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166 CHAPTER 9 Rhodiola imbricata
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• UDP-glucosyltransferase
• Anabolism reactionetransfers a glucosyl to tyrosol
(the aglycon) and forms salidroside
• Catabolism reactionebreakdown salidroside to
glucose and tyrosol
Shi et al. (2007)
9.7.2 Rosavin
• Phenylalanine is a derivative of the shikimicechorismic acid pathway and
Phenylalanine ammonia lyase is the enzyme that directs the carbon atoms toward the phenylpropanoid metabolites biosynthesis.
• Both salidroside as well as the cinnamyl alcohol glycosides are the products of
phenylpropanoid metabolism which is derived from phenylalanine.
• Cinnamyl-CoA ester is synthesized from cinnamic acid by hydroxycinnamate:
CoA ligase.
• Cinnamy l-CoA reductase reduces cinnamyl-CoA ester to cinnamaldehyde.
• Cinnamy l alcohol dehydrogenase further reduces the cinnamaldehyde to cinnamyl alcohol.
• Rosin, which is the simplest glycoside of rose root, is formed by one glucose
transfer.
• Rosavin is formed from rosin by the addition of an arabinose, and rosarin is
formed from rosin by the addition of an arabinofuranose (Grech-Baran et al.,
2015).
9.8 Conclusion and future prospects
The recent years observed the peaked interests of scientific community and the
governmental agencies in conserving the traditional pharmacopoeia of the TransHimalayan regions. The myriad of assorted botanical flora of the region has been
extensively researched with respect to their prophylactic and therapeutic potential,
specially plants like R. imbricata. The species has been explored for its hidden treasures in solving multiple human diseases, for which it has also been referred as
“magical concoction” or “Sanjeevani” by a number of renowned scientists over
and over again. R.a imbricata has unprecedented capabilities in curing mankind
of diseases such as Alzheimer’s, cancer, renal cysts, edema of limb, burns, etc.,
along with the compounds offering immunostimulant, antifatigue, antidepressant,
antihypoxia, antioxidant, antiradioactive, and hepatoprotective properties. Researchers are not only striving toward achieving best for pharmaceutical sector
but also ensuring use of well-equipped tissue culture labs and high-end technologies
for sustainable use of the vegetation in order to conserve the endangered species.
Rhodiola holds strong foundation in food and lifestyle of local communities as an
easy answer to all health ailments and a part of many delicacies. Hence, in order

References 167
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to avoid its overexploitation or overharvesting by the farmers, the need of the hour is
to ascertain effective in situ conservation of the plant. Moreover, immediate aid and
assistance through collaborative projects between indigenous people and associated
experts of the national and international agencies for attaining choicest results are
also desired. All in all, balance is the key for achieving best out of this plant while
corroborating its presence in the wild and natural ecosystem in the future to come.
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CHAPTER
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Saussurea lappa
10
Ira Vashisht
Crop Genetics & Informatics Group, School of Computational and Integrative Sciences (SCIS),
Jawaharlal Nehru University, New Delhi, India
10.1 Introduction
Saussurea lappa (syn. Saussurea costus) C.B. Clarke is an endangered medicinal
plant, best known for its antiinflammatory, anticancerous, antiulcerogenic, and hepatoprotective properties (Butola and Samant, 2010; Gautam and Asrani, 2018;
Zahara et al., 2014). Traditionally known as “kuth” or “kushtha,” this high-
altitude perennial herb forms an integral part of Ayurvedic, Unani, Siddha, and
Tibetan (Amchi) medicine (Butola and Samant, 2010). Mention of S. lappa goes
back to writings from Mesopotamia where evidence of medicinal herbs has been
documented by Indus Valley people (Shah, 2019). R. Campbell Thompson was
successful in deciphering the multiple names of plants, drugs, and minerals used
in Assyrian botany (Mesopotamia). A clear mention of S. lappa import from India
has been observed for treatment of jaundice (Thompson, 1949). Its ancient role in
panduroga (jaundice) has been shown to echo in India as well (Dwivedi, 1963). It
is a member of the large Asteraceae or Compositae family, comprising of approximately 410 globally distributed species which are mostly native to cold and
temperate regions of Asia, Europe, and North America. Highest diversity has
been observed to inhabit alpine regions of Central Asia and Himalayas (Butola
and Samant, 2010).
S. lappa has become endangered due to overexploitation of its natural habitat for
diverse medicinal and commercial purposes. It is one of main ingredients in closely
71 drug formulations documented in The Handbook of Traditional Tibetan Drugs
(Tsarong, 1986), and in several popular polyherbal drug formulations like Chandra
Kalka, Ashtamangal Ghrita , and Sharkaradi Kalka (Chamara et al., 2018; Singh,
2019). Bitter roots having a sweet and strong aromatic odor are the most valued
part of S. lappa plants which are harvested for extraction of bioactive constituents.
Costus oil is the main compound extracted from roots which is constituted by a
number of secondary metabolites which have been prevalently used in medicine
and perfumery (Table 10.1)(Butola and Samant, 2010). Most of the research in
S. lappa is focus ed on investigation of bioactive constituents and corresponding therapeutic applications. A few scattered reports exist on attempts at large-scale cultivation and in vitro propagation of S. lappa for alleviating the endangered status of the
Himalayan Medicinal Plants. https://doi.org/10.1016/B978-0-12-823151-7.00012-X
Copyright © 2021 Elsevier Inc. All rights reserved.
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174 CHAPTER 10 Saussurea lappa
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Table 10.1 Constituents of Saussurea lappa essential oil or costus oil
(Zahara et al., 2014; Abdelwahab et al., 2019; Maurer and Grieder, 1997;
Dhillon et al., 1987).
S.
No. Compound
1 Dehydrocostus
lactone
2 Costunolide 21 (þ)-g-costol
3 8-cedren-13-ol 22 ()-elema-1,3,11 (13)-trien-12-ol
4 a-curcumene 23 ()-a-costal
5 b-costol 24 (þ)-g-costal
6 d-elemene 25 ()-Elema-1,3,11(13)-trien-12-al
7 a-selinene 26 ()-(E)-trans-bergamota-2,12-dien-14-al
8 b-selinene 27 ()-ar-curcumene
9 a-costol 28 ()-caryophyllene oxide
10 4-terpinol 29 12-methoxy dihydrodehydro costus lactone
11 Elemol 30 Eudesma-5,11(13)-dien-8,12-olide
12 a-ionone 31 Phenanthrenone
13 b-elemene 32 9,12-octadecadienoic acid (Z,Z)
14 ()-g-elemene 33 Cyclohexane
15 p-cymene 34 Germacra-1(10),4,11(13)-trien-12-oic acid,6a`-
16 2-b-pinene 35 Androstan-17-one, 3-ethyl-3-hydroxy-, (5a`)
17 ()-a-selinene 36 Bicyclo[10.1.0]tridec-1-ene
18 (þ)-selina-4,11-
diene
19 ()-a-trans-
bergamotene
S.
No. Compound
20 ()-a-costol
hydroxy-,c¸ -lactone, (E,E)
37 Naphthalene
38 4a,8-dimethyl-2-(prop-1-en-2-yl)-1,2,3,4,4a,5,6,7-
octahydronaphthalene
plant. Low germination potential has limited the application of breeding approaches
in this herb; however, promising OMICS strategies are being explored for understanding molecular networks underlying biosynthesis of industrially valuable
secondary metabolites. This chapter provides an elaborate account of botanical,
biochemical, and therapeutic characteristics of S. lappa along with a discussion
on challenges and efforts pertaining to conservation status and techniques which
can open avenues for yield improvement and engineering of valuable secondary
metabolites.
10.2 Botanical identification and classification
First botanical nomenclature for S. lappa was provided by Ainsile (Ainsile, 1813)
who attempted to identify koostum, which was known in various vernaculars as

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koot (Sanskrit), kust (Arabian), kostum (Tamil), and Sippuday (Malayalam) among
many others. Later, Royle (1839) discovered roots of this plant in Indian “bazar” and
went on to call it “koot” exported from Cashmere where it thrives on mountains in
areas surrounding Cashmere. Dr. Hugh Falconer encountered the plant in Drass,
Kashmir, during his expedition in 1834e38 and corresponded with Royale regarding
the medicinal plant, while naming it as a new genus, Costia (Shah, 2019). Later in
1845, Falconer went on to publish its botanical name as Aucklandia costus Falc.
which was well accepted for quite a while. Further, Decne changed its genus to
Saussurea and named it as S. lappa (Decne) Sch. Bip 1846 for the first time. Finally,
in 1964, its name was accepted as Saussurea costus (Falc) Lipch. However,
International Rules of Botanical Nomenclature have retained its multiple names
as Aplotaxis lappa (Decne) (1843), A. costus Falc. (1845), Aucklandia lappa Decne
(1875), and Theodorea costus Kuntze (1891) (The Plant List, 2013). The current
widely accepted nomenclature is S. lappa ((Decne.) C. B. Clarke.) as described
by the famous botanist Charles Baron Clarke (1876) (Waly, 2009).
10.3 Botany of Saussurea lappa
S. lappa is a tall and erect perennial herb having a stout and upright stem which is
1e2 m high (Pandey et al., 2007)(Fig. 10.1). Leaves are lobate, auricled at base, and
FIGURE 10.1
(a) Saussurea lappa plant in its natural habitat (Ladakh region). Scale bar ¼ 15 cm
(Warghat et al., 2016). (b, c) Dried roots and root powder of S. lappa (Amara et al., 2017).

176 CHAPTER 10 Saussurea lappa
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irregularly toothed which are scaberulous above, while glabrate from beneath. Basal
leaves are very long having winged stalks, whereas upper leaves are smaller, shortly
petioled or subsessile having small lobes which almost clasp the stem (Pandey et al.,
2007; Amara et al., 2017). The plant contains both cauline and radical leaves where
former are small, irregularly toothed, and pubescent, while latter are relatively quite
large and triangular having a long-winged petiole (Shah, 2019). While in the field,
another plant, Arctium lappa has often been mistaken for S. lappa owing to exhibition of similar large leaves (Shah, 2019). The plant is headed by bluish-purple to
black stalkless flowers which are clustered together in either terminal or axil part
of leaves. It has many ovate-lanceolate and pointed involucral bracts and receptacle
bristles are quite long (Pandey et al., 2007). Anther tails are fimbriate and corolla is
about 2 cm long. Achnes are small, curved, and compressed with narrowed tip and
pappus hair are brown and feathery (Pandey et al., 2007; Shah, 2019), while fruits
are cupped, compressed, and curved (Pandey et al., 2007). Most valuable part of
the plant is the root which is 15e30 cm thick and 40e60 cm long (Hajra et al.,
1995) having a camphoraceous aroma (Shah, 2019)(Fig. 10.1). Secondary roots
are often tubular and usually wrinkled and ridged (Shah, 2019). Roots are diced
into small pieces and dried which appear muddy gray to creamy in color and possess
a bitter taste. These dri ed roots of S. lappa are used as the crude drug available in the
market (Fig. 10.1).
10.4 Origin and geographical distribution
S. lappa is a prominent member of the extensive Himalayan flora, found at elevation
of 2700e4000 m amsl (Kaul, 1997; Nandkarni, 1954). Globally, the herb inhabits
the cool arctic and temperate regions of Asia, North America, and Europe (Shah.
2006; Hajra et al., 1995). It has been an indigenous constituent of traditional med-
icine in India, China, and Pakistan (Shah, 2019). In India, S. lappa has a natural
habitat in Jammu and Kashmir, Himachal Pradesh, and Uttarakhand (Kaul, 1997;
Nandkarni 1954). Kashmir and adjoining areas harbor the majority of its native
habitat where it can be found in Jhelum, Sonmarga, Drass, Kistwar, Zanskar valley
(Ladakh), Chenab valley, and Kishenganga (Shah, 2019). Further, to quench the
ever-increasing industrial demands, it is widely cultivated in Kashmir, Uttar
Pradesh, and Tamil Nadu (Shah, 1982; Kamalpreet et al., 2019). In Pakistan,
S. lappa can be observed on the moist, open hill slopes of the Himalayan region
in Leepa, Neelam, and Kaghan valleys, while a scattered presence can be seen in
valleys of Siran and Gurez along with Poonch and Bagh districts (Shah, 2019).
10.5 Biochemical/analytical properties
Roots are the primary source of the tremendous therapeutic potential of S. lappa.
Studies on the biochemical properties of the herb date back to 1950s which have
led to discovery of various act ive ingredients like terpenes, sesquiterpenoids,
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