ArticlePDF Available

Abstract and Figures

Armoracia rusticana (horseradish), a member of the Brassicaceae family, has been known since ancient times as a folk medicinal herb and as a plant of nutritional value and culinary interest. Currently horseradish is cultivated for its thick, fleshy and white roots which have a delicious intense pungency and for its tender leaves which are frequently used for salad mixed to other vegetables. The traditions to use horseradish plant for medicinal purpose are still applied in many countries. Horseradish is a rich source of a number of bioactive compounds such as glucosinolates (GLSs) and their breakdown products. Sinigrin is the dominant glucosinolate in both leaves and roots. Recent studies have shown that crude plant extracts have a complex profile of naturally occurring GLSs, with particular regard to sprouts. The increasing interest in these secondary metabolites, associated to the long and diffuse tradition of using horseradish in food preservation and as condiment in many parts of the world, is generating new applications of this plant in several agro-industrial and pharmaceutical sectors and is encouraging the use of its roots and leaves in functional food and medicine for human health. A bibliography review is discussed on ethnobotanical aspects and uses of this plant, as well as knowledge about its flavour compounds and GLS content and composition. This study summarizes also the updated information concerning the influence of the genotype and environment on GLS profile in horseradish.
Content may be subject to copyright.
REVIEW PAPER
Horseradish (Armoracia rusticana), a neglected medical
and condiment species with a relevant glucosinolate
profile: a review
Rosa Agneta Christian Mo
¨llers
Anna Rita Rivelli
Received: 20 March 2013 / Accepted: 28 May 2013 / Published online: 14 August 2013
ÓSpringer Science+Business Media Dordrecht 2013
Abstract Armoracia rusticana (horseradish), a
member of the Brassicaceae family, has been known
since ancient times as a folk medicinal herb and as a
plant of nutritional value and culinary interest. Cur-
rently horseradish is cultivated for its thick, fleshy and
white roots which have a delicious intense pungency
and for its tender leaves which are frequently used for
salad mixed to other vegetables. The traditions to use
horseradish plant for medicinal purpose are still
applied in many countries. Horseradish is a rich
source of a number of bioactive compounds such as
glucosinolates (GLSs) and their breakdown products.
Sinigrin is the dominant glucosinolate in both leaves
and roots. Recent studies have shown that crude plant
extracts have a complex profile of naturally occurring
GLSs, with particular regard to sprouts. The increasing
interest in these secondary metabolites, associated to
the long and diffuse tradition of using horseradish in
food preservation and as condiment in many parts of
the world, is generating new applications of this plant
in several agro-industrial and pharmaceutical sectors
and is encouraging the use of its roots and leaves in
functional food and medicine for human health. A
bibliography review is discussed on ethnobotanical
aspects and uses of this plant, as well as knowledge
about its flavour compounds and GLS content and
composition. This study summarizes also the updated
information concerning the influence of the genotype
and environment on GLS profile in horseradish.
Keywords Brassicaceae Ethnobotany
Ethnopharmacognosy Glucosinolates
Horseradish Isothiocyanates
Introduction
Horseradish, Armoracia rusticana G. Gaertn., B. Mey.
et Scherb., is an extremely hardy perennial plant,
member of the Brassicaeae family (Weber 1949;
Shehata et al. 2009). Its root and leaves have been used
in antiquity as both a medicinal herb and a condiment
(Rosengarten 1969), and the latter use is the principal
nowadays. It is currently cultivated for its thick, fleshy
and white roots that have a mix of delicious intense
pungency and cooling taste which is caused by sulfur
compounds, namely glucosinolates (GLSs) (Weber
1949; Balasinska et al. 2005; Walters and Wahle 2010).
R. Agneta (&)
Doctoral School of Crop Systems, Forestry and
Environmental Sciences, University of Basilicata,
Via dell’Ateneo Lucano, 85100 Potenza, PZ, Italy
e-mail: rosa.agneta@unibas.it
C. Mo
¨llers
Department of Crop Sciences, Georg-August-Universita
¨t
Go
¨ttingen, Von Siebold-Str. 8, D-37075 Go
¨ttingen,
Germany
A. R. Rivelli
School of Agricultural, Forest, Food and Environmental
Sciences, University of Basilicata, Via dell’Ateneo
Lucano, 85100 Potenza, PZ, Italy
123
Genet Resour Crop Evol (2013) 60:1923–1943
DOI 10.1007/s10722-013-0010-4
The species has become naturalized in many parts
of the world. Horseradish can be found in various
environments including fields, home gardens, weedy
areas, farmland, roadsides, ditches and disturbed areas
throughout Europe (Al-Shehbaz 1988; Hammer et al.
1992; Luczaj and Szymanski 2007; Sampliner and
Miller 2009). In open fields, horseradish is cultivated
as an annual or perennial crop. As annual crop it
requires a long growing season with high temperatures
during the summer and fall to enhance root develop-
ment; as a perennial it may stay productive for
10–20 years or more, thus requiring a careful field
management (Walters and Wahle 2010).
The main areas of commercial cultivation are found
in Europe and in North America (California, Illinois,
and Wisconsin) (Sampliner and Miller 2009). In
European countries, horseradish is especially pro-
duced in Germany, Hungary, and Poland (Shehata
et al. 2009). Particularly, large scale productions can
be found in certain regions of Germany, like Upper
Franconia, Thuringia and the Spreewald, firstly doc-
umented in 1569 (Hanelt 2001). In the Serbian
province of Vojvodina, the collection of wild horse-
radish began a long time ago, and the particular
ecotype collected, called ‘‘Novosadski’’, was highly
sought after by the Austro-Hungarian Empire court
thanks to its exceptional quality (Perlaki and Djurovka
2009). In Italy this plant grows in the Northern part
and in the Lazio and Basilicata regions (Hammer et al.
1990,1992; La Rocca and Chisci 2005; Guarrera
2006; Sarli et al. 2012). Although the hypothesis needs
to be confirmed by more detailed historic-botanical
studies (Pieroni et al. 2005), it is probable that
horseradish arrived in villages of inland Lucania
(viz. Basilicata) via migrants from Swabia during the
thirteenth century. Hammer et al. (2011) reported that
it was possibly introduced as a traditional spice plant
from Albania with Albanian immigrants to South
Italy, where today is particularly diffused in the
Arbe
¨reshe
¨areas of Basilicata, far away from North
Italy. A. rusticana is cultivated also in Russia, the
Caucasus, Asia and in some regions 1,000 m above
sea level in tropical countries (Sampliner and Miller
2009). In some regions of Russia, it may still be
possible to find horseradish landraces (Vetela
¨inen
et al. 2009). Recently, China has also begun commer-
cial production of horseradish, although much of the
crop is produced by small farmers (Shehata et al.
2009).
In this review we first present a botanical descrip-
tion of A. rusticana and its traditional uses in food and
medical fields due to its peculiar chemical composi-
tion. Afterwards, we review knowledge on flavour
constituents and profile, concentration and chemical
form of GLSs and their breakdown products in roots
and leaves. This review also stresses the influence of
the genetic variability and agronomic management on
GLS content and composition of horseradish plants.
Botanical description and origin
Horseradish has well defined botanical characteristics
which clearly separate the species from other
Brassicaceae. Various Latin names have been used
in the course of time by several authors for this
species. Taxonomically, horseradish has been placed
in various genera by different botanists (Weber 1949).
In 1969, Courter and Rhodes have recorded the names
for classification of horseradish during three historical
periods, as shown in Table 1. It was classified as
Nasturtium armoracia Fries, Roripa armoracia
Hitchc., Radicula armoracia Robinson, Cochlearia
armoracia L., Armoracia armoracia Britton, and A.
rusticana Gaertn. (Weber 1949). Specific epithets
used were rivini,rusticana,armoracia or lapathifolia.
Although A. lapathifolia is favored by some taxono-
mists, Lawrence (1953) and Fosberg (1965) have
pointed out that A. rusticana should have preference
(Courter and Rhodes 1969). The first valid published
scientific name of horseradish which resulted from the
generic name Armoracia was A. rusticana in 1800 by
Gaertner, Meyer, and Scherbius: this binomial is used
today (Shehata et al. 2009). Even more complex is the
variety of common names used to designate this
species whose polynomial is due to different lan-
guages and dialectical differentiation in different
territories. Horseradish has its gene center in Eastern
Europe and the south western parts of Russia
(Wedelsba
¨ck Bladh and Olsson 2011). In fact, early
records indicate that horseradish is a native of the
temperate regions of Eastern Europe and western Asia,
where wild types are found growing from Finland and
Poland to the regions around the Caspian Sea and the
desert of Cumania (now Romania) and Turkey. From
here, horseradish spread to Western Europe and across
the Atlantic to the New World by early settlers. Today,
horseradish has become naturalized in many parts of
1924 Genet Resour Crop Evol (2013) 60:1923–1943
123
the world and can be found both cultivated and
growing wild (De Candolle 1890; Shehata et al. 2009).
According to De Candolle (1890) the most probable
place of origin of A. rusticana was the temperate
region of Eastern Europe because of the word chren,
common in Slavic languages which was the most
primitive name for horseradish. Chren was introduced
into German and French dialects in the forms of kren,
kreen,cran and cranson.Meerretig,meer-radys and
meridi, which literally mean sea-radish, are other
words for horseradish that are common in several
western European languages, but these words are less
primitive than the name chren. The names horseradish
in English, raifort in French, and pepperrot in Swedish
are also of more recent origin than chren (Courter and
Rhodes 1969). The first use of the term horse radish
was made by John Gerard in his famous English herbal
(1597) that contains a lengthy entry with a woodcut
and clear description of the plant. Some believe that
the English called the plant horseradish in reference to
its propensity to rapidly spread in a ‘galloping’
behavior (i.e., it is difficult to control once it has been
introduced in an area) (Shehata et al. 2009). It is also
possible that it was called harsh radish because it was
so bitter on the tongue (McCann 2004). The word
armoracia has been commonly used as a generic name
or specific epithet: Armoracia is formed from the
Celtic: ar ‘near’, mor ‘the sea’, rich ‘against’, viz., a
plant growing near the sea (Courter and Rhodes 1969).
Armoracia belongs to the Cardamineae, which
comprise 10 genera and more than 340 species that
grow in moist habitats. The three species within the
genus (A. macrocarpa,A. rusticana,A. sisymbrioides)
have an affinity for wet habitats and commonly spread
through rhizomes (Al-Shehbaz et al. 2006; Sampliner
and Miller 2009). Brzezinski (1909) believes that
horseradish is not a natural species but a hybrid
(Weber 1949). Sampliner and Miller (2009) hypoth-
esized that A. rusticana is a species known only from
cultivation, and that cultivated A. rusticana popula-
tions were derived from natural populations of either
A. macrocarpa or A. sisymbrioides.
Horseradish is a large-leaved, hardy, and glabrous
perennial herb that grows to a height of up to 120 cm
(Fig. 1). Cauline leaves vary in shape and margin
depending on the habitat in which they are growing.
The leaves are long-petioled, oblong-ovate, cordate at
the base, unevenly crenate, and they grow to a length
of 30 cm up to 100 cm. (Sampliner and Miller 2009;
Shehata et al. 2009). Moreover, cultivars with an acute
leaf-base have smooth leaves, while those with a
cordate leaf-base have crinkled leaves; some geno-
types have intermediate leaf-types (Courter and
Rhodes 1969). The leaves on stalk are smaller in size,
have narrow bases and mostly sessile, are alternate,
lanceolate, and unevenly serrate to entire-margined. It
is interesting to note, as reported by Shehata et al.
(2008), that horseradish leaf morphology varies
through the season from entire (laminate) in the
summer to divided (pinnate or fern leaf) in the fall;
intermediate types of leaves are visible throughout the
season. The same authors (Shehata et al. 2008) have
shown in their work a photo with the different types of
leaves. The unstable leaf morphology of horseradish is
of considerable interest to botanists. In general, the
change from entire to pinnate in most obvious in the
fall as plants approach dormancy, suggesting environ-
mental influence (Shehata et al. 2008).
Table 1 Classification of horseradish during three periods of
history
Name Source
Ancient
Persicum sinapi Dioscorides
Sinapi persicum Dioscorides
Thlaspi cratevae Dioscorides
Persicon napy Pliny
Renaissance
Thlaspi alterum Turner
Raphanus vulgaris Matthioli
Raphanus magna Dodoens
Raphanus rusticanus Gerard
Raphanus sylvestris Bauhin J.
Linnaeus to modern
Armoracia rivini Ruppins
Cochlearia armoracia Linnaeus
Cochlearia rusticana Lamarck
Armoracia lapathifolia Gilibert
Armoracia rusticana Gaertner, Meyer
and Scherbius
Nasturtium armoracia Fries
Roripa rusticana Gren. & Godr.
Roripa armoracia Hitchoock
Radicula armoracia Robinson
Armoracia armoracia Britton
Table from Courter and Rhodes (1969) reproduced with
permission
Genet Resour Crop Evol (2013) 60:1923–1943 1925
123
The plant may or may not form a flower stalk
depending on environmental conditions (Weber
1949). In natural habitats, horseradish blooms very
profusely and for a long period of time, until mid-
August (Winiarczyk and Bednara 2008). Shehata et al.
(2009) reported that A. rusticana produces numerous
fragrant flowers with 5–7 cm long upright pedicels
borne on racemes that have four sepals, four petals,
and six tetradynamous stamens: their sepals are
2.5–3 mm long, broadly ovate and have membranous
white margins; petals are white, 5–7 mm long, and
broadly obovate; the inner stamens are 2.5 mm long
and the outer ones are 1.5 mm long; the stigma is
broad, round, and gently two-lobed. Horseradish bears
4–6 mm long, globose to obovate siliques with
persistent style on 20 mm long, upright-spreading
stems (Shehata et al. 2009). A. rusticana produces few
if any seeds, usually none, at most 6 seeds per pod
(Sampliner and Miller 2009). The seeds are smooth
and brown when mature (Shehata et al. 2009).
Winiarczyk and Bednara (2008) observed that, a
dozen or so days after the fall of the perianth of the
inflorescence, siliques started to form (Fig. 2): most of
the flowers were withered, but on every inflorescence,
there were a few dozen fruits, the majority of which,
however, were shriveled and deformed; in every
silique, most of the ovules were aborted, some were
much smaller and deformed, but sporadically there
were also large, properly developed young seeds. As a
matter of fact, many early botanists reported that
horseradish rarely produced viable seed and, until the
early part of the twentieth century, the plant was
considered to be highly sterile (Courter and Rhodes
1969). Weber (1949) observed partial pairing of
chromosomes and other irregularities during both
microsporogenesis and megasporogenesis. Wini-
arczyk and Bednara (2008) noted on the stigma of A.
rusticana morphological symptoms of the reaction of
self-incompatibility, typical for some species of the
Brassicacea family. This could explain the fact that
most A. rusticana plants are incapable of producing
viable seeds. In addition, the plants growing in a given
area, i.e. constituting a certain population, are often
Fig. 1 Armoracia rusticana plant grown in Basilicata Region
(South Italy). Personal photo
Fig. 2 Typical silique of a horseradish plant 2-year-old:
aFruits-silique started to form at the end of the vegetative
period; bsingle silique; cdevelopment of seeds in silique.
Images captured with stereo microscope (Carl Zeiss) at 96.3 by
a camera Olympus 8 Megapixel. The bar in Fig. 2bis2mm
1926 Genet Resour Crop Evol (2013) 60:1923–1943
123
genetically identical with the parent plant, as a
consequence of their vegetative reproduction (Wini-
arczyk and Bednara 2008). In fact, horseradish is
asexually propagated by planting root sections col-
lected from the previous year’s crop (Walters and
Wahle 2010) through the formation of adventitious
buds (Kamada et al. 1995). In particular, lateral roots
are separated from the healthy taproots for next year’s
planting stock (Shehata et al. 2009). The root system
of horseradish consists of a long, white, cylindrical or
tapering main root that can grow to about 60 cm in
loose soils; several thin lateral roots are formed around
the main root and near the collar of the crown;
undisturbed, the root system can reach a depth of
3–4 m with a lateral spread of about 1 m (Shehata
et al. 2009). Horseradish roots exhibit distinct polarity
with a proximal end (or point of attachment to the
main root that sprouts producing new vegetation) and
distal end (Walters and Wahle 2010).
According to Sampliner and Miller (2009) the
clonal propagation of A. rusticana may have resulted
in increased sterility due to (1) the accumulation of
deleterious mutations in gene regions associated with
seed production; (2) reproduction of sterile individuals
derived from an interspecific hybridization; or (3) self-
incompatibility mechanisms precluding fertilization
between widespread, genetically-identical plants. Fur-
thermore, research aimed at obtaining horseradish
plants capable of sexual reproduction could result in
better adaptation to various environmental conditions
and pest resistance (Winiarczyk et al. 2007).
Traditional uses as a spice and as folk medicinal
plant
The use of horseradish as food or folk medicinal plant
has spread from the Eastern and Mediterranean areas
both north- and westward during the Middle Ages (De
Candolle 1890; Wedelsba
¨ck Bladh and Olsson 2011).
It is believed that horseradish became popular as a
condiment in old Europe because there was no
refrigeration and its sharp spiciness covered the taste
of tainted meat (Shehata et al. 2009). In Bulgaria,
Romania and Russia, the whole root, grated root, or
piece of cut root are still used to flavour, help ferment,
aerate pickling liquid, and conserve cabbage for the
winter; leaves were also cut and used to conserve cut
vegetables for the winter (Sampliner and Miller 2009).
Horseradish was used by European Jews as a
symbolic food in the ritual meal as bitter herb of
Passover because its bitterness remembered the
suffering of their ancestors of the exodus from Egypt
(Schaffer 1981). By the late 1600s the English people
added horseradish to beef and oysters, and particularly
in spring, the tender leaves frequently were used for
greens mixed with other wild plants (Shehata et al.
2009); for Germans, horseradish roots were an excel-
lent addition to meat and fish when cut in very small
pieces, then crushed and mixed with salt and vinegar
(Courter and Rhodes 1969). These latter authors wrote
that horseradish was also mixed in catsup used to
flavour ground beets or added to mustard as a
seasoning. As reported by Shehata et al. (2009),
horseradish was often consumed boiled as a pot herb;
it was boiled, the water was drained, and it was boiled
a second time to eliminate bitter or harmful substances
before consumption.
The main portions of this plant are still used today
to prepare traditional dishes in many countries around
the world. In Poland, roots and leaves are used for
seasoning or as preservatives: the roots, usually
collected from the wild plants, were utilized as a
condiment, with pickled cucumbers, grated with
chopped boiled eggs, soups or meat dishes, often used
at Easter. The leaves placed in the oven under baking
bread, partly to prevent the bread from sticking and
partly to flavour the bread, are still widely used
(Łuczaj and Szyman
´ski, 2007). In Bulgaria, Romania
and Russia, grated roots are used as a condiment for
any cooked meat (e.g., lamb, pig, and chicken); they
are often combined with vinegar, salt, oil, cream, or
yogurt (Sampliner and Miller 2009). Particularly, in
Romania, grated roots, with or without oil, are eaten
with potatoes or polenta, or are grilled and eaten with
cream accompanying lamb and chicken; a mix of
grated horseradish, apples, salt, sugar, and vinegar is a
garnish used for meals; small pieces of cut A.
rusticana root are eaten in soups; leaves can also be
put into bread dough, which is then grilled (Sampliner
and Miller 2009). In South Italy there are many
traditional culinary uses, particularly, in the Basilicata
area this plant is typical base for preparing dishes
during the carnival period (Sarli et al. 2012): horse-
radish is grated as condiment for pasta or home-made
noodles (‘ferriciedd’), and it is used in fried eggs paste
to prepare a traditional dish (‘rafanat’) during the
carnival period; pieces of roots are also used for
Genet Resour Crop Evol (2013) 60:1923–1943 1927
123
aromatization of hand-made sausages; it is used raw
grated as condiment for cooked pork meat or for
lamb and chicken where the pieces of roots are
often combined with oil, salt and vinegar; the leaves
are used for salad, mixed to other vegetable species
(Pieroni et al. 2005; Sarli et al. 2012). Grated roots
are often combined with oil and used all year round,
also smeared on grilled bread (Sarli et al. 2012).
The root is still popular in Europe and North
America and is appreciated freshly grated, mixed
with vinegar or in a sauce to form a condiment often
used with meats or fish and as a flavour in other
recipes in salads and soups, on sandwiches, and also
in drinks such as the Bloody Mary (Wedelsba
¨ck
Bladh and Olsson 2011).
In addition to be used as food and condiment,
horseradish has a long history of utilization for
medicinal purposes in the traditional medicine (Sheh-
ata et al. 2009). In a recent review, Wedelsba
¨ck Bladh
and Olsson (2011) have well documented the histor-
ical uses of horseradish as a medicinal plant in
different parts of the world, and have shown also a
table with the different uses in various historical
periods. Since the ancient Greeks horseradish was
used as an aphrodisiac and as a rub to alleviate pain in
the back. Material medica’s of ancient physician and
the herbal of early botanists record many medical
virtues of horseradish (Courter and Rhodes 1969).
Early records, believed to be about horseradish are
found in ‘‘Naturalis Historia’’ by the Roman naturalist
and philosopher Gajus Plinius Secundus (AD 23-79)
that recommended the plant freshly grated for the
digestion after a heavy meal, and in ‘‘De Materia
Medica’’ by the Greek physician Pedanius Dioscorides
(AD 40-90) that described the plant as a diuretic and
very hot herb (Bostock and Riley 1856; Courter and
Rhodes 1969; Wedelsba
¨ck Bladh and Olsson 2011).
Hildegard of Bingen (1098–1179), a German abbess,
in her book about medicinal plants, recommended
horseradish mixed with warm vine or water as a
treatment for lung diseases; to cure heartache or heart
diseases, dry and pulverized horseradish could be
mixed with Alpinia galanga, a Chinese herb often used
as a stimulant and drug (Wedelsba
¨ck Bladh and
Olsson 2011). In 1597, the botanist and herbalist John
Gerard claimed that horseradish reduced pain from
sciatica, relieved colic, increased urination, and killed
worms in children (Courter and Rhodes 1969). It was
stated to be an expectorant, soothing for respiratory
problems, and may help to relieve rheumatism by
stimulating blood flow to inflamed joints (Shehata
et al. 2009). The most common use of horseradish was
as a remedy for scurvy, as also recommended by
herbalist William Coles (1626–1662), especially for
seamen on long voyages that had not access to fresh
vegetables or fruit, so the lack of vitamin C in the diet
often caused scurvy outbreaks on board (Wedelsba
¨ck
Bladh and Olsson 2011). In 1880, Bentley and Trimen
reported that grated horseradish root was used mixed
with honey and warm water for influenza, and it could
be used as poultice by adding cornstarch to fresh
horseradish and applying it to the affected areas in a
gauze bandage. Native Americans used horseradish to
treat toothaches, as a urinary aid for gravel (kidney
stones), as a diuretic, as a gastrointestinal aid to
improve digestion, and as a respiratory aid to treat
asthma, cough, and bronchitis (Moerman 1998;
McCann 2004; Shehata et al. 2009). The tradition to
use the plant for medicinal purpose is still applied in
countries like Bulgaria, Romania, and some part of
Russia: grated roots or leaves are put in a cloth
(sometimes with alcohol or vinegar) and applied to
the skin to ease pain, or a paste made from the root is
applied to the throat to ease breathing problems.
Mixtures with honey from Robinia pseudoacacia are
used for coughs and bronchitis, while in combination
with vinegar, salt, and sugar it is used for lowering
blood pressure (Sampliner and Miller 2009). Further-
more, in an ethnobotanical study about a community
of Russlanddeutsche (Germans from Russia) living in
Ku
¨nzelsau (Germany), it was reported that Russian
sauerkraut, used as an important home medicine for
treating flu and liver disease, contained among other
herbs, also horseradish leaves (Pieroni and Gray
2008). In the Basilicata region (South Italy), horse-
radish leaves and roots have been traditionally used
as a remedy for rheumatism, headaches, sinuses,
coughs and bronchitis. Added to water, vinegar, salt,
sugar, or aromatised with Anethum graveolens and
Laurus nobilis leaves, this plant is also used as a
remedy for healing drunkenness. Particularly, in
Castelmezzano (Basilicata), the Ukrainian women
have introduced the functional use of pickled toma-
toes (with dill, bay and horseradish leaves), con-
sumed to recover from a drunken state. Leaves, added
to dog food, are used because of their antimicrobial
activity (Pieroni et al. 2004; Pieroni and Quave 2005;
Sarli et al. 2012).
1928 Genet Resour Crop Evol (2013) 60:1923–1943
123
Also, according to Wedelsba
¨ck Bladh and Olsson
(2011), the traditional knowledge of using horseradish
in food preservation at home is generating new
applications for the plant as a natural anti-bacterial
component in food (e.g., to reduce coli-bacteria or
other microbial growth) and the new knowledge of
bioactive components such as GLSs and their break-
down products, combined with traditional knowledge
of medicinal properties, is encouraging the use of
horseradish roots and leaves in functional food and
medicine to inhibit different cancer forms or gastric
lesions.
Glucosinolates, the main flavour constituents
of horseradish
Horseradish tissues, when unbroken, are inodorous.
The intense pungent aroma and the lachrymatory
odour of horseradish results from crushing, grinding,
or chewing the cells (Courter and Rhodes 1969; Jiang
et al. 2006) and therefore it is specially cultivated to
supply a hot spicy flavour (Sultana et al. 2003). The
characteristic taste and odour is due to compounds
such as GLSs and their mostly volatile breakdown
products (Mucete et al. 2006; Redovnikovic et al.
2008a). GLSs are hydrolysed by the enzyme com-
monly known as myrosinase to a variety of com-
pounds as isothiocyanates, nitriles, thiocyanates,
epithionitriles, and oxazolidines. The most common
products are isothiocyanates (Bones and Rossiter
2006). GLSs are compounds mainly found in plants
belonging to the order of Capparales, with particular
reference to several species of the Brassicacee family
(radish, cabbage, cauliflower, broccoli, mustard, tur-
nip, oilseed rape) including horseradish (Fahey et al.
2001; Blazevic and Mastelic 2009). These secondary
metabolites are characterized by a core sulfated
isothiocyanate group, which is conjugated to thioglu-
cose, and a further R-group derived from amino acids
(Clarke 2010). GLSs can be divided into three classes
based on the structure of different amino acid precur-
sors: (1) aliphatic GLSs derived from methionine,
isoleucine, leucine or valine; (2) aromatic GLSs
derived from phenylalanine or tyrosine; and (3) indole
GLSs derived from tryptophan (Wittstock and Halkier
2002; Redovnikovic et al. 2008b). The number of
reported GLSs is now approaching 200 (Clarke 2010).
A number of plants contain only a single GLS, the
majority contain 2–5, while 34 individual GLSs are
reported in the seeds and leaves of a collection of
ecotypes of Arabidopsis thaliana (Kliebenstein et al.
2001; Clarke 2010). GLS concentrations in plants,
although highly variable, are around 1 % of dry weight
in some Brassica vegetables (Rosa et al. 1997; Clarke
2010). The GLS content is higher in black mustard
seed (Brassica nigra) and horseradish roots (over
10 % by dry weight) than in the other constituent parts
of the Brassicaceae (Li and Kushad 2004; Mucete
et al. 2006). However, very little is known about the
concentration and biochemical composition of horse-
radish, especially the distribution of GLSs in the
different organs of the plants (Li and Kushad 2004).
A first study about the identification of GLSs of
horseradish roots by GC–MS (gas chromatography-
mass spectrometry) was published by Grob and Matile
(1980). They identified 30 GLSs (Table 2) which
exceeds the number of GLSs detected before in any
other species. The authors reported that a close
examination of the chromatograms may indicate that
the number of structures present in horseradish may be
even larger, although today it is necessary to investi-
gate and confirm all individual molecules listed by
Grob and Matile (1980). Several methods have been
developed to detect either intact GLSs or their desul-
foglucosinolates (Lee et al. 2006; Cataldi et al. 2007).
The structural identity of GLSs is known and
described only for those which can be isolated and
identified by traditional methods. With increasing
complexity of GLS composition and decreasing con-
centration of individual GLSs in the sample, more
sensitive and selective analytical methods are required
for their identification (Agneta et al. 2012; Lelario et al.
2012). Hence, for their accurate structural identifica-
tion, in addition to traditional methods, sensitive and
selective analytical methods are required, especially
when GLSs are present in trace amounts only. Mevy
et al. (1997) investigated the occurrence of GLSs in
various differentiated horseradish tissues (Table 3)by
HPLC analysis of desulphoglucosinolates. Particu-
larly, the individual GLSs present in regenerated
plantlets were compared with those of embryoids,
suspension cells and calli in order to investigate
whether the distribution of these compounds resulted
from cellular differentiation. Moreover, the authors
revealed the presence of two GLSs, i.e. 2-phenylethyl-
and 2-propenyl-, which appeared to be confined to
roots and leaves, respectively, while the highest
Genet Resour Crop Evol (2013) 60:1923–1943 1929
123
content of indole-3-methyl-GLS (glucobrassicin) was
found in calli. Their data clearly demonstrated that the
occurrence of indole GLSs in embryoids, suspensions
cells and calli relates to their age. It has also been
reported that younger and growing tissues synthesize
more indole GLSs than older and senescent tissue, so
their results raise the question of the role of GLSs in
healthy plants (Mevy et al. 1997). Mevy et al. (1999),
by comparison of immobilized and suspension cells of
horseradish, showed that GLS production may be
improved by cell immobilization technique. More-
over, the chromatograms obtained revealed that gluco-
brassicin occurred earlier than its hydroxylated form in
cells, therefore their data suggest that hydroxygluco-
brassicin might result directly from modification of the
side chain of glucobrassicin.
Table 2 Glucosinolates of
horseradish roots as
identified by their
corresponding mustard oils
(Table modified from Grob
and Matile 1980) and odour
description (reported by
Sultana et al. 2003)
The common names are
those generally accepted
(Fahey et al. 2001;
Bellostas et al. 2007a;
Clarke 2010)
a
Concentration is
expressed as lg/g fresh
weight
b
Odour description of the
corresponding
isothiocyanates
GLS common name GLS systematic name lg/g
a
Odour description
b
Glucoviorylin 2-Methylthioethyl- 2
Glucoputranjivin Iso-propyl- 1 Chemical, weak mustard
like
Sinigrin Allyl- 240 Strongly pungent, mustard-
like, lachrymatory, bitter
Glucoarabidopsithalin 2-Hydroxypropyl- 3
Glucoiberverin 3-Methylthiopropyl- 5 Strongly radish-like, pungent
Glucoiberin 3-Methylsulfinylpropyl- 3
Glucocheirolin 3-Methylsulfonylpropyl- 3
n-butyl- 5
sec-butyl- 22 Chemical, weak mustard like
Glucoconringianin iso-butyl- 6 Sweet, chemical
Gluconapin 3-Butenyl- 14 Green, pungent, aroma
Glucocappariflexin 3-Hydroxybutyl- 0.5
Glucoerucin 4-Methylthiobutyl- 0.5
Glucokohlrabiin n-pentyl- 1
3-Methylbutyl- 8
2-Methylbutyl- 0.5
Glucobrassicanapin 4-Pentenyl- 20 Green, pungent, acrid,
fragrant leaf
Glucomoracialapathin 2-Hydroxypentyl- 0.1
3-Hydroxy-4-pentenyl- 0.8
Glucoberteroin 5-Methylthiopentyl- 0.2 Radish-like, pickle-like
3-Hydroxy-5-
methylthiopentyl-
0.2
Iso-hexyl- structure
unknown
0.4
Glucowasabiamin 5-Hexenyl- 2 Green, pungent, fatty
Glucolesquerellin 6-Methylthiohexyl- 0.1 Radish-like, sweet, fatty
Iso-heptyl- structure
unknown
0.05
Glucotropaeolin Benzyl- 4 Strongly radish-like, pungent,
strong watercress aroma,
tingling sensation
Gluconasturtiin 2-Phenylethyl- 55
Glucoarmoracialapicin 3-Phenylpropyl- 0.1
Glucoarmoracialafolicin 4-Phenylbutyl- 0.5
Methoxybenzyl- 0.5
1930 Genet Resour Crop Evol (2013) 60:1923–1943
123
Redovnikovic et al. (2008a) analyzed horseradish
plantlets leaves by reversed-phase high-performance
liquid chromatography (RP-HPLC) analysis of des-
ulphoglucosinolates and showed that the aliphatic
GLS 2-propenyl-GLS (sinigrin) accounted for more
than 80 % of total GLSs. Lower amounts of phenyl-
ethyl-GLS (gluconasturtiin) and of the three indole
compounds 3-indolylmethyl-GLS (glucobrassicin),
4-methoxy-3-indolylmethyl-GLS (4-methoxygluco-
brassicin) and 4-hydroxy-3-indolylmethyl-GLS (4-
hydroxyglucobrassicin) were found. Li and Kushad
(2004) evaluated roots from 27 horseradish accessions
and leaves from 9 accessions for their GLS content by
RP-HPLC as desulphoglucosinolates. In roots, total
GLS concentration ranged from 2 to 296 lmol g
-1
of
dry weight (DW), with sinigrin representing on
average 83 % of the total GLSs, followed by glu-
conasturtiin with 11 %, and glucobrassicin with 1 %.
In some of the accessions, other GLSs as progoitrin,
gluconapin, 4-hydroxyglucobrassicin, and 4-meth-
oxyglucobrassicin, were detected in minor concentra-
tions (\1 %). According to the same authors (Li and
Kushad 2004), the origin of plant did not seem to have
an effect on total GLSs level; furthermore, the average
total GLS concentration in most horseradish roots
accessions was similar to the value for brown and
oriental mustard seeds, but significantly higher than
what the authors had previously reported for broccoli
(13 lmol g
-1
of DW), brussels sprouts (25 lmol g
-1
of DW), cabbage (11 lmol g
-1
of DW), cauliflower
(15 lmol g
-1
of DW), or kale (15 lmol g
-1
of DW).
The total GLS levels in the leaves, instead, ranged
from 34 to 201 lmol g
-1
of DW. Similar to the roots,
sinigrin was the dominant GLSs in the leaves,
representing on average 92 % of the total. Leaves
also contained gluconasturtiin, but at a much lower
concentration (0.2 lmol g
-1
of DW) than in roots
(10.3 lmol g
-1
of DW). Conversely, leaf tissues
contained neoglucobrassicin (2.5 % of total) instead
of glucobrassicin.
Recently, Alnsour et al. (2012), referring to the
GLS content, confirmed previous results of Redov-
nikovic et al. (2008a) and Mevy et al. (1997) showing
that the overall GLS concentration (maximum
40 lmol g
-1
of DW) in the in vitro cultivated
Armoracia plants is much lower than that found by
Li and Kushad (2004) in soil-grown ones (in average
100 lmol g
-1
of DW). Surprisingly, in contrast to soil
grown plants, where the GLS concentration in roots
and leaves were quite similar, in the in vitro plantlets
the concentration of GLS in the root was much lower
than in the leaves (Alnsour et al. 2012).
The distribution of the GLSs varies among plant
organs, with both quantitative and qualitative differ-
ences between roots, leaves, stems and seeds; in any
case the plant age is a major determinant of the
qualitative and quantitative GLS composition in the
plants (Fahey et al. 2001; Velasco et al. 2007;
Cleemput and Becker 2011). Li and Kushad (2004)
reported differences in GLS content between young
Table 3 Glucosinolate contents of various horseradish tissue
Materials Medium
a
Glucosinolate contents (lg/g dry wt.)
b
Total
contents
IMG OH-IMG PhG PrG
Callus (28 days) MS 4.95 (0.30)
c
0.32 (0.03) 5.27
Suspension cells (21 days) MS 0.09 (0.02) – – – 0.09
Suspension cells (21 days) D-MS 0.31 (0.04) – – – 0.31
Embryoids (28 days) D-MS 0.71 (0.02) – – – 0.71
Root of r. p.
d
(3 months) H-MS 2.35 (0.40) 2.35
Small leaves of r. p. (3 months) H-MS – – 16.4 (4.00) 16.4
Large leaves of r. p. (3 months) H-MS – – 28.3 (5.00) 28.3
Table modified from Mevy et al. (1997)
a
Murashige and Skoog basal salts with organic substances, MS; 2,4-D-free-MS, D-MS; hormone-free-MS, H-MS
b
Indole-3-methylglucosinolte, IMG; 4-hydroxy-indole-3-methylglucosinolate, OH-IMG; 2-phenylethylglucosinolate, PhG;
2-propenylglucosinolate, PrG
c
Mean values of three independent extractions ±(SD)
d
r. p. regenerated plantlets
Genet Resour Crop Evol (2013) 60:1923–1943 1931
123
and fully developed roots and leaves of horseradish
(Table 4), noting that the major differences were
observed regarding indole GLSs, in fact the younger
tissue of roots and leaves tended to accumulate more
glucobrassicin and neoglucobrassicin, respectively,
than fully developed organs ([60-fold higher in young
leaves than fully developed leaves). Due to the
complexity of this vegetal matrix, further studies are
required to better understand the accumulation of
various GLSs during different growth stages also by
comparing of different genotypes. For the model plant
Arabidopsis thaliana, a member of the Brassicaceae, it
was reported that GLS accumulation varies signifi-
cantly among different organs and developmental
stages, with regards to both composition and concen-
tration (Brown et al. 2003; Yang and Quiros 2010).
Particularly, dormant and germinating seeds had the
highest concentration (2.5–3.3 % by dry weight);
inflorescences and siliques had the next highest levels
(0.6–1.2 %) followed by roots, stems and cauline
leaves, and rosette leaves (0.3–1.0 %) (Fahey et al.
2001; Brown et al. 2003). The GLSs are plant defense
compounds that accumulate preferentially in the
reproductive organs as seeds, flowers, fruits and young
sprouts (Fahey et al. 2001; Brown et al. 2003;
Bellostas et al. 2007b). The low GLS concentration
in the leaves and stems in comparison with the seeds
could be attributed to the dilution of GLSs during plant
growth (Clossais-Besnard and Larher 1991; Cleemput
and Becker 2011). Recently, Agneta et al. (2012) gave
a complex profile of naturally occurring intact GLSs in
sprouts and roots of a horseradish accession largely
diffused in southern Italy by using reversed-phase
liquid chromatography coupled with electrospray
ionization and a hybrid quadrupole linear ion trap
and Fourier transform ion cyclotron resonance mass
spectrometry (LC-ESI-FTICR MS). In sprouts 16
and in roots 11 GLSs were identified (Table 5).
The authors confirmed the presence of sinigrin, 4-hy-
droxyglucobrassicin, glucobrassicin, gluconasturtiin,
and 4-methoxyglucobrassicin and identified glucoiber-
in, gluconapin, glucocochlearin, glucoconringianin,
glucosativin, glucoibarin, 5-hydroxyglucobrassicin,
glucocapparilinearisin or glucobrassicanapin, glucotro-
paeolin, and glucoarabishirsutain, not previously char-
acterized in horseradish. Particularly notable was the
presence of the putative 2-methylsulfonyl-oxo-ethyl-
GLS, not reported before. In addition, horseradish
Table 4 Change in total, individual and residual glucosinolates in two accession of horseradish roots and leaves as a function of
growth stage
Accession Growth stage Sinigrin Glucobrassicin Gluconasturtiin Total GLSs
a
Residual GLSs
b
lmol g
-1
of DW
Roots
810-A Small 58.0b
c
13.4b 0.2c 74.7b 3.1c
810-A Mature 258.0a 2.8c 20.1a 295.8a 14.9a
1573 Small 77.3b 33.9a 0.2c 119.1b 7.7ab
1573 Mature 47.5b 0.7c 2.0b 55.5c 5.3bc
Accession Growth stage Sinigrin Neoglucobrassicin Gluconasturtiin Total GLSs Residual GLSs
lmol g
-1
of DW
Leaves
810-A Small 77.6ab 66.2a 0.5a 150.2a 2.3a
810-A Mature 114.8a 1.2b 0.1a 126.0a 1.1a
1573 Small 60.3b 69.7a 0.3a 129.2a 2.4a
1573 Mature 48.6c 1.1b 0.4a 52.7b 1.2a
Table modified from Li and Kushad (2004)
a
Total glucosinolates in roots and leaves represent the sum of sinigrin, glucobrassicin, and gluconasturtiin plus residual
glucosinolates and the sum of sinigrin, neoglucobrassicin and gluconasturtiin plus residual glucosinolates, respectively
b
Residual glucosinolates is the sum of at least four additional glucosinolates
c
Values represent the means of four replicates per accession. Mean separation within each column by Duncan’s multiple-range test,
P=0.05
1932 Genet Resour Crop Evol (2013) 60:1923–1943
123
sprouts were found to be richer in GLSs than roots.
Further studies are required to quantify the content of
these molecules that are responsible for the typical
flavour of this plant.
Normally, the components that cause the intense
pungency are physically separated from one another
(Shehata et al. 2009). GLSs are stored in the vacuole
(Grob and Matile 1979; Helmlinger et al. 1983;
Koroleva et al. 2000) while the myrosinase enzyme
that causes their degradation (Mucete et al. 2006)is
confined within specialized cells known as myrosin
cells (Kissen et al. 2009; Borgen et al. 2010). Upon
crushing horseradish roots or leaves allyl and 2-phen-
ylethyl isothiocyanate are produced from sinigrin
and 2-phenylethyl-GLS. Allyl isothiocyanate is
reported to be a lachrymatory compound, bitter in
taste with a strong, pungent smell, while 2-phenylethyl
isothiocyanate had no pungency and lachrymatory role
at all, like allyl isothiocyanate (Sultana et al. 2003;
Shehata et al. 2009). Both isothiocyanates are largely
responsible for the typical, characteristic flavour and
pungency in mustard as well as in horseradish (Mithen
et al. 2000; Sultana et al. 2003; Kosson and Horbowicz
2008).
Recently, eighteen compounds that contribute to
the flavour of horseradish were identified in roots
immediately after harvesting (Table 6) by D’Auria
et al. (2004), using SPME–GC–MS (solid phase
microextraction coupled with gas chromatography
mass spectrometry). The main compounds were allyl
isothiocyanate, 4-isothiocyanato-1-butene, and 2
phenylethyl isothiocyanate. Allyl isothiocyanate was
the largest peak in the chromatogram and accounted
for 81 % of the overall integrated peak area. It is
noteworthy that methyl, ethyl, and isopropyl isothio-
cyanate were not found. By the same authors (D’Auria
et al. 2004) the different volatile organic compounds
listed in Table 6were identified in cut horseradish
kept at 5 °C for 12 h. The concentration of most of the
volatile compounds declined rapidly and 2-phenyl-
ethyl isothiocyanate compounds became the most
abundant compound. The fact that the flavour com-
position of cut horseradish samples changes is relevant
when specifying the treatment of horseradish to obtain
mustard.
To preserve the pungency and quality, the ground
product should be consumed quickly or refrigerated
to minimize loss of volatile flavour compounds.
Table 5 Glucosinolates identified in different portions of horseradish plants by LC-ESI-FTICR MS: common name, retention time
(t
R
), molecular formulae, monoisotopic value as [M-H]
-
ion (m/z), mass error (ppm)
GLS common name t
R
(min) Molecular
formulae
Monoisotopic calculated
value as [M-H]
-
ion (m/z)
Mass error
(ppm)
a
Glucoiberin 4.3 C
11
H
21
NO
10
S
3
422.02549 -1.30
Sinigrin 4.4 C
10
H
17
NO
9
S
2
358.02720 -0.59
2-Methylsulfonyl-oxo-ethyl-GLS 4.6 C
10
H
17
NO
12
S
3
437.98402 -1.21
Gluconapin 5.5 C
11
H
19
NO
9
S
2
372.04285 -1.30
Glucocochlearin 6.2 C
11
H
21
NO
9
S
2
374.05850 1.00
Glucoconringianin 6.4 C
11
H
21
NO
9
S
2
374.05850 -0.70
Glucosativin 6.5 C
11
H
21
NO
9
S
3
406.03057 1.35
Glucoibarin 7.3 C
15
H
29
NO
10
S
3
478.08808 -0.10
4-Hydroxyglucobrassicin 7.4 C
16
H
20
N
2
O
10
S
2
463.04866 -1.50
5-Hydroxyglucobrassicin 7.5 C
16
H
20
N
2
O
10
S
2
463.04866 -1.70
Glucocapparilinearisin or
Glucobrassicanapin
7.8 C
12
H
21
NO
9
S
2
386.05850 -0.20
Glucotropaeolin 8.5 C
14
H
19
NO
9
S
2
408.04285 -1.10
Glucobrassicin 10.3 C
16
H
20
N
2
O
9
S
2
447.05375 -1.50
Gluconasturtiin 12.1 C
15
H
21
NO
9
S
2
422.05850 -1.20
4-Methoxyglucobrassicin 12.7 C
17
H
22
N
2
O
10
S
2
477.06431 -1.20
Glucoarabishirsutain 16.7 C
15
H
29
NO
9
S
3
462.09317 -1.20
Table modified from Agneta et al. (2012)
a
Mass error in part per million, ppm =10
6
9(accurate mass-exact mass)/exact mass
Genet Resour Crop Evol (2013) 60:1923–1943 1933
123
However, ground horseradish slowly losses its pun-
gency, becomes dark, and develops off flavours even
under refrigeration; this quality loss can be slowed by
adding a fat or oil, such as cream (Courter and Rhodes
1969). Kosson and Horbowicz (2009) showed that
higher storage temperature (2, 8 and 18 °C were
tested) caused faster decline of isothiocyanates con-
centration in horseradish cream. Therefore, to main-
tain the human health-promoting compounds in
processed horseradish, the authors suggested storing
horseradish cream under cold conditions. Further-
more, also dehydrated horseradish in diced, flaked or
powder form has a great potential as a flavour
ingredient in sauces, dressings, and seasoning formu-
lations (Sahasrabudhe and Mullin 1980). Saha-
srabudhe and Mullin (1980) recommended that in
processing horseradish to a dried product, one must
process the cleaned roots immediately after dicing or
crushing and maintain the temperature below 65 °C, to
retain viability of the myrosinase to obtain a product
with the desired flavour intensity and acceptable color.
In addition, Sahasrabudhe and Mullin (1980) evalu-
ated the odour intensity of the powders by sniffing,
after adding water: they noted that freeze dried
Table 6 Volatile organic
compounds found in the
fresh horseradish root and
in the cut of roots kept at
5°C for 12 h
Table modified from
D’Auria et al. (2004)
Compound r.t. (min) Fresh 5 °C for 12 h
ppm ppm
Thiobismethane 1.76 0.8 ±0.2 1.4 ±0.1
Carbon disulphide 1.81 0.8 ±0.2 –
3-Butenenitrile 2.38 7.8 ±0.1 –
3-Methylbutanal 2.73 0.4 ±0.2 0.6 ±0.1
2-Ethylfuran 2.78 1.2 ±0.1 5.8 ±0.3
3-Methyl-1-butanol 3.16 1.4 ±0.1
Toluene 3.69 0.8 ±0.1
Hexanal 4.32 4.5 ±0.1 3 ±0.3
E-2-hexenal 5.42 1.6 ±0.1 3 ±0.2
Heptanal 6.45 0.3 ±0.1
Allyl isothiocyanate 6.64 3,300 ±2.0 175 ±2.0
Benzaldehyde 7.78 0.5 ±0.1
Isobutyl isothiocyanate 7.85 15 ±0.3 –
4-Isothiocyanato-1-butene 8.45 63 ±1.0 –
2-Pentylfuran 8.45 3 ±0.3
Butyl isothiocyanate 8.74 13 ±0.3 –
2-Ethyl-1-hexanol 9.24 1.8 ±0.2
Phenylacetaldehyde 9.60 0.3 ±0.1
3-Methylbutyl isothiocyanate 9.98 8.6 ±0.2 –
Pentyl isothiocyanate 10.78 3.7 ±0.2 –
Nonanal 10.85 1.6 ±0.2 1.4 ±0.2
4-Ethylbenzaldehyde 12.01 0.4 ±0.1
4-Methylpentyl isothiocyanate 12.02 0.4 ±0.1 –
L-(-)-Menthol 12.20 0.2 ±0.1
Naphthalene 12.43 0.2 ±0.1
Decanal 12.76 0.5 ±0.1
Trans,trans-nona-2,4-dienal 12.95 0.2 ±0.1
Benzenepropanenitrile 13.49 0.8 ±0.1 0.2 ±0.1
Benzyl isothiocyanate 15.67 14 ±0.2 8.8 ±2.0
Junipene 16.43 0.2 ±0.1
Italicene 17.24 0.8 ±0.1
2-Phenylethyl isothiocyanate 17.44 400 ±1.0 432 ±3.0
1934 Genet Resour Crop Evol (2013) 60:1923–1943
123
samples and samples processed at 65 °C had the same
odor intensity as fresh samples while samples pro-
cessed at 90 °C lacked odor, as a consequence of
deactivation of myrosinase at temperatures higher
than 70 °C.
Further investigations are required to quantify the
content of the molecules that are responsible for the
typical flavour, which is one of the main characteris-
tics for making a high-quality product and to achieve a
high marketable yield, as requested by the agro-
industrial and pharmaceutical sectors.
Biological activities of Armoracia rusticana: human
health, antimicrobial and insecticidal effects
When plant tissue is disrupted or crushed, myrosinase
hydrolyzes GLSs to variety of molecules. These
products, especially isothiocyanates, are reported to
have biological activities including anticarcinogenic
activity (Verhoeven et al. 1997; Murillo and Mehta
2001; Balasinska et al. 2005). GLSs and their break-
down products may act as inhibitors of microbial
growth (Tierens et al. 2001), deterrents for herbivores,
and as attractants of specialist insects (Redovnikovic
et al. 2008b); moreover, these secondary metabolites
suppress soil-borne organisms such as bacteria, fungi,
viruses, nematodes and weeds (Tedeschi et al. 2011).
Particularly, the activity of isothiocyanates such as
sulforaphane against numerous human pathogens (e.g.
Escherichia coli,Salmonella typhimurium,Candida
sp.) could contribute to the medicinal properties
ascribed to Brassicaceae vegetables, such as cabbage
and mustard, which have been used as wound poul-
tices and antitumor agents for centuries (Hartwell
1982; Fahey et al. 2001). More recent interest has been
focussed on the potential anticarcinogenic activity of
GLS degradation products (Mithen 2001). The reason
of this increasing interest is due to the strong
correlation found between the consumption of Brass-
icaceae vegetables and the decreased risk for pancreas,
lung, stomach, prostate, and breast cancer, as recently
reported by Velasco et al. (2010). Particularly, several
studies have shown that allyl isothiocyanate may
inhibit different kinds of human prostate cancer
(Srivastava et al. 2003) or could contribute to the
lower incidence of bladder cancer (Munday 2002).
Horseradish contains more than 10 times as much
GLSs as broccoli. The high levels of sinigrin in
horseradish (Balasinska et al. 2005), easily hydrolyzed
by myrosinase to allyl isothiocyanate when the roots
are crushed (Matsuda et al. 2007), makes this root
interesting as a possible cancer-protecting component
in the diet (Talalay and Fahey 2001; Wedelsba
¨ck
Bladh and Olsson 2011). As reported by Shehata et al.
(2009), some German studies have investigated the
effects of horseradish on nonspecific urinary tract
infections and the antibacterial action of its essential
oils. A preparation with horseradish containing sini-
grin or allyl isothiocyanate was patented in 1995 for
topical application as a spray or applied with a swab
onto the affected mucosa, for treatment of nasal and
sinus dysfunction (Friedman 1995). Horseradish has
been approved in Germany for the treatment of
infections of the respiratory tract and as supportive
treatment in urinary tract infections. In the United
States, the root is the active ingredient of Rasapen
Ò
,a
urinary antiseptic drug (Shehata et al. 2009).
Recently, reported health benefits of horseradish
and its constituents include plasma cholesterol reduc-
tion in mice (Balasinska et al. 2005; Hara et al. 2008),
tumor cell proliferation and cyclooxygenase activity
prevention (Weil et al. 2005; Hara et al. 2008), as well
as an antimicrobial effect on Vibrio parahaemolyticus
by horseradish extract (Yano et al. 2006; Hara et al.
2008). Matsuda et al. (2007) showed that allyl
isothiocyanate provided significant protection against
ethanol-induced gastric lesions. Majewska et al.
(2004) have evaluated some antioxidant properties of
leaf and root extracts and mustard oil from four
different types of horseradish growing under different
environmental conditions: although leaf and root
extracts did not exhibit strong antioxidant properties,
the different environmental conditions affected these
properties significantly; also, volatile oil obtained
from horseradish roots revealed stronger antioxidant
properties than pure allyl isothiocyanate.
According to Mari et al. (2003) allyl isothiocyanate,
a naturally occurring flavour compound in mustard
and horseradish, has a well-documented antimicrobial
activity, therefore this volatile substance can be
employed successfully in modified atmosphere pack-
aging or as a gaseous treatment before storage. Also,
horseradish distillate is a potentially useful antimicro-
bial adjunct in packaged, refrigerated precooked
meats, in fact, essential oils of Brassicaceae plants,
including horseradish, have long been known to
possess antibacterial activity versus pathogenic
Genet Resour Crop Evol (2013) 60:1923–1943 1935
123
bacteria such as Staphylococcus aureus,Escherichia
coli and Serratia grimesii (Ward et al. 1998).
Khomvilai et al. (2006) observed the fungicidal
activities of allyl isothiocyanate against Saprolegnia
parasitica, a fish-pathogen oomycete, and determined
both minimum inhibitory concentration for mycelia
growth (68 mg/l with 60 min exposure) and zoospore
germination (42.5 mg/l with 5 min exposure).
Recently, first results of antifungal tests demonstrated
that horseradish ethanol extracts also present a fungi-
static activity against Sclerotium rolfsii Sacc., Fusar-
ium oxysporum Schlecht and Fusarium culmorum
(Wm. G. Sm.) Sacc. (Tedeschi et al. 2011).
Moreover, Park et al. (2006) have tested essential
oils from 40 plant species for their insecticidal
activities against larvae of Lycoriella ingenue (Du-
four) using a fumigation bioassay: horseradish showed
the most potent insecticidal activity among the plant
essential oils (only 1.25 ll/l caused 100 % mortality)
the most toxic compound to L. ingenue was allyl
isothiocyanate. In addition, also the results obtained
by Wu et al. (2009) suggest good insecticidal efficacy
of allyl isothiocyanate obtained from A. rusticana as
alternative fumigant against four major pest species of
stored products, maize weevil Sitophilus zeamais
(Motsch.), lesser grain borer Rhizopertha dominica
(F.), Tribolium ferrugineum (F.) and book louse
Liposcelis entomophila (Enderlein). Furthermore,
interesting and significant insecticidal activity against
larvae of Aedes albopictus (Skuse) (LC
50
value of
2.34 g/l) was obtained by using horseradish extract
prepared from fresh plants in a solution of ethanol
80 % (Tedeschi et al. 2011). Recently, Chen et al.
(2011) have assessed the fumigant toxicity of essential
oil from horseradish against Plodia interpunctella
(Hu
¨bner) and Sitophilus zeamais Motschulsky. The
essential oil of A. rusticana showed fumigant bioac-
tivity against all stages of P. interpunctella and adults
of S. zeamais. It caused high mortality of all stages of
P. interpunctella when exposed to 32 ll/l gas vapour
of A. rusticana. The oil appeared not to have much
impact on the adults of S. zeamais when the gas vapour
was lower than 16 ll/l, however, at 32 ll/l, the
percentage mortality was 100 %. GC–MS showed
that the main component of the essential oil from A.
rusticana was allyl isothiocyanate (98 %).
Armoracia rusticana oil or its major constituents
could be efficient fumigants and also could be
integrated with other pest management procedures
(Chen et al. 2011). For the practical use of horseradish
oil and their constituents as novel fumigants, further
study is necessary on the safety of these materials to
humans and on development of formulation to
improve efficacy and stability, and to reduce cost
(Park et al. 2006). Although higher concentrations of
these natural anti-fungal and insecticidal products
should be used, their intrinsically low toxicity towards
humans makes them an interesting alternative to
current chemicals. All these researches demonstrated
that horseradish extracts could be promising alterna-
tives to synthetic agrochemical products although
more work is needed (Tedeschi et al. 2011).
Another suggested use for horseradish includes
weed suppression via the allelopathic response of
GLSs in soil. GLSs and their byproducts can act as
biofumigants when used as green manure, thus
showing potential for weed control in organic pro-
duction (Shehata et al. 2009).
Genetic variability and influence of agronomic
management on glucosinolate content
The GLS content and composition in planta can be
strongly influenced by genotype and environment
(Rosa et al. 1997,2005; Sarikamis¸ et al. 2009;
Kabouw et al. 2010).
The genetic variability may play a primary role in
determining the amount of functional metabolites,
such as GLSs (Brown et al. 2002; De Pascale et al.
2007). Only few studies have investigated GLS
variation within different horseradish genotypes.
Among these, Li and Kushad (2004) have evaluated
the GLS levels in root and leaf samples of horseradish
accessions of a germplasm collection of the Vegetable
Research Farm, University of Illinois, Urbana-Cham-
paign, IL. They found that GLS content varied widely
among all accessions. According to Faltusova
´et al.
(2011), genotypes with high GLS contents will be
further utilized as a potential genetic source for
breeding. To date, information on genetic diversity
between different accessions of horseradish, are
scarce. Although molecular marker analyses are
successfully used for characterization of genetic
resources of many species, bio-agronomic character-
ization is the first step for the description and
classification of germplasm (Lotti et al. 2008; Sarli
et al. 2012).
1936 Genet Resour Crop Evol (2013) 60:1923–1943
123
Rhodes et al. (1969) have classified 20 genotypes of
horseradish, grown at the University of Illinois
Horticultural Farm, chosen from a gene pool contain-
ing over 400 genotypes cultivars to represent the
extreme and intermediate forms of morphological
variability found in horseradish. Two methods of
classification were compared. One classification was
based on 2 highly diagnostic characters (basal angle
and rugose index) that showed the extreme and
intermediate limits of the germplasm in form of a
scatter plot. The other classification was based on 40
characters. For this last classification, methods of
numerical taxonomy were used to show the germ-
plasm diversity by scatter diagrams and phenograms.
The two classifications appeared to be equal in
defining the extreme limits of the genetic variability,
although minor differences were found among the
relative positions of some genotypes. Sarli et al.
(2012) have recorded and statistically analyzed data
regarding the variation of 26 morphological descrip-
tors by using modified UPOV (International Union for
the Protection of New Varieties of Plants) guidelines
for performing tests for distinctness, uniformity and
stability for horseradish. In this study morphological
traits of 30 horseradish accessions from Basilicata
region (South Italy) were characterized and compared
to two European genotypes: the resulting cluster
displayed four major groups of genotypes, on the base
of the characteristics of rhizome and leaves. The
results showed a significant degree of variability
among accessions. Also, the results of univariate and
multivariate analysis showed that the investigated
accessions cannot be clearly separated according to
their place of origin, assuming that a distribution of an
accession is facilitated by territorial continuity and
similarity. According to Sarli et al. (2012), this
preliminary screening of a horseradish germplasm
collection is quite useful to perform future programs of
conservation in situ and ex situ, in order to expand and
improve the cultivation of A. rusticana. In any case,
there is a lack of basic biological and genetic
information about horseradish (Shehata et al. 2009).
The first studies, as reported above, could be a good
starting point to develop methods to measure genetic
diversity in different accessions of horseradish. The
genetic relationships among cultivars, breeding lines,
and wild accessions need to be determined, so that the
greatest possible gain can be made in specific crosses
(Shehata et al. 2009). Until today, only few studies
were performed on the characterization of genetic
diversity of horseradish. Various molecular marker
systems applied in plants belonging to the Brassica-
ceae family (Margale
´et al. 1995; Farnham 1996;
Rabbani et al. 1998; Das et al. 1999, Negi et al. 2004;
Faltusova
´et al. 2011) may be useful for a first
detection of polymorphisms in this species. However,
the morphological and genetic variation that seems to
exist among different horseradish accessions could be
compared with the GLS content of various horseradish
accessions for developing improved cultivars. In fact,
there is a growing interest in the metabolic engineering
of plants for the production of high-value bioactive
compounds, because the availability of phytochemi-
cals in their natural sources is often limited (Møldrup
et al. 2011). Therefore the information on genetic
diversity could be utilized in order to identify geno-
types with desirable traits for future breeding
programs.
There are even less studies on the effects of
environment and cultivation techniques on yield and
GLS content of this plant. Some researchers have
demonstrated that GLS profile can be manipulated by
cultural management (Rosen et al. 2005). About the
crop management, fertilization have been shown to
significantly affect GLS concentration in plant, partic-
ularly sulphur (S) and nitrogen (N) (Kim et al. 2002;
Rosen et al. 2005), but some controversy still exists on
such effects (Aires et al. 2006). Horseradish is a crop
that needs a lot of nutrients. Its powerful root system
drains large amounts of soil moisture and nutrient
reserves (Perlaki and Djurovka 2009). However, until
today, there is little information about the GLS content
in relation to S and N fertilization, with particular
regard to sprouts, young leaves and roots. S and N are
necessary for the synthesis of amino acids, proteins,
and various other cellular components, including thiol
compounds and the so-called secondary sulphur com-
pounds, which have a significant bearing on protection
of plants against stress and pests (Anjum et al. 2012).
GLS content is highly correlated with the S content
(Koroleva et al. 2000), therefore, there is a strong
potential for modifying GLS accumulation in crop
plants by altering S nutrition (Falk et al. 2007). Several
reports indicated that S fertilization may affect the GLS
level more than N fertilization (Rosen et al. 2005;De
Pascale et al. 2007). In fact, each GLS molecule
contains at least two sulfur atoms. One is part of the
sulfate group originating from 30-phosphoadenosine
Genet Resour Crop Evol (2013) 60:1923–1943 1937
123
50-phosphosulfate (PAPS), while the other, originating
from cysteine, is part of an S-linked glucose residue. In
addition, many GLSs are synthezised from methionine
and so may have a third sulfur atom in their structures
(Falk et al. 2007). S fertilization leads to increases in
GLS content in most cases. In some cases an over
10-fold increase was observed, as reported by Falk
et al. (2007) in their review about the effect of S supply
on GLS content.
Alnsour et al. (2012) showed recently that in vitro
grown horseradish plantlets contained increasing GLS
concentrations when the culture media was supple-
mented with additional sulfate. They concluded that
GLS concentrations in the stalks and leaves of
Armoracia in vitro plants could be modulated 20-fold
by varying the sulfate concentration in the medium,
while the increase of sulfate concentration had nearly
no impact on the GLS concentration in the roots.
The GLS content of a plant is dependent not only on
S nutrition, but also on the N level of the plant.
Particularly, through handling of fertility manage-
ment, the experimental results of Rosen et al. (2005)
clearly demonstrate that GLSs in cabbage were
maximized at low N and high S application rates. As
the ratio of N to S increases, GLS content declines,
probably because vegetative growth outpaces GLS
biosynthesis, diluting the content of these metabolites
(Kim et al. 2002; Rosen et al. 2005; Falk et al. 2007).
To quantify the effects of supply and combination of
these nutrients on GLS composition, Li et al. (2007)
showed as the total GLS concentration in turnip roots
was strongly influenced by N and S supply: with low S
supply (10 and 20 kg ha
-1
), total GLS concentrations
were 0.8–2.3 times higher at 80 kg N ha
-1
compared
to treatments with 160 and 240 kg N ha
-1
; however,
at the high S level (60 kg ha
-1
), increasing N supply
(160, 240, and 320 N ha
-1
) did not affect total GLS
concentration. In broccoli, Omirou et al. (2009) found
that the GLS concentration was low in the 50 kg ha
-1
N treatments in all plant organs, but it did not respond
to N application above 250 kg ha
-1
, while total GLS
concentrations, clearly responded to increasing S
applications within the whole wide range of S
applications (from 10 to 150 kg ha
-1
), indicating that
S is main determinant of the concentration of total
GLSs in the plant organs.
All these studies suggest that manipulating the N
and S supply might be one means of altering GLS level
in plants, thereby providing an opportunity to enhance
organoleptic properties and health benefits of horse-
radish, or its value as biofumigant (Falk et al. 2007;Li
et al. 2007).
To maintain proper nutrition, fertilizer is applied
based on results from chemical soil analysis, soil type,
cultivar, and cropping history (Shehata et al. 2009). As
reported in the review of Walters and Wahle (2010), in
A. rusticana a typical fertilizer program includes about
168 kg ha
-1
nitrogen (N), with phosphorus (P) and
potassium (K) rates based on soil type or soil test
results, with applications generally ranging from 56 to
280 kg ha
-1
. In addition, about 1–2.8 kg ha
-1
boron
and 17–28 kg ha
-1
sulfur is made with the initial N–
P–K broadcast application. Excessive nitrogen rates
should be avoided, as this can result in excessive foliar
growth and highly branched, irregular root formation
(Shehata et al. 2009). For the best response to
fertilizers, a pH range of 6–7.5 is ideal. The horserad-
ish plant thrives in deep loam or sandy soil types, well-
drained; organic matter is often added to maintain a
good soil structure, while shallow soils or those with
hard pans are not suitable as they compromise strong
root development and thus may curtail yield (Shehata
et al. 2009; Walters and Wahle 2010). Fields are
prepared for planting with ridgers to create raised beds
that increases yield of high-quality roots by ensuring
that the soil is loose so that large roots can develop.
Planting is done either by hand or using modified
transplanters that place sets (about 10 cm deep for
hand planting) in the beds at a 45-degree angle; the
spacing adopted is 40–60 cm between plants and
75–90 cm between rows, giving a plant population of
20,000–25,000 plants ha
-1
(Shehata et al. 2009).
Irrigation during drought conditions, especially during
late summer and fall, is sometimes done to improve
marketable yields because most root-sizing occurs
during this period (Walters and Wahle 2010). Usually,
horseradish is harvested in late October–November
once the foliage has been killed by frost, continuing
through the winter and early spring months (Shehata
et al. 2009; Walters and Wahle 2010).
Finally, to preserve the traditional knowledge
related to crop management, it is important to
underline that in regions like Bavaria (Germany) or
Hajdu
´sa
´g (Hungary), some cultivation practices (e.g.
removing sprouts after planting to obtain a root with a
single crown), even today, take place mainly by hand.
More studies are required for a better understanding
particularly of the effects of irrigation, fertilization
1938 Genet Resour Crop Evol (2013) 60:1923–1943
123
and time of harvest on yield and quality of horseradish
to optimize crop management.
Conclusion
Over recent years, the GLSs, a large group of nitrogen-
and sulfur-containing secondary metabolites, particu-
larly abundant in plants belonging to the Brassicaceae
and related families, have attracted the attention of the
scientific community for their fungicidal, bactericidal,
nematicidal, allelopathic and anticancerogenic proper-
ties. Among the Brassicaceae vegetables, horseradish is
particularly interesting because it is an exceptional rich
source of GLSs which could be employed for agro-
industrial and pharmaceutical purposes.
Considering this interest, more studies are needed
on qualitative and quantitative characterization of
GLSs, especially in relation to the variability of the
currently available horseradish germplasm, growing
conditions and crop management, plant phenological
phases and harvest dates.
In addition, it is important to stress that recently the
interest in natural substances has increased and
numerous studies on the biocidal activity of a wide
range of secondary metabolites have been reported, as
for example for the allyl isothiocyanate produced from
sinigrin with a well-documented antimicrobial activ-
ity. As reported by Mari et al. (2003), in Japan,
synthetic allyl isothiocyanate is registered as a food
additive and preliminary data showed that food
preserved with this compound contains only very
low residue of allyl isothiocyanate (Isshiki et al. 1992).
Therefore, natural extracts of horseradish could be
used as alternative to synthetic molecules.
For the great heritage about the gastronomic and
folk medical uses of horseradish, the species should be
valorized to prevent extinction of genotypes of high
nutritional value and specific taste and flavour and
farmers should continue to cultivate landraces in
traditional cultivation areas, maintaining the links
with territory and its history. In fact, the clonal
propagation of A. rusticana endangers its genetic
diversity, so efforts are required to establish well-
documented collections and to analyze genetic diver-
sity in all accessions available. In addition, legal
protection of geographic identity of horseradish land-
races could contribute to conservation of genetic
resources, and at same time, could increase the
viability of little farms. For example, by following a
centuries-old tradition, horseradish is today grown by
farmers in Bavaria and Hajdu
´sa
´g regions as an PGI
(Protected Geographical Indication) and PDO (Pro-
tected Designation of Origin), respectively, to preserve
the historical link with the territory. For small scale
growers, horseradish has potential marketing outlets
through fresh sales to local restaurants, grocery and
specialty food stores, and to specialized food process-
ing companies. They can also make sales at farmers
markets and roadside stands. For large scale produc-
tion, it is common for condiment processors to work on
a contract basis with growers. These markets can be
competitive and difficult to break into (Bratsch 2009).
Today, further investigations on the chemical,
biological and molecular aspects of the GLSs and
the composition and partitioning in the plant are
needed to achieve the goal of efficient biodiversity
conservation to prevent the genetic erosion of local
genotypes of horseradish and the risk to lose traits that
can be useful in future breeding programs.
Acknowledgments The authors are grateful to Mr. Vito
Agneta for assistance in graphic design images and Prof.
Ippolito Camele of the University of Basilicata for providing the
images in Fig. 2. The authors are also thankful to the Editors of
Economic Botany, Phytochemistry and Journal of Agricultural
and Food Chemistry for: (1) the use of the Table 1; (2) the adapt
of the Table 2—Copyright (1980) Elsevier Ltd and Table 3
Copyright (1979) Elsevier Ireland Ltd; (3) the permission to
adapted Tables 4and 5—Copyright (2004) American Chemical
Society, respectively.
References
Agneta R, Rivelli AR, Ventrella E, Lelario F, Sarli G, Bufo SA
(2012) Investigation of glucosinolate profile and qualita-
tive aspects in sprouts and roots of horseradish (Armoracia
rusticana) using LC-ESI -hybrid linear ion trap with
Fourier transform ion cyclotron resonance mass spec-
trometry and infrared multiphoton dissociation. J Agric
Food Chem 60(30):7474–7482. doi:10.1021/jf301294h
Aires A, Rosa E, Carvalho R (2006) Effect of nitrogen and sulfur
fertilization on glucosinolates in the leaves and roots of
broccoli sprouts (Brassica oleracea var. italica). J Sci Food
Agric 86(10):1512–1516. doi:10.1002/jsfa.2535
Alnsour M, Kleinwa
¨chter M, Bo
¨hme J, Selmar D (2012) Sulfate
determines the glucosinolate concentration of horseradish
in vitro plants (Armoracia rusticana Gaertn., Mey. &
Scherb.). J Sci Food Agric. doi:10.1002/jsfa.5825
Al-Shehbaz IA (1988) The genera of Arabideae (Cruciferae;
Brassicaceae) in the Southeastern United States. J Arnold
Arbor 69:85–166
Genet Resour Crop Evol (2013) 60:1923–1943 1939
123
Al-Shehbaz IA, Beilstein MA, Kellogg EA (2006) Systematics
and phylogeny of the Brassicaceae (Cruciferae): an over-
view. Plant Syst Evol 259(2–4):89–120. doi:10.1007/
s00606-006-0415-z
Anjum NA, Gill SS, Umar S, Ahmad I, Duarte AC, Pereira E
(2012) Improving growth and productivity of oleiferous
brassicas under changing environment: significance of
nitrogen and sulphur nutrition, and underlying mecha-
nisms. Sci World J. doi:10.1100/2012/657808
Balasinska B, Nicolle C, Gueux E, Majewska A, Demigne C,
Mazur A (2005) Dietary horseradish reduces plasma cho-
lesterol in mice. Nutr Res 25(10):937–945. doi:10.1016/j.
nutres.2005.09.015
Bellostas N, Sørensen AD, Sørensen JC, Sørensen H (2007a)
Genetic variation and metabolism of glucosinolates. Adv
Bot Res 45:369–415. doi:10.1016/S0065-2296(07)45013-3
Bellostas N, Kachlicki P, Sørensen JC, Sørensen H (2007b)
Glucosinolate profiling of seeds and sprouts of B. oleracea
varieties used for food. Sci Hortic 114(4):234–242. doi:10.
1016/j.scienta.2007.06.015
Blazevic I, Mastelic J (2009) Glucosinolate degradation prod-
ucts and other bound and free volatiles in the leaves and
roots of radish (Raphanus sativus L.). Food Chem
113:96–102. doi:10.1016/j.foodchem.2008.07.029
Bones AM, Rossiter JT (2006) The enzymic and chemically
induced decomposition of glucosinolates. Phytochemistry
67(11):1053–1067. doi:10.1016/j.phytochem.2006.02.024
Borgen BH, Thangstad OP, Ahuja I, Rossiter JT, Bones AM
(2010) Removing the mustard oil bomb from seeds:
transgenic ablation of myrosin cells in oilseed rape
(Brassica napus) produces MINELESS seeds. J Exp Bot
61(6):1683–1697. doi:10.1093/jxb/erq039
Bostock J, Riley HT (1856) The natural history of Pliny, vol IV.
H.G. Bohn, London
Bratsch A (2009) Specialty crop profile: horseradish. Virginia
cooperative extension. http://pubs.ext.vt.edu/438/438-104/
438-104.html
Brown AF, Yousef GG, Jeffery EH, Klein BP, Wallig MA,
Kushad MM, Juvik JA (2002) Glucosinolate profiles in
broccoli: variation in levels and implications in breeding
for cancer chemoprotection. J Am Soc Hortic Sci 127(5):
807–813
Brown PD, Tokuhisa JG, Reichelt M, Gershenzon J (2003)
Variation of glucosinolate accumulation among different
organs and developmental stages of Arabidopsis thaliana.
Phytochemistry 62:471–481. doi:10.1016/S0031-9422(02)
00549-6
Cataldi TRI, Rubino A, Lelario F, Bufo SA (2007) Naturally
occurring glucosinolates in plant extracts of Rocket Salad
(Eruca sativa L.) identified by liquid chromatography
coupled with negative ion electrospray ionization and
quadrupole ion-trap mass spectrometry. Rapid Commun
Mass Spectrom 21(14):2374–2388
Chen H, Akinkurolere RO, Zhang H (2011) Fumigant activity of
plant essential oil from Armoracia rusticana (L.) on Plodia
interpunctella (Lepidoptera: Pyralidae) and Sitophilus
zeamais (Coleoptera: Curculionidae). Afr J Biotechnol
10(7):1200–1205. doi:10.5897/AJB10.2023
Clarke B (2010) Glucosinolates, structures and analysis in food.
Anal Methods 2:310–325. doi:10.1039/B9AY00280D
Cleemput S, Becker HC (2011) Genetic variation in leaf and
stem glucosinolates in resynthesized lines of winter rape-
seed (Brassica napus L.). Genet Resour Crop Evol. 59:
539–546
Clossais-Besnard N, Larher F (1991) Physiological role of
glucosinolates in Brassica napus. Concentration and dis-
tribution pattern of glucosinolates among plant organs
during a complete life cycle. J Sci Food Agric 56:25–38.
doi:10.1002/jsfa.2740560104
Courter JW, Rhodes AM (1969) Historical notes on horseradish.
Econ Bot 23:156–164. doi:10.1007/BF02860621
Das S, Rajagopal J, Bhatia S, Srivastava PS, Lakshmikumaran
M (1999) Assessment of genetic variation within Brassica
campestris cultivars using amplified fragment length
polymorphism and random amplification of polymorphic
DNA markers. J Biosci 24(4):433–440
D’Auria M, Mauriello G, Raciotti R (2004) SPME-GC-MS
analysis of horseradish (Armoracia rusticana). Ital J Food
Sci 4(16):487–490. ISSN: 1120-1770
De Candolle A (1890) Origin of cultivated plants. Appleton and
Co., New York
De Pascale S, Maggio A, Pernice R, Fogliano V, Barbieri G
(2007) Sulphur fertilization may improve the nutritional
value of Brassica rapa L. subsp. sylvestris. Eur J Agron
26:418–424. doi:10.1016/j.eja.2006.12.009
Fahey JW, Zalcmann AT, Talalay P (2001) The chemical
diversity and distribution of glucosinolates and isothiocy-
anates among plants. Phytochemistry 56(1):5–51. doi:10.
1016/S0031-9422(00)00316-2
Falk KL, Tokuhisa JG, Gershenzon J (2007) The effect of sulfur
nutrition on plant glucosinolate content: physiology and
molecular mechanisms. Plant Biol 9:573–581. doi:10.
1055/s-2007-965431
Faltusova
´Z, Kuc
ˇera L, Ovesna
´J (2011) Genetic diversity of
Brassica oleracea var. capitata gene bank accessions
assessed by AFLP. Electron J Biotechnol 14(3):1–10.
doi:10.2225/vol14-issue3-fulltext-4
Farnham MW (1996) Genetic variation among and within
United States collard cultivars and landraces as determined
by randomly amplified polymorphic DNA markers. J Am
Soc Hortic Sci 121:374–379
Friedman WH (1995) Horseradish preparation for the treatment
of nasal and sinus dysfunction. US Paatent 5385734
Grob K, Matile PH (1979) Vacuolar location of glucosinolatea
in horseradish root cells. Plant Sci Lett 14(4):327–335.
doi:10.1016/S0304-4211(79)90281-5
Grob K, Matile PH (1980) Capillary GC of glucosinolate-
derived horseradish constituents. Phytochemistry 19(8):
1789–1793. doi:10.1016/S0031-9422(00)83814-5
Guarrera PM (2006) Usi e tradizioni della Flora Italiana. Med-
icina popolare ed etnobotanica. Aracne editrice s.r.l, Rome
Hammer K, Knu
¨pffer H, Perrino P (1990) A checklist of the
South Italian cultivated plants. Kulturpflanze 38:91–310
Hammer K, Knu
¨pffer H, Laghetti G, Perrino P (1992) Seeds from
the past. A catalogue of crop germplasm in South Italy and
Sicily. Germplasm Institute of C.N.R (ed). Bari, Italy, p 173
Hammer K, Laghetti G, Pignone D (2011) The meeting of two
cultures and their agricultures. In: Linguistic Islands and
plant genetic resources—the case of Arbe
¨reshe
¨. Aracne ed.
s.r.l., Rome, pp 253–271
1940 Genet Resour Crop Evol (2013) 60:1923–1943
123
Hanelt P, Institute of Plant Genetics and Crop Plant Research
(eds) (2001) Mansfeld’s encyclopedia of agricultural and
horticultural crops. Springer, Berlin, pp 1419–1420
Hara M, Oda M, Yogo T, Sumi T, Arai R, Kuboi T, Etoh H
(2008) Detection of horseradish (Armoracia rusticana)
myrosinase genes in samples containing horseradish. Food
Sci Technol Res 14(4):389–394. doi:10.3136/fstr.14.389
Hartwell JL (1982) Plants Against Cancer: A Survey. Quar-
terman Publications, Lawrence
Helmlinger J, Rausch T, Hilgenberg W (1983) Localization of
newly synthesized indoIe-3-methylglucosinolate (= gluco-
brassicin) in vacuoles from horseradish (Armoracia rusti-
cana). Physiol Plant 58(3):302–310. doi:10.1111/j.1399-
3054.1983.tb04185.x
Isshiki K, Tokuora K, Mori R, Chiba S (1992) Preliminary
examination of allyl isothiocyanate vapor for food preser-
vation. Biosci Biotechnol Biochem 56:1476–1477
Jiang ZT, Li R, Yu JC (2006) Pungent components from thio-
glucosides in Armoracia rusticana grown in China,
obtained by enzymatic hydrolysis. Food Technol Bio-
technol 44(1):41–45. ISSN 1330-9862
Kabouw P, Biere A, Van der Putten WH, Van Dam NM (2010)
Intra-specific differences in root and shoot glucosinolate
profiles among white cabbage (Brassica oleracea var.
capitata) cultivars. J Agric Food Chem 58(1):411–417.
doi:10.1021/jf902835k
Kamada H, Tachikawa Y, Saitou T, Harada H (1995) Effects of
light and growth regulators on adventitious bud formation
in horseradish (Armoracia rusticana). Plant Cell Rep
14(10):611–615. ISSN: 0721-085X
Khomvilai C, Kashiwagi M, Yoshioka M (2006) Fungicidal
activities of horseradish extract on a fish-pathogen oomy-
cetes, Saprolegnia parasitica. Bull Fac Bioresources
33:1–7. ISSN: 0915-0471
Kim SJ, Matsuo T, Watanabe M, Watanabe Y (2002) Effect of
nitrogen and sulphur application on the glucosinolates
content in vegetable turpin rape (Brassica rapa L.). Soil Sci
Plant Nutr 48:43–49. doi:10.1080/00380768.2002.
10409169
Kissen R, Rossiter J, Bones A (2009) The ‘mustard oil bomb’:
not so easy to assemble?! Localization, expression and
distribution of the components of the myrosinase enzyme
system. Phytochem Rev 8(1):69–86. doi:10.1007/s11101-
008-9109-1
Kliebenstein DJ, Kroyman J, Brown P, Figuth A, Pedersen D,
Gershenzon J, Mitchell-Olds T (2001) Genetic control of
natural variation in Arabidopsis glucosinolate accumulation.
Plant Physiol 126:811–825. doi:10.1104/pp.126.2.811
Koroleva OA, Davies A, Deeken R, Thorpe MR, Tomos AD,
Hedrich R (2000) Identification of a new glucosinolate-rich
cell type in Arabidopsis flower stalk. Plant Physiol
124:599–608. doi:10.1104/pp.124.2.599
Kosson R, Horbowicz M (2008) Effect of long term storage on
some nutritive components and isothiocyanates content in
roots of two horseradish types. Veg Crops Res Bull
69:155–164. doi:10.2478/v10032-008-0030-3
Kosson R, Horbowicz M (2009) Some quality characteristics
including isothiocyanates content of horseradish cream as
affected by storage period. Veg Crops Res Bull 71:123–
132. doi:10.2478/v10032-009-0033-8
La Rocca F, Chisci G (2005) Il Cren. Rafano Rusticano o Bar-
baforte, Libreria Editrice Fiorentina. ISBN 88-89264-61-6
Lee KC, Cheuk MW, Chan W, Ming AW, Zhao ZZ, Jiang ZH,
Cai Z (2006) Determination of glucosinolates in traditional
Chinese herbs by high-performance liquid chromatography
and electrospray ionization mass spectrometry. Anal Bio-
anal Chem 386:2225–2232. doi:10.1007/s00216-006-
0882-7
Lelario F, Bianco G, Bufo SA, Cataldi TRI (2012) Establishing
the occurrence of major and minor glucosinolates in
Brassicaceae by LC–ESI-hybrid linear ion-trap and Fou-
rier-transform ion cyclotron resonance mass spectrometry.
Phytochemistry 73:74–83. doi:10.1016/j.phytochem.2011.
09.010
Li X, Kushad MM (2004) Correlation of glucosinolate content
to myrosinase activity in horseradish (Armoracia rustica-
na). J Agric Food Chem 52:6950–6955. doi:10.1021/
jf0401827
Li S, Schonhof I, Krumbein A, Li L, Stu
¨tzel H, Schreiner M
(2007) Glucosinolate concentration in turnip (Brassica
rapa ssp. rapifera L.) roots as affected by nitrogen and
sulfur supply. J Agric Food Chem 55:8452–8457. doi:10.
1021/jf070816k
Lotti C, Marcotrigiano AR, De Giovanni C, Resta P, Ricciardi
A, Zonno V, Fanizza G, Ricciardi L (2008) Univariate and
multivariate analysis performed on bio-agronomical traits
of Cucumis melo L. germplasm. Genet Resour Crop Evol
55:511–522. doi:10.1007/s10722-007-9257-y
Łuczaj Ł, Szyman
´ski WM (2007) Wild vascular plants gathered
for consumption in the Polish countryside: a review.
J Ethnobiol Ethnomed 3:17. doi:10.1186/1746-4269-3-17
Majewska A, Bałasin
´ska B, Da˛browskaB (2004) Antioxidant
properties of leaf and root extract and oil from different
types of horseradish (Armoracia rusticana Gaertn.). Folia
Horticulturae 16(1):15–22
Margale
´E, Herve
´Y, Hue J, Quiros CF (1995) Determination of
genetic variability by RAPD markers in cauliflower, cab-
bage and kale local cultivars from France. Genet Resour
Crop Evol 42(3):281–289. doi:10.1007/BF02431263
Mari M, Bertolini P, Pratella GC (2003) Non-conventional
methods for the control of post-harvest pear diseases.
J Appl Microbiol 94:761–766. doi:10.1046/j.1365-2672.
2003.01920.x
Matsuda H, Ochi M, Nagatomo A, Yoshikawa M (2007) Effects
of allylisothiocyanate from horseradish on several experi-
mental gastric lesions in rats. Eur J Pharmacol
561(1–3):172–181. doi:10.1016/j.ejphar.2006.12.040
McCann J (2004) The horseradish plant. www.globalgourmet.
com/food/egg/egg1296/horsplnt.html
Mevy JP, Rabier J, Quinsac A, Krouti M, Ribaillier D (1997)
Glucosinolate contents of regenerated plantlets from em-
bryoids of horseradish. Phytochemistry 44(8):1469–1471.
doi:10.1016/S0031-9422(96)00759-5
Mevy JP, Rabier J, Quinsac A, Ribaillier D (1999) Sucrose
metabolism and indoleglucosinolate production of immo-
bilized horseradish cells. Plant Cell, Tissue Organ Cult
57(3):163–171. doi:10.1023/A:1006325222785
Mithen R (2001) Glucosinolates—biochemistry, genetics and
biological activity. Plant Growth Regul 34(1):91–103.
doi:10.1023/A:1013330819778
Genet Resour Crop Evol (2013) 60:1923–1943 1941
123
Mithen RF, Dekker M, Verkerk M, Rabot S, Johnson I (2000)
The nutritional significance, biosynthesis and bioavail-
ability of glucosinolates in human foods. J Sci Food Agric
80:967–984. doi:10.1002/(SICI)1097-0010(20000515)80:
7\967:AID-JSFA597[3.0.CO;2-V
Moerman DE (1998) Native American etnobotany. Timber
Press, Portland
Møldrup ME, Geu-Flores F, Olsen CE, Halkier BA (2011)
Modulation of sulfur metabolism enables efficient gluco-
sinolate engineering. BMC Biotechnol 11:12. doi:10.1186/
1472-6750-11-12
Mucete D, Radu F, Poinana M, Jianu I (2006) Myrosinase
activity in Armoracia rusticana. Bull USAMV-CN 62:
88–93
Munday CM (2002) Selective induction of phase II enzymes in
the urinary bladder of rats by allyl isothiocyanate, a com-
pound derived from brassica vegetables. Nutr Cancer
44(1):52–59. doi:10.1207/S15327914NC441_7
Murillo G, Mehta R (2001) Cruciferous vegetables and cancer
prevention. Nutr Cancer 41:17–28. doi:10.1080/01635581.
2001.9680607
Negi MS, Sabharwal V, Bhat SR, Lakshmikumaran M (2004)
Utility of AFLP markers for the assessment of genetic
diversity within Brassica nigra germplasm. Plant Breed
123(1):13–16. doi:10.1046/j.0179-9541.2003.00926.x
Omirou MD, Papadopoulou KK, Papastylianou I, Constantinou
M, Karpouzas DG, Asimakopoulos I, Ehaliotis C (2009)
Impact of nitrogen and sulfur fertilization on the compo-
sition of glucosinolates in relation to sulfur assimilation in
different plant organs of broccoli. J Agric Food Chem
57:9408–9417. doi:10.1021/jf901440n
Park IK, Choi KS, Kim DH, Choi IH, Kim LS, Bak WC, Choi
JW, Shin SC (2006) Fumigant activity of plant essential
oils and components from horseradish (Armoracia rusti-
cana), anise (Pimpinella anisum) and garlic (Allium sati-
vum) oils against Lycoriella ingenua (Diptera: Sciaridae).
Pest Manag Sci 62(8):723–728. doi:10.1002/ps.1228
Perlaki Z, Djurovka M (2009) Horseradish root yield depending
on organic and mineral fertilizers. Contemp Agric 58:
106–111. ISSN 0350-1205
Pieroni A, Gray C (2008) Herbal and food folk medicines of the
Russlanddeutschen living in Ku
¨nzelsau/Tala
¨cker, South-
Western Germany. Phytother Res 22:889–890. doi:10.1002/
ptr.2410
Pieroni A, Quave CL (2005) Traditional pharmacopoeias and
medicines among Albanians and Italians in southern Italy:
a comparison. J Ethnopharmacol 101:258–270. doi:10.
1016/j.jep.2005.04.028
Pieroni A, Quavec CL, Santorod RF (2004) Folk pharmaceutical
knowledge in the territory of the Dolomiti Lucane, inland
southern Italy. J Ethnopharmacol 95:373–384. doi:10.
1016/j.jep.2004.08.012
Pieroni A, Nebel S, Santoro RF, Heinrich M (2005) Food for two
seasons: culinary uses of non-cultivated local vegetables
and mushrooms in a south Italian village. Int J Food Sci
Nutr 56(4):245–272. doi:10.1080/09637480500146564
Rabbani MA, Murakami Y, Kuginuki Y, Takayanagi K (1998)
Genetic variation in radish (Raphanus sativus L.) germ-
plasm from Pakistan using morphological traits and
RAPDs. Genet Resour Crop Evol 45:307–316. doi:10.
1023/A:1008619823434
Redovnikovic IR, Peharec P, Rasol MK, Delonga K, Brkic K,
Vorkapic-Furac J (2008a) Glucosinolate profiles, myrosi-
nase and peroxidase activity in horseradish (Armoracia
lapathifolia Gilib.) Plantlets, Tumour and Teratoma Tis-
sues Food Technol Biotechnol 46(3):317–321. ISSN
1330-9862
Redovnikovic IR, Glivetic T, Delonga K, Vorkapic-Furac J
(2008b) Glucosinolates and their potential role in plant.
Periodicum Biologorum 110(4):297–309. ISSN 0031-5362
Rhodes AM, Carmer SG, Courter J (1969) Measurement and
classification of genetic variability in horseradish. J Am
Soc Hortic Sci 94:98–102
Rosa AS, Heaney R, Fenwick G, Portas C (1997) Glucosinolates
in crop plants. In: Janick J (ed) Horticultural reviews, vol
19. USA, pp 99–215
Rosen CJ, Fritz VA, Gardner GM, Hecht SS, Carmella SG,
Kenney PM (2005) Cabbage yield and glucosinolate con-
centrations as affected by nitrogen and sulfur fertility.
HortScience 40:1493–1498
Rosengarten F Jr (1969) The book of the spices. Livingston Pub.
Co., Wynnewood
Sahasrabudhe MR, Mullin WJ (1980) Dehydration of horse-
radish roots. J Food Sci 45:1440–1441. doi:10.1111/j.
1365-2621.1980.tb06577.x
Sampliner D, Miller A (2009) Ethnobotany of horseradish (Ar-
moracia rusticana, Brassicaceae) and its wild relatives
(Armoracia spp.): reproductive biology and local uses in
their native ranges. Econ Bot 63(3):303–313. doi:10.1007/
s12231-009-9088-1
Sarikamis¸ G, Balkaya A, Yanmaz R (2009) Glucosinolates within
a collection of white head cabbages (Brassica oleracea var.
capitata subvar. alba) from Turkey. Afr J Biotechnol
8(19):5046–5052. doi:10.2225/vol14-issue3-fulltext-4
Sarli G, De Lisi A, Agneta R, Grieco S, Ierardi G, Montemurro
F, Negro D, Montesano V (2012) Collecting horseradish
(Armoracia rusticana, Brassicaceae): local uses and mor-
phological characterization in Basilicata (Southern Italy)
Genet Resour Crop Evol 59(5): 889–899. doi:10.1007/
s10722-011-9730-5
Schaffer A (1981) The history of horseradish as the bitter herb of
Passover. Gesher 8:217–237
Shehata AM, Skirvin RM, Norton MA (2008) Leaf morphology
affects horseradish regeneration in vitro. Int J Veg Sci
15(1):24–27. doi:10.1080/19315260802446385
Shehata AM, Mulwa RMS, Babadoost M, Uchanski M, Norton
MA, Skirvin R, Walters SA (2009) Horseradish: botany,
horticulture, breeding, in horticultural reviews. In: Janick J
(ed) vol 35. Wiley, Hoboken. doi:10.1002/978047
0593776.ch4
Srivastava SK, Xiao D, Lew KL, Hershberger P, Kokkinakis
DM, Johnson CS, Trump DL, Singh SV (2003) Allyl iso-
thiocyanate, a constituent of cruciferous vegetables,
inhibits growth of PC-3 human prostate cancer xenografts
in vivo. Carcinogenesis 24(10):1665–1670. doi:10.1093/
carcin/bgg123
Sultana T, Savage GP, McNeil DL, Porter NG, Clark B (2003)
Comparison of flavour in wasabi and horseradish. J Food
Agric Environ 1(2):117–121
Talalay P, Fahey JW (2001) Phytochemicals from cruciferous
plants protect against cancer by modulating carcinogen
metabolism. J Nutr 131(11):3027S–3033S
1942 Genet Resour Crop Evol (2013) 60:1923–1943
123
Tedeschi P, Leis M, Pezzi M, Civolani S, Maietti A, Brandolini
V (2011) Insecticidal activity and fungitoxicity of plant
extracts and components of horseradish (Armoracia rusti-
cana) and garlic (Allium sativum). J Environ Sci Health B
46(6):486–490. doi:10.1080/03601234.2011.583868
Tierens KFMJ, Thomma BPHJ, Brouwer M, Schmidt J, Kistner
K, Porzel A, Mauch-Mani B, Cammue BPA, Broekaert WF
(2001) Study of the role of antimicrobial glucosinolate-
derived isothiocyanates in resistance of Arabidopsis to
microbial pathogens. Plant Physiol 125:1688–1699.
doi:10.1104/pp.125.4.1688
Velasco P, Cartea ME, Gonza
´lez C, Vilar M, Orda
´s M (2007)
Factors affecting the glucosinolate content of kale (Bras-
sica oleracea acephala group). J Agric Food Chem
55(3):955–962. doi:10.1021/jf0624897
Velasco P, Francisco M, Cartea ME (2010) Glucosinolates in
Brassica and cancer. In: Watson RR, Preedy VR (eds)
Bioactive foods and extracts. Cancer treatment and pre-
vention, 1st edn. CRC Press, USA, pp 3–29. doi:10.1201/
b10330-3
Verhoeven DTH, Verhagen H, Goldbohm RA, Brant PAVD,
Poppel GV (1997) A review of mechanisms underlying
anticarcinogenicity by brassica vegetables. Chem Biol
Interact 103:79–129. doi:10.1016/S0009-2797(96)03745-3
Vetela
¨inen M, Negri V, Maxted N (2009) European landraces:
on-farm conservation, management and use. Bioversity
Technical Bulletin No. 15. Bioversity International, Rome,
Italy. ISBN 978-92-9043-805-2
Walters SA, Wahle EA (2010) Horseradish production in Illi-
nois. HorTecnology 20:267–276
Ward SM, Delaquis PJ, Holley RA, Mazza G (1998) Inhibition
of spoilage and pathogenic bacteria on agar and pre-cooked
roast beef by volatile horseradish distillates. Food Res Int
31(1):19–26. doi:10.1016/S0963-9969(98)00054-4
Weber WW (1949) Seed production in horseradish. J Hered
40:223–227
Wedelsba
¨ck Bladh K, Olsson KM (2011) Introduction and use
of horseradish (Armoracia rusticana) as food and medicine
from antiquity to the present: emphasis on the Nordic
countries. J Herbs Spices Med Plants 17(3):197–213.
doi:10.1080/10496475.2011.595055
Weil MJ, Zhang Y, Nair MG (2005) Tumor cell proliferation
and cyclooxygenase inhibitory constituents in horseradish
(Armoracia rusticana) and wasabi (Wasabia japonica).
J Agric Food Chem 53(5):1440–1444. doi:10.1021/jf
048264i
Winiarczyk K, Bednara J (2008) The progamic phase and seed
formation in Armoracia rusticana. Plant Breed 127(2):
203–207. doi:10.1111/j.1439-0523.2007.01439.x
WiniarczykK, Tcho
`rzewska D, Bednara J (2007) Development of
the male gametophyte of an infertile plant Armoracia rusti-
cana Gaertn. Plant Breed 126(4):433–439. doi:10.1111/j.
1439-0523.2007.01365.x
Wittstock U, Halkier BA (2002) Glucosinolate research in the
Arabidopsis era. Trends Plant Sci 7(6):263–270. doi:10.
1016/S1360-1385(02)02273-2
Wu H, Zhang GA, Zeng S, Lin KC (2009) Extraction of allyl
isothiocyanate from horseradish (Armoracia rusticana)
and its fumigant insecticidal activity on four stored-product
pests of paddy. Pest Manag Sci 65(9):1003–1008. doi:10.
1002/ps.1786
Yang B, Quiros CF (2010) Survey of glucosinolate variation in
leaves of Brassica rapa crops. Genet Resour Crop Evol
57(7):1079–1089. doi:10.1007/s10722-010-9549-5
Yano Y, Satomi M, Oikawa H (2006) Antimicrobial effect of
spices and herbs on Vibrio parahaemolyticus. Int J Food
Microbiol 111(1):6–11. doi:10.1016/j.ijfoodmicro.2006.
04.031
Genet Resour Crop Evol (2013) 60:1923–1943 1943
123
... Sinigrin is the predominant glucosinolate found in horseradish and accounts for approximately 74% of the total glucosinolate content of the plant. The individual isothiocyanate in horseradish root includes allyl isothiocyanate (AITC, 96 mg/100 g), n-butyl isothiocyanate (0.45 mg/100 g), 2-phenylethyl isothiocyanate (22 mg/100 g), 3-butenyl isothiocyanate (0.85 mg/100 g), 4-pentenyl isothiocyanate (0.1 mg/100 g) (Agneta et al., 2013, Nguyen et al., 2013. AITC and 2-phenylethyl isothiocyanate are the most versatile and functionally important compounds present in horseradish. ...
... The glucosinolate content in nearly all the crucifers can be significantly enhanced by the enrichment of the growth medium with nitrogen and sulfur-containing chemicals. However, sulfur fertilization has been reported to increase the glucosinolate concentration more effectively than nitrogen fertilization (Agneta et al., 2013). Isothiocyanates are efficient anticarcinogenic agents that inhibit the development of cancer cells, eliminate developed cancer cells, inhibit carcinogen-activating enzymes, induce carcinogen-detoxifying enzymes, arrest the cell cycle progression in G1, S, G2-M phases, reduce oxidative stress by elevating and maintaining cellular antioxidants, retard or eliminate the clonal expansion of initiated, transformed, and/or neoplastic cells, inhibit P450 monooxygenase, activate caspase-3, -8, -9, or -12 leading to increased apoptosis (Zhang, 2004). ...
... Isothiocyanates are efficient anticarcinogenic agents that inhibit the development of cancer cells, eliminate developed cancer cells, inhibit carcinogen-activating enzymes, induce carcinogen-detoxifying enzymes, arrest the cell cycle progression in G1, S, G2-M phases, reduce oxidative stress by elevating and maintaining cellular antioxidants, retard or eliminate the clonal expansion of initiated, transformed, and/or neoplastic cells, inhibit P450 monooxygenase, activate caspase-3, -8, -9, or -12 leading to increased apoptosis (Zhang, 2004). Some other traditional applications of the crop are also well reported, for example, in the US, the root is the active ingredient of Rasapen, a commercial urinary antiseptic drug (Agneta et al., 2013). ...
Chapter
The sedentary lifestyle coupled with continuously changing food habits and search for nutrient-dense enriched protective foods has resulted in increased demand and consumption of natural foods. Among natural dietary ingredients, vegetables are the daily consumed dietary ingredients packed with vitamins, minerals, antioxidants, and an array of bioactive phytochemicals. Amongst vegetable crops, cruciferous vegetables like broccoli, cabbage, cauliflower, arugula, horseradish, mustard green, bok choy, brussels sprouts, etc. are the crops which are perceived far important than the mere table items for daily consumption owing to their rich functional bioactive profile. Presence of the sulfur-rich compounds (methyl cysteine sulfoxide and glucosinolates), coloring pigments (carotenoids, anthocyanins), minerals (Se, Fe, K, Ca), vitamins (B complex and C), dietary fiber, and other bioactive compounds (phytoalexins, terpenes, tocopherols, hydroxycinnamic acid, chlorogenic acid, ferulic acid, synapic acid and flavonols) give them the distinctive nutraceutical status with well documented therapeutic benefits. Most of the research effort in the last decade has been directed to effectively find out the exact mode of action of these bioactive compounds on health with their minimum effective concentration, means to ensure their effective delivery to the target organs, and increased bioavailability of these compounds. Though, there is substantial evidence based on in vivo and in vitro findings that scientifically demonstrate their benefits more research needs to be conducted with an exploration of the unknown beneficial activities as well as the unwanted effects. Future research should be directed towards the functional enrichment either through genetic modifications or through regulation of pathways for ensuring the national and health security to the general population and the health-conscious people.
... The pungency of horseradish roots results from the sulfur-containing glucosinolates in the tissues that break down into isothiocyanates [6,7], although this plant contains many other nutraceutical compounds. The intense pungency of horseradish roots is primarily caused by isothiocyanate compounds (mostly sinigrin and 2phenylethylglucosinolate) that result from the hydrolysis of glucosinolates by the naturally occurring enzyme myrosinase [6,8]. ...
... The pungency of horseradish roots results from the sulfur-containing glucosinolates in the tissues that break down into isothiocyanates [6,7], although this plant contains many other nutraceutical compounds. The intense pungency of horseradish roots is primarily caused by isothiocyanate compounds (mostly sinigrin and 2-phenylethylglucosinolate) that result from the hydrolysis of glucosinolates by the naturally occurring enzyme myrosinase [6,8]. ...
... Sulfur-containing glucosinolates provide the bitter flavor and pungent characteristic of horseradish, as a result of their breakdown into isothiocyanates [14][15][16]. Singirin accounts for about >80% of the total glucosinolates content in horseradish roots [7,15]. Once horseradish root tissues are ground or crushed, sinigrin (or other glucosinolates) mix with myrosinase, and pungent volatile allyl compounds (isothiocyanates) are produced [1,16]. ...
Article
Full-text available
Horseradish is a flavorful pungent herb that has been used for centuries to enhance the flavor of food, aid in digestion, and improve human health. Horseradish is a neglected and underutilized plant species (NUS), especially concerning the potential benefits to improve human health. The roots of this plant have been known for centuries to provide effective treatments for various human health disorders and has a long history of use in traditional medicine. Horseradish is a source of many biologically active compounds and its richness in phytochemicals has encouraged its recent use as a functional food. The medicinal benefits of horseradish are numerous, and this plant should be promoted more as being beneficial for human health. Glucosinolates or their breakdown products, isothiocyanates, are responsible for most of the claimed medicinal effects. Recent studies have suggested that glucosinolates provide prevention and inhibitory influences on different types of cancer, and horseradish contains high amounts of these compounds. Other medicinal benefits of horseradish include its well-known antibacterial properties that are also attributed to isothiocyanates, and its high content of other antioxidants that benefit human health. Additionally, horseradish contains enzymes that stimulate digestion, regulate bowel movement, and reduce constipation. Horseradish is a species that is vastly underexploited for its abilities as a medicinal plant species for improving human health. The health promoting effects of horseradish are numerous and should be used in an extensive marketing campaign to improve consumption habits. Consumers need to be made more aware of the tremendous health benefits of this plant, which would most likely increase consumption of this valuable NUS. Although horseradish is a highly versatile plant species and holds great potential for improving human health, this plant can also be used to enhance biodiversity in landscapes and food systems, which will also be briefly discussed.
... Armoracia rusticana has been extensively used since ancient times for its nutritional value and as a culinary spice [43]. Traditionally, fresh and dried roots have been consumed as a condiment as a paste or sauce, containing grated root, acetic acid from vinegar, and salt. ...
... However, due to its strong organoleptic properties-which can lead to serious side effects-the use of high concentration of horseradish root is limited [46]. Moreover, A. rusticana is well known as a folk medicinal herb [43]. In Europe, it was traditionally used to treat gout, kidney stones, asthma, and bladder infections. ...
... Among them, isothiocyanates (ITCs) are the dominant components, followed by nitriles and/or thiocyanates. Thus, the typical sharp taste and smell of horseradish roots is mainly attributed to sinigrin-derived allyl isothiocyanate (AITC) Figure 1(8) and gluconasturtiinderived 2-phenylethyl isothiocyanate (PEITC) Figure 1 (9) [39,43,47]. ...
Article
Full-text available
Since ancient times, plant roots have been widely used in traditional medicine for treating various ailments and diseases due to their beneficial effects. A large number of studies have demonstrated that—besides their aromatic properties—their biological activity can often be attributed to volatile constituents. This review provides a comprehensive overview of investigations into the chemical composition of essential oils and volatile components obtained from selected aromatic roots, including Angelica archangelica, Armoracia rusticana, Carlina sp., Chrysopogon zizanioides, Coleus forskohlii, Inula helenium, Sassafras albidum, Saussurea costus, and Valeriana officinalis. Additionally, their most important associated biological impacts are reported, such as anticarcinogenic, antimicrobial, antioxidant, pesticidal, and other miscellaneous properties. Various literature and electronic databases—including PubMed, ScienceDirect, Springer, Scopus, Google Scholar, and Wiley—were screened and data was obtained accordingly. The results indicate the promising properties of root-essential oils and their potential as a source for natural biologically active products for flavor, pharmaceutical, agricultural, and fragrance industries. However, more research is required to further establish the mechanism of action mediating these bioactivities as well as essential oil standardization because the chemical composition often strongly varies depending on external factors.
... Similarly, very little information is available for watercress, with only four papers published in the last 40 years [17][18][19][20]. Horseradish is the most well characterised of these four species, but still, only six studies of note have been published in the last 50 years [6,[21][22][23][24][25]. ...
... The compound was also present in wasabi at a medium intensity. The compound has been previously reported in horseradish as having green, chemical, and mustard like aromas [21], and it is known to activate the human Transient Receptor Potential Ankyrin 1 (TRPA1). This receptor is known to act in response to environmental irritants, and several ITCs identified in this study are known to activate it to varying degrees (isopropyl ITC, 3; isobutyl ITC, 10; allyl ITC, 5/6; 3-butenyl ITC, 11; 4-pentenyl ITC, 14; benzyl ITC, 22; phenylethyl ITC, 24; [72]). ...
... As discussed in previous sections, compounds such as methyl thiocyanate (2) may be produced from degradation of other hydrolysis products. Others such as the presence of sec-butyl ITC (9) in rocket, butyl ITC (12) in wasabi, and octyl ITC (21) in watercress cannot be so easily explained. There are several explanations with varying levels of likelihood: firstly, the most likely is that the ITCs and other GSL hydrolysis products have been identified incorrectly, and that they belong to other parent GSLs present within the analysed tissues. ...
Article
Full-text available
It is widely accepted that the distinctive aroma and flavour traits of Brassicaceae crops are produced by glucosinolate (GSL) hydrolysis products (GHPs) with other non-GSL derived compounds also reported to contribute significantly to their aromas. This study investigated the flavour profile and glucosinolate content of four Brassicaceae species (salad rocket, horseradish, wasabi, and watercress). Solid-phase microextraction followed by gas chromatography-mass spectrometry and gas chromatography-olfactometry were used to determine the volatile compounds and odorants present in the four species. Liquid chromatography-mass spectrometry was used to determine the glucosinolate composition, respectively. A total of 113 compounds and 107 odour-active components were identified in the headspace of the four species. Of the compounds identified, 19 are newly reported for ‘salad’ rocket, 26 for watercress, 30 for wasabi, and 38 for horseradish, marking a significant step forward in understanding and characterising aroma generation in these species. There were several non-glucosinolate derived compounds contributing to the ‘pungent’ aroma profile of the species, indicating that the glucosinolate-derived compounds are not the only source of these sensations in Brassicaceae species. Several discrepancies between observed glucosinolates and hydrolysis products were observed, and we discuss the implications of this for future studies.
... The present research is focused on horseradish, Armoracia rusticana, which belongs to the Brassicaceae family. The root of A. rusticana, a pungent spice, is used worldwide as a flavoring agent [12]. These roots contain various isothiocyanates, which are known for their antifungal, antibacterial, anticancer, and insecticidal properties [13][14][15][16]. ...
Article
Full-text available
Phenethyl isothiocyanate isolated from Armoracia rusticana root oil and its derivatives were tested at different doses in a bioassay designed to evaluate repellency against individual Haemaphysalis longicornis nymphs. Among the tested compounds, benzyl isothiocyanate exhibited repellency against H. longicornis nymphs at the lowest dose of 0.00625 mg/cm2, followed by phenethyl isothiocyanate (0.0125 mg/cm2) and phenyl isothiocyanate (0.025 mg/cm2). The behavioral responses of H. longicornis nymphs exposed to benzyl isothiocyanate and phenethyl isothiocyanate indicated that the mode of action of these compounds can be mainly attributed to the vapor phase. Encapsulated benzyl isothiocyanate showed repellency up to 120 min post-application at 0.1 mg/cm2, whereas pure benzyl isothiocyanate showed repellency up to 60 min post-application at 0.1 mg/cm2. The present study suggests that benzyl isothiocyanate is a potential repellent for protection against H. longicornis nymphs, and encapsulation in yeast cells may enhance the repellency effect.
... & Scherb.). This is a well-known species with a rich content of biologically active compounds (Agneta et al., 2013;Tomsone et al., 2020). It is a well-known plant with biological activities such as antioxidant (Calabrone et al., 2015;Ivanišová et al., 2020), and antimicrobial ones (Park et al., 2013) that is also widely used as a food plant with a pungent smell, intense lachrymatory odor and bitter taste (Rivelli et al., 2017). ...
Article
Full-text available
Background: The search for new plant raw material as a potential source of antioxidants is still ongoing. This study aimed to evaluate the antioxidant and antimicrobial capacity of the plant raw material of Crambe spp. during vegetation. Methods: The free radical scavenging activity and molybdenum reducing power of the extracts were used to determine antioxidant activity. The quantification of polyphenol compounds was conducted with Folin-Ciocalteu reagent. Flavonoids and phenolic acids were also determined. The disc diffusion method was used to determine antimicrobial activity. Results: It was determined that the free radical scavenging activity, assessed using the DPPH-method, was 4.38-8.20 mg TE/g DW, the molybdenum reducing power of the extracts was 40.07-129.12 mg TE/g DW, total polyphenol content was 20.24-70.88 mg GAE/g DW, total flavonoid content was 5.73-29.92 mg QE/g DW, and phenolic acid content was 3.00-10.63 mg CAE/g DW. Antimicrobial activity depended on the stage of growth and the part of the plant used. Conclusions: Crambe spp. possess the antioxidant and antimicrobial potential to mean that they could be used in pharmaceutical studies and the food industry.
Article
Crucifers have long been documented for their potential therapeutic food properties and are also referred to as the super-foods. They include the crops like kale, collards, kohlrabi, and brussels sprouts. These constitute the major source of secondary metabolites viz. flavonoids, anthocyanins, carotenoids, polyphenols, vitamins, minerals, coumarins, antioxidant enzymes, terpenes. Long-term and regular consumption of these vegetables helps to fight against obesity, cancer, atherosclerosis, inflammation, metabolic syndrome, and reduces the risk of several diseases. In the raw form, the availability and bioactivity of these phytonutrients have been reported to the highest levels. The scientific confirmation of these functional benefits though needs systematic clinical documentation and most of the research efforts in the last decade are focused in this direction. Further, processing affects the availability and activity of the bioactive compounds adversely thus, appropriate methods should be adopted for processing and cooking these vegetables. In order to promote their development and sustainable conservation, methods like micropropagation, cryopreservation, in vitro conservation, tissue culture, somatic hybridization, DNA banks, mycorrhization and genetic engineering shall be adopted. Several biotic and abiotic stresses have been reported to increase the availability of the secondary metabolites significantly which can be standardized to boost their functionality. Molecular breeding approaches such as marker-assisted selection, marker-assisted back cross, generation sequencing and gene editing also hold tremendous potential for improving the yield and quality of these super-foods. This review has been conceptualized to provide in-depth knowledge about the health benefits, effects of processing, strategies to improve and conserve the underutilized cruciferous vegetables, and the developments made in the area of production, processing, and value addition of the crucifers in the last decade.
Book
Horseradish roots, due to the content of many valuable nutrients and substances with healing and pro-health properties, are used more and more in medicine, food industry and cosmetics. In Poland, the cultivation of horseradish is considered minor crops. In addition, its limited size causes horseradish producers to encounter a number of unresolved agrotechnical problems. Infectious diseases developing on the leaves and roots during the long growing season reduce the size and quality of root crops. The small range of protection products intended for use in the cultivation of horseradish generates further serious environmental problems (immunization of pathogens, low effectiveness, deterioration of the quality of raw materials intended for industry, destruction of beneficial organisms and biodiversity). In order to meet the problems encountered by horseradish producers and taking into account the lack of data on: yielding, occurrence of infectious diseases and the possibility of combating them with methods alternative to chemical ones in the years 2012–2015, rigorous experiments have been carried out. The paper compares the impact of chemical protection and its reduced variants with biological protection on: total yield of horseradish roots and its structure. The intensification of infectious diseases on horseradish leaves and roots was analyzed extensively. Correlations were examined between individual disease entities and total yield and separated root fractions. A very important and innovative part of the work was to learn about the microbial communities involved in the epidemiology of Verticillium wilt of horseradish roots. The effect was examined of treatment of horseradish cuttings with a biological preparation (Pythium oligandrum), a chemical preparation (thiophanate-methyl), and the Kelpak SL biostimulator (auxins and cytokinins from the Ecklonia maxima algae) on the quantitative and qualitative changes occurring in the communities of these microorganisms. The affiliation of species to groups of frequencies was arranged hierarchically, and the biodiversity of these communities was expressed by the following indicators: Simpson index, Shannon–Wiener index, Shannon evenness index and species richness index. Correlations were assessed between the number of communities, indicators of their biodiversity and intensification of Verticillium wilt of horseradish roots. It was shown that the total yield of horseradish roots was on average 126 dt · ha–1. Within its structure, the main root was 56%, whereas the fraction of lateral roots (cuttings) with a length of more than 20 cm accounted for 26%, and those shorter than 20 cm for 12%, with unprofitable yield (waste) of 6%. In the years with higher humidity, the total root yield was higher than in the dry seasons by around 51 dt · ha–1 on average. On the other hand, the applied protection treatments significantly increased the total yield of horseradish roots from 4,6 to 45,3 dt · ha–1 and the share of fractions of more than 30 cm therein. Higher yielding effects were obtained in variants with a reduced amount of foliar application of fungicides at the expense of introducing biopreparations and biostimulators (R1, R2, R3) and in chemical protection (Ch) than in biological protection (B1, B2) and with the limitation of treatments only to the treatment of cuttings. The largest increments can be expected after treating the seedlings with Topsin M 500 SC and spraying the leaves: 1 × Amistar Opti 480 SC, 1 × Polyversum WP, 1 × Timorex Gold 24 EC and three times with biostimulators (2 × Kelpak SL + 1 × Tytanit). In the perspective of the increasing water deficit, among the biological protection methods, the (B2) variant with the treatment of seedlings with auxins and cytokinins contained in the E. maxima algae extract is more recommended than (B1) involving the use of P. oligandrum spores. White rust was the biggest threat on horseradish plantations, whereas the following occurred to a lesser extent: Phoma leaf spot, Cylindrosporium disease, Alternaria black spot and Verticillium wilt. In turn, on the surface of the roots it was dry root rot and inside – Verticillium wilt of horseradish roots. The best health of the leaves and roots was ensured by full chemical protection (cuttings treatment + 6 foliar applications). A similar effect of protection against Albugo candida and Pyrenopeziza brassicae was achieved in the case of reduced chemical protection to one foliar treatment with synthetic fungicide, two treatments with biological preparations (Polyversum WP and Timorex Gold 24 EC) and three treatments with biostimulators (2 × Kelpak SL, 1 × Tytanit). On the other hand, the level of limitation of root diseases comparable with chemical protection was ensured by its reduced variants R3 and R2, and in the case of dry root rot, also both variants of biological protection. In the dry years, over 60% of the roots showed symptoms of Verticillium wilt, and its main culprits are Verticillium dahliae (37.4%), Globisporangium irregulare (7.2%), Ilyonectria destructans (7.0%), Fusarium acuminatum (6.7%), Rhizoctonia solani (6.0%), Epicoccum nigrum (5.4%), Alternaria brassicae (5.17%). The Kelpak SL biostimulator and the Polyversum WP biological preparation contributed to the increased biodiversity of microbial communities associated with Verticillium wilt of horseradish roots. In turn, along with its increase, the intensification of the disease symptoms decreased. There was a significant correlation between the richness of species in the communities of microbial isolates and the intensification of Verticillium wilt of horseradish roots. Each additional species of microorganism contributed to the reduction of disease intensification by 1,19%.
Book
Full-text available
Protective treatments shaping the yielding, healthiness and diversity of microorganisms associated with Verticillium wilt of horseradish roots (Armoracia rusticana Gaertn.) (PDF) Zabiegi ochronne kształtujące plonowanie, zdrowotność oraz różnorodność mikroorganizmów związanych z czernieniem pierścieniowym korzeni chrzanu (Armoracia rusticana Gaertn.). Available from: https://www.researchgate.net/publication/354764440_Zabiegi_ochronne_ksztaltujace_plonowanie_zdrowotnosc_oraz_roznorodnosc_mikroorganizmow_zwiazanych_z_czernieniem_pierscieniowym_korzeni_chrzanu_Armoracia_rusticana_Gaertn#fullTextFileContent [accessed Sep 22 2021]. Summary Horseradish roots, due to the content of many valuable nutrients and substances with healing and pro-health properties, are used more and more in medicine, food industry and cosmetics. In Poland, the cultivation of horseradish is considered minor crops. In addition, its limited size causes horseradish producers to encounter a num�ber of unresolved agrotechnical problems. Infectious diseases developing on the leaves and roots during the long growing season reduce the size and quality of root crops. The small range of protection products intended for use in the cultivation of horseradish generates further serious environmental problems (immunization of pathogens, low effectiveness, deterioration of the quality of raw materials intended for industry, destruction of beneficial organisms and biodiversity). In order to meet the problems encountered by horseradish producers and tak�ing into account the lack of data on: yielding, occurrence of infectious diseases and the possibility of combating them with methods alternative to chemical ones in the years 2012–2015, rigorous experiments have been carried out. The paper compares the impact of chemical protection and its reduced variants with biological protection on: total yield of horseradish roots and its structure. The intensification of infectious diseases on horseradish leaves and roots was analyzed extensively. Correlations were examined between individual disease entities and total yield and separated root fractions. A very important and innovative part of the work was to learn about the microbial communities involved in the epidemiology of Verticillium wilt of horseradish roots. The effect was examined of treatment of horseradish cuttings with a biological preparation (Pythium oligandrum), a chemical preparation (thiophanate-methyl), and the Kelpak SL biostimulator (auxins and Summary 135 cytokinins from the Ecklonia maxima algae) on the quantitative and qualitative changes occurring in the communities of these microorganisms. The affiliation of species to groups of frequencies was arranged hierarchically, and the biodiversity of these communities was expressed by the following indicators: Simpson index, Shannon–Wiener index, Shannon evenness index and species richness index. Correlations were assessed between the number of communities, indicators of their biodiversity and intensification of Verticillium wilt of horseradish roots. It was shown that the total yield of horseradish roots was on average 126 dt · ha–1. Within its structure, the main root was 56%, whereas the fraction of lat�eral roots (cuttings) with a length of more than 20 cm accounted for 26%, and those shorter than 20 cm for 12%, with unprofitable yield (waste) of 6%. In the years with higher humidity, the total root yield was higher than in the dry seasons by around 51 dt · ha–1 on average. On the other hand, the applied protection treatments sig�nificantly increased the total yield of horseradish roots from 4,6 to 45,3 dt · ha–1 and the share of fractions of more than 30 cm therein. Higher yielding effects were obtained in variants with a reduced amount of foliar application of fungicides at the expense of introducing biopreparations and biostimulators (R1, R2, R3) and in chemical protection (Ch) than in biological protection (B1, B2) and with the limitation of treatments only to the treatment of cuttings. The largest increments can be expected after treating the seedlings with Topsin M 500 SC and spraying the leaves: 1 × Amistar Opti 480 SC, 1 × Polyversum WP, 1 × Timorex Gold 24 EC and three times with biostimulators (2 × Kelpak SL + 1 × Tytanit). In the perspective of the increasing water deficit, among the bio�logical protection methods, the (B2) variant with the treatment of seedlings with auxins and cytokinins contained in the E. maxima algae extract is more recom�mended than (B1) involving the use of P. oligandrum spores. White rust was the biggest threat on horseradish plantations, whereas the fol�lowing occurred to a lesser extent: Phoma leaf spot, Cylindrosporium disease, Alternaria black spot and Verticillium wilt. In turn, on the surface of the roots it was dry root rot and inside – Verticillium wilt of horseradish roots. The best health of the leaves and roots was ensured by full chemical protection (cuttings treatment + 6 foliar applications). A similar effect of protection against Albugo candida and Pyrenopeziza brassicae was achieved in the case of reduced chemical protection to one foliar treatment with synthetic fungicide, two treatments with biological prepa�rations (Polyversum WP and Timorex Gold 24 EC) and three treatments with bi�ostimulators (2 × Kelpak SL, 1 × Tytanit). On the other hand, the level of limitation of root diseases comparable with chemical protection was ensured by its reduced variants R3 and R2, and in the case of dry root rot, also both variants of biological protection. In the dry years, over 60% of the roots showed symptoms of Verticillium wilt, and its main culprits are Verticillium dahliae (37.4%), Globisporangium irregu�lare (7.2%), Ilyonectria destructans (7.0%), Fusarium acuminatum (6.7%), Rhizoctonia 136 Zabiegi ochronne kształtujące plonowanie, zdrowotność oraz różnorodność mikroorganizmów... solani (6.0%), Epicoccum nigrum (5.4%), Alternaria brassicae (5.17%). The Kelpak SL biostimulator and the Polyversum WP biological preparation contributed to the increased biodiversity of microbial communities associated with Verticillium wilt of horseradish roots. In turn, along with its increase, the intensification of the disease symptoms decreased. There was a significant correlation between the richness of species in the communities of microbial isolates and the intensification of Verticillium wilt of horseradish roots. Each additional species of microorganism contributed to the reduction of disease intensification by 1,19%
Article
To enhance the quality or, in other words, increase the variety of horseradish sauce, two-step fermentation using lactic acid bacteria (LAB) and Saccharomyces cerevisiae is proposed for the production of the sauce. Total phenolics, acids and amino acid nitrogen contents were determined and used to assess the fermentation efficiency, while antioxidant activities, profiles of amino acids and volatile flavor compounds was used to indicate the quality of the sauce. Molecular structure determination as well as electronic tongue and sensory analyses were also conducted. Adding S. cerevisiae after 48 h of LAB fermentation resulted in higher fermentation efficiency. The sauce was noted to be fermented more fully, with increased content of amino acids by 23.10% compared with unfermented samples; its DPPH radical scavenging ability and reducing power increased by 26.26% and 33.33%, respectively. The contents of alcohols, esters and acids increased in the sauce. The electronic tongue and sensory evaluation showed that the sauce had good sensory quality.
Article
Full-text available
This paper is an ethnobotanical review of wild edible plants gathered for consumption from the end of the 18th century to the present day, within the present borders of Poland. 42 ethnographic and botanical sources documenting the culinary use of wild plants were analyzed. The use of 112 species (3.7% of the flora) has been recorded. Only half of them have been used since the 1960s. Three species: Cirsium rivulare, Euphorbia peplus and Scirpus sylvaticus have never before been reported as edible by ethnobotanical literature. The list of wild edible plants which are still commonly gathered includes only two green vegetables (Rumex acetosa leaves for soups and Oxalis acetosella as children's snack), 15 folk species of fruits and seeds (Crataegus spp., Corylus avellana, Fagus sylvatica, Fragaria vesca, Malus domestica, Prunus spinosa, Pyrus spp., Rosa canina, Rubus idaeus, Rubus sect. Rubus, Sambucus nigra, Vaccinium myrtillus, V. oxycoccos, V. uliginosum, V. vitis-idaea) and four taxa used for seasoning or as preservatives (Armoracia rusticana root and leaves, Carum carvi seeds, Juniperus communis pseudo-fruits and Quercus spp. leaves). The use of other species is either forgotten or very rare. In the past, several species were used for food in times of scarcity, most commonly Chenopodium album, Urtica dioica, U. urens, Elymus repens, Oxalis acetosella and Cirsium spp., but now the use of wild plants is mainly restricted to raw consumption or making juices, jams, wines and other preserves. The history of the gradual disappearance of the original barszcz, Heracleum sphondylium soup, from Polish cuisine has been researched in detail and two, previously unpublished, instances of its use in the 20th century have been found in the Carpathians. An increase in the culinary use of some wild plants due to media publications can be observed. Poland can be characterized as a country where the traditions of culinary use of wild plants became impoverished very early, compared to some parts of southern Europe. The present use of wild plants, even among the oldest generation, has been almost entirely restricted to fruits.
Article
Full-text available
The fumigant toxicity of essential oil from horseradish plant, Armoracia rusticana (L.), was assessed against Plodia interpunctella (Hübner) and Sitophilus zeamais Motschulsky. A. rusticana oil was active against different life stages of P. interpunctella and adults of S. zeamais. The LC50 value for adults was the lowest and that of pupa was the highest. The major compound found by gas chromatography-mass spectrometry was allyl isothiocyanate (97.81%). These results indicate that it may be possible to achieve toxicity levels similar to those of standard chemical fumigants through the applications of essential oils from A. rusticana.
Article
Full-text available
Glucosinolates are sulfur- and nitrogen-containing plant secondary metabolites common in the Brassicaceae and related plant families. In the plant, they coexist with an endogenous β-thioglucosidase (EC 3.2.3.1) called myrosinase, though glucosinolates are stored in the vacuoles of so-called S-cells and myrosinase in separate but adjacent cells. Upon plant tissue disruption, glucosinolates are released at the damage site and become hydrolyzed by myrosinase. The chemical nature of the hydrolysis products depends on the structure of the glucosinolate side chain, plant species and reaction conditions. Biosynthesis of glucosinolates comprises three phases: (i) amino acid chain elongation, in which additional methylene groups are inserted into the side chain, (ii) conversion of the amino acid moiety to the glucosinolate core structure, (iii) and subsequent side chain modifications. Glucosinolate pattern differs between species and ecotype as well as between and even within individual plants, depending on developmental stage, tissue and photoperiod. A number of environmental conditions such as light plant, nutritional status, fungal infection, wounding and insect damage can alter the glucosinolate pattern significantly. The change of the glucosinolate profile by several environmental factors has brought forward different theories regarding their potential roles in the plant. However, the most accepted theory is that the glucosinolate-myrosinase system is involved in defense against herbivores and pathogens. This review summarized recent progress in glucosinolate biosynthesis, degradation and organization of the myrosinase-glucosinolate system. Furthermore, current knowledge of the potential role of glucosinolates in the plant, especially in plant defense, is discussed.
Book
Alphonse de Candolle (1806-93) was a French-Swiss botanist who was an important figure in the study of the origins of plants and the reasons for their geographic distribution. He also created the first Code of Botanical Nomenclature. Despite initially studying law, he took over both the chair of botany at the University of Geneva, and the directorship of Geneva's botanical gardens from his father Augustin de Candolle (1778-1841). He published numerous botanical books, and edited ten volumes of the Prodromus, a seventeen-volume reference text intended to cover the key properties of all known seed plants. This work, reissued in the second edition of the English translation of 1886, is his most famous and influential book, tracing the geographic origins of plants known to have been cultivated by humans. It is one of the earliest studies of the history of crop domestication, and an important contribution to phytogeography.