. This information allows us to understand the biosynthetic pathway of steroidal sapogenins (diosgenin) in Dioscorea composita (Wang et al. 2015a, b, c).

Asparagus racemosus

Steroidal saponins in Asparagus racemosus mainly accumulates in the root system. The dried root of Asparagus is used as an ayurvedic medicine. The saponin present in the root of Asparagus collectively known as shatavarins. Leaf versus root transcriptome of Asparagus racemosus using Illumina sequencing approach revealed that majority of transcripts related to steroidal saponin biosynthesis including P450 and UDP-glucosyltransferase was found to be upregulated in root tissue as compared to leaf. This will lead to the hypothesis that steroidal saponin accumulated in the roots of Asparagus racemosus, which is mainly responsible for its pharmacological properties (Upadhyay et al. 2014).

Chlorophytum borivilianum

Safed musli has high saponin and polysaccharide content but the main active compound known as borivilianosides is found in roots. Steroidal saponins, borivilianosides (spirostanol type), are the major type of saponins present in Chlorophytum borivilianum. Earlier phytochemical studies indicated that initial reactions of saponins biosynthetic pathway occur in leaves, while later step modifications and storage occurs in roots (Kalra et al. 2013). Illumina HiSeq 2000 sequencing analysis of root tissue of this plant helps in the identification of candidate genes that encode enzymes controlling later stages of the saponin biosynthetic pathway and modifications (Kumar et al. 2016; Kalra et al. 2013).

Paris polyphylla

The rhizome as well as leaves and stem of Paris polyphylla is an important source of steroidal saponins. The root tissue is the main site of steroid saponin biosynthesis in Paris polyphylla so comparative transcriptome analysis was performed between 8-year-old and 4-year-old roots of Paris polyphylla. Comparative saponin biosynthesis in different aged roots showed that genes including P450 and UGTs involved in saponin biosynthesis were upregulated in the 8-year-old root, an understandable finding considering that the amount of saponin is higher in older roots (Liu et al. 2016a, b).

Trigonella foenum-graecum

The seeds and leaves of Trigonella foenum-graecum (fenugreek) are used as ayurvedic medicine in the Indian subcontinent to treat gastrointestinal ailments, high cholesterol, diabetes, wounds and inflammation. Leaves and seeds of this plant are edible and can be used as condiments (Chevallier 2000). Representational difference analysis (RDA) of cDNA of was performed using transcriptome user-friendly analysis (TRUFA, a webserver platform dedicated to RNA-seq analysis) was used to identify up-regulated genes in fenugreek in response to MeJA, cholesterol and squalene (Kornobis et al. 2015). MeJA mainly act as elicitor in plant defense under biotic and abiotic stresses. Majority of the transcripts involved in diosgenin biosynthesis including various P450 was found to be upregulated after MeJA treatment (Ciura et al. 2017) that further confirms the involvement of MeJA in the upregulation of secondary biosynthetic pathway related gene.

Trillium govanianum

Diosgenin that accumulates in rhizome as “Trillarin” is considered as the main bioactive constituent of Trillium govanianum. In addition to diosgenin govanoside, borassoside, and pennogenin were also reported to be present in the rhizome of Trillium govanianum (Rahman et al. 2015). Spatial differential expression of genes associated with steroidal saponin biosynthesis in rhizome, stem, leaf and fruit tissues of Trillium govanianum were studied using Illumina GAIIx sequencing platform. The expression analysis revealed that maximum numbers of genes are involved in saponin biosynthesis and found to be highly expressed in fruit and leaf tissues, indicating active biosynthesis of steroidal saponin takes place in fruit and leaf tissues. The downstream genes of steroidal saponin biosynthesis in leaf (SQS, CPI1, CyP5G1, FK, HDY1 and SMO2) and fruit (SQLE, SMT1, SMO1, DWF5, UGT80B1 and β-glucosidase) also showed higher expression level. These results indicate that steroidal saponins biosynthesis mainly occur in leaf and fruit tissues (Singh et al. 2017).

Allium species

Steroidal saponins were reported in more than 40 different species of allium. Tigogenin, diosgenin, alliogenin, gitogenin, agigenin, and β-chlorogenin are among the most common spirostanol sapogenins identified in the plants of allium species (Sobolewska et al. 2016). High-throughput RNA-Seq of the root, bulb and leaf tissues of Allium fistulosum with extra chromosome 2A from shallot (FF2A), monosomic addition lines (MALs) and Allium cepa aggregatum group (AA) was carried out using Illumina’s HiSeq 2500 platform to gain molecular insight into the Allium saponin biosynthesis pathway. It was observed that in FF2A line acetyl-CoA-acetyltransferase and SQS are highly expressed transcripts that are related to saponin biosynthesis. Further oxidation, hydroxylation and glycosylation steps of the saponin backbone via P450 and UGT family transcripts, respectively, were also found to be remarkably up-regulated in the FF2A line (Abdelrahman et al. 2017).

Production of steroidal saponin in culture media

Due to the limited availability of medicinal plants and slow production of medicinally important secondary metabolites, it is necessary to develop in vitro techniques for the production of bioactive compounds that can be useful for pharmacological purposes. Advances in functional genomics and application of multidisciplinary approaches help to discover new plant saponins that have different pharmacological properties. Plant cell and tissue culture methods have been explored as potentially more efficient alternatives for the production of saponin and other plant derivatives. These methods were started many years ago but the optimization of culture condition was still going on to increase the saponin production for commercial use.

In vitro culture methods to obtain steroidal saponin can be possible by optimizing plant growth regulator/plant hormone concentration in culture media, light treatment, growth conditions, precursor feeding and other parameters to obtain better results. For example, plant hormone auxin was used to induce the production of steroidal saponin solanine, solasodine and solanidine in the tissue and suspension culture of Solanum lyratum (Kuo et al. 2012). Similarly, steroidal saponin (aculeatiside A, aculeatiside B) production was observed in considerable amount in the hairy root culture of Solanum aculeatissimum under continuous light treatment for 8 weeks (Ikenaga et al. 1995). Epicotyls of Solanum aculeatissimum were used to induce callus culture and it was observed that it was able to produce steroidal saponins aculeatiside A and B (Ikenaga et al. 2000). Attempts were also made for the production of steroidal saponins from the roots of Asparagus racemosus and its callus and suspension cultures were established to produce high levels of saponins both intracellularly and extracellularly. Being an endangered species it is necessary to propagate this plant through tissue culture because traditional method of propagation through seeds is not an efficient method and it takes a lot of time. Using in vitro culture methods 20 times higher levels of saponins (shatavarins) were produced that has found to stimulate immunological response (Pise et al. 2015).

Root cultures are considered as reliable and alternative sources for the production of valuable secondary metabolites. It is easy to culture root tissues of dicot plant but in the case of monocot plants where root forms modified structures such as bulb, corm or tubers the root culture seems to be difficult but the tuberous root culture of Chlorophytum borivilianum was developed and considerable amount of saponin was quantified (Basu and Jha 2015). Effect of growth regulators was determined in the adventitious root culture of Dioscorea nipponica and it was observed that dioscin and prosapogenin C content in indole-3-butyric acid (IBA) treated roots were significantly higher as compared to 1-naphthaleneacetic acid (NAA) treated roots (Ahn et al. 2005). In in vitro shoot culture of Bacopa monnieri it was observed that application of abiotic elicitors (MeJA, salicylic acid, and CuSO4) enhances bacoside production and among these elicitors, CuSO4 was found to accumulate bacoside in a large amount (Sharma et al. 2015).

Somatic embryogenesis is a valuable technique in medicinal plant improvement programs, such as for propagation and genetic transformation. Embryogenic callus culture derived from the leaf explants of Tribulus terrestris L. accumulated steroidal saponin, i.e., diosgenin. Saponin content in embryogenic callus culture of Tribulus terrestris was found to be similar to that of seeds of naturally growing plants after the application of casein hydrolysate (CH) in growth media (Nikam et al. 2009). Fagonia indica burm was cultured in vitro on MS medium supplemented with 2,4-d for rapid regeneration of plant. Somatic embryogenesis was induced by the elimination of 2,4-dichlorophenoxyacetic acid (2,4-d) and a slight increase of 6-benzylaminopurine (BA) concentration (Ebrahimi and Payan 2013). Shoot organ culture of Yucca schidigera accumulated all kinds of sapogenins as in plant shoot but in different concentrations and their glycoside pattern was similar to that of saponins present in mature Yucca schidigera rhizome (Kaneda et al. 1987).

Steroidal saponin production in cell and tissue culture of certain plant species can also be increased by the addition of hormone or biotic and abiotic growth regulators like the production of diosgenin was stimulated by light or fungal elicitation in the cell suspension cultures of Dioscorea galeottiana when cells were grown in the dark condition. Application of low phosphate and sucrose in the growth medium results in highest production of diosgenin (Rojas et al. 1999). Diosgenin accumulation in the tissue culture of Dioscorea deltoidea was enhanced in the presence of 2,4-d (Marshall and Staba 1976).

Microbial transformation of saponin present in Dioscorea zingiberensis tubers was difficult due to the association of saponin with starch so it cannot be able to make contact with microorganisms so a combined technology of enzymatic saccharification for excluding starch from raw herb and the microbial transformation was established to prepare diosgenin from the tubers of Dioscorea zingiberensis. Using these approach saponins from Dioscorea zingiberensis was converted into diosgenin using the fungal culture of Trichoderma reesei (Zhu et al. 2010). A strain of Gibberella intermedia was also used for conversion of saponins into diosgenin by optimizing culture conditions, and using this method three times higher diosgenin yield was observed as compared to original culture media (Zhang et al. 2013a, b, c). In addition to this, nine steroidal saponins were obtained by the fermentation of total furostanol saponins from Dioscorea zingiberensis tubers incubated with the culture of a fungus Absidia coerulea. Two compounds among them show induced platelet aggregation activity due to change in their chemical structure (Pang et al. 2015). Biotransformation of total steroidal saponins of Agave sisalana into tigogenin was achieved using a rod-shaped bacterium obtained from soil samples of karst area of Guilin, China. The microbial transformation is an eco-friendly approach in the production of tigogenin (Wang et al. 2014a, b, c).

Rhizomes and roots of Ruscus aculeatus contain steroidal saponins (ruscogenins) that have several pharmacological properties. Due to its slow growth rate, complicated cultivation methods and pollination failure make this plant a vulnerable species. Ruscogenin biosynthesis in Ruscus aculeatus is greatly influenced by its clonal origin and culture type that forms the basis of its ex situ conservation (Ivanova et al. 2015). In vitro culture of Ruscus aculeatus has a potential capacity to biosynthesize steroidal saponin. Rhizome segments of this plant are used to generate calli. It was observed that young regenerated plants of about 6 weeks old have a greater capacity to biosynthesize and accumulate saponins than calli cultures and the neoruscogenin contents were found to be greater than those of ruscogenin in all samples, i.e.,plantlets and calli (Palazón et al. 2006).

It is challenging to produce saponins in the heterologous system, i.e., in microbes, and only limited successful pathway engineering attempts have been reported. The effects of several previously unexplored gene knockout targets of Saccharomyces cerevisiae were assessed for the heterologous production and to improve the production capabilities of this saponin production platform. Dramatic expansion of the endoplasmic reticulum stimulates the production of recombinant triterpene biosynthetic enzymes through disruption of the phosphatidic acid phosphatase-encoding PAH1 of Saccharomyces cerevisiae through CRISPR/Cas9 which results in the accumulation of triterpenoids and triterpene saponin (Arendt et al. 2017). Transgenic yeast strains expressing β-amyrin synthase (bAS), P450 reductase, CyP93E2 and CyP72A61v2 were able to produce soyasapogenol B, and yeast strain over expressing bAS, P450 reductase, CyP716A12 and CyP72A68v2 produce gypsogenic acid. Additionally, P450 s that seemed not to work together in planta were combinatorially expressed in transgenic yeast for the production of saponins (Fukushima et al. 2013). Combinatorial synthetic biology program was also successful for the synthesis of monoglycosylated saponins in yeast by combining CyP716Y1 with oxidosqualene cyclase, P450, and glycosyltransferase genes from different plant species (Moses et al. 2014a, b). Another transgenic yeast strain expressing dammarenediol-II synthase and protopanaxadiol synthase genes of Panax ginseng, together with a NADPH-cytochrome P450 reductase gene of Arabidopsis thaliana, were able to produce protopanaxadiol (Dai et al. 2013) and bioactive ginsenosides Rh2 or Rg3 were produced from simple sugars by microbial fermentation (Wang et al. 2015a, b, c). These platforms will provide an alternative approach to replace the traditional method of ginsenosides extraction from Panax plants. Elicitation process is another one of the most effective strategies to enhance metabolites biosynthesis and accumulation in biotechnological systems. By adopting this elicitation strategy, yeast extract can be used to elicit ginsenoside production in hairy root cultures of Panax quinquefolium (Kochan et al. 2017). Reports were available, that employ synthetic biology approach for heterologous production of triterpenoidal saponin but this technique was not reported in context of steroidal saponin production. Efforts will be made for the development of engineered yeast system that will provide an efficient system for large-scale production of steroidal saponins in near future.

Future prospects

Steroidal saponins possess a wide range of medicinal and biological properties due to the presence of structural diversity in them. To find out new plant sources of steroidal saponins and to identify the presence of new steroidal saponins in the known medicinal plants will be the area of interest of many researchers. The anti-cancerous properties of most of the steroidal saponins were already being explored widely and is still going on but the other pharmacological properties associated with this compound is yet to be investigated.

The emergence of NGS platform has revolutionized our understanding related to saponin biosynthesis especially the modification of saponin backbone. Most of the P450 s and UGTs that were involved in terpenoidal saponin biosynthesis have been characterized but identification and characterization of these two enzymes that will convert obtusifoliol to furostanol and spirostanol type of steroidal saponin are yet to be needed as they are responsible for providing different biological properties to steroidal saponins. Transcriptome data of non-model plant will be helpful in discovering new candidates of P450 s and UGTs that may be involved in the saponin backbone modification but they need to be characterized further.

The advancement in the synthetic biotechnology tools like CRISPR/Cas9-based approaches will be beneficial to engineer medicinal plants regarding secondary metabolites production and to functionally characterize the biosynthetic (backbone modifying) and regulatory enzymes related to steroidal saponin biosynthesis. Due to the lack of availability of functional mutants in most of the medicinal plants, the genome editing technology based on type II CRISPR/Cas system can be utilized to establish mutants for gene function analysis. This technique relies on Agrobacterium tumefaciens mediated transformation technology and can also be combined with Agrobacterium rhizogenes-mediated transformation. Agrobacterium rhizogenes-based Ri transformed hairy roots show the characteristics of rapid growth, reduced apical dominance, high branching, and enhanced stable production of secondary metabolites, making them a promising system for investigating biosynthesis of various secondary metabolites, especially in medicinal plants. CRISPR/Cas9 system has a wide range of applications, with more accuracy and precision as compared to other genome editing tools. This approach can be applied for converting saponin-producing plants into biofactories for mass production of saponin by diverting the flux towards steroidal biosynthetic pathway. For example, by blocking the bAS gene of triterpenoidal saponin biosynthetic pathway, phytoene synthase gene of carotenoid biosynthetic pathway or geraniol synthase gene of alkaloid biosynthesis, metabolic flux can be shifted towards steroidal saponin production. Additionally this technique will be beneficial for elucidating the function of novel P450 and UGTs. As the life cycle of saponin metabolite producing plants is generally high so Cas9 based system can further be employed to control the spatiotemporal patterns of gene expression in plants and modulating life cycles of various economically useful plants to obtain useful metabolites in short duration of time.

Saponin molecules are synthesized in plants in very trace amount that will not satisfy the needs of its commercial use and the plant sources are also very limited. So there will be the need for the production of saponins using in vitro culture methods. Tissue culture-based techniques were already being developed and need to be further optimized. Combinatorial biosynthesis of steroidal saponins in yeast system will be a part of future research and research is going on to generate semi-synthetic steroidal saponin.

Author contribution statement

SU has compiled and written the manuscript. GSJ and S have edited the manuscript. RKS has conceptualized, supervised and edited the manuscript.


Swati, Gajendra, and Shikha acknowledge CSIR-UGC for fellowship. Shukla RK would like to acknowledge SCIENCE & ENGINEERING RESEARCH BOARD EMR/2016/005256 (GAP397) for funding.


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  • Swati Upadhyay
    • 1
  • Gajendra Singh Jeena
    • 1
  • Shikha
    • 1
  • Rakesh Kumar Shukla
    • 1
    Email author
  1. 1.Biotechnology Division (CSIR-CIMAP)Central Institute of Medicinal and Aromatic Plants, (CSIR-CIMAP) P.O. CIMAP (a laboratory under Council of Scientific and Industrial Research, India)LucknowIndia

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