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218 CHAPTER 11 Stevia rebaudiana
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Kong, M.K., Kang, H.J., Kim, J.H., Oh, S.H., Lee, P.C., 2015. Metabolic engineering of the
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Brandle, J.E., Gijzen, M., Starratt, A., 2000. Stevia rebaudiana, Its Biological, Chemical and
Agricultural Properties (No. 00991).
Lucho, S.R., do Amaral, M.N., Lo´pez-Orenes, A., Kleinowski, A.M., do Amarante, L.,
Ferrer, M.A´., Caldero´n, A.A., Braga, E.J.B., 2019. Plant growth regulators as potential
elicitors to increase the contents of phenolic compounds and antioxidant capacity in Stevia plants. Sugar Tech. 21 (4), 696e702.
Wang, Y.H., Avula, B., Tang, W., Wang, M., Elsohly, M.A., Khan, I.A., 2015. Ultra-HPLC
method for quality and adulterant assessment of steviol glycosides sweetenerseStevia
rebaudiana and Stevia products. Food Addit. Contam. 32 (5), 674e685.
Swertia chirayita
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CHAPTER
12
Vijay Singh1, Vikrant Jaryan2, Vikas Sharma3, Himanshu Sharma4, Indu Sharma2,
1
Department of Botany, Mata Gujri College, Fatehgarh Sahib, Punjab, India;2Department of
Botany, Sant Baba Bhag Singh University, Khiala, Jalandhar, Punjab, India;
Molecular Biology and Genetic Engineering, Lovely Professional University, Jalandhar, Punjab,
4
India;
Agri-Biotechnology Division, National Agri-Food Biotechnology Institute, Mohali, Punjab,
Vikas Sharma
3
Department of
India
12.1 Introduction
Swertia chirayita Roxb. ex Fleming commonly known as Chirata is one of the most
important medicinal plants distributed from temperate to the alpine region of the Indian Himalayas, ranging across the altitudinal gradient from 1200 to 3000 m above
sea level (Clarke, 1885; Tandon et al., 2010; Pradhan and Badola, 2015). Sometimes
it is also found up to 3400 m above sea level (Bora and Singh, 2016). It is an indigenous species of the Himalaya and distributed from Kashmir to Sikkim. Besides, this
species is also found to grow in Khasi hills at an altitude from 1200 to 1500 m
(Tandon et al., 2010). It is also found to grow in Nepal, Bhutan, Pakistan, and
Afghanistan (Pradhan and Badola, 2015). This plant species belong to family Gentianaceae and is one of the important ingredients of Ayurvedic, Unani, Siddha, and
modern medicines (Bora and Singh, 2016). It is a synonym of Gentiana chirayita
Roxb. ex Flem.; G. chirayita Wall. It is an old traditional herb and its medicinal
value was known from earlier times. It was known Kiratatikta in Sanskrit, while
Nilavaembu in Tamil (Selvam, 2012). The plant is native of the Himalaya and highly
threatened due to its high demand as a medicinal plant and its occurrence in narrow
range of specialized habitat. In this chapter, we will discuss the occurrence, habitat,
morphology, cytology, ecology, biochemical constituents and genetic diversity of
the S. chirayita and its comparative account to related species of Swertia.
2
12.1.1 Morphology of plant
S. chirayita is an annual or biennial herb with 60e125 cm in height or sometimes
reaches up to 1.5 m in height (Aleem and Kabir, 2018; Kumar and Van Staden,
2016; Selvam, 2012). Its stem is sometimes robust, branched, cylindrical below,
four angled upward containing a large pith (Selvam, 2012). Stem is yellowish or
purplish in color (Aleem and Kabir, 2018). Its leaves are opposite, sessile, broadly
ovate or lanceolate, decussate five nerved with round base, acuminate at apex,
Himalayan Medicinal Plants. https://doi.org/10.1016/B978-0-12-823151-7.00016-7
Copyright © 2021 Elsevier Inc. All rights reserved.
223
224 CHAPTER 12 Swertia chirayita
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glabrous, cordate or obtuse at base with margins entire, and three to seven prominent
lateral veins. Plant flowers generally from July to August in rainy season. Its inflorescence has large leafy panicles of solitary axillary or axillary with three to five
flowers arranged in terminal clusters. Flowers are generally yellowish-green outside
and purple inside. They are tetramerous, drooping, or erect. Fruit is a capsule which
is ovoid or ellipsoid. Seeds are numerous, minute, globose, and brownish.
12.1.2 Habitat and climatic conditions
S. chirayita generally grows in subtropical to temperate forests in open forest margins. It prefers cool and moist places with shady moist slopes and tall grasses (Sel-
vam, 2012). Many studies confirm that the species performance varies among
microhabitats (Shrestha and Jha, 2010; Pradhan and Badola, 2012 ). Present climate
change is the biggest threat for the survival of this threatened species because of its
narrow range of specialized habitat and has a greater risk of instability which can
lead to the extinction of this species (Brys et al., 2005; Samant et al., 1996). Therefore, there is great demand to understand the climatic condition of these microhabitats to conserve narrow range of this specialized habitat species which can be
helpful for planning its conservation and management (Hegland et al., 2001; Colling
and Matthies, 2006; Kalliovirta et al., 2006).
This plant prefers sandy (light), loamy (medium), as well as clay (heavy) soil
conditions. The plant flourishes well in acidic, neutral, as well as alkaline soils.
The plant prefers semishade or woodland conditions or needs humid or damp soil.
The Chirata plants can withstand temperatures as low as 15
to grow well (Kumar et al., 2010).
C and still continue
12.2 Cytological studies of genus Swertia
Cytology is one of the most important fields in biology and deals with the study of
the cells, especially their structure, function, and chemistry with a focus on the chromosome. The chromosomes are thread-like structures in which DNA molecule is
packaged tightly and cemented with the help of histone proteins and carry the
genetic information from parents to offspring through meiosis. But the genetic
information is carried out through the generation by the process of meiosis only
by chromosomes of germline cells. However, the somatic cells help only in the
regeneration and maintenance within the generation by the process of mitosis.
Therefore, meiosis is required to maintain a genetic balance over generations,
although it produces variations via independent assortment and crossing over between nonsister chromatids of homologous chromosomes. Meiosis is best known
for its reliability over sexual life cycles, as it maintains genome stability through
many events (cellular and molecular), such as DNA/chromosome replication in
G1 phase, chromosome pairing and recombination in prophase-1, chromosomal
segregation in anaphase-I/II and cytokinesis. Further, the process of meiosis involves
12.2 Cytological studies of genus Swertia 225
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reductional and equational division and results in four haploid gametes (half genome
compared to the parent cell). Homologous recombination through crossing over in
the pachytene stage (prophase-I) is a dynamic process by which DNA sequences/
strands are exchanged, hence induce structural aberrations and ultimately cause
the evolution of gene/genome.
It is a widely accepted view that cytotaxonomy is one of the key determinants in
the plant systematics as chromosome numbers significantly contribute to grouping
plant species/taxa. The first ever report of the cytological work in the flowering
plants (orchids) was given by Strasburger in 1882 (cf. Fedorov, 1969), while the first
chromosomal report for Indian populations was made by Johnson in 1910 (cf. Dar-
lington and Wylie, 1955). The determination of chromosome numbers coupled with
detailed meiotic studies is always a valuable effort that provides a future platform for
the researchers in applied sciences. The chromosomes through their numerical
(euploid or aneuploid cytotype) and structural changes have played a key role in
chromosomal evolution by producing reduced or doubled gametes and the phenomenon is quite common in Angiosperms (Stebbins, 1950; Ornduff et al., 1963;
Heywood et al., 1977; Robinson et al., 1981).
In the family Gentianaceae, at world level, a number of prominent cytologists
have explored the family, which includes Favarger (1952), Skalinska (1952), Lo¨ve
(1953), etc. In Indian populations, the previous detailed chromosome work in the
family was undertaken from the Western Himalayas (Mehra and Gill, 1968; Vasude-
van, 1975), South India (Subramanyam and Kamble, 1966), and Kashmir Himalaya
(Khoshoo and Tandon, 1963; Khoshoo et al., 1966; Koul and Gohil, 1973; Gohil
et al., 1981; Jee et al., 1985, 1989). Recently, the meiotic study was undertaken in
the three genera covering nine species of the family and revealed first ever chromosome counts in Gentiana aprica (2n ¼ 20), Gentiana argentea var. albescens
(2n ¼ 20), and Gentiana pygmaea (2n ¼ 20) and new/varied chromosome counts
in Swertia ciliata (2n ¼ 26) and Swertia purpurascens (2n ¼ 18) at the global level
(Bala et al., 2015). Recently, three species of Swertia viz. S. ciliata, S. cordata, and
S. petiolata have been worked out cytologically by our group. The results of this
investigation are given in Fig. 12.1 and discussed below:
S. ciliata (D. Don ex G. Don) B. L. Burtt. Meiotic analysis in the species
revealed a diploid cytotype with 2n ¼ 26 showing 13 bivalents at metaphase-I
and 13:13 equal distribution of chromosomes at each pole in anaphase-I
(Fig. 12.1; 1a,b). The present chromosome count of 2n ¼ 26 is in line with the
previous reports from India. Besides, the species is also known to exist with
2n ¼ 18, 20, and 24 in Indian accessions (Table 12.1).
S. cordata Wall. The species is reported to exist at diploid level with 2n ¼ 26
(Fig. 12.1; 2a), which is in line with the previous report from India (Table 12.1).
S. petiolata Royle. Cytological study of the species revealed a diploid cytotype
with 2n ¼ 26 showing 13:13 distribution of chromosomes at anaphase-I
(Fig. 12.1; 3a). The species is known with a single chromosome count of
2n ¼ 26, which has been confirmed in the present study as well (Table 12.1).
226 CHAPTER 12 Swertia chirayita
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FIGURE 12.1
1: Swertia ciliata, 1a: PMC at M-I showing 13 bivalents connections, 1b: PMC at A-I
showing 13 chromosomes at one pole. 2: Swertia cordata; 2a: PMC at A-I showing 13:13
distribution of chromosomes at both poles. 3: Swertia petiolata; 3a: PMC at A-I showing
13:13 distribution of chromosomes at each pole.
Out of a total of 170 taxonomically known species, the chromosome numbers
have been reported in 52 species with a range from 2n ¼ 14e60, with 12 different
chromosome numbers in between, which depict its polybasic nature (x ¼ 7, 8, 9, 10,
11, 12, 13), of which x ¼ 10, 12, and 13 are prominent ones. In India, 22 species are
cytologically known so far and chromosome number ranges from 2n ¼ 16e26, of
which 2n ¼ 26 being the most common, although x ¼ 8, 9, and 10 are also reported
in Indian accessions.
Table 12.1 Table showing names of taxons and chromosome numbers in cytologically worked out species at world level.
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Chromosome number in cytologically worked out species
Outside India India
Name of the taxon
1. Swertia abyssinica Hochst. / 18þ2B (Morton, 1993)
2. Swertia alata (Royle ex D. Don)
C.B. Clarke
3. Swertia albicaulis var.
albicaulis
Douglas ex Kuntze
4. Swertia angustifolia Burkill // 13 (Vasudevan,
5. Swertia beddomei C.B. Clarke /// 26 (Mallikarjuna,
6. Swertia bifolia Batalin / c.26 (Huang et al., 1996)
7. Swertia bimaculata (Siebold
and Zucc.) Hook. F.and
Thomson ex C.B. Clarke
8. Swertia calycina Franch. / 20 (Auquier and Renard, 1975)
9. Swertia chinensis Franch. / 20 (Wada, 1966)
10. Swertia chirata B. ham ex.
Wall.
n2n n2n
20 (Nemomisa, 1998)
13 (Khatoon
and Ali, 1993)
26 (Chambers
et al., 1998)
26 (Shigenobu 1982, 1983)11(Vasudevan,
/
/ 20, 24 (Wada, 1966)26(Khoshoo and
/ 13 (Vasudevan,
28 (He et al., 1999)
28 (Kawakami, 1930)
//
1975)
1975)
1975)
13 (Sharma, 1970)
Tandon 1963)
/
26 (Mallikarjuna,
1985)
1985; Mallikarjuna
et al., 1987)
/
/
13 ( Vasudevan, 1975)
Continued
12.2 Cytological studies of genus Swertia 227
Table 12.1 Table showing names of taxons and chromosome numbers in cytologically worked out species at world level.dcont’d
https://t.me/med1917
Chromosome number in cytologically worked out species
Outside India India
Name of the taxon
11. Swertia ciliata (D. Don) B. L.
Burtt
12. Swertia cordata (Wall. ex G.
Don) C.B. Clarke
13. Swertia corymbosa Wight ex
Griseb.
14. Swertia crassiuscula Gilg / 20 (Thulin, 1970) //
15. Swertia cuspidata (Maxim.)
Kitag.
16. Swertia densifolia
(Grisebach) Kashyapa
17. Swertia diluta (Turcz.) Benth.
and Hook. f.
18. Swertia emarginata Schrenk / 16 (Ma et al., 1990) //
19. Swertia engleri Gilg 10
20. Swertia fimbriata (Hochst.)
Cufod.
21. Swertia franchetiana HarrySm/ 20 (He et al., 1999) //
n2n n2n
// 09 (Bala and Gupta,
// 13 (Vasudevan,
// 26 (Mallikarjuna,
/ 28 (Shigenobu, 1983) //
//
/ 20 (Yuan and Ku¨pfer, 1993) //
(Nemomissa,
1998)
/ 26 (Nemomissa, 1998) //
13 (Mallikarjuna,
///
2011)
10 (Vasudevan,
1975)
12 (Mehra and Gill
1968)
13 (Malik et al., 2011)
1975)
1985)
1985)
2(Khoshoo and
Tandon, 1963)
24 (Mehra and Gill,
1968)
/
/
228 CHAPTER 12 Swertia chirayita
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