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15.5 Major challenges in ashwagandha 281
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Table 15.1 List of some important therapeutic properties of Ashwagandha and their
effects’ studies.dcont’d
Therapeutic
potential
Potential to treat
male infertility
(improvement of
semen quality)
Prophylactic and
neuroprotective
potential
Experimental
model Effects of treatment References
Spermatozoa
and seminal
plasma from
infertile men
Male Sprague
Dawley rats
W. somnifera
treatment significantly
lowered ROS levels in
oligozoospermic
(n ¼ 25) and
asthenozoospermic
(n ¼ 25) men; lowered
the apoptosis in
normozoospermic
(n ¼ 25) and
oligozoospermic
(n ¼ 25) men. Also,
with treatment the
concentrations of
essential metal ions
were increased in
seminal plasma of
infertile men. Thus,
W. somnifera
treatment improved
semen quality and
could be used to treat
male infertility.
Ashwagandha (root
extracts) treatments
modulated the levels
of nitric oxide,
acetylcholine, the
activity of
acetylcholine
esterase, and
enhanced number of
pyknotic cells as well
as levels of
corticosterone in
hippocampal region of
rats. Therefore, the
root extracts of
Ashwagandha
ameliorated the
memory impairment
and
neurodegeneration
induced by hypobaric
hypoxia in male
Sprague Dawley rats.
Shukla et al. (2011)
Baitharu et al.
(2013)
Continued
282 CHAPTER 15 Withania somnifera
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Table 15.1 List of some important therapeutic properties of Ashwagandha and their
effects’ studies.dcont’d
Therapeutic
potential
Antioxidant and
antiinflammatory
agent
Cardiorespiratory
potential and
improvement of
quality of life
Neuroprotective
and antistress
potential; prolonged
life expectancy
Experimental
model Effects of treatment References
Rats Ashwagandha root
Male/female
athletic adults
Caenorhabditis
elegans
powder (600 mg/kg)
significantly reduced
the severity of arthritis
through suppression
of its symptoms and
effective restoration of
motor activity in the
arthritic rats.
Root extract of
Ashwagandha
enhanced the levels of
maximum oxygen
consumption
max) resulting in
(VO
2
improved
cardiovascular
dynamics and
enhanced
cardiorespiratory
endurance.
Furthermore, the
quality of life was
improved in healthy
male and female adult
athletics.
A bioactive
component of
W. somnifera,
withanolide A, has
antiaging, antistress,
neuroprotective
potential and
enhanced life span,
health span, neuron
functionality through
stimulation of insulin/
IGF-1 signaling
pathway in C. elegans.
Furthermore, it also
enhanced the
transcriptional level of
HSF-1 (heat-shock
transcription factor)
and SKN-1
(antioxidant response
transcription factor).
Gupta and Singh
(2014)
Choudhary et al.
(2015)
Akhoon et al. (2016)
15.5 Major challenges in ashwagandha 283
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Table 15.1 List of some important therapeutic properties of Ashwagandha and their
effects’ studies.dcont’d
Therapeutic
potential
Somnogenic (sleepinducing) potential
Neuroprotective
potential
Antistress potential
and moodenhancing
properties
Experimental
model Effects of treatment References
Male C57BL/6
mice
Transgenic mice
carrying G93A
sod1 mutant
Healthy male
and female
adults aged
between 18 and
65 years
Triethylene glycol
present in water
extracts of
W. somnifera induced
rapid and nonrapid
eye movement sleep in
mice. The quality of
sleep was also
enhanced and the
sleep-promoting
potential of triethylene
glycol has been
reported in mice.
Neuroprotective
properties of
W. somnifera root
extracts have been
reported on transgenic
mouse model
(expressing G93A
sod1 mutant) of
amyotrophic lateral
sclerosis with TDP-43
proteinopathy.
Ashwagandha
treatment resulted in
enhanced longevity,
better motor
coordination,
regulated expression
of cellular chaperons,
and improved number
of motor
neurons situated in the
lumbar spinal cord of
G93A
SOD1
Study on efficacy and
tolerability of Shoden
(standardized
Ashwagandha extract)
revealed that Shoden
treatments relieved the
anxiety, hormone
production, and stress
in healthy male and
females under mild
stress.
mice.
Kaushik et al. (2017)
Dutta et al. (2018)
Lopresti et al.
(2019)
Continued
284 CHAPTER 15 Withania somnifera
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Table 15.1 List of some important therapeutic properties of Ashwagandha and their
effects’ studies.dcont’d
Therapeutic
potential
Nephroprotective
potential
Anticancer potential Human prostate
Experimental
model Effects of treatment References
Albino Wistar
rats
cancer cell lines
(22Rv1 and
LNCaP)
Levels of serum urea
and creatinine
(biomarkers studied to
check the renal
toxicity) were
decreased with the
treatment of
W. somnifera root
extracts in nephrotoxic
(cisplastin-induced)
rats. This study
emphasized on the
nephroprotective
property of
Ashwagandha to treat
renal dysfunctions.
The ethanolic root
extracts of
W. somnifera inhibited
the fatty acid synthesis
in human prostate
cancer cell lines
through
downregulation of
expression of fatty
acid metabolism
erelated proteins.
Withania root extracts
also suppressed the
intracellular amounts
of total free fatty acids
and decreased the
levels of acetyl-CoA in
prostate cancer cell
lines, revealing the
anticancer efficacy of
Ashwagandha.
Rasheed et al.
(2020)
Kim et al. (2020)
15.5.1 Low seed viability
For the commercial crop production under field conditions the high rates of seed
germination, seed viability, and seedling survival are required. Although there is
an increasing demand of Ashwagandha in the herbal drug industry, the traditional
cultivation of Ashwagandha through its seeds has been limited due to low percentage
15.5 Major challenges in ashwagandha 285
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of seed viability, poor germination, and seedling survival; and moreover the
dormancy in the seeds is also a limiting factor in its cultivation (Niyaz and Siddiqui,
2014; Shanmugaratnam et al., 2013; Vakeswaran and Krishnasamy, 2003). In addi-
tion to these factors, the dependence of germination on both light and temper ature
(Kambizi et al., 2006; Khanna et al., 2013) further affects its commercial production.
To overcome these problems, several researchers tried and optimized various physicochemical treatments, microbial treatments, and the storage conditions to overcome the problem of low seed germination in Ashwagandha. The hydropriming
treatment has been found to be effective in enhancing the rate of seed germination
(Mehta and Raina, 2016). Siderophore producing Alcaligenes faecalis broth promoted the seed germination by 75% (Sayyed et al., 2007). It was also found that
inoculation of seed with plant growthepromoting rhizobacteria (PGPR)-selected
isolate not only enhanced seed germination under normal circumstances but also under UV-B exposure (Rathaur et al., 2012).
However, the low germination percentage of W. somnifera may be attributed to
seed dormancy but various treatments have been tried to break the dormancy.
Among the different treatments tried to break the seed dormancy including overnight
soaking in water and hot water soaking (Shanmugaratnam et al., 2013), a higher
germination percentage recorded after hot water soaking (52.75%) than soaking
overnight in water (33.75%). Himangini and Thakur (2018) tried various storage
conditions and recorded that storing the seeds at 0
nation. In another experiment performed to investigate the effect of different physicochemical treatments, storage, temperature, photoperiod, and growth regulators on
the seed germinability of Ashwagandha, it was recorded that the GA
(150 mg/mL) was most effective. However, the optimum conditions favoring germination were recorded as 25
C of temperature with continuous light, suggesting a significant role of photoperiod on seed germination (Khanna et al., 2013). Whereas
Niyaz and Siddiqui (2014) found that maximum increase in seed germination was
at 500 mg/L treatment of GA
and the heat treatment at 50C reduced the germina-
3
tion drastically. The prechilling treatment inhibits the seed germination in Ashwagandha and the content temperature of 25, 35, and 45
inhibits it (Kambizi et al., 2006).
When the effect of various nutritional sources was evaluated on seed germination
of W. somnifera, the nitrates of sodium, calcium, and potassium were much effective
in increasing the rate of seed germination (Ingle and Kareppa, 2012).
C for 4 months is best for germi-
treatment
3
C in dark completely
15.5.2 Pests and diseases
Despite the immense therapeutic properties of Ashwagandha, it is prone to a number
of pests and diseases which not only limit its production but have some implications
in the quality of the produce too. The biotic agents pose a big challenge to the quality
and quantity of the produce and could be bacterium, fungi, virus, nematode, insect,
mite, etc.
286 CHAPTER 15 Withania somnifera
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One of the most important pathogens is Alternaria alternata (Fr.) Keissler, that
causes leaf spot disease in W. somnifera and one of its related species W. coagulans
(Pati et al., 2008; Sharma et al., 2013b). This fungal pathogen not only induces some
biochemical, physiological, and ultrastructural changes in Ashwagandha but also affects the production of withanolides by lowering the expression of some key genes
of withanolide pathway (Sharma et al., 2011, 2014; Singh et al., 2017a,b). Therefore,
a sensitive detection system for the presence of A. alternata is highly desirable
(Sharma, 2013a) for quality control of herbal medicines prepared their off. Since
a variability exists in W. somnifera germplasm for the resistance toward
A. alternata (Meena et al., 2019; Mohammad and Shabbir, 2016), the resistant lines
can be used as a breeding material to transfer this trait in the elite chemotypes. When
different isolates of A. alternata were screened for their virulence, one of the isolates, i.e., Chempatti (I6) was found to be most virulent. Further, among the seven
fungicides tested, mancozeb (0.2%) was found to be most effective (Kalieswari
et al., 2016). Ashwagandha plants treated with the com bination of two species of
endophytic bacteria viz., Bacillus amyloliquefaciens and Pseudomonas fluorescens,
showed the lowest plant mortality rate in the presence of A. alternata stress (Mishra
et al., 2018).
Besides A. alternata, several diseases of W. somnifera have also been reported
that are caused by various classes of plant pathogens viz., bacteria, fungi, phytoplasma, viruses, nematodes, etc., that are also listed in Table 15.2.
Table 15.2 List of some common diseases of Withania somnifera, caused by
various types of plant pathogens.
Nature of
the
pathogen Pathogen Disease References
Fungi Alternaria alternata Leaf spot Pati et al. (2008);
Alternaria
dianthicola
Alternaria
chlamydospora
Alternaria solani e Alwadi and Baka
Fusarium
oxysporum
Fusarium solani Root rot and wilt/leaf
Cercospora
withaniae
Pseudoercospora
withaniae
Leaf blight disease Maiti et al. (2007,
Leaf blight disease Vanitha et al. (2006)
Wilt Sharma and Trivedi
spot
Leaf spot Chavan and Korekar
Leaf spot Singh (2012)
Sharma et al. (2011)
2012)
(2001)
(2010)
Chavan and Korekar
(2011); Gupta et al.
(2004)
(2011)
15.5 Major challenges in ashwagandha 287
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Table 15.2 List of some common diseases of Withania somnifera, caused by various
types of plant pathogens.dcont’d
Nature of
the
pathogen Pathogen Disease References
Pseudocercospora
fuligena
Pithomyces
chartarum
Colletotrichum
gloeosporioides
Puccinia withaniae Yellow rust El-Ariqi et al. (2009)
Choanephora
cucurbitarum
Myrothecium
roridum
Chaetomium
globosum
Phytoplasma Phytoplasma (16S
rVI group)
Virus Begomovirus
(Jatropha mosaic
India virus)
Tobacco leaf curl
virus
Eggplant mottled
dwarf virus
Nematode Meloidogyne
incognita
Meloidogyne
javanica
Black leaf spot Saroj et al. (2014)
Leaf spot Verma et al. (2008)
Leaf spot Solanki and
Wet rot Saroj et al. (2012)
Leaf spot Mahrshi (1986)
Black leaf lesions Shah and Daniel
Witches-broom disease,
little leaf symptoms
Yellow mosaic disease,
vein clearing, yellow net
or mild leaf curl
symptoms
Vain clearing, leaf rolling,
and vein banding
e Al-Musa and
Root knot disease Pandey and Kalra
Root knot disease Bhatti et al. (1974)
Basudeb (2017)
(2004)
Khan et al. (2006);
Samad et al. (2006);
Zaim and Samad
(1995)
Baghel et al. (2010,
2012)
Pathak and
Raychaudhuri
(1967); Singh and
Kumar (1998)
Lockhart (1990)
(2003); Saikia et al.
(2013)
In addition to these pathogens, many pest species are also known to affect the
health of W. somnifera plants. One of the major insect pests is 28-spotted beetle
Henosepilachna vigintioctopunctata. The larvae of this plant rapidly feed on its
leaves, leaving behind a fibrous skeleton, reducing the commercial value of the
plants (Sharma and Pati, 2011b). Efforts have been made to identify different parasitoids and fungal biocontrol agents for the eco-friendly management of this pest
(Jamwal et al., 2017; Sharma et al., 2012b; Venkatesha, 2006). Cowbug, Oxyrachis
288 CHAPTER 15 Withania somnifera
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tarandus, is also an important pest which mainly affects apical portions of the stems,
turning them brown, rough, and woody and the apical leaves are shed in severe infestation gradually dried and apical leaves were shed off (Sharma and Pati, 2011a). An
invasive mealybug, Phenacoccus solenopsis, was also found as one of the important
pests of W. somnifera, affecting mainly its leaves that can be deformed and shed prematurely (Sharma and Pati, 2013). The carmine red spider mite, Tetranychus urticae,
is the most impo rtant mite species affecting Ashwagandha. It is commonly found on
the aerial and more commonly on the apical parts of the infested shoots, commonly
feeding on the leaves, turning them shiny white in color, which gradually turn
brown, and are shed off at a later stage (Sharma and Pati, 2012).
Besides these important pests mentioned above, several other pests are known to
affect the cultivation of Ashwagandha as listed in Table 15.3.
Table 15.3 List of some major pest species infesting Withania somnifera.
Pest species
Henosepilachna
vigintioctopunctata
(Coleoptera: Coccinellidae)
Helicoverpa armigera
(Lepidoptera: Noctuidae)
Tetranychus urticae
(Trombidiformes:
Tetranychidae)
Phalantha phalantha Drary
(Nymphalidae: Lepidoptera)
Phenacoccus solenopsis
(Hemiptera:
Pseudococcidae)
Acherontia styx
(Sphingidae: Lepidoptera)
Oxyrachis tarandus
(Hemiptera: Membracidae)
Acherontia atropos
(Lepidoptera: Sphingidae)
Spilarctia oblique
(Lepidoptera: Arctiidae)
Tricentrus bicolor
(Membracidae: Hemiptera)
Eutetranychus orientalis
(Acarina: Tetranychidae)
Tricentrus sp.
(Membracidae: Hemiptera)
Coccidohystrix insolitus
(Pseudococcidae:
Hemiptera)
Affected plant part/
symptoms References
Leaves are fed by both larvae
and adults
Fruit borer Rehaman and Pradeep
Feed on leaves, turning them
shiny white in color
Young leaves are fed by larvae Gorain et al. (2012)
Leaves are deformed and shed
prematurely
Leaves are affected by
caterpillars
Apical portions of the stem
affected
Leaves are affected by
caterpillars
e Chandra (2004);
e Kumar et al. (2009c)
e Gupta and Karmakar
Tender stems are affected,
adults and nymphs both suck
the plant sap
Leaves are affected Ravikumar et al. (2008)
Sharma and Pati
(2011b); Sharma et al.
(2012b)
(2016); Rehaman et al.
(2018)
Sharma and Pati (2013)
Sharma and Pati (2013)
Singh and Kumar
(1998)
Sharma and Pati
(2011a)
Akkuzu et al. (2007)
Kumar et al. (2009a)
(2011)
Singh and Kumar
(1998)
15.5 Major challenges in ashwagandha 289
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Table 15.3 List of some major pest species infesting Withania somnifera.dcont’d
Pest species
Drosicha mangiferae (green)
(Pseudococcidae:
Hemiptera)
Nezara viridula Linn.
(Heteroptera:
Pentatomidae)
Plautia sp. (Hemiptera:
Pentatomidae)
Dysdercus cingulatus
(Fabr.) (Hemiptera:
Pyrrhocoridae)
Trialeurodes vaporariorum
(Westwood) (Homoptera:
Aleyrodidae)
Diacrisia oblique (Arctidae:
Lepidoptera)
Lygeus equistris
(Lygaeidae: Hemiptera)
Hieroglyphus banian
(Acrididae: Orthoptera)
Trilophida annulata
(Acrididae: Orthoptera)
Aphis craccivora
(Aphididae: Hemiptera)
Coccus sp. (Coccidae:
Hemiptera)
Oxycarenus hyalinipennis
(Lygaeidae: Hemiptera)
Pseudococcus sp.
(Pseudococcidae:
Hemiptera)
Agromyza sp.
(Agromyzidae: Diptera)
Nisotra striatipennis
(Chrysomelidae:
Coleoptera)
Polyphagotarsonemuslatus
sp. (Tarsonemidae: Acari)
Myzus persicae (Hemiptera:
Aphididae)
Planococcus citri (Risso)
(Hemiptera:
Pseudococcidae)
Affected plant part/
symptoms References
e Bhagat (2004); Kumar
e Kumar et al. (2009b)
e Kumar et al. (2009b)
e Kumar et al. (2009b)
e Kumar et al. (2009b)
Leaves are affected by
caterpillars
Leaves are affected, adults and
nymphs both suck the plant sap
Leaves are eaten by adults and
nymphs both
Leaves are eaten by adults and
nymphs both
Leaves and tender stems are
affected, insects suck the plant
sap
Leaves are affected, insects
suck the plant sap
Leaves are affected. Both
adults and nymphs suck the
plant sap
Leaves and other tender shoots
are affected. Insects suck the
plant sap
Leaves are affected (mined) Singh and Kumar
Leaves are fed by adults Singh and Kumar
Leaves are affected and
defoliation occurs
e Kumar et al. (2009c)
e Attia and Awadallah
et al. (2009b)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
Singh and Kumar
(1998)
(1998)
(1998)
Singh and Kumar
(1998)
(2016)
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15.5.3 Quality control in herbal medicine
Because of the use of W. somnifera in several herbal drug formulations (as singleand multiherbal products), the stringent measures to ensure the quality of the herbal
drugs and the adoption of good manufacturing practices for the preparation of herbal
drugs are a challenge. The quality control of medicinal plants has emerged as an
important aspect in herbal drug industry and there are strict international guidelines
for this too (WHO, 2007).
A study was conducted to analyze the major bioactive constituents like withaferin A, in commercially available herbal formulation derived of Ashwagandha,
in the Indian market, a huge variation (more than 70-fold) in the daily intake of withaferin A have been recorded (Sangwan et al., 2004). Age of the plant also significantly affects the metabolic contents of the plant; therefore, the harvest at correct
stage is also very important to get a consistent performance of the herbal drugs
derived their off and could be responsible for the huge variation in the amount of
bioactives. Withaferin A content increases in both the leaves and the roots with
the age of plant (up to 12e18 week old), but withaferin A content (along with
some other withanolides) is declined on the overmaturation (Dhar et al., 2013 ; Pal
et al., 2011). Further the microbial pathogens like A. alternata can reduce the con-
centration of bioactives present in Ashwagandha too (Pati et al., 2008); therefore, the
management of the disease and development of an efficient detection system for the
disease becomes even more important.
15.6 Phytochemicals
The phytochemistry of Ashwagandha has been investigated extensively. There are
several families of plant secondary metabolite which have been found in it, including
the alkaloids, flavonoids, steroidal lactones, tannin, etc. Out of these the steroidal lactones with a basic C-28 framework are the signature bioactive molecules, identified in
W. somnifera, called as withanolides. The basic structure of withanolides is represented
as Fig. 15.1a and structures of three key withanolides (withanolide A, withaferin A,
and withanone) are represented as Fig. 15.1bed. The steroidal lactones present in
W. somnifera are ashwagandhanolide, withanolide AeY, withaferin A, withasomidie-
non, withasomniferin, withanone, withasomniferols, and a novel chlorinated withanolide namely 6a-chloro-5b,17a dihydroxywithaferin A (Dar et al., 2016; Singh et al.,
2015; Tripathi et al., 2018). The major phenolics present in Ashwagandha include
quercetin, kaempferol, cholorogenic acid, gallic acid, ellagic acid, tannic acid, caffic
acid, and rutin. Besides, steroidal lactones and phenolics, W. somnifera has
glycowithanolides-like sitoindoside (VIIeX), sterols, withanol, somnitol, somnisol,
cholesterol, diosgenin, stigmastadien, stigmasterol, b-sitosterol, alkaloids, and
flavonol glycosides (Ba¨hr and Ha¨nsel, 1982; Bhattacharya et al., 1997; Chatterjee
et al., 2010; Chaurasiya et al., 2009, 2012; Ghosal et al., 1988; Mirjalili et al., 2009;
Sangwan et al., 2008; Singh et al., 2015; Sivanandhan et al., 2013; Xu et al., 2011).
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