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Abstract

Common mullein (Verbascum thapsus L.) is a medicinal plant readily found in roadsides, meadows and pasture lands and has been used to treat pulmonary problems, inflammatory diseases, asthma, spasmodic coughs, diarrhoea and migraine headaches. Although it has been used medicinally since ancient times, the popularity of common mullein has been increasing commercially for the past few years. Today, the dried leaves and flowers, swallow capsules, alcohol extracts and the flower oil of this plant can easily be found in health stores in the United States. The use of common mullein extracts in folk medicine begun recently to be supported by an increasing number of research studies. This paper thoroughly reviews all the scientific research related to Verbascum thapsus L. including plant tissue cultures and the biological properties of this plant.
COMMON MULLEIN (VERBASCUM THAPSUS L.) 733
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
Copyright © 2005 John Wiley & Sons, Ltd.
PHYTOTHERAPY RESEARCH
Phytother. Res. 19, 733– 739 (2005)
Published online in Wiley InterScience (www.interscience.wiley.com). DOI: 10.1002/ptr.1653
REVIEW ARTICLE
Common Mullein (Verbascum thapsus L.):
Recent Advances in Research
Arzu Ucar Turker and Ekrem Gurel*
Abant Izzet Baysal University, Faculty of Science and Arts, Department of Biology, 14280 Bolu, Turkey
Common mullein (Verbascum thapsus L.) is a medicinal plant readily found in roadsides, meadows and
pasture lands and has been used to treat pulmonary problems, inflammatory diseases, asthma, spasmodic
coughs, diarrhoea and migraine headaches. Although it has been used medicinally since ancient times, the
popularity of common mullein has been increasing commercially for the past few years. Today, the dried
leaves and flowers, swallow capsules, alcohol extracts and the flower oil of this plant can easily be found in
health stores in the United States. The use of common mullein extracts in folk medicine begun recently to be
supported by an increasing number of research studies. This paper thoroughly reviewes all the scientific
research related to Verbascum thapsus L. including plant tissue cultures and the biological properties of this
plant. Copyright © 2005 John Wiley & Sons, Ltd.
Keywords: Verbascum thapsus L.; mullein; medicinal herb; Scrophulariaceae.
Received 25 February 2004
Accepted 14 December 2004
* Correspondence to: Dr E. Gurel, Abant Izzet Baysal University, Faculty
of Science and Arts, Department of Biology, 14280 Bolu, Turkey.
E-mail: gurel_e@ibu.edu.tr
INTRODUCTION
The Scrophulariaceae family is an important family
of plants comprising over 200 genera and about 2500
species. They occur mostly in temperate and subtrop-
ical regions, and many of them produce flowers of great
beauty in either a garden setting or as roadside ‘weeds’.
The family includes Mimulus, Penstemon, Digitalis,
Veronica and Verbascum (Grieve, 1981). A number of
the Scrophulariaceae are, or have been, valued for their
curative properties and are widely employed both in
domestic and regular medicine. At least 250 species
of Verbascum are known. Of the species formerly used
in medicine, the most important is Verbascum thapsus
L. Common names include mullein, common mullein,
great mullein, wooly mullein, candlewick plant, velvet
plant, blanket leaf, white mans footsteps, Aaron’s
rod, Jacob’s staff, hedge taper, high taper, old man’s
flannel, lady’s foxglove and sı˝ır kuyru˝u (in Turkish)
(Strange, 1977).
The generic name of this plant, Verbascum, is
believed to be a corruption of barbascum, from the
Latin barba, meaning a beard, referring to the shaggy
appearance of the genus, while thapsus, its specific
name, may refer to the Greek island of that name,
where the species originally thrived. The word ‘mullein’
comes from the Middle English moleyne and the Old
French moleine, and originally from the Latin mollis,
meaning ‘soft’ and referring to the leaves (Strange,
1977).
BOTANY
Verbascum thapsus L. is a biennial, or rarely an annual
plant, with a deep tap root. In its first year, it produces
a low vegetative rosette up to 60 cm in diameter, which
overwinters and is followed in the succeeding growing
season by a stout flowering stem 0.3–2.0 m tall (Gross
and Werner, 1978). The basal leaves are oblong-obovate
to obovate-lanceolate and 1040 cm long including the
petiole.
Cauline leaves are elliptic-lanceolate, decurrent,
gradually reduced up the stem and densely woolly with
branched hairs (Millspaugh, 1974). The leaf system is
so arranged that the smaller leaves above drop the rain
onto the larger ones below, which direct the water to
the roots. This is a necessary arrangement since mullein
grows mostly on dry soils. The stellately branched hairs,
which cover the leaves so thickly, act as a protective
coat, thus reducing moisture loss, and also providing a
defence. They prevent attack by creeping insects and
set up an intense irritation in the mucus membrane of
any grazing animals that may attempt to browse upon
them. The hairs are not confined to the leaves alone,
but are also on every part of the stem, on the calyces
and on the outside of the corollas, so that the whole
plant appears whitish or grey (Gross and Werner, 1978;
Whitson, 1991). According to Muzik (1970) these epi-
dermal hairs hold away the droplets from the leaf sur-
face so that they can protect the leaves from an aqueous
solution of 2,4-D. The homely but valuable ‘mullein
tea’, a remedy of the greatest antiquity for coughs and
colds, must always be strained through fine muslin to
remove any hairs that may be floating in the hot water,
which was poured over the flowers or leaves. They can
cause intolerable itching in the mouth (Grieve, 1981).
The inflorescence is a spike-like raceme 20–50 cm
long and approximately 3 cm in diameter. It is usually
very dense; rare axillary racemes may arise from the
734 A. U. TURKER AND E. GUREL
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
upper leaves. The sessile flowers are usually one per
axil with pedicels less than 2 mm and slightly irregular
with rotate corollas (Millspaugh, 1974). Individual
flowers of mullein are ephemeral, opening before dawn
and closing before mid-afternoon of the same day. They
are protogynous, the style maturing first and then bend-
ing downward once the anthers appear. The flowers
are also autogamous, self-pollination occurring at the
end of the day if cross-pollination has not occurred.
The style returns to its original position and the corolla
closes, pushing the still receptive stigma against the
anthers. The calyx consists of five lanceolate or ovate
sepals, 7–9 mm long with caudate tips. The corolla
is 20–25 mm broad consisting of five yellow (rarely
white) petals. Stamens are irregular and attached
to the corolla, three upper filaments are shorter and
densely white-villous, the lower two are longer and
glabrous. Anthers are larger and coloured. The ovary
is superior and two-celled. The fruit is an ovoid, stellate-
pubescent, capsule 36 mm long, longer than the calyx
and splits into two valves at maturity. There are numer-
ous brown seeds, 0.5–1.0 mm long which are six-sided
and have angular lateral surfaces with rows of pits
(Abrams, 1951; Davis, 1965–1985; Munz and Keck, 1973;
Gross and Werner, 1978; Radford et al., 1968). Chro-
mosome counts from plants collected in Ottawa and
British Colombia gave 2n = 36 (Packer, 1964; Mulligan,
1961). Other counts from European material have given
2n = 34 and 2n = 36 (Darlington and Wylie, 1955; Love
and Love, 1961) and n = 9, 11, 17 (Love and Love, 1961).
DISTRIBUTION
V. thapsus L. is native to Europe and Asia. It was prob-
ably introduced into North America several times as a
medicinal herb. It was introduced in the mid-1700s to
Virginia as a piscicide (fish poison) and spread rapidly
(Semenza et al., 1978). In Turkey, common mullein is
distributed in Black Sea region (Kastamonu, Ordu,
Trabzon, Rize, Çoruh) and commonly found in river-
sides, forests, Corylus and Quercus scrub and volcanic
tuff (Davis, 1965–1985). V. thapsus L. grows wild
on stony ground in wasteland, woodland clearings
and roadsides. It does well in shallow, well-drained,
nitrogen-rich soils (Semenza et al., 1978). It occurs in
areas where the mean annual precipitation is 50–150 cm
and the growing season is at least 140 days (Gross and
Werner, 1978).
MEDICINAL OR HISTORICAL USES
Historically, mullein has been used as a remedy for the
respiratory tract, particularly in cases of irritating coughs
with bronchial congestion (Hoffman, 1988). Mullein
leaves and flowers have expectorant and demulcent
properties which are used by herbalists to treat respira-
tory problems such as bronchitis, dry coughs, whooping
cough, tuberculosis, asthma and hoarseness (Grieve,
1981; Mabey, 1988; Berk, 1996). The flowers are mildly
diuretic and have a soothing and antiinflammatory
effect on the urinary tract (Mabey, 1988). The leaves
are also diuretic, helping to reduce inflammation of the
urinary system and to counter the irritating effect of
acid urine (Ambasta, 1986; Tyler, 1993; 1994). Some
herbal texts extend the therapeutic use to pneumonia
and asthma (Grieve, 1981).
The leaves, roots and the flowers are also anodyne,
antiseptic, antispasmodic, astringent, emollient, nervine,
vulnerary, analgesic, antihistaminic, anticancer, anti-
oxidant, antiviral, bacteristat, cardiodepressant, oestro-
genic, fungicide, hypnotic and sedative (Lucas, 1969;
Harris, 1972; Null and Null, 1972; Grieve, 1981).
The demulcent and emollient properties come from
the polysaccharide mucilage and gums that soothe the
irritated tissue. The expectorant property is the result
of saponins that stimulate fluid production. The anti-
inflammatory property is due to iridoid glycosides and
flavonoids that decrease inflammation (Grieve, 1981).
The mullein combines the expectorant action of its
saponins with the soothing effect of its mucilage, mak-
ing this a most useful herb for the treatment of hoarse-
ness, tight coughs, bronchitis, asthma and whooping
cough (Mabey, 1988). The dried leaves are sometimes
smoked in an ordinary tobacco pipe to relieve the irrita-
tion of the respiratory mucus membrane, and will
completely control the hacking cough of consumption.
The leaves are employed with equal benefit when made
into cigarettes, for asthma and spasmodic coughs. The
flowers placed in a bottle and set in the sunshine are
said to yield a fatty matter valuable as a cure for haem-
orrhoids. Fomentations and poultices of the leaves have
been found serviceable in haemorrhoidal complaints.
Mullein is said to be of much value in diarrhoea, from
its combination of demulcent with astringent properties,
this combination strengthening the bowels at the same
time (Grieve, 1981; Mabey, 1988). In Europe, a sweet-
ened infusion of the flowers strained in order to separ-
ate the rough hairs is used as a domestic remedy for
mild catarrhs and colic. A conserve of the flowers has
also been employed against ringworm, and a distilled
water of the flowers was long reputed to be a cure for
burns and erysipelas (Millspaugh, 1974; Grieve, 1981).
A decoction of leaves was used as a hearth stimulant.
A decoction of roots febrifuge is used to alleviate tooth-
ache and also to relieve cramps, convulsions and
migraines. The juice of the plant and powder made
from the dried roots is said to quickly remove rough
warts when rubbed on them (Tyler, 1993; 1994). An oil
produced by macerating mullein flowers in olive oil,
stored in a corked bottle during prolonged exposure
to the sun, or by keeping it near the fire for several
days, is used as a local application in country districts
in Germany for piles and other mucus membrane
inflammations, and also for frost bite and bruises.
Mullein oil is recommended for earache and discharge
from the ear, and for any eczema of the external ear
and its canal (Mabey, 1988; Yarnell, 1997). Mullein oil
is a valuable destroyer of disease germs (Chopra et al.,
1956; Milspaugh, 1974). The fresh flowers, steeped for
21 days in olive oil, are said to make an admirable
bactericide. An alcohol tincture is prepared by homoeo-
pathic chemists, from the fresh herb with spirits of wine,
which has proved beneficial for migraines or sick head-
aches of long standing, with oppression of the ear
(Bianchini and Corbetta, 1977; Lewis and Elvin-Lewis,
1977). The seeds of mullein are said to be toxic and
should not be used in any of these preparations (Berk,
1996). The seeds when thrown into the water are said
COMMON MULLEIN (VERBASCUM THAPSUS L.) 735
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
to intoxicate fish, and are used by poachers for that
purpose, being slightly narcotic. Major toxic elements
affecting the circulatory, respiratory and central nervous
systems of the fish include saponin, rotenone and glyco-
side. The common mullein causes fish to have difficulty
in breathing (Wilhelm, 1974).
Swedish settlers called it wild tobacco and tied
the leaves around their feet and arms when they
had the ague. Some prepared a tea from the leaves
for dysentery. A decoction of the roots was injected
into the wounds of cattle when afflicted with worms,
which caused them to die and fall out. Also some
American Indian tribes (Mohegans, Penobscots,
Catawbas, Chochtaws, Creeks, Forest Potawatomis and
Menominees) used common mullein as a medicinal
herb (Moerman, 1986). The Mohegans smoked them
to relieve asthma and sore throat, and the Penobscots
smoked the dried and powdered leaves for asthma. The
Catawbas boiled the root and sweetened it to make
syrup for croup in children. The leaves were mashed
and applied as a poultice for pain and swelling, sprains,
bruises and wounds. The Chochtaws put the leaves on
the head as a headache poultice. The Creeks boiled the
roots with those of Button Willow for a drink used for
coughs. The leaves were also boiled and the patient
bathed in the infusion while it was hot. The Forest
Potawatomis smoked the dried leaves for asthma, but
it is not certain whether they learned the practice from
the whites or vice versa. A smoke smudge was made of
the leaves and the fumes inhaled for catarrh and to
revive an unconscious patient. The Menominees smoked
the root for pulmonary diseases. Whites smoked the
leaves for asthma and bronchitis and that the flowers
were believed to be diuretic and had been used for
tuberculosis (Moerman, 1986; Vogel, 1990).
The flowering stem was used dried by Greeks and
Romans as a taper dipped in tallow for light. Mullein
torches were said to repel witches. There is evidence
that at one time it was a ‘magical plant’ of the ancients.
Agrippa, a general and minister under Caesar Augustus,
claimed that the scent from the leaves had an over-
powering effect on demons. Mullein was thought to be
an ingredient in brews and love potions, and mentioned
in incantations used by witches during the Middle Ages.
The women of Rome also infused the flowers and mixed
the resulting liquid with lye, using it as a wash to turn
their hair golden yellow (Strange, 1977).
BIOLOGICAL STUDIES
Williams and Kemp (1976) showed that seedlings of
V. thapsus L. collected from a range of cold to warm
temperature habitats (based on altitude and latitude)
exhibit similar rates of photosynthesis within a tem-
perature range 20°–35 °C. Only at the highest tempera-
ture tested, 40 °C, the seedlings from the warmest
habitat (low altitude and latitude) exhibit higher photo-
synthetic rates than those from the coolest habitat.
They concluded that the ability of an individual plant
to photosynthesize over a broad range of temperature
has contributed to mullein’s success across a diversity
of habitats (Gross and Werner, 1978).
Williams et al. (1975) reported a CO2 compensation
point of 58 vpm CO2 for V. thapsus L. and on this basis
concluded that the species had a C3 photosynthetic
pathway. Wuenscher (1970) shaved the dense trichomes
of the leaves of V. thapsus L. and found that the
unshaved half of the leaf was consistently warmer than
the shaved half. The hairs must, then, affect the leaf
energy exchange, since two halves of the same leaf,
differing only in the presence of hairs, reached dif-
ferent equilibrium temperatures. Convection and the
latent heat loss are reduced by the hairs although
the hairs have little effect on radiation absorption. The
transpiration rate is reduced in hairy leaves. All of these
effects are explained by an increase in boundary layer
thickness caused by the hairs. The boundary layer forms
above the surface of the hair coating rather than di-
rectly over the leaf surface. This thickens the boundary
layer by the distance to which the hairs extend from
the leaf surface. Increased transpiration resistance has
obvious ecological significance for V. thapsus L., which
grows on dry, exposed sites. The dense hair coating
is an efficient water-conserving mechanism. Parkhurst
(1976), evaluating the work of Wuenscher (1970),
showed by calculation that the main effect of the
trichomes was to increase stomatal resistance with only
a slight increase in boundary layer resistance.
Lortie and Aarssen (1997) demonstrated that
clipping the shoot apex of V. thapsus L. resulted in
significantly more branches, and the branching inten-
sity could not be increased by greater resource levels
in mullein when the apical meristem was intact. Branch-
ing was stimulated by the addition of nutrients only
when the shoot apex was damaged. These results indi-
cated that nutritional status does not solely determine
the degree of branching expressed in mullein.
Glier and Caruso (1973) demonstrated that decreas-
ing temperatures were observed to induce starch
degradation in the roots of mullein. Virtually no starch
remains in the roots when the acclimation period of
decreasing temperature was followed by an extended
period of exposure to 4 °C. The breakdown of starch
which was presumed to occur in late autumn under
field conditions may provide cryoprotective chemicals
for the over-wintering rosette. Such chemicals might
also serve as an energy source for the bolting pro-
cess, which occurs during the following spring or early
summer. Later, they showed that the starch content in
roots of V. thapsus L. was reduced when rosettes were
exposed to low temperatures because of the increased
activities of the starch degradative enzymes (Glier and
Caruso, 1974). They also (Glier and Caruso, 1977b)
reported that two nonspecific enzymes, namely, acid
and alkaline phosphatase increased their activities in
Verbascum roots exposed to decreasing temperatures;
alkaline phosphatase was of more importance than acid
phosphatase in over-wintering rosettes. It was known
that mullein had a cold-requirement for bolting and
that applied gibberellin substituted for this requirement
(Caruso and Glier, 1970). Glier and Caruso (1977a)
also described the influence of gibberellin on the act-
ivities of starch degradative enzymes and phosphatase.
There was a sharp decrease in starch in the roots of
Verbascum as a result of treating rosettes with GA3
and all three starch degradative enzymes revealed
increases in their activities. Also, application of GA3
resulted in an increase in the activity of alkaline
phosphatase in roots and rosettes showed an early
response to applied gibberellin.
736 A. U. TURKER AND E. GUREL
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
Gross (1981) showed that the rosette sizes of V.
thapsus L. gives a reliable estimate of an individual’s
fate the following year. A minimum size must be
reached before a plant is capable of flowering and the
probability of flowering increases steadily with rosette
size. For mullein, all rosettes with a diameter greater
than 41 cm flower the subsequent year and rosettes less
than 9 cm in diameter do not flower. Conversely, the
probability of death decreases with increasing rosette
size.
According to Reinartz (1980; 1984a), plant size has
a large impact on plant fitness; within V. thapsus
L. populations large plants develop faster, flower ear-
lier and produce more seed. Compared with other
monocarpic species, V. thapsus L. has an extremely low
reproductive effort (11%–23% of biomass) and seed
output (4%–8%). The reproductive efforts and seed
output of other monocarpic species average 40%
and 25%, respectively (Reinartz, 1984b). Differences
between cohorts of mullein in plant size and the year
of flowering were primarily environmentally induced;
however, the variation between cohorts in a number of
other characters appeared to have a genetic basis
(Reinartz, 1984c).
The seeds of V. thapsus L. are contained in a capsule
with two cells. Salisbury (1942) reported that the mean
number of capsules per explant was 226 ± 42 (SD; n =
37), with an average of 596 ± 30 (SD; n = 16) seeds per
capsule. This gives an approximate average of 10 000
180 000 seeds per individual plant, each seed averaging
0.067 mg (Gross, 1980; Gross and Werner, 1982). The
seeds possess no specialized morphological adaptations
for dispersal by wind or animals. The capsules split open
along the longitudinal axis when mature, and move-
ment of the stalk by wind or a large animal is required
to release the seeds from the parent. Seeds are dis-
persed as far as 11 m, although 93% of them fall within
5m and 75% of them fall within 1 m of the parent
plant. Darlington and Steinbauer (1961) recorded that
the seeds of mullein remain viable for up to 35 years
but Kivilaan and Bandurski (1981) noticed that seeds
may remain viable for over 100 years and in Denmark
viable seeds of V. thapsus were collected from soil
samples archaeologically dated from 1300 A.D.
(Ødum, 1965).
Mullein seeds may germinate under a wide variety
of environmental conditions. Germination is completely
inhibited below 10 °C and at constant temperatures
above 40 °C (Semenza et al., 1978). Chilling during
the winter lowers the temperature requirement for
germination; thus, seeds brought to the surface in the
autumn by soil disturbance are able to germinate early
the next spring (Baskin and Baskin, 1981). Semenza
et al. (1978) found that only 35% of seeds germinated
in the dark, compared with 93% germination in the
light. This light sensitivity varies seasonally, but on
the whole, only those seeds, which lie at, or near the
soil surface (0.5 cm or less) will be able to germinate. If
seed burial occurs due to subsequent disturbance or
heavy rains rapidly sifting the seeds below the soil
surface, germination may be reduced or prevented
through light deprivation (Gross, 1980).
There are some natural enemies (insects and patho-
gens) of common mullein such as the curculionid
weevil (Gymnaetron tetrum Fab.) which is specific to
V. thapsus L. and was introduced to North America
from Europe (Burcham, 1937). The larvae mature in
the capsules (Sleeper, 1954) and destroy up to 50% of
the seeds. Weevils were responsible for the predation
of 57% of mullein capsules (Pottmeyer, 1985). The other
insect, the mullein moth (Cucullia verbasci) feeds and
develops on mullein species. There are some patho-
gens which cause disease in common mullein such
as Erysiphe cichoracearum (powdery mildew) and
Phymatotrichum omnivorum (root rot) (Gross and
Werner, 1978).
Although extracts from mullein can inhibit the growth
of wheat seedlings, it is not a serious agricultural weed,
since it can be controlled by cultivation. In overgrazed
or poor pastures, the presence of common mullein rep-
resents a further degradation of the pasture because
grazing animals avoid eating mullein. This species is
not allelopathic, allergenic or poisonous to humans
(Gross and Werner, 1978).
Filippini et al. (1990) examined the commercial pow-
dered samples of V. thapsus L. flowers and powdered
samples of V. thapsus L. cultivated in the Botanical
Garden of the University of Padua by using SEM.
Contamination of a commercial powdered sample of
V. thapsus L. flowers with plant material not belonging
to a Verbascum species was observed.
Vegetative reproduction does not occur in V. thapsus
L. (Gross and Werner, 1978; Gross, 1980).
Caruso (1971) showed that excised internodal seg-
ments of flowering specimens of V. thapsus L. with
vascular tissues grew and formed numerous buds on a
simple nutrient medium which lacked added growth
regulator. Pith explants without vascular tissues became
brown in a matter of 2 to 3 weeks with no visible sign
of growth. Endogenous growth regulators supplied by
vascular tissues were believed to be major factors in
bud formation in excised internodal segments of this
species.
An in vitro culture protocol for common mullein
was established by Turker et al. (2001). Explants (leaf
discs, petioles and roots) were cultured on Murashige
and Skoog minimal organics (MSMO) medium with
benzyladenine (BA) or kinetin (KIN). The best shoot
proliferation was obtained from leaf discs and petiole
explants with 3 mg /L BA. Leaf discs were cultured
on MSMO medium with 3 mg /L BA in combina-
tion with naphthalene acetic acid (NAA) or 2,4-
dichlorophenoxyacetic acid (2,4-D). More shoot
development was obtained with 3 mg / L BA and
1mg/L NAA. Shoots were transferred to rooting
media containing different levels of NAA or 2,4-D. Most
shoots developed roots on medium with 1 mg/L NAA.
Plants were transferred to vermiculite and subsequently
to potting media and maintained in a greenhouse.
ACTIVE INGREDIENTS OF V. THAPSUS L.
The constituents of V. thapsus L. include poly-
saccharides; iridoid glycosides including harpago-
side; harpagide and aucubin (especially in the leaf);
flavonoids, including 3 methylguercitin, hesperedin and
verbascoside; saponins and volatile oils (Pascual Teresa
et al., 1978a; 1978b; 1980; Hattori and Hatanaka, 1958;
Khuroo et al., 1988; Mehrotra et al., 1989; Warashina
et al., 1991; 1992).
COMMON MULLEIN (VERBASCUM THAPSUS L.) 737
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
Pascual Teresa et al. (1978a) isolated veratric acid
and
α
-spinasterol from the benzene extract of capsules
of V. thapsus L. From the hydrolysed ethanol extract,
the triterpene A, saikogenin A, benzyl alcohol and
methylfurfural were isolated. They also showed that
the seed oil from mullein (benzene extract) had the
following components: fatty acids: palmitic, steriac, oleic,
linoleic, linolenic, arachic and behenic. Unsaponifiable
matter:
β
-sitosterol and ergosta-7-en-3-
β
-ol (Pascual
Teresa et al., 1978b). Phenylethanoid and lignan
glycosides (Warashina et al., 1992), sterones, iridoid
glycosides and sesquiterpene acid (Khuroo et al., 1988;
Warashina et al., 1991) and verbacoside, a new luteolin
glycoside (Mehrotra et al., 1989) were obtained from
whole plants of V. thapsus L. Bourquelot and Bridel
(1910) discovered the verbascose, an oligosaccharide in
the root of the mullein. Hattori and Hatanaka (1958)
demonstrated the oligosaccharides in V. thapsus L. and
distribution of mono- and oligosacharides in various
organs of the plant in various stages was studied.
Pascual Teresa et al. (1980) isolated four saponins
from the capsules of V. thapsus L., thapsuine A,
thapsuine B, hydroxythapsuine A and hydroxythapsuine
B with a chromatographic separation on silica gel (TLC).
Crushed capsules (5 kg) were extracted with benzene
and ethanol and subjected to chromatography on silica
gel and sephadex. They obtained 300 mg thapsuine A
and 460 mg thapsuine B.
Bom et al. (1998) purified
α
-galactosidase from the
roots of V. thapsus L. using hybrid affinity chromato-
graphy with good recovery.
An extraction and analytical protocol for saponins
was established for Verbascum thapsus L. (Turker
et al., 2003). Four different kinds of plant sample were
analysed; (i) field-grown, (ii) in vitro cultured, (iii) com-
mercially obtained leaves and (iv) field-grown capsules.
A cleanup procedure with octadecyl (C18) solid phase
extraction columns was used prior to HPLC (high pres-
sure liquid chromatography) analysis. Ilwensisaponin
A was used as an external standard and digitoxin as an
internal standard. C18 reverse phase column and gradi-
ent elutions (acetonitrile with 0.1% orthophosphoric
acid and water with 0.1% orthophosphoric acid) were
used for HPLC analysis. Commercially obtained leaves
had a higher concentration of saponin (0.215 mg/g
tissue) than other leaves (0.081–0.198 mg/g tissue) or
the capsule samples (0.016 mg/g tissue).
SCREENING STUDIES
McCutcheon et al. (1992) used the disc diffusion method
to show the antibiotic activity of common mullein leaves
and demonstrated that a methanol extract of leaves
had antibacterial activity against Escherichia coli
[inhibition zone (iz); 8.0–10.0 mm], Mycobacter phlei
(iz; 8.0–10.0 mm) and Staphylococcus aureus methicillin-
resistant (iz; 10.1–15.0 mm).
The methanol extract of common mullein leaves
was found to partially inhibit the cytopathic effects of
bovine herpesvirus type 1, two double-stranded DNA
viruses, which causes respiratory, genital, conjunctival
or encephalitic infections which become latent in the
trigeminal ganglion (McCutcheon et al., 1995). More-
over, this extract had antifungal activity against Micro-
sporum cookerii (iz; 8.0–10.0 mm) and M. gypseum (iz;
8.0–10.0 mm) (McCutcheon et al., 1994).
Pardo et al. (1998) isolated iridoid glycosides
lateroside 1, harpagoside 2, ajugol 3 and aucubin 4
from an ethanol extract of the roots of mullein that
exhibits antigermination activity on the seeds of barley
(Hordeum vulgare). Bioassays indicated that at 3 mm
concentration, compounds 1, 2 and 4 showed moderate
inhibition of seed germination. Aucubin 4 was the most
active against root elongation and ajugol 3 showed no
activity in the bioassays on barley seed germination
and growth.
Biological activities of common mullein extracts
and commercial mullein products were assessed using
selected bench-top bioassays (Turker and Camper,
2002). Bioassays included antibacterial, antitumour, and
two toxicity assays (i.e. brine shrimp and radish seed
bioassay). Extracts were prepared in water, ethanol and
methanol and their biological activities assessed. For
extractions, three different sources of leaves (field-
grown, in vitro cultured and commercially obtained)
and capsules (from field-grown plants) were used.
Commercial products of common mullein (tea bags,
swallow capsules, an alcohol extract and flower oil) and
purified saponins were also assessed. The disc diffusion
assay (Kirby-Bauer method) and six different bacterial
strains (Escherichia coli, Pseudomonas aeruginosa
and Klebsiella pneumoniae which are gram-negative
bacteria, and Streptococcus pyogenes, Staphylococcus
aureus and Staphylococcus epidermidis which are gram-
positive bacteria) were used for antimicrobial activity
assessment. In general, aqueous extracts were the most
effective and showed antibacterial activity against K.
pneumoniae and S. aureus. Methanol extracts from leaf
materials showed little antibacterial activity only against
K. pneumoniae. Generally, commercial products and
purified saponins showed little or no antibacterial
activity against these bacteria (except for the flower oil
sample showing inhibition to K. pneumoniae, E. coli, P.
aeruginosa and S. aureus). Agrobacterium tumefaciens-
induced tumours in potato tissue were inhibited by all
common mullein extracts, commercial products and
purified saponins. Methanol extracts of in vitro-grown
leaves and commercially obtained leaves inhibited
tumour formation better than the other methanol,
ethanol and water extracts. The results of brine shrimp
bioassay showed that extracts of mullein were toxic
at higher doses (around 1000 mg / L). Among aqueous
extractions, decoctions had more toxicity (LC50 <
1000 mg/L) than the infusion extracts. In a radish seed
bioassay, at high doses (10 000 mg/L), all extracts
affected the root length. However, at 1000 mg / L, the
extracts did not affect the root length as much. Seed
germination was also inhibited by mullein extracts at
7500 mg /L. Low doses (1000 mg/L) affected germina-
tion, but not as much as higher doses. Ethanol extracts
inhibited seed germination more than other types of
extracts.
CONCLUSION
Verbascum thapsus L. (common mullein) is a herb with
a long history of use in folk medicine. The commercial
popularity of this plant has been increasing for the past
738 A. U. TURKER AND E. GUREL
Copyright © 2005 John Wiley & Sons, Ltd. Phytother. Res. 19, 733– 739 (2005)
respiratory disorders, tumour formation, urinary tract
infection and certain skin diseases. With these results,
V. thapsus L. has some scientific justification as a
medicinal herb. In the future more different bioassays
related to the biological activity of common mullein
can be performed and a more scientific rationale for
this plant may be obtained.
few years with the growing interest in herbs and pre-
ferring the ‘greener’ lifestyle. Today in health food
stores, one can easily find dried leaves and flowers,
swallow capsules, alcohol extracts and flower oil of
mullein in USA. The screening studies (bioassays
results) mentioned here confirm the popular utilization
of mullein extracts in folk medicine as a remedy for
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... Historically, mullein has been employed as an antidote for the respirational tract, mostly in cases of annoying coughs with bronchial blocking (Hoffmann and Manning 2002). Mullein leaves and flowers have expectorant and demulcent effects, which are employed by herbalists to cure respiratory complications for example bronchitis, dry coughs, whooping cough, tuberculosis, asthma, and harshness (Turker and Gurel 2005;Berk 1996). The flowers are gently diuretic and have a gentle and anti-inflammatory properties on the urinary area. ...
... The leaves are also diuretic, potion to reduce inflammation of the urinary system, and oppose the irritating effect of acid urine (Ambasta 1986;Turker and Camper 2002). Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). ...
... Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). The demulcent and emollient properties originate from the polysaccharide adhesive and gums that mitigate the bothered tissue. ...
Chapter
More than half of the population in developing nations depends on natural medication for treatment of different sicknesses and problems. Among them, Achillea millefolium from Asteraceae family is one restoratively significant plant called as “yarrow” and revealed as being utilized in folklore medication for sicknesses, for example, skin irritations, convulsive, hepatobiliary, and gastrointestinal issues. Monoterpenes are the most delegate metabolites, establishing 90% of the fundamental oils comparable to the sesquiterpenes, and a wide scope of chemical compounds have likewise been found. Distinctive pharmacological examinations in numerous in vitro and in vivo models have demonstrated the capability of A. millefolium with anti-inflammatory, antiulcer, anticancer activities, and so forth loaning help to the reasoning behind various of its conventional uses. Because of the essential pharmacological activities, A. millefolium will be a superior alternative for new medication discovery. Our chapter extensively gathers late phytochemical and pharmacological activities of A. millefolium, and should, accordingly, act as an appropriate reference for future investigation into the plant’s phytochemical profiling and by and large pharmacological assessment.
... Historically, mullein has been employed as an antidote for the respirational tract, mostly in cases of annoying coughs with bronchial blocking (Hoffmann and Manning 2002). Mullein leaves and flowers have expectorant and demulcent effects, which are employed by herbalists to cure respiratory complications for example bronchitis, dry coughs, whooping cough, tuberculosis, asthma, and harshness (Turker and Gurel 2005;Berk 1996). The flowers are gently diuretic and have a gentle and anti-inflammatory properties on the urinary area. ...
... The leaves are also diuretic, potion to reduce inflammation of the urinary system, and oppose the irritating effect of acid urine (Ambasta 1986;Turker and Camper 2002). Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). ...
... Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). The demulcent and emollient properties originate from the polysaccharide adhesive and gums that mitigate the bothered tissue. ...
... Historically, mullein has been employed as an antidote for the respirational tract, mostly in cases of annoying coughs with bronchial blocking (Hoffmann and Manning 2002). Mullein leaves and flowers have expectorant and demulcent effects, which are employed by herbalists to cure respiratory complications for example bronchitis, dry coughs, whooping cough, tuberculosis, asthma, and harshness (Turker and Gurel 2005;Berk 1996). The flowers are gently diuretic and have a gentle and anti-inflammatory properties on the urinary area. ...
... The leaves are also diuretic, potion to reduce inflammation of the urinary system, and oppose the irritating effect of acid urine (Ambasta 1986;Turker and Camper 2002). Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). ...
... Some herbal texts extend the remedial use to pneumonia and asthma (Turker and Gurel 2005). The leaves, roots, and the flowers are also anodyne, antibacterial, antispasmodic, astringent, emollient, nerving, vulnerary, analgesic, antihistaminic, anticancer, antioxidant, antiviral, bacteriostat, cardiodepressant, estrogenic fungicide, hypnotic, and calming (Harris 1972;Lucas 1969;Turker and Gurel 2005). The demulcent and emollient properties originate from the polysaccharide adhesive and gums that mitigate the bothered tissue. ...
Chapter
Verbascum thapsus is annual or biennial herb, which belongs to the family Scrophulariaceae. It has becomes naturalized in most temperature regions of the world, where it can be found in abundance on roadsides, meadows, and pasture lands. The plant V. thapsus is commonly known as “Mullein.” V. thapsus has a broad native range including Europe, North Africa, Western and Central Asia. It has been introduced to Japan, Sri Lanka, the United States of America, Australia, and New Zealand. The herb has been utilized as a medicinal herb since ancient times, and has a great potential to treat a number of ailments. Mullein is supposed to be loaded with significant number of bioactive constituents including triterpene, tetraglycosides, saponins, terpenes, flavonoids, carotenoids, tannins, carbohydrates, phenolic acid, sugars, proteins, and minerals. Due to the presence of these potent phytoconstituents, it has been traditionally used as a folk medicine for a majority of locals in different parts of world. Reported literature of the plant available from primary and secondary search engines unveil a number of pharmacological activities of the plant, including antitumor, cardiovascular, anti-inflammatory, hepatoprotective, antibacterial, antiviral, nephroprotective, anthelmintic, analgesic activity, etc. The aim of this chapter is to provide a comprehensive review of the various therapeutic activities of the plant along its phytochemical constituents which are responsible for its medicinal status. Keywords: Verbascum thapsus · Mullein · Scrophulariaceae · Phytochemical moieties · Pharmacological profile
... Whole plant are utilized traditionally to cure various diseases like fresh leaves is used to cure migraine attacks (Bianchini and Corbetta, 1977;Lewis and Elvin-Lewis, 1977), flowers and leaves are used against inflammatory ailments in respiratory tract and others (Rodriguez-Fragoso et al., 2008), seeds are used to cure fever due to poison (Gurmet et al.,2020), roots are source for treatment of the symptoms of diabetes and its complications (McCune and Johns, 2002) aerial part are used against wound healing, urinary disease, edema (Rajbhandari et al., 2009), whole plant act as a remedy in the treatment of tuberculosis (Allen and Hatfield, 2004). Furthermore, modern scientific advancement in medicine and therapeutic potential of V thapsus has been reviewed (Turker andGurel 2005 andRaiz et al., 2013). Verbascum thapsus is used in traditional Sowa-Rigpa system of medicine of Ladakh. ...
... Whole plant are utilized traditionally to cure various diseases like fresh leaves is used to cure migraine attacks (Bianchini and Corbetta, 1977;Lewis and Elvin-Lewis, 1977), flowers and leaves are used against inflammatory ailments in respiratory tract and others (Rodriguez-Fragoso et al., 2008), seeds are used to cure fever due to poison (Gurmet et al.,2020), roots are source for treatment of the symptoms of diabetes and its complications (McCune and Johns, 2002) aerial part are used against wound healing, urinary disease, edema (Rajbhandari et al., 2009), whole plant act as a remedy in the treatment of tuberculosis (Allen and Hatfield, 2004). Furthermore, modern scientific advancement in medicine and therapeutic potential of V thapsus has been reviewed (Turker andGurel 2005 andRaiz et al., 2013). Verbascum thapsus is used in traditional Sowa-Rigpa system of medicine of Ladakh. ...
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Trans-Himalayan region of India is known for its unique and valuable medicinal plants and Ladakh is major part of trans-himalayan. Verbascum thapsus is an important medicinal plant for traditional medicine. Present study was initiated to develop an agrotechnique for cultivation of Verbascum thapsus in three different environmental conditions that is in polycarbonate house, trench and open condition. To prepare any kind of drug, availability of raw material is indispensible. It was observed that significantly higher production of Verbascum thapsus was recorded in polycarbonate condition 1.95kg/plant fresh weight (whole plant weight) followed by trench 420g/plant and then in open air condition 370 g/plant. Similarly, other agro-morphological features were also observed to be enhanced in polycarbonate as compared to other two environments. Consequently, greenhouse condition is recommended as most suitable condition for large scale production the plant in trans-himalayan region of Ladakh. Similarly, in comparison to some reported data production of the plant was higher in open air condition in trans-himalayan cultivation may also be performed in open air condition after raising seedling in greenhouse.
... On the other hand, common mullein (V. thapsus) is a plant used in traditional medicine to treat inflammatory diseases, bronchial congestion, coughs, diarrhea, and snakebite [18]. R. communis leaf extract was used for the green synthesis of spherical gold nanoparticles with diameters between 40 and 80 nm and probed against Hela and HepG2 cells, obtaining 58.64% and 42.74% inhibitory effects, respectively [19], while V. thapsus was used for the synthesis of metallic nanoparticles, e.g., AgNPs for antibacterial activity [20], CuO nanoparticles for photocatalytic degradation of methylene blue [21], and Iron nanoparticles for Cr (VI) reduction [22]. ...
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Gold nanoparticles (AuNPs) are promising nanomaterials exhibiting anti-cancer effects. Green AuNPs synthesis using plant extracts can be used to achieve stable and beneficial nanoparticles due to their content of bioactive compounds. This research aimed to synthesize and evaluate the antiproliferative and caspase-3 activity induction of green AuNPs synthesized with common mullein (V. thapsus) flowers (AuNPsME) and castor bean (R. communis) leaves (AuNPsCE) ethanolic extracts in human HT29 and SW480 colorectal cancer cells. Their effect was compared with chemically synthesized AuNPs (AuNPsCS). The extracts mainly contained p-coumaric acid (71.88–79.93 g/g), ferulic acid (19.07–310.71 g/g), and rutin (8.14–13.31 g/g). The obtained nanoparticles presented typical FT-IR bands confirming the inclusion of polyphenols from V. thapsus and R. communis and spherical/quasi-spherical morphologies with diameters in the 20.06–37.14 nm range. The nanoparticles (20–200 g/mL) showed antiproliferative effects in both cell lines, with AuNPsCE being the most potent (IC50 HT29: 110.10 and IC50SW480: 64.57 g/mL). The AuNPsCS showed the lowest intracellular reactive oxygen species (ROS) generation in SW480 cells. All treatments induced caspase 3/7 activity to a similar or greater extent than 30 mM H2O2-treated cells. Results indicated the suitability of V. thapsus and R. communis extracts to synthesize AuNPs, displaying a stronger antiproliferative effect than AuNPsCS.
... Urtica dioica reported here against rheumatism has also been reported against the same disease in other folk medicines (Zouari Bouassida et al., 2017). In our study, Verbascum thapsus has been reported to cure asthma, cough and inflammation which is also evidenced by other ethnomedicinal reports (Turker & Gurel, 2005). Besides a number of plants, which are indigenous or endemic to the region, and introduced and naturalized from other parts of the world were also found. ...
Article
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The native communities of Himachal Pradesh are largely dependent on natural resources for their sustenance and have rich knowledge regarding indigenous uses, but require proper documentation for their long-time conservation. Diverse biological diversity in the Ban oak forests causes immense miscellany of widely used medicinal plants, thus an ethnobotanical approach was adopted to file the undocumented indigenous wealth and folkloric therapeutics of the communities against different human disorders. Folkloric practice provides an alternative to the needy rural population for the treatment of various ailments. The present study was conducted to document the medicinal plant diversity of ban oak forests during 2018-2019 and recorded 182 species of medicinal plants used by the local inhabitants. These are listed along with their uses, parts used, mode of administration and threat status.
... The main toxicants affecting the circulatory, respiratory, and nervous systems of fish are saponins, rotenone, and glucosides. Mullein, when thrown crushed into lentic waters, causes fish to have breathing difficulties (165). Among the most important Verbascum species that poison fish are V. phlomoides L., V. sinuatum, and V. thapsus. ...
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In this study, we document the practices of ethnoveterinary medicine and ethnopharmacology in the context of traditional transhumance routes that cross Castilla La Mancha from north to south. Transhumance is a type of grazing system that allows advantage to be taken of winter pastures (wintering places) and summer pastures by seasonal movement, twice a year, of cattle and their shepherds. Our study is based on over 200 interviews (from 1994 to 2021) conducted in 86 localities along eight major transhumance routes "cañadas reales" and 25 other minor transhumance routes, and involved 210 informants, 89 single and 121 groups, and 562 individuals, of which the majority were men. Sixty-three recorded pathologies and their treatments are discussed. Two hundred and two species and substances, belonging to 92 different families, have been recorded from the interviews, of which most are plants. Amid the toxic plant species, the most cited in the interviews are Erophaca baetica (L.) Boiss., Lupinus angustifolius L., and Oenanthe crocata L. Some of the species reported as toxic were reservoirs of pathogens or markers for dangerous areas. One of the fields most widely covered in our study is that of prevention, protection, and control of endo-and ectoparasites. This control is carried out mainly by means of aromatic plants. As a polyvalent species, Daphne gnidium L. is outstanding, and it contributes one-tenth of the records of our study. Among the species of fundamentally therapeutic use, Cistus ladanifer L. stands out by far. Principal Coordinate Analysis (PCoA) based on the repertories of ingredients, separates the routes whose most important sections run through siliceous terrain with its characteristic flora, especially in the provinces of Ciudad Real and Toledo, from the routes that run through the limestone terrain of Albacete and Cuenca, and link the Eastern Mancha and the "Serranía de Cuenca" with Andalusia and the Spanish Levant.
... Verbascum phlomoides L., orange mullein, is an evergreen species in the Scrophulariaceae family, native to Europe and Asia Minor and it is known as a pollinator attracting plant. V. phlomoides is biennial, but mulleins can be annual, biennial, or perennial (Turker & Gurel, 2005). Its height can reach up to 1.8 m, and it has dense yellow flowers from early to late summer. ...
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Orange mullein is a biennial plant belonging to the figwort (Scrophulariaceae) family. The vivid yellow flowers are arranged in spikes located on the top of the stem. It is a drought and cold-tolerant plant requiring much sunlight that grows on pastures, roadsides, and in dry weed associations. The aim of this study was to assess the effect of different solvents (water, 70% acetone, 70% methanol, and 70% ethanol) on the assessment of antioxidant capacity of leaves and flowers of orange mullein. Total phenolics are higher in the leaves and reach a value of up to 15.70 mg of gallic acid per one g of dry leaf weight, while flavonoids are more dominant in flowers and reach a value of 5.82 mg of quercetin per one g of dry flower. Correlation was performed between total phenolics, total tannins, total flavonoids, and antioxidant tests. It is the flavonoids that are mainly responsible a the high antioxidant activity establishing a correlation with all the tests performed, including FRAP, ABTS, DPPH, total antioxidant activity, total reduction capacity, and NBT reduction test.
... Both leaves and flowers are on the FDA GRAS and German Commission E lists. 135 Historically, a wider range of closely related Verbascum species was used to manage respiratory, gastrointestinal, and cardiovascular pathologies. 136 The plant is a good source of monoterpene iridoids such as aucubin (Figure 2, 8) and phenylpropanoids such as verbascoside. ...
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Medicinal plants represent rich sources of traditional medicines and numerous currently used medicines are either directly or indirectly derived from plants. Verbascum thapsus L. (great mullein or common mullein), a medicinal herb indigenous to northern Africa, western and central Asia, and Europe, has been brought to the Americas and Australia. It has been used as a medicine for lung, skin and throat disorders and has a long history of therapeutic importance, particularly as an astringent and calming agent. Presently, the dried leaves, flowers, various plant extracts and flower oil are used in several formulations within Indian traditional medicine. An extract taken from the roots is useful in minimizing toothache, and it also relieves stiffness and seizures. V. thapsus contains a wide variety of phytoconstituents, such as flavonoids, iridoid, phenylethanoid and phenylpropanoid glycosides, saponins, as well as vitamin C and minerals. The most valuable constituents are coumarin and hesperidin, which possess healing properties. Emerging literature based on experimental studies on V. thapsus demonstrates various biological and pharmacological properties, including antiviral, antioxidant, analgesic, sedative, anti-inflammatory, hypnotic, antibacterial, antifungal, as well as anticancer activities. The present review provides an updated, comprehensive, and critical evaluation of various health-promoting and disease-mitigating properties of V. thapsus.
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Darlington, H. T., and G. P. Steinbauer. (Michigan State U., East Lansing.) The eighty-year period for Dr. Beal's seed viability experiment. Amer. Jour. Bot. 48 (4): 321–325. 1961.—Seeds of 20 species of plants were buried 80 years ago on the Michigan State University campus. Bottles have been dug up periodically in the intervening years and the seeds tested for viability. A bottle representing the 80-year period of burial was dug up this spring and the contents distributed uniformly on the surface of sterilized soil in the greenhouse. Of the 20 species whose seeds were buried in 1879, only 3 have survivors as viable seeds. These are the same 3 that had survived the 70-year burial period, namely: curled dock, Rumex crispus; evening primrose, Oenothera biennis; and moth mullein, Verbascum blattaria. The low percentage survival of curled dock (2%) indicates that the longevity of the buried seeds of this species is nearing the end. The situation is more favorable for evening primrose, with 10% survival. The third, moth mullein, has about as many survivors as 10 years ago, a remarkably high figure of 70% viability. Since there has been no significant drop in viability of the buried seeds over the last 30 years, it seems highly probable that longevity of buried seeds of this species may extend over a century. A brief summary is given of the current status of the behavior of buried seeds, based on this and other buried seed experiments.
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The vascular tissues of the leafless form of the “lanceolate” mutant in tomato resemble those of the normal hypocotyl in the lower portions. A divergence in development occurs in the middle region of the mutant hypocotyl: two xylary strands split acropetally into five to seven units, while in the normal hypocotyl four main vascular bundles are established. The three to four week old mutant hypocotyl shows a marked proliferation of cells in its central cylinder when it is grown on a simple nutrient medium; proliferation also occurs in older specimens grown in soil. This condition is averted if the mutant is grafted beneath an actively growing normal shoot tip. Instead of having numerous, random cell divisions in the central cylinder, the grafted mutant shows large pith parenchyma cells and an active vascular cambium. These features characterize the normal hypocotyl and they have not been observed in the intact leafless mutant. The observed behavior of intact and grafted mutant hypocotyls can be interpreted in terms of the normal shoot tip acting both as physiological sink, with respect to cell-division stimuli produced mainly in the root system, and physiological source, with respect to cambial activating hormones.
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After 100 years of burial in moist well aerated sand, 21 seeds of Verbascum blattaria, one seed of Verbascum thapsus and one seed of Malva rotundifolia, germinated and produced normal plants. Six additional seeds, of the original 50 seeds of each of 20 varieties, germinated and died before identification was possible.
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Excised internodal segments of flowering specimens of Verbascum thapsus, with and without vascular tissues, were grown on a nutrient medium which lacked added growth regulators such as auxin and cytokinin. Stem segments which contained vascular tissues formed numerous buds in superficial layers of newly formed tissue within two weeks after transfer of the explants to the medium. Tangential cell divisions in the cambium and phloem are thought to be primarily responsible for the newly formed tissue. A regenerated bud displays dendritic trichomes on its first leaf primordium, whereas the earliest that such trichomes appear on intact seedlings is on the second foliage leaf primordium. Pith explants without vascular tissues become brown in a matter of two to three weeks with no visible sign of growth. Endogenous growth regulators supplied by vascular tissues are believed to be major factors in bud formation in excised internodal segments of this species.
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Germination of common mullein (Verbascum thapsus L.) in relation to constant and alternating temperatures, and light quality and duration was studied. In the dark, optimum germination occurred with 8-h warm periods of 30 to 40 C alternating with 16-h cold periods of 15 to 35 C. In the dark, no germination occurred below 10 C. With light during the 8-h warm periods, germination occurred at temperatures from 2 to 40 C, alternating with 0 C cold periods. The range of optimum temperatures for germination was also greatly expanded with light. Exposures to 24-h periods of red light, followed by incubation in the dark, produced the same germination response as 8 h of white light daily. Common mullein seeds germinated on the surface of the soil even though the seeds did not have mucilage.