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22. Swertia hickinii Burkill / 20 (He et al., 1999) //
https://t.me/med1917
23. Swertia iberica Fisch. and
C.A. Mey.
24. Swertia japonica (Schult.)
Makino
25. Swertia kilimandscharica
Engl.
26. Swertia komarovii Pissjauk / 26 (Krogulevich, 1978) //
27. Swertia lawii Burkill /// 26 (Mallikarjuna,
28. Swertia lugardiae 10
29. Swertia lurida Royle ex D.
Don
30. Swertia macrosepala Gilg / 26 (Hedberg and Hedberg, 1977) / -/
31. Swertia minor (Grisebach)
Knobl.
32. Swertia nervosa (Wall. ex G.
Don) C.B. Clarke
33. Swertia paniculata Wall. 8 (Khatoon
34. (Bala and Gupta,
35. Swertia perennis L. 14 (Post,
36. Swertia petiolata D. Don // 13 (Jee et al., 1989) e
/ 26 (Davlianidze, 1984; Gagnidze and
/ 20, 21 (Shigenobu, 1983)
/ 26 (Hedberg and Hedberg, 1977) //
(Nemomissa,
1998)
// 13 (Vasudevan,
/// 20 (Mallikarjuna,
/// 26 (Sharma, 1970)
and Ali, 1993)
1983)
Gviniaschvili, 1984; Gvinianidze and
Avazneli, 1982)
///
/ 8(Vasudevan, 1975)
28 (Love and Love, 1986; Dawe and
Murray, 1979)
/
/
1975; Mehra and
Vasudevan, 1972)
2011)
//
/
1985)
/
1985)
/
Continued
12.2 Cytological studies of genus Swertia 229
Table 12.1 Table showing names of taxons and chromosome numbers in cytologically worked out species at world level.dcont’d
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Chromosome number in cytologically worked out species
Outside India India
Name of the taxon
37. Swertia pseudochinensis H.
Hara
38. Swertia punctata Baumg. / 28 ( Tam and Vladimirov, 2001) //
39. Swertia speciosa Wall. // 13 ( Vasudevan,
40. Swertia stenopetala Pissjauk. / 14 (Zhukova et al., 1973) //
41. Swertia subnivalis T.C.E. Fr. / 26 (Hedberg and Hedberg, 1977) //
42. Swertia swertopsis Makino / 52 (Shigenobu, 1982, 1983) //
43. Swertia tashiroi Makino / 60 (Shigenobu, 1982,
44. Swertia tetragona R. H. Miao /// 18 (Vasudevan,
45. Swertia tetrandra Hochst. 10
46. Swertia tetraptera Maxim. / 14 (He et al., 1999) //
47. Swertia thomsonii C. B.
Clarke
48. Swertia trichotoma Wall. // 26 (Mallikarjuna et al.,
49. Swertia tongluensis Burkill /// 18 (Sharma and
50. Swertia uniflora Mildbr. / 26 (Hedberg and Hedberg, 1977) //
51. Swertia veratroides Maxim.
ex Kom.
Swertia wolfgangiana Gru¨ning
n2n n2n
/ 20 (Shigenobu, 1983) //
1975)
1983) //
(Nemomissa,
1998)
// 13 (Vasudevan,
/
/
///
1975; Khoshoo and
Tandon, 1963)
1987)
26 (Probatova, 2006)
28 (He et al., 1999)
/
/
/
1975; Khoshoo and
Tandon, 1963)
/
/
Sarkar, 1967e68
/
/
230 CHAPTER 12 Swertia chirayita
)
12.3 Genetic diversity studies 231
https://t.me/med1917
The family Gentianaceae represe nts a lot of polyploidy and dysploidy in its species, as a wide range of base numbers exist, i.e., x ¼ 5, 6, 7, 8, 9, 10, 11, 12, 13, 15,
17, 19, or higher. But it is dominated by x ¼ 9 in most of the genera of the family.
According to Favarger (1952), the original base number of the family is x ¼ 5, and
that the base number of x ¼ 11 might be a result of polyploidy followed by fragmentation. The same explanation may be given to another base number x ¼ 13 which
may have originated from ¼ 7 by polyploidization and fusion. The same idea was
adopted by Skalinska (1952) while studying chromosome number in Gentiana frig-
ida (2n ¼ 24) and reveals that x ¼ 11 (5 þ 6) and x ¼ 13 (6 þ 7) have been evolved
from x ¼ 5, 6, and 7.
12.3 Genetic diversity studies
Evaluation of diversity in a plant is important for knowing its life dynamics and
sustainability in future. Estimates of DNA polymorphism show the capability of
adaptive behavior in changing climates and evolutionary history of that plant species. However, studies of DNA polymorphism in S. chirayita are severely lacking
and only two reports are available which used only limited accessions of this species
because of its declining populations in nature. Joshi and Dhawan (2007a ,b) analyzed
13 accessions of S. chirayita using ISSR markers and revealed high genetic diversity
among analyzed samples. In a recent study, Kaur et al. (2019) explored the interspecific genetic diversity of five Swertia species including five accessions of
S. chirayita. They used ISSR and RAPD markers and concluded that S. chirayita
showed low diversity. There is urgent need of studies with more numbers of
populations and accessions which should be analyzed using better marker systems
such as Simple Sequence Repeat markers so that accurate estimates can be drawn
which can help in designing conservation and management strategies in future.
12.3.1 Genes and metabolite association studies
Molecular data related to the biosynthesis of secondary metabolites of S. chirayita
are lacking. However, few workers have tried to explore the pathways and relative
contents of secondary metabolite and their association to regulatory genes and miRNAs (Vaidya et al., 2013; Fan et al., 2014; Kumar et al., 2014; Padhan et al., 2015,
2016; Koul et al., 2016; Liu et al., 2017; Pal et al., 2018; Padhan, 2018). Padhan et al.
(2015) reported the expression profiling of swertiamarin, amarogentin, and mangi-
ferin biosynthesis pathway genes and their correlation with the respective metabolites content in different tissues of S. chirayita. They observed that root tissues of
greenhouse-grown plants contained the maximum amount of swertiamarin, whereas
maximum accumulation of mangiferin in floral organs. Ten genes of the secoiridoids
biosynthesis pathway and five genes of mangiferin biosynthesis identified and
correlated to corresponding metabolite contents. Their results can be important in
its genetic improvement works. Liu et al. (2017) generated transcriptome sequences
from the root, leaf, stem, and flower tissues of S. mussotii to understand the
232 CHAPTER 12 Swertia chirayita
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secoiridoid biosynthesis pathway and identified 39 candidate transcripts encoding
the key enzymes for secoiridoid biosynthesis. Their study revealed that the levels
of three bioactive compounds, i.e., sweroside, swertiamarin, and gentiopicroside,
were variable in different tissues. They found no significant correlation with the
expression profiles of key genes and suggested complex biological behaviors in
the coordination of metabolite biosynthesis and accumulation. Pal et al. (2018)
generated comparative transcriptomes of S. chirayita to decipher the genes and other
regulatory components related to secondary metabolites biosynthesis. They found
that 19 genes from primary metabolism showed higher in silico expression indicating their involvement in regulating the central carbon pool. When validated by
qRT-PCR, 10 genes showed similar expression pattern across both the methods.
The authors identified differentially expressed transcription factors and ABC-type
transporters putatively associated with secondary metabolism in S. chirayita.
12.4 Bioactivity and medicinal uses
This plant is used as expectorant, laxative, antispasmodic, antioxidant, antidiabetic,
antipyretic, antitussive, stomachic, anthelmintic, and antidiarrhea. It has many
medicinal properties such as antiinflammatory, hypoglycemic, hepatoprotective,
antibacterial and wound healing (Laxmi et al., 2011; Tabassum et al., 2012;
Mahmood et al., 2014; Alam et al., 2009). Different ailments cured by
S. chirayita are described in Table 12.2. Besides, many medicinal compounds
were isolated from natural herbs which have anticancer, antitumor, and anti-AIDS
properties (Sultana and Ahmed, 2013).
Table 12.2 Ailments cured by using different procedure and parts of Swertia
chirayita.
S.
No. Ailments Parts used Procedure
1 Malaria Whole plant Overnight dipped plants in
2 Digestive organs Whole plant Decoction as tonic
3 Bronchial asthma, cough, cold,
headache and fever, diarrhea,
constipation, dyspepsia,
burning of the body, and skin
diseases
4 Liver diseases and urinary
disorders
5 Joint pain Roots Roots crushed and paste
6 Boils and scabies Leaves Leaves warmed and paste
Whole plant Paste of plant
Roots Root juice
water and juice extracted
rubbed over joints
prepared with mustard oil
applied over boils and scabies
12.5 Tissue culture studies in Swertia chirayita 233
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12.4.1 Active principles
Swertia contain many bioactive compounds in the form of phenolics compounds,
alkaloids, and other secondary metabolites with wide therapeutic importance and
potential applications (Singh et al., 2012). The major classes of phytochemicals
include many Xanthone compounds and other phytochemicals specifically swerchirin, swertiamarin, swertanone, swertenol, episwertinol, chiratenol, gammacer16-en-3b-ol, 21-a-H-hop-22(29)-en-3b-ol, taraxerol, oleanolic acid, ursolic acid,
ophelic acid, and 1,3,6,7-tetrahydroxyxanthone-C-2-b-D-glucoside (mangiferin)
which have shown different therapeutic effects (Singh, 2005; Ghosal et al., 2006;
Iqbal et al., 2006; Selvam, 2012; Singh et al., 2012).
12.5 Tissue culture studies in Swertia chirayita
Plant tissue culture methods have been used in conservation of medicinal plants for
past many years. Over time, these methods have been developed and modified for
exploiting medicinal plant potential and helping in establishment of numerous plant
productsebased industries like pharmaceutical, nutrition, and life style. IHR (IndoHimalayan region) is well known for its biodiversity, flora, and fauna. S. chirayita is
critically endangered and listed in Red Data book. There have been so many conservation measures taken to conserve the plant and to increase its natural populations.
But these programs were not efficient in achieving the goal as natural propagation in
this plant is not good because of seed dormancy, seed viability, and natural pathogen
attack or excessive grazing in its natural habitats. Vegetative propagation programs
are also highly discouraged because of unavailability of quality planting materials,
difficult to establish root stocks; moreover, it cannot be propagated through cutting
or grafting. Therefore, this plant has remained quite important among the plant
biotechnologists and has been studied extensively for establishment of in vitro propagation protocols across the globe. S. chirayita is having high commercial potential
because of its therapeutic properties and variety of bioactives. The various plant
tissue culture strategies such as tissue and organ cultures, suspension cultures/callus
cultures, somatic embryogenesis, and authentication of micropropagated plantlets
and more recently in vitro co-culture, have been studied in this plant. Therefore,
current study has been done to compile these studies like micropropagation,
caulogenesis, rhizogenesis, callogenesis, somatic emb ryogenesis, and their
advancements.
Seeds, nodal segments, axillary buds, and leaves as explants have been used from
field-grown young juvenile plants, and explants from micropropagated plants have
been tested for enhanced multiplication of S. chirayita. Nodal segments or nodal
explants have been found more suitable for enhanced shoot multiplication and production of genetically true-to-type plants among all explants tested (Sharma et al.,
2013, 2016). Almost all studies in this plant have reported the use of MS as basal
tissue culture media; Gamborg B5 have also been reported in some studies with
modifications for in vitro bud break and culture establishment. Optimum shoot
234 CHAPTER 12 Swertia chirayita
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development and maturation have been observed in MS medium (Sharma et al.,
2013, 2016; Wang et al., 2009; Chaudhuri et al., 2007; Wawrosch et al., 1999).
The type of carbon source and its concentration plays a very significa nt role in
micropropagation of plants. Optimum caulogenesis has been reported when medium
was fortified with sucrose as carbon source at a concentration of 2.5%e3% (Sharma,
2011). In vitro propagation of S. chirayita has been reported by many researchers
using different explants, like shoot tips and axillary buds (Wawrosch et al., 1999;
Joshi and Dhawan, 2007a,b; Ahuja et al., 2003), leaf segments (Chaudhuri et al.,
2008; Wang et al., 2009), cultured root explant (Pant et al., 2010), and nodal segment
(Sharma et al., 2013, 2015, 2016; Chaudhuri et al., 2007).
One of the safer approaches to decrease the somaclonal incidences and confirming genetic fidelity of micropropagated plantlets is to use axillary buds and nodal
segments. The importance of PGRs (auxins and cytokinins) in breaking axillary
bud dormancy, or shoot multiplication, as well as their effect alone or in combination
for shoot proliferation, multiplication, and elongation has been well established in
S. chirayita.
The effect of BAP with respect to induction and shoot enhancement in initial
phase of culture establishment has been reported by many workers. Also the synergistic effect of BAP and IAA or additives like adenine sulfate in enhancing the shoot
multiplication rate and reducing the use of other cytokinins has been obtained in
S. chirayita (Sharma et al., 2013, 2016). Cytokinins BAP; Kinetin; 2-iP (alone or
in combination with Kinetin or auxins IAA; IBA; NAA) have been used. BAP alone
or in combination with other cytokinin or auxins in lower proportions has been found
optimum in shoot multiplication from nodal segments and leaf explants (Sharma
et al., 2016; Wang et al., 2009). One recent study on micropropagation of Withania
somnifera involving the use of in vitro co-culture with PGPRs has resulted in shoot
proliferation with faster rates, enhanced and vigorous rhizogenesis, and use of very
reduced concentration of PGRs. Reduced use of phytohormones results in lesser
incidences of basal callusing during shoot proliferation and thus reduces the chances
of som aclonal variations and confirms the clonal identity (Sharma et al., 2015).The
in vitro co-culture can also serve as a bioassay for screening of positive plant
microbe interactions that can be further exploited for various other beneficial
outputs. Addition of growth factors like polyamines, reduced nitrogen sources
like adenines, and coconut milk has profound effect on in vitro caulogenesis
(Sharma et al., 2013, 2016; Sharma, 2011). Indirect organogenesis via callus formation is also a good means of mass multiplication; somatic embryogenesis has been
studied in S. chirayita using nodal segment, leaf explant, in vitro grown seedlings,
and internode (Chaudhuri et al., 2009; Sharma, 2011). The superiority of 2,4-D in
establishing the callus and its maturation has been reported by the workers. The suitability of leaf explant for inducing embryogenic callus over intermodal segments has
also been reported. It has been suggested that it is better to take leaf explants from
micropropagated plantlets as they bypass the additional steps of sterilization and
give better response. Root emergence, development, and elongation are vital steps
12.6 Ecological status 235
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in every tissue culture study. Same has been extensively studied in Chirata also. Influence of phytohormones and media strength has morphogenic effects on the events
of rooting. Auxins alone or in combination with other auxins in rooting the microshoots were effective in S. chirayita (Joshi and Dhawan, 2007a,b; Sharma et al.,
2016; Wang et al., 2009). Direct organogenesis via leaf explant has also been
reported in S. chirayita (Chaudhuri et al., 2008). One study has also reporte d the
in vitro flowering in S. chirayita through tissue culture (Sharma et al., 2014).
12.6 Ecological status
As S. chirayita is a highly important medicinal plant and used as an ingredient in
Ayurvedic, Unani, Siddha, and modern medicines, it has become vulnerable in its
natural state in forest due to its overextractions. It is in great demand in pharmaceutical industries both nationally and internationally and, therefore, its overexploitation, unsustainable harvesting, overgrazing, and illegal trading has put this plant
under critical endangered status (Kumar and Van Staden, 2016; Badola and Pal,
2002). Destruction of its microhabitat and overexploitation leads to drastic reduction
in its natural population all over the Himalayan region (Bhatt et al., 2005). Its low
seed viability, long gestation period, and low seed germination percentage are
also the major causes for its dwindling natural population (Samant et al., 1998; Joshi
and Dhawan, 2005). Exceeding developmental activities, building of new roads and
infrastructure, unorganized urbanization, and increasing anthropogenic pressure all
over the Himalayan region are the main driving forces for its habitat destruction. It
can leads to the loss of biodiversity and extinction of species from its natural habitat.
The extinction rate of plant species was estimated to 100e1000 times faster than the
natural speed of extinction (Kumar and Van Staden, 2016). Recently, ministry of
environment and forestry has banned the export of this plant along with many other
Himalayan medicinal plants. Efforts were made for its in situ and ex situ conservations (Nishteswar, 2014). The main focus is to conserve the habitat of the S. chirayita
by maintaining the open spaces with moist slopes around the forest and stopping the
surrounding shrubs to encroach the habitat. The surrounding shrubs overtake the
habitat of S. chirayita and modify the microhabitat resulting in the decrease in its
population. Maintaining the microhabitat, allowing the sustainable harvest, and
clearance of shrubs are helpful in preserving the microhabitat of this species as an
excellent strategy for in situ conservation. Approaches have also started to conserve
the species through in vitro tissue culture techniques. Mass multiplication has been
carried out by tissue culture technique via direct shoot multiplication through leaf
explants Chuadhury et al. (2008). Restoration of natural population can be carried
out by planting the multiplied material through tissue culture. Cultivation is another
approach for the ex situ conservation of natural population of S. chirayita and can
save it from the overexploitation from natural habitat (Shukla et al., 2017). Species
Distribution Modeling and Ecological Niche Modeling can be some excellent
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approaches for identification of the suitable niche (microclimatic region) for restoration of the natural population of S. chirayita in the Himalayan region (Gaikwad
et al., 2011; Jaryan et al., 2013).
12.7 Conclusion
S. chirayita is an important medicinal plant which yields many important active
principles. However, declining populations due to habitat destruction and human
disturbances is the major concern. The plant has been explored to some extent
including chromosomal, genetic, genomic, biochemical, tissue culture, and ecological levels but more research works are required to make its best use in a sustainable
way. The different ecotypes if any present in natural populations should be explored
and documented. Genetic diversity of available germplasm using molecular tools
needs to be assessed to make insights into past and future population dynamics of
the species. The elite accession having high contents of active principles present
in the natural populations needs to be identified and multiplied either through
conventional methods or using tissue culture technology. More insights into biosynthetic pathways of active principles of the plants are required so that desired genetic
manipulations can be done in future to increase the yield. Hence, these are some
major research fields where the future works can be focused for the conservation
and utilization of this incredible plant species.
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